pursuant to French law, the sections of theBy-laws relating to the number of directors and election and removal of a director from office may only be modified by a resolution adopted by atwo-thirds majority vote of our shareholders present, represented by a proxy or voting by mail at the meeting.

Holders of our ADSs do not directly hold our ordinary shares.

As an ADS holder, you are not treated as one of our shareholders and you do not have ordinary shareholder rights. French law governs shareholder rights. The depositary, Citibank, N.A., is the holder of the ordinary shares underlying your ADSs. As a holder of ADSs, you have ADS holder rights. The deposit agreement among us, the depositary and you, as an ADS holder, and all other persons directly and indirectly holding ADSs, sets out ADS holder rights, as well as the rights and obligations of the depositary.

Holders of our ADSs may not be able to exercise their right to vote the ordinary shares underlying such ADSs.

Holders of ADSs may exercise voting rights with respect to the ordinary shares represented by the ADSs only in accordance with the provisions of the deposit agreement and not as a direct shareholder. The deposit agreement provides that, upon receipt of notice of any meeting of holders of our ordinary shares, the depositary will fix a record date for the determination of ADS holders who shall be entitled to give instructions for the exercise of voting rights. Upon timely receipt of notice from us, if we so request, the depositary shall distribute to the holders as of the record date (1) the notice of the meeting or solicitation of consent or proxy sent by us and (2) a statement as to the manner in which instructions may be given by the holders.

Holders of ADSs may instruct the depositary of the ADSs to vote the ordinary shares underlying such ADSs. Otherwise, holders of our ADSs will not be able to exercise their right to vote, unless they withdraw the ordinary shares underlying such ADSs. However, holders of our ADSs may not know about the meeting far enough in advance to withdraw those ordinary shares. If we ask for instructions, the depositary, upon timely notice from us, will notify holders of our ADSs of the upcoming vote and arrange to deliver our voting materials to such holders. We cannot guarantee that holders of our ADSs will receive the voting materials in time to ensure that they can instruct the depositary to vote such ordinary shares or to withdraw such ordinary shares so as to vote them directly. If the depositary does not receive timely voting instructions from holders of our ADSs, it may give a proxy to a person designated by us to vote the ordinary shares underlying such ADSs in accordance with the recommendation of our board of directors. In addition, the depositary and its agents are not responsible for failing to carry out voting instructions or for the manner of carrying out voting instructions. This means that holders of our ADSs may not be able to exercise their right to vote, and there may be nothing such holders can do if the ordinary shares underlying such ADSs are not voted as requested.

The right of holders of our ADSs to participate in any future preferential subscription rights or to elect to receive dividends in shares may be limited, which may cause dilution to holders of ADSs.

According to French law, if we issue additional shares or securities for cash giving right, immediately or in the future, to new shares, current shareholders will have preferential subscription rights for these securities proportionally to their shareholding in our company unless they waive those rights at an extraordinary meeting of our shareholders (by atwo-thirds majority vote) or individually by each shareholder. However, our ADS holders in the United States will not be entitled to exercise or sell such rights unless we register the rights and the securities to which the rights relate under the Securities Act or an exemption from the registration requirements is available. In addition, the deposit agreement for our ADSs provides that the depositary will not make rights available to holders of our ADSs unless the distribution to ADS holders of both the rights and any related securities are either registered under the Securities Act or exempted from registration under the Securities Act. Further, if we offer holders of our ordinary shares the option to receive dividends in either cash or shares, under the deposit agreement the depositary may require satisfactory assurances from us that extending the offer to holders of ADSs does not require registration of any securities under the Securities Act before making the option available to holders of ADSs. We are under no obligation to file a registration statement with respect to any such

rights or securities or to endeavor to cause such a registration statement to be declared effective. Moreover, we may not be able to establish an exemption from registration under the Securities Act. Accordingly, ADS holders may be unable to participate in our rights offerings or to elect to receive dividends in shares and may experience dilution in their holdings. In addition, if the depositary is unable to sell rights that are not exercised or not distributed or if the sale is not lawful or reasonably practicable, it will allow the rights to lapse, in which case holders of our ADSs will receive no value for these rights.

Holders of our ADSs may be subject to limitations on the transfer of such ADSs and the withdrawal of the underlying ordinary shares.

ADSs, which may be evidenced by American Depositary Receipts, or ADRs, are transferable on the books of the depositary. However, the depositary may close its books at any time or from time to time when it deems expedient in connection with the performance of its duties. The depositary may refuse to deliver, transfer or register transfers of ADSs generally when our books or the books of the depositary are closed, or at any time if we or the depositary think it is advisable to do so because of any requirement of law, government or governmental body, or under any provision of the deposit agreement, or for any other reason subject to an ADS holders’ right to cancel such ADSs and withdraw the underlying ordinary shares. Temporary delays in the cancellation of such ADSs and withdrawal of the underlying ordinary shares may arise because the depositary has closed its transfer books or we have closed our transfer books, the transfer of ordinary shares is blocked to permit voting at a shareholders’ meeting or we are paying a dividend on our ordinary shares. In addition, holders of our ADSs may not be able to cancel such

ADSs and withdraw the underlying ordinary shares when such holders owe money for fees, taxes and similar charges and when it is necessary to prohibit withdrawals in order to comply with any laws or governmental regulations that apply to ADSs or to the withdrawal of ordinary shares or other deposited securities.

As a foreign private issuer, we are exempt from a number of rules under the U.S. securities laws and are permitted to file less information with the SEC than a U.S. company. This may limit the information available to holders of ADSs.

We are a “foreign private issuer,” as defined in the SEC’s rules and regulations and, consequently, we are not subject to all of the disclosure requirements applicable to public companies organized within the United States. For example, we are exempt from certain rules under the Exchange Act that regulate disclosure obligations and procedural requirements related to the solicitation of proxies, consents or authorizations applicable to a security registered under the Exchange Act, including the U.S. proxy rules under Section 14 of the Exchange Act. In addition, our officers and directors are exempt from the reporting and “short-swing” profit recovery provisions of Section 16 of the Exchange Act and related rules with respect to their purchases and sales of our securities. Moreover, while we currently make annual and quarterly filings with the SEC, we are not required to file periodic reports and financial statements with the SEC as frequently or as promptly as U.S. domestic public companies and are not required to file quarterly reports on Form10-Q or current reports on Form8-K under the Exchange Act. Accordingly, there may be less publicly available information concerning our company than there would be if we were a U.S. domestic issuer.

As a foreign private issuer, we follow certain home country practices in relation to corporate governance matters that differ significantly from Nasdaq corporate governance standards. These practices may afford less protection to shareholders than they would enjoy if we complied fully with Nasdaq’s corporate governance standards.

As a foreign private issuer listed on the Nasdaq Global Market, we are subject to corporate governance standards. However, Nasdaq’s rules permit a foreign private issuer like us to follow the corporate governance practices of its home country. Certain corporate governance practices in France, which is our home country, may differ significantly from corporate governance standards of the Nasdaq. For example, neither the corporate laws of France nor ourBy-laws require a majority of our directors to be independent and our independent directors are

not required to hold regularly scheduled meetings at which only independent directors are present. In addition, home country practice in France does not require us to maintain a nominating and corporate governance committee or to maintain a compensation committee composed entirely of independent directors. Currently, we follow home country practice in certain key respects. Therefore, our shareholders may be afforded less protection than they otherwise would have under corporate governance listing standards applicable to U.S. domestic issuers. A discussion of our corporate governance practices is set forth in the section titled “Management—Corporate Governance Practices.”

We may lose our foreign private issuer status in the future, which could result in significant additional cost and expense.

While we currently qualify as a foreign private issuer, the determination of foreign private issuer status is made annually on the last business day of our most recently completed second fiscal quarter and, accordingly, the next determination will be made with respect to us on June 30, ~~2018.~~2019.

In the future, we would lose our foreign private issuer status if we to fail to meet the requirements necessary to maintain our foreign private issuer status as of the relevant determination date. For example, if more than 50% of our securities are held by U.S. residents and more than 50% of our executive officers or members of our board of directors are residents or citizens of the United States, we could lose our foreign private issuer status.

The regulatory and compliance costs to us under U.S. securities laws as a U.S. domestic public company would be significantly more than costs we currently incur as a foreign private issuer. If we lost our foreign private issuer status, we would be required to file periodic reports on Form10-Q and current reports on Form8-K, to file periodic reports and registration statements on U.S. domestic issuer forms with the SEC, which are more detailed and extensive in certain respects than the forms available to a foreign private issuer. We would be required under current SEC rules to prepare our financial statements in accordance with U.S. GAAP, rather than IFRS, in U.S. dollars rather than euros. Such conversion of our financial statements to U.S. GAAP would involve significant time and cost. In addition, we would lose our ability to rely upon exemptions from certain corporate governance requirements of the Nasdaq that are available to foreign private issuers, such as the ones described above, and we would be required to modify certain of our policies to comply with corporate governance practices associated with U.S. domestic issuers. Moreover, we would lose our ability to rely upon exemptions from procedural requirements related to the solicitation of proxies.

It may be difficult to enforce civil liabilities against our company and directors and senior management and the experts named in this Annual Report.

Certain members of our board of directors and senior management and those of our subsidiaries, arenon-residents of the United States, and all or a substantial portion of our assets and the assets of such persons are located outside the United States. As a result, it may not be possible to serve process on such persons or us in the United States or to enforce judgments obtained in U.S. courts against them or us based on civil liability provisions of the securities laws of the United States. Additionally, it may be difficult to assert U.S. securities law claims in actions originally instituted outside of the United States. Foreign courts may refuse to hear a U.S. securities law claim because foreign courts may not be the most appropriate forums in which to bring such a claim.

Even if a foreign court agrees to hear a claim, it may determine that the law of the jurisdiction in which the foreign court resides, and not U.S. law, is applicable to the claim. Further, if U.S. law is found to be applicable, the content of applicable U.S. law must be proved as a fact, which can be a time-consuming and costly process, and certain matters of procedure would still be governed by the law of the jurisdiction in which the foreign court resides. In particular, there is some doubt as to whether French courts would recognize and enforce certain civil liabilities under U.S. securities laws in original actions or judgments of U.S. courts based upon these civil liability provisions. In addition, awards of punitive damages in actions brought in the United States or elsewhere may be unenforceable in France. An award for monetary damages under the U.S. securities laws would be considered punitive if it does not seek to compensate the claimant for loss or damage suffered but is intended to

punish the defendant. French law provides that a shareholder, or a group of shareholders, may initiate a legal action to seek indemnification from the directors of a corporation in the corporation’s interest if it fails to bring such legal action itself. If so, any damages awarded by the court are paid to the corporation and any legal fees relating to such action are borne by the relevant shareholder or the group of shareholders.

The enforceability of any judgment in France will depend on the particular facts of the case as well as the laws and treaties in effect at the time. The United States and France do not currently have a treaty providing for recognition and enforcement of judgments (other than arbitration awards) in civil and commercial matters.

Provisions in our collaboration agreement with Servier may prevent or delay a change in control.

Our collaboration agreement with Servier provides that if any third party begins to control us, directly or indirectly, by any means, or in the event that we engage in a change of control transaction, including, but not limited to, the sale of all or substantially all of our assets or all or substantially all of our assets that are material to the performance of our obligations under the collaboration agreement, then Servier has the right tobuy-out our interest in thepre-candidate products, product candidates, and products as described under the collaboration agreement. We refer to this right to acquire such interest as thebuy-out. In the event we fail to agree with Servier on the amount of payment for our interest in thepre-candidate products, product candidates or products within twenty days following Servier’s provision of abuy-out notice, then thebuy-out payment would be determinedby-third party valuators.

Thebuy-out may have the effect of delaying or preventing a change in control transaction involving us, or may reduce the number of companies interested in acquiring us. If Servier were to exercise thebuy-out, it would gain exclusive development and marketing rights to thepre-candidate products, product candidates and products developed under the collaboration agreement. Were this to happen, our successor would not receive milestone payments or royalty payments on net sales of any of the products sold to Servier in connection with thebuy-out. These provisions could have the effect of delaying or preventing a change in control transaction involving Cellectis, or could reduce the number of companies interested in acquiring us.

The rights of shareholders in companies subject to French corporate law differ in material respects from the rights of shareholders of corporations incorporated in the United States.

We are a French company with limited liability. Our corporate affairs are governed by ourBy-laws and by the laws governing companies incorporated in France. The rights of shareholders and the responsibilities of members of our board of directors are in many ways different from the rights and obligations of shareholders in companies governed by the laws of U.S. jurisdictions. For example, in the performance of its duties, our board of directors is required by French law to consider the interests of our company, its shareholders, its employees and other stakeholders, rather than solely our shareholders and/or creditors. It is possible that some of these parties will have interests that are different from, or in addition to, the interests of our shareholders. See the sections of this Annual Report titled “Memorandum and Articles of Association” and “Corporate Governance.”

ITEM 4.

INFORMATION ON THE COMPANY

A.	History and Development of the Company

Our legal and commercial name is Cellectis S.A. We were incorporated as asociété anonyme, or S.A., under the laws of the French Republic on January 4, 2000 for a period of 99 years. We are registered at the Paris Registre du Commerce et des Sociétés under the number 428 859 052. Our principal executive offices are located at 8, rue de la Croix Jarry, 75013 Paris, France, and our telephone number is +33 1 81 69 16 00. Our agent for service of process in the United States is Cellectis, Inc. located at 430 East 29^th29th Street, New York, New York 10016. We also maintain a website at www.cellectis.com. The reference to our website is an inactive textual reference only and the information contained in, or that can be accessed through, our website is not a part of this Annual Report.

~~Our actual capital expenditures~~The acquisition of intangible assets, property, plant and ~~additions to intangible assets~~equipment for the years ended December 31, ~~2015,~~ 2016, 2017 and ~~2017~~2018 together amounted to ~~$4.4million,~~ $14.6 million, $2.6 million and ~~$2.6~~$4.9 million, respectively. These expenditures primarily consisted of the acquisitions of industrial and laboratory equipment and fittings required to conduct our research ~~programs.~~programs, the improvements of Calyxt’s and Cellectis’ sites and the first investments to start the construction of our new manufacturing facilities in Paris and in the United States. We expect our capital expenditures to increase in absolute terms in the near term as we continue to advance our research and development programs and grow our operations. We anticipate our capital expenditure in ~~2018~~2019 to be financed from our cash and cash equivalents on hand. Primarily, these capital expenditures will be made both in France and in the United States, where our research and development facilities are currently located.

On January 21, 2015, we signed a lease for an administrative and research facility in New York, New York to enhance our presence in the United States. This facility, which includes largestate-of-the-art research laboratories, opened on April 8, 2015, and essentially supports ~~the development of~~ our~~CAR-T~~ ~~pipeline.~~ innovation activities. In March 2016, we entered into a lease for a 26,928 square-foot space in Montvale, New Jersey, which we intend to discontinue before its scheduled expiration in September 2026.

In March 2016, Calyxt acquired10-acres in 2 parcels in Roseville, Minnesota, and a headhouse and a greenhouse were built and they have been in operations since September 2016. In the third quarter of 2017, Calyxt entered into a sale and lease-back agreement related to these land and buildings. See Item 4D for further information.

In March 2019, we entered into a lease agreement for a 82,000 square foot commercial-scale manufacturing facility, called the IMPACT site, which stands for “Innovative Manufacturing Plant for Allogeneic Cellular Therapies”. The IMPACT facility is located in Raleigh, North Carolina. The new manufacturing facility is being designed to provide GMP manufacturing for clinical supply and commercial product upon potential regulatory approval. The facility is planned to be operational by 2021.

For information on the SEC’s website and our website, please refer to “Item 10.H.Documents on Display”.

B.	Business Overview

We are a clinical stage biotechnological company, employing our core proprietary technologies to developbest-in-class products in the field of immuno-oncology. Our product candidates, based on gene-editedT-cells that express chimeric antigen receptors, or CARs, seek to harness the power of the immune system to target and eradicate cancer cells. We believe thatCAR-based immunotherapy is one of the most promising areas of cancer research, representing a new paradigm for cancer treatment. We are designing next-generation immunotherapies that are based on gene-edited CART-cells. Our gene-editing technologies allow us to create allogeneic CART-cells, meaning they are derived from healthy donors rather than the patients themselves. We believe that the production of allogeneic CART-cells will allow us to develop cost-effective,off-the-shelf products that are capable of being cryopreserved, stored and distributed worldwide. Our gene-editing expertise also enables us to develop product candidates that feature additional safety and efficacy attributes, including control properties designed to prevent them from attacking healthy tissues, to enable them to tolerate standard oncology treatments, and to equip them to resist mechanisms that inhibit immune-system activity. In addition to our focus on immuno-oncology, we are exploring the use of our gene-editing technologies in other therapeutic applications, as well as to develop healthier food products for a growing population.

Cancer is the second-leading cause of death in the United States and accounts for approximately one in four deaths. Immuno-oncology seeks to harness the power of the body’s immune system to target and kill cancer. A key to this effort is a type of white blood cell known as theT-cell, which plays an important role in identifying and killing cancer cells. Unfortunately, cancer cells often develop mechanisms to evade the immune system. CARs, which are engineered receptors that can be expressed on the surface ofT-cells, provide theT-cells with a specific targeting mechanism, thereby enhancing its ability to seek, identify, interact with and destroy tumor cells bearing a selected antigen. Research and development of CART-cell immunotherapies currently focuses on two approaches: autologous and allogeneic therapies. Autologous CART-cell immunotherapies modify a patient’s ownT-cells to target specific antigens that are located on cancer cells. This type of therapy requires an individualized immunotherapy product for each patient and is currently being tested in clinical trials by several academic institutions, ~~and~~as well as biotechnology and pharmaceutical companies. In contrast, an allogeneic CART-cell immunotherapy is an approach by which a cancer patient is infused with a mass-produced,off-the-shelf immunotherapy product derived from a healthyT-cell donor. Our initial focus is on developing allogeneic treatments, and we believe that we are the leading company pursuing this approach.

Limitations of Current Autologous Treatments and Key Benefits of our UCART approach

68Many of the CART-cell immunotherapy treatments currently under development are created through an autologous approach in which the patient’s ownT-cells are engineered to fight cancer cells. Part of our scientific basis for pursuing allogeneic approaches rests in the recognized limitations of autologous approaches, including:

Autologous treatments must be specifically manufactured for each patient and the resulting engineered cells may have different properties due to significantpatient-to-patient variability in the quality of theT-cell;

•

Autologous treatments can bear high costs due to the necessity of producing a bespoke treatment for each patient and the effort consumed in modifying and growing each patient’sT-cells (by way of example in the United States, the list price of Kymriah^® for pediatric use is $475,000, and the list price of Yescarta^® and Kymriah^® for adult use is $373,000); and

At this time, autologous treatments cannot be mass produced, may involve significant delay in production time if the number of patients exceeds the number of productions that can be made in parallel, and require patients be treated at select advanced facilities.

Although some autologous approaches to CART-cell have recently demonstrated encouraging clinical data, we believe ourCAR-T approach and manufacturing process has the potential to provide the following benefits:

•

Market access. Enable products to be shipped globally, thereby reducing deployment obstacles and providing accessibility to a broad patient population;

•

Cost-effectiveness and Scalable Manufacturing. Streamlined manufacturing process has the potential to reduce costs, with approximatively hundreds of doses per batch;

•

Novel Features. Develop products with specific safety and control properties, through a CAR linked to a suicide switch;

•

Engraftment. Avoid graft-versus-host disease (GvHD) through the inactivation of theT-cell receptor (TCR).

•

Persistence. Manage rejection and persistence of the UCART product candidate, through notably the option to inactivate CD52 and beta2-microglobulin (ß2M) genes respectively;

A key enabler of the allogeneic approach is our gene editing technology, relying on a particular class of proteins derived from transcription activator-like effectors fused to the nuclease domain of a type II restriction endonuclease (TALEN). Gene editing is a type of genetic engineering in which DNA is inserted, deleted, repaired or replaced from a precise location in the genome. The most fundamental challenge of gene editing is the need to specifically and efficiently target a precise DNA sequence within a gene. Our proprietary nuclease-based gene-editing technologies, combined with 18almost 20 years of genome engineering experience, allow us to edit any gene with highly precise insertion, deletion, repair and replacement of DNA sequences. Our nucleases, including ~~a particular class of proteins derived from transcription activator-like effectors~~TALEN, act like DNA scissors to edit genes at precise target sites and allow us to design allogeneic CART-cells. Our patented PulseAgile electroporation technology allows us to efficiently deliver our clinical grade nucleases into human cells while preserving cell viability, making it particularly well-suited for a large-scale manufacturing process. We believe these technologies will enable our ~~clinical grade~~clinical-grade drug therapeutic products to be manufactured, cryopreserved, stored, distributed broadly and infused into patients in anoff-the-shelf approach.

Our candidate products

We are developing ~~products~~product candidates internally and through strategic alliances with ~~Pfizer~~Allogene Therapeutics, Inc. (“Allogene”) and ~~Servier.~~Les Laboratoires Servier (“Servier”). We believe that our alliances with ~~Pfizer~~Allogene and Servier have validated our technology platform, our strong expertise in the allogeneic CART-cells field and the strength of our intellectual property portfolio. Our strategic alliances include potential milestone payments to us of up to $3.9 billion and royalties on future sales.

Under the Allogene License Agreement (see below), Allogene has exclusive rights to pursue development and commercialization of products for a total of fifteen targets of their choice, which have been selected, including BCMA(ALLO-715), FLT3(ALLO-819), CD70, and DLL3.

In 2016, Servier commenced two Phase I clinical studies for UCART19, one in adult Acute Lymphoblastic Leukemia (ALL), the CALM study, and one in pediatric ALL, the PALL study. The CALM study is commenced in the United Kingdom, the United States, and France, and the PALL study is commenced in the United Kingdom, Belgium and France. We refer in this Annual Report to the CALM and the PALL studies, collectively, as the UCART19 Clinical Studies. In November 2015, when we exclusively licensed the rights to UCART19 to Servier, Servier also announced that it had granted Pfizer (subsequently transferred to Allogene) the exclusive rights for the development and the commercialization of UCART19 in the United States. Consequently, Servier’s CALM study in the United States is conducted in collaboration with ~~Pfizer.~~Allogene. In ~~addition to early data~~December 2018, Servier presented ~~by Servier and Pfizer during a meeting at the National Institutes of Health’s Recombinant DNA Advisory Committee (or “RAC”) held on December 14, 2016, in December 2017, Servier presented~~pooled intermediate results from the UCART19 Clinical Studies during the American Society for Hematology (ASH) annual conference. ~~Such intermediate results show 83%~~After UCART19 infusion, 82% (14/17) of patients who received a lymphodepletion regimen (consisting of fludarabine, cyclophosphamide and alemtuzumab, an anti-CD52 monoclonal antibody) achieved a complete remission, ~~rate across~~or “CR”, or complete remission with incomplete blood cell recovery (or “Cri”) by day 28 or day 42 after infusion. Within responder patients, 71% (10/14) of them were ‘minimum residual disease’ (MRD) negative(MRD- stands for less than 1 leukemic cell among 10E4 normal cells) assessed by flow or qPCR. When considering all treated patients, 67% (14/21) of them did achieve CR/CRi. Regarding safety considerations, there was no serious adverse events (grade³3) for graft versus host disease (GvHD) and neurological events. Grade3-4 toxicities did only regard events of cytokine release syndrome (14%, 3/21), prolonged cytopenia (29%, 6/21) and viral infections (24%, 5/21). In January 2019, Allogene announced, in collaboration with Servier, that the ~~two studies: five out of seven patients achieving minimal residual disease (MRD) negativity~~Food and Drug Administration (FDA) approved the Investigational New Drug (IND) for Phase 1 clinical study forALLO-501, corresponding to UCART19 which we exclusively licensed to Servier and which was sublicensed to Allogene by Servier in US, in relapsed/refractoryNon-Hodgkin Lymphoma (NHL, the CALM study at 28th day after the infusion and all five children achieving MRD negativity in the PALL study at the 28th day after the infusion. Additional data will be presented on March 21, 2018 during the European Society for Blood and Marrow Transplantation Annual Meeting.“ALPHA Study”).

With respect to UCART123, we obtained the unanimous approval of the ~~RAC~~National Institute of Health’s Recombinant DNA Advisory Committee (RAC) on December 14, 2016 to start two proposed studies in the United States. In December 2016, we submitted an ~~Investigational New Drug (IND)~~IND application for UCART123 with respect to two proposed Phase I studies to be conducted, one in Acute Myeloid Leukemia (AML) and one in Blastic Plasmacytoid Dendritic Cell Neoplasm (BPDCN). In February 2017, the FDA approved the IND and the first patients were enrolled during 2017. Due to a death in the BPDCN study, the FDA placed a clinical hold on both trials in September 2017 which was lifted by the FDA in November 2017, based on revised protocols. The Phase I clinical study in AML is performed by Weill Cornell and MD Anderson Cancer Center, and the Phase I clinical study in BPDCN is performed by MD Anderson Cancer Center. We refer in this Annual Report to the AML and the BPDCN studies, collectively, as the UCART123 Clinical Studies.

With respect to UCART22, in April 2018, we submitted an IND application with respect to a proposed Phase I study to be conducted in ALL. In May 2018, the FDA approved the IND, and we expect to enroll the first patient in this study in 2019. We refer in this Annual Report to this study as the UCART22 Clinical Study.

With respect to UCARTCS1, on December 28, 2018, we submitted an IND application with respect to a proposed Phase I study to be conducted in Multiple Myeloma (MM). On January 25, 2019, the FDA approved the IND, and the trial is expected to start in 2019. We refer in this Annual Report to this study as the UCARTCS1 Clinical Study.

Additionally, we are pursuing proprietarypre-clinical programs (for ~~UCARTCS1, for UCART22, and for~~ other UCART product ~~candidates)~~candidates, including in solid tumors). Our objective is to file, directly or indirectly, one ~~Investigational New Drug, or~~ IND application (or foreign equivalent), per year from our maturing product ~~candidates~~candidate portfolio.

Our vision in the biopharmaceutical industry is to continue to leverage the potential of gene editing to deliver revolutionary products that address unmet medical needs. Our initial focus is to apply our leadership in gene-editing technology to develop and commercialize~~best-in-class~~ ~~allogeneic CART-cell~~ ~~therapeutic products in the area of immuno-oncology.~~

Until July 2017, we fully owned Calyxt, Inc. ~~On July 25, 2017,~~ Calyxt ~~closed its initial public offering~~is focused on ~~the Nasdaq Global Market, selling an aggregate of 8,050,000 shares of common stock at a price of $8.00 per~~

share (including the full exercise by the underwriters of their over-allotment option). The Company received net proceeds of approximately $58.0 million, after deducting underwriting discounts and commissions and offering expenses. As part of the IPO, we purchased 2,500,000 shares of common stock for a value of $20.0 million, which is included in the net proceeds that Calyxt received. Calyxt used $5.7 million of the proceeds from ususing TALENgene editing technology to ~~cover a portion of the outstanding obligations owed~~provide healthier food ingredients to ~~Cellectis.~~consumers. As of December 31, ~~2017,~~2018, Cellectis owns approximately ~~79.7%~~69.5% of Calyxt’s common stock. In connection with Calyxt’s initial public offering, we and Calyxt entered into certain agreements that related to our relationship with Calyxt prior to the IPO or that provide a framework for our ongoing relationship with Calyxt.

Our Strategy

Our strategy is to leverage the transformative potential of our unique gene-editing technologies and expertise through our cell therapy platform.

The key elements of our strategy are to:

Advance our self-owned allogeneic UCART portfolio of product candidates up to the Biologics License Application (BLA) and commercialize them;

Build a self-owned manufacturing capacity to produce commercial-grade UCART products for clinical use, as well as nucleic acids and vectors as starting material of the UCART product candidates;

Structure a commercial launch plan for our wholly-owned product candidates;

Prepare our next innovative project throughan hematopoietic stem cells (HSC) platform;

Utilize our gene-editing platform to develop and commercialize plant products, through our 69.5% (as of December 31, 2018) ownership in Calyxt, for the multibillion dollar agricultural-biotechnology market. Calyxt is ~~focused on using TALEN gene editing technology~~applying our gene-editing technologies to ~~provide healthier~~create healthy food ~~products~~ingredients. By selecting and inactivating target genes in selected crops, we believe Calyxt can produce unique variants with consumer benefits. Calyxt is developing a pipeline of traits for ~~a growing population across the world.~~soybeans, wheat, and alfalfa, and intends to conduct further development programs to build upon this current pipeline.

~~Our Immuno-oncology~~UCART Pipeline

We are developing a series of product candidates for advanced hematologic cancers. Our lead immuno-oncology product candidates, which we refer to as ~~UCARTs, or universal CARTs,~~Universal CART-cells (UCARTs), are ~~all~~ allogeneic CART-cells engineered to be used as an“off-the-shelf” treatment for ~~treating~~ any patient with a particular cancer type. Each UCART product candidate ~~targets~~is designed to target a selected antigen expressed on tumor cells and bears specific engineered attributes, such as inhibition of alloreactivity and compatibility with specific medical regimens that cancer patients may undergo. UCART is the first therapeutic product line that we are developing ~~using~~with our gene-editing platform to address unmet medical needs in oncology. We are focusing our initial internal pipeline in the hematologic ~~malignancies~~cancer space, targeting diseases with high unmet needs such as ~~acute lymphoblastic leukemia (ALL), acute myeloid leukemia (AML), multiple myeloma (MM)~~ALL, AML, NHL, MM and ~~different~~other types of ~~lymphomas.~~ cancers.

In December 2016, we filed an IND for our lead product candidate, UCART123 in AML and in BPDCN, and in February 2017, we received FDA approval to initiate UCART123 clinical studies. With respect to UCART22, in April 2018, we submitted an IND application with respect to a proposed Phase I study to be conducted in ALL, and in May 2018, the FDA approved the IND. Although we have not yet commenced ~~UCART123 Clinical Studies.~~the UCART22 clinical study, the trial is expected to start in 2019. On December 28, 2018, we submitted an IND application with respect to a proposed Phase I study to be conducted for UCARTCS1 in MM. On January 25, 2019, the FDA approved the IND, and the trial is expected to start in 2019. All of our other ~~fully-controlled~~ product candidates are currently in thepre-clinical~~proof-of-concept~~ phase, and the following chart highlights ~~some of these~~our key product candidates:

LOGO

*	~~or foreign equivalent~~ under Servier License Agreement

**	~~UCART19 is exclusively licensed to Servier and~~ under ~~a joint clinical development agreement between Servier and Pfizer~~Allogene License Agreement

~~With respect~~Targeted Indications

Acute Lymphoblastic Leukemia (ALL)

ALL is a heterogeneous hematologic disease characterized by the proliferation of immature lymphoid cells in the bone marrow, peripheral blood, and other organs. The proliferation and accumulation of blast cells in the marrow results in suppression of hematopoiesis and, thereafter, anemia, thrombocytopenia, and neutropenia. Extramedullary accumulations of lymphoblasts may occur in various sites, especially the meninges, gonads, thymus, liver, spleen, or lymph nodes. Data from the Surveillance, Epidemiology, and End Results (SEER) database have shown anage-adjusted incidence rate of ALL in the United States of 1.7 per 100000 individuals per year, with approximately 5960 new cases and 1,470 deaths estimated in 2018. The median age at diagnosis for ALL is 15 years with 55.4% of patients diagnosed at younger than 20 years of age. In contrast, 28% of cases are diagnosed at 45 years or older and only 12.3% of patients are diagnosed at 65 years or older. ALL represents 75% to ~~our two~~80% of acute leukemia among children, making it the most ~~advanced~~common form of childhood leukemia; by contrast, ALL represents approximately 20% of all leukemia among adults. The cure rates and survival outcomes for patients with ALL have improved dramatically over the past several decades, primarily among children. Improvements are largely owed to advances in the understanding of the molecular genetics and pathogenesis of the disease, the incorporation of risk-adapted therapy, and the advent of new targeted agents. Despite great progress in the development of curative therapies, ALL remains a leading cause of pediatric cancer-related mortality for patients presenting with a relapsed or refractory disease. New therapies are needed to overcome chemotherapy resistance and reduce~~pre-clinical~~non-specific ~~product candidates, we intend~~treatment associated side effects.

Acute Myeloid Leukemia (AML)

AML is a form of cancer that is characterized by infiltration of the bone marrow, blood, and other tissues by proliferative, clonal, abnormally and/or poorly differentiated cells of the hematopoietic system called blast cells. These cells interfere with normal

hematopoiesis, thus contributing to ~~file~~the bone marrow failure which is the most common underlying cause of death. AML is the most common type of acute leukemia in adults with an ~~IND~~age-adjusted incidence rate in the United States of 4.3 per 100000 individuals per year, with approximately 19,520 new cases and 10,670 deaths estimated in 2018. Although it can occur in children and adults, AML is primarily a disease of the elderly. The median age at onset is 68 years and only 16.2% of patients are younger than 45 years of age at diagnosis. While complete response rates can be as high as 80% in patients undergoing initial induction cytotoxic chemotherapy, the majority of AML patients will ultimately be diagnosed with relapsed or refractory disease with a poor prognosis. The outcome in older patients who are unable to receive intensive chemotherapy without unacceptable side effects remains dismal, with a median survival of only 5 to 10 months. CD123 is highly expressed on AML leukemic stem cells and blast cells, as well as in other hematologic malignancies, and constitutes an attractive target for ~~our UCART22 product candidate~~AML.

Multiple Myeloma (MM)

MM is a clonal plasma cell malignant neoplasm that is characterized by the proliferation of a single clone of plasma cells producing a monoclonal immunoglobulin. This clone of plasma cells proliferates in the bone marrow and often results in extensive skeletal destruction with osteolytic lesions, osteopenia, and/or pathologic fractures. Additional disease-related complications include hypercalcemia, renal insufficiency, anemia, and infections. MM accounts for approximately 10% of hematologic malignant disorders. The annual incidence,age-adjusted to the US population, is 6.7 per 100000, resulting in over 30770 new patients in the United States in 2018. The median age at onset is 69 years, and only 3.3% of patients are younger than 45 years of age at diagnosis. Several drugs have been approved over the ~~first~~last few years for the treatment of MM, substantially expanding the number of treatment regimens available for patients in all stages of the disease. In the last decade, survival of MM patients has markedly improved with a median survival of approximately 7 to 10 years but with major variation depending on host factors, stage of the disease, cytogenetic abnormalities, and response to therapy. However, despite this progress, patients with disease refractory to both immunomodulatory drugs (IMiDs) and proteasome inhibitors have a median overall survival (OS) of only 9 to 13 months.

Non-Hodgkin Lymphoma (NHL)

NHL is a heterogeneous disease resulting from the malignant transformation of lymphocytes with distinctive morphologic, immunophenotypic, genetic, and clinical features. NHL is more common than the other general type of lymphoma, Hodgkin lymphoma (HL). The past several decades have seen a steady increase in incidence rates of NHL, with overall rates in the United States nearly doubling over the period 1975 to 2008. In 2018, there were 74680 estimated new cases with 19910 estimated deaths. In 2015, there were an estimated 686042 people living with NHL in the United States. Many different subtypes ofnon-Hodgkin’s lymphoma exist. The most common NHL subtypes include diffuse largeB-cell lymphoma (DLBCL) and follicular lymphoma (FL).

Blastic Plasmacytoid Dendritic Cell Neoplasm (BPDCN)

BPDCN is a rare and aggressive hematological neoplasm classified among AML in the 2008 World Health Organization (WHO) classification of hematologic malignancies, and reclassified under myeloid neoplasms, acute leukemia in the 2016 WHO classification. BPDCN is a rare myeloid disease characterized by the clonal proliferation of precursors of plasmacytoid dendritic cells. There are no formal studies on the incidence of BPDCN in the general population. The few available data reported indicate that its overall incidence is extremely low, accounting for 0.44% of all hematologic malignancies and 0.7% of cutaneous lymphomas. Moreover, the leukemic form of disease is a rare phenomenon, representing fewer than 1% of cases of acute leukemia. The disease may occur at any age, but most patients are elderly men who present with skin lesions and/or involved lymph nodes, spleen, and bone marrow. Given its rarity and only recent recognition as a distinct clinico-pathological entity, no standardized therapeutic approach has been established for BPDCN and the optimal therapy remains to be defined. Although transient responses are seen to combination chemotherapy regimens used to treat acute leukemia or lymphoma, most patients relapse with drug-resistant disease with a median OS rate of 9 to 13 months, irrespective of the initial presentation of the disease.

UCART19 for ALL

UCART19 is an allogeneic,off-the-shelf product candidate designed to exhibit high efficacy in fighting hematological malignancies bearing theB-lymphocyte antigen CD19. In November 2015, Servier acquired the exclusive rights to UCART19 from Cellectis. Servier and Allogene collaborate on a joint clinical development program for UCART19, and Allogene has the exclusive rights from Servier to develop and commercialize UCART19 in the United States.

Product Features

UCART19 is an allogeneicT-cell product intended for the treatment of CD19-expressing hematologic malignancies such as ALL.

UCART19 is designed to become active, proliferate, secrete cytokines and kill CD19-bearingB-cell malignancies upon contact with such cells, following administration to patients. Activation of UCART19 is driven by contact between its anti-CD19 CAR and the CD19 protein on the surface of tumor cells.

UCART19 cells bear a CAR targeting the CD19 antigen that drives their capacity to kill CD19-bearing cells. Moreover, as all UCART product candidates, UCART19 lacks the TCR responsible for recognition ofnon-self antigens by theT-cells, which allows use of healthy donorT-cells to produce UCART19, with reduced potential for GvHD. In addition, some UCART19 cells lack CD52, a protein expressed on the cell surface that makesT-cells sensitive to alemtuzumab. This feature permits the use of UCART19 in patients recently treated or being treated with the immunosuppressing/lymphodepleting agent alemtuzumab.

UCART19 activation could potentially lead to eradication of CD19-expressing cancer cells throughT-cell mediated killing of such cancer cells and potentiallypro-inflammatory immune system production as well as CART-cell amplification.

Clinical Findings

In December 2018, Servier presented pooled intermediate results from the UCART19 Clinical Studies during the ASH annual conference. After UCART19 infusion, 82% (14/17) of patients who received a lymphodepletion regimen (consisting of fludarabine, cyclophosphamide and alemtuzumab, an anti-CD52 monoclonal antibody) achieved a complete remission (CR) or complete remission with incomplete blood cell recovery (CRi) by day 28 or day 42. Within responder patients, 71% (10/14) of them were ‘minimum residual disease’ (MRD) negative(MRD- stands for less than 1 leukemic cell among 10E4 normal cells) assessed by flow or qPCR. When considering all treated patients (lymphodepleted or not), 67% (14/21) of them did achieve CR/CRi. Regarding safety considerations, there was no serious adverse events (grade³3) for graft versus host disease (GvHD) and neurological events. Grade3-4 toxicities did only regard events of cytokine release syndrome (14%, 3/21), prolonged cytopenia (29%, 6/21) and viral infections (24%, 5/21). We are encouraged by these promising preliminary results reported for the UCART19 Clinical Studies.

Development Status

In 2016, Servier commenced the UCART19 Clinical Studies—a Phase I clinical study in pediatric ALL, the PALL study, and a Phase I clinical study in adult patients with ALL, the CALM study, each of which was approved by the MHRA.

The PALL Study is commenced in the United Kingdom at UCL Great Ormond Hospital (London), in Belgium at Het Kinderziekenhuis Prinses Elisabeth (Gent), in France at Hôpital Robert-Debré (Paris) and in United States at the Children’s Hospital of Philadelphia (Pennsylvania).

The CALM Study is commenced in the United Kingdom at King’s College Hospital NHS Foundation Trust (London), in United States at the Hospital of the University of Pennsylvania (Philadelphia, Pennsylvania), at University of Texas MD Anderson Cancer Center (Houston, Texas) and at the Massachusetts General Hospital (Boston, Massachusetts), in France at Hôpital Saint-Antoine (Paris) and Hopital Saint-Louis.

UCART19, orALLO-501, for NHL

ALLO-501 (or UCART19 corresponding to the product candidate which we exclusively licensed to Servier and which was sublicensed to Allogene by Servier in the United States) is an allogeneic engineeredT-cell product intended for the treatment of CD19-expressing hematologic malignancies.ALLO-501 is a product candidate under the license agreement between Servier and Cellectis.

Development Status

On January 28, 2019, Allogene announced, in collaboration with Servier, that the FDA approved the IND for Phase 1 clinical study forALLO-501 in relapsed/refractory NHL (the “ALPHA Study”).

UCART123 for AML and BPDCN

UCART123 is an allogeneic engineeredT-cell product designed for the treatment of hematologic malignancies expressing the alpha chain of theinterleukin-3 receptor (IL3RA), or CD123, and is currently being developed for the treatment of AML and BPDCN.

Product Features

UCART123 is an allogeneicT-cell product candidate intended for the treatment of CD123-expressing hematologic malignancies.

UCART123 is designed to become active, proliferate, secrete cytokines and kill CD123 expressing cells. UCART123 bears a CAR targeting the CD123 antigen, providing specificity for CD123 expressing cells. In addition, as with all UCART products, UCART123 lacks the TCR and is intended to be used in an allogeneic context. UCART123 activity could potentially lead to eradication of CD123-expressing cancer cells throughT-cell mediated killing,pro-inflammatory cytokine production as well as CART-cell amplification. New versions of UCART123 produced during the fourth quarter of 2018 have, in addition of the suppression of the TCRµ gene, the suppression of the CD52 gene in order to eventually induce resistance to alemtuzumab preconditioning. This product might get into the clinic during the year 2019.

Pre-clinical Findings

UCART123 has been evaluated bothin vitro and in animal studies, with promising results.

In vitro studies demonstrated efficient killing of human CD123-bearing cell lines by UCART123. In addition, UCART123 has also demonstrated efficient killing of human CD123-expressing cells derived from AML and BPDCN patients. Animal studies were conducted in mice injected both with UCART123 and human CD123-bearing tumor cells, and have shown anti-tumor activity in an immunodeficient mouse model. In addition, in another animal model, limited toxicity against normal, healthy cells, has been observed.

UCART123 was also tested for its potential to induce GvHD. Mice receiving unmodifiedT-cells from a human donor showed GvHD, while mice receiving the UCART123 cells that lack the TCR showed no sign of GvHD.Pre-clinical and translational activities on UCART123 in AML and BPDCN were performed in collaboration with Weill Cornell and MD Anderson Cancer Center, respectively.

Development Status

On December 14, 2016, Cellectis received unanimous approval from the RAC for two proposed Phase I protocols for UCART123. In February 2017, the FDA granted Cellectis an IND approval to conduct a Phase I clinical study with UCART123 in AML and BPDCN. The initial patients were enrolled during 2017, at Weill Cornell Medical College for AML and at MD Anderson Cancer Center for BPDCN. Due to a death in the BPDCN study, both trials were put on hold in September 2017 but then cleared to resume by the FDA, in November 2017, based on revised protocols.

On September 14, 2018, Cellectis received approval from the MHRA in the United Kingdom for a proposed Phase I protocol for UCART123 in first-line adverse genetic risk AML.

UCART22 for ALL

UCART22 is an allogeneic engineeredT-cell product candidate designed for the treatment of ALL.

Product Features

UCART22 is an allogeneic engineeredT-cell product candidate intended for the treatment of CD22-expressing hematologic malignancies. UCART22 is designed to become active, proliferate, secrete cytokines and kill CD22 expressing cells (i.e. either CD22 positive tumor cells ornon-malignant CD22-positive B lineage cells). UCART22 bears a CAR targeting the CD22 antigen, providing specificity for CD22 expressing cells. As with all UCART products, UCART22 lacks the TCR and is intended to be used in an allogeneic context. In addition, some UCART22 cells lack CD52, a protein expressed on the cell surface that makesT-cells sensitive to alemtuzumab, a drug often used to treat CLL patients. This feature should allow for improved engraftment of the cells in conjunction with a potential alemtuzumab treatment.

UCART22 activity could potentially lead to eradication of CD22-expressing cancer cells throughT-cell mediated killing,pro-inflammatory cytokine production as well as CART-cell amplification.

Pre-clinical findings

UCART22 has been evaluated bothin vitro and in animal studies, with promising results.

In vitro studies demonstrated efficient killing of human CD22-bearing cell lines by UCART22. In addition, UCART22 has also demonstrated efficient killing of human CD22-expressing cells derived from ALL patients. Animal studies were conducted in mice injected both with UCART22 and human CD22-bearing tumor cells, and have shown anti-tumor activity in an immunodeficient mouse model. Furtherin vitro andin vivo studies are ongoing to further investigate the safety and the activity of UCART22.

Pre-clinical and translational activities on UCART22 in ALL are performed in collaboration with MD Anderson Cancer Center.

Development Status

In April 2018, we filed an IND and in May 2018, the FDA granted Cellectis an IND approval to conduct a Phase I clinical study with UCART22 in ALL. Manufacturing of clinical-grade UCART22 at large scale in accordance with GMP is still ongoing and should be completed in 2019.

UCARTCS1 for MM

UCARTCS1 is an allogeneic engineeredT-cell product candidate designed for the treatment ofCS1-expressing hematologic malignancies which is being developed in MM.

Product Features

UCARTCS1 is an allogeneicT-cell drug candidate intended for the treatment of CS1 (also known as SLAMF7)-expressing hematologic malignancies, in particular MM. UCARTCS1 is designed to become active, proliferate, secrete cytokines and kill CS1 expressing cells. As CS1 is strongly expressed on the cell surface of CD8T-cells but also mildly expressed on CD4 cells, B cells, NK cells and macrophages, CS1 will be inactivated in UCART cells prior to transduction with a viral vector encoding ananti-CS1 CAR. The inactivation of the CS1 gene may improve the production and activity of UCARTCS1 by preserving the balance between CD8 and CD4T-cell population. In addition, as with all UCART products, UCARTCS1 lacks the TCR and is intended to be used in an allogeneic context. We believe that UCARTCS1 might have a potential lymphodepleting activity by attacking the immune cells of the patient expressing CS1.

As compared to other targets frequently addressed byCAR-T candidates in MM, such as BCMA, CS1 expression has been observed to be higher and more uniform. In certain mouse models, CS1CAR-T therapy is showing deeper response than what is seen with BCMACAR-T therapy.

Pre-clinical Findings

In vitro studies demonstrated efficient killing of humanCS1-bearing cell lines by UCARTCS1. In addition, UCARTCS1 has also demonstrated efficient killing of humanCS1-expressing cells derived from MM patients. Furthermore, whilenon-gene-editedT-cells expressing ananti-CS1 CAR display limited cytolytic activityin vitro against MM cell lines and result in a progressive loss of CD8T-cells,CS1-gene-edited CAR cells (UCARTCS1) display significantly increased cytotoxic activity, with the percentage of CD8T-cells remaining unaffected. Experiments in an orthotopic MM mouse model showed that UCARTCS1 was able to mediate anin vivo anti-tumoral activity. Furtherin vitro andin vivo studies are ongoing to further investigate the safety and the activity of UCARTCS1.

Pre-clinical and translational activities for UCARTCS1 in MM are performed in collaboration with the MD Anderson Cancer Center.

Development Status

During the second half of 2018, ~~and~~a first set of batches of the manufacturing campaign of UCARTCS1 have been manufactured at CELLforCURE, our third-party manufacturing contractor.

On December 28, 2018, Cellectis filed an IND, which was approved by the FDA on January 25, 2019 by the FDA, in order to conduct a Phase I clinical study with UCARTCS1 in MM.

UCARTCLL1 for ~~UCART CS1 in 2019. All the above products are~~AML

UCARTCLL1 is an allogeneic engineeredT-cell product candidate designed for the treatment of AML.

Product Features

UCARTCLL1 is an allogeneicT-cell drug candidate intended for the treatment of CLL1 (also known as CLEC12A)-expressing hematologic malignancies, in particular AML. UCARTCLL1 is designed to become active, proliferate, secrete cytokines and kill CLL1 expressing cells. In addition, as with all UCART products, UCARTCLL1 lacks the TCR and is intended to be used in an allogeneic context. In our UCARTCLL1 product, theß2-microglobulin (ß2M) gene will also be inactivated, which is expected to result in betterCAR-T persistence (see section on “Next-generation products” above). Furthermore, while we are using lentiviral (LV) vectors to express the CAR in all our current UCART products, UCARTCLL1 will benefit from targeted insertion of the CAR at the TCR locus through an adeno-associated viral (AAV) vector. This is expected to result in targeted integration of our CAR gene in theT-cell and more predictable expression profile of the CAR gene, which will be under endogenous regulatory elements.

UCART programs for solid tumors

We are currently applying our UCART platform to developCAR-Tcandidates ~~that bear CARs that seek~~targeting solid tumors, currently in preclinical phase.

Other gene editing programs

Beyond ourCAR-T programs, we are leveraging our TALEN gene editing platform to ~~kill cells expressing targets (e.g., CD123, CS1,~~pursue additional development opportunities, both internally and ~~CD22), which are found~~ in ~~other hematologic tumors, such as acute myeloid leukemia,~~collaboration with third party companies and academics centers. We aim to enter the clinic with one or ~~AML, and multiple myeloma, or MM, as well as acute lymphoblastic leukemia, or ALL.~~more gene editing programs beyond UCARTs in the near future.

Our Strategic Alliances

In addition to the development of our own portfolio of product candidates targeting tumor-associated antigens, we have pursued a strategy of forging strong pharmaceutical alliances. ~~We believe that our strategic alliances~~

License Agreement with ~~Servier~~Allogene

In June 2014, we entered into a Research Collaboration and License Agreement (the “Collaboration and License Agreement”) with Pfizer, ~~validated our technology platform, our strong expertise~~Inc. (“Pfizer”) pursuant to which we agreed to collaborate to conduct discovery andpre-clinical development activities to generate CART-cells directed atPfizer- and Cellectis-selected targets in the ~~allogeneic CAR~~field of human oncology. We granted Pfizer an exclusive, worldwide, royalty-bearing, sublicensable license, on a~~T-cells~~target-by-target ~~field, our approach to CART-cell~~ ~~development and the strength~~basis, under certain of our intellectual property ~~portfolio. Our strategic alliances include~~to make, use, sell, import, and otherwise commercialize products directed at thePfizer-selected targets in the field of human oncology. Pursuant to the Collaboration and License Agreement, Pfizer made an upfront,non-refundable $80.0 million payment to us, concurrent with Pfizer’s €25.8 million equity investment in our company. On April 3, 2018, Pfizer and ~~potential~~Allogene Therapeutics, Inc. (“Allogene”), a new company started by former Kite Pharma executives Dr. Arie Belldegrun and Dr. David Chang, announced that they entered into an asset contribution agreement, pursuant to which Allogene purchased Pfizer’s portfolio of assets related to allogeneic CART-cell therapy (the “Asset Contribution Transaction”). Pursuant to the Asset Contribution Transaction, effective as of April 6, 2018, Allogene purchased Pfizer’s portfolio of assets related to allogeneic CART-cell Therapy, including the Collaboration and License Agreement.

During the ASH annual conference in December 2018, Allogene presented data related toALLO-715 andALLO-819, which are product candidates directed at two of the fifteen targets (BCMA and FLT3 respectively) that are exclusively licensed to Allogene from Cellectis.ALLO-715 is an allogeneic BCMACAR-T therapy for the treatment of Multiple Myeloma andALLO-819 is an allogeneic FLT3CAR-T therapy for the treatment of AML. Each ofALLO-715 andALLO-819 is designed to possess anoff-switch for safety. Allogene holds the exclusive development and commercial rights for these product candidates.

On March 7, 2019, we and Allogene agreed to terminate the Collaboration and License Agreement and entered into a new license agreement (the “Allogene License Agreement”) to reflect the relationship between us and Allogene following the Asset Contribution Transaction. The Allogene License Agreement establishes the rights and obligations of Cellectis and Allogene with respect to their collaboration program.

Pursuant to the Allogene License Agreement, we granted to Allogene an exclusive, worldwide, royalty-bearing, license, on atarget-by-target basis, with sublicensing rights under certain conditions, under certain of our intellectual property, including our TALEN and electroporation technology, to make, use, sell, import, and otherwise exploit and commercialize chimeric antigen receptor (CAR) T products directed at certain targets, including BCMA, FLT3, DLL3 and CD70, for human oncologic therapeutic, diagnostic, prophylactic and prognostic purposes. In addition, the Allogene License Agreement accommodates an exclusive global license and collaboration agreement under which Allogene has obtained from Servier exclusive rights to develop and commercialize

UCART19 in the United States. Further, Allogene granted us anon-exclusive, worldwide, royalty-free, perpetual and irrevocable license, with sublicensing rights under certain conditions, under certain of Allogene’ s intellectual property, to make, use, sell, import and otherwise commercialize CAR T products directed at certain targets.

The Allogene License Agreement provides for development and sales milestone payments ~~to us~~by Allogene of up to ~~$3.9 billion~~$185.0 million, with aggregate potential development and ~~high single digit~~sales milestone payments totaling up to $2.8 billion. Allogene expects to pay us $5.0 million upon the dosing of the first patient in its Phase 1 clinical trial ofALLO-715. We are also eligible to receive tiered royalties on ~~future sales.~~annual worldwide net sales of any products that are commercialized by Allogene that contain or incorporate, are made using or are claimed or covered by, our intellectual property licensed to Allogene under the Allogene License Agreement at rates in the high single-digit percentages.

Unless earlier terminated in accordance with the agreement, our agreement with Allogene will expire on aproduct-by-product andcountry-by-country basis, until the later of (1) the expiration of the last to expire of the licensed patents covering such product; (2) the loss of regulatory exclusivity afforded such product in such country, and (3) the tenth anniversary of the date of the first commercial sale of such product in such country; however, in no event shall the term extend, with respect to a particular licensed product, past the twentieth anniversary of the first commercial sale for such product. At any time after the first anniversary of the effective date of the agreement, Allogene will have the right to terminate the agreement at will upon 60 days’ prior written notice, either in its entirety or on atarget-by-target basis. Either party may terminate the agreement, in its entirety or on atarget-by-target basis, upon 90 days’ prior written notice in the event of the other party’s uncured material breach. The agreement may also be terminated upon written notice by Allogene at any time in the event that we become bankrupt or insolvent or upon written notice within 60 days of a consummation of a change of control of Cellectis.

~~Collaboration~~License, Development and Commercialization Agreement with Servier

In February 2014, we entered into a ~~strategic collaboration agreement~~Research, Product Development, Option, License and Commercialization Agreement (as amended, the “Prior Servier Agreement”) with ~~Servier to develop and commercialize certain product candidates.~~Servier. Pursuant to the ~~agreement,~~Prior Servier ~~made~~Agreement, we were responsible for the research and development up to and including the Phase I clinical trial of candidate products directed against five targets, including the UCART19 product candidate. Pursuant to the Prior Servier Agreement, we granted Servier options to obtain exclusive, worldwide licenses, each on a productcandidate-by-product candidate basis, with respect to each product candidate selected by Servier and developed under the Prior Servier Agreement.

On March 6, 2019, we and Servier entered into a License, Development and Commercialization Agreement (the “Servier License Agreement”). The Servier License Agreement supersedes the Prior Servier Agreement to establish the terms of our continuing collaboration and to reflect the status of products in development. Among other things, the Servier License Agreement updates the targets covered by the Prior Servier Agreement. Servier exercised such option with respect to the UCART19 and continues to have an ~~upfront payment~~exclusive license to this product candidate.

Pursuant to the Servier License Agreement, upon Servier’s exercise of ~~$8.2 million. In addition,~~each license option, Servier will pay us a lump sum license fee, and we will grant Servier an exclusive, worldwide, royalty-bearing license, with right to sublicense, under certain of our patents andknow-how covering the ~~strategic alliance,~~relevant product candidate. This license will cover the development, manufacture and commercialization of such product in the field of anti-tumor adoptive immunotherapy. At such time, Servier will assume responsibility for the further clinical development, manufacture and commercialization of such product.

The Servier License Agreement restricts us from researching, developing, or commercializing any product directed against a target that is used for the same purpose as ~~amended in November 2015, provides for~~it is used with a product candidate developed under the agreement.

We are eligible to receive aggregate ~~additional~~ payments from Servier of up to $1,064 million pursuant to the Servier License Agreement, comprising payments upon the exercise of ~~each option~~options granted to Servier under the agreement and payments upon the occurrence of certain specified ~~development~~milestones (development and commercial ~~milestones.~~ in nature), as well as reimbursement for development costs.

We are also eligible to receive tiered royalties ranging in the high single-digit percentages based on annual net sales of commercialized products. ~~This~~

The agreement ~~covers~~will expire upon the ~~development~~expiration of the last sale of products licensed pursuant to the agreement. The parties also may terminate the agreement at any time by mutual consent. At its sole discretion, Servier has the right to terminate the agreement in its entirety or with respect to specific products, upon three months’ prior written notice to us.

In addition, either party may terminate the agreement following the other party’s uncured material breach upon 90 days’ prior written notice to the breaching party, or 30 days’ notice if such breach relates to a payment obligation. Servier may terminate the agreement at any time for product-related safety reasons. Either party may terminate the agreement in the event of the other party’s bankruptcy or insolvency.

In the event that Servier does not exercise its option to license a product candidate, we may independently pursue all activities related to such product candidate and/or license such product candidate and the ~~potential commercialization of the lead product candidate, UCART19, as well as other product candidates directed at four other targets. Under the terms of the agreement,~~associated intellectual property to a third party. For such purpose, Servier granted us anon-exclusive license, with right to sublicense, under any such Servier-controlled intellectual property for which we will ~~be responsible for~~pay tiered royalties on annual net revenues at rates ranging in the low single-digit percentages.

Collaboration with research and clinical centers

Alliance with The University of Texas M.D. Anderson Cancer Center

On September 1, 2015, Cellectis and the University of Texas MD Anderson Cancer Center (the MD Anderson Cancer Center) entered into a research and development alliance (the Strategic Alliance Agreement) aimed at bringing novel cellular immunotherapies to patients suffering from different types of ~~certain product candidates through~~liquid tumors, particularly MM, ALL,T-cell ALL(T-ALL) and BPDCN. Under this strategic alliance, the ~~end~~MD Anderson Cancer Center and Cellectis have agreed to collaboratively conduct severalpre-clinical studies on candidate products: UCART123 in BPDCN, UCARTCS1 for multiple myeloma, UCART38 forT-ALL and UCART22 for ALL. Cellectis has agreed to provide funding and other support for these studies. The objective of ~~their respective Phase I clinical trials. We granted Servier an exclusive option~~the studies is to ~~obtain an exclusive, worldwide license~~build on ~~a productcandidate-by-product~~ ~~candidate basis, with respect to each target selected by Servier~~complementary expertise from the MD Anderson Cancer Center and developed under the agreement, to further develop, manufacture and commercialize such product in the field of anti-tumor adoptive immunotherapy. Upon exercising each such option, Servier will assume responsibilityCellectis for the ~~further clinical~~ development ~~manufacture and commercialization~~ of the relevant product candidate. In November 2015, we entered into an amendment to our initial collaboration agreement with Servier, which allowed for an early exercise of Servier’s option with respect to UCART19 and other product candidates. ~~Pursuant to this amendment, Servier has exercised its option to acquire the exclusive worldwide rights to further develop~~The MD Anderson Cancer Center and ~~commercialize UCART19. In addition, Pfizer and Servier have entered into an exclusive global license and collaboration agreement, under which Pfizer has obtained exclusive rights~~Cellectis will work together to develop and ~~commercialize UCART19~~implement improvements to the research plan for the programs under joint direction of the MD Anderson Cancer Center and Cellectis’ investigators. The objective of the studies is to demonstrate the functionalities and specificity of the UCART candidate products listed above, define thepre-clinical package required for clinical trial applications, prepare a clinical trial protocol and the regulatory documents required for interactions with FDA and the clinical trial applications. Pursuant to the alliance, Cellectis is responsible for generation and manufacturing of the UCART candidate products and some of the in vitro and in vivopre-clinical work. The MD Anderson Cancer Center is responsible for evaluation of the ~~United States.~~candidate products against primary patient samples and for some activities to be performed in animal models. The alliance also includes the possibility for Cellectis and the MD Anderson Cancer Center to collaborate on one or more early phase clinical studies on the same product candidates.

~~Collaboration~~Pursuant to the Strategic Alliance Agreement, we entered in March 2017 into a study order with ~~Pfizer~~MD Anderson Cancer Center, according to which MD Anderson Cancer Center performs the UCART123 clinical study for BPDCN.

In ~~June 2014,~~January 2018, we entered into a ~~global strategic collaboration agreement~~new study order with ~~Pfizer pursuant to which we will collaborate to conduct discovery andpre-clinical~~ ~~development activities to generate CART-cells~~ ~~directed at targets selected by Pfizer or us in the field of oncology. Pursuant to the agreement, Pfizer made an upfront,non-refundable~~ $80.0 million payment to us, concurrent with Pfizer’s equity investment in our Company. In addition, the strategic alliance provides for up to $2.8 billion in potential clinical and commercial milestone payments. We are also eligible to receive tiered royalties ranging in the high single-digit percentages based on annual net sales of commercialized products. We may also receive funding for research and development costs associated with the~~Pfizer-selected~~ targets and for four Cellectis-selected targets within the alliance. Pfizer has exclusive rights to pursue development and commercialization of products for a total of fifteen targets of their choice, which have all been selected.

~~Our Strategy~~

~~Our strategy is to leverage the transformative potential of our unique gene-editing technologies and expertise through our cell engineering platform.~~

~~The key elements of our strategy are to:~~

~~Accelerate our clinical operations~~MD Anderson Cancer Center in order to ~~accumulate data~~expand the performance of the UCART123 clinical study in AML to MD Anderson Cancer Center.
In May 2018, we entered into a new study order with MD Anderson Cancer Center according to which MD Anderson Cancer Center performs the UCART22 clinical study for ALL.
Research Collaboration Agreement with Cornell University, for and on behalf of its Joan & Standford I. Weill Medical College
On February 20, 2019, Cellectis and Cornell University, for and on behalf of its Joan & Standford I. Weill Medical College, entered into a Research Collaboration agreement. Under this Research Collaboration Agreement, Cornell University performs pre-clinical evaluation regarding certain of our product candidates in pre-clinical development.

Clinical collaborations with Weill Cornell, Dana Farber Cancer Institute and ~~prove their value.~~ ~~Clinical data will be~~H. Lee Moffitt Cancer Center

In April 2017, we entered into a clinical trial agreement with Cornell University acting for and on behalf of its Joan and Sanford I. Weill Medical College and The New York Presbyterian Hospital (collectively referred to as Weill Cornell) under which Weill Cornell performs the ~~lever~~UCART123 clinical study for AML.

In August 2018, we entered into a new clinical study agreement with Dana Farber Cancer Institute and H. Lee Moffitt Cancer Center in order to ~~confirm~~expand the ~~efficacy and value~~performance of the ~~allogeneic CART-cell~~ ~~approach~~UCART123 clinical study in AML to Dana Farber Cancer Institute and ~~bring breakthrough innovation to patients.~~

H. Lee Moffitt Cancer Center.

~~Continue to leverage our cell-engineering platform to develop additional UCART product~~~~candidates~~~~and to expand our clinical pipeline of CART-cell~~ ~~product candidates in the coming years.~~

~~Leverage our existing and potential future alliances to advance our research and to bring products to market~~~~. Our strategic alliances for the development of CART-cell~~ applications in oncology provide us with funding for research and development, and may provide milestone payments and royalties on sales. We may enter into additional strategic alliances to facilitate our development and commercialization of CAR~~T-cell~~ ~~immunotherapy products.~~

~~Expand our product pipeline to other therapeutic indications with unmet medical needs~~. We intend to continue using our gene-editing technologies in therapeutic applications beyond immuno-oncology, including the treatment of chronic infectious diseases, autoimmune diseases and allergic diseases.

~~Utilize our gene-editing platform to develop plant products, through our 79.3% (as of February~~ ~~28, 2018) ownership in Calyxt, for the multibillion dollar agricultural-biotechnology market~~. Calyxt is applying our gene-editing technologies to create food products with consumer health benefits, adaptations for climate change or nutritional enhancements that address the needs of a growing population. By selecting and inactivating target genes in certain agricultural crops, we believe Calyxt can produce unique variants with consumer benefits. For example, Calyxt is developing a diversified portfolio spanning across five core crops and a multitude of product candidates, which include innovative consumer-centric product candidates as well as innovative, farmer-centric solutions.

Immunotherapy: Turning the Immune System into “Smart Drugs”

The immune system has evolved to protect the body from invading pathogens or external harmful materials by identifying these foreign bodies through“non-self” antigens, which are molecular signatures that they carry and are foreign to the body. A central function of the immune system is to discriminate between “self,” which is recognized through antigens normally present in the body and borne by cells, proteins, sugars or lipids, and“non-self”, which is detected through abnormal or foreign antigens. Cancer cells thrive, in part, because they trick the immune system into treating them as self, even though they express abnormal antigens, and thus immune tolerance occurs when the immune system fails to recognize and attack tumors. Breaking immune tolerance is an important aspect of most immuno-oncology-based therapeutics because it enables the immune system to recognize and treat tumors asnon-self and lead to tumor destruction.

The immune system recognizesnon-self danger signals and responds to threats at a cellular level. The immune system may be conceptualized as comprising two arms. The first arm, known as the innate immune system, recognizesnon-specific signals of infection or abnormalities as a first line of defense. The innate immune system is the initial response to an infection, and the response

is the same every time regardless of prior exposure to the infectious agent. The second arm, known as the adaptive immune system, is composed of highly specialized cells and provides long-term specific recognition and protection from infectious agents and abnormal processes such as cancer. The adaptive immune response is further subdivided into antibody-based responses and cellular responses, which includeT-cell-based immune responses. The most significant components of the cellular aspect of the adaptive immune response areT-cells, which are specialized cells that generally mature in the thymus.T-cells are involved in sensing and killing infected or abnormal cells, as well as coordinating the activation of other cells and mounting an immune response.

Although the immune system is designed to identify and destroy foreign or abnormal protein-bearing tumor cells, this process is often defective in cancer patients. Additionally, cancer cells employ a number of mechanisms to escape immune detection and attack to suppress the effect of the immune response.

Immunotherapy is a type of treatment that modifies, stimulates, orre-directs certain parts of the immune system to fight diseases, such as cancer. Immunotherapy works by stimulating a patient’s own immune system or by turning its attacks towards harmful targets, such as cancer cells. Immunotherapy can also be pursued by giving patients engineered immune cells, such as CART-cells to target certain cells. Immunotherapy is playing an increasingly large role in treating cancer, chronic infectious diseases, autoimmune diseases and allergic diseases.

T-cells andT-cell Receptors (TCRs)

T-cells are a class of white blood cells that carry a specific TCR at their surface that allows them to recognize and kill other cells that express antigens foreign to the individual. Normal cells express a set of specific

molecules, called human leukocyte antigen, or HLA, at their surface. HLA is associated with small fragments, or peptides of the proteins expressed inside the cell or processed from the extracellular body fluids. Abnormal or foreign proteins (viruses, for example) can attach to HLAs and be presented at the cell’s surface and be recognized byT-cells through theseHLA-peptide complexes and identified as foreign antigens. This recognition triggers the activation of theT-cells, which destroy the foreignHLA-peptide complex-bearing cell, secrete specific cytokines attracting other immune-competent cells to their location, and start multiplying to establish a full immune response.

~~An activatedT-cell~~ ~~can multiply tens of thousands of times, so long as there remains a presence of the foreign antigen in the body.~~ Unlike antibodies that mainly diffuse passively through the body and its circulating fluids,T-cells actively leave blood vessels or lymphoid organs and travel through the tissues of the body where they can attack foreign antigens. Once the antigen is eliminated from the body, theT-cells run out of stimulation and die off, with only a fraction surviving as “memoryT-cells,” which can react promptly should the antigen reappear in the body.

There is a high variability of HLA molecules in the population. Therefore, if a cell is introduced into a person and originally comes from another individual that is notHLA-matched, it will bear, at its surface,HLA-peptide complexes that are recognized as foreign and will be killed by theT-cells of the recipient. This mechanism of graft rejection has been a major limitation to transplanting patients with allogeneic tissues. Reciprocally, ifT-cells are grafted from one individual to another and start recognizing as foreign the normalHLA-peptide complexes at the surface of all tissues of the grafted individual, then they may attack and kill those healthy tissues, leading to Graft-versus-Host disease (GvHD), which can be very severe, and potentially fatal, if left untreated. ~~GvHD has been a major limitation to the use of allogeneicT-cells~~ ~~when treating patients.~~

Cancerous cells express abnormal antigens and can be killed byT-cells. However, cancer may grow and spread to various organs whenT-cells with cancer-specific receptors are in low numbers, of poor quality, or rendered inactive by suppressive mechanisms employed by tumor tissues.T-cells are a key armament when fighting cancers. They play a particularly significant role if they are tailored to target tumors, and potentially even more so if their genes are edited to overcome tumor defenses, to makeT-cells compatible with other anti-cancer drugs that can be combined with them, and to prevent GvHD, which would allow the use of allogeneicT-cells.

Chimeric Antigen Receptor ~~(CARs)~~(CAR)

CARs are engineered molecules that, when present at the surface ofT-cells, enable them to recognize specific proteins or antigens that are present on the surface of other cells. These receptors are typically used to graft the specificity of an antibody derived from a single cell, or a monoclonal antibody, onto aT-cell and provide it with a specific targeting mechanism to seek, identify, interact with and destroy the tumor cells bearing a selected antigen associated with that tumor also known as tumor-associated antigen, or TAA and tumor-specific antigens, or TSA. The expression of some genes, or combinations of genes, can be associated with certain classes of cancers. It is sometimes possible to identify TAAs that are expressed at various levels by tumor cells from a given cancer type. These TAAs may also be normally expressed by other tissues at different stages of development.

T-cells with CARs are referred to as CART-cells. Whereas naturalT-cell receptors, or TCRs, only recognize antigens bound to an HLA molecule at a cell’s surface, a CAR is able to directly recognize antigens that are present at the targeted cell’s surface. It is believed that uponcell-to-cell contact between a CART-cell and an antigen-bearing targeted cell, antigen recognition by the CAR “activates” the CART-cell, triggering it to multiply, attack and kill its target through the release of “hole-forming” proteins, known

as perforins, and “degradation enzymes,” known as granzymes, that enter the targeted cell through the perforin-formed holes and carry out the killing. The activation of aT-cell through a CAR results in a target-associated “kill and amplify” chain reaction that eradicates the tumor.

CARs are constructed by assembling components, or domains, from different proteins, including:

In the extracellular space, one or more target binding domains, coming from ligands, such as antibodies or receptors, that can recognize their targets on the outside of theT-cell;

A hinge that helps position the target binding domains relative to their targets;

Trans-membrane domains that anchor the CAR at theT-cell’s surface relative to theT-cells; and

A set of activating or signaling domains, which are located within theT-cell’s interior, that deliver appropriate signals to theT-cells leading toT-cell activation or repression according to theT-cell environment. Such signals may induce tumor cell killing, cytokine secretion and CART-cell multiplication.

The following diagram shows the mechanism by which a CART-cell is believed to attack a tumor cell:

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Recent immuno-oncology advancements have supported the potential to cure certain cancers by harnessing the body’s immune system to fight cancer cells. ~~For example:~~In particular, two autologousCAR-T cell therapies targeting the antigen CD19 have been approved in 2017:

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In August 2017, the FDA approved tisagenlecleucel (Kymriah^®) for the treatment of patients up to 25 years of age withB-cell precursor ALL that is refractory or in second or later relapse. ~~This approval was based on~~In May 2018, the ~~results~~FDA approved a label extension for Kymriah^® for the treatment of ~~an open-label, multicenter~~adult patients with relapsed or refractory large~~single-arm~~B-cell ~~trial. Of the 63 evaluable patients, 52 responded, including 40 patients (63%) with a CR within 3 months~~lymphoma after ~~infusion, and 12 (19%) with a CR but with incomplete blood count recovery. Bone marrow from all patients demonstrated minimum residual disease-negative status.~~two or more lines of systemic therapy.Q1-Q3 2018 sales of Kymriah^® were $48 million.

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In October 2017, the FDA approved axicabtagene ciloleucel (Yescarta^®) for the treatment of ~~adults~~adult patients with ~~diffuse~~relapsed or refractory largeB-cell lymphoma ~~(DLBCL) who have not responded to~~after two or ~~who have relapsed after at least two other kinds~~more lines of ~~treatment. The approval was based on the results~~systemic therapy.Q1-Q3 2018 sales of ~~an open label trial in refractory aggressivenon-Hodgkin~~ ~~lymphoma (DLBCL, primary mediastinalB-cell~~ lymphoma, or transformed follicular lymphoma), which enrolled 111 patients, 101 of whom received Yescarta. The objective response rate (ORR) was 82% and at a median follow up of 8.7 months, 44% of patients had responded, and 39%Yescarta were ~~in CR.~~$183 million.

Based on these and other advancements, immuno-oncology has become a new frontier for treatment, and we believe it is one of the most promising areas of development within oncology.

~~Limitations of Current Autologous Treatments~~

~~Many of the CART-cell~~ ~~immunotherapy treatments currently under development are created through an autologous approach in which the patient’s ownT-cells~~ ~~are engineered to fight cancer cells. Part of our scientific basis for pursuing allogeneic approaches rests in the recognized limitations of autologous approaches, including:~~

~~Autologous treatments must be specifically manufactured for each patient and the resulting engineered cells may have different properties due to significantpatient-to-patient~~ ~~variability in the quality of theT-cell;~~

~~Autologous treatments can bear high costs due to the necessity of producing a bespoke treatment for each patient and the effort consumed in modifying and growing each patient’sT-cells;~~ ~~and~~

~~Although some autologous approaches to CART-cell~~ ~~have recently demonstrated encouraging clinical data, we believe our allogeneic approach provides developmental benefits.~~

Our Gene-Editing Approach to Allogeneic CART-cell Therapy

The most fundamental challenge of genome engineering is the need to specifically and efficiently target a precise DNA sequence within a complex genome. Our founder, Chairman and CEO, Dr. André Choulika, was one of the pioneers and first researchers in nuclease-based genome engineering in the early 1990s and has been integral in the development and advancement of gene-editing tools.

Our proprietary gene-editing platform relies on our capacity to custom designDNA-sequence specific cutting enzymes, or nucleases, for any chosen gene we need to modify and our capability to introduce such custom-made nucleases into the living cells we want to engineer. Our platform relies on precisely chosen protein families that can specifically recognize unique DNA sequences and can be tailored to target such sequences in any chosen gene or genetic region.

We are currently developing an allogeneic CART-cell therapy approach based on our technology platform that combines ~~single or multi-chain~~ CARs, TALEN and PulseAgile, our electroporation technology, to address the opportunities for improvement discussed above. Our approach aims to deliver anoff-the-shelf product with the following benefits:

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Market ~~Access~~access. Enable products to be shipped globally, thereby reducing deployment obstacles and providing accessibility to a broad patient population;

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Cost-effectiveness and Scalable Manufacturing. Streamlined manufacturing process has the potential to reduce ~~costs;~~costs, with approximatively hundreds of doses per batch;

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Novel Features. Develop products with specific safety and control ~~properties;~~properties, through a CAR linked to a suicide switch;

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~~Compatibility~~Engraftment.~~. Develop products taking into consideration~~ Avoid graft-versus-host disease (GvHD) through the ~~current standards~~inactivation of ~~cancer care; and~~theT-cell receptor (TCR).

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~~Consistency~~Persistence. ~~Qualify~~Manage rejection and ~~develop cancer products that are designed for optimal dosage, while reducingbatch-to-batch~~ ~~variability.~~persistence of the UCART product candidate, through notably the option to inactivate CD52 and beta2-microglobulin (ß2M) genes respectively;

TALEN—Proprietary Gene-editing Technology

The flagship nuclease structure we use for gene editing is based on a class of proteins derived from transcription activator-like effectors, or TALE. TALEN products are designed by fusing theDNA-cutting domain of a nuclease to TALE domains, which can be tailored to specifically recognize a unique DNA sequence. These fusion proteins serve as readily targetable “DNA scissors” for genome engineering applications that enable us to perform targeted genome modifications such as sequence insertion, deletion, repair and replacement in living cells.

The following diagram shows the structure of a TALEN. The DNA binding domain of TALEN is composed of DNA binding units (repeat variabledi-residues or RVDs) that each individually recognize a single base pair, and that are assembled to collectively recognize a DNA sequence. The specificity of this RVD single base pair recognition is mediated by two of the amino-acids in the RVD (NN, NI, NG, or HD), the RVDs that directly interact with the base of the DNA.

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We believe the key benefits of TALEN technology are:

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Precision. It is possible to design a TALEN that will cleave at any selected region in any gene, giving us the ability to achieve the desired genetic outcome with any gene in any living species.

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Specificity and Selectivity. TALEN may be designed to limit its DNA cleavage to the desired sequence and to reduce the risk of cutting elsewhere in the genome. This parameter is essential, especially for therapeutic applications, because unwanted genomic modifications potentially could lead to harmful effects for the patient. In addition, gene editing requires only a transient presence of TALEN, thus preserving the integrity and functionality of theT-cell’s genome.

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Efficiency. A large percentage of cells treated by the nuclease bear the desired genomic modification after treatment is completed. In our routine gene-editing processes, around 70% of theT-cells treated by TALEN to inactivate one gene copy bear the desired genomic modification. We believe TALEN’s high efficiency will be important to the cost-effectiveness of a manufacturing process involving the generation of gene-editedT-cells.

The following diagram shows the ~~mechanism~~various gene editing mechanisms enabled by ~~which TALEN inactivates, or knocks out, a gene:~~TALEN:

LOGO

We are able to assemble long arrays of modular domains with predictable specificity for a chosen sequence of DNA unique within a genome. When a TALEN is present, its TALE domains recognize its target DNA sequence and thereby direct the enzyme to the proper chromosomal location. Once bound to its target DNA sequence, the DNA cleaving-domain of the TALEN induces a DNA break at the targeted location to induce permanent DNA modifications. We believe TALEN stands out among nucleases as exceptionally precise, accurate and efficient to perform gene inactivation.

Other Types of Gene Editing Technologies

We have developed a strong expertise and capacity in meganuclease technologies, which involve enzymes capable of recognizing very large unique DNA sequences. In addition, using the flexibility of the TALE domain, we have developed new classes of custom-designed nucleases, such as compact TALEN and mega-TALE nucleases that combine meganucleases and TALEN technology. Compact-TALEN is built with a single TALE molecule fused to a fragment of a chosen meganuclease that carries limited DNA sequence recognition functionality but fully functionalDNA-cleaving activity. These chimeric proteins are smaller in size than

classical TALEN, which can facilitate their delivery to cells. In contrast, ~~mega-TALEs~~mega-TALE use afull-size meganuclease to enhance their DNA sequence recognition capacities, while demonstrating enhanced precision. We also have

discovered a new class of nuclease that we named BurrH nucleases, also based on arrays of singleDNA-base recognizing modular domains. Recently, we announced the issuance of two US CRISPR (clustered regularly interspaced short palindromic repeats) patents, covering certain uses ofRNA-guided endonucleases, such as Cas9 or Cpf1, for the genetic engineering ofT-cells.

PulseAgile—Electroporation Technology

In order to perform gene editing, we use our proprietary PulseAgile electroporation technology to introduce nucleases inside the targetT-cell where they can access the cell’s DNA. Electroporation allows messenger RNA, or mRNA, molecules coding for the nuclease to enter into the cell, where it is translated into the nuclease protein that can cut into the cell’s DNA. The mRNA molecules are rapidly degraded by the cell, which means that the nuclease is only expressed for a short time.

PulseAgile electroporation uses a unique electrical field wave-form that, in combination with a proprietary buffer solution, enables molecules, such as nucleases, to enter efficiently into the cell while maintaining a high percentage of viable cells. PulseAgile technology is particularly effective due to the shape of the electrical field that includes high voltage peaks, which are optimized to create transient holes in the cell membrane, followed by lower voltage pulses that help mRNA (for example TALEN-encoding mRNA) migrate into the cells. In addition, PulseAgile is optimized to preserve high cell viability and thus suited for large-scale manufacturing.

~~Next-Generation Products Based on Multi-chain CAR~~

Historically, CAR components have been assembled into a linear CAR molecule, known as a “single-chain” CAR. We have developed another architecture, which we call “multi-chain” CAR that is currently based on the structure of the high-affinity IgE receptor, which is normally absent in~~T-cells.~~ ~~The multi-chain CAR is composed of several membrane-bound proteins that naturally assemble at the cell’s surface and, as described below, have several benefits.~~

~~CAR architectures generally utilize a single polypeptide chain, which requires multiple appending of domains to enable a configuration that provides both theT-cell~~ ~~activation andco-stimulation~~ ~~needed for optimalT-cell~~ responses. Typically, the target-binding domain of a CAR consists of a single-chain variable fragment of an antibody comprising variable domains of large polypeptide subunits, or heavy chains, and small polypeptide subunits, or light chains, joined by a short linker peptide. This structure allows the expression of the CAR as a single-chain protein. One limitation of single-chain CAR architecture is the small number of features that it is able to add to a cell. When such protein domains are not located close to the transmembrane structure, they may be less stable or active, which limits the number of domains that can be added.

~~We have developed a novel multi-subunit CAR architecture that overcomes these structural issues and expands the functional possibilities that can be brought to aT-cell.~~ ~~Our multi-chain CAR is currently based on the architecture of a specific high affinity IgE receptor, that is normally absent fromT-cells.~~ This architecture offers the potential for inclusion of multiple intra-cellular signaling domains, optimally located at their natural distance from the membrane. In other words, our multi-chain CAR offers additional positions close to the cell membrane so that domains can be more flexibly located relative to the cell’s transmembrane structure. This also facilitates the potential implementation of multiple recognition domains, possibly allowing the recognition of not only a single antigen, but also of patterns of multiple antigen expression. This novel multi-subunit architecture would allow the construction of CARs with both improved activity and specificity and thus with an expanded range of applications.

~~CARs, whether single or multi-chain, are means to redirectT-cell~~ ~~activity toward cells bearing selected antigens. Our platform allows us to design CARs and to optimize their design depending on where and how their target is expressed on the surface of cancer cells.~~

~~The following diagram shows the distinction between single-chain CARs and multi-chain CARs:~~

Nuclease Technology andT-cells: The Design Process

OurT-cell gene-editing process involves two engineering rounds:

Step 1: Gene Editing to add Genes, such as a CAR

In the first round, genetic material is added to theT-cell’s genome using a viral vector—a benign modified virus that cannot replicate autonomously but can efficiently deliver such genetic material into a cell with which it is in contact. In particular, we use ~~allow~~ targeted integration. The genetic material added includes a ~~gene-coding~~gene coding for a CAR, which becomes a new receptor at theT-cell’s surface that allows it to recognize and bind to a target molecule that is present at the surface of other cells. At this stage, we can also add ~~additional~~other genes to these cells that confer specific properties. For example, we add suicide genes, which code for proteins that can makeT-cells susceptible to certain drugs and enable us to deplete our engineeredT-cells at our discretion by administering a drug to the patient. This system can also be integrated within the CAR itself.

Step 2: Gene Editing to Inactivate Genes, such as the TCRαµ,~~PD-1,~~ and CD52

In the second round, we use our PulseAgile electroporation technology to introduce specific TALEN mRNA into theT-cells to inactivate a number of genes that are naturally present in the genome of theseT-cells.

TCRs at the surface ofT-cells allow them to recognize cells that express foreign,non-self, antigens (for example, cells infected by a virus or cells coming from another individual).Non-modified allogeneicT-cells bear

functional TCRs and, if injected into a patient, can potentially recognizenon-self on that patient’s tissues and start to attack them. For this reason, all of our UCART product candidates undergo the inactivation of a gene coding for ~~TCRα~~TCRµ, a key component of the natural antigen receptor ofT-cells, to suppress their alloreactivity. The engineeredT-cells lack functional TCRs and are no longer capable of recognizing foreign antigens. As a result, when injected into a patient, the engineeredT-cell would not recognize the tissues of the host patient as foreign and thus would avoid attacking the patient’s tissues. This could avoid the GvHD that can sometimes be observed when allogeneicTCR-positiveT-cells are infused into some patients.

The figure below depicts the suppression of alloreactivity inT-cells engineered to lack functional TCRs. The figure summarizes experiments in which we injected mice withT-cells engineered for the inactivation of ~~TCRα~~TCRµ while injecting other mice withnon-engineeredT-cells with functional TCRs. We then measured the effects of such injections on mean body weight, which serves as a proxy for the impact of GvHD.

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During the manufacturing process, the T cells from a healthy donor are first engineered. The CAR gene is transduced and cell attributes like the TCR alpha gene are knocked out by TALEN. Then, theT-cells of our UCART products are amplified. The desired~~TCR-alpha~~ TCR alpha deleted are finally purified from the cells that may still bear a TCR, and are finally frozen. We perform a battery of specialized testing techniques and various quality assurance and quality control assays to further validate cellular functional integrity following gene editing.

The lack of a TCR at the surface of our UCART product candidates is a key feature that allows them to be used as allogeneicoff-the-shelf products. Other genes can also be inactivated in this round to confer additional specific attributes to theT-cells. They can be made resistant, and therefore compatible, with specific medical regimens used during the course of cancer treatments. For example, we inactivate the CD52 gene, which codes for the target of alemtuzumab, a monoclonal antibody sometimes used in CLL patients, that would otherwise destroy our engineeredT-cells. Likewise, we believe we can inactivate the deoxycytidine kinase (dCK) or glucocorticoid receptor (GR) genes in order to make ourT-cells respectively resistant to purine nucleotide analogs (e.g., fludarabine, clofarabine or cytarabine) or to corticoids that are used for several types of cancer patients.

The following diagram shows the key stages in our engineering of ~~UCART19:~~UCARTCS1:

LOGO

Next-Generation Products – Inactivate Additional Genes, such as ß2M andPD-1

The allogenic CART-cell approach developed by Cellectis aims at increasing accessibility to treatment for patients by using healthy donor cells to manufacture CART-cells. As evidenced above, the inactivation of the TRAC gene reduces the risk of graft vs. host disease, and the lymphodepletion regimen of the patients aims at supporting early engraftment of the candidate product.

We are investigating the inactivation of the beta2-microglobulin (ß2M) gene to increase persistence of allogenic cells in this context. ß2M is necessary for presentation of antigens on class I major histocompatibility complex (MHC) to cytotoxicT-cells. Allogenic TRAC/ß2M doubleknock-out CART-cells infused into a patient are expected not to be recognized by the patient’s ownT-cells and therefore to show prolonged survival after patients’T-cells recover following lymphodepletion.

We developed severalß2M-specific TALEN allowing high efficiency of gene inactivation in combination with TRAC TALEN (up to 88% doubleknock-out). We have shown on human cells and on mouse cells that ß2M inactivation improves allogenic cell survival in the presence of alloreactiveT-cells, and we are pursuing the ß2M inactivation approach for our preclinical candidate UCARTCLL1.

Our engineeredT-cell could also be made insensitive to inhibition signals, which diminishes immune system activity, that may be present within the tumor microenvironment and that usually blockT-cell attacks. For example, we inactivate the programmed cell death 1~~PD-1~~(PD-1) gene in our engineeredT-cells ~~so that they would no longer be subject~~in order to suppress the checkpoint regulator inhibition by tumors expressing~~PDL-1,~~PD-L1, a common anti-immune defense mechanism found in cancer.

The following diagram shows the inactivation of thePD-1 gene to suppress checkpoint inhibition in theT-cell:

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Using our ability to add and to inactivate genes, our platform has the ~~flexibility~~potential to deliver smartT-cells designed for specific indications and purposes.

~~Key Benefits~~Next-Generation Products – Armored CARs

While CART-cell therapies have led to complete remission in previously untreatable diseases such relapse/refractory ALL, not all patients respond, and even among those that respond, a fraction end up relapsing. There is therefore a need to investigate strategies to make CART-cells even more effective, such as boosting their activity by overexpression of ~~our UCART Approach~~

~~We believe our~~an immunomodulatory molecule (i.e. a cytokine or a costimulatory receptor). In order to limit toxicity effects due to immunostimulatory molecules being produced uncontrollably and systemically, we have developed strategies exploiting cellular endogenous pathways to restrict expression of a gene of interest only when CAR ~~T approach~~T-cells are activated. This is made possible by inserting genes of interest at a desired position in the genome by combining a locus-specific nuclease and ~~manufacturing process has the potential to provide the following benefits:~~

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~~Market Access~~~~. Enable products to be shipped globally, thereby reducing deployment obstacles.~~

~~Our UCART product candidates~~a donor template vectorized with an adeno-associated viral (AAV) vector. SincePD-1 and CD25 are intended to enable universal market access driven by an allogeneic approach. Current autologous treatments require dedicated infrastructure, which could limit their availability to only a few select sites. Because our UCART product candidates are designedknown to be ~~frozen~~upregulated uponT-cell activation, inserting certain cytokine coding sequence under the control ofPD-1 or CD25 genetic regulatory elements allows secretion of that certain cytokine only upon activation of the CART-cells and ~~availableoff-the-shelf,~~ ~~they~~enhancesin vitro antitumor activity.

This strategy could be ~~shipped globally at any time and administered immediately to patients when needed, including in local clinics.~~

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~~Cost-effectiveness~~~~. Streamlined manufacturing process has the potential to reduce costs.~~

~~Our manufacturing process is a benefit to our UCART product candidate line that could contribute~~extended to the ~~design~~use of ~~a reasonably priced product. Our manufacturing process produces UCART product candidates from healthy, selected, screened and tested donorT-cells.~~ ~~Moreover, because our process is powered by our nucleases and our proprietary PulseAgile electroporation systems, we expect~~various genetic loci to ~~be able to inactivate~~express genes ~~in a highly efficient manner that avoids harmingT-cells~~ ~~during processing, which could allow us to manufacture quality UCART products~~with therapeutic benefits at ~~high yields. This could enable us to manufacture in bulk, and we expect thatT-cells~~ from one healthy donor, and one manufacturing run of UCART, could be used to create hundreds of doses of product. These efficiencies could allow us to reduce costs to patients and produce competitive gross profit margins.

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~~Novel Features~~~~. Develop products~~desirable expression level or with ~~specific safety and control properties.~~

~~We aim to engineerT-cells~~ ~~for specific clinical results that enhance safety and provide greater control over cellular activity. For example, our research includes disablingT-cells~~ ~~from attacking a patient’s healthy tissues, designingT-cells~~ ~~to be compatible with standard oncology treatments, enabling our engineeredT-cells~~ ~~to surpass key immune checkpoint regulators that can protect tumors from the immune system, and building into our products a suicide gene that directs the natural clearance of allogeneicT-cells~~ ~~with the addition of a drug.~~

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~~Compatibility~~~~. Develop products taking into consideration the current standards of cancer care.~~

~~Our research also aims at allowingT-cells~~ to resist and be compatible with compounds that cancer patients are exposed to, including standard oncology treatments, such as specific monoclonal antibody therapies, corticoids, or other relevant chemotherapies. We are pursuing treatment options that would allow patients to be treated with our engineered~~T-cells~~ ~~after or during treatment with traditional approaches that would impairT-cell~~ ~~function or viability.~~

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~~Consistency~~~~. Qualify and develop cancer products that are designed for optimal dosage, while reducingbatch-to-batch~~ ~~variability.~~

~~Our frozen,off-the-shelf~~ UCART product candidates are intended to be produced pursuant to cGMP and are extensively controlled. Like other pharmaceutical products, we expect that their quality will be controlled to ensure consistency over time and from batch to batch. This is a significant difference from autologous approaches currently reported to be in clinical development. In these autologous settings, where the donor of~~T-cells~~ ~~is the very patient who will receive theCAR-bearing~~ ~~cells,~~ a specific ~~batch ofT-cells~~ ~~must be made for each patient and in some cases, sufficient T cells may not be available from the patient to create the autologous product.~~temporal or regional expression pattern.

UCART Manufacturing: How can we turn a procedure into a ~~large scale,~~large-scale, widely available drug?

AutologousCAR-T cell approaches are therapeutic procedures conducted for each patient, which involve the engineering ofT-cells by addition of a transgene coding for a chimeric antigen receptor into the patient’s own T cells. Our UCART approach goes one step further in engineering and also in moving the CAR concept from apatient-by-patient therapeutic procedure to anoff-the-shelf widely available pharmaceutical compound.

The manufacturing process of our allogeneic CART-cell product line, Universal CARTs or UCARTs, yields frozen,off-the-shelf, allogeneic, engineered CART-cells. UCARTs are meant to be readily available CART-cells for a large patient population. The specificity of those allogeneic therapies is thatT-cells from healthy donors are genetically edited with our proprietary technology, TALEN, to seek and destroy cancer cells. TALEN-based gene editing is designed to suppressT-cell alloreactivity (and, for certain UCART product candidates, to confer resistance to alemtuzumab) to theT-cells.

Our UCARTs are designed and manufactured through a common platform that relies on defined unit operations and technologies combined into a single process adapted to each individual UCART. The process is gradually developed from small to larger scales, incorporating elements that are eventually used in GMP conditions. Notwithstanding this central unit operations based model, each product is unique and for each new UCART, a developmental phase is necessary to individually customize each engineering step and to create a robust procedure that can later be implemented in a GMP environment to ensure the production of clinical batches. This work is performed in our research & development environment to evaluate and assess variability in each step of the process in order to define the most reliable experimental conditions.

The following diagram summarizes the generic UCART production process made of distinct unit operations. The engineering steps for transduction and electroporation can take place one before another (and several times), depending on the product.

LOGO

On October 28, 2015, we announced that we completed a series of three production runs of UCART19 confirming the transfer of Cellectis’ manufacturing process into clinical grade, GMP conditions. This important milestone showed that UCARTs can be manufactured in GMP conditions and demonstrated the industrial production of UCART19, as well as the capacity of Cellectis’ pipeline of UCART product candidates to be manufactured for clinical investigations. ~~Since then, UCART19 manufacturing of clinical supplies has been conducted by CELLforCURE to support the two clinical trials opened with UCART19 in the United Kingdom.~~ On November 15, 2016, we announced that we completed a series of production runs of UCART123 at CELLforCURE, our CMO, to support the UCART123 Clinical Trials for which we filed an IND, the FDA approval of which was announced on February 6, 2017. ~~This year,~~In November 2017, we ~~also proceeded~~started a series of production runs of UCART22 at CELLforCURE, to support the UCART22 Clinical Trial. In November 2018, we started a series of production runs of UCARTCS1 at CELLforCURE to support the UCARTCS1 Clinical Trial, and started a new campaign of UCART123 in November 2018 at CELLforCURE

On July 27, 2017, we announced the signature of a capacity production agreement with MolMed, a biopharmaceutical company based in Milan, Italy that has Contract Manufacturing Operations facilities for the production of GMP grade CART-cells. At the end of 2018, we completed the technology transfer to MolMed for the production of UCART123, and it is our intention to start production of UCART products at MolMed in 2019.

In June 2018, CELLforCURE informed us that GMP grade batches of dimethyl-sulfoxide (DMSO) and Dulbecco’s phosphate-buffered saline (dPBS) used during manufacturing of UCART123 and UCART22 contained particles, and that some batches of final product had been produced using this DMSO and dPBS batches. We informed the FDA and voluntarily decided not to use the batches that could potentially contain particles until receiving clearance from the FDA. Subsequently, CELLforCURE informed us that the water for injection (WFI) they had used during manufacturing contained glass particles and that some batches of UCART123 and UCART22 had been manufactured with this WFI. We informed the FDA and voluntarily decided not to use the batches that could potentially contain glass particles until receiving clearance from the FDA. We have been, since then, working with the FDA to assess the risk-benefit profile of vials produced during these campaigns. The batches of UCARTCS1 and UCART123 that were made during the fourth quarter of 2018 did not use these DMSO, dPBS and WFI batches and were successfully manufactured.

We aim to continuously improve our manufacturing processes for ~~UCARTCS1~~better safety and ~~UCART22~~robustness of our product lines.

Towards manufacturing autonomy: building twostate-of-the-art plants

In order to achieve manufacturing autonomy, we have started the construction of two manufacturing facilities, dedicated to starting materials for clinical supply and clinical & commercial UCART products, respectively.

SMART (Starting MAterial Realization forCAR-T products) will be a ~14,000 sq. ft.in-house manufacturing in Paris, France. SMART will be dedicated to the ~~same CMO.~~

~~UCART Pipeline~~

production of certain raw material for clinical supply, with the potential to supply commercial starting material. We ~~are developing a series of product candidates for advanced hematologic cancers.~~

~~Our lead immuno-oncology product candidates, which we refer to as Universal CART-cells~~ ~~(UCARTs), are allogeneic CART-cells~~ ~~engineered~~expect this manufacturing facility to be ~~used as an“off-the-shelf”~~ ~~treatment~~operational in 2020.

IMPACT (Innovative Manufacturing Plant for ~~any patient with~~Allogeneic Cellular Therapies) will be a ~~particular cancer type. Each UCART product candidate targets a selected antigen expressed on tumor cells and bears~~

specific engineered attributes, such as inhibition of alloreactivity and compatibility with specific medical regimens that cancer patients may undergo. UCART is the first therapeutic product line that we are developing with our gene-editing platform to address unmet medical needs in oncology. We are focusing our initial internal pipeline~80,000 sq. ft. facility located in the hematologic cancer space, targeting diseases with high unmet needs such as acute lymphoblastic leukemia (ALL), acute myeloid leukemia (AML), blastic plasmacytoid dendritic cell neoplasm (BPDCN), multiple myeloma (MM) and different types of lymphomas. In December 2016, we filed an IND for our lead product candidate, UCART123 in AML and in BPDCN and in February 2017, we received FDA approval to initiate UCART123 Clinical Studies. All of our other product candidates are currently in the late~~pre-clinical~~ ~~/ early manufacturing phase, and the following chart highlights some of these product candidates:~~

*	~~or foreign equivalent~~

**	~~UCART19 is exclusively licensed to Servier and under a joint clinical development program between Servier and Pfizer.~~

~~UCART19 for Acute Lymphoblastic Leukemia~~

~~UCART19 is an allogeneic,off-the-shelf~~ ~~product candidate designed to exhibit high efficacy in fighting hematological malignancies bearing theB-lymphocyte~~ antigen CD19, or CD19. In November 2015, Servier acquired the exclusive rights to UCART19 from Cellectis. Servier and Pfizer collaborate on a joint clinical development program for UCART19, and Pfizer has acquired exclusive rights from Servier to develop and commercialize UCART19 in the United States.

~~Targeted Indications~~

~~Acute Lymphoblastic Leukemia (ALL)~~

ALL is a heterogeneous hematologic disease characterized by the proliferation of immature lymphoid cells in the bone marrow, peripheral blood, and other organs. The proliferation and accumulation of blast cells in the marrow results in suppression of hematopoiesis and, thereafter, anemia, thrombocytopenia, and neutropenia. Extramedullary accumulations of lymphoblasts may occur in various sites, especially the meninges, gonads, thymus, liver, spleen, or lymph nodes. The~~age-adjusted~~ incidence rate of ALL in the United States is 1.7 per 100,000 individuals per year, with approximately 5970 new cases and 1440 deaths estimated in 2017. The median age at diagnosis for ALL is 15 years with 56.1% of patients diagnosed at younger than 20 years of age. In contrast, 28% of cases are diagnosed at 45 years or older and only 11.7 % of patients are diagnosed at 65 years or older. ALL represents 75% to 80% of acute leukemia among children, making it the most common form of childhood leukemia; by contrast, ALL represents approximately 20% of all leukemia among adults. The cure rates and survival outcomes for patients with ALL have improved dramatically over the past several decades, primarily among children. Improvements are largely owed to advances in the understanding of the molecular genetics and pathogenesis of the disease, the incorporation of risk-adapted therapy, and the advent of new targeted agents. Data from the Surveillance, Epidemiology, and End Results (SEER) database have shown a 5-year overall survival (OS) of 86% to 89% for children. Adults have the poorest~~5-year~~ OS rate of 24.1% for patients between the ages of 40 and 59 years and an even lower rate of 17.7% for patients between the ages of 60 and 69 years. Despite great progress in the development of curative therapies, ALL remains a leading cause of pediatric cancer-related mortality for patients

~~presenting with a relapsed or refractory disease. New therapies are needed to overcome chemotherapy resistance and reducenon-specific~~ ~~treatment associated side effects.~~

~~Product Features~~

~~UCART19 is an allogeneicT-cell~~ ~~product intended for the treatment of CD19-expressing hematologic malignancies such as ALL.~~

~~UCART19 is designed to become active, proliferate, secrete cytokines and kill CD19-bearingB-cell~~ malignancies upon contact with such cells, following administration to patients. Activation of UCART19 is driven by contact between its anti-CD19 CAR and the CD19 protein on the surface of tumor cells.

UCART19 cells bear a CAR targeting the CD19 antigen that drives their capacity to kill CD19-bearing cells. Moreover, as all UCART product candidates, UCART19 lacks the TCR responsible for recognition of~~non-self~~ ~~antigens by theT-cells,~~ ~~which allows use of healthy donorT-cells~~ ~~to produce UCART19, with reduced potential for GvHD. In addition, some UCART19 cells lack CD52, a protein expressed on the cell surface that makesT-cells~~ ~~sensitive to alemtuzumab. This feature permits the use of UCART19 in patients recently treated or being treated with the immunosuppressing/lymphodepleting agent alemtuzumab.~~

~~UCART19 activation could potentially lead to eradication of CD19-expressing cancer cells throughT-cell~~ ~~mediated killing of such cancer cells and potentiallypro-inflammatory~~ ~~immune system production as well as CART-cell~~ ~~amplification.~~

~~Clinical Findings~~

~~In December 2017, preliminary results from the first twelve patients in the UCART19 Clinical Studies were presented at the 59~~^th ~~American Society of Hematology (ASH) Annual Meeting. Thesefirst-in-human~~ data show 83% complete remission rate across the adult and pediatric patient population. Cytokine release syndrome (CRS) was mild and manageable except in one adult patient treated with UCART19 at the first dose level, who developed CRS Grade 4 and neutropenic sepsis leading to death at day 15. Only two cases of Grade 1 acute graft versus host disease (GvHD) occurred and no severe neurotoxicity was observed.

~~We are encouraged by these promising preliminary results reported for the UCART19 Clinical Studies.~~

~~Development Status~~

In 2016, Servier commenced the UCART19 Clinical Studies—a Phase I clinical study in pediatric acute lymphoblastic leukemia (ALL), the PALL study, and a Phase I clinical study in adult patients with ALL, the CALM study, each of which was approved by the MHRA.

~~The PALL Study is commenced in United Kingdom at UCL Great Ormond Hospital (London), in Belgium at Het Kinderziekenhuis Prinses Elisabeth (Gent), and in France at Hôpital Robert-Debré (Paris).~~

The CALM Study is commenced in United Kingdom at King’s College Hospital NHS Foundation Trust (London), in United States at the Hospital of the University of Pennsylvania (Philadelphia, Pennsylvania) and at University of Texas MD Anderson Cancer Center (Houston, Texas), and in France at Hôpital Saint-Antoine (Paris).

~~UCART123 for Acute Myeloid Leukemia (AML) and Blastic Plasmacytoid Dendritic Cell Neoplasm (BPDCN)~~

~~UCART123 is an allogeneic engineeredT-cell~~ ~~product designed for the treatment of hematologic malignancies expressing the alpha chain of theinterleukin-3~~ ~~receptor, or CD123, and is currently being developed for the treatment of acute myeloid leukemia (AML) and blastic plasmacytoid dentric cell neoplasm (BPDCN).~~

~~Targeted Indications~~

~~Acute Myeloid Leukemia (AML)~~

Acute myeloid leukemia (AML) is a form of cancer that is characterized by infiltration of the bone marrow, blood, and other tissues by proliferative, clonal, abnormally and/or poorly differentiated cells of the hematopoietic system called blast cells. These cells interfere with normal hematopoiesis, thus contributingU.S. East Coast. IMPACT will be dedicated to the ~~bone marrow failure which is the most common underlying cause~~production of ~~death. AML is the most common type of acute leukemia in adults with anage-adjusted~~ incidence rate in the United States of 4.2 per 100,000 individuals per year, with approximately 21,380 new cases and 10,590 deaths estimated in 2017. Although it can occur in children and adults, AML is primarily a disease of the elderly. The median age at onset is 68 years and only 16.5% of patients are younger than 45 years of age at diagnosis While complete response rates can be as high as 80% in patients undergoing initial induction cytotoxic chemotherapy, the majority of AML patients will ultimately be diagnosed with relapsed or refractory disease with a poor prognosis. The outcome in older patients who are unable to receive intensive chemotherapy without unacceptable side effects remains dismal, with a median survival of only 5 to 10 months. CD 123 is highly expressed on acute myeloid leukemia (AML) leukemic stem cells and blast cells, as well as in other hematologic malignancies, and constitutes an attractive target for AML.

~~Blastic Plasmacytoid Dendritic Cell Neoplasm (BPDCN)~~

Blastic plasmacytoid dendritic cell neoplasm (BPDCN) is a rare and aggressive hematological neoplasm classified among AML in the 2008 World Health Organization (WHO) classification of hematologic malignancies, and reclassified under myeloid neoplasms, acute leukemia in the 2016 WHO classification. BPDCN is a rare myeloid disease characterized by the clonal proliferation of precursors of plasmacytoid dendritic cells. There are no formal studies on the incidence of BPDCN in the general population. The few available data reported indicate that its overall incidence is extremely low, accounting for 0.44% of all hematologic malignancies and 0.7% of cutaneous lymphomas. Moreover, the leukemic form of disease is a rare phenomenon, representing fewer than 1% of cases of acute leukemia. The disease may occur at any age, but most patients are elderly men who present with skin lesions and/or involved lymph nodes, spleen, and bone marrow. Given its rarity and only recent recognition as a distinct clinico-pathological entity, no standardized therapeutic approach has been established for BPDCN and the optimal therapy remains to be defined. Although transient responses are seen to combination chemotherapy regimens used to treat acute leukemia or lymphoma, most patients relapse with drug-resistant disease with a median overall survival rate of 9 to 13 months, irrespective of the initial presentation of the disease.

~~Product Features~~

~~UCART123 is an allogeneicT-cell~~ ~~product candidate intended for the treatment of CD123-expressing hematologic malignancies.~~

~~Pre-clinical~~ ~~Findings~~

~~UCART123 has been evaluated both in vitro and in animal studies, with promising results.~~

~~In vitro~~ ~~studies demonstrated efficient killing of human CD123-bearing cell lines by UCART123. In addition, UCART123 has also demonstrated efficient killing of human CD123-expressing cells derived from~~

AML and BPDCN patients. Animal studies were conducted in mice injected both with UCART123 and human CD123-bearing tumor cells, and have shown anti-tumor activity in an immunodeficient mouse model. In addition, in another animal model, limited toxicity against normal, healthy cells, has been observed.

~~UCART123 was also tested for its potential to induce GvHD. Mice receiving unmodifiedT-cells~~ ~~from a human donor showed GvHD, while mice receiving the UCART123 cells that lack the TCR showed no sign of GvHD.Pre-clinical~~ ~~and translational activities on UCART123 in AML and BPDCN were performed in collaboration with Weill Cornell and MD Anderson Cancer Center, respectively.~~

~~Development Status~~

On December 14, 2016, Cellectis received unanimous approval from the National Institute of Health’s Recombinant DNA Advisory Committee (“ RAC”) for two proposed Phase I protocols for UCART123. In February 2017, the FDA granted Cellectis an Investigational New Drug (IND) approval to conduct a Phase I clinical study with UCART123 in AML and BPDCN. The initial patients were enrolled during 2017, at Weill Cornell Medical College for AML and at MD Anderson Cancer Center for BPDCN. Due to a death in the BPDCN study, both trials were put on hold in September 2017 but then cleared to resume by the FDA, in November 2017, based on revised protocols.

~~UCART22 for Acute Lymphoblastic Leukemia (ALL)~~

~~UCART22 is an allogeneic engineeredT-cell~~ ~~product candidate designed for the treatment of Acute Lymphoblastic Leukemia (ALL).~~

~~See “Item 4.B.—Business Overview—UCART Pipeline—UCART19 for Acute Lymphoblastic Leukemia—Targeted Indications—Acute Lymphoblastic Leukemia” for more information on ALL.~~

~~Product Features~~

~~UCART22 is an allogeneic engineeredT-cell~~ product candidate intended for the treatment of CD22-expressing hematologic malignancies. UCART22 is designed to become active, proliferate, secrete cytokines and kill CD22 expressing cells (i.e. either CD22 positive tumor cells or~~non-malignant~~ CD22 positive B lineage cells). UCART22 bears a CAR targeting the CD22 antigen, providing specificity for CD22 expressing cells. As with all UCART products, UCART22 lacks the TCR and is intended to be used in an allogeneic context. In addition, some UCART22 cells lack CD52, a protein expressed on the cell surface that makes~~T-cells~~ ~~sensitive to alemtuzumab, a drug often used to treat CLL patients. This feature should allow for improved engraftment of the cells in conjunction with a potential alemtuzumab treatment.~~

~~UCART22 activity could potentially lead to eradication of CD22-expressing cancer cells throughT-cell~~ ~~mediated killing,pro-inflammatory~~ ~~cytokine production as well as CART-cell~~ ~~amplification.~~

~~Pre-clinical~~ ~~findings~~

~~UCART22 has been evaluated both~~~~in vitro~~ ~~and in animal studies, with promising results.~~

~~In vitro~~ studies demonstrated efficient killing of human CD22-bearing cell lines by UCART22. In addition, UCART22 has also demonstrated efficient killing of human CD22-expressing cells derived from ALL patients. Animal studies were conducted in mice injected both with UCART22 and human CD22-bearing tumor cells, and have shown anti-tumor activity in an immunodeficient mouse model. Further~~in vitro~~ ~~and~~~~in vivo~~ ~~studies are ongoing to further investigate the safety and the activity of UCART22.~~

~~Pre-clinical~~ ~~and translational activities on UCART22 in ALL are performed in collaboration with MD Anderson Cancer Center.~~

~~Development Status~~

~~UCART22 is at the latepre-clinical~~ / early manufacturing stage of development. The manufacturing process has been transferred to CELLforCURE, our CMO. Manufacturing of clinical grade UCART22 at large scale in accordance with GMP was initiated at the end of 2017.

~~UCARTCS1 for Multiple Myeloma (MM)~~

~~UCARTCS1 is an allogeneic engineeredT-cell~~ ~~product candidate designed for the treatment ofCS1-expressing~~ ~~hematologic malignancies which is being developed in multiple myeloma (MM).~~

~~Targeted Indication Multiple Myeloma~~

Multiple myeloma (MM) is a clonal plasma cell malignant neoplasm that is characterized by the proliferation of a single clone of plasma cells producing a monoclonal immunoglobulin. This clone of plasma cells proliferates in the bone marrow and often results in extensive skeletal destruction with osteolytic lesions, osteopenia, and/or pathologic fractures. Additional disease-related complications include hypercalcemia, renal insufficiency, anemia, and infections. MM accounts for approximately 10% of hematologic malignant disorders. The annual incidence,~~age-adjusted~~ to the US population, is 6.6 per 100,000, resulting in over 30,000 new patients in the United States each year. The median age at onset is 69 years, and only 3.5 % of patients are younger than 45 years of age at diagnosis. Nine drugs have been approved over the past fifteen years for the treatment of MM, substantially expanding the number of treatment regimens available for patients in all stages of the disease. In the last decade, survival of multiple myeloma (MM) patients has markedly improved with a median survival of approximately 5 to 7 years but with major variation depending on host factors, stage of the disease, cytogenetic abnormalities, and response to therapy. However, despite this progress, patients with disease refractory to both immunomodulatory drugs (IMiDs) and proteasome inhibitors have a median overall survival (OS) of only 9 months.

~~Product Features~~

~~UCARTCS1 is an allogeneicT-cell~~ drug candidate intended for the treatment of CS1 (also as known as SLAMF7)-expressing hematologic malignancies, in particular MM. UCARTCS1 is designed to become active, proliferate, secrete cytokines and kill CS1 expressing cells. As CS1 is expressed on the cell surface of CD8~~T-cells,~~ ~~CS1 will be inactivated inT-cells~~ ~~prior to transduction with a viral vector encoding ananti-CS1~~ CAR. The inactivation of CS1 may improve the production and activity of UCARTCS1 by preserving the CD8 T cell population. In addition, as with all UCART products, UCARTCS1 lacks the TCR and is intended to be used in an allogeneic context.

~~Pre-clinical~~ ~~Findings~~

~~In vitro studies demonstrated efficient killing of humanCS1-bearing~~ ~~cell lines by UCARTCS1. In addition, UCARTCS1 has also demonstrated efficient killing of humanCS1-expressing~~ ~~cells derived from MM patients. Furthermore, whilenon-gene-editedT-cells~~ ~~expressing ananti-CS1~~ ~~CAR display limited cytolytic activity in vitro against MM cell lines and result in a progressive loss of CD8T-cells,CS1-gene-edited~~ ~~CAR cells (UCARTCS1) display significantly increased cytotoxic activity, with the percentage of CD8T-cells~~ ~~remaining unaffected. Experiments in an orthotopic MM mouse model showed that UCARTCS1 was able to mediate an~~~~in vivo~~ ~~anti-tumoral activity. Further in vitro and in vivo studies are ongoing to further investigate the safety and the activity of UCARTCS1.~~

~~Pre-clinical~~ ~~and translational activities for UCARTCS1 in MM are performed in collaboration with the MD Anderson Cancer Center.~~

~~Development Status~~

~~UCARTCS1 is at the latepre-clinical~~ ~~/ early manufacturing stage of development.~~

~~Manufacturing process transfer has been completed to CELLforCURE, our third-party manufacturing contractor, for clinical supplies of UCARTCS1.~~

~~Our Strategic Alliances~~

~~We have signed collaboration agreements with Pfizer and Servier, which we believe validate our research and approach toCAR-T.~~ ~~Our strategic alliances include potential milestone payments to us of up to $3.9 billion and royalties on future sales.~~

~~Research Collaboration and License Agreement with Pfizer~~

~~In June 2014, we entered into a Research Collaboration and License Agreement with Pfizer pursuant to which we collaborate to conduct discovery andpre-clinical~~ ~~development activities to generate CART-cells~~ ~~directed atPfizer-~~ ~~and Cellectis-selected targets in the field of human oncology. We granted Pfizer an exclusive, worldwide, royalty-bearing, sublicensable license, on atarget-by-target~~ ~~basis, under certain of our intellectual property to make, use, sell, import, and otherwise commercialize products directed at thePfizer-selected~~ ~~targets in the field of human oncology.~~

~~Under the agreement, we are obligated to use commercially reasonable efforts to develop, for each Cellectis-selected target, at least one product candidate. Pfizer granted us anon-exclusive,~~ worldwide, royalty-free license, with sublicensing rights under certain conditions, under certain of its intellectual property to conduct research, and to make, use, sell, import and otherwise commercialize products directed at Cellectis-selected targets.

~~Pursuant to the agreement, Pfizer made an upfront,non-refundable~~ $80.0 million payment to us, concurrent with Pfizer’s €25.8 million equity investment in our company. In addition, the strategic alliance provides for payments of up to $185.0 million per product that is directed against a~~Pfizer-selected~~ ~~target, with aggregate potentialpre-clinical,~~ clinical and commercial ~~milestone payments totaling up to $2.8 billion.~~UCART products. We are also eligible to receive from Pfizer tiered royalties on annual net sales of any products that are commercialized by Pfizer that contain or incorporate certain of our intellectual property at rates in the high single-digit percentages.

Except as required of us by our collaboration agreement with Servier, until the earlier of (1) the completion or termination of a four-year term or (2) the filing by Cellectis of an IND for certain targets to which we retain rights, we and our affiliates may not grant rights under certain of our intellectual property and intellectual property developed in the course of the collaboration to develop or commercialize CAR~~T-cells~~ ~~in the field of human oncology, other than certain specifiednon-commercial~~ ~~collaborations.~~

~~Unless earlier terminated in accordance with the agreement, our agreement with Pfizer will expire on aproduct-by-product~~ ~~andcountry-by-country~~ basis, until the later of (1) the expiration of the last to expire of the licensed patents covering such product; (2) the loss of regulatory exclusivity afforded such product in such country, and (3) the tenth anniversary of the date of the first commercial sale of such product in such country; however, in no event shall the term extend, with respect to a particular licensed product, past the twentieth anniversary of the first commercial sale for such product. At any time after the first anniversary of the effective date of the agreement, Pfizer will have the right to terminate the agreement at will upon 60 days’ prior written notice, either in its entirety or on a~~target-by-target~~ ~~basis. Either party may terminate the agreement, in its entirety or on atarget-by-target~~ basis, upon 90 days’ prior written notice in the event of the other party’s uncured material breach. The agreement may also be terminated upon written notice by Pfizer at any time in the event that we become bankrupt or insolvent.

~~Research, Product Development, Option, License and Commercialization Agreement with Servier~~

~~In February 2014, we entered into a Research, Product Development, Option, License and Commercialization Agreement with Servier. Pursuant to~~expect this agreement, we are responsible for the research and development up to and including the Phase I clinical trial of candidate products directed against five targets, including the UCART19 product candidate. Pursuant to the agreement, we granted Servier the right to exercise an exclusive option to obtain an exclusive, worldwide license, on a product~~candidate-by-product~~ candidate basis, with respect to each product candidate selected by Servier and developed under the agreement. Upon Servier’s exercise of each option, we shall grant Servier an exclusive, worldwide, royalty-bearing, sublicensable license under certain of our patents and~~know-how~~ covering the respective product candidate to develop, manufacture and commercialize such product in the field of anti-tumor adoptive immunotherapy, and Servier will assume responsibility for the further clinical development, manufacture and commercialization of such product. During the term of the agreement, we are prohibited from researching, developing, or commercializing any product directed against a target that is used for the same purpose as it is used with a product candidate developed under the agreement.

~~Pursuant to the agreement, Servier made an upfront payment of $10.0 million and, upon its exercise of each license option provided for in the agreement, Servier will pay us a lump sum license fee.~~

In November 2015, we entered into an amendment to our initial collaboration agreement with Servier, which allowed for an early exercise of Servier’s option with respect to UCART19 and other product candidates. Pursuant to this amendment, Servier has exercised its option to acquire the exclusive worldwide rights to further develop and commercialize UCART19. In connection with the entry into the amendment to the collaboration agreement, Servier made an upfront payment of $38.5 million, excluding taxes.

As of December 31, 2017, we are eligible to receive from Servier aggregate additional payments of up to $1,064 million, comprising payments upon the exercise of options granted to Servier under the agreement and payments upon the occurrence of certain specified development and commercial milestones. Pursuant to the agreement, we are also eligible to receive tiered royalties ranging in the high single-digit percentages based on annual net sales of commercialized products.

Unless earlier terminated, the agreement will expire upon the expiration of the last to expire of the patents covering a product licensed pursuant to the agreement. Either party may terminate the agreement for the other party’s uncured material breach upon 90 days’ prior written notice to the breaching party, or 30 days’ notice if such breach relates to a payment obligation. The parties may also terminate the agreement by mutual consent. The agreement immediately and automatically terminates upon the expiration of Servier’s last license option in the event Servier has not exercised any option to license in accordance with the agreement prior to such expiration. Servier has the right, at its sole discretion, to terminate the agreement in its entirety or with respect to specific products, upon three months’ prior written notice to us. Servier may also terminate the agreement at any time for product-related safety reasons. Either party may terminate the agreement in the event of the other party’s bankruptcy or insolvency. In the event that Servier does not exercise its option to license a product candidate, we may independently pursue all activities related to such product candidate and/or license such product candidate and the associated intellectual property to a third party. For such purpose, Servier granted us a~~non-exclusive,~~ ~~sublicensable license under any such Servier-controlled intellectual property for which we will pay tiered royalties on annual net revenues at rates ranging in the low single-digit percentages.~~

~~Collaboration with Cornell University and MD Anderson Cancer Center~~

~~In 2015, we entered into alliances with Cornell University and the MD Anderson Cancer Center to accelerate the development of our lead candidate products.~~

~~Alliance with Cornell University~~

On June 2, 2015, Cornell University and Cellectis entered into a strategic research alliance to accelerate the development of a targeted immunotherapy for patients with acute myeloid leukemia (AML). Under our alliance

with Cornell University and Cellectis will conduct research and develop clinical strategies with the objective of implementing and conducting one or more clinical trials at Weill Cornell on UCART123 and potentially other product candidates in AML. Cellectis is responsible for funding the research programs, and Cornell University and Cellectis will work together to develop and implement improvements to the research plan for the programs. The objectives of the collaboration are to demonstrate functionalities and specificity of UCART123 in vitro and in vivo, define the~~pre-clinical~~ package required for a clinical trial application, prepare the clinical trial protocol and the regulatory and other study-specific documents, and discovery research for the identification of novel targets in AML patients potentially enabling the development of additional CARs for AML.

~~In connection with the alliance, Cellectis was responsible for generating and~~ manufacturing ~~UCART123 and performing in vitro and in vivopre-clinical~~ activities on tumor cell lines and in animal models. Cornell University was responsible for evaluating UCART123 activity against primary AML samples and in animal models, as well as evaluating toxicity against HSCs in animal models. Cornell University is also working on the development and implementation of correlative studies. In addition, Cornell University and Cellectis collaborate on the preparation of the different versions of the clinical trial protocols. Finally, Cellectis and Cornell University are working on target discovery in the AML area, in order to identify new potential targets for AML and generate new potential candidate products for AML patients.

On April 2017, we enter into a clinical trial agreement with Cornell University acting for and on behalf of its Joan and Sanford I. Weill Medical College and The New York Presbyterian Hospital (collectively referred to as Weill Cornell) under which Weill Cornell performs the UCART123 clinical study for AML.

~~Alliance with The University of Texas M.D. Anderson Cancer Center~~

Pursuant to the Strategic Alliance Agreement, we entered, in March 2017, into a study order with MD Anderson Cancer Center, according to which MD Anderson Cancer Center performs the UCART123 clinical study for BPDCN.

~~On January 2018, we entered into a new study order with MD Anderson Cancer Center in order to expand the performance of the UCART123 clinical study in AML to MD Anderson Cancer Center.~~

Raw Materials

We are currently dependent on specialized third parties, who are subject to stringent manufacturing requirements and regulations, for the supply of various critical and biological ~~materials—~~materials – such as cells, chemicals, water, cytokines, vectors, nucleic acids, ~~antibodies—that~~antibodies, medium, serum, buffers —that are necessary to produce our product candidates. We source these raw and starting materials through ~~master~~ service agreements ~~and/or on a purchase order basis~~ and do not systematically have long-term supply contracts in place. However, we believe that

competitive pricing is achieved because there are a number of potential long-term replacements to each of our suppliers. Generally, the prices of the principal biological raw and starting materials that we purchase are stable or fluctuate within a limited range. To the extent that we are exposed to price fluctuations, we generally do not expect, in the near term, to be able to pass on cost increases because of the early development stage of our product candidates. However, with the SMART facility project, we expect to become independent for the supply of the most critical raw and starting materials.

Applications of Our Technology in Agriculture

Calyxt was incorporated in the State of Delaware in the United States in 2010. Calyxt is a consumer-centric, food- and agriculture-focused company that is combining its leading gene-editing technology and technical expertise with an innovative commercial strategy to pioneer a paradigm shift to deliver healthierhealthy food ~~ingredients for consumers and agriculturally advantageous traits for farmers.~~ ingredient company.

Before its initial public offering, which closed on July 25, 2017, Calyxt was a ~~wholly owned~~wholly-owned subsidiary of ours. As of December 31, ~~2017,~~2018, we owned approximately ~~79.7%~~69.5% of Calyxt’s outstanding common stock. Calyxt’s common stock is listed on the Nasdaq market under the ticker symbol “CLXT”.

~~Calyxt’s commercial strategy is centered on two core elements: developing healthier specialty food ingredients to enable~~Calyxt leverages proprietary intellectual property, technical expertise and anend-to-end supply chain toward its mission of “Making the ~~food industry to address evolving consumer trends and developing agriculturally advantageous traits, such as herbicide tolerance, for farmers. Its~~Food You Love a Healthier Choice^TM”. Calyxt’s first product ~~candidate, which is expected to be commercialized by the end of 2018,~~ is a ~~high oleic soybean~~High Oleic Soybean designed to produce a healthier oil that has increased heat stability with zero trans ~~fats.~~fats per serving. Calyxt received its first order for its High Oleic Soybean Oil in the first quarter of 2019. Among ~~Calyxt’s~~its other product candidates are other soybean products and a high fiber ~~wheat and herbicide tolerant~~ wheat. We believe each of these three Calyxt product candidates addresses a multibillion dollar market opportunity. Moreover, while the traits that enable these characteristics may occur naturally and randomly through evolution—or under a controlled environment through traditional agricultural technologies—those processes are imprecise and take many years, if not decades. Calyxt’s technology enables it to cost effectively edit a plant genome with precision and specificity in order to elicit the desired traits and characteristics, resulting in a final product that has no foreign DNA.

~~Gene Editing in Agricultural Biotechnology~~

While plant breeders have been crossbreeding varieties and selecting advantageous traits for thousands of years, the modern agriculture industry has relied primarily on two methods of crop improvement: genetic modification, which involves the use of genetic technologies to randomly insert foreign genetic material into a plant’s genome for the development of seeds in which the inserted genes express specific traits, and chemical mutagenesis, in which mutagenesis is induced in plants using agents and chemicals. We believe these traditional approaches can no longer effectively meet societal demands for innovative solutions demanded by the consumer and the farmer. We believe that the proprietary technologies deployed by Calyxt will bridge this divide because it enables Calyxt to precisely and specifically edit a plant genome to elicit a desired trait and characteristic and to do so more quickly and cost effectively than traditional methods.

~~Market Dynamics~~

The agriculture industry has historically been burdened by high infrastructure costs in a market that has focused on price and market share resulting in commoditization. A highly segmented supply chain has also resulted in the legacy agriculture companies focusing on increasing margins and market share through increased yields and consolidation, and on passing along maximum value to the growers, thereby keeping pace with the

growing demand for food globally. Over the past few decades the agriculture industry has seen a consolidation of over 200 seed companies, leaving the industry with only a handful of large, dominant players such as Bayer AG, Monsanto Co., DowDuPont Inc., AgReliant and Syngenta AG. In addition, development at these legacy agriculture companies has been significantly limited by time and cost constraints. Genetic modification, a primary method of these companies to improve crops, involves a lengthy and expensive process to progress a new crop from the discovery stage through commercialization. Innovations have primarily achieved increases in yields and food production volumes through the creation of herbicide tolerance and insect resistance, using genetically modified traits that in many cases contain bacterial DNA. We believe these industry dynamics explain the inability for the agricultural industry to evolve to a consumer- and farmer-focused approach, and thereby effectively meet their demands as societal trends shift and provide new market opportunities.

~~TALEN Technology Platform in Agricultural Biotechnology~~

Using Calyxt’s proprietary technologies and expertise, including TALEN gene-editing technology exclusively licensed to Calyxt ~~edits~~in the ~~genome~~field of ~~food crops by using “molecular scissors”~~agriculture, Calyxt precise cuts to ~~precisely cut~~ DNA in a single plant cell ~~use~~using the plant’s natural repair ~~machinery~~machinery. This allows Calyxt to make aour desired genome edit and ~~finally~~ regenerate the single cell into a full ~~plant. Calyxt~~plant that includes this gene edit.

Market and Industry Overview

Consumers have developed an increasingly heightened awareness of the role that dietary habits play in long-term wellness. This trend is ~~able~~especially prevalent in wealthier, developed nations where consumers have greater access to ~~develop targeted traits—some of which would be nearly impossible~~information that is helping to ~~develop using traditional trait-development methods—quicker, more efficiently~~shift their consumption habits. In the United States and ~~more cost effectively than would be possible using traditional trait-development methods. Calyxt’s technology also puts it in a position to assess the probability of success early on~~other developed nations buying habits have been creating dynamic shifts in the ~~research~~grocery aisle. Consumers now view food as a key to good health. More food products are being launched that go beyond basic nutrition to support health, digestive health, and ~~development process, potentially eliminating excess cost associated~~higher energy levels. Locally sourced foods with adirect-to-consumer model are becoming more attractive and the demand for transparency in food sourcing, production and labeling is gaining traction. Calyxt believes that as consumers continue the shift from a traditional ~~trait-development methods~~production-driven food culture to a modern demand-driven food culture, they will continue to demand more information and ~~further reducing~~accountability about how ingredients are sourced and processed, how “real” their food products are, and how responsive they are to consumers’ desire for choice and customization.

Regulatory agencies are also playing a larger role in monitoring which food ingredients reach consumers. Beginning in 2018, the ~~risk~~FDA banned certain uses of partially hydrogenated oils, the primary artificial source of trans fat in processed foods. Following the passage of the ~~product development process.~~Healthy, Hunger-Free Kids Act of 2010, the USDA gained significant oversight of the federal school lunch program and holds the authority to set new, healthier standards for food sold in U.S. schools. These healthier food mandates include minimum serving requirements for fiber, fruits and vegetables and maximum allowable content standards for fat, sugar and sodium. Consumers’ rising demand for healthier food presents an opportunity for us to provide our innovative solutions for customers and consumers and the food industry.

TALEN Technology in Agricultural Biotechnology

Calyxt’s ~~proprietary technologies and intellectual property portfolio enable it~~gene-editing platform relies on Calyxt’s capacity to ~~edit DNA in living plant cells by knocking out genes or creating precise gene edits. Calyxt’s scientists are capable of~~ custom ~~designing~~ designDNA-sequence specific cutting enzymes, or nucleases, for any chosen gene ~~that they need~~it intends to edit and its capability to introduce such custom-made nucleases into the living plant cells ~~they want~~it wants to edit. Calyxt’s platform also relies on precisely chosen protein families that can specifically recognize unique DNA sequences and can be tailored to target such sequences in any chosen gene or genetic region.

Calyxt’s proprietary technologies and intellectual property portfolio enable Calyxt to edit the plant genome by knocking out genes or making precise gene edits. Calyxt takes advantage of ~~deep~~its knowledge about plant gene function to ~~create~~produce novel genetic variation that results in traits of value.

A ~~feature that distinguishes~~key difference between Calyxt’s gene-edited products ~~from those~~and products created through genetic modification (GMO) is that GMOs insert foreign DNA into crops and Calyxt’s ~~crop varieties lack foreign DNA. As such, for~~gene-editing does not. For each of ~~the six~~Calyxt’s product candidates ~~Calyxt has~~that it submitted to ~~date,~~the U.S. Department of Agriculture (USDA), the USDA ~~has~~ confirmed that the ~~products~~product candidates are not regulated ~~articles, which represents regulatory cost savings for the development of these products.~~articles. This determination decreased Calyxt’s costs and increased speed to market.

~~In addition, by utilizing~~Using Calyxt’s proprietary technologies and expertise, including TALEN gene-editing ~~technologies~~technology exclusively licensed to Calyxt in the field of agriculture, Calyxt develops food crops with targeted traits quickly and ~~leveraging Calyxt’s expertise and innovative supply chain, we~~more cost effectively than traditional methods. We believe that Calyxt is able to identify a consumer ~~or farmer~~ need and develop a

product from “concept to fork” ~~or “concept to field”~~ in ~~approximately three to~~cycles as short as six years ~~which would make it possible to effectively respond to evolving consumer preferences and farmer needs.~~by utilizing these proprietary technologies.

Calyxt Agricultural Biotechnology Products

~~Calyxt has an extensive~~Calyxt’s product pipeline ~~as set forth~~includes a variety of traits for soybeans, wheat, and alfalfa, and Calyxt intends to conduct further development programs to build upon our current pipeline. In the future, Calyxt may expand our product pipeline to include other food crops. Calyxt’s initial focus is on its High Oleic Soybean and High Fiber Wheat product candidates. Calyxt received its first order for its High Oleic Soybean product candidate in the ~~table below, that is intended to address the potential market opportunities Calyxt has identified to date.~~

~~Calyxt categorizes the stages~~first quarter of~~pre-commercial~~ development from Phase I to Phase III. Prior to entering Phase I, in Discovery, Calyxt identifies genes of interest. In Phase I, Calyxt edits the identified genes of interest, targets edits that it wishes to make, and produces an initial seed that contains the desired edit. Phase II is trait validation, where Calyxt performs small-scale and large-scale tests to confirm phenotype and ingredient functionality. In this phase Calyxt also performs replicated, multi-location field testing, after confirming that the product is not a regulated article by the USDA. In Phase III, Calyxt develops the first commercial-scale pilot production, begins to build out the supply chain and inventory and performs customer testing prior to commercialization.

~~Calyxt’s current main product candidates are:~~ 2019.

High Oleic Soybean ~~(Consumer Trait)~~

Soybean oil has historically been partially hydrogenated to enhance its oxidative stability in order to increase shelf life and improve frying characteristics. This process, however, creates trans-unsaturated fatty acids, or trans fats, which have been demonstrated to raiselow-density lipoprotein (LDL) cholesterol levels and

lower high-density lipoprotein (HDL)~~, both of which contribute~~ cholesterol levels. High LDL and low HDL have been tied to increased risk for cardiovascular disease. The discovery that dietary trans fats increase the risk of several adverse health issues led the FDA to rule in 2003 that manufacturers be required to include ~~trans fat~~trans-fat content information on the “Nutrition Facts” label of foods. In 2015, the FDA took a further step and banned the use of partially hydrogenated oils, the primary dietary source of artificial ~~trans fat~~trans-fat in processed foods, by all food manufacturers beginning in 2018. ~~After the FDA’s 2003 ruling, commodity soybean oil—which leads to high trans fats in foods—lost approximately 20% market share to other vegetable oils, such as palm oil and canola oil.~~

Monounsaturated fats, such as oleic acid, have been linked to reducing LDL cholesterol and triglycerides and raising HDL cholesterols. Diets rich in monounsaturated acids are associated with lower fat mass and decreased blood pressure. High levels of oleic acids can be found in olive, canola, sunflower and safflower oils.

Calyxt developed a soybean trait that has produced oils with a fatty acid profile that contains 80% oleic acid, 20% less saturated fatty acids compared to commodity soybean oil and zero ~~transfats.~~ grams of trans fats per serving.

Oil ~~created~~produced from Calyxt’s ~~high oleic soybean product candidate~~High Oleic Soybean has multiple desirable characteristics as an ingredient for the food industry. The high level of oleic acid in Calyxt’s soybean oil enhances oxidative stability more than fivefold when compared to commodity oil. This eliminates the need for partial hydrogenation, and thus little to no trans fats are produced during oil production. Furthermore, Calyxt’s ~~high oleic soybean oil~~High Oleic Soybean Oil offers additional potential benefits, including reduced saturated fats, a threefold increase infry-life, and reduced polymerization upon frying at high temperatures. Soybean oil is also neutral in flavor, odorless and colorless, and is therefore ~~highly~~ desired as a food ingredient because it has limited impact on the sensory characteristics of the final food product.

Calyxt’s ~~high oleic soybean product candidate~~High Oleic Soybean was ~~created~~developed using ~~our~~ TALEN gene-editing technology. WeCalyxt designed a TALEN to specifically target two fatty acid desaturase genes (designatedFAD2-1Aand~~FAD2-1B~~)FAD2-1B). These genes convert oleic acid (a mono-unsaturated fatty acid) to ~~linoleic~~linolenic acid (a polyunsaturated fat). By specifically inactivating both theFAD2-1A andFAD2-1B genes, ~~by removing DNA with TALEN,~~ oleic acid accumulates in the seed—increasing from about 20% to 80%. By key measures, including yield, ~~this high oleic soybean~~Calyxt’s High Oleic Soybean variety performs comparably to its unedited conventional(non-GMO)counterpart. Further, this improved soybean variety does not contain any foreign DNA. Because the TALEN technology is so precise and can target genes with well-known functions in the plant, Calyxt has not, to date, detected any other changes as a result of the gene-editing process or undesired effects in the product.

Inmid-2015, Calyxt received a letter from the USDA indicating that ~~its high oleic soybean variety~~Calyxt’s High Oleic Soybean Oil is not a regulated article under the Plant Protection Act. ~~This allowed Calyxt to test the performance of its soybean variety in the field.~~ In November 2015, Calyxt announced the completion of the second year of multi-location field trials in Minnesota and South Dakota. The agronomic and yield performance of ~~its high oleic soybeans~~Calyxt’s High Oleic Soybean is on par with thenon-GMO variety used to ~~create~~develop this product.

~~Calyxt’s soybean product candidate~~Soybeans are a crop that is ~~in Phase III~~photoperiod sensitive, meaning the crop is sensitive to the length of the ~~development process,~~day. Plant breeders have developed soybean varieties that are adapted to distinct latitudes and agroclimatic conditions—which are referred to as maturity groups. Different growing regions in the United States require different maturity groups. For example, the northern United States requires soybeans in the1.5- 2.2 maturity group ranges, with farmers typically planting a few varieties within this range to diversify their risk.

Calyxt hasin-licensed over thirtynon-genetically modified soybean varieties with commercial rights. Calyxt initially introduced Calyxt’s High Oleic Soybean in the northern United States on one variety within a particular maturity group range. Over time, Calyxt intends to expand to additional varieties for farmers in the same region to provide farmers with the ability to diversify their risk. Calyxt expects to continue this expansion over time by adding regions in other maturity groups and varieties. Over the next five to ten years Calyxt envisions expanding its germplasm portfolio to early, middle and late maturity groups. Calyxt believes this strategy will enable it to expand its supply chain through additional crushing plants within target growing regions and at the same time lower potential premiums and production costs as it introduces into the supply chain additional varieties that meet the diversity of seed variety that farmers need to mitigate their risk.

Calyxt plans to focus its efforts on farmers in the upper Midwest. Calyxt currently has growers in South Dakota and Minnesota and will be launching new soybean varieties in 2020 and 2021 that will enable it to expand to other growing locations. The High Oleic Soybean seed is sold either directly or through distributors to farmers. Calyxt believes that it can generate a successful supply chain by contracting with fewer than 2,000 farmers.

In 2018 Calyxt contracted more than 17,000 acres with nearly 80 growers in South Dakota and Minnesota to grow this High Oleic Soybean. The harvest of these High Oleic Soybean acres has been completed and Calyxt ~~is making preparations~~has taken delivery of a portion of the harvested crop. Calyxt has contracted with third parties to ~~create the capacity to~~ crush and refine Calyxt’s High Oleic Soybean on a commercial scale, ~~which will enable~~enabling sales of High Oleic Soybean Oil and High Oleic Soybean Meal to customers. Calyxt has commercialized its High Oleic Soybean product ~~testing by potential food company customers. During~~candidate in the ~~third~~first quarter of 2017, Calyxt completed the harvest of its high oleic soybean and produced more high-quality seeds than needed to meet its commercialization target date. Calyxt is currently completing its commercialization plan and anticipates commercialization by the end of 2018.2019.

In December 2017, Calyxt announced its partnership with Farmers Business Network (FBN) to expand the distribution and grower base of its identity-preserved high oleic soybeans. FBN will distribute its high oleic soybean seeds to growers in its network and register farmers for our Producer Premium Program. This partnership will help develop a dedicated, high-quality grower base and bolster supply chain operations in the upper Midwest.

~~In addition, Calyxt’s high oleic soybean product candidate can be stacked with other soybean traits.~~

High Fiber Wheat ~~(Consumer Trait)~~

Fiber is the indigestible portion of food that is essential for healthy digestion. Research has shown that ~~fiber—the indigestible portion of food—~~fiber may play a large role in maintaining bowel health, lowering cholesterol, stabilizing blood glucose levels and controlling weight gain. A high fiber diet has the potential to lower the rate of glucose entry into circulation, thus decreasing the risk of food-related chronic diseases, such as coronary artery disease and diabetes. The average American adult consumes approximately15-18 grams of fiber daily, only half of the amount recommended by the U.S. Department of Health’s dietary guidelines based on the average caloric intake. In recent years, the awareness of the health benefits of high fiber diets has ~~increased, which~~increased. This has translated to a strong growth in demand for high fiber food ~~products.~~products, with approximately 35% of grocery shoppers now seeking high fiber foods.

Calyxt is developing ~~high fiber wheat traits~~its High Fiber Wheat product candidate that could be used to produce white flour with up to three times more dietary fiber than standard white ~~flour. Calyxt anticipates that by~~flour while maintaining the same flavor and convenience of use. By altering the proportion of certain slower digested carbohydrates in the wheat grain, Calyxt has developed a product candidate that it believes will ~~increase~~exhibit increased dietary fiber. This would allow consumers to reach their daily value of fiber without changing their existing food preferences. ~~These~~

Calyxt believes its High Fiber Wheat flour will be able to be incorporated into many food products—from pasta to bread. Whereas a single serving of whole wheat flour can provide 49% of an individual’s daily fiber needs, a single serving of Calyxt’s High Fiber Wheat flour may provide up to 100% of the recommended daily requirement thereby allowing food manufacturers to make high fiber ~~wheat product candidates will not contain any foreign DNA.~~products sought after by many consumers.

This product candidate is currently in Phase III of Calyxt’s development process and ~~may launch~~is expected to be commercialized as early as ~~2020 – 2021. The gene-edited wheat line has been identified~~2022. In March 2018 Calyxt received confirmation from USDA that its High Fiber Wheat is deemednon-regulated pursuant to APHIS’s regulatory procedures. In October 2018 Calyxt successfully harvested High Fiber Wheat in field trials. In the next year, Calyxt intends to further confirm the product concept in field conditions and ~~grown and the fiber levels are under examination in grain derived from greenhouse grown plants.~~to complete food application studies.

In addition to Calyxt’s ~~high fiber wheat~~High Fiber Wheat product candidate, ~~Calyxt~~the company is also developing other consumer traits in ~~its~~our wheat pipeline, including a reduced gluten product candidate.

~~Herbicide Tolerant Wheat (Farmer Trait)~~

Weed control is one of the greatest challenges farmers face in producing crops. Herbicide tolerance is a plant’s ability to withstand a particular chemical herbicide. Herbicide tolerance traits in crops can provide additional crop protection chemistry alternatives to control weeds and increase crop yields.

Herbicide tolerant traits may offer farmers a vital tool in managing weeds effectively during crop production. The deployment of herbicide tolerant traits in wheat significantly lags other major crops and wheat production is constantly faced with yield-robbing weeds that can result in lower yield and higher dockage costs at the elevator. In the United States, no broad acre GMO herbicide tolerant trait has been developed for wheat, largely due to the complexity of the wheat genome.

Calyxt is pioneering the development of herbicide tolerant traits in wheat without the use of foreign DNA, built using the TALEN gene-editing technology. Calyxt aims to achieve herbicide tolerance by specifically making a subtle repair to prevent herbicides from being able to recognize and block functions of certain plant-encoded proteins, such that the edited plant survives the application of the herbicide. In September 2017, Calyxt successfully advanced its herbicide tolerant wheat program from the Discovery phase into Phase I development.

~~Powdery Mildew Resistant Wheat (Farmer Trait)~~

In addition to herbicide tolerance, Calyxt is developing traits that are advantageous to the farmer including a variety of wheat that confers resistance to a fungal pathogen, namely powdery mildew. Powdery mildew resistant wheat advanced to Phase II in 2017 after successfully conducting the first field trial to assess agronomic and trait performance of the powdery mildew resistance trait line. We believe powdery mildew resistance may increase yield and reduce the need for the use of costly fungicides.

~~Improved Oil Composition Canola (Consumer Trait)~~

Calyxt’s improved oil composition canola is its first canola product candidate. The development of this first canola product expands Calyxt’s improved oils franchise, in line with its mission to create healthier specialty ingredients and become a preferred partner of the food industry. In September 2017, Calyxt successfully advanced its improved oil composition canola program from Discovery stage into Phase I development.

~~Other Products in Our Development Pipeline~~

Calyxt’s extensive product pipeline includes a variety of consumer- and farmer-centric traits for soybean, wheat, canola, alfalfa and potato. As of December 31, 2017, Calyxt had a total of nine product candidates in Phase I or higher across its five crops, which is reflective of its rapidly advancing product pipeline.

Intellectual Property

We seek to protect and enhance proprietary technology, inventions, and improvements that are commercially important to the development of our business by seeking, maintaining, and defending patent rights, whether developed internally or licensed from third parties. We will also seek to rely on regulatory protection afforded through orphan drug designations, data exclusivity, market exclusivity and patent term extensions where available.

To achieve this objective, we maintain a strategic focus on identifying and licensing key patents that provide protection and serve as an optimal platform to enhance our intellectual property and technology base.

Historical Perspectives

Cellectis was founded in early 2000. In June 2000, Institut Pasteur provided us with exclusive rights to its gene-editing patent portfolio. This patent portfolio includes patents relating to homologous recombination and rare-cutting endonucleases (also named meganucleases), respectively, for genetic engineering in living cells.

Since 2002, we have filed a large number of patent applications, many now issued as patents, for custom-made meganucleases, and uses thereof, that specifically target a desired genetic sequence in a genome. In 2014, we entered into a cross-licensing agreement with Precision Biosciences, Inc., or Precision, in settlement of patent litigation and patent proceedings related to this technology. Pursuant to this cross-license, we licensed our patents and patent applications in this area to Precision, and Precision licensed its relevant patents and patent applications to us.

In 2010, we acquired a portfolio of patents and patent applications relating to electroporation methods and devices. In 2011, we entered into an exclusive license agreement with the Regents of the University of Minnesota (UMN) pursuant to which wein-licensed one patent family related to ~~a new generation of~~ customized rare-cutting endonucleases, in connection with which we have registered the trademark TALEN in certain ~~jurisdictions outside of the United States.~~jurisdictions. This patent portfolio comprises ~~five~~six patents in the United States and ~~one~~two European ~~patent, which is under opposition before the European Patent Office.~~patents,. In addition, in 2014, we entered into a series of agreements with Life Technologies Corporation ~~(now~~(controlled by Thermo Fisher Scientific Inc.) pursuant to which we received anon-exclusive sublicense under certain patents and patent applications related to the research and therapeutic uses of ~~TALEN~~TALE-nucleases and we granted certain rights to Life Technologies under our TALEN technology. In addition, we entered into a license agreement with Calyxt, pursuant to which Calyxt has been granted certain rights in connection with our gene editing and plant intellectual property portfolio.

Since 2012, we have filed about 5060 new patent applications related to the CART-cell technology. Included in this patent portfolio are patent applications relating to manufacturing allogeneic immune cells and to CAR design, including multi-subunit CARs and conditional expression CARs. In addition, we have filed a number of patent applications related to new TALEN structures (for example, compact TALEN, methylation TALEN) and alternatives to the TALEN structure (BurrH, CRISPR-Cas9). On December 2018, an anonymous third-party filed an opposition to two European patents of the patent family entitled “A Method for producing precise DNA cleavage using CAS9 – Double nickase”.

In October 2014 and March 2014, we exclusivelyin-licensed two patent portfolios from Ohio State Innovation Foundation and University College London, respectively. The Ohio State Innovation Foundation patent portfolio includes ~~an international~~ patent ~~application~~applications relating to CARs directed to cancer marker CS1. The University College London patent portfolio includes patent applications relating to a polypeptide expressing the suicide gene RQR8, and uses thereof.

Current Intellectual Property Portfolio

As a result of the licensing opportunities described below and our continuing research and development efforts, our intellectual property estate now contains patent applications that cover our products, including claims that cover:

methods central to genome engineering and gene editing, including methods of homologous recombination, nuclease-based gene targeting, replacement, insertions and/orknock-out;

the main products we use in the manufacturing process, including nucleases;

manufacturing steps, including cell electroporation, transformation and genetic modifications;

engineered cells;

single-chain and multi-subunit CARs expressed at the surface ofT-cells;

specific gene inactivation and suicide gene expression;

allogeneic and autologous treatment strategies using ourT-cell products; and

plant traits and methods for gene editing plant cells.

The most relevant issued patents in our portfolio consist of approximately 2733 Cellectis-owned and 4718in-licensed U.S. patents, 1632 Cellectis-owned and 134in-licensed European patents, and 47 Cellectis-owned and 13in-licensed patents in other jurisdictions, ~~including~~such as Australia, Canada, China, Hong Kong, India, Israel, Japan, Korea, Mexico and Singapore.

The most relevant pending patent applications in our portfolio consist of approximately ~~115~~88 Cellectis-owned and 204in-licensed U.S. patent applications, 8068 Cellectis-owned and 192in-licensed European patent applications, ~~and 436~~412 Cellectis-owned and 6728in-licensed patent applications pending in other jurisdictions, ~~including~~such as Australia, Brazil, Canada, China, Hong Kong, India, Israel, Japan, Korea, Mexico and Singapore.

Our most relevant portfolio includes a total of ~~165~~146 owned andin-licensed granted patents, and ~~737~~596 owned andin-licensed patent applications.

Our UCART product candidates rely for each product candidate upon one or more patent rights protecting various aspects of the technologies, including rights relating to:

the genetic editing ofT-cells, using TALEN technology, covered by approximately twelve Cellectis-owned patent families and threein-licensed patent families;

the insertion of transgenes intoT-cells using electroporation of mRNA, covered by approximately five Cellectis-owned patent families;

the appending of attributes toT-cells, covered by approximately eight Cellectis-owned patent families and onein-licensed patent family;

the molecular structure of CARs, covered by approximately six Cellectis-owned patent families; and

specific CARs that target selected antigen markers are covered by approximately fifteen Cellectis-owned patent applications and onein-licensed patent family.

For additional information, see “—Gene-Editing Platform” below.

Similarly, our most advanced agricultural product candidates each rely upon one or more patent rights relating to:

the genetic editing of plants using TALEN technology, covered by approximately six Cellectis-owned patent families and twoin-licensed patent families;

the genetic editing of plants using meganuclease technology, covered by approximately eight Cellectis-owned patent families and onein-licensed patent family;

the genetic editing of plants using CRISPR-Cas9 technology, covered by approximately two Cellectis-owned patent families and threein-licensed patent families; and

specific plant traits, which are covered by approximately twelve Cellectis-owned patent families.

Individual patent terms extend for varying periods of time, depending upon the date of filing of the patent application, the date of patent issuance, and the legal term of patents in the countries in which they are obtained. In most countries in which we file patent applications, including the United States, the patent term is 20 years from the date of filing of the firstnon-provisional application to which priority is claimed. In certain instances, a patent term can be extended under certain circumstances. For example, in the United States, the term of a patent that covers anFDA-approved drug may be eligible for a patent term restoration of up to five years to effectively compensate for the patent term lost during the FDA regulatory review process, subject to several limitations discussed below under “—Our Intellectual Property Strategy.” Also, in the United States, a patent’s term may be lengthened by patent term adjustment, which compensates a patentee for administrative delays by the U.S. Patent and Trademark Office in granting a patent, or may be shortened if a patent is terminally disclaimed over an earlier-filed patent. Our issued patents will expire on dates ranging from 2019 to 2035. If patents are issued on our pending patent applications, the resulting patents are projected to expire on dates ranging from 2023 to 2035. However, the actual protection afforded by a patent varies on aproduct-by-product basis, fromcountry-to-country, and depends upon many factors, including the type of patent, the scope of its coverage, the availability of regulatory-related extensions, the availability of legal remedies in a particular country, and the validity and enforceability of the patent.

The patent portfolio for our most advanced product candidates, UCART19 and UCART123, are summarized below.

Gene Editing Platform

Each of our UCART product candidates relies upon our gene-editing platform andT-cell and CAR technology platforms. The patent portfolio covering these platforms and technologies, includes approximately 30 issued patents or pending patent ~~applications.~~applications in various countries. These issued patents and pending patent applications, which expire between 2019 and 2033, cover product claims or process claims relevant to each of our product candidates, including UCART19 and UCART123.

Our gene-editing platform and each of our UCART product candidates benefits from the protections of several patents and patent applications in our patent portfolio. As a result of this broad range of patent protection, very few individual patents in our portfolio are critical to our ability to effectively conduct our product development activities. Although certain patents relating to our electroporation technology will expire in 2019, other patents covering this technology remain in force or are expected to be granted under patent applications, and additional patents protect the nucleases delivered by our electroporation technology, as well as the methods to modify the cells by use of such nucleases. As a result of the breadth of our patent protection and the integration of patented technologies, compositions and methods of use within our gene-editing,T-cell and CAR technology platforms, we do not expect that the expiration of these patents in 2019, individually or in the aggregate, will have a material effect on our future operations or financial position.

UCART19

In addition to the patent portfolio relating to our platform and technologies, described above, our patent portfolio relating specifically to UCART19 includes pending patent applications from the patent family WO2014184143 (CD19 Specific Chimeric Antigen Receptor and Uses Thereof). We believe these pending patent applications, which, if issued, would expire in 2034, include claims to cover the composition of matter of UCART19, methods of manufacture of UCART19, and methods to use UCART19 in treatment.

UCART123

In addition to the patent portfolio relating to our platform and technologies, described above, our patent portfolio relating specifically to UCART123 includes pending patent applications from the patent family WO2015140268 (CD123 Specific Chimeric Antigen Receptors for Cancer Immunotherapy). We believe these pending patent applications, which, if issued, would expire in 2034, include claims to cover the composition of matter of UCART123, methods of manufacture of UCART123, and methods to use UCART123 in treatment.

In each case, some of the issued patents and pending patent applications, if issued, may be eligible for patent term extension and patent term adjustment, thereby extending their terms, as described above.

UCARTCS1

Our patent portfolio relating specifically to UCARTCS1 includes pending patent applications from the patent family WO2014179759(CS1-SPECIFIC CHIMERIC ANTIGEN RECEPTOR ENGINEERED IMMUNE EFFECTOR CELLS1) licensed exclusively from the Ohio State University, which, if issued, would expire in 2034. This patent family is directed to composition of matters including a CARanti-CS1 per se. Our patent portfolio also includes patent applications filed by Cellectis from the family WO2015166056 (CS1 SPECIFIC MULTI CHAIN CHIMERIC ANTIGEN RECEPTOR) and WO2015121454(T-CELLS FOR IMMUNOTHERAPY ENGINEERED FOR TARGETING ANTIGEN PRESENT BOTH ONT-CELLS AND PATHOLOGICAL CELLS), which, if issued, would expire on 2035. Both families relate to the use of CAR anti CS1 in allogeneicT-cells, methods of manufacture of UCARTCS1, and methods to use UCARTCS1 in cell therapy treatment.

Material Exclusive Licenses Granted to Cellectis

Licenses from Institut Pasteur

In 2000, we entered into ~~three~~series of license agreements with L’Institut Pasteur, or Pasteur, pursuant to which wein-licensed a substantial portion of Pasteur’s gene-editing patent portfolio. The details of ~~the two~~these license agreements ~~that remain in effect~~which are still effective are provided below.

In June 2000, we entered into an agreement with Pasteur, later amended in 2003 (collectively, the “Second June 2000 Agreement”), acting on its own behalf and on the Boston Children Hospital’s behalf, pursuant to which it granted to us an exclusive, worldwide, royalty-bearing, sublicenseable license under certain patents andknow-how owned by Pasteur and the Boston Children’s Hospital relating to certain chimeric endonucleases for chromosomal gene editing by homologous recombination in cells to use, manufacture, and sell products and to practice processes covered by such patents. The license granted under the Second June 2000 Agreement isnon-exclusive, however, with respect to the licensed processes applied to human gene therapy. In the event that Pasteur has the possibility to grant exploitation rights for applications to human gene therapy, it must immediately inform us, and we may amend our agreement with Pasteur to obtain such exploitation rights. ~~One of the licensed patents is under third partyex-parte~~ ~~reexamination.~~

The exclusivity of each of the licenses granted under the Second June 2000 Agreement is further contingent upon our continued diligence in designing, developing, and obtaining the required regulatory authorizations necessary to sell the respective licensed products and processes.

In October 2000, we entered into an agreement with Pasteur, later amended in 2003, 2004, 2005, and 2007 (collectively, the “October Agreement”), pursuant to which we obtained an exclusive, worldwide, royalty-bearing, sublicenseable license under certain patents andknow-how owned by Pasteur relating to a method of homologous recombination to make, use, and sell products and to implement processes covered by such patents. The exclusivity of the license granted under the October Agreement is subject to a license granted to a third party under the licensed patents in the domain of genes that encode for Erythropoietin.

We may only grantsub-licenses under our Pasteur agreements with Pasteur’s prior approval, which is deemed to have been given if Pasteur does not object to a proposedsub-license within a specified period of time from notice of the proposed sublicense and which may only be withheld for serious cause.

Pursuant to the terms of each of the Second June 2000 Agreement and the October Agreement, we made cash payments to Pasteur in an aggregate amount of 600,000 French Francs with respect to the entry into the agreement and the reimbursement of license fees. Pursuant to the terms of the October Agreement, we made cash payments to Pasteur in the aggregate amount of 500,000 French Francs with respect to the entry into the agreement and the reimbursement of license fees and 250,000 Euros in connection with the execution of amendments. Under the Second June 2000 Agreement, we are also required to pay Pasteur an ongoing royalty fee equal to alow- tomid-single digit percentage of our net income with respect to licensed products under the respective agreement. With respect to sublicenses granted under the Second June 2000 Agreement, we are also required to pay Pasteur a percentage of

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all payments received under such sublicenses, subject in certain cases to minimum payment amounts based on net revenues of the applicable sublicensee. Under the October Agreement, we are also required to pay Pasteur an ongoing royalty fee equal to alow-single digit percentage of our net income with respect to licensed products under the October Agreement. With respect to sublicenses granted under the October Agreement, we are required to pay Pasteur a tiered percentage of all compensation received by us during the applicable year under the sublicense agreement, subject in certain cases to minimum payment amounts based on net revenues of the applicable sublicensee.

The terms of each of our agreements with Pasteur will expire upon the expiration of thelast-to-expire of the respective patents licensed to us pursuant to the applicable agreement. We expect the last to expire patent under the Second June 2000 Agreement to expire on February 3, 2020 and the last to expire patent under the October Agreement to expire on ~~April 3,~~March 4, 2020. Pasteur and we may each terminate any of our agreements with Pasteur in the event of the other party’s breach of an obligation under the applicable agreement, which remains uncured for 90 days following receipt of notice of such breach from the terminating party. Pasteur may immediately terminate such agreements if we challenge or contest the validity of any of the licensed patents under the respective agreement before a court or patents office. In addition, Pasteur and we may terminate any of the agreements, upon 60 days’ prior notice, in connection with certain insolvency-related judicial proceedings instituted against the other party. Further, we have the right to terminate any of these agreements for any reason immediately upon notice to Pasteur.

License from Regents of the University of Minnesota

In January 2011, we entered into an exclusive license agreement with Regents of the University of Minnesota, or UMN. Pursuant to this agreement, as amended in 2012, 2014, and 2015 we and our affiliates were granted an exclusive, worldwide, royalty-bearing, sublicenseable license, under certain patents and patent applications owned by UMN, to make, use, sell, import, and otherwise dispose of products covered by the licensed patents, for all fields of use. These licensed patents relate to TALEN molecules and their use in gene editing. Pursuant to the agreement, we are required to achieve certain specified research- and sales-related milestones.

Pursuant to the terms of the agreement, we paid UMN an upfront license fee in the amount of $250,000 upon the effective date of the license agreement, and a second upfront payment in the amount of $1,000,000 following execution of the third amendment. In thenon-agricultural field we are also required to pay to UMN low single digit percentage royalties on net sales of licensed products, as well as a percentage of all revenues received by us under sublicenses. Pursuant to the agreement, UMN is entitled to a minimum annual ~~royalties~~royalty of $30,000 per year. In the agricultural field, no royalties are due on net sales of licensed products, but an annual fee of $150,000 per year is due to UMN and commercial milestones are due upon the occurrence of certain commercial sale milestones. We are also required to pay UMN milestone payments up to a total of $290,000 in the aggregate upon the occurrence of specified events and to pay certain patent-related expenses incurred under the agreement for prosecuting and maintaining the licensed patents. If we undergo a change of control and wish to assign our rights and duties under the agreement, we will be required to pay UMN an additional transfer fee.

The license agreement will expire upon the expiration of the last to expire valid claim of the licensed patents. UMN may terminate the agreement upon advance written notice in the event of our insolvency or bankruptcy, and immediately upon written notice in the event that we challenge the validity or enforceability of any licensed patent in a court or other applicable authority. UMN and we may terminate the agreement by written notice in the event of the other party’s breach that has not been cured within a specified number of days after receiving notice of such breach.

License from Ohio State Innovation Foundation

In October 2014, we entered into an exclusive license agreement with Ohio State Innovation Foundation. Pursuant to this agreement, we were granted an exclusive, worldwide, royalty-bearing, sublicenseable license

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under certain patents and patent applications owned by Ohio State Innovation Foundation to use, make, distribute, sell, lease, loan or import products or process covered by the licensed patents, for any and all activities relating to cancer immunotherapy. The licensed portfolio includes an international patent application relating to CAR directed to cancer marker CS1. Pursuant to the agreement, we must use diligence and commercially reasonable efforts to commercialize licensed products or processes, including achieving certain milestone events by specified deadlines, subject to our ability to extend such deadlines upon payment of certain fees.

Pursuant to the terms of the agreement, we paid Ohio State Innovation Foundation an upfront license fee in the amount of $100,000. We are required to pay an annual license maintenance fee of $20,000 from 2015 onward until our first sale of a licensed product. We are also required to pay to Ohio State Innovation Foundation low single-digit percentage royalties on net sales of licensed products and licensed processes by us and are subject to minimum annual royalties due to Ohio State Innovation Foundation of $100,000. We are also required to pay Ohio State Innovation Foundation a percentage of royalties paid to us by sublicensees. We are also required to pay Ohio State Innovation Foundation milestone payments up to a total of $1,950,000 in the aggregate upon the occurrence of certain development-related events prior to deadlines specified in the agreement.

Unless earlier terminated, the license agreement will expire upon the expiration of the last to expire valid claim of the licensed patents, which we expect will be on May 2, 2034. We may terminate the agreement at our option by giving 90 days’ written notice. Ohio State Innovation Foundation may immediately terminate the agreement, any part of the licensed patent rights or the agreement’s exclusivity if we fail to make required payments under the agreement and such breach continues for sixty days after delivery of written notice from Ohio State Innovation Foundation or if we breach any other provision of the agreement and fail to cure such breach within 60 days after delivery of written notice from Ohio State Innovation Foundation. Ohio State Innovation Foundation may also terminate the agreement if we or our affiliate initiates any proceeding or action challenging the validity, enforceability or scope of any of the patent rights or assists a third party in such a proceeding or action. The agreement automatically terminates if we file for bankruptcy or become bankrupt or insolvent, our board of directors elects to liquidate our assets or dissolve our business, we cease business operations, we make an assignment for the benefit of creditors or if we are otherwise placed in the hands of a receiver, assignee or trustee, whether by our voluntary act or otherwise.

~~Other Exclusive Licenses Granted to Cellectis~~

With respect to our plant sciences business, we have filed nine patent applications in connection with new soybean, tobacco and potato traits. These new traits have been obtained mainly by using TALEN, which is protected by the UMN and the Cellectis TALEN patent portfolios.

~~License Agreements from Regents of the University of Minnesota~~

In December 2014, Calyxt entered into an exclusive license with the Regents of the University of Minnesota, or UMN, pursuant to which Calyxt was granted an exclusive, worldwide, sublicensable license under a specified patent application owned by UMN relating to the use of the CRISPR-Cas9 technology to make use, and commercialize products covered by the licensed patents in any field of use. Pursuant to the terms of the agreement, Calyxt must use commercially reasonable efforts to commercialize the licensed technology and to manufacture, offer to sell, and sell licensed products as soon as practicable and to maximize sales. Calyxt must also achieve certain sales- and patent-related milestones.

Pursuant to the terms of the agreement, Calyxt paid UMN an upfront license fee payment in the amount of $130,000 in connection with entering into the agreement. Calyxt is also required to pay UMN a tiered annual fee that increases from $20,000 to $225,000 based on the occurrence of certain specified events, including the grant of a sublicense to a third party, as well as patent-related expenses incurred under the agreement in prosecuting and maintaining the licensed patents. Calyxt is also required to pay UMN a percentage of all revenues received by it under sublicenses. If Calyxt undergoes a change of control and wishes to assign all of its rights and duties under the agreement, Calyxt will have to pay UMN a specified transfer fee.

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Unless earlier terminated, the agreement will continue in effect until no licensed patent is active and until no licensed patent application is pending, which we expect will be on March 24, 2034. UMN may terminate the agreement for Calyxt’s uncured breach of the agreement upon 90 days’ prior written notice, or 60 days’ prior written notice if the breach relates to Calyxt’s payment obligations under the agreement. UMN may also terminate the agreement, upon 10 days’ prior written notice, if Calyxt files for bankruptcy or becomes insolvent. UMN may also immediately terminate the agreement if Calyxt or its agents or representatives commences or maintains an action in any court or before any governmental agency asserting or alleging the invalidity or unenforceability of the licensed patent rights. Calyxt may terminate the agreement for UMN’s uncured breach of the agreement upon 90 days’ prior written notice. Calyxt may also terminate the agreement at any time upon 60 days’ prior written notice.

~~Other Exclusive License Granted to Calyxt~~

In April 2015, we entered into an exclusive license agreement with Plant Bioscience Limited, or PBL, pursuant to which we were granted an exclusive, worldwide license to use and exploit certain gene-editing technologies related to wheat with endogenous resistance to powdery mildew, and to provide commercial development and technical services to third parties. The technology licensed to us was originally exclusively licensed to PBL by the Institute of Genetics and Developmental Biology, or IGDB, which holds certain rights to which our exclusive license from PBL is subject. Pursuant to the agreement, we are required to use commercially reasonable efforts to achieve certain milestones in a specified development and commercial program, and to exploit the license so as to maximize the sublicensing of the licensed technology and the development and commercial use of the licensed products.

The agreement will expire on the later of the tenth anniversary of the date that sales of the licensed products first occurred or the last expiration of a valid claim of a licensed patent. We may terminate the agreement at any time for any reason by giving PBL written notice. PBL may terminate the agreement upon our material breach of the agreement that has not been cured within a specified number of days after receiving notice of such breach. The agreement will also automatically terminate if we become insolvent or bankrupt.

In December 2017, we entered into an Evaluation License and Option Agreement with PBL to evaluate gene-editing technologies related to wheat with altered grain size and structure. The licensed technology was originally licensed to PBL by IGDB, which holds certain rights to which our evaluation license from PBL is subject. Pursuant to the agreement, we have the ability to evaluate the technology~~in-house~~ ~~with the option of acquiring a worldwide exclusive right. The agreement will endure for a period of three years from the effective date.~~

Our Intellectual Property Strategy

We believe our current layered patent estate, together with our efforts to develop and patent next generation technologies, provides us with substantial intellectual property protection. However, the area of patent and other intellectual property rights in biotechnology is an evolving one with many risks and uncertainties.

Our strategy is also to develop and obtain additional intellectual property covering innovative manufacturing processes and methods for genetically engineeringT-cells expressing new constructs and for genetically engineering plants expressing new traits. To support this effort, we have established expertise and development capabilities focused in the areas ofpre-clinical research and development, manufacturing and manufacturing processscale-up, quality control, quality assurance, regulatory affairs and clinical trial design and implementation. Thus, we expect to file additional patent applications to expand this layer of our intellectual property estate.

The term of individual patents depends upon the legal term of the patents in the countries in which they are obtained. In most countries in which we file, the patent term is 20 years from the date of filing of the first

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non-provisional application to which priority is claimed. In the United States, a patent’s term may be lengthened by patent term adjustment, which compensates a patentee for administrative delays by the U.S. Patent and Trademark Office in granting a patent, or may be shortened if a patent is terminally disclaimed over an earlier-filed patent. The term of a patent that covers anFDA-approved drug may also be eligible for a patent term restoration of up to five years under the Hatch-Waxman Act, which is designed to compensate for the patent term lost during the FDA regulatory review process. The length of the patent term restoration is calculated based on the length of time the drug is under regulatory review. A patent term restoration under the Hatch-Waxman Act cannot extend the remaining term of a patent beyond a total of 14 years from the date of product approval and only one patent applicable to an approved drug may be restored. Moreover, a patent can only be restored once, and thus, if a single patent is applicable to multiple products, it can only be extended based on one product. Similar provisions are available in Europe and certain other foreign jurisdictions to extend the

term of a patent that covers an approved drug. When possible, depending upon the length of clinical trials and other factors involved in the filing of a BLA, we expect to apply for patent term extensions for patents covering our product candidates and their methods of use.

Our commercial success may depend in part on our ability to obtain and maintain patent and other proprietary protection for commercially important technology, inventions andknow-how related to our business; defend and enforce our patents; preserve the confidentiality of our trade secrets; and operate without infringing the valid enforceable patents and proprietary rights of third parties. Our ability to stop third parties from making, using, selling, offering to sell or importing our products may depend on the extent to which we have rights under valid and enforceable patents or trade secrets that cover these activities. With respect to both licensed and company-owned intellectual property, we cannot be sure that patents will be granted with respect to any of our pending patent applications or with respect to any patent applications filed by us in the future, nor can we be sure that any of our existing patents or any patents that may be granted to us in the future will be commercially useful in protecting our commercial products and methods of manufacturing the same.

We may rely, in some circumstances, on trade secrets to protect our technology. However, trade secrets can be difficult to protect. We seek to protect our proprietary technology and processes, in part, by entering into confidentiality agreements with our employees, consultants, scientific advisors and contractors. We also seek to preserve the integrity and confidentiality of our data and trade secrets by maintaining physical security of our premises and physical and electronic security of our information technology systems. While we have confidence in these individuals, organizations and systems, agreements or security measures may be breached, and we may not have adequate remedies for any breach. In addition, our trade secrets may otherwise become known or be independently discovered by competitors. To the extent that our consultants, contractors or collaborators use intellectual property owned by others in their work for us, disputes may arise as to the rights in related or resultingknow-how and inventions.

Competition

The biotechnology and pharmaceutical industries put significant resources toward developing novel and proprietary therapies for the treatment of cancer, which often incorporate novel technologies and incorporate valuable intellectual property. We compete with companies in the immunotherapy space, as well as companies developing novel targeted therapies for cancer. In addition, our products will compete with existing standards of care for the diseases that our product candidates target. We anticipate that we will face intense and increasing competition from many different sources, including new and established biotechnology and pharmaceutical companies, academic research institutions, governmental agencies and public and private research institutions.

The immuno-oncology cell therapy competitive landscape is increasing, with the main approaches includingCAR-T cells (autologous and allogeneic) and autologousT-cell receptors (TCRs).

The most advanced autologousCAR-T cell programs are:

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two products approved in United States (see below for more details): Kymriah^®, commercialized by Novartis (targeting CD19 inB-cell Acute Lymphoblastic Lymphoma and largeB-cell lymphoma) and Yescarta^® commercialized by Kite (targeting CD19 inB-cell Non Hodgkin Lymphoma);

3 product candidates in pivotal clinical trials in the United States: Juno’s product candidate JCAR017 (targeting CD19 inB-cell Non Hodgkin Lymphoma), bluebird’s product candidate bb2121 (targeting BCMA in Multiple Myeloma; in partnership with Celgene), Legend Bio’s LCAR-B38M (target BCMA in Multiple Myeloma; in partnership with Janssen).

Our competitors include:

Gene-editing space: CRISPR Therapeutics, Inc., Editas Medicine, Inc., Intellia Therapeutics, Inc., ~~Caribou Biosciences,~~ Precision BioSciences, Inc. and Sangamo BioSciences, Inc.

Autologous and Allogeneic CART-cell space: ~~Bellicum Pharmaceuticals, Inc.,~~ Juno Therapeutics, Inc. ~~(acquired~~(in collaboration with Editas Medicine Inc.), acquired by Celgene ~~Corporation),~~Corporation and acquired since by Bristol-Myers Squibb; Bluebird bio, Inc., (in collaboration with Celgene Corporation); Ziopharm Oncology Inc. (in collaboration with Intrexon Corporation); Kite Pharma Inc. (in collaboration with Amgen Inc. and with Sangamo Therapeutics Inc.), ~~Kite~~acquired by Gilead Sciences Inc.; Novartis AG (in collaboration with Intellia Inc.); Johnson & Johnson (in collaboration with Transposagen Biopharmaceuticals Inc.); Precision Biosiences (in collaboration with Shire Plc, asset acquired since by Servier), Regeneron Pharmaceuticals Inc. (in collaboration with Adicet Bio Inc); Fate Therapeutics Inc.; CRISPR Therapeutics Inc. (in collaboration with Bayer AG and Vertex Inc.).

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~~Pharma, Inc. (in collaboration with Amgen and acquired by Gilead Sciences, Inc.), Novartis AG and Johnson & Johnson (in collaboration with Transposagen), and Autolus Limited.~~

Cell-therapy space: Adaptimmune Ltd, ~~Lion Biotechnologies, Inc.,~~Iovance Biotherapeutics, Unum Therapeutics, Inc., NantKwest, Inc., Celyad S.A., Atara Biotherapeutics, Inc., and Immunocore Ltd.

Agricultural biotechnology space:

Companies developing plants with enhanced properties: Arcadia Biosciences, ~~Inc., Chromatin~~ Inc., Cibus Global, Ltd., Evogene Ltd., Danzinger Innovation Ltd., Keygene N.V. ~~and~~ Precision PlantSciences, Inc., Pairwise, Inari, Bension Hill.

Large Agricultural Biotechnology, Seed and Chemical Companies: BASF SE, Bayer AG, ~~DowDuPont, Monsanto Co., Syngenta AG,~~Corteva, ChemChina, and Takii & Company, LTD.

Specialty Food Ingredient Companies: International Flavors & Fragrances Inc., Givaudan, Keery Group plc, CSM N.V., FMC Corporation, CP Kelco, Novzymes, Ingredion Incorporated and Royal DSM N.V.

In August 2017, Novartis AG has obtained approval from the FDA to commercialize the firstCAR-T cell therapy, Kymriah^®, for children and young adults withB-cell ALL that is refractory or has relapsed at least twice, and has recently been granted US FDA priority review for adults with refractory/relapsed DLBCL, priced at $475,000. In May 2018, the FDA approved a label extension for Kymriah^® for the treatment of adult patients with relapsed or refractory largeB-cell lymphoma after two or more lines of systemic therapy, priced at $373,000. In October 2017, Kite Pharma, Inc. (acquired by Gilead Sciences, Inc.), acquired since by Bristol-Myers Squibb) has obtained approval from the FDA to commercialize Yescarta^®, the firstCAR-T cell therapy for the treatment of adult patients with relapsed or refractory largeB-cell lymphoma after two or more lines of systemic therapy, priced at $373,000. Due to the promising therapeutic effect ofT-cell therapies in clinical exploratory trials, we anticipate substantial direct competition from other existing and new competitors developing these therapies. In particular, we expect to compete with therapies with tumor infiltrating lymphocytes, or TILs, that are naturally occurring tumor-reactiveT-cells harvested, propagatedex vivo andre-infused into patients. We also expect to compete with therapies using genetically engineeredT-cells, rendered reactive against tumor-associated antigens prior to their administration to patients. However, we believe that most of our competitors are currently focused on autologous therapies, and we believe that we are the most advanced company developing the allogeneicCAR-T cell approach. In addition, we differentiate ourselves by using our gene-editing capabilities to add specific features to ourT-cell products, such as cancer drug resistance or resistance to checkpoint inhibition.

We also face competition fromnon-cell based treatments offered by companies such as Amgen Inc., AstraZeneca plc, Bristol-Myers Squibb Company, Incyte Corporation, Merck & Co., Inc., and F.Hoffman-La Roche AG. Immunotherapy is further being pursued by several biotech companies as well as bylarge-cap pharmaceuticals. Many of our current or potential competitors, either alone or with their collaboration partners, have significantly greater financial resources and expertise in research and development, manufacturing,pre-clinical testing, conducting clinical trials, and marketing approved products than we do. Mergers and acquisitions in the pharmaceutical, biotechnology and gene therapy industries may result in even more resources being concentrated among a smaller number of our competitors. Smaller or early-stage companies may also prove to be significant competitors, particularly through collaborative arrangements with large and established companies. These competitors also compete with us in recruiting and retaining qualified scientific and management personnel and establishing clinical trial sites and patient registration for clinical trials, as well as in acquiring technologies complementary to, or necessary for, our programs.

Our commercial opportunity could be reduced or eliminated if our competitors develop and commercialize products that are safer, more effective, have fewer or less severe side effects, are more convenient or are less expensive than any products that we may develop. Our competitors also may obtain FDA or other regulatory approval for their products more rapidly than we may obtain approval for ours, which could result in our competitors establishing a strong market position before we are able to enter the market or make our development more complicated. The key competitive factors affecting the success of all of our programs are likely to be their efficacy, safety, and convenience.

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Government Regulation and Product Approval

Government Regulation of Biological Products

We are subject to extensive regulation. ~~We expect our cell~~Our product candidates ~~to be~~are regulated as ~~biologics.~~biologics, gene therapies. Governmental authorities, including the FDA and comparable regulatory authorities in other countries, regulate the design, development, production / manufacturing, testing, safety, efficacy, labeling, storage, record-keeping, advertising, promotion and marketing of pharmaceutical products, including biologics.Non-compliance with applicable requirements can result in fines and other judicially imposed sanctions, including product seizures, import restrictions, injunctive actions and criminal prosecutions of both companies and individuals. In addition, administrative remedies can involve requests to recall violative products; the refusal of the government to enter into supply contracts; or the refusal to approve pending product approval applications until manufacturing or other alleged deficiencies are brought into compliance. The FDA and similar authorities around the world also have the authority to cause the withdrawal of approval of a marketed product, to impose labeling restrictions or to require that we redo somenon-clinical and/or clinical studies.

The FDA categorizes human cell- or tissue-based products as either minimally manipulated or more than minimally manipulated, and has determined that more than minimally manipulated products require clinical trials to demonstrate product safety and efficacy and the submission of a BLA for marketing authorization.

Our product candidates must be approved by the FDA before they may be legally marketed in the United States and by the appropriate foreign regulatory agencies before they may be legally marketed in foreign countries. Generally, our activities in foreign countries will be subject to regulation that is similar in nature and scope as that imposed in the United States, although there can be important differences. Additionally, some significant aspects of regulation in Europe are addressed in a centralized way, but country-specific regulation remains essential in many respects. The process for obtaining regulatory marketing approvals and the subsequent compliance with appropriate federal, state, local and foreign statutes and regulations require the expenditure of substantial time and financial resources.

Ethical, social and legal concerns about gene therapy, gene modifications, genetic testing and genetic research could result in additional regulations restricting or prohibiting the processes we may use. Federal and state agencies, congressional committees and foreign governments have expressed interest in further regulating biotechnology. More restrictive regulations or claims that our products are unsafe or pose a hazard could prevent us from commercializing any products in one or more jurisdictions. New government requirements may be established that could delay or prevent regulatory approval of our product candidates under development. It is impossible to predict whether legislative changes will be enacted, regulations, policies or guidance changed, or interpretations by agencies or courts changed, or what the impact of such changes, if any, may be.

Set forth below is a description of the process of obtaining U.S. government approval for biological product development. Similar processes apply in other jurisdictions.

U.S. Biological Product Development

In the United States, the FDA regulates biologics under the Federal Food, Drug, and Cosmetic Act, or FDCA, and the Public Health Service Act, or PHSA, and their implementing regulations. Biologics are also subject to other federal, state and local statutes and regulations. The process required by the FDA before biologic product candidates may be marketed in the United States and the subsequent compliance with appropriate federal, state, local and foreign statutes and regulations require the expenditure of substantial time and financial resources. Failure to comply with the applicable U.S. requirements at any time during the product development process, approval process or after approval, may subject an applicant to administrative or judicial sanctions. These sanctions could include, among other actions, the FDA’s refusal to approve pending applications, withdrawal of an approval, a clinical hold, untitled or warning letters, product recall requests or withdrawals from the market, labeling restrictions,non-clinical and/or clinical studies to be performed again, product seizures,

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total or partial suspension of production or distribution injunctions, import restrictions, fines, refusals of government contracts, restitution, disgorgement, or civil or criminal penalties for both companies and individuals. Any agency or judicial enforcement action could have a material adverse effect on us.

Our biological product candidates must be approved by the FDA through the Biologics License Application, or BLA, process before they may be legally marketed in the United States. The process required by the FDA before a biologic may be marketed in the United States generally involves the following:

completion of extensive nonclinical, sometimes referred to aspre-clinical laboratory tests,pre-clinical animal studies and formulation studies in accordance with applicable regulations, including the FDA’s GLP regulations;

production and testing of clinical products according to the current Good Manufacturing Practices (cGMP) and possible FDA product specific requirements

submission to the FDA of an IND, which must become effective before clinical trials may begin and must be updated at least annually;

performance of adequate and well-controlled clinical trials in accordance with applicable IND and other clinical trial-related regulations, sometimes referred to as Good Clinical Practices, or GCPs, to establish the safety and efficacy of the proposed product candidate for each proposed indication;

submission to the FDA of a BLA;

satisfactory completion of an FDApre-approval inspection of the manufacturing facility or facilities where the active pharmaceutical ingredient, or API, and finished product are manufactured to assess compliance with the IND/BLA and FDA’s cGMP requirements to assure that the facilities, methods and controls are adequate to preserve the product’s identity, strength, quality, purity and potency;

FDA review and approval of the BLA prior to any commercial marketing or sale of the product in the United States.

The data required to support a BLA is generated in three distinct development stages: manufacturing,pre-clinical and clinical. The manufacturing development stage generally involves laboratory evaluations of drug chemistry and biology properties, formulation and stability, thepre-clinical stage generally involves studies to evaluate pharmacology and toxicity in animals, which support subsequent clinical testing. The conduct of the manufacturing andpre-clinical studies must comply with federal regulations, including GMPs and GLPs for the main Toxicology Studies.

Where a trial involving the deliberate transfer of (1) recombinant DNA or (2) DNA or RNA derived from recombinant DNA into one or more human research participants (including recombinant DNA molecules, and/or organisms and viruses containing recombinant DNA molecules) (a gene therapy trial) is conducted at, or sponsored by, institutions receiving NIH funding for recombinant DNA research, a protocol and related documentation is submitted to and the trial is registered with the NIH Office of Biotechnology Activities, or OBA, prior to the submission of an IND to the FDA. This is done pursuant to the NIH Guidelines for Research Involving Recombinant or Synthetic Nucleic Acid Molecules, or NIH Guidelines. Compliance with the NIH Guidelines is mandatory for investigators at institutions receiving NIH funds for research involving recombinant DNA, however many companies and other institutions not otherwise subject to the NIH Guidelines voluntarily follow them. The NIH is responsible for convening the RAC, a federal advisory committee, which discusses protocols that raise novel or particularly important scientific, safety or ethical considerations at one of its quarterly public meetings. The OBA will notify the FDA of the RAC’s decision regarding the necessity for full public review of a gene therapy protocol. RAC proceedings and reports are posted to the OBA web site and may be accessed by the public.

The sponsor must submit the results of thepre-clinical studies, together with manufacturing information, analytical data, any available clinical data or literature and a proposed clinical protocol, to the FDA as part of an

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IND before any clinical testing may proceed. An IND is a request for authorization from the FDA to administer an investigational drug product to humans. The IND must become effective before clinical trials may begin. The IND is automatically approved 30 days after receipt by the FDA, unless during that time the FDA raises concerns or questions regarding the proposed clinical trials. In such a case, the FDA may place the IND on clinical hold and the IND sponsor and the FDA must resolve any outstanding concerns before the clinical trial can begin. With gene therapy protocols, if the FDA allows the IND to proceed, and the RAC decides that full public review of the protocol is warranted but did not take place before the IND review is complete, the FDA will request at the completion of its IND review that sponsors delay initiation of the protocol until after completion of the RAC review process. The FDA may also impose clinical holds on a product candidate at any time before or during clinical trials due to safety concerns ornon-compliance. Accordingly, we cannot be sure that submission of an IND will result in the FDA allowing clinical trials to begin, or that, once begun, issues will not arise that could cause the trial to be suspended or terminated.

Before the IND becomes active, the Clinical Protocol will also need to be approved by the relevant Institutional Review Boards (IRBs) and Institutional Biosafety Committees (IBCs), which are the cornerstone of institutional oversight of recombinant DNA clinical research.

Clinical trials involve the administration of the investigational product to human subjects under the supervision of qualified investigators, generally physicians not employed by or under the trial sponsor’s control, in accordance with GCPs, which include the requirement that all research subjects provide their informed consent for their participation in any clinical trial. Clinical trials are conducted under protocols detailing, among other things, the objectives of the clinical trial, dosing procedures, subject selection and exclusion criteria, and the parameters to be used to monitor subject safety and assess efficacy. Each protocol, and any subsequent amendments to the protocol, must be submitted to the FDA as part of the IND. Further, each clinical trial must be reviewed and approved by an independent institutional review board, or IRB, at or servicing each institution at which the clinical trial will be conducted. An IRB is charged with protecting the welfare and rights of trial participants and considers such items as whether the risks to individuals participating in the clinical trials are minimized and are reasonable in relation to anticipated benefits. The IRB also approves the informed consent form that must be provided to each clinical trial subject or his or her legal representative and must monitor the clinical trial until completed.

All gene therapy experiments and clinical trials are also subject to review and oversight by an institutional biosafety committee, or IBC, a local institutional committee that reviews and oversees basic and clinical research conducted at that institution. The IBC assesses the safety of the research and identifies any potential risk to public health or the environment.

There are also requirements governing the reporting of ongoing clinical trials and completed clinical trial results to public registries. Sponsors of clinical trials ofFDA-regulated products, including biologics, are required to register and disclose certain clinical trial information, which is publicly available at www.clinicaltrials.gov. Information related to the product, patient population, phase of investigation, study sites and investigators, and other aspects of the clinical trial is then made public as part of the registration. Sponsors are also obligated to discuss the results of their clinical trials after completion. Disclosure of the results of these trials can be delayed until the new product or new indication being studied has been approved.

Human clinical trials are typically conducted in three sequential phases. However, these phases may overlap or be combined:

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Phase I. The biological product candidate is initially introduced into healthy human subjects and tested for safety. In the case of some products for severe or life-threatening diseases, especially when the product may be too inherently toxic to ethically administer to healthy volunteers, ifpre-clinical testing warrants, the initial human testing may be conducted in patients with the condition of interest.

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Phase II. The biological product candidate is evaluated in a limited patient population with the condition of interest to identify possible adverse effects and safety risks, to preliminarily evaluate the

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efficacy of the product for specific targeted diseases and to determine dosage tolerance, optimal dosage and dosing schedule.

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Phase III. Clinical trials are undertaken to further evaluate dosage, clinical efficacy, potency and safety in an expanded patient population with the condition of interest at geographically dispersed clinical trial sites. These clinical trials are intended to establish the overall risk to benefit ratio of the product and provide an adequate basis for approval, including appropriate product labeling.

Post-approval clinical trials, sometimes referred to as “Phase 4” clinical trials, may be conducted after initial marketing approval. These clinical trials are used to gain additional experience from the treatment of patients in the intended therapeutic

indication, particularly for long-term safetyfollow-up. The FDA recommends that sponsors observe subjects for potential gene therapy-related delayed adverse events for a15-year period, including a minimum of five years of annual examinations followed by ten years of annual queries, either in person or by questionnaire, of study subjects.

During all phases of clinical development, regulatory agencies require extensive monitoring and auditing of all clinical activities, clinical data, and clinical trial investigators. Annual progress reports detailing the results of the clinical trials must be submitted to the FDA. Written IND safety reports must be promptly submitted to the FDA, ~~the NIH,~~ IRB, and the investigators for serious and unexpected adverse events, any findings from other studies, tests in laboratory animals orin vitro testing that suggest a significant risk for human patients, or any clinically important increase in the rate of a serious suspected adverse reaction over that listed in the protocol or investigator brochure. The sponsor must submit an IND safety report within 15 calendar days after the sponsor determines that the information qualifies for reporting. The sponsor also must notify the FDA of any unexpected fatal or life-threatening suspected adverse reaction within seven calendar days after the sponsor’s initial receipt of the information.

Phase I, Phase II and Phase III clinical trials may not be completed successfully within any specified period, if at all. The FDA or the sponsor or its data safety monitoring board may suspend or terminate a clinical trial at any time on various grounds, including a finding that the research patients are being exposed to an unacceptable health risk, including risks inferred from other unrelated immunotherapy trials. Similarly, an IRB can suspend or terminate approval of a clinical trial at its institution if the clinical trial is not being conducted in accordance with the IRB’s requirements or if the biological product has been associated with unexpected serious harm to patients.

Human immunotherapy products and gene therapy products are a new category of therapeutics. Because this is a relatively new and expanding area of novel therapeutic interventions, there can be no assurance as to the length of the trial period, the number of patients the FDA will require to be enrolled in the trials in order to establish the safety, efficacy, purity and potency of immunotherapy products, or that the data generated in these trials will be acceptable to the FDA to support marketing approval.

Concurrently with clinical trials, companies usually complete additional animal studies and must also develop additional information about the biological and physical characteristics of the biological product as well as finalize a process for production and testing the product in commercial quantities in accordance with cGMP requirements. To help reduce the risk of the introduction of adventitious agents with use of biological products, the PHSA emphasizes the importance of manufacturing control for products whose attributes cannot be precisely defined. The manufacturing process must be capable of consistently producing quality batches of the product candidate and, among other things, the sponsor must develop and validate methods for testing the identity, strength, quality, potency and purity of the final biological product. Additionally, appropriate packaging must be selected and tested and stability studies must be conducted to demonstrate that the biological product candidate does not undergo unacceptable deterioration over its shelf life.

U.S. Review and Approval Processes for Biological Product Candidates

After the completion of clinical trials,non-clinical and manufacturing activities of a biological product candidate, FDA approval of a BLA must be obtained before commercial marketing of the biological product. The

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BLA must include results of product development, laboratory and animal studies, human trials, information on the manufacture and composition of the product, proposed labeling and other relevant information. The approval processes require substantial time and effort and there can be no assurance that the FDA will accept the BLA for filing and, even if filed, that any approval will be granted on a timely basis, if at all.

Under the Prescription Drug User Fee Act, or PDUFA, as amended, each BLA must be accompanied by a significant user fee. The FDA adjusts the PDUFA user fees on an annual basis. PDUFA also imposes an annual product fee for biological products and an annual establishment fee on facilities used to manufacture prescription biological products. Fee waivers or reductions are available in certain circumstances, including a waiver of the application fee for the first application filed by a small business. Additionally, no user fees are assessed on BLAs for products designated as orphan drugs, unless the product also includes anon-orphan indication.

Within 60 days following submission of the application, the FDA reviews a BLA submitted to determine if it is substantially complete before the agency accepts it for filing. The FDA may refuse to file any BLA that it deems incomplete or not properly reviewable at the time of submission and may request additional information. In this event, the BLA must be resubmitted with the additional information. The resubmitted application also is subject to review before the FDA accepts it for filing. Once the submission is accepted for filing, the FDA begins anin-depth substantive review of the BLA. The FDA reviews the BLA to determine, among other things, whether the proposed product is safe and potent, or effective, for its intended use, and has an acceptable purity profile, and whether the product is being manufactured in accordance with cGMP regulations to assure and preserve the product’s identity, safety, strength, quality, potency and purity. The FDA may refer applications for novel biological products or biological products that present difficult questions of safety or efficacy to an advisory committee, typically a panel that includes clinicians and other experts, for review, evaluation and a recommendation as to whether the application should be approved and under what conditions. The FDA is not bound by the recommendations of an advisory committee, but it considers

such recommendations carefully when making decisions. During the biological product approval process, the FDA also will determine whether a Risk Evaluation and Mitigation Strategy, or REMS, is necessary to assure the safe use of the biological product candidate. A REMS may be imposed to ensure safe use of the drug, and could include medication guides, physician communication plans, or elements to assure safe use, such as restricted distribution methods, patient registries and other risk minimization tools. If the FDA concludes a REMS is needed, the sponsor of the BLA must submit a proposed REMS. The FDA will not approve a BLA without a REMS, if required.

Before approving a BLA, the FDA will inspect the facilities at which the product candidate isand the associated vector are manufactured. The FDA will not approve the product candidate unless it determines that the manufacturing processes and facilities are in compliance with cGMP requirements and adequate to assure consistent production of the product within required specifications. For cell based immunotherapy products, the FDA also will not approve the product if the manufacturer is not in compliance with the current good tissue practice, or GTP requirements, to the extent applicable. These requirements are set out in FDA regulations and guidance documents and govern the methods used in, and the facilities and controls used for, the manufacture of human cells, tissues, and cellular and tissue based products, or HCT/Ps, which are human cells or tissue intended for use in implantation, transplantation, infusion, or transfer into a human recipient. The primary intent of the GTP requirements is to ensure that cell and tissue based products are manufactured in a manner designed to prevent the introduction, transmission and spread of communicable disease. FDA regulations also require tissue establishments to register and list their HCT/Ps with the FDA and, when applicable, to evaluate donors through screening and testing. Additionally, before approving a BLA, the FDA may inspect one or more clinical sites to assure that the clinical trials were conducted in compliance with IND trial requirements and GCP requirements. To assure cGMP, GTP and GCP compliance, an applicant must incur significant expenditure of time, money and effort in the areas of training, record keeping, production, and quality control.

Notwithstanding the submission of relevant data and information, the FDA may ultimately decide that the BLA does not satisfy its regulatory criteria for approval and deny approval. Data obtained from clinical trials are not always conclusive and the FDA may interpret data differently than we interpret the same data. If the agency

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decides not to approve the BLA in its submitted form, the FDA will issue a complete response letter that describes all of the specific deficiencies in the BLA identified by the FDA. The deficiencies identified may be minor, for example, requiring labeling changes, or major, for example, requiring additional clinical trials. Additionally, the complete response letter may include recommended actions that the applicant might take to place the application in a condition for approval. If a complete response letter is issued, the applicant may either resubmit the BLA, addressing all of the deficiencies identified in the letter, or withdraw the application.

If a product candidate receives regulatory approval, the approval may be significantly limited to specific diseases and dosages or the indications for use may otherwise be limited, which could restrict the commercial value of the product.

Further, the FDA may require that certain contraindications, warnings or precautions be included in the product labeling. The FDA may impose restrictions and conditions on product distribution, prescribing, or dispensing in the form of a REMS, or otherwise limit the scope of any approval. In addition, the FDA may require post marketing clinical trials, sometimes referred to as Phase 4 clinical trials, or additional studies like safety studies, designed to further assess a biological product’s safety and effectiveness, and testing and surveillance programs to monitor the safety of approved products that have been commercialized.

In addition, unless a waiver is granted, under the Pediatric Research Equity Act, or PREA, a BLA or supplement to a BLA must contain data to assess the safety and effectiveness of the product for the claimed indications in all relevant pediatric subpopulations and to support dosing and administration for each pediatric subpopulation for which the product is safe and effective. The Food and Drug Administration Safety and Innovation Act, or FDASIA, requires that a sponsor who is planning to submit a marketing application for a drug or biological product that includes a new active ingredient, new indication, new dosage form, new dosing regimen or new route of administration submit an initial Pediatric Study Plan, or PSP, within sixty days after anend-of-Phase II meeting or as may be agreed between the sponsor and FDA. The initial PSP must include, among other things, an outline of the pediatric study or studies that the sponsor plans to conduct, including to the extent practicable study objectives and design, age groups, relevant endpoints and statistical approach, or a justification for not including such detailed information, and any request for a deferral of pediatric assessments or a full or partial waiver of the requirement to provide data from pediatric studies along with supporting information. FDA and the sponsor must reach agreement on the PSP. A sponsor can submit amendments to an agreed-upon initial PSP at any time if changes to the pediatric plan need to be considered based on data collected from nonclinical studies, early phase clinical trials, and/or other clinical development programs. The FDA may grant deferrals for submission of data or full or partial waivers. Unless otherwise required by regulation, PREA does not apply to any product for an indication for which orphan designation has been granted. However, if only one indication for a product has orphan designation, a pediatric assessment may still be required for any applications to market that same product for thenon-orphan indications.

One of the performance goals agreed to by the FDA under the PDUFA is to review 90% of standard BLAs in 10 months and 90% of priority BLAs in six months, whereupon a review decision is to be made. The FDA does not always meet its PDUFA goal

dates for standard and priority BLAs and its review goals are subject to change from time to time. The review process and the PDUFA goal date may be extended by three months if the FDA requests or the BLA sponsor otherwise provides additional information or clarification regarding information already provided in the submission within the last three months before the PDUFA goal date.

Orphan Drug Designation

Under the Orphan Drug Act, a sponsor may request and the FDA may grant orphan designation to a drug or biologic intended to treat a rare disease or condition, which is generally a disease or condition that affects fewer than 200,000 individuals in the United States, or if it affects more than 200,000 individuals in the United States and there is no reasonable expectation that the cost of developing and making available in the United States drug or biologic for this type of disease or condition will be recovered from sales in the United States for that product.

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Orphan drug designation must be requested before submitting a BLA. After the FDA grants orphan drug designation, the generic and trade name, if any, of the drug or biologic and the rare disease or condition for which orphan-drug designation was granted are disclosed publicly by the FDA. The orphan drug designation does not convey any advantage during, or shorten the duration of, the regulatory review or approval process.

If a product that has orphan drug designation subsequently receives the first FDA approval for the disease or condition for which it has such designation, the product is entitled to orphan product exclusivity, which means that the FDA may not approve any other applications, including a full BLA, to market the same drug or biologic for the same indication for seven years, except in limited circumstances, such as a showing of clinical superiority to the product with orphan drug exclusivity. Orphan drug exclusivity does not prevent FDA from approving a different drug or biologic for the same disease or condition, or the same drug or biologic for a different disease or condition. Among the other benefits of orphan drug designation are tax credits for certain research and a waiver of the BLA application user fee.

A designated orphan drug may not receive orphan drug exclusivity if it is approved for a use that is broader than the indication for which it received orphan designation. Orphan exclusivity also could block the approval of one of our products for seven years if a competitor obtains approval of the same drug or biologic as defined by the FDA or if our product candidate is determined to be contained within the competitor’s product for the same indication or disease. In addition, exclusive marketing rights in the United States may be lost if the FDA later determines that the request for designation was materially defective or if the manufacturer is unable to assure sufficient quantities of the product to meet the needs of patients with the rare disease or condition.

The criteria for designating an “orphan medicinal product” in the EU are similar to those in the United States. Such designation can be requested in the case of products that are intended for the diagnosis, prevention or treatment of a life-threatening or chronically debilitating condition and either (a) such condition affects no more than five in 10,000 persons in the EU when the application is made, or (b) the product, without the benefits derived from orphan status, would unlikely generate sufficient return in the EU to justify the necessary investment. Moreover, in order to obtain orphan designation it is necessary to demonstrate that there exists no satisfactory method of diagnosis, prevention or treatment of such condition authorized for marketing in the EU, or if such a method exists, the product will be of significant benefit to those affected by the condition.

In the EU, orphan medicinal products are eligible for financial incentives as well as specific regulatory assistance and scientific advice. Products receiving orphan status in the EU can receive ten years of market exclusivity, during which time no similar medicinal product for the same indication may be placed on the market. An orphan product can also obtain an additional two years of market exclusivity in the EU for pediatric studies. No extension to any supplementary protection certificate can be granted on the basis of pediatric studies for orphan indications.

However, the10-year market exclusivity may be reduced to six years in certain circumstances, including for example if, at the end of the fifth year, it is established that the product is sufficiently profitable not to justify maintenance of market exclusivity.

There can be no assurance that we will receive orphan drug designation for any product candidates in the United States, in the EU or in any other market. There can be no assurance that an Orphan exclusivity from a competitor could not block the approval of one of our products for a certain period of time, in the United States, in the EU or in any other market.

Expedited Development and Review Programs

The FDA has a Fast Track program that is intended to expedite or facilitate the process for reviewing new drugs and biological products that meet certain criteria. Specifically, new products are eligible for Fast Track designation if they are intended to treat a serious or life-threatening disease or condition and demonstrate the

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potential to address unmet medical needs for the disease or condition. Fast Track designation applies to the combination of the product and the specific indication for which it is being studied. The sponsor of a new drug or biologic may request the FDA to designate the drug or biologic as a Fast Track

product candidate at any time during the clinical development of the product candidate. Unique to a Fast Track product, the FDA may consider the review of sections of the BLA on a rolling basis before the complete application is submitted, if the sponsor provides a schedule for the submission of the sections of the BLA, the FDA agrees to accept sections of the BLA and determines that the schedule is acceptable, and the sponsor pays any required user fees upon submission of the first section of the BLA.

Any product candidate, submitted to the FDA for approval, including a product with a Fast Track designation, may also be eligible for other types of FDA programs intended to expedite development and review, such as priority review and accelerated approval. A product candidate is eligible for priority review if it has the potential to provide safe and effective therapy where no satisfactory alternative therapy exists or a significant improvement in the treatment, diagnosis or prevention of a disease compared to marketed products. The FDA will attempt to direct additional resources to the evaluation of an application for a new product candidate designated for priority review in an effort to facilitate the review, and aims to review such applications within six months as opposed to ten months for standard review. Additionally, a product candidate may be eligible for accelerated approval. Product candidates studied for their safety and effectiveness in treating serious or life-threatening illnesses and that provide meaningful therapeutic benefit over existing treatments may receive accelerated approval which means that they may be approved on the basis of adequate and well-controlled clinical trials establishing that the product candidate has an effect on a surrogate endpoint that is reasonably likely to predict clinical benefit, or on the basis of an effect on a clinical endpoint other than irreversible morbidity or mortality or other clinical benefit, taking into account the severity, rarity, or prevalence of the condition and the availability or lack of alternative treatments. As a condition of approval, the FDA may require that a sponsor of a drug or biological product candidate receiving accelerated approval perform adequate and well-controlled post-marketing clinical trials. In addition, the FDA currently requires as a condition for accelerated approvalpre-approval of promotional materials, which could adversely impact the timing of the commercial launch of the product. Fast Track designation, priority review and accelerated approval do not change the standards for approval but may expedite the development or approval process.

Breakthrough Therapy / Regenerative Medicine Advanced Therapy Designation

Under the provisions of the Food and Drug Administration Safety and Innovation Act, or FDASIA, enacted in 2012, the FDA established a Breakthrough Therapy Designation which is intended to expedite the development and review of products that treat serious or life-threatening conditions. A breakthrough therapy is defined as a drug that is intended, alone or in combination with one or more other drugs, to treat a serious or life-threatening disease or condition, and preliminary clinical evidence indicates that the drug may demonstrate substantial improvement over existing therapies on one or more clinically significant endpoints, such as substantial treatment effects observed early in clinical development. The designation includes all of the features of Fast Track designation, as well as more intensive FDA interaction and guidance. The Breakthrough Therapy Designation is a distinct status from both accelerated approval and priority review, but these can also be granted to the same product candidate if the relevant criteria are met.

The FDA must take certain actions, such as holding timely meetings and providing advice, intended to expedite the development and review of an application for approval of a breakthrough therapy. All requests for breakthrough therapy designation will be reviewed within 60 days of receipt, and FDA will either grant or deny the request.

In addition, as described in Section 3033 of the 21st Century Cures Act, signed into law in December 2016, a drug is eligible for Regenerative Medicine Advanced Therapy (RMAT) Designation if:

the drug is a regenerative medicine therapy, which is defined as a cell therapy, therapeutic tissue engineering product, human cell and tissue product, or any combination product using such therapies or products, except for those regulated solely under Section 361 of the Public Health Service Act and part 1271 of Title 21, Code of Federal Regulations;

the drug is intended to treat, modify, reverse, or cure a serious or life-threatening disease or condition; and

preliminary clinical evidence indicates that the drug has the potential to address unmet medical needs for such disease or condition.

The RMAT designation carries all of benefits of Breakthrough and Fast Track therapy designations, including: intensive interaction with FDA on an efficient drug development program beginning as early as phase 1, organizational commitment involving senior FDA personnel, and rolling BLA review. RMAT designees are also eligible foraccelerated approval and priority review if relevant criteria are met.

Where applicable, we plan to request Fast Track and/or Breakthrough Therapy Designation for our product candidates. Even if we receive one of these designations for our product candidates, the FDA may later decide that our product candidates no longer meet the conditions for qualification. In addition, these designations may not provide us with a material commercial advantage.

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Post-Approval Requirements

Maintaining compliance with applicable federal, state, and local statutes and regulations requires the expenditure of substantial time and financial resources. Rigorous and extensive FDA regulation of biological products continues after approval, particularly with respect to cGMP and pharmacovigilance requirements as well as post marketing commitments. Any products for which we receive FDA approval will be subject to continuing regulation by the FDA, including, among other things, record-keeping requirements, reporting of adverse experiences with the product, providing the FDA with updated safety and efficacy information, product sampling and distribution requirements, and complying with FDA promotion and advertising requirements, which include, among others, standards fordirect-to-consumer advertising, restrictions on promoting products for uses or in patient populations that are not described in the product’s approved uses (known asoff-label use), limitations on industry-sponsored scientific and educational activities, and requirements for promotional activities involving the internet. Although physicians may prescribe legally available products foroff-label use that they deem to be appropriate in their professional medical judgment, manufacturers may not market or promote suchoff-label uses.

Other post-approval requirements applicable to biological products include reporting of cGMP deviations that may affect the identity, potency, purity and overall safety of a distributed product, record-keeping requirements, reporting of adverse effects, reporting updated safety and efficacy information, and complying with electronic record and signature requirements. After a BLA is approved, the product may also be subject to official lot release. In this case, as part of the manufacturing process, the manufacturer is required to perform certain tests on each lot of the product before it is released for distribution. If the product is subject to official release by the FDA, the manufacturer submits samples of each lot of product to the FDA together with a release protocol showing a summary of the history of manufacture of the lot and the results of all of the manufacturer’s tests performed on the lot. The FDA also may perform certain confirmatory tests on lots of some products before releasing the lots for distribution by the manufacturer. In addition, the FDA conducts laboratory research related to the regulatory standards on the safety, purity, potency, and effectiveness of biological products.

In addition, we and any third-party manufacturers of our products will be required to comply with applicable requirements in the cGMP regulations, including quality control and quality assurance and maintenance of records and documentation. We rely, and expect to continue to rely, on third parties for the production of clinical and commercial quantities of our products in accordance with cGMP regulations. cGMP regulations require among other things, quality control and quality assurance as well as the corresponding maintenance of records and documentation and the obligation to investigate and correct any deviations from cGMP. Manufacturers and other entities involved in the manufacture and distribution of approved products are required to register their establishments with the FDA and certain state agencies, and are subject to periodic announced and unannounced inspections by the FDA and certain state agencies for compliance with cGMP and other The FDA also may require post-marketing studies, known as Phase 4 studies, and surveillance to monitor the effects of an approved product. laws. Accordingly, manufacturers must continue to expend time, money, and effort in the area of production and quality control to maintain cGMP compliance. Discovery of problems with a product after approval may result in restrictions on a product, manufacturer, or holder of an approved BLA, including, among other things, recall or withdrawal of the product from the market. In addition, changes to the manufacturing process are strictly regulated, and depending on the significance of the change, may require prior FDA approval before being implemented. Other types of changes to the approved product, such as adding new indications and claims, are also subject to further FDA review and approval.

~~The FDA also may require post-marketing studies, known as Phase 4 studies, and surveillance to monitor the effects of an approved product.~~ Discovery of previously unknown problems with a product or the failure to comply with applicable FDA requirements can have negative consequences, including adverse publicity, judicial or administrative enforcement, warning letters from the FDA, mandated corrective advertising or communications with doctors, and civil or criminal penalties, among others. Newly discovered or developed safety or effectiveness data may require changes to a product’s approved labeling, including the addition of new warnings and contraindications, and also may require the implementation of other risk management measures.

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Also, new government requirements, including those resulting from new legislation, may be established, or the FDA’s policies may change, which could delay or prevent regulatory approval of our products under development.

U.S. Patent Term Restoration and Marketing Exclusivity

The Biologics Price Competition and Innovation Act, or BPCIA, amended the PHSA to authorize the FDA to approve similar versions of innovative biologics, commonly known as biosimilars. A competitor seeking approval of a biosimilar must file an application to establish its molecule as highly similar to an approved innovator biologic, among other requirements. The BPCIA, however, bars the FDA from approving biosimilar applications for 12 years after an innovator biological product receives initial marketing approval. This12-year period of data exclusivity may be extended by six months, for a total of 12.5 years, if the FDA requests that the innovator company conduct pediatric clinical investigations of the product.

The first biological product submitted under the abbreviated approval pathway that is determined to be interchangeable with the reference product has exclusivity against other biologics submitting applications under the abbreviated approval pathway for the lesser of (1) one year after the first commercial marketing, (2) 18 months after approval if there is no legal challenge, (3) 18 months after the resolution in the applicant’s favor of a lawsuit challenging the biologic’s patents if an application has been submitted, or (4) 42 months after the application has been approved if a lawsuit is ongoing within the42-month period.

Depending upon the timing, duration and specifics of the FDA approval of the use of our product candidates, some of our U.S. patents, if granted, may be eligible for limited patent term extension under the Drug Price Competition and Patent Term Restoration Act of 1984, commonly referred to as the Hatch-Waxman Act. The Hatch-Waxman Act permits a patent restoration term of up to five years, as compensation for patent term lost during product development and the FDA regulatory review process. However, patent term restoration cannot extend the remaining term of a patent beyond a total of 14 years from the product’s approval date. The patent term restoration period is generallyone-half the time between the effective date of an IND and the submission date of a BLA plus the time between the submission date of a BLA and the approval of that application. Only one patent applicable to an approved product is eligible for the extension and the application for the extension must be submitted prior to the expiration of the patent. The U.S. Patent and Trademark Office, in consultation with the FDA, reviews and approves the application for any patent term extension or restoration. In the future as applicable, we may apply for restoration of patent term for one of our currently owned or licensed patents seeking restored patent life beyond its current expiration date, depending on the expected length of the clinical trials and other factors involved in the filing of the relevant BLA.

Pediatric exclusivity is another type of regulatory market exclusivity in the United States. Pediatric exclusivity, if granted, adds six months to existing exclusivity periods and patent terms. Thissix-month exclusivity, which attaches to and runs from the end of other exclusivity protection or patent term, may be granted based on the voluntary completion of a pediatric trial in accordance with anFDA-issued “Written Request” for such a trial.

Other U.S. Healthcare Laws and Compliance Requirements

In the United States, our activities are subject to regulation by various federal, state and local authorities in addition to the FDA, including but not limited to, the Centers for Medicare and Medicaid Services, or CMS, other divisions of the U.S. Department of Health and Human Services (e.g., the Office of Inspector General), the U.S. Department of Justice, or DOJ, and individual U.S. Attorney offices within the DOJ, and state and local governments. For example, sales, marketing and scientific/educational grant programs must comply with the anti-fraud and abuse provisions of the Social Security Act, the false claims laws, the privacy provisions of the Health Insurance Portability and Accountability Act, or HIPAA, and similar state laws, each as amended.

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The federal Anti-Kickback Statute prohibits, among other things, any person or entity, from knowingly and willfully offering, paying, soliciting or receiving any remuneration, directly or indirectly, overtly or covertly, in cash or in kind, to induce or in return for purchasing, leasing, ordering or arranging for the purchase, lease or order of any item or service reimbursable under Medicare, Medicaid or other federal healthcare programs. The term remuneration has been interpreted broadly to include anything of value. The Anti-Kickback Statute has been interpreted to apply to arrangements between pharmaceutical manufacturers on one hand and prescribers, purchasers, and formulary managers on the other. There are a number of statutory exceptions and regulatory safe harbors protecting some common activities from prosecution. The exceptions and safe harbors are drawn narrowly and practices that involve remuneration that may be alleged to be intended to induce prescribing, purchasing or recommending may be subject to scrutiny if they do not qualify for an exception or safe harbor. Failure to meet all of the requirements of a particular applicable statutory exception or regulatory safe harbor does not make the conduct per se illegal under the Anti-Kickback Statute. Instead, the legality of the arrangement will be evaluated on acase-by-case basis based on a cumulative review of all of its facts and circumstances. Our practices may not in all cases meet all of the criteria for protection under a statutory exception or regulatory safe harbor.

Additionally, the intent standard under the Anti-Kickback Statute was amended by the ACA to a stricter standard such that a person or entity no longer needs to have actual knowledge of the statute or specific intent to violate it in order to have committed a violation. In addition, the ACA codified case law that a claim including items or services resulting from a violation of the federal Anti-Kickback Statute constitutes a false or fraudulent claim for purposes of the federal False Claims Act (discussed below).

The civil monetary penalties statute imposes penalties against any person or entity who, among other things, is determined to have presented or caused to be presented a claim to a federal health program that the person knows or should know is for an item or service that was not provided as claimed or is false or fraudulent.

The federal False Claims Act prohibits, among other things, any person or entity from knowingly presenting, or causing to be presented, a false claim for payment to, or approval by, the federal government or knowingly making, using, or causing to be made or used a false record or statement material to a false or fraudulent claim to the federal government. As a result of a modification made by the Fraud Enforcement and Recovery Act of 2009, a claim includes “any request or demand” for money or property presented to the U.S. government. Recently, several pharmaceutical and other healthcare companies have been prosecuted under these laws for allegedly providing free product to customers with the expectation that the customers would bill federal programs for the product. Other companies have been prosecuted for causing false claims to be submitted because of the companies’ marketing of the product for unapproved, and thusnon-reimbursable, uses.

HIPAA created new federal criminal statutes that prohibit knowingly and willfully executing, or attempting to execute, a scheme to defraud or to obtain, by means of false or fraudulent pretenses, representations or promises, any money or property owned by, or under the control or custody of, any healthcare benefit program, including private third-party payors and knowingly and willfully falsifying, concealing or covering up by trick, scheme or device, a material fact or making any materially false, fictitious or fraudulent statement in connection with the delivery of or payment for healthcare benefits, items or services.

Also, many states have similar fraud and abuse statutes or regulations that apply to items and services reimbursed under Medicaid and other state programs, or, in several states, apply regardless of the payor.

We may also be subject to data privacy and security regulations by both the federal government and the states in which we conduct our business. HIPAA, as amended by the Health Information Technology for Economic and Clinical Health Act, or HITECH, and its implementing regulations, imposes requirements relating to the privacy, security and transmission of individually identifiable health information. Among other things, HITECH makes HIPAA’s privacy and security standards directly applicable to business associates independent contractors or agents of covered entities that receive or obtain protected health information in connection with providing a service on behalf of a covered entity. HITECH also created four new tiers of civil monetary

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penalties, amended HIPAA to make civil and criminal penalties directly applicable to business associates, and gave state attorneys general new authority to file civil actions for damages or injunctions in federal courts to enforce the federal HIPAA laws and seek attorneys’ fees and costs associated with pursuing federal civil actions.

In addition, state laws govern the privacy and security of health information in specified circumstances, many of which differ from each other in significant ways and may not have the same effect, thus complicating compliance efforts.

Additionally, the federal Physician Payments Sunshine Act, enacted as part of the ACA, and its implementing regulations, require certain manufacturers of drugs, devices, biological and medical supplies for which payment is available under Medicare, Medicaid or the Children’s Health Insurance Program (with certain exceptions) to report information related to certain payments or other transfers of value made or distributed to physicians and teaching hospitals, or to entities or individuals at the request of, or designated on behalf of, the physicians and teaching hospitals and to report annually certain ownership and investment interests held by physicians and their immediate family members.

In order to distribute products commercially, we will need to comply with state laws that require the registration of manufacturers and wholesale distributors of drug and biological products in a state, including, in certain states, manufacturers and distributors who ship products into the state even if such manufacturers or distributors have no place of business within the state. Some states also impose requirements on manufacturers and distributors to establish the pedigree of product in the chain of distribution, including some states that require manufacturers and others to adopt new technology capable of tracking and tracing product as it moves through the distribution chain. Several states have enacted legislation requiring pharmaceutical and biotechnology companies to establish marketing compliance programs, file periodic reports with the state, make periodic public disclosures on sales, marketing, pricing, clinical trials and other activities, and/or register their sales representatives, as well as to prohibit pharmacies and other healthcare entities from providing certain physician prescribing data to pharmaceutical and biotechnology companies for use in sales and marketing, and to prohibit certain other sales and marketing practices. All of our activities are also potentially subject to federal and state consumer protection and unfair competition laws.

If our operations are found to be in violation of any of the federal and state healthcare laws described above or any other governmental regulations that apply to us, we may be subject to penalties, including without limitation, civil, criminal and/or administrative penalties, damages, fines, disgorgement, exclusion from participation in government programs, such as Medicare and Medicaid, injunctions, private “qui tam” actions brought by individual whistleblowers in the name of the government, or refusal to allow us to enter into government contracts, contractual damages, reputational harm, administrative burdens, diminished profits and future earnings, and the curtailment or restructuring of our operations, any of which could adversely affect our ability to operate our business and our results of operations.

Coverage, Pricing and Reimbursement

Sales of our products will depend, in part, on the extent to which our products, if approved, will be covered and reimbursed by third-party payors, such as government health programs, commercial insurance and managed healthcare organizations. These third-party payors are increasingly reducing reimbursements for medical products and services. The process for determining whether a third-party payor will provide coverage for a drug product typically is separate from the process for setting the price of a drug product or for establishing the reimbursement rate that a payor will pay for the drug product once coverage is approved. Third-party payors may limit coverage to specific drug products on an approved list, also known as a formulary, which might not include all of the approved drugs for a particular indication.

In order to secure coverage and reimbursement for any product candidate that might be approved for sale, we may need to conduct expensive pharmacoeconomic studies in order to demonstrate the medical necessity and

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cost-effectiveness of the product candidate, in addition to the costs required to obtain FDA or other comparable regulatory approvals. Whether or not we conduct such studies, our product candidates may not be considered medically necessary or cost-effective. A third-party payor’s decision to provide coverage for a drug product does not imply that an adequate reimbursement rate will be approved. Further, one payor’s determination to provide coverage for a product does not assure that other payors will also provide coverage for the product. Even if coverage is obtained from third party payors, reimbursement may not be sufficient to enable us to maintain price levels high enough to realize an appropriate return on our investment in product development.

The containment of healthcare costs has become a priority of federal, state and foreign governments, and the prices of drugs have been a focus in this effort. The U.S. government, state legislatures and foreign governments have shown significant interest in implementing cost-containment programs, including price controls, restrictions on reimbursement and requirements for substitution of generic products. Similar policies and laws have been adopted by many EU Member States. Adoption of price controls and cost-containment measures, and adoption of more restrictive policies in jurisdictions with existing controls and measures, could further limit our net revenue and results. Decreases in third-party reimbursement for our product candidate or a decision by a third-party payor to not cover our product candidate could reduce physician usage of the product candidate and have a material adverse effect on our sales, results of operations and financial condition.

In addition, in some foreign countries, the proposed pricing for a drug must be approved before it may be lawfully marketed. The requirements governing drug pricing vary widely from country to country. For example, the EU provides options for its Member States to restrict the range of medicinal products for which their national health insurance systems provide reimbursement and to control the prices of medicinal products for human use. A Member State may approve a specific price for the medicinal product or it may instead adopt a system of direct or indirect controls on the profitability of the company placing the medicinal product on the market. For example, in France, effective access to the market assumes that our future products will be supported by the hospital (through an agreement for local communities) or reimbursed by social security. The price of medications is negotiated with the Economic Committee for Health Products, or CEPS. There can be no assurance that any country that has price controls or reimbursement limitations for pharmaceutical products will allow favorable reimbursement and pricing arrangements for any of our product candidates. Historically, products launched in the EU do not follow price structures of the United States and generally tend to be significantly lower.

Healthcare Reform and Subsequent Legislation

In March 2010, President Obama enacted the ACA, which has the potential to substantially change healthcare financing and delivery by both governmental and private insurers, and significantly impact the pharmaceutical and biotechnology industry. The ACA will impact existing government healthcare programs and will result in the development of new programs.

Among the ACA’s provisions of importance to the pharmaceutical and biotechnology industries, in addition to those otherwise described above, are the following:

an annual, nondeductible fee on any entity that manufactures or imports certain specified branded prescription drugs and biologic agents apportioned among these entities according to their market share in some government healthcare programs;

an increase in the statutory minimum rebates a manufacturer must pay under the Medicaid Drug Rebate Program to 23.1% and 13% of the average manufacturer price for most branded and generic drugs, respectively and a cap on the total rebate amount for innovator drugs at 100% of the Average Manufacturer Price, or AMP;

a new Medicare Part D coverage gap discount program, in which manufacturers must agree to offer 50%point-of-sale discounts off negotiated prices of applicable brand drugs to eligible beneficiaries during their coverage gap period, as a condition for the manufacturers’ outpatient drugs to be covered under Medicare Part D;

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extension of manufacturers’ Medicaid rebate liability to covered drugs dispensed to individuals who are enrolled in Medicaid managed care organizations;

expansion of eligibility criteria for Medicaid programs by, among other things, allowing states to offer Medicaid coverage to additional individuals and by adding new mandatory eligibility categories for individuals with income at or below 133% of the federal poverty level, thereby potentially increasing manufacturers’ Medicaid rebate liability;

expansion of the entities eligible for discounts under the Public Health Service pharmaceutical pricing program; and

a new Patient-Centered Outcomes Research Institute to oversee, identify priorities in, and conduct comparative clinical effectiveness research, along with funding for such research.

We anticipate that, absent to further legislative changes, the ACA will result in additional downward pressure on coverage and the price that we receive for any approved product in the United States, and could seriously harm our business. Any reduction in reimbursement from Medicare and other government programs may result in a similar reduction in payments from private payors. The implementation of cost containment measures or other healthcare reforms may prevent us from being able to generate revenue, attain profitability, or commercialize our product candidates, if approved. In addition, it is possible that there will be further legislation or regulation that could change parts of the ACA that affect public and private healthcare coverage. Those changes could harm our business, financial condition, and results of operations.

Other legislative changes have been proposed and adopted in the United States since the ACA was enacted. On August 2, 2011, the Budget Control Act of 2011 among other things, created measures for spending reductions by Congress. A Joint Select Committee on Deficit Reduction, tasked with recommending a targeted deficit reduction of at least $1.2 trillion for the years 2013 through 2021, was unable to reach required goals, thereby triggering the legislation’s automatic reduction to several government programs. This includes aggregate reductions to Medicare payments to providers of 2% per fiscal year, which started in April 2013. On January 2, 2013, President Obama signed into law the American Taxpayer Relief Act of 2012, or the ATRA, which, among other things, also reduced Medicare payments to several providers, including hospitals, imaging centers and cancer treatment centers, and increased the statute of limitations period for the government to recover overpayments to providers from three to five years. Further, in January 2017, President Trump signed an Executive Order directing federal agencies with authorities and responsibilities under the ACA to waive, defer, grant exemptions from, or delay the implementation of any provisions of the ACA that would impose a fiscal or regulatory burden on states, individuals, healthcare providers, health insurers, or manufacturer of pharmaceutical products. Congress may also consider subsequent legislation to replace elements of the ACA that are repealed. As a result, the full impact of the ACA, any law repealing and/or replacing elements of it, and the political uncertainty surrounding any repeal or replacement legislation remains unclear.

Additional Regulation

In addition to the foregoing, state and federal laws regarding environmental protection and hazardous substances, including the Occupational Safety and Health Act, the Resource Conservancy and Recovery Act and the Toxic Substances Control Act, affect our business. These and other laws govern our use, handling and disposal of various biological, chemical and radioactive substances used in, and wastes generated by, our operations. If our operations result in contamination of the environment or expose individuals to hazardous substances, we could be liable for damages and governmental fines. We believe that we are in material compliance with applicable environmental laws and that continued compliance therewith will not have a material adverse effect on our business. We cannot predict, however, how changes in these laws may affect our future operations.

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European Union Drug Development

In the EU, our future product candidates may also be subject to extensive regulatory requirements. As in the United States, medicinal products can only be marketed if a marketing authorization from the competent regulatory agencies has been obtained.

Similar to the United States, the various phases ofpre-clinical and clinical research in the European Union are subject to significant regulatory controls. Although the EU Clinical Trials Directive 2001/20/EC has sought to harmonize the EU clinical trials regulatory framework, setting out common rules for the control and authorization of clinical trials in the EU, the EU Member States have transposed and applied the provisions of the Directive differently. This has led to significant variations in the Member State regimes. To improve the current system, a new Regulation No. 536/2014 on clinical trials on medicinal product candidates for human use, which will repeal Directive 2001/20/EC, was adopted on April 16, 2014 and published in the European Official Journal on May 27, 2014. The new Regulation aims at harmonizing and streamlining the clinical trials authorization process, simplifying adverse event reporting procedures, improving the supervision of clinical trials, and increasing their transparency. The new Regulation entered into force on June 16, 2014 and was set to apply not earlier than May 28, 2016. Based on announcements of the European Medicines Agency, it is now expected that the mentioned Regulation will not enter into force before ~~October 2018.~~2020. Until then the Clinical Trials Directive 2001/20/EC will continue to apply. In addition, the transitional provisions of the new Regulation offer, under certain conditions, the clinical trial sponsors the possibility to choose between the requirements of the Directive and the Regulation for a limited amount of time.

Under the current regime, before a clinical trial can be initiated it must be approved in each of the EU Member States where the trial is to be conducted by two distinct sets of bodies: the National Competent Authority, or NCA, and one or more Ethics Committees, or ECs. Under the current regime all suspected unexpected serious adverse reactions, or SUSARs, to the investigational product that occur during the clinical trial have to be reported to the NCA and ECs of the Member State where they occurred as well as in the European safety database, EudraVigilance.

~~European Union Drug Review and Approval~~

In the EU, medicinal products can only be commercialized after obtaining a Marketing Authorization, or MA. The same rules also apply in the EFTA Member States (Norway, Iceland and Liechtenstein). There are two types of marketing authorizations:

The Community MA, which is issued by the European Commission through the Centralized Procedure, based on the opinion of the Committee for Medicinal Products for Human Use, or CHMP, of the European Medicines Agency, or EMA, and which is valid throughout the entire territory of the EU. The Centralized Procedure is mandatory for certain types of products, such as biotechnology medicinal products, orphan medicinal products, and medicinal products containing a new active substance indicated for the treatment of AIDS, cancer, neurodegenerative disorders, diabetes, auto-immune and viral diseases. The Centralized Procedure is also mandatory forso-called Advance Therapy Medicinal Products (or ATMPs). ATMPs comprise gene therapy, somatic cell and tissue engineered products. In this regard, on May 28, 2014, the EMA issued a recommendation that Cellectis’ UCART19 be considered a gene therapy product under Regulation (EC) No 1394/2007 on ATMPs. The Centralized Procedure is optional for products containing a new active substance not yet authorized in the EEA, or for products that constitute a significant therapeutic, scientific or technical innovation or which are in the interest of public health in the EU. Under the Centralized Procedure, the CHMP serves as the scientific committee that renders opinions about the safety, efficacy and quality of human products on behalf of the EMA. The CHMP is composed of experts nominated by each Member State’s national drug authority, with one of them appointed to act as Rapporteur for theco-ordination of the evaluation with the possible assistance of a further member of the Committee acting as aCo-Rapporteur. The CHMP has 210 days to adopt an opinion as to whether a marketing authorization should be granted. The process usually takes longer as

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additional information is requested, which triggers clock-stops in the procedural timelines. Based on the CHMP’s opinion the European Commission will adopt a decision on the granting of the marketing authorization. In case of ATMPs, the EMA’s Committee for Advanced Therapies, a multidisciplinary committee of experts on ATMPs, will prepare a draft opinion which will be submitted to the CHMP before the latter adopts its final opinion.

Under the above-described procedure, before granting the MA, the EMA makes an assessment of the risk-benefit balance of the product on the basis of scientific criteria concerning its quality, safety and efficacy.

In the EU, new products authorized for marketing (i.e., reference products) qualify for eight years of data exclusivity and an additional two years of market exclusivity upon marketing authorization. The data exclusivity period prevents generic applicants from relying on thepre-clinical and clinical trial data contained in the dossier of the reference product when applying for a generic marketing authorization in the EU during a period of eight years from the date on which the reference product was first authorized in the EU. The market exclusivity period prevents a successful generic applicant from commercializing its product in the EU until ten years have elapsed from the initial authorization of the reference product in the EU. Theten-year market exclusivity period can be extended to a maximum of eleven years if, during the first eight years of those ten years, the marketing authorization holder obtains an authorization for one or more new therapeutic indications which, during the scientific evaluation prior to their authorization, are held to bring a significant clinical benefit in comparison with existing therapies.

The EU also provides opportunities for market exclusivity. For example, products receiving orphan designation in the EU can receive ten years of market exclusivity, during which time no similar medicinal product for the same indication may be placed on the market. An orphan product can also obtain an additional two years of market exclusivity in the EU for pediatric studies. No extension to any supplementary protection certificate can be granted on the basis of pediatric studies for orphan indications.

The criteria for designating an “orphan medicinal product” in the EU are similar in principle to those in the United States. Under Article 3 of Regulation (EC) 141/2000, a medicinal product may be designated as orphan if (1) it is intended for the diagnosis, prevention or treatment of a life-threatening or chronically debilitating condition; (2) either (a) such condition affects no more than five in 10,000 persons in the EU when the application is made, or (b) the product, without the benefits derived from orphan status, would not generate sufficient return in the EU to justify investment; and (3) there exists no satisfactory method of diagnosis, prevention or treatment of such condition authorized for marketing in the EU, or if such a method exists, the product will be of significant benefit to those affected by the condition, as defined in Regulation (EC) 847/2000. Orphan medicinal products are eligible for financial incentives such as reduction of fees or fee waivers and are, upon grant of a marketing authorization, entitled to ten years of market exclusivity for the approved therapeutic indication. The application for orphan drug designation must be submitted before the application for marketing authorization. The applicant will receive a fee reduction for the marketing authorization application if the orphan drug designation has been granted, but not if the designation is still pending at the time the marketing authorization is submitted. Orphan drug designation does not convey any advantage in, or shorten the duration of, the regulatory review and approval process.

The10-year market exclusivity may be reduced to six years if, at the end of the fifth year, it is established that the product no longer meets the criteria for orphan designation, for example, if the product is sufficiently profitable not to justify maintenance

of market exclusivity. Additionally, marketing authorization may be granted to a similar product for the same indication at any time if:

The second applicant can establish that its product, although similar, is safer, more effective or otherwise clinically superior;

The applicant consents to a second orphan medicinal product application; or

The applicant cannot supply enough orphan medicinal product.

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Food Oversight Responsibilities Between USDA and FDA, Including Transgenic or Genetically Modified Organisms

In the United States, the FDA and the USDA Food Safety Inspection Service, or FSIS, are primarily responsible for overseeing food regulation and safety, although as many as fifteen federal agencies also play a role in U.S. food regulation, including several other agencies within USDA.

FSIS is responsible for ensuring the safety, wholesomeness, and correct packaging and labeling of the nation’s commercial supply of meat, poultry, egg products, and catfish. The agency’s main authorizing statutes are the Poultry Products Inspection Act, Federal Meat Inspection Act, Agricultural Marketing Act, and the Egg Products Inspection Act. To carry out its mission, FSIS deploys almost 8,000 inspection program personnel to the more than 6,000 establishments around the country to ensure food manufactures are following the proper procedures to reduce the risks of food borne illnesses such asSalmonella, Escherichia coli, Listeria monocytogenes, andCampylobacter.

USDA has regulatory jurisdiction over transgenic crops through the Animal and Plant Health Inspection Service, or APHIS. Under the Plant Protection Act, USDA requires anyone who wishes to import, transport interstate, or plant a “regulated article” to apply for a permit or notify APHIS that the introduction will be made. Regulated articles are defined as “any organism which has been altered or produced through genetic engineering [...] which USDA determines is a plant pest or has reason to believe is a plant pest.” The petition process can be a multi-year process that varies based on a number of factors, including APHIS’ familiarity with similar products, the type and scope of the environmental review conducted, and the number and types of public comments received. APHIS conducts a comprehensive science-based review of the petition to assess, among other things, plant pest risk, environmental considerations pursuant to the National Environmental Policy Act of 1969, or NEPA, and any potential impact on endangered species. If, upon the completion of the review, APHIS grants the petition, the product is no longer deemed a “regulated article” and the petitioner may commercialize the product, subject to any conditions set forth in the decision. If APHIS does not determine the product to benon-regulated, the product may be subject to extensive regulation, including permitting requirements for import, handling, interstate movement, and release into the environment, and inspections.

Calyxt has submitted petition to APHIS for ~~six~~seven of ~~our~~its product candidates to date: ~~high oleic soybeans, high oleic/~~High Fiber Wheat, High Oleic Soybean, High Oleic/low ~~linoleic soybeans, cold storable potatoes, reduced browning potatoes, powdery mildew resistant wheat,~~Linolenic Soybean, Cold Storable Potato, Reduced Browning Potato, Powdery Mildew Resistant Wheat, and ~~improved quality alfalfa.~~Improved Quality Alfalfa. Calyxt has received confirmation from APHIS for all ~~six~~seven product candidates that APHIS does not consider such product candidate to be a “regulated article” under the Plant Protection Act. There can be no guarantee of the timing or success in obtaining nonregulated status from APHIS for other crops developed by Calyxt or that the governing regulations will not change. Government regulations, regulatory systems, and the policies that influence them vary widely among jurisdictions and change often.

As part of its National Organic Program, USDA also regulates GMOs, or genetically modified (“GM”) foods, to the extent that food manufacturers can use the “USDA Organic” label on their products. The use of genetic engineering, or GMOs, is prohibited in USDA organic products. According to USDA, this means, for example, that an organic farmer cannot plant GMO seeds, an organic cow cannot eat GMO alfalfa or corn, and an organic soup producer cannot put any GM ingredients into its soup. To label products with the USDA organic seal, farmers and food processors must show they are not using GMOs and that they are protecting their products from contact with GMOs (along with other prohibited substances) from farm to table.

FDA has jurisdiction to regulate more than 80 percent of the U.S. food supply. It derives its regulatory power from the Federal Food, Drug, and Cosmetic Act (“FDCA”), which has been amended over time by several subsequent laws. FDA’s oversight of food safety and security is primarily carried out by its Center for Food Safety and Applied Nutrition (“CFSAN”). To execute its responsibilities, FDA has a team of 900 investigators and 450 analysts in the foods program who conduct inspections and collect and analyze product samples. FDA typically does not performpre-market inspection for foods. FDA also regulates ingredients, packaging, and

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labeling of foods, including nutrition and health claims and the nutrition facts panel. Foods are typically not subject to premarket review and approval requirements, with limited exceptions.

For its part, FDA regulates foods made with GMOs under its 1992 “Statement of Policy: Foods Derived from New Plant Varieties.” Under this policy, FDA regulates foods derived from GM plant varieties consistent with the framework fornon-GM foods. In most cases, foods derived from GM plant varieties are not subject to premarket review and approval. In some cases,

however, such foods will be considered to contain “food additives” that require premarket review and approval. FDA offers a voluntary consultation process to determine whether foods derived from GM plant varieties will be subject to these more stringent regulatory requirements.

FDA does not currently require manufacturers to label foods made with GMOs as such, but permits voluntary labeling pursuant to a guidance document finalized in November 2015. The topic of GMO use and labeling has been of significant public interest; as political forces continue to work, and there can be no guaranty that it will not change in the future. Additionally, three states have passed, and nearly half of U.S. states have considered, mandatory GMO labeling laws to date.

FDA is currently evaluating its approach to the regulation of gene-edited plants. The FDA’s thinking on the use of genome editing techniques to produce new plant varieties that are used for human or animal food continues to evolve. To that end, in January 2017 FDA announced a Request for Comments (“RFC”) seeking public input to help inform its thinking about human and animal foods derived from new plant varieties produced using genome editing techniques. Among other things, the RFC asks for data and information in response to questions about the safety of foods from gene-edited plants, such as whether categories of gene-edited plants present food safety risks different from other plants produced through traditional plant breeding. If FDA enacts new regulations or policies with respect to gene-edited plants, such policies could result in additional compliance costs and/or delay the commercialization of Calyxt’s product candidates.

EU Regulation of GMOs and Genetically Modified Food and Feed Products

In the EU, genetically modified organisms (“GMOs”) and genetically modified food and feed products can only be sold in the market once they have been properly authorized. The procedures for evaluation and authorization of GMOs and genetically modified food and feed products are established by Regulation (EC) 1829/2003 on genetically modified food and feed (‘‘Regulation (EC) 1829/2003’’) and Directive 2001/18/EC on the release of GMOs into the environment (‘‘Directive 2001/18/EC’’). An application for authorization must be submitted under Directive 2001/18/EC if a company seeks to release GMOs for experimental purposes (e.g., field tests) and/or to sell GMOs, as such or in products, in the market (e.g., cultivation, importation or processing). In turn, an application for authorization must be submitted under Regulation (EC) 1829/2003 if a company seeks to sell GMOs in the market for food and feed use and/or food and feed products containing or produced from GMOs. At the national level, EU member states have the ability to restrict or prohibit GMO cultivation in their territories by invoking grounds such as environmental or agricultural policy objectives, town and country-planning, land use, coexistence, socio-economic impacts or public policy.

In addition, Directive 2001/18/EC, Regulation (EC) 1829/2003 and Regulation (EC) 1830/2003 establish specific labeling and traceability requirements for GMOs and products that contain or are produced from GMOs. Finally, Directives 2002/53/EC and 2002/55/EC require genetically modified varieties to be authorized in accordance with Directive 2001/18/EC and/or Regulation (EC) 1829/2003, as applicable, before they can be included in a ‘‘Common Catalogue of Varieties,’’ which would permit the seeds of such genetically modified varieties to be marketed in the EU.

A recent ruling of the European Court of Justice (‘‘ECJ’’) in July 2018 concluded that organisms obtained by new mutagenesis plant breeding techniques involving the use of genetic engineering are GMOs and therefore fall, in principle, under Directive 2001/18/EC described above. The ECJ found further that varieties obtained by modern forms of mutagenesis are genetically modified varieties covered by Directive 2002/53/EC, and are therefore subject to the obligations of such directive. The ECJ clarified that only mutagenesis techniques which (a) have been used in a number of applications and (b) have a long safety record, can be exempted from these requirements, although EU member states remain free to subject even such exempted organisms to the obligations under Directive 2001/18/EC, or to other obligations.

Other Regulatory Matters

French Pharmaceutical Company Status

To date, we do not have the status of pharmaceutical establishment, and therefore, cannot either manufacture the product candidates we develop or directly consider their marketing. Obtaining the pharmaceutical establishment license, either as distributor, operator, importer or as manufacturer, requires the submission of a request file specific to each of the mentioned qualifications with theAgence nationale de sécurité du médicament et des produits de santé (ANSM), which only grants it after review of this file and evaluation, usually after verification that the company has adequate premises, the necessary personnel and an adapted structure with satisfactory procedures for carrying out the proposed pharmaceutical activities.

We currently entrust CMOs with the manufacturing of clinical batches and intend to continue relying on CMOs for the production of the first commercial batches. We may consider internalizing production once our first product candidate is approved by regulatory authorities.

Legal Proceedings

From time to time, we may be involved in various claims and legal proceedings relating to claims arising out of our operations. We are not currently a party to any legal proceedings that, in the opinion of our management, are likely to have a material adverse effect on our business. Regardless of outcome, litigation can have an adverse impact on us because of defense and settlement costs, diversion of management resources and other factors.

C.	Organizational Structure

In 2015, Cellectis S.A. had two French subsidiaries (Cellectis Bioresearch and Ectycell) and three U.S. subsidiaries (Calyxt, Inc., Cellectis, Inc. and Cellectis Bioresearch Inc.).Non-controlling shareholders held a 24.5% interest in Cellectis Bioresearch, Cellectis Bioresearch Inc. and Ectycell until May 18, 2015.

The following internal reorganization was completed in 2015:

Ectycell was merged into, and absorbed by Cellectis Bioresearch in August 2015 with retroactive effect as at January 1, 2015 for French tax purposes;

Cellectis Bioresearch was merged into, and absorbed by, Cellectis ~~S.A~~S.A. in December 2015 with retroactive effect as at January 1, 2015 for French tax purposes;

Cellectis Bioresearch Inc. was merged into Cellectis Inc. in September 2015.

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Following the reorganization and as of December 31, 2016, the two remaining subsidiaries, Cellectis, Inc. and Calyxt, Inc., were wholly-owned by Cellectis.

Until July 25, 2017, Cellectis S.A. fully owned Calyxt, Inc. ~~was a wholly-owned subsidiary of Cellectis.~~ On July 25, 2017, Calyxt closed its IPO with $64.4 million in gross proceeds to Calyxt from the sale of 8,050,000 shares at $8 per share, including the full exercise of the underwriter’s over-allotment option ~~with $64.4 million in gross proceeds. Cellectis purchased~~and Cellectis’ purchase of $20.0 million of shares in the ~~course~~IPO. On May 22, 2018, Calyxt, Inc completed afollow-on offering of its common stock. Calyxt, Inc. sold an aggregate of 4,057,500 shares of common stock at a price of $15.00 per share, including 457,500 shares of common stock pursuant to the exercise of the ~~IPO,~~underwriters’ option to purchase additional shares. In the aggregate, Calyxt, Inc. received net proceeds from thefollow-on offering and exercise of the overallotment option of approximately $57.0 million, after deducting underwriting discounts and commissions of $3.2 million and offering expenses totaling approximately $0.7 million. As part of thefollow-on offering, Cellectis SA purchased 550,000 shares of common stock for a value of $8.3 million, the proceeds of which isare included in the net proceeds ~~that Calyxt received.~~of approximately $57.0 million.

As of December 31, 2018, Cellectis S.A. owns 100% of Cellectis, Inc. and approximately 69.5% of Calyxt’s outstanding shares of common stock. As of December 31, 2017, Cellectis S.A. owns 100% of Cellectis, Inc. and approximately 79.7% of Calyxt’s outstanding shares of common stock.

D.	Property, ~~Plants~~Plant and Equipment

Cellectis S.A. leases a ~~3,064~~3,820 square-meter facility in Paris for administrative and research and development activities. The lease commenced on April 1, 2011 and ~~has~~was amended on December 1, 2018 with a~~10-year~~9-year initial term expiring on ~~April 1, 2021.~~November 30, 2027.

Cellectis, Inc. leases a ~~12,052~~15,532 square-foot facility in New York, New York for administrative and research and development activities, which commenced on March 30, 2015 and ~~has~~was amended on December 1, 2018 with a~~66-month~~60-month initial term expiring on ~~September 30, 2020.~~December 31, 2023. In addition, in March 2016, Cellectis Inc. entered into a lease agreement for a 26,928 square-foot facility in Montvale, New Jersey. As of December 31, ~~2017,~~2018, Montvale facility is not operational and we ~~intend~~have the willingness to discontinue this lease before its ~~scheduled~~ termination (September 2026).

In March 2019, we entered into a lease agreement for a 82,000 square foot commercial-scale manufacturing facility, called the IMPACT site, which stands for “Innovative Manufacturing Plant for Allogeneic Cellular Therapies”. The IMPACT facility is located in ~~September 2026.~~Raleigh, North Carolina. The new manufacturing facility is being designed to provide GMP manufacturing for clinical supply and commercial product upon potential regulatory approval. The facility is planned to be operational by 2021.

Calyxt, Inc. ~~leases a 17,485 square-foot facility in New Brighton, Minnesota, which commenced operations on October 15, 2012. The lease for this facility expires on May 31, 2018.~~

~~In March 2016, Calyxt acquired10-acres~~ ~~in 2 parcels in Roseville (MN). A headhouse and a greenhouse were built on the Roseville property and have been in operation since September 2016.~~

~~In September 2017, Calyxt consummated~~entered into a sale-leaseback transaction ~~including a Lease Agreement dated~~on September 6, 2017 with a third party ~~with respect to Calyxt’s lease of certain real property~~for its corporate headquarters and ~~improvements located~~lab facility in Roseville, ~~Minnesota, for a~~Minnesota. Calyxt, Inc. committed to an initial lease term of twenty years, with four options to extend the term ~~for up to an additional twenty years. Pursuant to the purchase agreement, Calyxt received approximately $7 million in connection with the sale~~ of ~~the property. The property will be the new corporate headquarters for Calyxt and the landlord is constructing abuilt-to-suit~~ ~~facility to be composed of a nearly 40,000 square-foot office and lab building, with greenhouses and outdoor research plots. The corporate headquarters are expected to be ready in the spring of 2018.~~

~~Under~~ the Lease Agreement for five years each. The transaction also included a construction contract for Calyxt, ~~will~~Inc. 40,000 square-foot corporate headquarters which includes office, research laboratory space and outdoor growing plots. During the construction period, which ended in June 2018, Calyxt, Inc. initially ~~pay~~paid annual base rent of ~~$490,000~~$490 thousand until the ~~earlier of (i) the next day after issuance of a temporary certificate of occupancy or other permit to occupy the~~ property by the City of Roseville and (ii) the next day after the certification of substantial completion executed by landlord’s architect or contractor confirming that the work to be done on the property has beenwas substantially completed ~~(such date,~~in May, at which time, the ~~“Initial Term Commencement Date”). On~~lease commenced. Under the ~~Initial Term Commencement Date,~~lease, Calyxt, ~~will pay~~Inc. now pays an estimated annual base rent of 8% of the total project cost (“Annual Base Rent”) with scheduled increases in rent of 7.5% on the sixth, eleventh and sixteenth anniversaries of the initial term commencement date as well as on the first day of each Renewal Term (as defined in the Lease Agreement). Based on the initial cost of the project Calyxt will pay an estimated annual base rent of approximately ~~$1,480 thousand.~~

~~The Lease Agreement~~$1.4 million. Calyxt, Inc. is ~~a net lease and~~responsible for the other costs and expenses associated with the ~~property are to be paid for by Calyxt. In consideration~~use of ~~and as an inducement to,~~ the ~~landlord’s agreement to enter into the Lease Agreement,~~property. Cellectis entered into a Lease Guaranty with the landlord for the facilities, whereby Cellectis has guaranteed ~~all of Calyxt’s~~Calyxt, Inc.’s obligations under the Lease Agreement. Cellectis’ guarantee of Calyxt’s obligations under the sale-leaseback transaction will terminate at the end of the second consecutive calendar year in which ~~Calyxt’s~~Calyxt, Inc.’s tangible net worth exceeds $300 million, as determined in accordance with generally accepted accounting principles.

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~~On November 10, 2017,~~ Calyxt, Inc. agreed to indemnify Cellectis for any obligations incurred by Cellectis under the Lease Guaranty. This indemnification agreement ~~will become effective at such time as~~takes effect when Cellectis owns 50% or less of ~~Calyxt’s~~Calyxt, Inc.’s outstanding common stock.

In December 2018 Calyxt, Inc. consummated a sale-leaseback transaction with a third party to finance equipment. The lease has a term of four years and Calyxt, Inc. may add up to $1.1 million of future purchases to the financing agreement. Calyxt, Inc. was required to deposit cash into a restricted account in an amount equal to the future rent payments required by the lease.

ITEM 4A.

UNRESOLVED STAFF COMMENTS

Not applicable.

ITEM 5.

OPERATING AND FINANCIAL REVIEW AND PROSPECTS

The following Operating and Financial Review and Prospects should be read in conjunction with our audited consolidated financial statements and related notes included elsewhere in this Annual Report. In addition to historical consolidated financial information, this discussion also contains forward-looking statements, based on current expectations and related to future events and our future financial performance, that involve risks and uncertainties. Our actual results may differ materially from those anticipated in these forward-looking statements as a result of certain factors, including but not limited to those set forth under “Risk Factors” and elsewhere in this Annual Report.

Overview

We are a clinical stage biotechnological company, employing our core proprietary technologies to developbest-in-class products in the ~~emerging~~ field of immuno-oncology. Our product candidates, based on gene-editedT-cells that express chimeric antigen receptors, or CARs, seek to harness the power of the immune system to target and eradicate cancers. We believe thatCAR-based immunotherapy is one of the most promising areas of cancer research, representing a new paradigm for cancer treatment. We are designing next-generation immunotherapies that are based on gene-edited CART-cells. Our gene-editing technologies allow us to create allogeneic CART-cells, meaning they are derived from healthy donors rather than the patients themselves. We believe that the allogeneic production of CART-cells will allow us to develop cost-effective,“off-the-shelf” products and are capable of being stored and distributed worldwide. Our gene-editing expertise also enables us to develop product candidates that feature additional safety and efficacy attributes, including control properties designed to prevent them from attacking healthy tissues, to enable them to tolerate standard oncology treatments, and to equip them to resist mechanisms that inhibit immune-system activity. In addition to our focus on immuno-oncology, we are exploring the use of our gene-editing technologies in other therapeutic applications, as well as to develop healthier food products for a growing population.

We currently conduct our operations through two business segments, Therapeutics and Plants. Our Therapeutics segment is mainly focused on the development of products in the field of immuno-oncology. Our Plants segment focuses on applying our gene-editing technologies to develop new generation plant products in the field of agricultural biotechnology through its own efforts or through alliances with other companies in the agricultural market.

Since our inception in early 2000, we have devoted substantially all of our financial resources to research and development efforts. Our current research and development focuses primarily on our CART-cell immunotherapy product candidates, including preparing to conduct clinical studies of our product candidates, providing general and administrative support for these operations and protecting our intellectual property. In addition, by leveraging our plant-engineering platform and the transformative potential of gene editing, we aim to create food products with consumer health benefits, adaptations for climate change or nutritional enhancements that address the needs of a growing population. We do not have any products approved for sale and have not generated any revenues from immunotherapy or agricultural biotechnology product sales.

In February 2014, we entered into an alliance with Servier for the development of UCART19 and other product candidates directed at four additional molecular targets. In November 2015, we entered into an

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amendment to our initial collaboration agreement with Servier, which allowed for an early exercise of Servier’s option with respect to UCART19 and other product candidates. Pursuant to this amendment, Servier has exercised its option to acquire the exclusive worldwide rights to further develop and commercialize UCART19. In addition, Pfizer and Servier have entered into an exclusive global license and collaboration agreement under which Pfizer has obtained from Servier exclusive rights to develop and commercialize UCART19 in the United States. We entered into amendments to our collaboration agreements with each of Servier and Pfizer to facilitate this agreement between Servier and Pfizer. In connection with the entry into the amendment to the collaboration agreement, Servier made an upfront payment of $38.5 million, excluding taxes. As of December 31, ~~2017,~~2018, Cellectis was eligible to receive up to ~~$1,064~~$932.8 million in potential option exercise fees, development, clinical and sales milestones, in addition to royalties on sales and research and development costs reimbursements.

On March 6, 2019, we and Servier entered into the Servier License Agreement, which supersedes the prior collaboration agreement in order to modify the targets covered by the Servier License Agreement, to establishes the terms of our and Servier’s collaboration and to reflect the status of products in development.

Our alliance with Pfizer, which commenced in June 2014, ~~addresses~~addressed the development of other CART-cell immunotherapies in the field of oncology. Pfizer also purchased 10% of our then-outstanding equity in connection with this collaboration for €25.8 million. On April 3, 2018, Pfizer announced that it had assigned the agreement they had signed with Cellectis on June 18, 2014 to a newly formed biotechnology company named Allogene.

On March 7, 2019, we and Allogene agreed to terminate the prior collaboration agreement and entered into a new license agreement to reflect the relationship between us and Allogene.

This strategic alliance with Allogene is potentially worth up to $2.9 billion in payments by ~~Pfizer~~Allogene to us, including an $80 million upfront payment and $2.8 billion in potential clinical and commercial milestone payments. In addition, we invoice research and development costs assigned to our projects in common with ~~Pfizer. Pfizer also purchased 10% of our then-outstanding equity in connection with this collaboration for €25.8 million.~~ Allogene.

We believe that ~~both of these~~our strategic transactions with Allogene and Servier position us to compete in the promising field of immuno-oncology and add additional clinical and financial resources to our programs.

We have also entered into research and development alliances with each of Cornell University and the MD Anderson Cancer Center. Pursuant to these strategic alliances, we collaborate with these two centers to accelerate the development of our lead product candidates UCART123, UCARTCS1 ~~UCART22~~ and ~~UCART38~~UCART22 in AML, BPDCN, multiple myeloma,~~B-cell~~ and~~T-ALL,~~B-ALL, respectively. Under these agreements, we fund the research activities performed at Cornell University and the MD Anderson Cancer Center. In August 2018, we entered into a new clinical study agreement with Dana Farber Cancer Institute and H. Lee Moffitt Cancer Center in order to expand the performance of the UCART123 clinical study in AML to Dana Farber Cancer Institute and H. Lee Moffitt Cancer Center.

Our cash consumption is driven by our internal operational activities, as well as our outsourced activities, including the preclinical activities and the manufacturing activities of the requisite raw materials for the manufacturing of UCART123, UCART22 and ~~UCART22,~~ UCARTCS1, the technology transfer ~~of UCART22~~to CELLforCURE and ~~UCARTCS1 process to CELLforCURE,~~MolMed, and the GMP manufacturing of UCART123, UCART22 and ~~UCART22~~UCARTCS1 at CELLforCURE and MolMed. In addition, we incurred significant annual payment and royalty expenses related to ourin-licensing agreements with different parties including Institut Pasteur, LifeTechnologies and University of Minnesota. In addition, in 2017 and 2018, we initiated ~~our~~ clinical studies at Weill Cornell and the MD Anderson Cancer Center, leading to additional cash burn through payments to the clinical research centers, the Contract Research Organisation involved and the companies involved in the logistics and testing of the clinical sample material.

In addition to our cash generated by operations (including payments under our strategic alliances), we have funded our operations primarily through private and public offerings of our equity securities, grant revenues, payments received under intellectual property licenses, and reimbursements of research tax credits. Our ordinary shares have traded on the Euronext Growth market of Euronext in Paris since February 7, 2007. In March 2015, we completed our U.S. initial public offering of 5,500,000 American Depositary Shares on the Nasdaq Global Market for gross proceeds of $228.2 million. On April 10, 2018, Cellectis closed afollow-on offering of 5,646,000 ADS at a public offering price of $31.00 per ADS resulting in gross proceeds of $175 million. On May 11, 2018, in connection with the exercise by the underwriters of thefollow-on offering of their option to purchase additional shares, Cellectis closed the sale of an additional 500,000 ADS at the public offering price of $31.00 per ADS resulting in additional gross proceeds of $15.5 million.

On July 25, 2017, Calyxt completed an initial public offering of its common stock on the Nasdaq, selling an aggregate of 8,050,000 shares of common stock at a price of $8.00 per share (including 1,050,000 shares of common stock pursuant to the exercise by the underwriters of their option to purchase additional shares). Calyxt received net proceeds of approximately $58.0 million, after deducting underwriting discounts and commissions and offering expenses. As part of the Calyxt IPO, Cellectis purchased 2,500,000 shares of common stock for a value of $20.0 million, which is included in the net proceeds that Calyxt received. On May 22, 2018, Calyxt closed afollow-on offering with $60.9 million in gross proceeds, inclusive of $8.25 million from Cellectis’ purchase of shares in thefollow-on offering

In ~~2015,~~ 2016, 2017 and ~~2017,~~2018, we received respectively ~~$52.0 million,~~ $27.3 million, $8.1 million and ~~$8.1~~$4.7 million in payments pursuant to the ~~Pfizer~~Pfizer/Allogene and Servier collaborations.

Financial Operations Overview

We have incurred net losses in nearly each year since our inception. Substantially all of our net losses resulted from costs incurred in connection with our development programs and from selling, general and

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administrative expenses associated with our operations. As we continue our intensive research and development programs, we expect to continue to incur significant expenses and may again incur operating losses in future periods. We anticipate that such expenses will increase substantially if and as we:

progress the clinical trial of our wholly-controlled UCART123 product candidate and initiate additional clinical trials for other wholly-controlled product candidates;

continue to advance the research and development of our current and future immuno-oncology product candidates;

continue, through Calyxt, to advance the research and development of our current and future agricultural product candidates;

initiate additional clinical studies for, or additionalpre-clinical development of, our immuno-oncology product candidates;

conduct and multiply, though Calyxt, additional field trials of our agricultural product candidates;

further develop and refine the manufacturing process for our immuno-oncology product candidates;

change or add additional manufacturers or suppliers of biological materials;

build-up, commission and operate our own manufacturing capacity;

seek regulatory and marketing approvals for our product candidates, if any, that successfully complete development;

establish a sales, marketing and distribution infrastructure to commercialize any products for which we may obtain marketing approval;

seek to identify and validate additional product candidates;

acquire orin-license other product candidates, technologies, germplasm or other biological material;

make milestone or other payments under anyin-license agreements;

maintain, protect and expand our intellectual property portfolio;

secure manufacturing arrangements for commercial production;

seek to attract and retain new and existing skilled personnel;

create additional infrastructure to support our operations as a public company; and

experience any delays or encounter issues with any of the above.

We do not expect to generate material revenues from sales of our product candidates unless and until we successfully complete development of, and obtain marketing approval for, one or more of our product candidates, which we expect will take a number of years and is subject to significant uncertainty. Accordingly, we anticipate that we will need to raise additional capital prior to completing clinical development of any of our product candidates. Until such time that we can generate substantial revenues from sales of our product candidates, if ever, we expect to finance our operating activities through a combination of milestone payments received pursuant to our strategic alliances, equity offerings, debt financings, government or other third-party funding and collaborations, and licensing arrangements. However, we may be unable to raise additional funds or enter into such arrangements when needed on favorable terms, or at all, which would have a negative impact on our financial condition and could force us to delay, limit, reduce or terminate our development programs or commercialization efforts or grant to others rights to develop or market product candidates that we would otherwise prefer to develop and market ourselves. Failure to receive additional funding could cause us to cease operations, in part or in full.

Our consolidated financial statements for ~~2015,~~ 2016, 2017 and ~~2017~~2018 have been prepared in accordance with International Financial Reporting Standards, or IFRS, as issued by the International Accounting Standards Board, or IASB.

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Financial Operations Overview

Revenues and Other Income

Revenues

We currently derive all our revenues from payments pursuant to our collaboration agreements with ~~Pfizer~~Allogene and Servier, patent licensing arrangements and royalties on licensed technologies. Our collaboration agreements provide fornon-refundable

upfront payments that we received upon execution of the relevant agreement, milestone payments that we are entitled to receive when the triggering event has occurred and the performance obligation is achieved, research and development cost reimbursements that are recognized over the period of these services and royalty payments. The triggering event for a milestone payment may be the receipt of favorable scientific results, regulatory approval, or marketing of products developed pursuant to the agreement. Royalties are based on sales of licensed products or technologies. ~~They are recognized in accordance with the terms of the licensing agreement when sales can be determined reliably and there is reasonable assurance that the receivables from outstanding royalties~~Royalty revenues, if earned, will be ~~collected.~~recognized at the later of when (1) the subsequent sale or usage occurs; and (2) the performance obligation to which the sales-based or usage-based royalties relates has been satisfied.

Our ability to generate product revenues and become profitable depends upon our and our collaborators’ ability to successfully develop and commercialize products. If we fail to become profitable or are unable to sustain profitability on a continuing basis, then we may be unable to continue our operations at planned levels and be forced to reduce our operations.

Other Income

Government Grants

Due to the innovative nature of our product candidate development programs, we have benefited from a certain number of sources of assistance from the French government or local public authorities, intended to finance our research and development efforts or the recruitment of specific personnel. Government grants that offset expenses that we incur for those research programs are recognized as other income in the period in which the expenses that are reimbursable pursuant to the grant have been incurred.

Research Tax Credit

The main research tax credit that we benefit from is theCrédit d’Impôt Recherche, or CIR, which is granted to companies by the French tax authorities in order to encourage them to conduct technical and scientific research. Companies demonstrating that they have research expenditures that meet the required CIR criteria receive a tax credit that may be used for the payment of their income tax due on the fiscal year in which the expenditures were incurred and during the next three fiscal years. If taxes due are not sufficient to cover the full amount of the tax credit at the end of the three-year period, the difference is repaid to us in cash by the French tax authorities. We also satisfy certain criteria that qualify us as a small/middle size company and permit us to request immediate payment of the CIR. The expenditures taken into account for the calculation of the CIR only involve research expenses.

The main characteristics of the CIR are the following:

the CIR results in a cash inflow to us from the tax authorities;

a company’s corporate income tax liability does not limit the amount of the CIR; and

the CIR is not included in the determination of the corporate income tax.

We have concluded that the CIR meets the definition of a government grant as defined in IAS 20,Accounting for Government Grants and Disclosure of Government Assistance, and that the classification as other income within operating loss in our statement of operations is appropriate.

In December 2018, the French Tax Authority has initiated an audit related to the 2014, 2015, 2016 and 2017 French research tax credits. We do not believe that a provision should be recorded at this stage of this audit. As a result of such audit, the reimbursement of the French research tax credit related to 2017 is currently pending.

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Operating Expenses

Our operating expenses consist primarily of royalty expenses, research and development expenses and selling, general and administrative expenses.

Royalty Expenses

We have entered into several license agreements to obtain access to technology that we use in our product development efforts. Royalty expenses consist ofin-licensing costs, which reflect royalties we pay to use rights granted to us. Depending on the contractual provisions, royalty expenses are either proportional to revenues generated by using the patents or fixed annual royalties or conditioned by milestones.

Research and Development Expenses

We engage in substantial research and development efforts to develop innovative CART-cell immunotherapy and agricultural product candidates.

Research and development expenses consist primarily of:

personnel costs, including salaries, related benefits and share-based compensation, for our employees engaged in scientific research and development functions;

cost of third-party contractors such as contract research organizations, or CROs, and academic institutions involved inpre-clinical or clinical trials that we may conduct, or third-party contractors involved in field trials;

purchases and manufacturing of biological materials, real-estate leasing costs as well as conferences and travel costs; and

certain other expenses, such as expenses for use of laboratories and facilities for our research and development activities.

We classify personnel and other costs related to information technology, human resources, business development, legal, intellectual property and general management in research and development expense based on the time that employees spent contributing to research and development activities versus general and administrative activities.

Our research and development efforts are focused on our existing product ~~candidates:~~candidates, (i) ~~our~~ UCART123 product candidate, which entered into clinical trials in the United States in February 2017, (ii) ~~our~~ UCART22 ~~and~~product candidate, for which FDA granted in May 2018 Cellectis an IND approval to conduct a Phase I clinical study with UCART22 in ALL in the United States, (iii) UCARTCS1 product ~~candidates,~~candidate, for which ~~are~~FDA granted in January 2019 Cellectis an IND approval to conduct a Phase I clinical study with UCARTCS1 in MM in the ~~manufacturing phase,~~United States , and ~~(iii) our~~(iv) other undisclosed ~~targets~~target which are in the development phases. We use our employee and infrastructure resources across multiple research and development programs directed toward developing our cell-based platform and for identifying and developing product candidates. We manage certain activities such aspre-clinical and clinical research and manufacture of product candidates through our partner institutions or other third-party vendors. Due to the number of ongoing projects and our ability to use resources across several projects, we do not record or maintain information regarding the costs incurred for our research and development programs on a program-specific basis.

Our research and development efforts are central to our business and account for a significant portion of our operating expenses. We expect that our research and development costs will increase in the foreseeable future as we continue to implement our new clinical trials, manufacturepre-commercial clinical trial andpre-clinical study materials, expand our research and development and process development efforts, seek regulatory approvals for our product candidates that successfully complete clinical trials, as well as access and develop additional technologies, and hire additional personnel to support our research and development efforts. This is because product candidates in later stages of clinical development generally have higher development costs than those in

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earlier stages of development, primarily due to the increased size and duration of later-stage clinical trials. Likewise, in our plant products business, we expect our research and development expenses will continue to increase over the next several years as we develop new agricultural product candidates and advance them through field trials toward commercial proof of concept.

We cannot determine with certainty the duration and completion costs of our future clinical trials of our product candidates or if, when, or to what extent we will generate revenues from the commercialization and sale of any of our product candidates, or those of our collaborators, that might obtain regulatory approval. We also cannot determine with certainty the duration and completion costs of our future field trials of our agricultural product candidates or if, when, or to what extent we will generate revenues from the commercialization and sale of any of our agricultural product candidates that might obtain regulatory approval. We may never succeed in achieving regulatory approval for any product candidates. The duration, costs and timing of clinical trials and development of our product candidates will depend on a variety of factors, including:

the scope, rate of progress and expense of our ongoing as well as any additionalpre-clinical studies, clinical trials and other research and development activities;

clinical trial and early-stage results;

the terms and timing of regulatory approvals;

the expense of filing, prosecuting, defending and enforcing patent claims and other intellectual property rights;

the ability to market, commercialize and achieve market acceptance for any product candidate that we may develop in the future; and

the scope, rate of progress and expense of our ongoing as well as any additional studies for our agricultural product candidates, field trials and other research and development activities.

Selling, General and Administrative Expenses

Selling, general and administrative expenses consist primarily of employee-related expenses for executive, business development, intellectual property, finance, legal and human resource functions. Administrative expenses also include facility-related costs and service fees, other professional services and recruiting fees.

We anticipate that our selling, general and administrative expenses will increase in the future as we increase our headcount to support the expected growth in our research and development activities and the potential commercialization of our product candidates. We also expect to continue to incur significant expenses associated with Cellectis S.A. and Calyxt, Inc. being a public company in the United States, including costs related to audit, legal, regulatory andtax-related services associated with maintaining compliance with U.S. exchange listing and SEC requirements, director and officer insurance premiums, and investor relations costs.

Financial Gain (Loss)

Financial gain (loss) mainly consists of interest income related to our savings accounts and bank deposits, exchange gains and losses associated with transactions in foreign currencies and fair value of our financial assets and derivative instruments. Significant transactions in foreign currencies are translated into euros at the exchange rates effective at the transaction dates, while the average rate for the previous month is used fornon-significant transactions. Monetary assets and liabilities denominated in foreign currencies at the reporting date are translated

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into euros using the exchange rate effective at that date. The resulting exchange gains or losses are recorded in the consolidated statements of income as financial income or expense. Financial gain (loss) reflects the net impact of financial income and financial expenses.

Critical Accounting Policies and Estimates

Some of the accounting methods and policies used in preparing our financial statements under IFRS are based on complex and subjective assessments by our management or on estimates based on past experience and assumptions deemed realistic and reasonable based on the circumstances concerned. The actual value of our assets, liabilities and shareholders’ equity and of our losses could differ from the value derived from these estimates if conditions changed and these changes had an impact on the assumptions adopted. We believe that the most significant management judgments and assumptions in the preparation of our financial statements are named below. For further details, see Notes to our consolidated financial statements.

Revenue Recognition: Collaboration Agreements and Licenses, Sales of Products and Services (Note 3.1)

Research Tax Credit (Note 3.1)

Share-Based Compensation (Note 15)

A.	Operating Results

The following table sets forth our selected consolidated statement of income data:

   For the year ended December 31,
   2015    2016    2017
   ($ in thousands)
Revenues and other income

Revenues
   55,864    44,808    25,188
Other income
   6,701    11,637    8,528






Total revenues and other income
   62,565    56,444    33,715






Operating expenses

Royalty expenses
   (2,746 )    (1,777 )    (2,620 )
Research and development expenses
   (58,154 )    (78,458 )    (79,227 )
Selling, general and administrative expenses
   (30,223 )    (43,413 )    (44,750 )
Other operating income (expenses)
   (2,425 )    (99 )    232






Operating income (loss)
   (30,984 )    (67,302 )    (92,650 )






Financial income
   10,253    7,147    7,262
Financial expenses
   (1,876 )    (7,101 )    (18,294 )






Financial gain (loss)
   8,378    46    (11,032 )






Income tax
   —      —      —
Net income (loss)
   (22,606 )    (67,255 )    (103,683 )






Attributable to shareholders of Cellectis
   (22,796 )    (67,255 )    (99,368 )
Attributable tonon-controlling interests
   190    —      (4,315 )

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	For the year ended December 31,
	2016			2017			2018
				($ in thousands)
Revenues and other income
Revenues		44,808			25,188			12,731
Other income		11,637			8,528			8,701

Total revenues and other income		56,444			33,715			21,432

Operating expenses
Royalty expenses		(1,777	)		(2,620	)		(2,739	)
Research and development expenses		(78,458	)		(79,227	)		(76,567	)
Selling, general and administrative expenses		(43,413	)		(44,750	)		(47,248	)
Other operating income (expenses)		(99	)		232			31

Operating income (loss)		(67,302	)		(92,650	)		(105,091	)

Financial income		7,147			7,262			20,572
Financial expenses		(7,101	)		(18,294	)		(3,813	)

Financial gain (loss)		46			(11,032	)		16,758

Income tax		—			—			—
Net income (loss)		(67,255	)		(103,683	)		(88,333	)

Attributable to shareholders of Cellectis		(67,255	)		(99,368	)		(78,693	)
Attributable tonon-controlling interests		—			(4,315	)		(9,640	)

Years Ended December 31, ~~2015,~~ 2016, 2017 and ~~2017~~2018

Revenues.


	For the year ended December 31,						% change						For the year ended December 31,						% change
	2015		2016		2017		2016 vs 2015			2017 vs 2016			2016		2017		2018		2017 vs 2016			2018 vs 2017
Collaboration agreements		53,580		41,891		22,821		-21.8	%		-45.5	%		41,891		22,821		10,497		-45.5	%		-54.0	%
Other revenues		2,284		2,916		2,367		27.7	%		-18.8	%		2,917		2,367		2,234		-18.8	%		-5.6	%
Revenues		55,864		44,808		25,188		-19.8	%		-43.8	%		44,808		25,188		12,731		-43.8	%		-49.5	%

The decrease in revenues of $12.5 million, or 49.5%, between the years ended December 31, 2017 and 2018 primarily reflects a decrease of $12.3 million in revenues under our collaboration agreements of which $7.8 million relates to a decrease in recognition of upfront fees already paid to Cellectis and $4.6 million relates to lower research and development cost reimbursements, and a decrease of $0.1 million of licensing revenues.

The decrease in revenues of $19.6 million, or 43.8%, between the years ended December 31, 2016 and 2017 primarily reflects a decrease of $19.1 million in revenues under our collaboration agreements, of which $8.5 million representsone-time milestone revenue received during the second quarter of 2016 with the first patient dosed in the Phase 1 clinical trial for UCART19, $6.0 million represents decreased recognition of upfront fees already paid to Cellectis, $1.9 million represents decreased research and development cost reimbursements (research services and ~~direct costs)~~costs incurred by Cellectis on Servier projects) and $2.8 million represents decreased revenue due to 2016one-time payments by Servier for the supply of raw materials and batches of UCART19 products, partially offset by the increase of $0.1 million in other services and products provided to Pfizer, and a $0.5 million decrease in licensing revenues.

Other income.


	For the year ended December 31,						% change
	2016		2017		2018		2017 vs 2016			2018 vs 2017
Research tax credit		10,038		8,327		8,561		-17.0	%		2.8	%
Other income		1,599		201		140		-87.4	%		-30.4	%
Other income		11,637		8,528		8,701		-26.7	%		2.0	%

The ~~decrease~~increase in ~~revenue~~other income of ~~$11.1~~$0.2 million, or ~~19.8%~~2.0%, between the years ended December 31, ~~2015~~2017 and ~~2016 is mainly due (i) to the decrease~~2018 reflects an increase of ~~$11.7~~$0.2 million in ~~collaboration revenues~~research tax credits, due to ~~revenue of $20.9 million recorded in 2015 in relation to~~higher research and development purchases and external expenses that are eligible for the early exercise by Servier of its option to acquire the exclusive worldwide rights to further develop and commercialize UCART19 partially offset by revenue totaling $13.2 million from an agreement to provide Servier with raw materials and additional batches of UCART19 products and the achievement of two milestones in 2016 and (ii) to the increase of $0.6 million in licenses fees.tax credit.

~~Other income.~~

For the year ended December 31,
   % change
2015      2016      2017    2016 vs
2015 2017 vs
2016
Research tax credit
5,591      10,038      8,327    79.5 % -17.0 %
Other income
1,110      1,599      201    44.1 % -87.4 %
Other income
6,701      11,637      8,528    73.7 % -26.7 %

The decrease in other income of $3.1 million, or 26.7%, between the years ended December 31, 2016 and 2017 reflects a decrease of $1.7 million in research tax credit, and a decrease of $1.4 million in research subsidies, resulting from settlements received after termination of research programs during the last quarter of the year ended December 31, 2016.

Royalty expenses.


	For the year ended December 31,									% change
	2016			2017			2018			2017 vs 2016			2018 vs 2017
Royalty expenses		(1,777	)		(2,620	)		(2,739	)		47.5	%		4.5	%

The increase in ~~other income~~royalty expenses of ~~$4.9~~$0.1 million, or ~~73.7%~~4.5%, between the years ended December 31, ~~2015~~2017 and ~~2016~~2018 primarily reflects ~~an increase of $4.4 million in research tax credit, and an increase of $0.5 million in research subsidies, resulting from settlements after termination of research programs.~~higher royalty expenses paid to our existing partners.

~~Royalty expenses.~~

For the year ended December 31,
% change
   2015    2016    2017 2016 vs
2015 2017 vs
2016
Royalty expenses
   (2,746 )    (1,777 )    (2,620 ) -35.3 % 47.5 %

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The increase in royalty expenses of $0.8 million, or 47.5%, between the years ended December 31, 2016 and 2017 primarily reflects higher payments to existing license providers.

~~The decrease in royalty expenses of $1.0 million, or 35.3%, between the years ended December 31, 2015 and 2016 primarily reflects lower expense to existing license providers.~~

Research and development expenses.


	For the year ended December 31,									% change						For the year ended December 31,									% change
	2015			2016			2017			2016 vs 2015			2017 vs 2016			2016			2017			2018			2017 vs 2016			2018 vs 2017
Personnel expenses		(39,341	)		(48,982	)		(37,906	)		24.5	%		-22.6	%		(48,982	)		(37,906	)		(34,608	)		-22.6	%		-8.7	%
Purchases, external expenses and other		(18,814	)		(29,476	)		(41,321	)		56.7	%		40.2	%		(29,476	)		(41,321	)		(41,959	)		40.2	%		1.5	%
Research and development expenses		(58,154	)		(78,458	)		(79,227	)		34.9	%		1.0	%		(78,458	)		(79,227	)		(76,567	)		1.0	%		-3.4	%

For the years ended December 31, 2017 and 2018 research and development expenses decreased by $2.6 million or 3.4%. Personnel expenses decreased by $3.3 million from $37.9 million in 2017 to $34.6 million in 2018 primarily due to a $5.8 million decrease innon-cash stock-based compensation expense and $1.0 million decrease in social charges on stock options offset by a $3.5 million increase in wages and salaries mainly explained by increase in R&D headcount in therapeutic activity. Purchases and external expenses increased by $2.0 million from $38.5 million in 2017 to $40.5 million in 2018, mainly due to an increase in costs to acquire grain prior to our achievement of commercial milestones at Calyxt. Other expenses, which relate to continuing leasing and other commitments, decreased by $1.4 million for the years ended December 31, 2017 and 2018.

During the years ended December 31, 2016 and 2017 research and development expenses increased by $0.8 million or 1.0%. Personnel expenses decreased by $11.1 million from $49.0 million in 2016 to $37.9 million in 2017, notably due to a $9.4 million decrease innon-cash ~~stock based~~stock-based compensation, a $2.8 million decrease in social charges on stock option grants, partly offset by a $1.1 million increase in wages and salaries. Purchases and external expenses increased by $10.7 million from $27.7 million in 2016 to $38.5 million in 2017 mainly due to increased expenses related to payments to third parties participating in product development, purchases of biological raw materials, expenses related to process development and expenses associated with the use of laboratories and other facilities. Research and development expenses in 2017 include manufacturing costs related to UCART123, UCART CS1 and UCART22 and expenses related to UCART123 clinical trials. Other expenses increased by $1.1 million mainly due to the impairment of assets for $0.8 million related to the vacant Montvale site recorded in 2017. Otherwise, other expenses include continuing leasing and other commitments.

During the years ended December 31, 2015 and 2016, research and development expenses increased by $20.3 million or 34.9%. Personnel expenses increased by $9.6 million from $39.3 million in 2015 to $49.0 million in 2016, notably due to a $1.8 million increase in wages and salaries, and a $12.6 million increase in~~non-cash~~ stock based compensation expense, partly offset by a $4.8 million decrease in social charges on stock option and free shares grants. Purchases and external expenses increased by $10.8 million from $16.9 million in 2015 to $27.7 million in 2016, mainly due to increased expenses related to UCART123 and other product candidates’ development, including payments to third parties, purchases of biological materials and expenses associated with the use of laboratories and other facilities. Expenses in 2016 also include costs related to preparation of UCART123 clinical trials. Other expenses relate to continuing leasing and other commitments and decreased by $0.1 million.

Selling, general and administrative expenses.


	For the year ended December 31,									% change						For the year ended December 31,									% change
	2015			2016			2017			2016 vs 2015			2017 vs 2016			2016			2017			2018			2017 vs 2016			2018 vs 2017
Personnel expenses		(21,735	)		(33,523	)		(34,486	)		54.2	%		2.9	%		(33,523	)		(34,486	)		(30,563	)		2.9	%		-11.4	%
Purchases, external expenses and other		(8,489	)		(9,890	)		(10,264	)		16.5	%		3.8	%		(9,890	)		(10,264	)		(16,685	)		3.8	%		62.6	%
Selling, general and administrative expenses		(30,223	)		(43,413	)		(44,750	)		43.6	%		3.1	%		(43,413	)		(44,750	)		(47,248	)		3.1	%		5.6	%

~~During~~For the years ended December 31, 2017 and 2018, the increase in selling, general and administrative expenses of $2.5 million, or 5.6%, primarily reflects (i) a $5.1 million increase in purchases and external expenses from $9.1 million in 2017 to $14.3 million in 2018, particularly linked to Calyxt’s increasing activity (from $4.3 million in 2017 to $7.8 million in 2018) explained by costs incurred to ramp up capabilities in advance of commercialization and increases in professional services expenses associated with being a public company, (ii) an increase of $1.3 million in other expenses related to taxes, various depreciation and amortization, and (iii) a decrease of $3.9 million in personnel expenses from $34.5 million to $30.6 million, attributable to a $7.4 million decrease innon-cash stock based compensation and $0.9 million decrease in social charges on stock options partially offset by a $4.4 million increase in wages and salaries, mainly explained by increase in SG&A headcount at Calyxt.

For the years ended December 31, 2016 and 2017 the increase in selling, general and administrative expenses of $1.3 million, or 3.1%, primarily reflects (i) an increase of $1.0 million in personnel expenses from $33.5 million to $34.5 million, attributable to a $2.0 million increase in wages and salaries, and an increase of $1.2 million ofnon-cash stock-based compensation expense, partly offset by a decrease of $2.2 million of social charges on stock options grants, and (ii) an increase of $0.3 million in purchases and external expenses. Other

~~133~~

expenses relate to taxes, various depreciation and amortization and other commitments and increased by $0.1 million, due to higher business taxes and higher provisions.

~~During~~Other operating income and expenses.


	For the year ended December 31,							% change
	2016			2017		2018		2017 vs 2016			2018 vs 2017
Other operating income (expenses)		(99	)		232		31		-334.2	%		-86.6	%

The decrease in other operating income and expenses between the years ended December 31, ~~2015~~2017 and ~~2016,~~2018 amounted to $0.2 million. During the ~~increase in selling, general~~year ended December 31, 2018, other operating income and ~~administrative~~ expenses primarily include reversal of ~~$13.2~~commercial litigation provision for $0.6 million, or 43.6%, primarily reflects (i) an increase of $11.8 million in personnel expenses from $21.7 million to $33.5 million, attributable to a $1.0 million increase in wages and salaries, and an increase of $12.6 million of~~non-cash~~ ~~stock-based compensation expense, partly~~partially offset by ~~a decrease of $1.8 million of~~ social charges paid on ~~stock options and free share grants, and (ii) an increase~~former employee compensation for $0.2 million, reversal of ~~$2.1~~subsidy for $0.2 million ~~in purchases and external expenses. Other expenses relate to taxes, various depreciation and amortization~~ and other ~~commitments and decreased by $0.7 million, due to lower business taxes and lower provisions.~~immaterial variances.

~~Other operating income (expenses).~~

For the year ended December 31,    % change
2015    2016    2017    2016 vs
2015 2017 vs
2016
Other operating income (expenses)
(2,425 )    (99 )    232    -95.9 % -334.2 %

The increase in other operating income and expenses between the years ended December 31, 2016 and 2017 amounted to $0.3 million. For the ~~year~~years ended December 31, 2017, other operating income primarily reflects (i) a receivable related to the refund of social charges paid on Cellectis free share grants that expired without being vested for $0.2 million, (ii) reversals of personnel litigation for a total amount of $0.1 million, and is partially offset by other operating expenses for $0.1 million related to social charges paid on former employee compensation.

Financial income.


	For the year ended December 31,						% change
	2016		2017		2018		2017 vs 2016			2018 vs 2017
Financial income		7,147		7,262		20,572		1.6	%		183.3	%

The ~~decrease~~increase in ~~net other operating expenses~~financial income of $13.3 million, or 183.3%, between the years ended December 31 ~~2015~~2017 and ~~2016 amounted~~2018 was mainly attributable to ~~$2.3~~an increase in foreign exchange realized and unrealized gain of $12.4 million ~~or 95.9%. Other operating expenses for the year ended December 31, 2016 included $0.4~~and an increase in interest income of $4.8 million ~~provisions for personnel~~ and ~~commercial litigations that have been partially~~partly offset by ~~other operating income, including (i)~~ a~~one-off~~ ~~tax reimbursement, (ii) the reversal~~ decrease in fair value of ~~lease incentive deferrals~~financial derivative instruments and ~~(iii) reversals~~current financial assets of reserves for personnel and commercial litigation. During the year ended December 31, 2015, other operating income and expenses included income from (i) the reversal of a subsidy provision, (ii) the reversal of lease incentives deferrals, and (iii) a gain from a settlement with a supplier, while other operating expenses reflected settlements signed in 2015.$3.9 million.

~~Financial income.~~

For the year ended December 31,
   % change
2015      2016      2017    2016 vs
2015 2017 vs
2016
Financial income
10,253      7,147      7,262    -30.3 % 1.6 %

The increase in financial income of $0.1 million, or 1.6%, between the years ended December 31 2016 and 2017 was mainly attributable to an increase in fair value of financial derivative instruments and current financial assets of $3.2 million and an increase in interest income of $0.5 million partly offset by a decrease in foreign exchange realized and unrealized gain of $3.6 million.

Financial expenses.


	For the year ended December 31,									% change
	2016			2017			2018			2017 vs 2016			2018 vs 2017
Financial expenses		(7,101	)		(18,294	)		(3,813	)		157.6	%		-79.2	%

The decrease in financial ~~income~~expense of ~~$3.1~~$14.5 million, or ~~30.3%~~79.2%, between the years ended December 31 ~~2015~~2017 and ~~2016,~~2018 was mainly attributable to ~~the~~a decrease ~~of $4.3 million of~~in foreign exchange realized and unrealized gain of $14.6 million and partly offset by an increase in ~~interest income~~other financial expense of ~~$0.5~~$0.1 million.

~~134~~

~~Financial expenses.~~

   For the year ended December 31, % change
   2015 2016 2017 2016 vs
2015 2017 vs
2016
Financial expenses
   (1,876 ) (7,101 ) (18,294 ) 278.6 % 157.6 %

The increase in financial expenses of $11.2 million, or 157.6%, between the years ended December 31 2016 and 2017 was mainly attributable to $13.5 million increase in foreign exchange realized and unrealized loss, partly offset by a decrease in fair value of financial derivative instruments and other current assets of $2.7 million.

Net Income / loss.


	For the year ended December 31,									% change
	2016			2017			2018			2017 vs 2016			2018 vs 2017
Net income (loss)		(67,255	)		(103,683	)		(88,333	)		54.2	%		-14.8	%

The ~~change~~decrease in ~~financial expenses~~net loss of ~~$5.2~~$15.4 million ~~or 278.6%,~~ between the years ended December 31, ~~2015~~2017 and ~~2016,~~2018 was mainly ~~attributable~~due to ~~$2.3~~(i) a $27.8 million increase in ~~foreign exchange realized~~financial result, (ii) a $13.2 million decrease innon-cash stock-based compensation expense and ~~unrealized loss~~(iii) a decrease of $1.8 million in social charges on stock options, partially offset by (a) a $12.3 million decrease in revenues and ~~$2.7~~other income, (b) a $7.8 million increase in ~~fair value of financial derivative instruments~~wages, (c) a $7.1 million increase in purchases, external expenses and ~~current financial assets.~~other, (d) a $0.2 million decrease in other operating income and (e) a $0.1 increase in royalty expenses.

~~Net Income (loss).~~

For the year ended December 31, % change
2015 2016 2017 2016 vs
2015 2017 vs
2016
Net income (loss)
(22,606 ) (67,255 ) (103,683 ) 197.5 % 54.2 %

The increase in net loss of $36.4 million, or 54.2%, between the ~~year~~years ended December 31, 2016 and 2017 was mainly due to (i) a $22.7 million decrease in revenues and other income, (ii) anthe $11.1 million decrease in financial gain (loss), (iii) ana $11.0 million increase in purchases and external expenses, (iv) a $3.1 million increase in wages, (v) a $1.1 million increase in other selling, general and administrative expenses and research and development expenses, ~~and~~ (vi) a $0.8 million increase in royalty expenses, partially offset by (a) a $8.2 million decrease innon-cash stock-based compensation expense, and (b) a $5.5 million decrease in social charges on stock options and free share grants. The remaining difference is mainly due to the decrease in other operating expenses.

~~The increase in net loss from continuing operations of $44.6 million, or 197.5%, between the year ended December 31, 2015 and 2016 was mainly due to (i) a $25.2 million increase innon-cash~~ stock-based compensation expense, (ii) a $12.9 million increase in purchases and external expenses, (iii) a $2.8 million increase in wages, (iv) a $6.1 million decrease in revenues and other income and (v) an $8.3 million decrease in financial gain (loss), partially offset by $6.6 decrease in social charges on stock options and free share grants. The remaining difference is mainly due to the decrease in other operating expenses.

~~Gain (loss)~~Gain/Loss attributable tonon-controlling interests.

   For the year ended December 31, % change
   2015      2016      2017 2016 vs
2015 2017 vs
2016
Gain (loss) attributable tonon-controlling interests
   190      —        (4,315 ) -100.0 % 0.0 %


	For the year ended December 31,								% change
	2016		2017			2018			2017 vs 2016		2018 vs 2017
Gain (loss) attributable tonon-controlling interests		0		(4,315	)		(9,640	)		n.a.		123.4	%

During the year ended December 31, 2018, we recorded $9.6 million in loss attributable tonon-controlling interests. The increase in net loss attributable tonon-controlling interests of $5.3 million is a result of increase in Calyxt’s net loss and an increase in % of Calyxt common shares being owned by minority shareholders after thefollow-on offering in May 2018.

During the year ended December 31, 2017, we recorded $4.3 million in loss attributable tonon-controlling interests. The change in net loss attributable tonon-controlling interests is a result of 20.3% of Calyxt common shares being owned by minority shareholders ~~following~~after its initial public offering.

~~During the year ended December 31, 2015, we recorded $0.2 million in gain attributable tonon-controlling~~ ~~interests.Non-controlling~~ ~~interests for 2015 include 25% of the income of Cellectis Bioresearch and its~~

~~135~~

~~subsidiaries until our repurchase from “~~~~Caisse des Dépôts et Consignations~~~~” on May 18, 2015 of its shares, which represented the entirety of thisnon-controlling~~ ~~interest. As such, there is no income (loss) attributable tonon-controlling~~ ~~interests in 2016.~~

Segment Results

Information related to each of our reportable segments is set out below. Segment revenues and Other income, Research and development expenses, Selling, general and administrative expenses, and Royalties and other operating income and expenses, and Adjusted net income (loss) attributable to shareholders of Cellectis (which does not includenon-cash stock-based expense) are used by the CODM to measure performance. The CODM does not review any asset or liability information by segment or by region.

Adjusted Net Income (Loss) attributable to shareholders of Cellectis is not a measure calculated in accordance with IFRS. Because Adjusted Net Income (Loss) attributable to shareholders of Cellectis excludesNon-cash stock based compensation expense—anon-cash expense, we believe that this financial measure, when considered together with our IFRS financial statements, can enhance an overall understanding of Cellectis’ financial performance. Moreover, our management views the Company’s operations, and manages its business, based, in part, on this financial measure.

There are inter-segment transactions between the two reportable segments, including the allocation of corporate general and administrative expenses by Cellectis S.A. and the allocation of research and development expenses among the reportable segments. With respect to corporate general and administrative expenses, Cellectis S.A. provides Calyxt, Inc. with general sales and administrative functions, accounting and finance functions, investor relations, intellectual property, legal advice, human resources, communication and information technology pursuant to a management agreement. Under the management agreement, Cellectis S.A. charges Calyxt, Inc. in euros at cost plus amark-up ranging between zero to 10%, depending on the nature of the service. Amounts due to Cellectis S.A. pursuant to inter-segment transactions bear interest at a rate of12-month Euribor plus 5% per annum.

The intersegment revenues represent the transactions between segments. Intra-segment transactions are eliminated within a segment’s results and intersegment transactions are eliminated in consolidation as well as in key performance indicators by reportable segment.

Years Ended December 31, ~~2015,~~ 2016, 2017 and ~~2017~~2018

The following table summarizes segment revenues and segment operating profit (loss) for the years ended December ~~31, 2015,~~ 2016, 2017 and ~~2017:~~2018:


	For the year ended December 31, 2016									For the year ended December 31, 2017									For the year ended December 31, 2018
$ in thousands	Plants			Therapeutics			Total reportable segments			Plants			Therapeutics			Total reportable segments			Plants			Therapeutics			Total reportable segments
External revenues		399			44,409			44,808			508			24,680			25,188			236			12,495			12,731
External other income		186			11,450			11,637			239			8,290			8,528			178			8,523			8,701

External revenues and other income		585			55,859			56,444			747			32,969			33,715			414			21,018			21,432

Royalty expenses		(468	)		(1,309	)		(1,777	)		(390	)		(2,230	)		(2,620	)		(595	)		(2,144	)		(2,739	)
Research and development expenses		(4,112	)		(74,345	)		(78,458	)		(6,057	)		(73,170	)		(79,227	)		(8,638	)		(67,929	)		(76,567	)
Selling, general and administrative expenses		(4,809	)		(38,603	)		(43,413	)		(13,143	)		(31,607	)		(44,750	)		(21,067	)		(26,180	)		(47,248	)
Other operating income and expenses		(6	)		(93	)		(99	)		6			225			232			(50	)		81			31

Total operating expenses		(9,395	)		(114,351	)		(123,746	)		(19,584	)		(106,782	)		(126,366	)		(30,351	)		(96,172	)		(126,523	)

Operating income (loss) before tax		(8,810	)		(58,492	)		(67,302	)		(18,837	)		(73,813	)		(92,650	)		(29,937	)		(75,154	)		(105,091	)

Financial gain (loss)		87			(41	)		46			—			(11,032	)		(11,032	)		1,420			15,339			16,758

Net income (loss)		(8,722	)		(58,533	)		(67,255	)		(18,837	)		(84,846	)		(103,683	)		(28,517	)		(59,816	)		(88,333	)

Net income (loss) from discontinued operations		—			—			—			—			—			—			—			—			—
Non controlling interests		—			—			—			4,315			—			4,315			9,640			—			9,640

Net income (loss) attributable to shareholders of Cellectis		(8,722	)		(58,533	)		(67,255	)		(14,522	)		(84,846	)		(99,368	)		(18,877	)		(59,816	)		(78,693	)

R&Dnon-cash stock-based expense attributable to shareholder of Cellectis		477			32,731			33,208			967			22,623			23,590			838			16,852			17,689
SG&Anon-cash stock-based expense attributable to shareholder of Cellectis		621			24,793			25,414			4,990			20,345			25,335			5,218			11,655			16,873

Adjustment of share-based compensation attributable to shareholders of Cellectis		1,098			57,524			58,622			5,957			42,968			48,925			6,056			28,507			34,563

Adjusted net income (loss) attributable to shareholders of Cellectis		(7,625	)		(1,009	)		(8,633	)		(8,565	)		(41,877	)		(50,442	)		(12,821	)		(31,309	)		(44,130	)
																		��
Depreciation and amortization		(345	)		(1,866	)		(2,211	)		(551	)		(2,820	)		(3,371	)		(637	)		(1,740	)		(2,377	)
Additions to tangible and intangible assets		10,410			4,164			14,573			792			1,849			2,642			1,871			3,040			4,911
Impairment of tangible assets		—			—			—			—			(798	)		(798	)		—			—			—

~~136~~Therapeutics segment—2017 vs. 2018

External revenues and other income in our Therapeutics segment decreased by $12.0 million, from $33.0 million for the year ended December 31, 2017 to $21.0 million for the year ended December 31, 2018. The decrease was primarily due to a decrease of $12.3 million in collaboration agreement revenues, partially offset by a $0.2 million increase in research tax credit, as described in sections “Revenues” and “Other income” under “Results of Operation” for the consolidated Group.

The decrease in total operating expenses of $10.6 million, from $106.8 million for the year ended December 31, 2017 to $96.2 million for the year ended December 31, 2018 resulted primarily from (i) lower personnel expenses, attributable, to a decrease of $14.5 million innon-cash stock-based compensation expenses and a decrease of $1.8 million in social charges on stock options, lower other operating expenses for $0.7 million partly offset by an increase of $4.7 million in personnel wages and salaries, and higher purchases, external expenses for $1.7 million.

Operating loss before tax for our Therapeutics segment increased by $1.4 million from $73.8 million for year ended December 31, 2017, to $75.2 million for year ended December 31, 2018.

   For the year ended December 31, 2015 For the year ended December 31, 2016 For the year ended December 31, 2017
$ in thousands    Plants Therapeutics Total
reportable
segments Plants Therapeutics Total
reportable
segments Plants Therapeutics Total
reportable
segments
Segment revenues and other income (1)
   49 65,159 65,208 716 59,458 60,173 914 35,584 36,498
Inter-segment revenues (1)
   —   (2,643 ) (2,643 ) (130 ) (3,599 ) (3,729 ) (167 ) (2,615 ) (2,782 )


External revenues and other income
   49 62,516 62,565 585 55,859 56,444 747 32,969 33,715


Research and development expenses
   (2,874 ) (55,280 ) (58,154 ) (4,112 ) (74,345 ) (78,458 ) (6,057 ) (73,170 ) (79,227 )
Selling, general and administrative expenses
   (1,834 ) (28,390 ) (30,223 ) (4,809 ) (38,603 ) (43,413 ) (13,143 ) (31,607 ) (44,750 )
Royalties and other operating income and expenses
   (272 ) (4,899 ) (5,171 ) (474 ) (1,402 ) (1,876 ) (384 ) (2,005 ) (2,389 )


Total operating expenses
   (4,980 ) (88,569 ) (93,549 ) (9,395 ) (114,351 ) (123,746 ) (19,584 ) (106,782 ) (126,366 )


Operating income (loss) before tax
   (4,931 ) (26,053 ) (30,984 ) (8,810 ) (58,492 ) (67,302 ) (18,837 ) (73,813 ) (92,650 )


Financial gain (loss)
   259 8,119 8,378 87 (41 ) 46 —   (11,032 ) (11,032 )


Net income (loss)
   (4,672 ) (17,934 ) (22,606 ) (8,722 ) (58,533 ) (67,255 ) (18,837 ) (84,846 ) (103,683 )


Non controlling interests
   —   (190 ) (190 ) —   —   —   4,315 —   4,315


Net income (loss) attributable to shareholders of Cellectis
   (4,672 ) (18,124 ) (22,796 ) (8,722 ) (58,533 ) (67,255 ) (14,522 ) (84,846 ) (99,368 )


Adjustment of share-based compensation
   789 32,613 33,402 1,098 57,524 58,622 5,957 42,968 48,925


Adjusted net income (loss) attributable to shareholders of Cellectis
   (3,883 ) 14,489 10,606 (7,625 ) (1,009 ) (8,633 ) (8,565 ) (41,877 ) (50,442 )


Depreciation and amortization
   (99 ) (1,838 ) (1,937 ) (345 ) (1,866 ) (2,211 ) (551 ) (2,820 ) (3,371 )
Additions to tangible and intangible assets
   526 3,886 4,413 10,410 4,164 14,573 792 1,849 2,642
Impairment of tangible assets
   —   —   —   —   —   —   —   (798 ) (798 )

Adjusted net loss attributable to shareholders of Cellectis for our Therapeutics segment decreased by $10.6 million from $41.9 million for year period ended December 31, 2017 to $31.3 million for the year ended December 31, 2018.

~~(1)~~

Please note that for prior periods our intersegment revenues and other income of Therapeutics segment for 2015 was reported at €0.8 million, or $0.9 million. In this Annual Report, Intersegment revenues and other income of Therapeutics segment for 2015 is reported as $2.6 million and comprise both management fees and direct costs reinvoicing. The counterpart of this change is the line Segment revenues and other income. It has no impact on External revenues of Therapeutics segment.

~~137~~

Therapeutics segment—2016 vs. 2017

External revenues in our Therapeutics segment decreased by $22.9 million, or 41.0%, from $55.9 million for the year ended December 31, 2016 to $33.0 million for the year ended December 31, 2017. The decrease was primarily due to a decrease of $19.1 million in collaboration agreement revenues and lower subsidies revenue and research tax credit, as described in sections “Revenues” and “Other income” of the ~~consolidated~~Group’s operating result analysis.

The decrease in total operating expenses of $7.6 million, or 6.6%, from the year ended December 31, 2016, to the year ended December 31, 2017 resulted primarily from lower personnel expenses, mainly attributable to the decrease innon-cash stock-based compensation expenses and social charges on stock options grants, and is partially offset by the increase in purchases and external expenses for product development.

~~Segment operating~~Operating loss before tax for ~~the~~our Therapeutics segment ~~increased~~decreased by $15.3 million ~~or 26.2%,~~ from ~~the~~$58.5 million for year ended December 31, 2016, to ~~the~~$73.8 million for year ended December 31, 2017.

~~Therapeutics segment—2015 vs. 2016~~

~~External revenues in~~Adjusted net loss attributable to shareholders of Cellectis for our Therapeutics segment ~~decreased~~increased by ~~$6.7~~$40.9 million ~~or 10.6%,~~ from ~~$62.5~~$1.0 million for year period ended December 31, 2016 to $41.9 million for the year ended December 31, ~~2015~~2018.

Plants segment—2017 vs. 2018

External revenues and other income in our Plants segment decreased by $333 thousand from $747 thousand for the year ended December 31, 2017 to ~~$55.9~~$414 thousand for the year ended December 31, 2018, the decrease was attributable to a strategic decision to focus onin-house development of product candidates and to reduce the amount of subcontracted R&D that we were performing for third parties.

The increase in total operating expenses of $10.8 million from $19.6 million for the year ended December 31, ~~2016. The decrease was primarily due~~2017 to a decrease of $11.7 million in collaboration agreement revenues and higher research tax credit, as described in sections “Revenues” and “Other income” of the consolidated operating result analysis.

~~The increase in operating expenses of $25.8 million, or 29.1%, from the year ended December 31,2015 to~~$30.4 for the year ended December 31, ~~2016~~2018 resulted primarily from ~~higher~~a significant increase in Calyxt’s activities, which contributed to (i) an increase of $4.4 million in personnel expenses, ~~attributable, among other things, to the~~that includes an increase of $1.3 million innon-cash stock-based compensation expenses ~~as well as the~~and an increase of $3.1 million in personnel wages and salaries, (ii) an increase of $6.1 million in purchases, external expenses ~~for product development.~~and other. Increase in these costs is mainly explained to acquire grain prior to our achievement of commercial milestones and to ramp up our capabilities in advance of commercialization and increases in professional services expenses associated with being a public company.

~~Segment operating~~Operating loss before tax for ~~the Therapeutics~~our Plants segment increased by ~~$32.4~~$11.2 million ~~or 124.5%s~~ from $18.8 million for the year ended December 31, ~~2015~~2017 to $30.0 million for the year ended December 31, ~~2016.~~2018.

Adjusted net loss attributable to shareholders of Cellectis for our Plants segment increased by $4.2 million from $8.6 million for the year ended December 31, 2017 to $12.8 million for the year ended December 31, 2017.

Plants segment—2016 vs. 2017

External revenues in our Plants segment increased by $0.2 million, or 27.6%, from $0.6 million for the year ended December 31, 2016 to $0.7 million for the year ended December 31, 2017.

The increase in operating expenses of $10.2 million, or 108.4%, from the year ended December 31, 2016 to the year ended December 31, 2017 resulted primarily from a significant increase in ~~Calyxt’s~~Calyxt, Inc. activities, higher professional fees related to the cost of becoming a Nasdaq listed company, as well as an increase innon-cash stock-based compensation expenses.

Segment operating loss before tax ~~for the Plants segment~~ increased by $10.0 million, or 113.8%, from $8.8 million for the year ended December 31, 2016 to $18.8 million for the year ended December 31, 2017.

~~Plants segment—2015 vs. 2016~~

~~External revenues in our Plants segment increased by $0.5 million, from $49 thousand for the year ended December 31, 2015 to $0.6 million for the year ended December 31, 2016.~~

The increase in operating expenses of $4.4 million, or 88.7%, from the year ended December 31, 2015 to the year ended December 31, 2016 resulted primarily from a significant increase in Calyxt’s activities, as well as an increase in~~non-cash~~ ~~stock-based compensation expenses.~~

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Segment operating loss before tax for the Plants segment increased by $3.9 million, or 78.7%, from $4.9 million for the year ended December 31, 2015 to $8.8 million for the year ended December 31, 2016.

B.	Liquidity and Capital Resources

Introduction

We have incurred losses and cumulative negative cash flows from operations since our inception in 2000, and we anticipate that we will continue to incur losses for at least the next several years. We expect that our research and development and selling, general and administrative expenses will continue to increase and, as a result, we will need additional capital to fund our operations, which we may raise through a combination of equity offerings, debt financings, other third-party funding, marketing and distribution arrangements and other collaborations, strategic alliances and licensing arrangements.

We have funded our operations since inception primarily through private and public offerings of our equity securities, grant revenues, payments received under patent licenses, reimbursements of research tax credit claims and payments under our strategic alliances with Pfizer and Servier. Our ordinary shares have been traded on the Euronext Growth market of Euronext in Paris since February 7, 2007 and our ADSs have traded on the Nasdaq Global Market in New York since March 30, 2015.

Key Financing Events

•

2014~~2014—~~—In March 2014, we completed a private placement of 4,000,000 ordinary shares to institutional investors for gross proceeds of €20.5 million. In July 2014, in connection with our collaboration agreement with Pfizer, we issued 2,786,924 ordinary shares, representing 10% of our then-outstanding ordinary shares, to Pfizer for gross proceeds of €25.8 million. In November 2014, we issued shares in connection with the exercise ofnon-employee warrants for gross proceeds of €13.4 million.

•

2015~~2015—~~—In March 2015, we completed our U.S. initial public offering of ADSs on the Nasdaq for gross of $228 million of gross proceeds, of which we received net proceeds of $209.6 million.

2017—In July 2017, Calyxt completed an initial public offering of its common stock on the Nasdaq, selling an aggregate of 8,050,000 shares of common stock at a price of $8.00 per share (including 1,050,000 shares of common stock pursuant to the exercise by the underwriters of their option to purchase additional shares). Calyxt received net proceeds of approximately $58.0 million, after deducting underwriting discounts and commissions and offering expenses. As part of the IPO, Cellectis purchased 2,500,000 shares of common stock for a value of $20.0 million, which is included in the net proceeds that Calyxt received. Then, the net proceeds amounted to $38 million onat the Group level.

On April 10, 2018, Cellectis closed a ~~consolidated basis.~~follow-on offering of 5,646,000 ADS at a public offering price of $31.00 per ADS resulting in gross proceeds of $175 million. On May 11, 2018, in connection with the exercise by the underwriters of thefollow-on offering of their option to purchase additional shares, Cellectis closed the sale of an additional 500,000 ADS at the public offering price of $31.00 per ADS resulting in additional gross proceeds of $15.5 million.

On May 22, 2018, Calyxt, Inc. completed afollow-on offering of its common stock. Calyxt, Inc. sold an aggregate of 4,057,500 shares of common stock at a price of $15.00 per share, including 457,500 shares of common stock pursuant to the exercise of the underwriters’ option to purchase additional shares. In the aggregate, Calyxt, Inc. received net proceeds from thefollow-on offering and exercise of the overallotment option of approximately $57.0 million, after deducting underwriting discounts and commissions of $3.2 million and offering expenses totaling approximately $0.7 million. As part of thefollow-on offering, Cellectis SA purchased 550,000 shares of common stock for a value of $8.3 million, the proceeds of which are included in the net proceeds of approximately $57.0 million.

Liquidity management

As of December 31, ~~2017,~~2018 we had current financial assets and cash and cash equivalents of ~~$297.0~~$451.9 million comprising cash and cash equivalents of ~~$256.4~~$451.5 million and current financial assets of ~~$40.6~~$0.4 million.

Cash in excess of immediate requirements is invested in accordance with our investment policy, primarily with a view to liquidity and capital preservation. Currently, our cash and cash equivalents are held in bank accounts, money market funds, fixed bank deposits primarily in France. The portion of cash and cash equivalent denominated in U.S. dollars ~~amounted to $233.9~~is $298.1 million as of December 31, ~~2017.~~2018. Current financial assets denominated in U.S. Dollars amounted to ~~$39.7~~$0.4 million as of December 31, ~~2017.~~2018.

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Historical Changes in Cash Flows

The table below summarizes our sources and uses of cash for the years ended December 31, ~~2015,~~ 2016, 2017 and ~~2017:~~2018:


										For the year ended December 31,
	For the year ended December 31,									2016			2017			2018
	2015			2016			2017			$ in thousands
Net cash flows provided by (used in) operating activities		3,591			(32,710	)		(52,327	)		(32,710	)		(52,327	)		(68,137	)
Net cash flows provided by (used in) investing activities		(7,728	)		(53,137	)		1,784			(53,137	)		1,784			35,623
Net cash flows provided by (used in) financing activities		220,591			485			41,266			485			41,266			236,494

Total		216,454			(85,362	)		(9,277	)		(85,362	)		(9,277	)		203,981

Effect of exchange rate changes on cash		(10,743	)		(2,181	)		11,089			(2,181	)		11,089			(8,860	)

Year Ended December 31, 2018

The net cash flows used in operating activities are mainly due to cash payments of $41.8 million to suppliers, wages and social expenses of $19.3 million, rent payments of $4.3 million and $20.3 million of other operating payments and payments to Calyxt suppliers, partially offset by $4.7 million of payments received from Servier and Pfizer/Allogene Therapeutics pursuant to our collaboration agreements, $1.5 million of payments received from licenses, $6.9 million of interest received and $4.5 million of VAT and other taxes reimbursement as well as other variances.

The net cash flows used in investing activities primarily reflects our investments in R&D equipment and building fittings in both the United States and France of $4.9 million included $0.2 million of intangible assets, offset by (i) the proceeds from disposal of $1.3 million included Calyxt’s sale and leaseback agreement transaction for certain equipment for $1.2 million, (ii) the reimbursement of $0.2 million related to the termination of a liquidity contract that we were party to with Natixis Securities and by (iii) the proceeds from current financial assets of $39.0 million.

The net cash flows provided by financing activities mainly reflects (i) the net proceeds after deducting underwriting discounts and commissions and offering expenses of $178.6 million from the Cellectisfollow-on offering, (ii) the net proceeds, after deducting underwriting discounts and commissions and offering expenses and the purchase price paid by Cellectis with respect to our purchase of 550,000 shares of Calyxt common stock purchased by Cellectis in the offering, of $48.8 million, (iii) the exercise of 322,068 Cellectis stock options during the period for $7.5 million, (iv) the exercise of 592,342 Calyxt stock options during the period for $2.4 million, (v) the subscription ofnon-employee warrants for $0.2 million and (vi) the reimbursement of $0.3 million related to the termination of our liquidity contract with Natixis Securities, partially offset by Cellectis’ purchase on June 14, 2018 of 63,175 shares of Calyxt common stock from employees and nonemployees of Calyxt and Cellectis at a price of $19.49 per share (the closing price reported on the Nasdaq Global Market on June 14, 2018) for $1.2 million.

Year Ended December 31, 2017

The net cash flows used in operating activities are mainly due to cash payments of $53.9 million to suppliers, wages and social expenses of $19.8 million and rent payments of $3.7 million, partially offset by $8.4 million of payments received from Servier and Pfizer pursuant to our collaboration agreements and $8.1 million of research tax credit as well as other variances.

The net cash flows used in investing activities primarily reflects, the proceeds from disposal of $7.0 million related to Calyxt’s sale and leaseback agreement, partially offset by our acquisition of $2.6 million of financial current assets at Cellectis and our investments in research and development equipment and intangible assets in both the United States and France of $2.7 million.

The net cash flows provided by financing activities mainly reflects the net proceeds of the Calyxt IPO of $38.1 million, on a consolidated basis, the subscription ofnon-employee warrants for $0.3 million, and exercises of 121,492 BSPCE warrants and ~~31,873~~31 873 stock options by employees during the period for $2.7 million.

Year Ended December 31, 2016

The net cash flows used in operating activities are mainly due to our cash expenditures related to research and development efforts, including the advancement of UCART123, for which an IND was filed in the United States in 2016, plus several advance payments made for manufacturing activities, partially offset by the payments received from Servier and Pfizer pursuant to our collaboration agreements. Cash outflows for 2016 also include $7.0 million of social charges on stock options.

The net cash flows used in investing reflects our use of $10.0 million for the acquisition of land by Calyxt and the building of its greenhouse, and the acquisition of $37.5 million of current financial assets at ~~Cellectis.~~Cellectis S.A.

The net cash flows provided by financing activities includes the exercise ofnon-employees warrants in January 2016 for $0.3 million and the subscription ofnon-employees warrants in September 2016 for $0.3 million partially offset by the decrease of financial lease debt for $0.1 million.

~~Year Ended December 31, 2015~~

We had net cash flows provided by operating activities of continuing operations of $3.6 million for the year ended December 31, 2015, primarily as a result of payments received from Servier and, to a lesser extent, Pfizer. In December 2015, we received an upfront payment of $40.0 million in connection with the amendment to our

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collaboration agreement with Servier. Our cash expenditures during 2015 related to our research and development efforts, including the advancement of UCART19, for which a CTA was filed in the United Kingdom in 2015. Cash outflows for 2015 also include $13.5 million of social charges on free shares and stock options grants.

The net cash flows used in investing activities of continuing operations of $7.8 million for the year ended December 31, 2015 primarily reflects our use of $4.3 million for the acquisition of industrial and laboratory equipment at Cellectis S.A. and Cellectis Inc., and our repurchase for $3.9 million of 25% of the minority shares of Cellectis Bioresearch.

The net cash flows provided by financing activities of continuing operations of $220.6 million for the year ended December 31, 2015 includes the net proceeds from our U.S. initial public offering on the Nasdaq Global Market in March 2015.

Operating capital requirements

To date, we have not generated any revenues from therapeutic or agricultural product sales. We do not know when, or if, we will generate any revenues from product sales. We do not expect to generate significant revenues from product sales unless and until we obtain regulatory approval of and commercialize one of our current or future product candidates. We anticipate that we will continue to generate losses for the foreseeable future, and we expect the losses to increase as we continue the development of, and seek regulatory approvals for, our product candidates, and begin to commercialize any approved products. We are subject to

all risks incident in the development of new gene therapy products, and we may encounter unforeseen expenses, difficulties, complications, delays and other unknown factors that may adversely affect our business. We are also subject to all risks incident in the development of new agricultural products, and we may encounter unforeseen expenses, difficulties, complications, delays and other unknown factors that may adversely affect our business. We also anticipate substantial expenses related to audit, legal, regulatory andtax-related services associated with our public company obligations in the United States and our continued compliance with applicable U.S. exchange listing and SEC requirements. We anticipate that we will need additional funding in connection with our continuing operations, including for the further development of our existing product candidates and to pursue other development activities related to additional product candidates.

We believe our cash and cash equivalents, our cash flow from operations (including payments we expect to receive pursuant to our collaboration agreements) and government funding of research programs will be sufficient to fund our operations ~~into 2020.~~through 2021. However, we may require additional capital for the further development of our existing product candidates and may also need to raise additional funds sooner to pursue other development activities related to additional product candidates.

Until we can generate a sufficient amount of revenues from our products, if ever, we expect to finance a portion of future cash needs through public or private equity or debt offerings. Additional capital may not be available on reasonable terms, if at all. If we are unable to raise additional capital in sufficient amounts or on terms acceptable to us, we may have to significantly delay, scale back or discontinue the development or commercialization of one or more of our product candidates. If we raise additional funds through the issuance of additional debt or equity securities, it could result in dilution to our existing shareholders, increased fixed payment obligations and these securities may have rights senior to those of our ordinary shares. If we incur indebtedness, we could become subject to covenants that would restrict our operations and potentially impair our competitiveness, such as limitations on our ability to incur additional debt, limitations on our ability to acquire, sell or license intellectual property rights and other operating restrictions that could adversely impact our ability to conduct our business. Any of these events could significantly harm our business, financial condition and prospects.

Our assessment of the period of time through which our financial resources will be adequate to support our operations is a forward-looking statement and involves risks and uncertainties, and actual results could vary as a

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result of a number of factors. We have based this estimate on assumptions that may prove to be wrong, and we could use our available capital resources sooner than we currently expect. Our future funding requirements, both near and long-term, will depend on many factors, including, but not limited to:

the initiation, progress, timing, costs and results ofpre-clinical and clinical studies for our product candidates;

the initiation, progress, timing, costs and results of field trials for our agricultural product candidates;

the capacity of manufacturing our products in France and in United States;

the outcome, timing and cost of regulatory approvals by U.S. andnon-U.S. regulatory authorities, including the possibility that regulatory authorities will require that we perform more studies than those that we currently expect;

the ability of our product candidates to progress through clinical development successfully;

the ability of our agricultural product candidates to progress through late stage development successfully, including through field trials;

the cost of filing, prosecuting, defending and enforcing any patent claims and other intellectual property rights;

our need to expand our research and development activities;

our need and ability to hire additional personnel;

our need to implement additional infrastructure and internal systems, including manufacturing processes for our product candidates;

the effect of competing technological and market developments; and

the cost of establishing sales, marketing and distribution capabilities for any products for which we may receive regulatory approval.

If we cannot expand our operations or otherwise capitalize on our business opportunities because we lack sufficient capital, our business, financial condition and results of operations could be materially adversely affected.

C.	Research and Development, Patents and Licenses, etc.

Our research and development teams utilize our deep expertise to contribute to the growth of our business. As of December 31, ~~2017,~~2018, we had 97113 employees engaged in research and development activities. In the years ended December 31, ~~2015,~~ 2016, 2017 and ~~2017~~2018 we spent ~~$58.2 million,~~ $78.5 million, $79.2 million and ~~$79.2~~$76.6 million, respectively, on research and development. For a discussion of our research and development activities, see “Item 4.B—Business Overview” and “Item 5.A—Operating Results.”

D.	Trend Information

For a discussion of trends, see “Item 4.B—Business Overview,” “Item 5.A—Operating Results” and “Item 5.B—Liquidity and Capital Resources.” Other than as disclosed in these sections, we are not aware of any trends, uncertainties, demands, commitments or events since December 31, 2015 that are reasonably likely to have a material adverse effect on our revenues, income, profitability, liquidity or capital resources, or that would cause the disclosed financial information to be not necessarily indicative of future operating results or financial condition.

E.	Off-Balance Sheet Arrangements.

We entered into ~~(i) financial derivative instruments agreements to minimize impacts from exchange rate fluctuations and (ii)~~ seed and grain production agreements with settlement value based on commodity market future pricing. Otherwise, we do not have anyoff-balance sheet arrangements as defined under SEC rules.

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F.	Tabular Disclosure of Contractual Obligations

The following table discloses aggregate information about material contractual obligations and periods in which payments were due as of December 31, ~~2017.~~2018. Future events could cause actual payments to differ from these estimates.


As of December 31, 2017	Total		Less than 1 year		1 - 3 years		3 - 5 years		More than 5 years
As of December 31, 2018											Total		Less than 1 year		1 - 3 years		3 - 5 years		More than 5 years
	$ in thousands										$ in thousands
Sale and lease-back agreement		30,114		1,076		2,689		2,676		23,673		31,668		1,858		3,748		3,154		22,908
Facility lease agreements		13,416		3,110		5,939		1,670		2,697		28,230		2,290		7,814		8,193		9,933
License agreements		18,413		1,238		2,476		2,476		12,223		18,093		1,237		2,474		2,474		11,907
Manufacturing agreements		7,679		7,679		—		—		—		10,293		10,293		—		—		—
Other agreements		1,702		1,702		—		—		—		12,356		7,754		4,602		—		—

Total contractual obligations		71,324		14,805		11,104		6,821		38,593		100,641		23,433		18,638		13,822		44,749

The commitment amounts in the table above are associated with contracts that are enforceable and legally binding and that specify all significant terms, including fixed or minimum services to be used, fixed, minimum or variable price provisions, and the approximate timing of the actions under the contracts. The table does not include obligations under agreements that we can cancel without a significant penalty. We have collaboration agreements whereby we are obligated to pay royalties and milestones based on future events that are uncertain and therefore they are not included in the table above.

We also provided Letters of Credit to the landlords of our ~~facility~~facilities in New ~~York and Calyxt’s facility in New Brighton.~~York.

For further detail regarding license and manufacturing agreement, please see Note 18 – Commitments of our consolidated financial statements.

G.	Safe Harbor

This Annual Report contains forward-looking statements within the meaning of Section 27A of the Securities Act and Section 21E of the Exchange Act and as defined in the Private Securities Litigation Reform Act of 1995. See “Special Note Regarding Forward-Looking Statements.”

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ITEM 6.

DIRECTORS, SENIOR MANAGEMENT AND EMPLOYEES

A.	Directors and Senior Management

The following table sets forth information regarding our executive officers and directors as of March ~~13, 2018.~~11, 2019. Unless otherwise stated, the address for our executive officers and directors is 8, rue de la Croix Jarry, 75013 Paris, France.


Name		Age			Position(s)
*Executive Officers:*
André Choulika, Ph.D.			5354			Chairman of the Board, Chief Executive Officer, andCo-Founder
~~Julia Berretta, Ph.D.~~	Elsy Boglioli		37			~~Vice President Business Development~~ Chief Operating Officer
~~Elsy Boglioli (1)~~ Philippe Duchateau, Ph.D.			3656			Chief ~~Operating~~Scientific Officer
~~Stéphane Depil, M.D. Ph.D.~~ Eric Dutang			4445			~~Senior Vice President Research and Development and~~ Chief ~~Medical~~Financial Officer
~~Philippe Duchateau, Ph.D.~~ Stephan Reynier			5550			Chief ~~Scientific~~Regulatory Officer
~~Eric Dutang~~
David Sourdive, Ph.D.			4452			~~Chief Financial Officer~~
~~Stephan Reynier~~			49			~~Chief Regulatory Officer~~
~~David Sourdive, Ph.D.~~			51			Director, Executive Vice President, Technical Operations andCo-Founder
Marie-Bleuenn Terrier			3637			General Counsel
~~Federico Tripodi~~					41		~~Chief Executive Officer, Calyxt, Inc.~~
*Non-Employee Directors:*
Laurent Arthaud			5556			Director
Pierre Bastid			6364			Director
Rainer Boehm,			57			~~Director~~
~~Alain Godard~~			72			~~Director~~
~~Hervé Hoppenot~~	M.D.		58			Director
~~Jean-Marie Messier~~ Alain Godard			6173			Director
Hervé Hoppenot		59		Director
Annick Schwebig, M.D.			6768			Director

~~(1)~~

~~In connection with the retirement of Dr. Mathieu Simon, Ms. Boglioli was appointed Chief Operating Officer on March 13, 2018.~~

André Choulika, Ph.D., is one of the founders of Cellectis and served as Chief Executive Officer since the company’s inception in 1999. He is Chairman of the Board of Directors since 2011 and President and Chairman of the board of directors of Calyxt, Inc. since August 2010. From 1997 to 1999, Dr. Choulika worked as a post-doctoral fellow in the Division of Molecular Medicine at Boston Children’s Hospital, where he was one of the inventors of nuclease-based genome editing technologies and a pioneer in the analysis and use of meganucleases to modify complex genomes. After receiving his Ph.D. in molecular virology from the University of Paris VI (Pierre et Marie Curie), he completed a research fellowship in the Harvard Medical School Department of Genetics. His management training is from HEC (Challenge +). Based on Dr. Choulika’s deep knowledge of the company and scientific experience, we believe Dr. Choulika has the appropriate set of skills to serve as our chief executive officer and a member of our board of directors.

~~Julia Berretta~~Elsy Boglioli~~, Ph.D.~~, joined Cellectis in 2010 in the scientific alliance and business development department. She has served as VP Business Development and Strategic Planning since 2014. Prior to joining Cellectis, she worked as a researcher at the CNRS in~~Gif-sur-Yvette.~~ ~~Julia Berretta received her Ph.D. in molecular biology from the Université Paris XI, and holds a specialized Master’s Degree in innovation management from Neoma Business School.~~

~~Elsy Boglioli~~ ~~joined Cellectis in~~ December ~~2017. Commencing in 2018, she serves as Chief Operating Officer. Prior to assuming the COO role, she served~~2017 as Executive Vice President, Strategy and Corporate

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Development. She was named Chief Operating Officer in March 2018. Prior to joining Cellectis, she worked at Boston Consulting Group (BCG) where she served as Partner and Managing Director, and leader of BCG’s biotech-focused business in Europe. At BCG, she was also a member of the global biopharma leadership team and global strategy practice management team and served as regional head of the strategy practice for Europe. Ms. Boglioli graduated from Ecole Polytechnique in Paris, France and holds a master’s degree in economy and management from Pompeu Fabra University in Barcelona, Spain.

~~Stéphane Depil~~, M.D., Ph.D., joined Cellectis in December 2017. He serves as Senior Vice President, Research & Development and Chief Medical Officer. In addition to his role at Cellectis, Stéphane Depil is an onco-hematologist at Léon Bérard Cancer Center and an adjunct Professor at University Claude Bernard Lyon 1, France. Prof. Depil is a Board-certified physician in hematology and has over 15 years of experience in oncology clinical development, both in hospital / university and pharmaceutical companies. He obtained a PhD in immunology at Institut de Biologie de Lille after working on a project concerning cancer vaccination. Prior to joining the Léon Bérard Cancer Center & Cancer Research Center of Lyon, France, as Medical Director of the Cancer Immunotherapy Program, he served as Chief Executive Officer at Netris Pharma, where he was responsible for the management of an oncology startup and preclinical development of a~~first-in-class~~ monoclonal antibody in Phase I. Prior to Netris, Stéphane Depil worked at Servier for 8 years in a variety of roles, including Director of Oncology Research and Development, where he managed 20 programs: 5 in the clinic, 7 at late preclinical stages, and 8 at early preclinical stages. He also directly supervised the evaluation of over 100 licensing opportunities.

Philippe Duchateau, Ph.D., joined Cellectis in 2001 to pioneer the field of genome engineering and has served as Chief Scientific Officer since 2012. After receiving his PhD in 1993 in biochemistry and molecular biology at the Institut Pasteur (Lille, France), he completed a research fellowship from 1993 to 2001 at the University of California, San Francisco (United States) within the Cardiovascular Research Institute. He isco-inventor of numerous patents in the field of nucleases and genome engineering andco-authors on more than 50 scientific publications andco-editor of one book entitled “Site-directed Insertion of Transgenes.” As head of Cellectis’s Research department since 2004, he helped to the development of the Cellectis Technologies.

Eric Dutang joined us as Deputy Chief Financial Officer in May 2015 and was appointed Chief Financial Officer in 2016. Eric began his career as financial auditor with KPMG, first in Paris for five years and then in New York for two years. He worked for listed companies in France and the United States such as Vivendi, Veolia Environnement or Cablevision. He then became a member of the transactions and advisory teams in Paris for seven years where he carried out acquisitions / disposals for listed companies and private equity funds. After serving at KPMG, he worked on international business developments for French public listed groups, including Air Liquide and Thales. Eric holds a Master of Finance and Executive MBA from HEC Paris (France)/Babson Massachusetts (USA).

Stephan Reynier,MSc, joined Cellectis in April 2011. He serves as Chief Regulatory and Compliance Officer after holding the position of Head of Programs at Ectycell, a former subsidiary of Cellectis, from April 2011 to 2014 with the mission of managing and coordinating internal and external collaborative programs. As Chief Regulatory and Compliance Officer, Mr. Reynier is in charge of ensuring a speedy and successful development of the UCART product family by establishing close interactions with regulatory agencies such as EMA and FDA, while securing compliance to applicable regulations, regulatory guidelines and quality assurance standards. Mr. Reynier has extensive experience, from his previous positions as Senior Director at Voisin Consulting Life Sciences and European Associate Director Medical Affairs at Gilead Sciences, in the design and implementation of regulatory strategies for the development of drugs and biologics, with a strong focus on cell and gene therapy. Mr Reynier graduated as Agro-Engineer in France and received a Master of Science in Chemical Engineering from the University of Toronto, Canada.

David Sourdive,Ph.D., is aco-founder of Cellectis and has held the position of Executive Vice President, Technical Operations since 2017. Prior to that date, Dr. Sourdive served as Executive Vice President, Corporate

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Development since 2008. Dr. Sourdive has also been a member of our board of directors since 2000. From 2009 to 2012, he served as President of Ectycell SAS, and since 2012, he has served on its supervisory committee. Since February 2014, Dr. Sourdive has also served on the board of directors of Mediterranean Institute for Life Sciences. Since June 2015, he has served on the board of directors of Eukarÿs SAS, and since 2017 Dr. Sourdive serves on the Board of Omics SAS. Since December 2018, he has served on the board of directors of Enobraq SAS. He previously served on the boards of directors of Cellectis AB, Medicen Paris Region and Seine Saint Denis Avenir. From 1998 to 2000, he directed the biotechnologies laboratory of the Centre d’Etudes du Bouchet for the French Ministry of Defense. From 1997 to 1998, Dr. Sourdive worked at one of the leading laboratories in viral immunology at Emory University in Atlanta, Georgia. His work there was focused on immunologicalT-cell memory. Dr. Sourdive graduated from the École Polytechnique and received his PhD in molecular virology at the Institut Pasteur. He also has management training from the HEC (Challenge +). We believe Dr. Sourdive’s extensive experience in business and the biotechnology industry qualifies him to serve on our board of directors.

Marie-Bleuenn Terrier joined Cellectis as Legal Counsel in 2008, and was appointed General Counsel in 2013. Prior to joining Cellectis, she worked as Legal Counsel for Pfizer from 2004 to 2006, and for Boehringer-Ingelheim from 2006 to 2008. Marie-Bleuenn Terrier has also served as Secretary of our board of directors since 2015. She holds a Master’s degree in Law from the Panthéon La Sorbonne University in Paris.

~~Federico Tripodi~~was appointed CEO of Calyxt in May 2016. He holds a Master of Business Administration degree from Washington University’s Olin Business School, as well as an agronomic engineering degree from Buenos Aires University, and has gathered extensive experience in agricultural R&D and product development during his nearly~~two-decade~~ career in the agricultural biotechnology and seeds industry. Prior to joining Calyxt, he worked as General Manager for Monsanto Company’s Sugarcane Division in Brazil for three years. He held other roles for Monsanto in Saint Louis, Mo., spanning Corporate Strategy (2011-2013),~~Omega-3~~ Program Lead (2009-2011), Oilseeds Global Quality Management Lead (2008-2009) and multiple other roles that involved managing multidisciplinary research teams in the technology organization between 2001 and 2008. During his tenure at Monsanto, Mr. Tripodi led or participated with early discovery and late commercialization phase product launches across the Americas, which included biotechnology consumer traits (improved composition soybean oils) and farmer traits (high yield, drought tolerance, insect protection and herbicide tolerance). Mr. Tripodi started his career in Argentina in 1998 in field research of biotechnology traits and chemistry formulations until he moved to Saint Louis in 2001. Mr. Tripodi also has experience as a director of a startup and served on the board of directors for a~~not-for~~ ~~profit.~~

Laurent Arthaud has served as a member of our board of directors since October 28, 2011. Mr. Arthaud has been the Managing Director of Life Sciences and Ecotechnologies for Bpifrance Investissement (formerly CDC Enterprises, a subsidiary of Caisse des Dépôts) since 2012. He currently serves on the boards of directors of Calyxt, Inc., Kurma Life Sciences Partners, ~~TxCell,~~ Adocia, Sparinvision, Aledia and ~~Sparinvision. From 2006 to 2016, he~~Ribogenics, Inc.. He served on the board of directors of TxCell from 2012 to 2018, on the board of directors of Emertee ~~Gestion.~~Gestion from 2006 to 2016, and on the board of directors of Scynexis, Inc. from 200 to 2015. From 2006 to 2012, Mr. Arthaud held the position of Deputy CEO at CDC Entreprises. Since 2009 Mr. Arthaud has also directed InnoBio, an investment fund managed by Bpifrance Investissement as part of the FSI France Investissement program. From 1999 to 2004 he served as Vice President of Aventis Capital, an investment subsidiary of the pharmaceuticals group Aventis, and as President of Pharmavent Partners from 2004 to 2006. Mr. Arthaud is a graduate of the École Polytechnique and the École Nationale de Statistique et d’Administration Économique. We believe Mr. Arthaud’s extensive investment experience in the biotechnology industry qualifies him to serve as a member on our board of directors.

Pierre Bastid has served as a member of Cellectis’ board of directors since 2011. Mr. Bastid has 25 years experience in turning around, developing and running technology businesses in Asia, Europe and the United States. In addition to Cellectis, Mr. Bastid is currently serving on the board of directors of Pharnext (a biotechnology company), ~~Hougou Finance S.A.~~ and ~~Hougou Développement S.A. (his own~~DCTV CenterNew-York, and of a series of his owned investment ~~company), ZAKA S.A.~~ and ~~GRID (his own Paris and New York-based real estate companies), Shango S.A. (his own~~ private equity ~~company), EVOK (his own hotel group), Nepteam S.A.S. (a shipbuilding company), Louise342-344~~ ~~S.A., Hebioso S.A., La Chartreuse B S.C., Batuque Hotelaria e Turismo S.A. and Casino Royal S.A.~~

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companies. Mr. Bastid also advises a number of investment and private equity firms. Mr. Bastid is a trustee of the Juilliard School of Music and othernon-profit organizations based in the United States. We believe Mr. Bastid’s extensive business experience qualifies him to serve as a member on our board of directors.

Rainer Boehm,M.D., has served as a member of Cellectis’ board of directors since 2017. In addition, Mr. Boehm is currently serving on the board of directors of Humanigen, Inc. and Nordic Nanovector ASA since 2018. Mr. Boehm spent 29 years at Novartis, working locally, regionally and globally in various Senior Management roles, after building his career in Marketing & Sales and Medical Affairs. He led all emerging markets regions as well as the United States and Canada, either for Oncology or the Pharmaceuticals division. His most recent assignments were Chief Commercial and Medical Affairs Officer globally for Novartis Pharma, as well as ad interim CEO and Division Head Pharma. Rainer launched and oversaw the commercialization of many brands during his career, amongst them Femara, Zometa and Glivec, as well as Cosentyx and Entresto. ~~His broad interests include leadership development, coaching and mentoring, access to medicines and new business models including solutions to leverage digital medicines and real world evidence.~~ Rainer has a medical degree from the University of Ulm in Germany, and a Master of Business Administration from Schiller University in France. We believe Mr. Boehm’s extensive business and medical experience in pharmaceutical industry qualifies him to serve as a member on our board of directors.

Alain Godard has served as a member of Cellectis’ board of directors since 2007. In addition, Mr. Godard is currently serving on the board of directors of Calyxt, Inc. He is a graduate of the Ecole Nationale Supérieure Agronomique de Toulouse and began his agronomy career in 1967 in Africa as a researcher at theInstitut de Recherche pour les Huiles et Oléagineux. He joined the French chemical groupRhône-Poulenc in 1975 where he held various management positions in France and abroad before becoming CEO of the agrochemical subsidiary in 1991. In 1999 he was directly involved in the merger ofRhône-Poulenc and Hoechst to create Aventis and was appointed CEO of the Aventis CropScience subsidiary with a significant involvement in seeds and agricultural biotechnology. He left Aventis in 2002 to create a consulting company, SARL Godard & Co., specialized in agriculture and biotechnology, where he has served as Chief Executive Officer since 2009. Until 2016, Mr. Godard also served on the board of directors of Fermentalg S.A. We believe Mr. Godard’s leadership and management expertise in the plant biotechnology field qualifies him to serve as a member of our board of directors.

Hervé Hoppenothas served as a member of Cellectis’ board of directors since 2017. He serves as President and Chief Executive Officer of Incyte Corporation since 2014, and was appointed Chairman of the Board of Directors in 2015. Incyte is one of the fastest growing biopharmaceutical companies in the U.S. Prior to joining Incyte, Mr. Hoppenot was the President of Novartis Oncology, which included $11 billion in global sales, the largest oncology pipeline in the industry and 8000 employees in 50 countries. Prior to joining Novartis in 2003, Mr. Hoppenot started his career in 1983 with Rhone Poulenc, later known as Aventis, where he served in several senior roles of increasing responsibility, including Vice President of Oncology and Head of the US Oncology business unit. He and his family are dual citizens of France and the United States, having moved to the U.S. in 1991. We believe Mr. Hoppenot’s business experience in the biotechnology industry qualifies him to serve as a member of our board of directors.

~~Jean-Marie Messier~~ ~~has served as a member of our board of directors since May 18, 2015. He isco-founder~~ and head of Messier Maris & Associés, an international investment banking firm. Mr. Messier has served on the board of directors of Rentabiliweb Group since May 2011. After graduating from the French university, Ecole Polytechnique, Mr. Messier attended the Ecole Nationale d’Administration, which trains civil servants. He became Managing Partner at Lazard Frères in 1988, a position he held for six years. Prior to this, he was responsible for the French Government’s Privatization plan. As such, he privatized companies among which Alcatel, Lagardère, Saint Gobain, Suez and Société Générale, now all part of the top twenty French corporations. Mr. Messier served as President of Vivendi Universal from 1994 to 2002. During these years, he founded the mobile firm Cegetel and turned Vivendi into a conglomerate focused on two core activities: utilities (water, power and transport) and communications (pay TV, telecoms and internet), selling off assets in other areas. Mr. Messier has been managing his own “boutique” since 2003, Messier Maris & Associés, an investment bank headquartered in Paris and New York representing more than 50 professionals.

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Annick Schwebig, M.D., has served as a member of our board of directors since October 28, 2011. In 2000, she founded the French subsidiary of Actelion, of which she is a Senior Advisor. She formerly served as the General Manager of Actelion from 2000 to 2015. Actelion is a ~~biopharmaceuticals~~biopharmaceutical company specializing in innovative treatments to serve unmet medical needs. She is also a director of Inventiva Pharma, a biopharmaceutical company, and ~~Inserm-Transfert SA, the private subsidiary of the French National Institute of Health and Medical Research.~~B Cell Design, a biotechnologies company. A graduate of the University of Paris medical school, Dr. Schwebig worked as a senior manager at the biopharmaceuticals company Bristol-Myers Squibb for 17 years from 1983 to 2000. We believe Dr. Schwebig’s extensive experience in the biopharmaceutical industry qualifies her to serve as a member on our board of directors.

Family Relationships

There are no family relationships among any of our executive officers or directors.

B.	Compensation

Compensation of Directors and Executive Officers

The aggregate cash compensation paid and benefits in kind granted by us to our current executive officers and directors, for the year ended December 31, ~~2017,~~2018, was ~~$3.6~~$5.7 million. For the year ended December 31, ~~2017, 715,000~~2018, 50,000 stock options with an exercise price of ~~€22.57~~€24.80 per ordinary share were issued to ~~directors and~~ executive officers as compensation under the ~~2017~~2018 Stock Option Plan and 40,000 free shares were issued to executive officers as compensation under the Second Free Shares 2018 Plan. The total amount set aside or accrued to provide pension, retirement or similar benefits was ~~$35,285~~$38,135 for the year ended December 31, ~~2017.~~2018.

In connection with the vesting on June 14, 2018 of Restricted Stock Units (RSU) granted to certain employees andnon-employees of Calyxt Inc. and Cellectis SA, Cellectis SA purchased 2,352 common shares of Calyxt and 1,470 Calyxt shares at a price of $19.49 per share (the closing price reported on the Nasdaq Global Market on June 14, 2018) directly from Mr. André Choulika and Mr. Jean Marie Messier respectively in connection with share purchase transactions dated June 13 2018.

Service Agreements

Mr. Godard, a member of our board of directors, entered into a service agreement with us and provided consultancy services in the area of global development strategy. We paid ~~€34~~€56 thousand (or ~~$38~~$66 thousand) in compensation for those services in fiscal year ~~2017.~~2018.

Change of Control Benefits

We seek to balance the potential costs of change of control provisions with the costs that would arise from fear of job loss and other distractions that may result from potential, rumored or actual changes of control. As a result, after careful evaluation of the implications and economics of a change of control plan, on September 4, 2014, our board of directors adopted a change of control plan. As amended on December 11, 2014, the change of control plan provides benefits for certain of our executive officers and several other senior employees of our company.

Pursuant to the change of control plan, the severance package shall be paid if, within the36-month period following a change of control of our company, one of the following events occurs:

•

non-renewal or dismissal other than for gross misconduct (faute lourde) of the employees or executives concerned; and

for Drs. Choulika and Sourdive only, resignation as a result of a significant reduction of their duties or compensation, or end ornon-renewal of their corporate appointments.

The severance package shall be equal to 24 months of compensation increased by an amount equal to the maximum target bonus to which the employees or executives concerned may be entitled for the year of their departure (or for Dr. Choulika only, two times such target bonus), or, in the absence of such a target bonus, 1.5 times the last annual bonus paid to them during the 12 months prior to their departure.

The severance package shall be in addition to any legal and conventional severance payments owed to the employees or executives concerned.

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A “change of control” is defined by reference to ArticleL.233-3 of the French Commercial Code, which provides that one or more persons acting alone or in concert are considered to control a company if (1) they have direct or indirect ownership of a majority of the voting rights or a proportion of the voting rights allowing de facto control of the decisions made by the shareholders, provided that such control is presumed if no shareholder holds a greater proportion of the voting rights; or (2) they have the power to appoint or dismiss a majority of the board of directors

Limitations on Liability and Indemnification Matters

Under French law, provisions ofBy-laws that limit the liability of directors and officers are prohibited. However, French law allowssociétés anonymes to contract for and maintain liability insurance against civil liabilities incurred by any of their directors and officers involved in a third-party action, provided that they acted in good faith and within their capacities as directors or officers of the company. Criminal liability cannot be indemnified under French law, whether directly by the company or through liability insurance.

We expect to maintain customary liability insurance coverage for our directors and executive officers, including insurance against liability under the Securities Act. With certain exceptions and subject to limitations on indemnification under French law, ~~these agreements~~

this insurance coverage will provide for indemnification for damages and expenses including, among other things, attorneys’ fees, judgments, fines and settlement amounts incurred by any of these individuals in any action or proceeding arising out of his or her actions in that capacity. We believe that this insurance ~~and these agreements are~~coverage is necessary to attract qualified directors and executive officers.

~~These contractual indemnification agreements~~This insurance coverage may discourage shareholders from bringing a lawsuit against our directors and executive officers for breach of their fiduciary duty. ~~These provisions~~It also may have the effect of reducing the likelihood of derivative litigation against directors and executive officers, even though such an action, if successful, might otherwise benefit us and our shareholders. Furthermore, a shareholder’s investment may be adversely affected to the extent we pay the costs of settlement and damage awards against directors and officers pursuant to ~~these~~this insurance ~~agreements.~~coverage.

Certain of ournon-employee directors may, through their relationships with their employers or partnerships, be insured against certain liabilities in their capacity as members of our board of directors.

Equity Incentives

We believe that our ability to grant equity awards is a valuable and necessary compensation tool that allows us to attract and retain the best available personnel for positions of substantial responsibility, provides additional incentives to employees and promotes the success of our business. In accordance with French corporate law and tax considerations, we have granted several different equity incentive instruments to our directors, executive officers, employees and other service providers. These are:

•

employee warrants (otherwise known asbons de souscription de parts de créateur d’entreprise or BSPCE), granted only to employees of Cellectis;

•

non-employee warrants (otherwise known asbons de souscription d’actions or BSA), granted only tonon-employee directors and other service providers or consultants not eligible for employee warrants;

•

restricted, or free, shares (otherwise known asactions gratuites); and

•

stock options (otherwise known asoptions de souscription d’actions).

Our board of directors’ authority to grant these equity incentive instruments and the aggregate number of shares authorized to be granted under these instruments must be approved by atwo-thirds majority of the shares held by our shareholders present, represented or voting by mail at the relevant extraordinary shareholders’ meeting. Such extraordinary general meeting shall determine the aggregate amount of equity incentive

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instruments to be granted and the period during which such authorization may be used by our board of directors, which cannot exceed 18 months fornon-employee warrants and employee warrants and 38 months for stock option and restricted (free) shares, in each case beginning from the date of the applicable shareholders’ approval. The authority of our board of directors to grant equity incentives may be extended or increased only by extraordinary shareholders’ meetings. As a result, we typically request that our shareholders authorize new pools of equity incentive instruments at every annual shareholders’ meeting.

Employee warrants andnon-employee warrants are usually granted under similar terms. They expire ten years after the date of grant if not exercised earlier according to their vesting schedule (see below). In general, employee warrants (BSPCE) andnon-employee warrants (BSA) no longer continue to vest following termination of the employment, office or service of the holder and all vested shares must be exercised within post-termination exercise periods set forth in the applicable equity award grant documents. In the event of certain changes in our share capital structure, such as a consolidation or share split or dividend, French law and applicable equity award grant documentation provide for appropriate adjustments of the numbers of shares issuable and/or the exercise price of the outstanding warrants or share options.

Employee Warrants (BSPCE)

Employee warrants were granted only to employees of Cellectis who are French tax residents, since these employee warrants carry favorable tax and social security treatment for French tax residents. Employee warrants may also be granted to corporate officers of the company having an employee tax status (chairman, general manager or deputy general manager). Similar to stock options, they entitle a holder to exercise the warrant for the underlying vested shares at an exercise price per share determined by our board of directors equal to the higher of (1) the fair market value of an ordinary share on the date of grant and (2) if the company has carried out a capital increase within six months prior to the attribution of employee warrants, the issue price of such capital increase.

Employee warrants may only be issued by growth companies meeting certain criteria, which we no longer meet. Most significantly, the issuer must have been registered for less than 15 years and 25% of the issuer’s share capital must have been continuously held since the company’s formation by natural persons or by holding companies, of which 75% of such holding company’s share capital is held by natural persons. The calculation of such threshold does not include venture capital mutual

investment fund (fonds commun de placement à risques), specialized professional funds (fonds professionnels spécialisés), private equity funds (fonds professionnels de capital investissement), local investment funds (fonds d’investissement de proximité) and innovation-focused mutual funds (fonds commun de placement dans l’innovation).

We are no longer eligible to issue employee warrants since we no longer satisfy the legal conditions necessary to issue such employee warrants.

Our outstanding employee warrants were generally granted (1) either subject to a three-year vesting schedule under whichone-third (1/3) of the employee warrants vest upon the first anniversary of grant andone-third (1/3) at the expiration of each year thereafter, subject to continued service, or (2) subject to a five-year vesting schedule under which 40% of the employee warrants vest upon the second anniversary of grant and 20% at the expiration of each year thereafter, subject to continued service. In each case, any warrant which is not exercised before the tenth anniversary of the date of grant will automatically lapse. Some of our employee warrants provide that in the event of a change in control, as defined in the relevant grant documents, unvested warrants will automatically vest in full.

The term of each employee warrant is 10 years from the date of grant or, in the case of death or disability of the beneficiary during suchten-year period, 6 or 9 months respectively from the death or disability of the beneficiary. An employee warrant shall remain exercisable for three months following a beneficiary’s termination of continuous status with the company.

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Employee warrants are not transferable and may not be sold, pledged, assigned, hypothecated, transferred or disposed of in any manner other than by will or by laws of descent or distribution and may be exercised, during the lifetime of the warrant holder, only by the warrant holder.

As of December 31, ~~2017, 21,569~~2018, 19,702 employee warrants exercisable for an aggregate of ~~22,403~~19,702 ordinary shares at a weighted average exercise price of ~~€7.81~~€7.97 per share, were outstanding.

Non-Employee Warrants (BSA)

Non-employee warrants are granted by our board of directors to third-party service providers, consultants and directors who are not eligible for employee warrants. In addition to any exercise price payable by a holder upon the exercise of anynon-employee warrant,non-employee warrants need to be subscribed for at a price at least equal to five percent (5%) of the average price for a company share weighted by volume on the market or markets on which the company shares are listed during the five (5) trading days prior to the date of the grant of saidnon-employee warrant by the board of directors (rounded up to the next euro cent, if necessary).

Pursuant to delegations granted at our annual shareholders’ meeting, our board of directors determines the recipients, dates of grant and exercise price ofnon-employee warrants, the number ofnon-employee warrants to be granted and the terms and conditions thereof, including their vesting schedule. The term of eachnon-employee warrant is generally 10 years from the date of grant.

Ournon-employee warrants are generally granted subject to a three-year vesting, subject to continued service.

As of December 31, ~~2017, 1,079,400~~2018, 899,225non-employee warrants exercisable for an aggregate of ~~1,079,400~~899,225 ordinary shares at a weighted average exercise price of ~~€27.62~~€28.32 per share, were outstanding, all of which are held by certain of our directors and some of our consultants and exercisable at the date hereof.

Free Shares

Under our 2012, 2013, 2014, 2015, 2018 and ~~2015~~Second 2018 Free Share Plans, we have granted free shares to certain of our employees and officers. Our current plan, the ~~2015~~Second 2018 Free Share Plan, was adopted by our board of directors on ~~May 18, 2015.~~August 1, 2018.

Free shares may be granted to any individual employed by us or by any affiliated company. Free shares may also be granted to our Chairman, and our Chief Executive Officer. However, no free share may be granted to a beneficiary holding more than 10% of our share capital or to a beneficiary who would hold more than 10% of our share capital as a result of such grant.

Our board of directors has the authority to administer the 2012, 2013, 2014, 2015, 2018 and ~~2015~~Second 2018 Free Share Plans. Subject to the terms of these Free Share Plans, our board of directors determines the recipients, the dates of grant, the number of free shares to be granted and the terms and conditions of the free shares, including the length of their vesting period (starting on the grant date, during which the beneficiary holds a right to acquire shares for free but has not yet acquired any shares) and

holding period (starting when the shares are issued and definitively acquired but may not be transferred by the recipient) within the limits determined by the shareholders. Our shareholders have determined that the vesting period must be at least two years from the date of grant and the holding period must be two years from the end of the vesting period, with no holding period applicable to beneficiaries for whom the vesting period was four years or longer for the 2012, 2013, 2014 and 2015 Free Shares Plans. For the 2018 Free Share Plans, our shareholders have determined that the vesting period must be at least one year from the date of grant and the holding period must be one year from the end of the vesting period, with no holding period applicable to beneficiaries for whom the vesting period was two years or longer.

The board of directors has the authority to modify awards outstanding under our Free Share Plans, subject to the consent of the beneficiary for any modification adverse to such beneficiary. For example, the board has the authority to release a beneficiary from the continued service condition during the vesting period after the termination of the employment.

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The free shares granted under the Free Share Plans will be definitively acquired at the end of the vesting period as set by our board of directors subject to continued service during the vesting period, except if the board releases a given beneficiary from this condition upon termination of his/her employment contract. At the end of the vesting period, the beneficiary will be the owner of the shares. However, the shares may not be sold, transferred or pledged during the holding period. In the event of disability before the end of the vesting period, the free shares shall be definitively acquired by the beneficiary on the date of disability. In the event the beneficiary dies during the vesting period, the free shares shall be definitively acquired at the date of the request of allocation made by his or her beneficiaries in the framework of the inheritance provided that such request is made within six months from the date of death.

15,600 free shares granted in 2015 under the 2015 Free Share Plan will be acquired on May 18, 2019 for theNon-French residents subject to the continued service of the beneficiaries.

Stock Options

On March 24, 2015, according to the authorization granted by our shareholders meeting held on February 16, 2015, our board of directors adopted the 2015 Stock Option Plan, on October 28, 2016, according to the authorization granted by our shareholders meeting held on May 17, 2018, our board of directors adopted the 2016 Stock Option Plan, on October 11, 2017, according to the authorization granted by our shareholders meeting held on June 26, 2017, our board of directors adopted our ~~2015~~2017 Stock Option ~~Plan, which will expire on April 16, 2018~~Plan; and on ~~October 28, 2016,~~August 1, 2018, according to the authorization granted by our shareholders meeting held on June 26, 2018, our board of directors adopted our ~~2016~~current 2018 Stock ~~Option Plan which will expire on July 18, 2019.~~Options Plan. The 2015 Stock Option Plan, the 2016 Stock Option Plan, ~~as amended~~the 2017 Stock Option Plan and the ~~2017~~2018 Stock Option Plan (collectively, the “Stock Option Plans”) follow the same rules. Stock Options issued pursuant to the Stock Option Plans provide the holder with the right to purchase a specified number of ordinary shares from the Company at a fixed exercise price payable at the time the Stock Option is exercised, as determined by our board of directors. The Stock Option Plans generally provides that the exercise price for any Stock Option will be no less than ninety-five percent (95%) of the average selling prices of a share at close of trading on said market quoted during the twenty trading days immediately preceding the day of the board of directors decision to ~~allocate~~grant the options. The maximum number of ordinary shares, which may be subject to stock options issued is ~~7,354,930 ordinary shares~~8,938,106, provided that our board of directors may decide of new grant of options only under ~~the 2015~~our current 2018 Stock Option Plan ~~is 3,217,861~~with a maximum of 2,890,546 additional ordinary ~~shares under the 2016 Stock Option Plan and is 3,541,547 ordinary shares under the 2017 Stock Option Plan.~~shares. Incentive Stock Options andNon-qualified stock options may be granted under the Stock Option Plans.

Stock Options may be granted to any individual employed by us or by any affiliated company. Stock Options may also be granted to our Chairman, our general manager and to our deputy general managers. Incentive Stock Options granted to owners of shares possessing 10% or more of the total voting power in the Company will be subject to limitations on their exercise price and term.

Our board of directors has the authority to administer and interpret the Stock Option Plans. Subject to the terms of the Stock Option Plans, our board of directors determines the recipients, the dates of grant, the exercise price of the stock options, the number of stock options to be granted and the terms and conditions of the stock options, including the length of their vesting period. Our board of directors is not required to grant stock options with vesting and exercise terms that are the same for every participant. The term of each stock option granted under the Stock Option Plans will generally be 10 years from the date of grant. Further, Stock Options will generally terminate on the earlier of when the beneficiary ceases to be an employee or the Company or upon certain transactions involving the Company.

The board of directors has the authority to modify awards outstanding under our Stock Option Plans, subject to the written consent of the beneficiary for any modification adverse to such beneficiary. For example, the board has the authority to extend a post-termination exercise period.

Stock Options granted under the Stock Option plans generally may not be sold, transferred or pledged in any manner other than by will or by the laws of descent or distribution. In the event of disability, unless otherwise resolved by our board of directors, the beneficiary’s right to exercise the vested portion of his or her option generally terminates six months after the last day of such

beneficiary’s service, but in any event no later than the expiration of the maximum term of the applicable stock options. In the event the beneficiary dies during the

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vesting period, then, unless otherwise resolved by our board of directors, the beneficiary’s estate or any recipient by inheritance or bequest may exercise any vested portion within the six months following the date of death, but in any event no later than the expiration of the maximum term of the applicable stock options.

~~As of~~During the ~~date of this Annual Report:~~year ended December 31, 2018:

Cellectis S.A.

~~82,123~~

43,000 free shares have been granted in October, 2018 under the ~~2012~~Second 2018 Free Share Plan ~~vested on September 18, 2014 are now exercisable, of which 30,981 shares are held by our directors and officers;~~

~~60,000 free shares granted under the 2013 Free Share Plan vested on March 19, 2015 are now exercisable, of which 26,000 free shares are held by our directors and officers;~~

~~98,000 free shares granted under the 2014 Free Share Plan vested on April 10, 2016~~ and are under the vesting period of one year and a holding period of one year for the French residents and vesting period of two years for the US residents, of which ~~8,000~~40,000 free shares have been granted to our ~~directors and~~ officers;

~~50,000 free shares~~

100,000 stock options have been granted in ~~2015 to one employee~~October, 2018 under ~~terms and conditions similar to~~ the ~~2014 Free Share~~2018 Stock Option Plan ~~vested on January 8, 2017,~~ and are under the ~~holding~~vesting period of ~~two years; and~~four year.

414,950 free shares granted under the 2015 Free Share Plan vested on May 18, 2017 and are under the holding period of two years, of which 170,000 free shares have been granted to our directors and officers.

As of the date of this Annual Report, a maximum of 3,541,547 stock options may be optioned and issued under the 2017 Stock Option Plan. This figure includes 1,220,000 stock options granted under the 2017 Stock Option Plan on October 11, 2017 with an exercise price of €22.57 per ordinary share, of which 715,000 were granted to certain of our directors and executive officers.

Calyxt, Inc.

InDuring the year ended December 2014, our subsidiary Calyxt granted options representing a 0.83% interest to a small group of its employees and two of our directors and executive officers, and it reserved an additional 0.1% for further grants. Calyxt made these grants to provide incentives for these employees that are directly linked to the performance of Calyxt, rather than Cellectis as a whole.

~~In September 2015, our subsidiary Calyxt granted options representing a 0.4% interest to a group of its employees and two of our directors and executive officers.~~

~~In April 2016, our subsidiary Calyxt granted options representing a 6.1% interest to a group of its employees and ten of our directors, executive officers, employees and consultants.~~

~~In June 2017,~~31, 2018, our subsidiary Calyxt granted options and restricted stock unit representing a ~~12.9%~~3.1% interest to a group of its employees, ~~and twenty of our~~ directors, executive officers ~~employees~~ and consultants.

C.	Board Practices

Board Composition

Under French law and ourBy-laws, our board of directors must be composed of between three and eighteen members. Within this limit, the number of directors is determined by our shareholders. Directors are elected,re-elected and may be removed at a shareholders’ general meeting with a simple majority vote of our shareholders. Pursuant to ourBy-laws, our directors are elected for three-year terms. In accordance with French law, ourBy-laws also provide that our directors may be removed with or without cause by the affirmative vote of the holders of at least a majority of the votes of the shareholders present, represented by a proxy or voting by mail at the relevant ordinary shareholders’ meeting, and that any vacancy on our board of directors resulting from the death or resignation of a director, provided there are at least three directors remaining, may be filled by vote

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of a majority of our directors then in office provided that there has been no shareholders meeting since such death or resignation. Directors chosen or appointed to fill a vacancy shall be elected by the board for the remaining duration of the current term of the replaced director. The appointment must then be ratified at the next shareholders’ general meeting. In the event the board would be composed of less than three directors as a result of a vacancy, the remaining directors shall immediately convene a shareholders’ general meeting to elect one or several new directors so there are at least three directors serving on the board, in accordance with French law.

We currently have ~~nine~~eight directors. The following table sets forth the names of our directors, the years of their initial appointment as directors and the expiration dates of their current term.

Name ⁽¹⁾
   Current Position    Year of Initial
Appointment    Term
Expiration Year
André Choulika, Ph.D.
   Chairman    2000    2018
David Sourdive, Ph.D.
   Director    2000    2018
Alain Godard
   Director    2007    2018
Pierre Bastid
   Director    2011    2020
Laurent Arthaud
   Director    2011    2020
Annick Schwebig, M.D.
   Director    2011    2020
Jean-Marie Messier
   Director    2015    2018
Hervé Hoppenot
   Director    2017    2020
Rainer Boehm
   Director    2017    2020

~~(1)~~

~~Dr. Mathieu Simon retired from our board of directory in October 2017.~~

~~In addition, French law required companies having at least 50 employees for a period of 12 months over the last three years set up a~~~~Comité d’Entreprise~~~~, or Works’ Council, and~~~~Délégués du personnel~~, or staff delegates, both being composed of representatives elected from among the employees. In conjunction with the French law then applicable to companies of 200 employees and less, we chose to have only one employee representative body, the~~Délégation Unique du Personnel. Our~~~~Délégation Unique du Personnel~~ ~~has been elected for a four year period, and our current~~~~Délégation Unique du Personnel~~ ~~was formed on June 19, 2015. Among these representatives, two are entitled to attend all meetings of the board of directors and shareholders, in an observer capacity.~~


Name	Current Position	Year of Initial Appointment	Term Expiration Year
André Choulika, Ph.D.	Chairman	2000	2021
David Sourdive, Ph.D.	Director	2000	2021
Alain Godard	Director	2007	2021
Pierre Bastid	Director	2011	2020
Laurent Arthaud	Director	2011	2020
Annick Schwebig, M.D.	Director	2011	2020
Hervé Hoppenot	Director	2017	2020
Rainer Boehm, M.D.	Director	2017	2020

Pursuant to new French regulations, which entered into effect on January 1, 2018, any company having more than 10 employees must, on or before January 1st, 2020, implement aComité Social et Économique or Social and Economic Committee, which replaces and regroups the various employee representative bodies, including theDélégation Unique du Personnel. initially in place at Cellectis. We ~~currently intend to comply~~proceeded with the election, for atwo-year term, of this ~~obligation before the end of~~Social and Economic Committee in September 21, 2018.

Director Independence

As a foreign private issuer, under the listing requirements and rules of Nasdaq, we are not required to have independent directors on our board of directors, except with respect to our audit and finance committee, for which the Nasdaq listing requirements permit specifiedphase-in schedules.

Our board of directors has determined that, applying the applicable rules and regulations of the SEC and the Nasdaq listing standards, all of our directors, except Drs. Choulika and Sourdive, qualify as “independent directors.” In making such determination, our board of directors considered the relationships that eachnon-employee director has with us and all other facts and circumstances our board of directors deemed relevant in determining the director’s independence, including the number of ordinary shares beneficially owned by the director and his or her affiliated entities.

Role of the Board in Risk Oversight

Our board of directors is primarily responsible for the oversight of our risk management activities and has delegated to the audit and finance committee the responsibility to assist our board of directors in this task. While

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our board of directors oversees our risk management, our management is responsible forday-to-day risk management processes. We believe this division of responsibilities is the most effective approach for addressing the risks we face. Our board of directors expects our management to consider risk and risk management in each business decision, to proactively develop and monitor risk management strategies and processes forday-to-day activities and to effectively implement risk management strategies adopted by the board.

Corporate Governance Practices

As a Frenchsociété anonyme, we are subject to various corporate governance requirements under French law. In addition, as a foreign private issuer listed on the Nasdaq Global Market, we will be subject to the Nasdaq corporate governance listing standards. However, the Nasdaq Global Market’s listing standards provide that foreign private issuers are permitted to follow home country corporate governance practices in lieu of the Nasdaq rules, with certain exceptions. Certain corporate governance practices in France may differ significantly from corporate governance listing standards. For example, neither the corporate laws of France nor ourBy-laws require that (i) a majority of our directors be independent, (ii) our compensation committee include only independent directors, or (iii) our independent directors hold regularly scheduled meetings at which only independent directors are present. Other than as set forth below, we currently intend to comply with the corporate governance listing standards of Nasdaq to the extent possible under French law. However, we may choose to change such practices to follow home country practice in the future.

Although we are a foreign private issuer, we are required to comply with Rule10A-3 of the Exchange Act, relating to audit committee composition and responsibilities. Rule10A-3 provides that the audit committee must have direct responsibility for the nomination, compensation and choice of our auditors, as well as control over the performance of their duties, management of complaints made, and selection of consultants. Under Rule10A-3, if the laws of a foreign private issuer’s home country require that any such matter be approved by the board of directors or the shareholders of the Company, the audit committee’s responsibilities or powers with respect to such matter may instead be advisory. Under French law, the audit committee may only have an advisory role and appointment of our statutory auditors, in particular, must be decided by our shareholders at our annual meeting.

In addition, Nasdaq rules require that a listed company specify that the quorum for any meeting of the holders of share capital be at least 33¹/₃% of the outstanding shares of the company’s common voting stock. We follow our French home country practice, rather than complying with this Nasdaq rule. Consistent with French Law, ourBy-laws provide that when first convened, general meetings of shareholders may validly convene only if the shareholders present or represented hold at least (1) 20% of the voting shares in the case of an ordinary general meeting or of an extraordinary general meeting where shareholders are voting on a capital increase by capitalization of reserves, profits or share premium, or (2) 25% of the voting shares in the case of any other extraordinary general meeting. If such quorum required by French law is not met, the meeting is adjourned. There is no quorum requirement under French law when an ordinary general meeting or an extraordinary general meeting where shareholders are voting on a capital increase by capitalization of reserves, profits or share premium is reconvened, but the reconvened meeting may consider only questions that were on the agenda of the adjourned meeting. When any other extraordinary general meeting is reconvened, the required quorum under French law is 20% of the shares entitled to vote. If a quorum is not met at a reconvened meeting requiring a quorum, then the meeting may be adjourned for a maximum of two months.

Further, Nasdaq rules require that listed companies have a nominations committee comprised solely of independent directors. We follow our French home country practice rather than complying with this Nasdaq rule.

Finally, Nasdaq rules require shareholder approval when a plan or other equity compensation arrangement is established or materially amended. While the Company may, from time to time, obtain shareholder approval of an equity compensation arrangement in order to obtain advantageous tax treatment or otherwise, as a general matter, we intend to follow our French home country practice, which does not require shareholder approval of such plans or arrangements, rather than complying with this Nasdaq rule.

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Board Committees

The board of directors has established an audit and finance committee and a compensation committee, each of which operates pursuant to a separate charter adopted by our board of directors. The board of directors has also established a scientific committee. The composition and functioning of all of our committees will comply with all applicable requirements of the French Commercial Code, the Exchange Act, the Nasdaq Global Market, and the rules and regulations of the SEC.

In accordance with French law, committees of our board of directors will only have an advisory role and can only make recommendations to our board of directors. As a result, decisions will be made by our board of directors taking into accountnon-binding recommendations of the relevant board committee.

Audit and Finance Committee. Our audit and finance committee reviews our internal accounting procedures, consults with and reviews the services provided by our independent registered public accountants and assists our board of directors in its oversight of our corporate accounting and financial reporting. Currently, our audit and finance committee is comprised of three members of the board of directors: Messrs. Bastid, Arthaud, and ~~Messier.~~Hoppenot.

The duties specifically assigned to the audit and finance committee by our board of directors include, but are not limited to:

with regard to our financial statements:

review on a preliminary basis and express its opinion on the draft annual and quarterly financial statements prior to the board of directors officially receiving the financial statements;

examine the critical accounting policies and practices of the Company, including their relevance and consistency used for the preparation of the Company’s consolidated financial statements and rectify any failure to comply with these policies and practices;

monitor the scope of consolidation and review, where necessary, any explanations in connection thereto;

interview, when necessary, the statutory auditors, the chairman of the board of directors, the chief executive officer, the chief financial officer, the employees in charge of our internal controls or any other management personnel; these discussions may take place, where required, without the presence of the chairman of our board of directors and the chief executive officer; and

examine—prior to their publication—the draft annual and interim financial statements, the draft annual report and any other draft financial statements (including projected financial statements) prepared for the needs of upcoming material transactions together with the related press releases;

with regard to internal controls:

assess the efficiency and quality of internal control systems and procedures within the consolidated Company;

examine, with the persons in charge of the internal audit, and, if necessary, outside of the presence of the chairman of the board of directors and the chief executive officer, the contingency and action plans with respect to internal audit, the findings following the implementation of these actions and the recommendations andfollow-up actions in connection therewith; and

entrust the internal audit department with any mission which the committee deems necessary;

with regard to external controls:

examine any question relating to the appointment, renewal or dismissal of our statutory auditors and their fees regarding the performance of their control review functions;

oversee the rules relating to the use of the statutory auditors for assignments other than the audit of the financial statements and, more generally, ensure that we comply with the principles guaranteeing the statutory auditors’ independence;

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~~financial statements and, more generally, ensure that we comply with the principles guaranteeing the statutory auditors’ independence;~~

at least annually, review and discuss the information provided by management and the auditors relating to the independence of the audit firm;

pre-approve any services entrusted to the statutory auditors which is outside of the scope of the annual audit;

review every year with the statutory auditors all fees paid to by the Company and its subsidiaries to any networks to which the auditors belong, their work plan, their findings and recommendations, as well as actions taken by us following such recommendations;

review and discuss with the statutory auditors their comments on internal controls over financial reporting and any matters that have come to the attention of the statutory auditors that lead them to believe that modification to our disclosures about changes in internal control over financial reporting is necessary for management’s certifications pursuant to Section 302 of the Sarbanes-Oxley Act;

discuss if necessary any points of disagreement between the statutory auditors and the officers of the Company that may arise within the scope of these operations; and

review and discuss with the statutory auditors the plans for, and the scope of, the annual audit and other examinations; and

with regard to risks:

review on a regular basis the financial situation, the cash position and the material risks and undertakings of the Company and its subsidiaries; and

review the risk management policy and the process implemented to evaluate and manage these risks.

Compensation Committee. Our compensation committee assists our board of directors in reviewing the compensation of our executive officers and directors and makes recommendations in respect thereof. Currently, our compensation committee is comprised of two members of the board of directors: Mr. Godard and Dr. Schwebig. The principal duties and responsibilities of our compensation committee include, but are not limited to:

review the compensation of our employees and managers of the Company and its subsidiaries (fixed and variable compensations, bonus, etc.) and make any recommendation to our board of directors in connection therewith;

review equity incentive plans(non-employee warrants, stock options, restricted (free) shares, etc.) and make recommendations to our board of directors in connection therewith;

make recommendations to our board of directors regarding the compensation, pension and insurance plans, benefits in kind and other various pecuniary rights, of officers, as well as the allocation of equity incentive instruments granted to executive officers and directors of the Company;

evaluate and make recommendations on the compensation policies and programs of executive officers and on the compensation of directors;

recommend the approval, adoption and amendment of all cash- and equity-based incentive compensation plans in which any of our executive officers or directors participate and all other equity-based plans;

review any proposed employment agreement with, and any proposed severance or retention plans or agreements applicable to, any of our executive officers;

review, at least annually, corporate goals and objectives relevant to the compensation of our executive officers; and

evaluate the performance of the executive officers in light of corporate goals and objectives and recommend compensation levels for these executive officers based on those evaluations and any other factors the compensation committee deems appropriate.

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Code of Business Conduct and Ethics

We have adopted a Code of Business Conduct and Ethics, or the Code of Conduct, that is applicable to all of our employees, executive officers and directors. The Code of Conduct is available on our website atwww.cellectis.com. Our board of directors will be responsible for overseeing the Code of Conduct and will be required to approve any waivers of the Code of Conduct for employees, executive officers and directors. We expect that any amendments to the Code of Conduct, or any waivers of its requirements, will be disclosed on our website.

Employees

As of December 31, ~~2017,~~2018, we had ~~135~~171 employees, ~~131~~161 of whom are full-time, 4762 of whom hold Ph.D. or M.D. degrees, 96113 of whom were engaged in research and development activities and 3958 of whom were engaged in business development, legal, finance, information systems, facilities, human resources or administrative support. As of December 31, ~~2017, 84~~2018, 85 of our employees were located in France and 5186 of our employees were located in the United States. None of our employees is subject to a collective bargaining agreement. We consider our relationship with our employees to be good.

E.	Share Ownership

For information regarding the share ownership of our directors and executive officers, see “Item 6.B—Compensation” and “Item 7.A—Major Shareholders.”

ITEM 7.

MAJOR SHAREHOLDERS AND RELATED PARTY TRANSACTIONS

A.	Major Shareholders.

The following table sets forth information with respect to the beneficial ownership of our ordinary shares as of ~~February 28,~~December 31, 2018 for:

each beneficial owner of more than 5% of our outstanding ordinary shares;

each of our directors and executive officers; and

all of our directors and executive officers as a group.

Beneficial ownership is determined in accordance with the rules of the SEC. These rules generally attribute beneficial ownership of securities to persons who possess sole or shared voting power or investment power with respect to those securities and include ordinary shares that can be acquired within 60 days of February 28, ~~2018.~~2019. The percentage ownership information shown in the table is based upon ~~35,960,062~~42,430,069 ordinary shares outstanding as of December 31, ~~2017.~~2018.

Except as otherwise indicated, all of the shares reflected in the table are ordinary shares and all persons listed below have sole voting and investment power with respect to the shares beneficially owned by them, subject to applicable community property laws. The information is not necessarily indicative of beneficial ownership for any other purpose.

In computing the number of ordinary shares beneficially owned by a person and the percentage ownership of that person, we deemed outstanding ordinary shares subject to options and warrants held by that person that are immediately exercisable or exercisable within 60 days of February 28, ~~2018.~~2019. We did not deem these shares outstanding, however, for the purpose of computing the percentage ownership of any other person. Beneficial ownership representing less than 1% is denoted with an asterisk (*). The information in the table below is based on information known to us or ascertained by us from public filings made by the shareholders in France. Except

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as otherwise indicated in the table below, addresses of the directors, executive officers and named beneficial owners are in care of Cellectis, 8, rue de la Croix Jarry, 75013 Paris, France.


Name of Beneficial Owner	Ordinary Shares Beneficially Owned					Ordinary Shares Beneficially Owned
	Number		Percentage			Number		Percentage
5% Shareholders:
Bpifrance Participations		2,879,500		8.01	%		2,879,500		6.79	%
Pfizer, Inc. (1)		2,817,455		7.84	%		2,789,252		6.57	%
FMR LLC (2)		3,592,818		10.05	%		4,227,093		9.96	%

Name of Beneficial Owner
   Ordinary Shares Beneficially
Owned
   Number    Percentage
Directors and Executive Officers:

André Choulika, Ph.D (3)
   1,431,732    3.98 %
David Sourdive, Ph.D. (4)
   1,368,535    3.81 %
Philippe Duchateau, Ph.D. (5)
   354,973    *
Eric Dutang (6)
   143,950    *
Julia Berretta (7)
   217,165    *
Marie-Bleuenn Terrier (8)
   292,609    *
Stephan Reynier (9)
   124,493    *
Elsy Boglioli
   0    *
Stéphane Depil
   0    *
Federico Tripodi
   0    *
Alain Godard (10)
   144,999    *
Pierre Bastid (11)
   3,422,394    9.52 %
Laurent Arthaud
   0    *
Annick Schwebig, M.D. (12)
   51,997    *
Jean-Marie Messier (13)
   110,117    *
Hervé Hoppenot
   0    *
Rainer Boehm
   0    *
All directors and executive officers as a group (17 persons)
   7,664,904    21.32 %


Name of Beneficial Owner	Ordinary Shares Beneficially Owned
	Number		Percentage
Directors and Executive Officers:
André Choulika, Ph.D (3)		1,702,903		4.01	%
David Sourdive, Ph.D. (4)		1,570,819		3.70	%
Philippe Duchateau, Ph.D. (5)		509,197			*
Eric Dutang (6)		257,373			*
Marie-Bleuenn Terrier (7)		451,724			*
Stephan Reynier (8)		190,973			*
Elsy Boglioli (9)		45,000			*
Alain Godard (10)		201,724			*
Pierre Bastid (11)		3,479,119		8.20	%
Laurent Arthaud		0			*
Annick Schwebig, M.D. (12)		162,115			*
Hervé Hoppenot (13)		13,333			*
Rainer Boehm, M.D. (14)		13,333			*
All directors and executive officers as a group (13 persons)		8,597,614		20.26	%

*	Represents beneficial ownership of less than one per cent.

(1)

The address of Pfizer, Inc. is 235 East 42nd Street, New York, New York 10017. Shares beneficially owned by Pfizer, Inc. were acquired by Pfizer OTC B.V. on July 31, 2014 in the context of a share capital increase in connection with the Research and Collaboration Agreement between Pfizer Inc. and Cellectis S.A., dated June 17, 2014.

(2)

Amounts beneficially owned by FMR LLC were reported pursuant to a Schedule 13G amendment filed with the SEC on February 13, 2018 by FMR LLC and Abigail P. Johnson. FMR LLC’s address is 245 Summer Street, Boston, Massachusetts 02210. FMR LLC is the parent company of Fidelity Management & Research Company (“FMR Co.”), which carries out the voting of shares owned by various Fidelity funds under written guidelines established by the Fidelity funds’ boards of trustees.

(3)

Includes ~~164,379~~219,173 ordinary shares that Mr. Choulika has the right to acquire pursuant to stock options granted in March 2015 under the 2015 Stock Option Plan, ~~125,000~~175,000 ordinary shares that Mr. Choulika has the right to acquire pursuant to stock options granted in September 2015 governed by the 2015 Stock Option Plan, ~~80,350~~120,525 ordinary shares that Mr. Choulika has the right to acquire pursuant to stock options granted in March 2016 under the 2015 Stock Option Plan, ~~and 84,929~~141,548 ordinary shares that Mr. Choulika has the right to acquire pursuant to stock options granted in ~~March~~October 2016 under the 2016 Stock Option Plan and 50,625 ordinary shares that Mr. Choulika has the right to acquire pursuant to stock options granted in October 2017 under the 2017 Stock Option Plan.

(4)

Includes ~~131,507~~175,343 ordinary shares that Mr. Sourdive has the right to acquire pursuant to stock options granted in March 2015 under the 2015 Stock Option Plan, ~~109,375~~153,125 ordinary shares that Mr. Sourdive has the right to acquire pursuant to stock options granted in September 2015 governed by the 2015 Stock Option

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Plan and ~~70,307~~105,460 ordinary shares that Mr. Sourdive has the right to acquire pursuant to stock options granted in March 2016 under the 2015 Stock Option Plan, ~~74,313~~123,855 ordinary shares that Mr. Sourdive has the right to acquire pursuant to stock options granted in October 2016 under the 2016 Stock Option Plan and 30,000 ordinary shares that Mr. Sourdive has the right to acquire pursuant to stock options granted in October 2017 under the 2017 Stock Option Plan. Includes 703,041 shares held by Viveoo SARL.

(5)

~~Includes 98,630~~

Includes131,508 ordinary shares that Dr. Duchateau has the right to acquire pursuant to stock options granted in March 2015 under the 2015 Stock Option Plan, ~~93,750~~131,250 ordinary shares that Dr. Duchateau has the right to acquire pursuant to stock options granted in September 2015 governed by the 2015 Stock Option Plan and ~~60,263~~90,394 ordinary shares that Dr. Duchateau has the right to acquire pursuant to stock options granted in March 2016 under the 2015 Stock Option Plan, ~~and 63,696~~106,161 ordinary shares that Dr. Duchateau has the right to acquire pursuant to stock options granted in October 2016 under the 2016 Stock Option Plan and 11,250 ordinary shares that Dr. Duchateau has the right to acquire pursuant to stock options granted in October 2017 under the 2017 Stock Option Plan.

(6)

Includes ~~9,375~~13,125 ordinary shares that Mr. Dutang has the right to acquire pursuant to stock options granted in September 2015 governed by the 2015 Stock Option Plan, ~~60,263~~90,394 ordinary shares that Mr. Dutang has the right to acquire pursuant to stock options granted in March 2016 under the 2015 Stock Option Plan, ~~and 74,313~~123,855 ordinary shares that Mr. Dutang has the right to acquire pursuant to stock options granted in October 2016 under the 2016 Stock Option ~~Plan.~~

~~(7)~~

~~Includes 65,752~~Plan and 30,000 ordinary shares that Mrs. Berretta has the right to acquire pursuant to stock options granted in March 2015 under the 2015 Stock Option Plan, 56,250 ordinary shares that Mrs. Berretta has the right to acquire pursuant to stock options granted in September 2015 governed by the 2015 the Stock Option Plan, 40,263 ordinary shares that Mrs. Berretta has the right to acquire pursuant to stock options granted in March 2016 under the 2015 Stock Option Plan and 31,131 ordinary shares that Mrs. BerrettaMr. Dutang has the right to acquire pursuant to stock options granted in October ~~2016~~2017 under the ~~2016~~2017 Stock Option Plan.

~~(8)~~(7)

Includes ~~65,752~~87,671 ordinary shares that Mrs. Terrier has the right to acquire pursuant to stock options granted in March 2015 under the 2015 Stock Option Plan, ~~56,250~~78,750 ordinary shares that Mrs. Terrier has the right to acquire pursuant to stock options granted in September 2015 governed by the 2015 Stock Option Plan, ~~70,306~~105,460 ordinary shares that Mrs. Terrier has the right

to acquire pursuant to stock options granted in March 2016 under the 2015 Stock Option Plan, ~~and 74,313~~123,855 ordinary shares that Mrs. Terrier has the right to acquire pursuant to stock options granted in October 2016 under the 2016 Stock Option Plan and 30,000 ordinary shares that Mrs. Terrier has the right to acquire pursuant to stock options granted in October 2017 under the 2017 Stock Option Plan.

~~(9)~~(8)

Includes ~~29,589~~39,452 ordinary shares that Mr. Reynier has the right to acquire pursuant to stock options granted in March 2015 under the 2015 Stock Option Plan, ~~25,000~~35,000 ordinary shares that Mr. Reynier has the right to acquire pursuant to stock options granted in September 2015 governed by the 2015 Stock Option Plan, ~~29,428~~44,142 ordinary shares that Mr. Reynier has the right to acquire pursuant to stock options granted in March 2016 under the 2015 Stock Option Plan, ~~and 25,353~~42,256 ordinary shares that Mr. Reynier has the right to acquire pursuant to stock options granted in October 2016 under the 2016 Stock Option Plan and 15,000 ordinary shares that Mr. Reynier has the right to acquire pursuant to stock options granted in October 2017 under the 2017 Stock Option Plan.

(9)	Includes 45,000 ordinary shares that Mrs. Boglioli has the right to acquire pursuant to stock options granted in October 2017 under the 2017 Stock Option Plan.

(10)

The ordinary shares include 50,000non-employee warrants which are exercisable since March 27, 2016, ~~33,6333~~50,000non-employee warrants, which are exercisable since September 8, 2016, ~~26,783~~40,175non-employee warrants, which are exercisable since March 14, 2017, 26,667non-employee warrants, which are exercisable since October 28, 2017 and 13,333non-employee warrants, which are exercisable since October ~~28, 2017.~~10, 2018.

(11)

The ordinary shares include 50,000non-employee warrants which are exercisable since March 27, 2016, ~~33,333~~50,000non-employee warrants, which are exercisable since September 8, 2016, ~~26,783~~40,175non-employee warrants, which are exercisable since March 14, 2017, ~~13,333~~26,667non-employee warrants, which are exercisable since October 28, 2017, 13,333non-employee warrants, which are exercisable since October 10, 2018 and includes 3,298,944 shares held by ~~Zaka Rendement S.A.~~Lohas SARL.

(12)

The ordinary shares include 30,000non-employee warrants which are exercisable since March 27, 2016, ~~33,333~~50,000non-employee warrants, which are exercisable since September 8, 2016, ~~26,783~~40,175non-employee warrants, which are exercisable since March 14, 2017, 26,667non-employee warrants, which are exercisable since October 28, 2017 and 13,333non-employee warrants, which are exercisable since October ~~28, 2017.~~10, 2018.

(13)	The ordinary shares include ~~50,000~~13,333non-employee warrants which are exercisable since ~~March 27, 2016, and 33,333~~October 10, 2018.

(14)	The ordinary shares include 13,333non-employee warrants which are exercisable since ~~September 8, 2016, and 26,783non-employee~~ ~~warrants, which are exercisable since March 14, 2017.~~October 10, 2018.

The significant changes in the percentage ownership held by our principal shareholders since January 1, 2014 are as a result of the transactions described in our prospectus dated March 26, 2015, filed with the SEC

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pursuant to Rule 424(b), under the heading “Related Party Transactions—Transactions with Our Principal Shareholders, Directors and Executive Officers” and the dilution resulting from our public offering.

None of our principal shareholders has voting rights different than our other shareholders.

As of ~~February 28,~~December 31, 2018, assuming that all of our ordinary shares represented by ADSs are held by residents of the United States, we estimate that approximately ~~40.2%~~52.10% of our outstanding ordinary shares were held in the United States by 64114 holders of record.

B.	Related Party Transactions

Since January 1, ~~2015,~~2012, we have engaged in the following transactions with our directors, executive officers and holders of more than 5% of our outstanding voting securities and their affiliates, which we refer to as our related-parties.

Transactions with Our Principal Shareholders, Directors and Executive Officers

Alliance agreement

Pfizer purchased 10% of our then-outstanding ordinary shares on July 31, 2014. The revenues booked for Pfizer in the years ended December 31, ~~2015,~~ 2016, 2017 and ~~2017~~2018 amount to ~~$10.2 million,~~ $25.3 million, $19.7 million and ~~$19.7~~$7.4 million respectively. As of December 31, ~~2017,~~2018, there no outstanding receivables and Pfizer had a ~~7.84%~~6.57% ownership in Cellectis.

Conditional advances and subsidies

Bpifrance, which is a shareholder of Cellectis, has granted us conditional advances and subsidies. There was no outstanding conditional advances and subsidies since December 31, 2016.

Agreements with Our Directors and Executive Officers

Director and Executive Officer Compensation

See “Item 6.B—Compensation of Directors and Executive Officers” for information regarding compensation of directors and executive officers

Equity Awards

Since January 1, ~~2017,~~2018, we have granted equity awards to certain of our directors and executive officers:

On October ~~11, 2017,~~8, 2018, we granted ~~715,000~~50,000 stock options to certain of our ~~Directors and~~ Executive Officers; and

On October ~~11, 2017,~~8, 2018, we granted ~~240,000non-employee~~ ~~warrants (BSA)~~40,000 free shares to our ~~Directors.~~Executive Officers.

See “Item. 7A—Major Shareholders” for information regarding equity awards to certain of our executive officers.

Indemnification Agreements

See “Item. 6B—Limitations on Liability and Indemnification Matters.”

Insofar as indemnification for liabilities arising under the Securities Act may be permitted to directors, officers or persons controlling us pursuant to the foregoing provisions, we have been informed that in the opinion of the SEC such indemnification is against public policy as expressed in the Securities Act and is therefore unenforceable.

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Transactions with subsidiaries: Calyxt IPO and Key Arrangements

On July 25, 2017, Calyxt completed an initial public offering on the Nasdaq Global Market, selling an aggregate of 8,050,000 shares of common stock at a price of $8.00 per share (including the full exercise by the underwriters of their over-allotment option). The Company received net proceeds of approximately $58.0 million, after deducting underwriting discounts and commissions and offering expenses. As part of the IPO, Cellectis purchased 2,500,000 shares of common stock for a value of $20.0 million, which is included in the net proceeds that Calyxt received. Calyxt used $5.7 million of the proceeds from us to cover a portion of the outstanding obligations owed to Cellectis. Following the initial public offering, ~~as of February 28, 2018,~~ Cellectis owns approximately ~~79.3%~~79.6% of Calyxt’s common stock.

In May, 2018, Calyxt closed afollow-on offering of 4,057,500 ADS at a public offering price of $15.00 per ADS resulting in gross proceeds of $60.9 million. Cellectis purchased 550,000 shares of common stock at the public offering price of $15.00. In addition, in connection with the vesting on June 14, 2018, of restricted stock units for certain employees and nonemployees of Calyxt and Cellectis, Cellectis purchased approximately 63,175 shares of common stock of Calyxt at a price of $19.49 per share (the closing price reported on the Nasdaq Global Market on June 14, 2018) directly from such employees and nonemployees in private transactions pursuant to share purchase agreements dated June 13, 2018.

In connection with Calyxt’s IPO, we and Calyxt entered into certain agreements that relate to our relationship with Calyxt prior to the IPO or that provide a framework for our ongoing relationship with Calyxt. The summaries of the most significant provisions of these agreements. These summaries are qualified in their entirety by reference to the full text of such agreements.

Management Services Agreement

We are party to a management services agreement dated January 1, 2016 and amended July 25, 2017 that we entered into with Calyxt and Cellectis, Inc., a Delaware corporation and our ~~wholly owned~~wholly-owned subsidiary (“Cellectis, Inc.”), pursuant to which we and Cellectis, Inc. provide certain services to Calyxt, including certain general management, finance, investor relations, communication, legal, intellectual property, human resources and information technology services. In consideration for such services, Calyxt pays to us and Cellectis, Inc. certain fees, consisting of reimbursement of all costs and expenses reasonably incurred by us in connection with the provision of such services, payment of amark-up corresponding to a percentage of certain of the costs and expenses, which range from zero to 10%, and reimbursement of costs and expenses of services that are subcontracted by us on Calyxt’s behalf.

The management services agreement is automatically renewed for one year periods starting on January 1st of each year. Either party has the right to terminate the agreement at the anniversary date of the agreement by giving three months prior notice. We also entered into an amendment to the agreement in connection with IPO to provide that the agreement may otherwise be terminated by us or by Calyxt in connection with certain material breaches by the other party upon prior written notice subject to limited cure periods, the sale of all or substantially all of the assets of either party, certain bankruptcy events or certain judgments.

During fiscal year ~~2016,~~2018, Calyxt made payments to us for services provided under the management services agreement of ~~$1.8~~$1,5 million, which excludes directre-invoicing and royalties paid to us.

Stockholders Agreement

On July 25, 2017 we entered into a stockholders agreement with Calyxt, which we refer to as the stockholders agreement. Pursuant to our stockholders agreement with Calyxt, we have certain contractual rights for so long as we beneficially own at least 50% of the then outstanding shares of Calyxt’s common stock, including:

to approve any modification to Calyxt’s or any future Calyxt subsidiary’s share capital (e.g., share capital increase or decrease), the creation of any subsidiary by Calyxt, any grant of stock-based compensation, any distributions or initial public offering, merger,spin-off, liquidation, winding up orcarve-out transactions;

to approve Calyxt’s annual business plan and annual budget and any modification thereto;

to approve any external growth transactions of Calyxt exceeding $500,000 and not included in the approved annual business plan and annual budget;

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to approve any investment and disposition decisions by Calyxt exceeding $500,000 and not included in the approved annual business plan and annual budget (it being understood that this clause excludes the purchase and sale of inventory as a part of the normal course of business);

to approve any related-party agreement and any agreement or transaction between the executives or shareholders of Calyxt, on the one hand, and Calyxt or any of its subsidiaries, on the other hand;

to approve any decision by Calyxt pertaining to the recruitment, dismissal/removal, or increase of the compensation of executives and corporate officers;

to approve any material decision by Calyxt relating to a material litigation;

to approve any decision by Calyxt relating to the opening of a social or restructuring plan orpre-insolvency proceedings;

to approve any buyback by Calyxt of its own shares;

to approve any new borrowings or debts of Calyxt exceeding $500,000 and early repayment of loans, if any (it being understood that we will approve the entering into of contracts for revolving loans and other short-term loans and the repayment of such for financing general operating activities, such as revolving loans for inventory or factoring of receivables);

to approve grants by Calyxt of any pledges on securities;

to develop new activities and businesses not described in the annual business plan and annual budget;

to approve entry into any material agreement or partnership; and

to approve any offshore and relocation activities of Calyxt.

In addition, we have the following rights for so long as we beneficially own at least 15% of the then outstanding shares of Calyxt’s common stock, including:

to nominate the greater of three members of Calyxt’s Board of Directors or a majority of the directors;

to designate the Chairman of Calyxt’s Board of Directors and one member to each of the audit committee of the Board of Directors, the compensation committee of the Board of Directors and the nominating and corporation governance committee of the Board of Directors;

to approve any amendments to Calyxt’ amended and restated certificate of incorporation or its amended and restatedby-laws that would change the name of Calyxt, its jurisdiction of incorporation, the location of its principal executive offices, the purpose or purposes for which Calyxt is incorporated or the Cellectis approval items set forth in the stockholders agreement;

to approve the payment of any regular or special dividends;

to approve the commencement of any proceeding for the voluntary dissolution, winding up or bankruptcy of Calyxt or a material subsidiary;

to approve any public or private offering, merger, amalgamation or consolidation of Calyxt or the spinoff of a business of Calyxt or any sale, conveyance, transfer or other disposition of Calyxt’s assets; and

to approve any appointment to Calyxt’s Board of Directors contrary to the stockholders agreement or Calyxt’s certificate of incorporation or Calyxt’sby-laws.

In addition, for so long as we beneficially own at least 15% of the then outstanding shares of Calyxt’s common stock, (i) we will be entitled to certain information rights, including the right to consult with and advise senior management, to receive quarterly and annual financial statements and to review Calyxt’s books and records and (ii) Calyxt will also be required to cooperate with us in connection with certain sales and pledges of Calyxt’s shares or grants of security interests in respect thereof, including in connection with margin loans.

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The stockholders agreement will also provide us with certain registration rights, including certain demand and piggyback registration rights. The registration rights will remain in effect with respect to any shares covered by the Stockholders Agreement until (i) all of our Calyxt shares have been sold pursuant to an effective registration statement under the Securities Act; (ii) all of our Calyxt shares have been sold to the public pursuant to Rule 144 under the Securities Act; or (iii) we own less than 10% of the then outstanding shares of Calyxt’s common stock.

Separation Agreement

On July 25, 2017, we entered into a separation agreement with Calyxt, which sets forth certain agreements between us and Calyxt that will govern the relationship between us and Calyxt following this offering, including with respect to the following matters:

guarantees;

insurance policies;

mutual releases and indemnification matters;

accounting, financial reporting and internal control issues;

confidentiality;

ability of the parties to compete with each other; and

settlement of intercompany accounts.

The separation agreement will terminate upon the earlier of (i) mutual written consent of us and Calyxt and (ii) the date on which we and our affiliates cease to hold at least 15% of the then outstanding shares of Calyxt’s common stock.

License Agreement with Calyxt

We are party to a license agreement with Calyxt pursuant to which Calyxt has been granted an exclusive, worldwide license (subject to existing licenses granted by us to third parties) to use, commercialize and exploit certain intellectual property in the field of researching, developing and commercializing agricultural and food products, including traits, seeds, and feed and food ingredients (excluding any application in connection with animals and animal cells), except that such license will benon-exclusive in such field for any activities relating to researching, developing or commercializing certain modified or mutatedI-CreI homing endonucleases. Calyxt has also been granted anon-exclusive license to use the TALEN trademark in connection with its exploitation of licensed products under the agreement. Any improvements Calyxt makes to the licensed intellectual property will be owned by Calyxt but licensed back to us on an exclusive basis for any use outside of Calyxt’s exclusive agricultural field of use.

In consideration for the license from us, Calyxt is required to pay to us, on aproduct-by-product andcountry-by-country basis, a royalty of 3% of net sales less costs for grain and seed of any products that are covered by the patents licensed from us. In addition, Calyxt will be required to pay us 30% of revenue Calyxt receives for sublicensing its rights under the agreement to third parties. Calyxt’s payment obligations to us will expire upon the expiration of thelast-to-expire valid claim of the patents licensed to Calyxt by us.

Under our license agreement with Calyxt, and as between the parties, we have the first right to control the prosecution, maintenance, defense and enforcement of the licensed intellectual property and Calyxt will have the right to step in and assume such control with respect to the patents owned by us and exclusively licensed to Calyxt under the agreement if we elect to not prosecute, maintain, defend or enforce such patents. In certain circumstances, if we elect to abandon any patents owned by us and exclusively licensed to Calyxt under the agreement, Calyxt will have the right to assume ownership of such patents. In addition, some of the intellectual

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property that will be licensed to Calyxt by us consists of an exclusive sublicense, subject to existing sublicenses granted by us to third parties, of intellectual property originally licensed to us by the University of Minnesota to exploit such intellectual property in Calyxt’s exclusive agricultural field of use. Therefore, as to such sublicensed intellectual property, Calyxt’s license from us will be subject to the terms and conditions of the license agreement between the University of Minnesota and us, and to the extent Calyxt’s activities under such sublicense violate any terms and conditions of the license agreement between us and the University of Minnesota, Calyxt will be responsible for any damages that we may incur. In

addition, Calyxt is required to reimburse us for any and all payments made by us to the University of Minnesota pursuant to the license agreement between the University of Minnesota and us to the extent that any such payments are required to be made as a result of Calyxt’s applicable activities. Under the license agreement between us and the University of Minnesota, the University of Minnesota has the first right to control the prosecution and maintenance of the licensed intellectual property.

Calyxt’s license agreement with us is perpetual. However it may be terminated upon the mutual written agreement of both parties, either party’s uncured material breach of the agreement, or upon certain bankruptcy and insolvency related events.

Related-Party Transactions Policy

We have adopted a related-party transaction policy that sets forth our procedures for the identification, review, consideration and approval or ratification of related-party transactions. The policy became effective immediately upon the completion of our initial public offering. For purposes of our policy only, a related-party transaction is a transaction, arrangement or relationship, or any series of similar transactions, arrangements or relationships, in which we and any related parties are, were or will be participants, which are not (1) in the ordinary course of business, (2) at arms’ length and (3) in which the amount involved exceeds $120,000. Transactions involving compensation for services provided to us as an employee or director are not covered by this policy. For purposes of this policy, a related party is any executive officer, director (or nominee for director) or beneficial owner of more than five percent (5%) of any class of our voting securities, including any of their respective immediate family members and any entity owned or controlled by such persons.

Under the policy, related-party transactions must be reported to us by all related parties. If a transaction has been identified as a related-party transaction, our management must present information regarding therelated-party transaction to our board of directors for review, consideration and approval. Certain transactions may be presented to the Audit and Finance Committee, which will determine whether the transaction is arelated-party transaction, in which case the related-party transaction will be submitted to our board of directors. The presentation will include a description of, among other things, the material facts, the interests in the transaction, direct and indirect, of the related parties, the benefits to us of the transaction and whether the transaction is on terms that are comparable to the terms available to or from, as the case may be, an unrelated third-party or to or from employees generally. In addition, under our Code of Business Conduct and Ethics, our employees and directors have an affirmative responsibility to disclose any transaction or relationship that reasonably could be expected to give rise to a conflict of interest. In considering related-party transactions, our board of directors, or to the extent permitted by applicable law an independent committee of our board of directors, will take into account the relevant available facts and circumstances including, but not limited to:

the benefits and perceived benefits to us;

the opportunity costs of alternative transactions;

the materiality and character of the related party’s interest;

the actual or apparent conflict of interest of the related party; and

the terms available to or from, as the case may be, unrelated third parties or to or from employees generally.

The policy requires that, in determining whether to approve, ratify or reject a related-party transaction, our board of directors, or if permitted by applicable law an independent committee of our board of directors, must

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consider, in light of known circumstances, whether the transaction is in, or is not inconsistent with, our best interests and those of our shareholders, as our board of directors, or if permitted by applicable law an independent committee of our board of directors, determines in the good faith exercise of its discretion.

D.	Interests of Experts and Counsel

Not applicable.

ITEM 8.

FINANCIAL INFORMATION

A.	Consolidated Statements and Other Financial Information

Our consolidated financial statements are appended at the end of this Annual Report starting at pageF-1, and form a part hereof.

Legal Proceedings

From time to time, we may be involved in various claims and legal proceedings relating to claims arising out of our operations. We are not currently a party to any legal proceedings that, in the opinion of our management, are likely to have a material adverse effect on our business or our cash flows. Regardless of outcome, litigation can have an adverse impact on us because of defense and settlement costs, diversion of management resources and other factors.

Dividend Distribution

Approval of Dividends. Pursuant to French law, our board of directors may propose a dividend and/or reserve distribution for approval by the shareholders at the annual ordinary general meeting related to the statutory financial statements of Cellectis S.A.

Upon recommendation of our board of directors, our shareholders may decide to allocate all or part of any distributable profits to special or general reserves, to carry them forward to the next fiscal year as retained earnings or to allocate them to the shareholders as dividends. However, dividends may not be distributed when as a result of such distribution, our net assets are or would become lower than the amount of the share capital plus the amount of the legal reserves which, under French law, may not be distributed to shareholders (the amount of our share capital plus the amount of our legal and other reserves which may not be distributed was equal to ~~$2.3~~$2.5 million on December 31, ~~2017)~~2018). Moreover, the statutory accumulated deficit is ~~$159.5~~$162.4 million as of December 31, ~~2017.~~2018.

Our board of directors may distribute interim dividends after the end of the fiscal year but before the approval of the financial statements for the relevant fiscal year when the interim balance sheet, established during such year and certified by an auditor, reflects that we have earned distributable profits since the close of the last financial year, after recognizing the necessary depreciation and provisions and after deducting prior losses, if any, and the sums to be allocated to reserves, as required by law or theBy-laws, and including any retained earnings. The amount of such interim dividends may not exceed the amount of the profit so defined.

Distribution of Dividends. Dividends are distributed to shareholders proportionally to their shareholding interests. In the case of interim dividends, distributions are made to shareholders on the date set by our board of directors during the meeting in which the distribution of interim dividends is approved. The actual dividend payment date is decided by the shareholders at an ordinary general shareholders’ meeting or by our board of directors in the absence of such a decision by the shareholders. Shareholders that own shares on the actual payment date are entitled to the dividend.

Dividends may be paid in cash or, if the shareholders’ meeting so decides, in kind, provided that all the shareholders receive a whole number of assets of the same nature paid in lieu of cash. OurBy-laws provide that, subject to a decision of the shareholders’ meeting taken by ordinary resolution, each shareholder may be given the choice to receive his dividend in cash or in shares.

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B.	Significant Changes

~~None.~~In March 2019, we entered into a lease agreement for a 82,000 square foot commercial-scale manufacturing facility, called the IMPACT site, which stands for “Innovative Manufacturing Plant for Allogeneic Cellular Therapies”. The IMPACT facility is located in Raleigh, North Carolina. The new manufacturing facility is being designed to provide GMP manufacturing for clinical supply and commercial product upon potential regulatory approval. The facility is planned to be operational by 2021.

ITEM 9.

THE OFFER AND LISTING

A.	Offer and Listing Details

Our ADS have been listed on Nasdaq Global Market under the symbol “CLLS” since March 24, 2015. Prior to that date, there was no public trading market for ADSs. Our ordinary shares have been trading on Euronext Growth market of Euronext Paris under the symbol “ALCLS” since February 7, 2007. Prior to that date, there was no public trading market for ADSs or our ordinary shares. Our initial public offering in March 2015 was priced at $41.50 per ADS on March 24, 2015. The following tables set forth for the periods indicated the reported high and low sale prices per ADS on Nasdaq in U.S. dollars and per ordinary share on Euronext Growth in euros.

~~Nasdaq~~

Period
   High    Low
Annual

2015 (beginning March 24, 2015)
   $ 47.66    $ 23.67
2016
   $ 33.64    $ 16.40
2017
   $ 35.01    $ 17.52
2018 (through March 12, 2018)
   $ 34.17    $ 27.78
Quarterly

First Quarter 2016
   $ 30.16    $ 18.77
Second Quarter 2016
   $ 33.64    $ 25.34
Third Quarter 2016
   $ 28.35    $ 24.07
Fourth Quarter 2016
   $ 23.24    $ 16.40
First Quarter 2017
   $ 24.37    $ 17.52
Second Quarter 2017
   $ 25.92    $ 21.96
Third Quarter 2017
   $ 32.18    $ 21.65
Fourth Quarter 2017
   $ 35.01    $ 23.33
First Quarter 2018 (through March 12, 2018)
   $ 34.17    $ 27.78
Monthly

September 2017
   $ 32.18    $ 24.96
October 2017
   $ 35.01    $ 27.51
November 2017
   $ 29.18    $ 23.33
December 2017
   $ 29.15    $ 23.70
January 2018
   $ 34.17    $ 27.78
February 2018
   $ 32.65    $ 28.53
March 2018 (through March 12, 2018)
   $ 34.07    $ 31.57

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~~Alternext~~

Period
   High    Low
Annual

2013
   € 7.70    € 2.19
2014
   € 14.00    € 2.40
2015
   € 40.90    € 13.26
2016
   € 29.71    € 14.99
2017
   € 29.61    € 16.33
2018 (through March 12, 2018)
   € 27.42    € 23.12
Quarterly

First Quarter 2016
   € 27.74    € 22.01
Second Quarter 2016
   € 29.71    € 22.67
Third Quarter 2016
   € 25.18    € 21.97
Fourth Quarter 2016
   € 20.81    € 14.99
First Quarter 2017
   € 22.49    € 16.33
Second Quarter 2017
   € 24.02    € 19.96
Third Quarter 2017
   € 26.84    € 18.23
Fourth Quarter 2017
   € 29.61    € 19.55
First Quarter 2018 (through March 12, 2018)
   € 27.42    € 23.12
Monthly

September 2017
   € 26.84    € 20.84
October 2017
   € 29.61    € 22.88
November 2017
   € 26.90    € 19.55
December 2017
   € 24.01    € 20.09
January 2018
   € 27.14    € 23.12
February 2018
   € 26.80    € 23.34
March 2018 (through March 12, 2018)
   € 27.42    € 25.30

B.	Plan of Distribution

Not applicable.

Markets

The ADS have been listed on Nasdaq Global Market under the symbol “CLLS” since March 24, 2015 and our ordinary shares have been listed on the Euronext Growth market of Euronext in Paris under the symbol “ALCLS” since February 7, 2007.

D.	Selling Shareholders

Not applicable.

Dilution

Not applicable.

F.	Expenses of the Issue

Not applicable.

ITEM 10.

ADDITIONAL INFORMATION

A.	Share Capital

Not applicable.

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B.	Memorandum and Articles of Association

Key Provisions of OurBy-laws and French Law

The description below reflects the terms of ourBy-laws, and summarizes the material rights of holders of our ordinary shares under French law. Please note that this is only a summary and is not intended to be exhaustive. For further information, please refer to the full version of ourBy-laws which was filed on January 9, 2018. In the event that our Bylaws are modified in connection with the creation of a specific category of preferred shares, the rights of holders of such preferred shares under our~~By-laws~~ ~~and French law will be described in the applicable prospectus supplement.~~is included as an exhibit to this Annual Report.

Corporate Purpose

Our corporate purpose, which is set forth in Article 3 of our Bylaws, in France and abroad includes:

all activities related to genetics and more specifically to genome engineering, in particular, research, development and invention, filing and use of patents and trademarks, sale and marketing, advising and assisting, in all areas, in particular in the agro-food, pharmaceutical, textile and environmental sectors; and

more generally, all industrial, commercial, financial and civil transactions and transactions involving real estate or movable property relating directly or indirectly to any of the aforementioned corporate purposes or any similar or related purpose.

Directors

Quorum and Voting. The board of directors may only deliberate if at least half of the directors attend the applicable meeting in the manner provided for in ourBy-laws. In particular, French law and the charter of the board of directors allow

directors to attend meetings of the board of directors in person or, to the extent permitted by applicable law, by videoconference or other telecommunications arrangements. In addition, ourBy-Laws allow a director to grant another director a proxy to represent him or her at a meeting of the board of directors, but no director can hold more than one proxy at any meeting. Decisions of the board of directors are adopted by the majority of the voting rights held by the directors present or represented, it being specified that in case of a vote-split, the Chairman of the board of directors shall have a deciding vote.

Directors’ Voting Powers on Proposal, Arrangement or Contract in which any Director is Materially Interested. Under French law, any agreement entered into (directly or through an intermediary) between us and any director that is not entered into (1) in the ordinary course of business and (2) under standard terms and conditions is subject to the prior authorization of the board of directors, excluding the vote of the interested director.

The foregoing requirements also apply to agreements between us and another company, provided that the company is not one of our wholly-owned subsidiaries, if one of our directors is the owner or a general partner, manager, director, general manager or member of the executive or supervisory board of the other company, as well as to agreements in which one of our directors has an indirect interest.

~~Decisions Subject to the Prior Approval of the Board of Directors~~~~. OurBy-laws~~ provide that none of the decisions listed below (whether they relate to us or to any of our subsidiaries) shall be made without the prior approval of our board acting, upon first notice, by a majority of its members in office and, upon second notice, by a majority of its members present or represented by proxy:

~~approval and modification to the business plan and budget;~~

~~any decision pertaining to the recruitment, dismissal/removal, or increase of the compensation of executives and corporate officers;~~

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~~any material decision relating to a material litigation in which we or any of our subsidiaries may have to pay an amount exceeding €2,000,000;~~

~~any decision relating to the opening of a social or restructuring plan orpre-insolvency~~ ~~proceedings;~~

~~dual listing of the company on a stock market located outside of France;~~

~~any repurchase of our own shares;~~

~~any new borrowings or debts exceeding €1,000,000 and early repayment of loans, if any;~~

~~grants of any pledges on securities exceeding €1,000,000;~~

~~offshore or relocate activities;~~

~~development of new activities and businesses not described in the budget; and~~

~~entry into any material agreement or partnership.~~

Directors’ ~~Compensation~~Compensation.. The aggregate amount of attendance fees (jetons de présence) of the board of directors is determined at the shareholders’ annual ordinary general meeting. The board of directors then divides all or part (at the board’s discretion) of this aggregate amount among some or all of its members by a simple majority vote. In addition, the board of directors may grant exceptional compensation (rémunérations exceptionnelles) to individual directors on acase-by-case basis for special and temporary assignments. The board of directors may also authorize the reimbursement of reasonable travel and accommodation expenses, as well as other expenses incurred by directors in the corporate interest.

Board of Directors’ Borrowing ~~Powers~~Powers.. There are currently no limits imposed by ourBy-laws on the amounts of loans or borrowings that the board of directors may approve.

Directors’ Age ~~Limits~~Limits.. The number of directors who are more than seventy (70) years old may not exceed one third of the directors in office.

Term of Director ~~Office~~Office.. OurBy-laws provide that members of our board of directors are elected for a tenure of three years.

Employee Director ~~Limits~~Limits.. The number of directors who are also party to employment contracts with the Company may not exceed one third of the directors in office.

Directors’ Share Ownership Requirements. None.

Rights, Preferences and Restrictions Attaching to Ordinary Shares

Dividends. We may only distribute dividends out of our ~~“distributable~~“distributable profits,” plus any amounts held in our reserves that the shareholders decide to make available for distribution, other than those reserves that are specifically required to be maintained by law. ~~“Distributable profits”~~“Distributable profits” consist of our unconsolidated net profit in each fiscal year, as increased or reduced by any profit or loss carried forward from prior years, less any contributions to the reserve accounts pursuant to French law (see below under “—Legal Reserve”).

Legal ~~Reserve~~Reserve.. Pursuant to French law, we must allocate at least 5% of our unconsolidated net profit for each year to our legal reserve fund before dividends may be paid with respect to that year. Such allocation is compulsory until the amount in the legal reserve is equal to 10% of the aggregate par value of our issued and outstanding share capital. This restriction on the payment of dividends also applies to our French subsidiaries on an unconsolidated basis.

Approval of ~~Dividends~~Dividends.. Pursuant to French law, our board of directors may propose a dividend and/or reserve distribution for approval by the shareholders at the annual ordinary general meeting.

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Our board of directors may distribute interim dividends after the end of the fiscal year but before the approval of the financial statements for the relevant fiscal year when the interim balance sheet, established during such year and examined by an auditor, reflects that we have earned distributable profits since the close of the last financial year, after recognizing the necessary depreciation and provisions and after deducting prior losses, if any, and the sums to be allocated to reserves, as required by law or theBy-laws, and including any retained earnings. The amount of such interim dividends may not exceed the amount of the profit so defined.

Distribution of ~~Dividends~~Dividends.. Dividends are distributed to shareholders proportionally to their shareholding interests. In the case of interim dividends, distributions are made to shareholders on the date set by our board of directors during the meeting in which the distribution of interim dividends is approved. The actual dividend payment date is decided by the shareholders at an ordinary general shareholders’ meeting or by our board of directors in the absence of such a decision by the shareholders. Shareholders that own shares on the actual payment date are entitled to the dividend.

Timing of ~~Payment~~Payment.. Pursuant to French law, dividends must be paid within a maximum period of nine months following the end of the relevant fiscal year. An extension of such timeframe may be granted by court order. Dividends that are not claimed within a period of five years after the payment date will be deemed to expire and revert to the French state.

Voting ~~Rights~~Rights.. Each of our ordinary shares entitles its holder to vote and be represented in the shareholders’ meetings in accordance with the provisions of French law and of ourBy-laws. The ownership of a share implies the acceptance of ourBy-laws and any decision of our shareholders.

In general, each shareholder is entitled to one vote per share at any general shareholders’ meeting. However, ourBy-Laws provide that all shares held in registered form ~~(actions nominatives)~~(actions nominatives) for more than two years will be granted double voting rights.

Under French law, treasury shares or shares held by entities controlled by us are not entitled to voting rights and are not taken into account for purposes of quorum calculation.

Rights to Share in Our ~~Profit~~Profit.. Under French law, each ordinary share entitles its holder to a portion of the corporate profits and assets proportional to the amount of share capital represented thereby.

Rights to Share in the Surplus in the Event of ~~Liquidation~~Liquidation.. If we are liquidated, any assets remaining after payment of our debts, liquidation expenses and all of our remaining obligations will first be used to repay in full the par value of our outstanding shares. Any surplus will then be distributed among shareholders proportionally to their shareholding in our company.

Repurchase and Redemption of ~~Shares~~Shares.. Under French law, we may acquire our own shares. Such acquisition may be challenged on the ground of market abuse regulations. However, Regulation 596/2014 of April 16, 2014 and its related delegated regulations (MAR) provides for safe harbor exemptions when the acquisition is made (i) under abuy-back program to be

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authorized by the shareholders in accordance with the provisions of Article L.225-209 of the French Commercial Code and with the General Regulations of theAutorité des marchés financiers or AMF and (ii) for one of the following purposes which shall be provided for in thebuy-back program:

to decrease our share capital;

to meet our obligations arising from debt financial instruments issued by us that are exchangeable into shares;

to meet our obligations arising from share option programs, or other allocations of shares, to our employees or to our managers ~~our~~or the employees or managers of our affiliate.

In addition, we benefit from a simple exemption when the acquisition is made under a liquidity contract complying with the general regulations of, and market practices accepted by, the AMF.

All other purposes, and especially sharebuy-backs made for external growth operations in pursuance ofArticle L.225-209 of the French Commercial Code, while not forbidden, must be pursued in strict compliance of market manipulation and insider dealing rules.

Under the Market Abuse Regulation 596/2014 of ~~August~~April 16, 2014 (MAR) and in accordance with the General Regulations of the AMF, a corporation shall report to the ~~competent authority of the trading value on which the shares have been admitted to trading or are traded,~~AMF, no later than by the end of the seventh daily market session following the date of the execution of the transaction, all the transactions relating to thebuy-back program, in a detailed form and in an aggregated form. In addition, we shall provide to the AMF, on a monthly basis, and to the public, on a quarterly basis, a summary report of the transactions made under a liquidity contract.

The decision to repurchase shares in order to decrease our share capital shall not be driven by losses and a purchase offer shall be made to all shareholders on a pro rata basis, with the approval of the shareholders at the extraordinary general meeting deciding the capital reduction; in this case, the shares repurchased must be cancelled within one month from their repurchase date.

When shares are repurchased in order to provide shares for distribution to employees or managers under a profit-sharing, free share or share option plan, the shares repurchased must be distributed within 12 months from their repurchase failing which they must be cancelled.

In any case, no such repurchase of shares may result in us holding, directly or through a person acting on our behalf, more than 10% of our issued share capital. Shares repurchased by us continue to be deemed “issued” under French law but are not entitled to dividends or voting rights so long as we hold them directly or indirectly, and we may not exercise the preemptive rights attached to them.

Sinking Fund ~~Provisions~~Provisions.. OurBy-laws do not provide for any sinking fund provisions.

Liability to Further Capital ~~Calls~~Calls.. Shareholders are liable for corporate liabilities only up to the par value of the shares they hold; they are not liable to further capital calls.

Requirements for Holdings Exceeding Certain ~~Percentages~~Percentages.. There are no such requirements, except as described under the section of this Annual Report titled “—Form, Holding and Transfer of Shares—Ownership of Shares and ADSs byNon-French Persons.”

Actions Necessary to Modify Shareholders’ Rights

Shareholders’ rights may be modified as allowed by French law. Only the extraordinary shareholders’ meeting is authorized to amend any and all provisions of ourBy-laws. It may not, however, increase any of the shareholders’ commitments without the prior approval of each shareholder.

Special Voting Rights of Warrant Holders

Under French law, the holders of warrants of the same class (i.e., warrants that were issued at the same time and with the same rights), including founders’ warrants, are entitled to vote as a separate class at a general

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meeting of that class of warrant holders under certain circumstances, principally in connection with any proposed modification of the terms and conditions of the class of warrants or any proposed issuance of preferred shares or any modification of the rights of any outstanding class or series of preferred shares.

Rules for Admission to and Calling Annual Shareholders’ Meetings and Extraordinary Shareholders’ Meetings

Access to, Participation in and Voting Rights at Shareholders’ ~~Meetings~~Meetings.. The right to participate in a shareholders’ meeting is granted to all the shareholders, regardless of the number of shares they hold, whose shares are fully paid up and for whom a right to attend shareholders’ meetings has been established by registration of their shares in the names or names of the authorized intermediary acting on their behalf on the second business day prior to the shareholders’ meeting at midnight (Paris time), either in the registered shares accounts held by the Company or in the bearer shares accounts held by the authorized intermediary.

Each shareholder may attend the meetings and vote (1) in person, or (2) by granting a proxy to his or her spouse, his or her partner with whom he or she has entered into a civil union, or another shareholder, for physical persons, or any person that they may choose, for legal entities, or (3) by sending a proxy to us without indication of the beneficiary (in which case such proxy shall be cast in favor of the resolutions supported by the board of directors), or (4) by correspondence, or (5) by videoconference or another means of telecommunication organized by the board of directors and allowing identification of the relevant shareholder in accordance with applicable laws.

Shareholders may, in accordance with legal and regulatory requirements, send their vote or proxy, either by hard copy or via telecommunications means. Such vote or proxy must be received (1) at least three days prior to the meeting, in the case of hard copies, (2) by 3:00 p.m. (Paris time) on the day before the meeting, in the case of, electronic votes by email, (3) by the date of the meeting, in the case of a proxy granted to a designated person, and (4) by 3:00 p.m. (Paris time) on the day before the meeting, in the case of proxies without a designated attorney and therefore granted to the chairman of the meeting.

Shareholders sending their vote within the applicable time limit, using the form provided to them by us for this purpose, are deemed present or represented at the shareholders’ meeting for purposes of quorum and majority calculation.

The voting by correspondence form addressed by a shareholder is only valid for a single meeting or for successive meetings convened with the same agenda. To better understand the voting rights of the ADSs, you should carefully read the section of the accompanying prospectus “Description of American Depositary Shares”.

Notice of Annual Shareholders’ ~~Meetings~~Meetings.. Shareholders’ meetings are convened by our board of directors, or, failing that, by our statutory auditors, or by a court appointed agent or liquidator in certain circumstances, or by the majority shareholder in capital or voting rights following a public tender offer or exchange offer or the transfer of a controlling block on the date decided by the board of directors or the relevant person. Meetings are held at our registered offices or at any other location indicated in the convening notice. A meeting notice (avis de réunion) is published in the French Journal of Mandatory Statutory Notices (BALO) at least 35 days prior to the date of the shareholders’ meeting.

Additionally, a convening notice (avis de convocation) is published at least fifteen days prior to the date of the meeting in a legal gazette of the department in which the registered office of the company is located and in the French Journal of Mandatory Statutory Notices (BALO). Further, shareholders having held registered shares (actions nominatives) for at least one month at the time of the convening notice must be convened individually, by regular letter (or by registered letter if requested by the relevant shareholder) sent to their last known address.

When the shareholders’ meeting cannot deliberate due to the lack of the required quorum, the second meeting must be called at least ten days in advance in the same manner as used for the first notice.

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All notices to the shareholders must further specify the conditions under which the shareholders may vote by correspondence.

Agenda and Conduct of Annual Shareholders’ ~~Meetings~~Meetings.. The agenda of the shareholders’ meeting shall appear in the notice to convene the meeting. The shareholders’ meeting may only deliberate on the items on the agenda except for the removal of directors and the appointment of their successors, which may be put to vote by any shareholder during any shareholders’ meeting. One or more shareholders representing the percentage of share capital required by French law (currently 5%), and acting in accordance with legal requirements and within applicable time limits, may request the inclusion of items or proposed resolutions on the agenda.

Shareholders’ meetings shall be chaired by the Chairman of the board of directors or, in his or her absence, by a director appointed for this purpose by the board of directors; failing which, the meeting itself shall elect a Chairman. Vote counting shall be performed by the two members of the meeting who are present and accept such duties, who represent, either on their own behalf or as proxies, the greatest number of votes.

Ordinary Shareholders’ ~~Meeting~~Meeting.. Ordinary shareholders’ meetings are those meetings called to make any and all decisions that do not result in a modification of ourBy-laws. An ordinary shareholders’ meeting shall be convened at least once a year within six months of the end of each fiscal year in order to approve the annual and consolidated accounts for the relevant fiscal year or, in case of postponement, within the period established by court order. Upon first notice, the meeting may validly deliberate only if the shareholders present or represented by proxy or voting by mail represent at leastone-fifth of the shares entitled to vote. Upon second notice, no quorum is required. Decisions are made by a majority of the votes held by the shareholders present, represented by proxy, or voting by mail. Abstentions will have the same effect as a “no” vote.

Extraordinary Shareholders’ ~~Meeting~~Meeting.. Only an extraordinary shareholders’ meeting is authorized to amend our ~~Bylaws.~~By-laws. It may not, however, increase shareholders’ commitments without the approval of each shareholder. Subject to the legal

provisions governing share capital increases from reserves, profits or share premiums, the resolutions of the extraordinary meeting will be valid only if the shareholders present, represented by proxy or voting by mail represent at leastone-fourth of all shares entitled to vote upon first notice, orone-fifth upon second notice. If the latter quorum is not reached, the second meeting may be postponed to a date no later than two months after the date for which it was initially called. Decisions are made by atwo-thirds majority vote of the shareholders present, represented by proxy, or voting by mail. Abstentions will have the same effect as a “no” vote.

In addition to the right to obtain certain information regarding us at any time, any shareholder may, from the date on which a shareholders’ meeting is convened until the fourth business day preceding the date of the shareholders’ meeting, submit written questions relating to the agenda for the meeting to our board of directors. Our board of directors is required to respond to these questions during the meeting, except if the answers of the board are posted on the website of the Company at the latest at the end of the shareholder’s meeting.

Provisions Having the Effect of Delaying, Deferring or Preventing a Change in Control of the Company

Provisions contained in ourBy-laws and the corporate laws of France, the country in which we are incorporated, could make it more difficult for a third-party to acquire us, even if doing so might be beneficial to our shareholders. In addition, provisions of French law and ourBy-laws impose various procedural and other requirements which could make it more difficult for shareholders to effect certain corporate actions. These provisions include the following:

a merger (i.e., in a French law context, astock-for-stock exchange after which our company would be dissolved without being liquidated into the acquiring entity and our shareholders would become shareholders of the acquiring entity) of our company into a company incorporated in the European Union would require the approval of our board of directors as well as a two thirds majority of the votes held by the shareholders present, represented by proxy or voting by mail at the relevant meeting;

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a merger of our company into a company incorporated outside of the European Union would require the unanimous approval of our shareholders;

under French law, a cash merger is treated as a share purchase and would require the consent of each participating shareholder;

our shareholders have granted and may grant in the future our board of directors broad authorizations to increase our share capital or to issue additional ordinary shares or other securities (for example, warrants) to our shareholders, the public or qualified investors, including as a possible defence following the launching of a tender offer for our shares;

our shareholders have preferential subscription rights proportional to their shareholding in our company on the issuance by us of any additional shares or securities giving the right, immediately or in the future, to new shares for cash or aset-off of cash debts, which rights may only be waived by the extraordinary general meeting (by atwo-thirds majority vote) of our shareholders or on an individual basis by each shareholder;

our board of directors has the right to appoint directors to fill a vacancy created by the resignation or death of a director, subject to the approval by the shareholders of such appointment at the next shareholders’ meeting, which prevents shareholders from having the sole right to fill vacancies on our board of directors;

our board of directors can only be convened by our chairman or our managing director, if any, or, when no board meeting has been held for more than two consecutive months, by directors representing at least one third of the total number of directors;

our board of directors’ meetings can only be regularly held if at least half of the directors attend either physically or by way of videoconference or teleconference enabling the directors’ identification and ensuring their effective participation in the board of directors’ decisions;

our shares take the form of bearer securities or registered securities, if applicable legislation so permits, according to the shareholder’s choice. Issued shares are registered in individual accounts opened by us or any authorized intermediary (depending on the form of such shares), in the name of each shareholder and kept according to the terms and conditions laid down by the legal and regulatory provisions;

•

under French law, anon-French resident as well as any French entity controlled bynon-French residents may have to file a declaration for statistical purposes with the Bank of France (Banque de France) following the date of certain ~~direct or indirect~~foreign investments in ~~us;~~us. Additionally, certain investments in a French

company relating to certain strategic industries by individual or entities not residents in a member State of the European Union are subject to the prior authorization of the French Ministry of Economy — see the section of this Annual Report titled “Ownership of Shares and ADSs byNon-French Persons”;

approval of at least a majority of the votes held by shareholders present, represented by a proxy, or voting by mail at the relevant ordinary shareholders’ general meeting is required to remove directors with or without cause;

advance notice is required for nominations to the board of directors or for proposing matters to be acted upon at a shareholders’ meeting, except that a vote to remove and replace a director can be proposed at any shareholders’ meeting without notice;

in the event where certain ownership thresholds would be crossed, a number of disclosures should be made by the relevant shareholder ~~and can impose~~in addition to certain obligations; see the section of this Annual Report titled “—Declaration of Crossing of Ownership Thresholds”;

transfers of shares shall comply with applicable insider trading rules; ~~and~~

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Declaration of Crossing of Ownership Thresholds

Subject to requirements of French law, ourBy-laws do not require any specified disclosure by shareholders that cross ownership thresholds with respect to our share capital, except as described under the section of this Annual Report titled “—Form, Holding and Transfer of Shares—Ownership of Shares and ADSs byNon-French Persons.”

The absence of specific requirement in ourBy-laws is without prejudice to the following disclosures which are applicable to us according to French legal and regulatory provisions, it being provided that the following is a summary which is therefore not intended to be a complete description of applicable rules under French law:

Shareholders must make a declaration to us no later than the fourth trading day after such shareholder crosses the following thresholds: 5%, 10%, 15%, 20%, 25%, 30%, 33.33%, 50%, 66.66%, 90% and 95%.

Shareholders must make a declaration to the AMF no later than the fourth trading day after such shareholder crosses the following thresholds: 50% and 95%.

The above obligations of declaration apply when crossing each of the above-mentioned thresholds in an upward or downward direction.

In case of failure to declare shares or voting rights exceeding the fraction that should have been declared, such shares shall be deprived of voting rights at shareholders’ meetings for any meeting that would be held until the expiry of a period of two years from the date of regularization of the notification in accordance with Article L.233-14 of the French Commercial Code. Additional sanctions may apply pursuant to Article L.621-15 of the French Monetary and Financial Code.

Subject to certain exemptions, any shareholder crossing, alone or acting in concert, the 50% threshold shall file a mandatory public tender offer.

Changes in Share Capital

Increases in Share ~~Capital~~Capital.. Pursuant to French law, our share capital may be increased only with shareholders’ approval at an extraordinary general shareholders’ meeting following the recommendation of our board of directors. The shareholders may delegate to our board of directors either the authority(délégation de compé~~tence~~tence)) or the power(délégation de ~~pouvoir~~pouvoir)) to carry out any increase in share capital in accordance with applicable laws.

Increases in our share capital may be effected by:

issuing additional shares;

increasing the par value of existing shares;

creating a new class of equity securities; and

exercising the rights attached to securities giving access to the share capital.

Increases in share capital by issuing additional securities may be effected through one or a combination of the following:

issuances in consideration for cash;

issuances in consideration for assets contributed in kind;

issuances through an exchange offer;

issuances by conversion of previously issued debt instruments;

issuances by capitalization of profits, reserves or share premium; and

subject to certain conditions, issuances by way of offset against debt incurred by us.

Decisions to increase the share capital through the capitalization of reserves, profits and/or share premium require shareholders’ approval at an extraordinary general shareholders’ meeting, acting under the quorum and majority requirements applicable to ordinary shareholders’ meetings. Increases in share capital effected by an increase in the par value of shares require unanimous approval of the shareholders, unless effected by capitalization of reserves, profits or share premium. All other capital increases require shareholders’ approval at an extraordinary general shareholders’ meeting acting under the regular quorum and majority requirements for such meetings.

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Reduction in Share Capital. Pursuant to French law, any reduction in our share capital requires shareholders’ approval at an extraordinary general shareholders’ meeting following the recommendation of our board of directors. The share capital may be reduced either by decreasing the par value of the outstanding shares or by reducing the number of outstanding shares. The number of outstanding shares may be reduced by the repurchase and cancellation of shares. Holders of each class of shares must be treated equally unless each affected shareholder agrees otherwise.

Preferential Subscription ~~Right~~Right.. According to French law, if we issue additional shares or securities giving right, immediately or in the future, to new shares for cash, current shareholders will have preferential subscription rights to these securities on apro ratabasis. Preferential subscription rights entitle the individual or entity that holds them to subscribe proportionally to the number of shares held by them to the issuance of any securities increasing, or that may result in an increase of, our share capital by means of a cash payment or aset-off of cash debts. The preferential subscription rights may be transferred and/or sold during the subscription period relating to a particular offering. Pursuant to French law, the preferential subscription rights will be transferable during a period starting two working days prior to the opening of the subscription period and ending two working days prior to the closing of the subscription period.

The preferential subscription rights with respect to any particular offering may be waived at an extraordinary general meeting by a two thirds vote of our shareholders or individually by each shareholder. Our board of directors and our independent auditors are required by French law to present reports to the shareholders’ meeting that specifically address any proposal to waive the preferential subscription rights.

Further, to the extent permitted under French law, we may seek, during an extraordinary general shareholders’ meeting, the approval of the shareholders to waive their preferential subscription rights in order to authorize the board of directors to issue additional shares and/or other securities convertible or exchangeable into shares.

Form, Holding and Transfer of Shares

Form of Shares. Pursuant to ourBy-laws, shares may be held in registered or bearer form, at each shareholder’s discretion.

Further, in accordance with applicable legal and regulatory provisions, we may request at any time from the authorized intermediary responsible for holding our shares the name or, in the case of a legal entity, the corporate name, nationality and address of holders of securities, giving immediate or future access to voting rights at our shareholders’ meetings, the number of securities they own and, where applicable, the restrictions attaching to such securities.

Holding of Shares. In accordance with French law concerning the “dematerialization” of securities, the ownership rights of shareholders are represented by book entries instead of share certificates. Shares are registered in individual accounts opened by us or any authorized intermediary, in the name of each shareholder and kept according to applicable legal and

regulatory provisions.

Ownership of Shares and ADSs byNon-French Persons. Neither the French Commercial Code nor ourBy-laws presently impose any restrictions on the right ofnon-French residents ornon-French shareholders to own and vote shares.

However,non-French residents must file a declaration for statistical purposes with the Bank of France (Banque de France) within twenty working days following the date of certain direct foreign investments in us, including any purchase of our ADSs. In particular, such filings are required in connection with investments exceeding €15,000,000 that lead to the acquisition of at least 10% of our Company’s share capital or voting rights or cross such 10% threshold. Violation of this filing requirement may be sanctioned by five years of imprisonment and a fine of up to twice the amount of the relevant investment. This amount may be increased fivefold if the violation is made by a legal entity.

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~~Moreover,~~Further, any investment (i) by an individual or entity located in a country that is not a member State of the European Union or of a member State of the European Economic Area having entered into a convention on administrative assistance against tax evasion and fraud with France, or by a French citizen not residing in France, and (ii) that will result in the relevant investor acquiring the control of, all or part of a business of, or more than 33.33% of the share capital or voting rights of, a company registered in France and developing activities in certain ~~foreign investments in companies incorporated under French laws are~~strategic industries, such as, energy, public health, telecommunications, artificial intelligence, cybersecurity, robotics, data collection ordual-use goods and technology is subject to the prior authorization ~~from~~by the French ~~Minister~~Ministry of Economy. In the ~~Economy, where all or part~~absence of such authorization, the ~~target’s business~~relevant investment shall be deemed null and ~~activity relate to a strategic sector, such as energy, transportation, public health, telecommunications, etc.~~void.

Assignment and Transfer of Shares. Shares are freely negotiable, subject to applicable legal and regulatory provisions (including, in particular, the prohibition on insider trading).

Listing

Our ADSs have been listed on the Nasdaq Global Market under the symbol “CLLS” and our ordinary shares have been listed on the Euronext Growth market of Euronext in Paris under the symbol “ALCLS”.

Transfer Agent and Registrar

The transfer agent and registrar for our ADSs is Citibank, N.A. The transfer agent and registrar for our ordinary shares is Société Générale Securities Services.

C.	Material Contracts

We entered into an underwriting agreement among Merrill Lynch, Pierce, Fenner & Smith Incorporated and Jefferies LLC, as representatives of the underwriters, on March 24, 2015, with respect to the ADSs sold in our initial public offering. We have agreed to indemnify the underwriters against certain liabilities, including liabilities under the Securities Act, and to contribute to payments the underwriters may be required to make in respect of such liabilities.

We entered into an underwriting agreement among Goldman Sachs & Co. LLC, Citigroup Global Markets Inc. and Barclays Capital, Inc. as representatives of the underwriters, on April 4, 2018, with respect to the ADSs sold in ourfollow-on offering. We have agreed to indemnify the underwriters against certain liabilities, including liabilities under the Securities Act, and to contribute to payments the underwriters may be required to make in respect of such liabilities.

For additional information on our material contracts, please ~~see Item~~refer to items 4 and ~~Item~~ 6 of this Annual Report.

D.	Exchange Controls

Under current French foreign exchange control regulations there are no limitations on the amount of cash payments that we may remit to residents of foreign countries. Laws and regulations concerning foreign exchange controls do, however, require that all payments or transfers of funds made by a French resident to anon-resident such as dividend payments be handled by an accredited intermediary. All registered banks and substantially all credit institutions in France are accredited intermediaries.

Taxation

Material U.S. Federal Income Tax Considerations

The following is a discussion of the material U.S. federal income tax consequences of owning and disposing of ADSs. This summary does not address any aspect of U.S. federalnon-income tax laws, such as U.S. federal estate and gift tax laws, or state, local ornon-U.S. tax laws, and does not purport to be a comprehensive description of all of the U.S. tax considerations that may be relevant to particular holders.

The discussion applies to you only if you hold the ADSs as capital assets for U.S. federal income tax purposes (generally, for investment). This section does not apply to you if you are a member of a special class of holders subject to special tax rules, including:

a broker;

a dealer in securities, commodities or foreign currencies;

a trader in securities that elects to use amark-to-market method of accounting for your securities holdings;

a bank or other financial institution;

atax-exempt organization;

an insurance company;

a real estate investment trust;

a controlled foreign corporation;

a passive foreign investment company;

a regulated investment company;

an investor who is a U.S. expatriate, former U.S. citizen or former long term resident of the United States;

a mutual fund;

an individual retirement or othertax-deferred account;

a holder liable for alternative minimum tax;

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a holder that actually or constructively owns 10% or more, by voting power, ~~or value,~~ of our voting stock;

a partnership or other pass-through entity for U.S. federal income tax purposes;

a holder that holds ADSs as part of a straddle, hedging, constructive sale, conversion or other integrated transaction for U.S. federal income tax purposes; or

a U.S. holder whose functional currency is not the U.S. Dollar.

This section is based on the Internal Revenue Code of 1986, as amended, or (the Code), existing and proposed income tax regulations issued under the Code, legislative history, and judicial and administrative interpretations thereof, all as of the date of this Annual Report. All of the foregoing are subject to change at any time, and any change could be retroactive and could affect the accuracy of this discussion. In addition, the application and interpretation of certain aspects of the passive foreign investment company, or PFIC, rules, referred to below, require the issuance of regulations which in many instances have not been promulgated and which may have retroactive effect. There can be no assurance that any of these regulations will be enacted or promulgated, and if so, the form they will take or the effect that they may have on this discussion. This discussion is not binding on the U.S. Internal Revenue Service, or IRS, or the courts. No ruling has been or will be sought from the IRS with respect to the positions and issues discussed herein, and there can be no assurance that the IRS or a court will not take a different position concerning the U.S. federal income tax consequences of an investment in the ADSs or that any such position would not be sustained.

~~INVESTORS~~YOU SHOULD CONSULT ~~THEIR~~YOUR OWN TAX ADVISORS CONCERNING THE U.S. FEDERAL, STATE, LOCAL ANDNON-U.S. TAX CONSEQUENCES OF OWNING AND DISPOSING OF THE ADSs IN ~~THEIR~~YOUR PARTICULAR SITUATIONS, INCLUDING ANY CONSEQUENCES UNDER THE RECENTLY ENACTED LEGISLATION ~~KNOWN~~KNOWS AS THE TAX CUTS AND JOBS ACT.

You are a “U.S. holder” if you are a beneficial owner of ADSs and you ~~are:~~are or are treated for U.S. federal income tax purposes as:

a citizen or resident of the United States;

a corporation ~~(or other entity treated as a corporation for U.S. federal income tax purposes)~~ created or organized under the laws of the United States, any state thereof, or the District of Columbia;

an estate whose income is subject to U.S. federal income tax regardless of its source; or

a trust if (1) a U.S. court can exercise primary supervision over the trust’s administration and one or more U.S. persons are authorized to control all substantial decisions of the trust or (2) has a valid election in effect under applicable U.S. Treasury regulations to be treated as a U.S. person for U.S. federal income tax purposes.

In addition, this discussion is limited to U.S. holders who are not resident in France for purposes of the Income Tax Treaty between the United States and France.

If a partnership (including for this purpose any entity treated as a partnership for U.S. federal income tax purposes) is a beneficial owner of the ADSs, the U.S. tax treatment of a partner in the partnership generally will depend on the status of the partner and the activities of the partnership. A holder of the ADSs that is a partnership and partners in such a partnership should consult their own tax advisors concerning the U.S. federal income tax consequences of owning and disposing of ADSs.

A“non-U.S. holder” is a beneficial owner of ADSs that is neither a U.S. holder nor a partnership for U.S. federal income tax purposes.

Generally, holders of ADSs should be treated for U.S. federal income tax purposes as holding the ordinary shares represented by the ADSs. Accordingly, no gain or loss will be recognized upon an exchange of ordinary

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shares for ADSs or an exchange of ADSs for ordinary shares. The U.S. Treasury has expressed concerns that intermediaries in the chain of ownership between the holder of an ADS and the issuer of the security underlying the ADS may be taking actions that are inconsistent with the claiming of foreign tax credits for U.S. holders of ADSs. Accordingly, the credibility of foreign taxes, if any, as described below, could be affected by actions taken by intermediaries in the chain of ownership between the holder of an ADS and the company.

PFIC Considerations

The Code provides special rules regarding certain distributions received by U.S. persons with respect to, and sales, exchanges and other dispositions, including pledges, of, shares of stock (including ordinary shares represented by ADSs) in a PFIC. Anon-U.S. corporation will be treated as a PFIC for any taxable year in which either: (1) at least 75 percent of its gross income is “passive income” or (2) at least 50 percent of its gross assets during the taxable year (based on the average of the fair market values of the assets determined at the end of each quarterly period) are “passive assets,” which generally means that they produce passive income or are held for the production of passive income. Passive income for this purpose generally includes, among other things, dividends, interest, rents, royalties, gains from commodities and securities transactions, and gains from assets that produce passive income. In determining whether a foreign corporation is a PFIC, a pro rata portion of the income and assets of each corporation in which it owns, directly or indirectly, at least a 25% interest (by value) is taken into account.

~~While we may have been a PFIC for previous taxable years and although~~Although the matter is not free from doubt, we do not believe that we were a PFIC for U.S. federal income tax purposes for the ~~2017~~2018 taxable ~~year and~~year. No assurances may be given at this time ~~do not anticipate becoming a~~as to our PFIC for ~~the current or future taxable years.~~2019. However, our PFIC status must be determined annually and therefore is subject to change. Because this determination is made annually at the end of each taxable year and is dependent upon a number of factors, some of which are beyond our control, including the amount and nature of our income, as well as on the market valuation of our assets and our spending schedule for our cash balances, and because certain aspects of the PFIC rules are not entirely certain, there can be no assurance that we are or are not a PFIC or that the IRS will agree with our conclusion regarding our PFIC status. If we are not a PFIC during any taxable year in which you hold ADSs, then the remainder of the discussion under “Taxation—Material U.S. Federal Income Tax Considerations,” outside of this ~~“PFIC~~“—PFIC Considerations” portion may be relevant to you. U.S. holders should consult their tax advisors as to the applicability of the PFIC rules.

A U.S. holder that holds ADSs during any taxable year in which we qualify as a PFIC is subject to special tax rules with respect to (a) any gain realized on the sale, exchange or other disposition of the ADSs and (b) any “excess distribution” by the corporation to the holder, unless the holder elects to treat the PFIC as a “qualified electing fund,” or QEF, or makes a“mark-to-market” election, each as discussed below. An “excess distribution” is that portion of a distribution with respect to ADSs that exceeds 125% of the annual average of such distributions over the preceding three-year period or, if shorter, the U.S. holder’s holding period for its ADSs. Excess distributions and gains on the sale, exchange or other disposition of ADSs of a corporation which was a PFIC at any time during the U.S. holder’s holding period are allocated ratably to each day of the U.S. holder’s holding period. Amounts allocated to the taxable year in which the disposition occurs and amounts allocated to any period in the shareholder’s holding period before the first day of the first taxable year that the corporation was a PFIC will be taxed as ordinary income (rather than capital gain) earned in the taxable year of the disposition. Amounts allocated to each of the other taxable years in the U.S. holder’s holding period are not included in gross income for the year of the disposition, but are subject to the highest ordinary income tax rates in effect for individuals or corporations, as applicable, for each such year and the interest charge

generally applicable to income tax deficiencies will be imposed on the resulting tax attributable to each year. The tax liability for amounts allocated to years before the year of disposition or “excess distribution” cannot be offset by any net operating losses for such years, and gains (but not losses) realized on the sale of the ADSs cannot be treated as capital, even if a U.S. holder held such ADSs as capital assets.

If we are a PFIC for any taxable year during which a U.S. holder holds ADSs, then we generally will continue to be treated as a PFIC with respect to the holder for all succeeding years during which such holder

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holds ADSs, even if we no longer satisfy either the passive income or passive asset tests described above, unless the U.S. holder terminates this deemed PFIC status by making a “deemed sale” election. If such election is made, a U.S. holder will be deemed to have sold the ADSs at their fair market value on the last day of the last taxable year for which we were a PFIC, and any gain from such deemed sale would be subject to the excess distribution rules as described above. After the deemed sale election, the ADSs with respect to which the deemed sale election was made will not be treated as shares in a PFIC unless we subsequently become a PFIC.

If we are or become a PFIC, the excess distribution rules may be avoided if a U.S. holder makes a QEF election effective beginning with the first taxable year in the holder’s holding period in which we are treated as a PFIC with respect to such holder. A U.S. holder that makes a QEF election with respect to a PFIC is required to include in income its pro rata share of the PFIC’s ordinary earnings and net capital gain as ordinary income and capital gain, respectively, subject to a separate election to defer payment of taxes, which deferral is subject to an interest charge. If a foreign corporation ceases to be a PFIC, the U.S. holder’s QEF election would no longer require an annual income inclusion. However, cessation of a foreign corporation’s status as a PFIC will not terminate a QEF election and if the corporation becomes a PFIC again, an annual income inclusion may be required.

In general, a U.S. holder makes a QEF election by attaching a completed IRS Form 8621 to a timely filed (taking into account any extensions) U.S. federal income tax return for the year beginning with which the QEF election is to be effective. In certain circumstances, a U.S. holder may be able to make a retroactive QEF election. A QEF election can be revoked only with the consent of the IRS. In order for a U.S. holder to make a valid QEF election, the corporation must annually provide or make available to the holder certain information. For any taxable year in which we are a PFIC, we will determine whether we will provide to U.S. holders the information required to make a QEF election.

As an alternative to making a QEF election, a U.S. holder may make a“mark-to-market” election with respect to its ADSs if the ADSs meet certain minimum trading requirements, as described below. If a U.S. holder makes a validmark-to-market election for the first taxable year in which such holder holds (or is deemed to hold) ADSs in a corporation and for which such corporation is determined to be a PFIC, such holder generally will not be subject to the PFIC rules described above in respect of its ADSs. Instead, a U.S. holder that makes amark-to-market election will be required to include in income each year an amount equal to the excess, if any, of the fair market value of the ADSs that the holder owns as of the close of the taxable year over the holder’s adjusted tax basis in the ADSs. The U.S. holder will be entitled to a deduction for the excess, if any, of the holder’s adjusted tax basis in the ADSs over the fair market value of the ADSs as of the close of the taxable year; provided, however, that the deduction will be limited to the extent of any netmark-to-market gains with respect to the ADSs included by the U.S. holder under the election for prior taxable years. The U.S. holder’s basis in the ADSs will be adjusted to reflect the amounts included or deducted pursuant to the election. Amounts included in income pursuant to amark-to-market election, as well as gain on the sale, exchange or other disposition of the ADSs, will be treated as ordinary income. The deductible portion of anymark-to-market loss, as well as loss on a sale, exchange or other disposition of ADSs to the extent that the amount of such loss does not exceed netmark-to-market gains previously included in income, will be treated as ordinary loss. If a U.S. holder makes a validmark-to-market election, any distributions made by us in a year in which we are a PFIC would generally be subject to the rules discussed below under “—Taxation of Dividends,” except the lower rate applicable to qualified dividend income would not apply. If we are not a PFIC when a U.S. holder has amark-to-market election in effect, gain or loss realized by a U.S. holder on the sale of our ADSs will be a capital gain or loss and taxed in the manner described below under “—Taxation of Sale, Exchange or other Disposition of ADSs.”

Themark-to-market election applies to the taxable year for which the election is made and all subsequent taxable years, unless the ADSs cease to meet applicable trading requirements (described below) or the IRS consents to its revocation. The excess distribution rules generally do not apply to a U.S. holder for taxable years for which amark-to-market election is in effect. If we are a PFIC for any year in which the U.S. holder owns ADSs but before amark-to-market election is made, the interest charge rules described above will apply to any

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mark-to-market gain recognized in the year the election is made. Generally, if a foreign corporation ceases to be a PFIC, the U.S. holder’smark-to-market election would no longer require the income inclusion described above. However, cessation of a foreign corporation’s status as a PFIC will not terminate amark-to-market election and if the corporation becomes a PFIC again,mark-to-market income inclusions may be required.

Amark-to-mark election is available only if the ADSs are considered “marketable” for these purposes. ADSs will be marketable if they are regularly traded on a national securities exchange that is registered with the SEC (such as the Nasdaq Global Market) or on anon-U.S. exchange or market that the IRS determines has rules sufficient to ensure that the market price

represents a legitimate and sound fair market value. For these purposes, ADSs will be considered regularly traded during any calendar year during which more than a de minimis quantity of the ADSs is traded on at least 15 days during each calendar quarter. Any trades that have as their principal purpose meeting this requirement will be disregarded. Each U.S. holder should ask its own tax advisor whether amark-to-market election is available or desirable.

If we are a PFIC for any year in which a U.S. holder holds ADSs, such U.S. holder must generally file an IRS Form 8621 annually. A U.S. holder must also provide such other information as may be required by the U.S. Treasury Department if the U.S. holder (1) receives certain direct or indirect distributions from a PFIC, (2) recognizes gain on a direct or indirect disposition of ADSs, or (3) makes certain elections (including a QEF election or amark-to-market election) reportable on IRS Form 8621.

Under attribution rules, if we are a PFIC, U.S. holders of our ADSs will be deemed to own their proportionate shares of our subsidiaries that are PFICs, if any. ~~In such case, assuming~~Like the determination of whether we are a PFIC, the determination of whether any of our subsidiaries is a PFIC is made annually at the end of each taxable year. Assuming a U.S. holder does not receive from ~~such~~a PFIC subsidiary the information that the U.S. holder needs to make a QEF election with respect to such a subsidiary, a U.S. holder generally will be deemed to own a portion of the shares of such lower-tier PFIC and may incur liability for a deferred tax and interest charge if we receive a distribution from, or dispose of all or part of our interest in, or the U.S. holder otherwise is deemed to have disposed of an interest in, the lower-tier PFIC, even though the U.S. holder has not received the proceeds of those distributions or dispositions directly. ~~However, as discussed under “Item 4.C—Organization Structure”, as of December 31, 2017, we~~We currently do not haveany non-U.S. ~~corporations as subsidiary. Accordingly, for the 2017 taxable year, none of our~~ subsidiaries ~~would have constituted lower-tier PFICs if we were a PFIC.~~that could be PFIC subsidiaries.

U.S. holders are urged to consult their tax advisors as to our status as a PFIC, and, if we are treated as a PFIC, as to the effect on them of, and the reporting requirements with respect to, the PFIC rules and the desirability of making, and the availability of, either a QEF election or amark-to-market election with respect to our ADSs.

Taxation of Dividends

U.S. Holders. Subject to the PFIC rules described above under “—PFIC Considerations,” if you are a U.S. holder, you must include in your gross income the gross amount of any distributions of cash or property (other than certainpro rata distributions of ADSs) with respect to ADSs, to the extent the distribution is paid out of our current or accumulated earnings and profits, as determined for U.S. federal income tax purposes. A U.S. holder must include the dividend as ordinary income at the time of actual or constructive receipt. The amount of any dividend income paid in Euro will be the U.S. dollar amount calculated by reference to the exchange rate in effect on the date of receipt, regardless of whether the payment is in fact converted into U.S. dollars. If the dividend is converted into U.S. dollars on the date of receipt, a U.S. holder should not be required to recognize foreign currency gain or loss in respect of the dividend income. A U.S. holder may have foreign currency gain or loss if the dividend is converted into U.S. dollars after the date of receipt. Distributions in excess of current and accumulated earnings and profits, as determined for U.S. federal income tax purposes, will be treated as anon-taxable return of capital to the extent of your basis in the ADSs and thereafter as capital gain from the sale or exchange of such ADSs. Notwithstanding the foregoing, we do not intend to maintain calculations of our earnings and profits as determined for U.S. federal income tax purposes. Consequently, distributions generally will be reported as dividend income for U.S. information reporting purposes. The dividend will not be eligible for the dividends-received deduction generally allowed to U.S. corporations in respect of dividends received from other U.S. corporations.

Subject to the PFIC rules described above under “—PFIC Considerations,” dividends paid by anon-U.S. corporation generally will be taxed at the preferential tax rates applicable to long-term capital gain of

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non-corporate taxpayers if (a) suchnon-U.S. corporation is eligible for the benefits of certain U.S. treaties or the dividend is paid by suchnon-U.S. corporation with respect to stock that is readily tradable on an established securities market in the United States, (b) the U.S. holder receiving such dividend is an individual, estate, or trust, (c) such dividend is paid on shares that have been held by such U.S. holder for at least 61 days during the121-day period beginning 60 days before the“ex-dividend date,” and (d) we are not a PFIC in the year of the dividend or the immediately preceding year. If the requirements of the immediately preceding ~~paragraph~~sentence are not satisfied, a dividend paid by anon-U.S. corporation to a U.S. holder, including a U.S. holder that is an individual, estate, or trust, generally will be taxed at ordinary income tax rates (and not at the preferential tax rates applicable to long-term capital gains). As discussed above under “PFIC Considerations,” ~~while we may have been a PFIC for previous taxable years and~~ although the matter is not free from doubt (and while we can give no assurances as to our PFIC status for 2019 or future taxable years), we do not believe that we were a PFIC for U.S. federal income tax purposes for the ~~2017~~2018 taxable ~~year and we do not anticipate becoming a PFIC for the current or future taxable years.~~year. The dividend rules are complex, and each U.S. holder should consult its own tax advisor regarding the dividend rules.

The amount of dividend will include any amounts withheld by the Company in respect of French taxes. Subject to applicable limitations, some of which vary depending upon the U.S. holder’s circumstances and subject to the discussion above regarding concerns expressed by the U.S. Treasury, French income taxes withheld from dividends on ADSs at a rate not exceeding the rate provided by the Treaty will be creditable against the U.S. holder’s U.S. federal income tax liability.

Dividends received generally will be income fromnon-U.S. sources, which may be relevant in calculating your U.S. foreign tax credit limitation. Suchnon-U.S. source income generally will be “passive category income,” or in certain cases “general

category income”, which is treated separately from other types of income for purposes of computing the foreign tax credit allowable to you. The rules with respect to the foreign tax credit are complex and involve the application of rules that depend upon a U.S. holder’s particular circumstances. You should consult your own tax advisor to determine the foreign tax credit implications of owning the ADSs. Instead of claiming a credit, a U.S. holder may, subject to generally applicable limitations, elect to deduct such French taxes, if any, in computing taxable income. An election to deduct foreign taxes instead of claiming foreign tax credits applies to all foreign taxes paid or accrued in the taxable year.

Non-U.S. Holders. If you are anon-U.S. holder, dividends paid to you generally will not be subject to U.S. income tax unless the dividends are “effectively connected” with your conduct of a trade or business within the United States, and the dividends are attributable to a permanent establishment (or in the case of an individual, a fixed place of business) that you maintain in the United States if that is required by an applicable income tax treaty as a condition for subjecting you to U.S. taxation on a net income basis. In such cases you generally will be taxed in the same manner as a U.S. holder (other than with respect to the Medicare Tax described below). If you are a corporatenon-U.S. holder, “effectively connected” dividends may, under certain circumstances, be subject to an additional “branch profits tax” at a 30% rate or a lower rate if you are eligible for the benefits of an income tax treaty that provides for a lower rate.

Taxation of Sale, Exchange or other Disposition of ADSs

U.S. Holders. Subject to the PFIC rules described above under “—PFIC Considerations,” if you are a U.S. holder and you sell, exchange or otherwise dispose of your ADSs, you generally will recognize capital gain or loss for U.S. federal income tax purposes equal to the difference between the value of the amount realized and your tax basis in your ADSs. Gain or loss recognized on such a sale, exchange or other disposition of ADSs generally will be long-term capital gain if the U.S. holder has held the ADSs for more than one year. Long-term capital gains of U.S. holders who are individuals (as well as certain trusts and estates) are generally taxed at preferential rates. The gain or loss will generally be income or loss from sources within the United States for foreign tax credit limitation purposes, unless it is attributable to an office or other fixed place of business outside the United States and certain other conditions are met. Your ability to deduct capital losses is subject to limitations. As discussed above under “—PFIC Considerations,” ~~while we may have been a PFIC for previous taxable years and~~ although the matter is not free from doubt (and while we can give no assurances as to our PFIC status for 2019 or future taxable years), we do not believe that we were a PFIC for U.S. federal income tax purposes for the ~~2017~~2018 taxable ~~year and do not anticipate becoming a PFIC for the current or future taxable years.~~year.

Non-U.S. Holders. If you are anon-U.S. holder, you will not be subject to U.S. federal income tax on gain recognized on the sale, exchange or other disposition of your ADSs unless:

the gain is “effectively connected” with your conduct of a trade or business in the United States, and the gain is attributable to a permanent establishment (or in the case of an individual, a fixed place of business) that you maintain in the United States if that is required by an applicable income tax treaty as a condition for subjecting you to U.S. taxation on a net income basis; or

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~~that you maintain in the United States if that is required by an applicable income tax treaty as a condition for subjecting you to U.S. taxation on a net income basis; or~~

you are an individual, you are present in the United States for 183 or more days in the taxable year of such sale, exchange or other disposition and certain other conditions are met.

In the first case, thenon-U.S. holder will be taxed in the same manner as a U.S. holder (other than with respect to the Medicare Tax described below). In the second case, thenon-U.S. holder will be subject to U.S. federal income tax at a rate of 30% on the amount by which suchnon-U.S. holder’s ~~U.S.—source~~U.S.-source capital gains exceed such~~non-U.S.—source~~non-U.S.-source capital losses.

If you are a corporatenon-U.S. holder, “effectively connected” gains that you recognize may also, under certain circumstances, be subject to an additional “branch profits tax” at a 30% rate or at a lower rate if you are eligible for the benefits of an income tax treaty that provides for a lower rate.

Medicare Tax

Certain U.S. holders who are individuals, estates or trusts are required to pay a 3.8% Medicare surtax on all or part of that holder’s “net investment income” (or, in the case of an estate or trust, undistributed “net investment income”), which includes, among other items, dividends on, and capital gains from the sale or other taxable disposition of, the ADSs, subject to certain limitations and exceptions. U.S. holders should consult their own tax advisors regarding the effect, if any, of this surtax on their ownership and disposition of the ADSs.

Information with Respect to Foreign Financial Assets

U.S. holders that are individuals (and, to the extent provided in regulations, certain entities) that own “specified foreign financial assets,” including possibly the ADSs, with an aggregate value in excess of $50,000 are generally required to file IRS Form 8938 with information regarding such assets. Depending on the circumstances, higher threshold amounts may apply.

Specified foreign financial assets include any financial accounts maintained by foreign financial institutions, as well as any of the following, but only if they are not held in accounts maintained by financial institutions: (i) stocks and securities issued bynon-U.S. persons, (ii) financial instruments and contracts held for investment that havenon-U.S. issuers or counterparties and (iii) interests innon-U.S. entities. If a U.S. holder is subject to this information reporting regime, the failure to timely file IRS Form 8938 may subject the U.S. holder to penalties. In addition to these requirements, U.S. holders may be required to annually file FinCEN Report 114, ~~(Report~~Report of Foreign Bank and Financial ~~Accounts)~~Accounts with the U.S. Department of Treasury. U.S. holders are thus encouraged to consult their U.S. tax advisors with respect to these and other reporting requirements that may apply to their acquisition of the ADSs.

Backup Withholding and Information Reporting

In general, information reporting requirements will apply to distributions made on our ADSs within the United States to anon-corporate U.S. holder and to the proceeds from the sale, exchange, redemption or other disposition of ADSs by anon-corporate U.S. holder to or through a U.S. office of a broker. Payments made (and sales or other dispositions effected at an office) outside the U.S. will be subject to information reporting in limited circumstances.

In addition, backup withholding of U.S. federal income tax may apply to such amounts if the U.S. holder fails to provide an accurate taxpayer identification number (or otherwise establishes, in the manner provided by law, an exemption from backup withholding) or to report dividends required to be shown on the U.S. holder’s U.S. federal income tax returns.

Backup withholding is not an additional income tax, and the amount of any backup withholding from a payment to a U.S. holder will be allowed as credit against the U.S. holder’s U.S. federal income tax liability provided that the appropriate returns are timely filed.

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Anon-U.S. holder generally may eliminate the requirement for information reporting and backup withholding by providing a properly completed and duly executed certification of its foreign status to the payor, under penalties of perjury, on IRS FormW-8BEN,W-8BEN-E or other appropriate~~W-8BEN-E,~~W-8, as applicable. You should consult your own tax advisor as to the qualifications for exemption from backup withholding and the procedures for obtaining the exemption.

The foregoing does not purport to be a complete analysis of the potential tax considerations relating to the ownership and disposition of the ADSs. ~~Investors~~Prospective investors should consult their own tax advisors as to the particular tax considerations applicable to them relating to the ownership and disposition of the ADSs, including the applicability of the U.S. federal, state and local tax laws ornon-tax laws, foreign tax laws, and any changes in applicable tax ~~including the Tax Cuts and Jobs Act,~~ laws and any pending or proposed legislation or regulations.

Material French Income Tax Considerations

The following describes the material French income tax consequences to U.S. Holders (as defined below) of purchasing, owning and disposing of the ADSs and, unless otherwise noted, this discussion is the opinion of Jones Day, our French tax counsel, insofar as it relates to matters of French tax law and legal conclusions with respect to those matters.

This discussion does not purport to be a complete analysis or listing of all potential tax effects of the acquisition, ownership or disposition of our securities to any particular investor, and does not discuss tax considerations that arise from rules of general application or that are generally assumed to be known by investors. All of the following is subject to change. Such changes could apply retroactively and could affect the consequences described below.

In 2011, France introduced a comprehensive set of new tax rules applicable to French assets that are held by or in foreign trusts. These rules, among other things, provide for the inclusion of trust assets in the settlor’s net assets for purpose of applying the French wealth tax, for the application of French gift and death duties to French assets held in trust, for a specific tax on capital on the French assets of foreign trusts not already subject to the French wealth tax and for a number of French tax reporting and disclosure obligations. The following discussion does not address the French tax consequences applicable to securities (including ADSs) held in trusts. If securities are held in trust, the grantor, trustee and beneficiary are urged to consult their own tax advisor regarding the specific tax consequences of acquiring, owning and disposing of securities.

The description of the French income tax and wealth tax consequences set forth below is based on the Convention between the Government of the United States of America and the Government of the French Republic for the Avoidance of Double Taxation and the Prevention of Fiscal Evasion with Respect to Taxes on Income and Capital of August 31, 1994 which came into force on December 30, 1995 (as amended by any subsequent protocols, including the protocol of January 13, 2009), and the tax guidelines issued by the French tax authorities in force as of the date of this Annual Report, or the Treaty.

For the purposes of this discussion, the term “U.S. Holder” means a beneficial owner of securities that is (1) an individual who is a U.S. citizen or resident for U.S. federal income tax purposes, (2) a U.S. domestic corporation or certain other entities created or organized in or under the laws of the United States or any state thereof, including the District of Columbia, or (3) otherwise subject to U.S. federal income taxation on a net income basis in respect of securities.

If a partnership holds securities, the tax treatment of a partner generally will depend upon the status of the partner and the activities of the partnership. If a U.S. Holder is a partner in a partnership that holds securities, such holder is urged to consult its own tax advisor regarding the specific tax consequences of acquiring, owning and disposing of securities.

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This discussion applies only to investors that hold our securities as capital assets that have the U.S. dollar as their functional currency, that are entitled to Treaty benefits under the “Limitation on Benefits” provision contained in the Treaty, and whose ownership of the securities is not effectively connected to a permanent establishment or a fixed base in France. Certain U.S. Holders (including, but not limited to, U.S. expatriates, partnerships or other entities classified as partnerships for U.S. federal income tax purposes, banks, insurance companies, regulated investment companies,tax-exempt organizations, financial institutions, persons subject to the alternative minimum tax, persons who acquired the securities pursuant to the exercise of employee share options or otherwise as compensation, persons that own (directly, indirectly or by attribution) 5% or more of our voting stock or 5% or more of our outstanding share capital, dealers in securities or currencies, ~~brokers, mutual funds, individual retirement or other tax-deferred accounts,~~ persons that elect to mark their securities to market for U.S. federal income tax purposes and persons holding securities as a position in a synthetic security, straddle or conversion transaction) may be subject to special rules not discussed below.

U.S. Holders are urged to consult their own tax advisors regarding the tax consequences of the purchase, ownership and disposition of securities in light of their particular circumstances, especially with regard to the “Limitations on Benefits” provision.

Estate and Gift Taxes and Transfer Taxes

In general, a transfer of securities by gift or by reason of death of a U.S. Holder that would otherwise be subject to French gift or inheritance tax, respectively, will not be subject to such French tax by reason of the Convention between the Government of the United States of America and the Government of the French Republic for the Avoidance of Double Taxation and the Prevention of Fiscal Evasion with Respect to Taxes on Estates, Inheritances and Gifts, dated November 24, 1978, unless the donor or the transferor is domiciled in France at the time of making the gift or at the time of his or her death, or the securities were used in, or held for use in, the conduct of a business through a permanent establishment or a fixed base in France.

Financial Transactions Tax

Pursuant to Article 235 ter ZD of the French Tax Code (Code général des impôts), or the FTC, purchases of certain securities issued by a French company, including ordinary shares and ADSs, which are listed on a regulated market of the EU or an exchange market formally acknowledged by the AMF (in each case within the meaning of the French Monetary and Financial Code, or the FMFC) are subject in France to a 0.3% tax on financial transactions, or the TFT, provided inter alia that the issuer’s market capitalization exceeds ~~€1.0~~€1 billion as of December 1 of the year preceding the taxation year.

A list of relevant French companies whose market capitalization exceeds €1.0 billion as of December 1 of the year preceding the taxation year within the meaning of Article 235 ter ZD of the FTC ~~used to be published annually by the French Ministry of Economy. It~~ is ~~now~~ published by the French tax authorities, and could be amended at any time. Pursuant to Regulations~~BOI-ANNX-000467-20171221~~BOI-ANNX-000467-20181217 issued on December ~~21, 2017,~~17, 2018, Cellectis is currently not included in such list. Please note that such list may be updated from time to time, or may not be published anymore in the future.

As a result, neither the ADSs nor the ordinary shares are currently within the scope of the TFT.

Purchases of ~~Cellectis~~Cellectis’s securities may however become subject to the TFT if ~~Cellectis~~Cellectis’s market capitalization exceeds €1.0 ~~billion ($1.2 billion at December 31, 2017 closing rate).~~billion.

Registration Duties

In the case where the TFT is not applicable, (1) transfers of shares issued by a French company which are listed on a regulated or organized market within the meaning of the FMFC are subject to uncapped registration

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duties at the rate of 0.1% if the transfer is evidenced by a written statement (acte) executed either in France or outside France, whereas (2) transfers of shares issued by a French company which are not listed on a regulated or organized market within the meaning of the FMFC are subject to uncapped registration duties at the rate of 0.1% notwithstanding the existence of a written statement (acte).

As ordinary shares of Cellectis are listed on Euronext Growth market of Euronext in Paris, which is an organized market within the meaning of the FMFC, their transfer should be subject to uncapped registration duties at the rate of 0.1% subject to the existence of a written agreement (acte).

Although there is neither case law nor official guidelines published by the French tax authorities ~~are silent~~ on this point, transfers of ADSs should remain outside of the scope of the aforementioned 0.1% registration duties.

Wealth Tax

The French wealth tax (impôt de solidarité sur la fortune) has been repealed by the finance bill for 2018 (loi de finances pour 2018) dated December 30, 2017. It used to apply only to individuals and did not generally apply to securities held by a U.S. Holder who is a resident ~~as defined~~ pursuant to the provisions of the Treaty, provided that such U.S. Holder ~~did~~does not own directly or indirectly more than 25% of the issuer’s financial ~~rights and that the securities did not form part of the business property of a permanent establishment or fixed base in France. It~~rights.

As from January 1, 2018, it has been replaced by a new real estate wealth tax (impôt sur la fortune immobilière) ~~as from January 1, 2018. The scope of such new tax is narrowed~~which applies only to individuals owning French real estate assets ~~(and certain assets deemed to be real estate assets)~~ or rights, directly or indirectly through one or more legal entities and whose net taxable assets amount to at least ~~€1,300,000. Cellectis’ securities owned by a U.S. Holder should not fall within the scope of the new~~1,300,000 euros.

French real estate wealth tax ~~provided that~~may only apply to a U.S. individual to the extent such ~~U.S. Holder does not own~~individual holds, directly or indirectly, financial rights into a ~~shareholding exceeding~~company the assets of which comprise French real estate assets that are not allocated to its operational activity. Such financial rights may be taxable for the fraction of their value representing the French real estate assets that are not allocated to an operational activity. In any case, pursuant to Article 965, 2° of the FTC, shares of an operating company holding French real estate assets in which the relevant individual holds, directly and indirectly, less than 10% of the ~~financial rights and~~share capital or voting rights ~~of Cellectis.~~are exempt from real estate wealth tax.

Taxation of Dividends

Dividends paid by a French corporation tonon-residents of France are generally subject to French withholding tax at a rate of 30%. Such withholding tax may be reduced to 12.8% for dividends paid tonon-resident individuals. Dividends paid by a French corporation in anon-cooperative State or territory, as defined in Article238-0 A of the FTC, will generally be subject to French withholding tax at a rate of 75%. However, eligible U.S. Holders entitled to Treaty benefits under the “Limitation on Benefits” provision contained in the Treaty who are U.S. residents, as defined pursuant to the provisions of the Treaty, will not be subject to this 12.8%, 30% or 75% withholding tax rate, but may be subject to the withholding tax at a reduced rate (as described below).

Under the Treaty, the rate of French withholding tax on dividends paid to an eligible U.S. Holder who is a U.S. resident as defined pursuant to the provisions of the Treaty and whose ownership of ~~the~~ ordinary shares or the ADSs is not effectively connected with a permanent establishment or fixed base that such U.S. Holder has in France, is generally reduced to 15%, or to 5% if such U.S. Holder is a corporation and owns directly or indirectly at least 10% of the share capital of the issuer; such U.S. Holder may claim a refund from the French tax authorities of the amount withheld in excess of the Treaty rates of 15% or 5%, if any.

For U.S. Holders that are not individuals but are U.S. residents, as defined pursuant to the provisions of the Treaty, the requirements for eligibility for Treaty benefits, including the reduced 5% or 15% withholding tax rates contained in the “Limitation on Benefits” provision of the Treaty, are complicated, and certain technical changes were made to these requirements by the protocol of January 13, 2009. U.S. Holders are advised to consult their own tax ~~advisors~~advisers regarding their eligibility for Treaty benefits in light of their own particular circumstances.

In the event that dividends are paid by Cellectis, dividends paid to an eligible U.S. Holder may immediately be subject to the reduced rates of 5% or 15% provided that such holder establishes before the date of payment

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that it is a U.S. resident under the Treaty by completing and providing the depositary with a treaty form (Form 5000). Otherwise, dividends paid to a U.S. Holder that is a legal person or another legal entity and has not filed the Form 5000 before the dividend payment date will be subject to French withholding tax at the rate of 30%, or 75% for any U.S. ~~holder~~Holder if paid in anon-cooperative State or territory (as defined in Article238-0 A of the FTC) (unless the Company proves that neither the purpose nor the effect of paying the dividend in that ~~state~~State or territory are that of allowing, with the intent of tax evasion or avoidance, their location in such a ~~state~~State or territory), and then reduced at a later date to 5% or 15%, provided that such holder duly completes and provides the French tax authorities with the treaty forms Form 5000 and Form 5001 before December 31 of the second calendar year following the year during which the dividend is paid.

Certain qualifying pension funds and certain othertax-exempt entities are subject to the same general filing requirements as other U.S. Holders except that they may have to supply additional documentation evidencing their entitlement to these benefits.

Form 5000 and Form 5001, together with appropriate instructions, will be provided by the depositary to all U.S. Holders registered with the depositary. The depositary will arrange for the filing with the French tax authorities of all such forms properly completed and executed by U.S. Holders of ordinary shares or ADSs and returned to the depositary in sufficient time so that they may be filed with the French tax authorities before the distribution in order to obtain immediately a reduced withholding tax rate.

Tax on Sale or Other Disposition

As a matter of principle, under French tax law, a U.S. Holder should not be subject to any French tax on any capital gain from the sale, exchange, repurchase or redemption by us of ordinary shares or ADSs, provided that all of the following apply to such holder:

it is not a French tax resident for French tax purposes; and,

•

it has not held more than 25% of our dividend rights, known as “droits aux bénéfices sociaux” at any time during the preceding five years, either directly or indirectly, and, as relates to individuals, alone or with relatives; and,

it has not transferred ordinary shares or ADSs as part of redemption by Cellectis, in which case the proceeds may under certain circumstances be partially or fully characterized as dividends under French domestic law and, as result, be subject to French dividend withholding tax. As an exception, a U.S Holder, established, domiciled or incorporated in anon-cooperative State or territory as defined in Article238-0 A of the FTC should be subject to a 75% withholding tax in France on any such capital gain, regardless of the fraction of the dividend rights it holds.

In ~~general, under~~case an applicable double tax treaty between France and the ~~Treaty,~~U.S. Holder country of residence contains more favorable provisions, a U.S. Holder may not be subject to any French income tax or capital gains tax in case of sale or disposal of any ordinary shares or ADSs of Cellectis even if one or more of the above mentioned statements are not applicable.

Particularly, a U.S. Holder who is a U.S. tax resident for purposes of the Treaty and is entitled to Treaty benefit will not be subject to French tax on any such capital gain, ~~from the redemption (other than redemption proceeds characterized as dividends under French domestic tax law or administrative guidelines), sale or exchange of ordinary shares or ADSs~~ unless the ordinary shares or the ADSs form part of the business property of a permanent establishment or fixed base that the U.S. Holder has in France.

U.S. Holders who own ordinary shares or ADSs through U.S. partnerships that are not residents for Treaty purposes are advised to consult their own tax advisors regarding their French tax treatment and their eligibility for Treaty benefits in light of their own particular circumstances.

A U.S. Holder that is not a U.S. resident for Treaty purposes or is not entitled to Treaty benefit (and in both cases is not resident, established or incorporated in anon-cooperative State or territory as defined in Article238-0 A of the FTC) and has held more than 25% of our dividend rights, known as “droits aux bénéfices sociaux” at any time during the preceding five years, either directly or indirectly, and, as relates to individuals, alone or with relatives will be subject to a levy in France at the rate of 31% (anticipated to be progressively decreased to 25% in the coming years), if such U.S. Holder is a legal person, or 12.8%, if such U.S. Holder is an individual.

Special rules apply to U.S. Holders who are residents of more than one country.

F.	Dividends and Paying Agents

Not applicable.

G.	Statement by Experts

Not applicable.

H.	Documents on Display

We are subject to the information reporting requirements of the Exchange Act applicable to foreign private issuers and under those requirements file reports with the SEC. Those reports may be inspected without charge at the locations described below. As a foreign private issuer, we are exempt from the rules under the Exchange Act related to the furnishing and content of proxy statements, and our officers, directors and principal shareholders are exempt from the reporting and short-swing profit recovery provisions contained in Section 16 of the Exchange Act. In addition, we are not required under the Exchange Act to file periodic reports and financial statements with the SEC as frequently or as promptly as United States companies whose securities are

registered under the Exchange Act. Nevertheless, we file with the U.S. Securities and Exchange Commission an Annual Report containing financial statements that have been examined and reported on, with and opinion expressed by an independent registered public accounting firm, and we submit quarterly interim consolidated financial data to the SEC under cover of the SEC’s Form6-K.

We maintain a corporate website at www.cellectis.com. We intend to post our Annual Report on our website promptly following it being filed with the SEC. Information contained on, or that can be accessed through, our website does not constitute a part of this Annual Report. We have included our website address in this Annual Report solely as an inactive textual reference.

~~188~~

You may also review a copy of this Annual Report, including exhibits and any schedule filed herewith, and obtain copies of such materials at prescribed rates, at the Securities and Exchange Commission’s Public Reference Room in Room 1580, 100 F Street, NE, Washington, D.C. 20549-0102. You may obtain information on the operation of the Public Reference Room by calling the Securities and Exchange Commission at1-800-SEC-0330. The Securities and Exchange Commission maintains a website (http://www.sec.gov) that contains reports, proxy and information statements and other information regarding registrants, such as Cellectis, that file electronically with the Securities and Exchange Commission.

With respect to references made in this Annual Report to any contract or other document of Cellectis, such references are not necessarily complete and you should refer to the exhibits attached or incorporated by reference to this Annual Report for copies of the actual contract or document.

I.	Subsidiary Information

Not ~~applicable~~applicable.

ITEM 11.

QUANTITATIVE AND QUALITATIVE DISCLOSURES ABOUT MARKET RISK

Foreign Currency Exchange Risk

We derive a significant portion of our revenues, including payments under our collaboration agreement with Pfizer, in U.S. dollars. Since the beginning of fiscal year 2015, we have been significantly expanding our activities in the United States, but there continues to be a currency mismatch in our cash flows since most of our expenses remain denominated primarily in Euros. If the average value of the U.S. Dollar had been 10% higher relative to the euro during ~~2017,~~2018, our collaboration revenues would have increased by ~~€2.0~~€0.2 million. Our exposure to currencies other than the U.S. dollar is negligible.

Our financial condition and results of operations are measured and recorded in the relevant local base currency and then translated into Euros for inclusion in our Consolidated Financial Statements. We translate balance sheet amounts at the exchange rates in effect on the date of the balance sheet, while income and cash flow items are translated at the average rate of exchange in effect for the relevant period. Our exposure to currencies other than the U.S. dollar is negligible.

For the year ended December 31, ~~2017,~~2018, our revenues denominated in U.S. dollars related to the ~~Pfizer~~Allogene collaboration agreement and revenues from our Plants segment. Our cash and cash equivalents and marketable securities denominated in ~~U.S~~U.S. dollars amounted to ~~$194.2~~$298.1 million as of December 31, ~~2017.~~2018. Current financial assets denominated in U.S. dollars amounted to ~~$39.7~~$0.4 million as of December 31, ~~2017.~~2018.

~~During the year ended December 31, 2017, we subscribed to zero premium accumulators ($5.0 million nominal value).~~ The net foreign exchange result for the fiscal year ~~2016~~2018 is a gain of ~~€0.6~~$10.5 million. We cannot rule out the possibility that a significant increase in our business, particularly in the United States, may result in greater exposure to exchange rate risk. We would then consider adopting an appropriate policy for hedging against these risks.

Interest Rate Risk

We seek to engage in prudent management of our cash and cash equivalents, mainly cash on hand and common financial instruments (typically short- andmid-term deposits). Furthermore, the interest rate risk related to cash, cash equivalents and common financial instruments is not significant based on the quality of the financial institutions with which we work.

Inflation Risk

We do not believe that inflation has had a material effect on our business, financial condition or results of operations. If our costs were to become subject to significant inflationary pressures, we may not be able to fully

~~189~~

offset such higher costs through price increases. Our inability or failure to do so could harm our business, financial condition and results of operations.

ITEM 12.

DESCRIPTION OF SECURITIES OTHER THAN EQUITY SECURITIES

A.	Debt Securities

Not applicable.

B.	Warrants and Rights

Not applicable.

C.	Other Securities

Not applicable.

D.	American Depositary Shares

Citibank, N.A., as depositary for our ADSs, registers and delivers ADSs. Each ADS represents one ordinary share (or a right to receiveone-half of one ordinary share) deposited with Citibank International Limited, located at EGSP 186, 1 North Wall Quay, Dublin 1 Ireland or any successor, as custodian for the depositary. Each ADS will also represent any other securities, cash or other property which may be held by the depositary in respect of the depositary facility. The depositary’s corporate trust office at which the ADSs will be administered is located at 388 Greenwich Street, New York, New York 10013.

A deposit agreement among us, the depositary and the ADS holders sets out the ADS holder rights as well as the rights and obligations of the depositary. New York law governs the deposit agreement and the ADSs. A copy of the Agreement is incorporated by reference as an exhibit to this Annual Report.

Fees and Charges

As an ADS holder, you will be required to pay the following fees under the terms of the depositary agreement:


Service		Fees
• Issuance of ADSs upon deposit of shares (excluding issuance as a result of distributions of shares)		Up to U.S. 5¢ per ADS issued

• Cancellation of ADSs		Up to U.S. 5¢ per ADS canceled

• Distribution of cash dividends or other cash distributions (i.e., sale of rights and other entitlements)		Up to U.S. 5¢ per ADS held


Service		Fees

• Distribution of ADSs pursuant to (1) stock dividends or other free stock distributions, or (2) exercise of rights to purchase additional ADSs		Up to U.S. 5¢ per ADS held

• Distribution of securities other than ADSs or rights to purchase additional ADSs (i.e.,spin-off shares)		Up to U.S. 5¢ per ADS held

• ADS Services		Up to U.S. 5¢ per ADS held on the applicable record date(s) established by the depositary

~~190~~

As an ADS holder you will also be responsible to pay certain fees and expenses incurred by the depositary and certain taxes and governmental charges such as:

taxes (including applicable interest and penalties) and other governmental charges;

the registration fees as may from time to time be in effect for the registration of ordinary shares on the share register and applicable to transfers of ordinary shares to or from the name of the custodian, the depositary or any nominees upon the making of deposits and withdrawals, respectively;

certain cable, telex and facsimile transmission and delivery expenses;

the expenses and charges incurred by the depositary in the conversion of foreign currency;

the fees and expenses incurred by the depositary in connection with the compliance with exchange control regulations and other regulatory requirements applicable to ordinary shares, ADSs and ADRs; and

the fees and expenses incurred by the depositary, the custodian, or any nominee in connection with the servicing or delivery of deposited property.

ADS fees and charges payable upon (1) deposit of ordinary shares against issuance of ADSs and (2) surrender of ADSs for cancellation and withdrawal of ordinary shares are charged to the person to whom the ADSs are delivered (in the case of ADS issuances) and to the person who delivers the ADS, for cancellation (in the case of ADS cancellations). In the case of ADSs issued by the depositary into DTC or presented to the depositary via DTC, the ADS issuance and cancellation fees and charges may be deducted from distributions made through DTC, and may be charged to the DTC participant(s) receiving the ADSs or the DTC participant(s) surrendering the ADSs for cancellation, as the case may be, on behalf of the beneficial owner(s) and will be charged by the DTC participant(s) to the account(s) of the applicable beneficial owner(s) in accordance with the procedures and practices of the DTC participant(s) as in effect at the time. ADS fees and charges in respect of distributions and the ADS service fee are charged to the holders as of the applicable ADS record date. In the case of distributions of cash, the amount of the applicable ADS fees and charges is deducted from the funds being distributed. In the case of (1) distributions other than cash and (2) the ADS service fee, holders as of the ADS record date will be invoiced for the amount of the ADS fees and charges and such ADS fees and charges may be deducted from distributions made to holders of ADSs. For ADSs held through DTC, the ADS fees and charges for distributions other than cash and the ADS service fee may be deducted from distributions made through DTC, and may be charged to the DTC participants in accordance with the procedures and practices prescribed by DTC and the DTC participants in turn charge the amount of such ADS fees and charges to the beneficial owners for whom they hold ADSs.

In the event of refusal to pay the depositary fees, the depositary may, under the terms of the deposit agreement, refuse the requested service until payment is received or may set off the amount of the depositary fees from any distribution to be made to the ADS holder. Certain ADS fees and charges (such as the ADS service fee) may become payable shortly after the closing of the ADS offering.

Note that the fees and charges you may be required to pay may vary over time and may be changed by us and by the depositary. You will receive prior notice of such changes. The depositary may reimburse us for certain expenses incurred by us in respect of the ADR program, by making available a portion of the ADS fees charged in respect of the ADR program or otherwise, upon such terms and conditions as we and the depositary agree from time to time.

Depositary Payments for ~~2017~~2018

From time to time, the Depositary may make payments to us to reimburse and/or share revenue from the fees collected from ADS holders, or waive fees and expenses for services provided, generally relating to costs and expenses arising out of establishment and maintenance of the ADS program. In performing its duties under the deposit agreement, the Depositary may use brokers, dealers or other service providers that are affiliates of the Depositary and that may earn or share fees or commissions.

~~191~~

For the year ended December 31, ~~2017,~~2018, Citibank, N.A., as Depositary, had made reimbursements to the Company of ~~$0.1 million.~~$151 thousand.

PART II

ITEM 13.

DEFAULTS, DIVIDEND ARREARAGES AND DELINQUENCIES.

Not applicable.

ITEM 14.

MATERIAL MODIFICATIONS TO THE RIGHTS OF SECURITY HOLDERS AND USE OF PROCEEDS.

Initial Public Offering

In March 2015, we sold 5,500,000 ADSs, each representing one ordinary share, nominal value €0.05, in our initial public offering at a price of $41.50 per ADS, for aggregate gross proceeds of approximately $228.3 million. We incurred aggregate underwriting discounts of approximately $16.0 million and expenses of approximately $2.7 million, resulting in net proceeds to us of approximately $209.6 million. No payments were made directly or indirectly to any director, officer, general partner of ours or to their associates, persons owning ten percent or more of any class of our equity securities, or to any of our affiliates. The offering commenced on March 24, 2015 and did not terminate before all of the securities registered in the registration statement were sold. The effective date of the registration statement, FileNo. 333-202205, for our initial public offering was March 24, 2015. Merrill Lynch, Pierce, Fenner & Smith Incorporated, Jefferies LLC, Piper Jaffray & Co., Oppenheimer & Co. Inc. and Trout Capital LLC acted as underwriters of the initial public offering.

A portion of the net proceeds from our initial public offering was used for general corporate purposes in connection with the development of our current proprietary immuno-oncology product candidates, for further research and development regarding cell attributes and to develop our manufacturing processes and cell engineering technologies, to pursue new human therapeutics outside of oncology and to advance our agricultural biotechnology business. The balance is held in cash and cash equivalents and current financial assets and is intended to also be used for general corporate purposes. None of the net proceeds of our initial public offering were paid directly or indirectly to any director, officer, general partner of ours or to their associates, persons owning ten percent or more of any class of our equity securities, or to any of our affiliates.

Follow-on Offering

On April 10, 2018, Cellectis closed afollow-on offering of 5,646,000 ADS at a public offering price of $31.00 per ADS resulting in gross proceeds of $175 million. On May 11, 2018, in connection with the exercise by the underwriters of thefollow-on offering of their option to purchase additional shares, Cellectis closed the sale of an additional 500,000 ADS at the public offering price of $31.00 per ADS resulting in additional gross proceeds of $15.5 million.

ITEM 15.

CONTROLS AND PROCEDURES.

(a)

Our management, with the participation of our Chief Executive Officer and Chief Financial Officer, after evaluating the effectiveness of our disclosure controls and procedures (as defined in Exchange Act Rule13a-15(e)) as of the end of the period covered by this Form20-F, have concluded that our disclosure controls and procedures were effective as of December 31, ~~2017.~~2018.

(b)

Report of Management on Internal Control Over Financial Reporting

Management of the Company is responsible for establishing and maintaining adequate internal control over financial reporting, as such term is defined in Exchange Act Rule13a-15(f). Management, with the participation of the Chief Executive Officer and the Chief Financial Officer, has assessed the effectiveness of internal control over financial reporting as of December 31, ~~2017.~~2018. Management’s assessment was based on the framework in “Internal Control – Integrated Framework” (2013 framework) issued by the Committee of Sponsoring Organizations of the Treadway Commission (COSO).

Based on that assessment, management concluded that, as of December 31, ~~2017,~~2018, the Company’s internal control over financial reporting was effective to provide reasonable assurance regarding the reliability of its financial reporting and the preparation of its financial statements for external purposes, in accordance with generally accepted accounting principles.

~~192~~

Due to its inherent limitations, internal control over financial reporting may not prevent or detect misstatements, and can only provide reasonable assurance regarding the reliability of financial reporting and the preparation of financial statements. Also, projections of any evaluation of effectiveness to future periods are subject to the risk that controls may become inadequate because of changes in conditions, or that the degree of compliance with the policies or procedures may deteriorate.

The effectiveness of the Company’s internal control over financial reporting has been audited by Ernst & Young et Autres, independent registered public accounting firm, as stated in their report on the Company’s internal control over financial reporting as of December 31, ~~2017,~~2018, which is included herein. See paragraph (c) of the present Item 15, below.

(c)

See report of Ernst & Young et Autres, independent registered public accounting firm, included under “Item 18. Financial Statements” on pageF-3.

~~(d)~~

~~We have not made any significant change in internal controls over financial reporting during the year ended December 31, 2017.~~

ITEM 15T.

CONTROLS AND PROCEDURES.

Not applicable.

ITEM 16.

RESERVED

Not applicable.

ITEM 16A.

AUDIT COMMITTEE FINANCIAL EXPERT

Our board of directors has determined that Mr. Pierre Bastid, Mr. Laurent Arthaud, and Mr. ~~Jean-Marie Messier~~Hervé Hoppenot are audit and finance committee financial experts as defined by the Securities and Exchange Commission rules and have the requisite financial sophistication under the applicable rules and regulations of the Nasdaq Stock Market. Mr. Pierre Bastid, Mr. Laurent Arthaud, and Mr. ~~Jean-Marie Messier~~Hervé Hoppenot are independent as such term is defined in Rule10A-3 under the Exchange Act and under the listing standards of the Nasdaq Stock Market.

ITEM 16B.

CODE OF ETHICS

We have adopted a Code of Business Conduct and Ethics, or the Code of Conduct that is applicable to all of our employees, executive officers and directors. Following the completion of our initial public offering, the Code of Conduct became available on our website atwww.cellectis.com. Our board of directors is responsible for overseeing the Code of Conduct and is required to approve any waivers of the Code of Conduct for employees, executive officers and directors. We expect that any amendments to the Code of Conduct, or any waivers of its requirements, will be disclosed on our website.

~~193~~

ITEM 16C.

PRINCIPAL ACCOUNTANT FEES AND SERVICES

Ernst & Young et Autres, or Ernst & Young, has served as our independent registered public accounting firm for ~~2016~~2017 and ~~2017.~~2018. Our accountants billed the following fees to us for professional services in each of those fiscal years:


	Year Ended December 31,					Year Ended December 31,
	2016		2017			2017			2018
	($, in thousands)					($, in thousands)
Audit Fees		779		1.751	*		1,751	*		847	**
Audit-Related Fees		42		—			—			—
Tax Fees		—		—			—			—
Other Fees		—		—			—			—
Total		821		1.751			1,751			847

~~(*) $539~~

(*)	$539 thousand for Cellectis and $1.212 thousand for Calyxt (of which $898 thousand related to the Nasdaq IPO)

(**)	$513 thousand for Cellectis and $334 thousand for Calyxt

“Audit Fees” are the aggregate fees billed for the audit of our annual financial statements. This category also includes services that generally the independent accountant provides, such as consents and assistance with and review of documents filed with the SEC.

“Audit-Related Fees” are the aggregate fees billed for assurance and related services that are reasonably related to the performance of the audit and are not reported under Audit Fees.

“Tax Fees” are the aggregate fees billed for professional services rendered by the principal accountant for tax compliance, tax advice and tax planning related services.

“Other Fees” relate to services provided with respect to our registration statement for our initial public offering.

There were no “Audit Related Fees,” “Tax Fees” either billed or paid during ~~2016~~2017 or ~~2017.~~2018.

Audit andNon-Audit ServicesPre-Approval Policy

The audit and finance committee has responsibility for appointing, setting compensation of and overseeing the work of the independent registered public accounting firm. In recognition of this responsibility, the audit and finance committee has adopted a policy governing thepre-approval of all audit and permittednon-audit services performed by our independent registered public accounting firm to ensure that the provision of such services does not impair the independent registered public accounting firm’s independence from us and our management. Unless a type of service to be provided by our independent registered public accounting firm has received generalpre-approval from the audit and finance committee, it requires specificpre-approval by the audit and finance committee. The payment for any proposed services in excess ofpre-approved cost levels requires specificpre-approval by the audit and finance committee. All audit andnon-audit services rendered by our independent registered public accounting firm in ~~2016~~2018 werepre-approved by the audit and finance committee.

Pursuant to itspre-approval policy, the audit and finance committee may delegate its authority topre-approve services to the chairperson of the audit and finance committee. The decisions of the chairperson to grantpre-approvals must be presented to the full audit and finance committee at its next scheduled meeting. The audit and finance committee may not delegate its responsibilities topre-approve services to the management.

The audit and finance committee has considered thenon-audit services provided by Ernst & Young as described above and believes that they are compatible with maintaining Ernst & Young’s independence as our independent registered public accounting firm.

~~194~~

ITEM 16D.

EXEMPTIONS FROM THE LISTING STANDARDS FOR AUDIT COMMITTEES

Not applicable.

ITEM 16E.

PURCHASES OF EQUITY SECURITIES BY THE ISSUER AND AFFILIATED PURCHASERS

Not applicable.

ITEM 16F.

CHANGE IN REGISTRANT’S CERTIFYING ACCOUNTANT

Not applicable.

ITEM 16G.

CORPORATE GOVERNANCE

In addition, Nasdaq rules require that a listed company specify that the quorum for any meeting of the holders of share capital be at least 33¹/₃% of the outstanding shares of the company’s common voting stock. We intend to follow our French home country practice, rather than complying with this Nasdaq rule. Consistent with French Law, ourBy-laws provide that when first convened, general meetings of shareholders may validly convene only if the shareholders present or represented hold at least (1) 20% of the voting shares in the case of an ordinary general meeting or of an extraordinary general meeting where shareholders are voting on a capital increase by capitalization of reserves, profits or share premium, or (2) 25% of the voting shares in the case of any other extraordinary general meeting. If such quorum required by French law is not met, the meeting is adjourned. There is no quorum requirement under French law when an ordinary general meeting or an extraordinary general meeting where shareholders are voting on a capital increase by capitalization of reserves, profits or share premium is reconvened, but the reconvened meeting may consider only questions that were on the agenda of the adjourned meeting. When any other extraordinary general meeting is reconvened, the required quorum under French law is 20% of the shares entitled to vote. If a quorum is not met at a reconvened meeting requiring a quorum, then the meeting may be adjourned for a maximum of two months. See the section of the prospectus filed with the Commission on March 26, 2015 titled “Description of Share Capital—Key Provisions of OurBy-laws and French Law Affecting Our Ordinary Shares.”

~~195~~

Further, Nasdaq rules require that listed companies have a nominations committee comprised solely of independent directors. We intend to follow our French home country practice, as described under “—Board Composition,” rather than complying with this Nasdaq rule.

Board Committees

Audit and Finance Committee. Our audit and finance committee reviews our internal accounting procedures, consults with and reviews the services provided by our independent registered public accountants and assists our board of directors in its oversight of our corporate accounting and financial reporting. Currently, our audit and finance committee is comprised of three members of the board of directors: Messrs. Bastid, Arthaud and ~~Messier.~~Hoppenot.

The duties specifically assigned to the audit and finance committee by our board of directors include, but are not limited to:

with regard to our financial statements:

review on a preliminary basis and express its opinion on the draft annual and quarterly financial statements prior to the board of directors officially receiving the financial statements;

monitor the scope of consolidation and review, where necessary, any explanations in connection thereto;

with regard to internal controls:

assess the efficiency and quality of internal control systems and procedures within the consolidated Company;

entrust the internal audit department with any mission which the committee deems necessary;

~~196~~

with regard to external controls:

examine any question relating to the appointment, renewal or dismissal of our statutory auditors and their fees regarding the performance of their control review functions;

at least annually, review and discuss the information provided by management and the auditors relating to the independence of the audit firm;

pre-approve any services entrusted to the statutory auditors which is outside of the scope of the annual audit;

discuss if necessary any points of disagreement between the statutory auditors and the officers of the Company that may arise within the scope of these operations; and

review and discuss with the statutory auditors the plans for, and the scope of, the annual audit and other examinations; and

with regard to risks:

review on a regular basis the financial situation, the cash position and the material risks and undertakings of the Company and its subsidiaries; and

review the risk management policy and the process implemented to evaluate and manage these risks.

Compensation Committee. Our compensation committee assists our board of directors in reviewing the compensation of our executive officers and directors and makes recommendations in respect thereof. Currently, our compensation committee is comprised of two members of the board of directors: Mr. Godard and Dr. Schwebig. The principal duties and responsibilities of our compensation committee include, but are not limited to:

review equity incentive plans(non-employee warrants, stock options, restricted (free) shares, etc.) and make recommendations to our board of directors in connection therewith;

evaluate and make recommendations on the compensation policies and programs of executive officers and on the compensation of directors;

~~197~~

review any proposed employment agreement with, and any proposed severance or retention plans or agreements applicable to, any of our executive officers;

review, at least annually, corporate goals and objectives relevant to the compensation of our executive officers; and

ITEM 16H.

MINE SAFETY DISCLOSURE

Not applicable.

PART III

ITEM 17.

FINANCIAL STATEMENTS

See pagesF-1 through ~~F-54~~F-58 of this Annual Report.

ITEM 18.

FINANCIAL STATEMENTS

Not applicable.

ITEM 19.

EXHIBITS

The following exhibits are filed as part of this Annual Report:


Exhibit Number	Description of Exhibit	Schedule/ Form	File Number	Exhibit	File Date
~~1.1#~~
1.1	By-laws (status) of the registrant (English translation)	~~6-K~~	~~001-36891~~	~~99.1~~	~~January 9, 2018~~Filed herewith

2.1#	Form of Deposit Agreement	F-1	333-202205	4.1	March 10, 2015

2.2#	Form of American Depositary Receipt (included in Exhibit 2.1)	F-1	333-202205	Included in 4.1	March 10, 2015

~~4.1~~ 4.1#	~~Reserved~~Patent License Agreement#C-00061901 between L’Institut Pasteur and Cellectis S.A., dated June 19, 2000 (English translation)				Filed herewith

4.2#* 4.1.1	Amendment No. 1 to Patent License Agreement#C-00061901 between L’Institut Pasteur and Cellectis S.A., dated December 20, 2002 (English translation)				Filed herewith

4.1.2	Amendment No. 2 to Patent License Agreement#C-00061901 between L’Institut Pasteur and Cellectis S.A., dated September 8, 2003 (English translation)				Filed herewith

4.1.3#	Amendment No. 3 to Patent License Agreement#C-00061901 between L’Institut Pasteur and Cellectis S.A., dated February 26, 2008	F-1	333-202205	10.1.3	March 12, 2015

4.1.4#	Amendment No. 4 to Patent License Agreement#C-00061901 between L’Institut Pasteur and Cellectis S.A., dated April 11, 2013 (English translation)	F-1	333-202205	10.1.4	March 12, 2015

4.2	Patent License Agreement#C-00061906 between L’Institut Pasteur and Cellectis S.A., dated October 19, 2000 (English translation)	~~F-1~~	~~333-202205~~	~~10.2~~	~~March 12, 2015~~Filed herewith

4.2.1#* 4.2.1	Amendment No. 1 to Patent License Agreement#C-00061906 between L’Institut Pasteur and Cellectis S.A., dated September 8, 2003 (English translation)	~~F-1~~	~~333-202205~~	~~10.2.1~~	~~March 12, 2015~~Filed herewith

4.2.2#* 4.2.2	Amendment No. 2 to Patent License Agreement#C-00061906 between L’Institut Pasteur and Cellectis S.A., dated June 24, 2004 (English translation)	~~F-1~~	~~333-202205~~	~~10.2.2~~	~~March 12, 2015~~Filed herewith

4.2.3	Amendment No. 3 to Patent License Agreement#C-00061906 between L’Institut Pasteur and Cellectis S.A., dated August 24, 2005 (English translation)				Filed herewith

4.2.4	Amendment No. 4 to Patent License Agreement#C-00061906 between L’Institut Pasteur and Cellectis S.A., dated December 27, 2007 (English translation)				Filed herewith

~~198~~


Exhibit Number	Description of Exhibit	Schedule/ Form	File Number	Exhibit	File Date

4.2.3#*	~~Amendment No. 3 to Patent License Agreement#C-00061906~~ ~~between L’Institut Pasteur and Cellectis S.A., dated August 24, 2005 (English translation)~~	~~F-1~~	~~333-202205~~	~~10.2.3~~	~~March 12, 2015~~

4.2.4#*	~~Amendment No. 4 to Patent License Agreement#C-00061906~~ ~~between L’Institut Pasteur and Cellectis S.A., dated December 27, 2007 (English translation)~~	~~F-1~~	~~333-202205~~	~~10.2.4~~	~~March 12, 2015~~

4.3#* 4.3	Patent License Agreement#C-00061905 between L’Institut Pasteur and Cellectis S.A., dated June 19, 2000 (English translation)	~~F-1~~	~~333-202205~~	~~10.3~~	~~March 12, 2015~~Filed herewith

4.3.1#* 4.3.1	Amendment No. 1 to Patent License Agreement#C-00061905 between L’Institut Pasteur and Cellectis S.A., dated September 8, 2003 (English translation)	~~F-1~~	~~333-202205~~	~~10.3.1~~	~~March 12, 2015~~Filed herewith

4.4#* 4.4	~~Research and Collaboration Agreement between Pfizer Inc. and Cellectis S.A., dated June 17, 2014~~	~~F-1~~	~~333-202205~~	~~10.4~~	~~March 12, 2015~~

4.4.1*	~~Amendment to the Research Collaboration and License Agreement, by and between the Company and Pfizer Inc., dated as of December 1, 2016~~[Reserved]				~~Filed Herewith~~

4.5#* 4.4.1	Research, Product Development, Option, License and Commercialization Agreement, among Les Laboratoires Servier SAS, Institut de Recherches Internationales Servier SAS and Cellectis S.A., dated February 17, 2014	~~F-1~~	~~333-202205~~	~~10.5~~	~~March 12, 2015~~
4.5.1#*	Amendment to the Product Development, Option, License and Commercialization Agreement, among Les Laboratoires Servier SAS, Institut de Recherches Internationales Servier SAS and Cellectis S.A., dated November 18, 2015	~~20-F~~	~~001-36891~~	~~4.5.1~~	~~March 21, 2016~~

4.5.2*	~~Amendment No. 2 to the Product Development, Option, License and Commercialization Agreement, by and between the Company and Les Laboratoires Serviers, dated as of November 28, 2016~~[Reserved]				~~Filed Herewith~~

4.5	[Reserved]

4.5.1	[Reserved]

4.6#*	Exclusive Patent License Agreement between Regents of the University of Minnesota and Cellectis S.A., dated January 10, 2011	F-1	333-202205	10.6	March 12, 2015

4.6.1#*	First Amendment to the Exclusive Patent License Agreement between Regents of the University of Minnesota and Cellectis S.A., dated May 24, 2012	F-1	333-202205	10.6.1	March 12, 2015

~~199~~


~~Exhibit~~ ~~Number~~	~~Description of Exhibit~~	~~Schedule/~~ ~~Form~~	~~File Number~~	~~Exhibit~~	~~File Date~~

4.6.2#*	Second Amendment to the Exclusive Patent License Agreement between Regents of the University of Minnesota and Cellectis S.A., dated April 1, 2014	F-1	333-202205	10.6.2	March 12, 2015

4.6.3*	Third Amendment to the Exclusive Patent License Agreement between Regents of the University of Minnesota and Cellectis S.A., dated December 16, 2015	20-F	001-36891	4.6.3	~~Filed Herewith~~March 13, 2018

4.7#* 4.7	Patent & Technology License Agreement between Ohio State Innovation Foundation and Cellectis S.A., dated October 23, 2014				Filed herewith

4.8#	Warrants Issue Agreement between Cellectis S.A. and Kepler Capital Markets SA, dated December 20, 2012 (English translation)	F-1	333-202205	~~10.7~~10.8	March ~~12,~~10, 2015

~~4.8†~~ 4.8.1#	First Amendment to Warrants Issue Agreement between Cellectis S.A. and Kepler Capital Markets SA, dated June 6, 2013 (English translation)	F-1	333-202205	10.8.1	March 10, 2015

4.8.2#	Second Amendment to Warrants Issue Agreement between Cellectis S.A. and Kepler Capital Markets SA, dated October 7, 2013 (English translation)	F-1	333-202205	10.8.2	March 10, 2015

4.9#	Warrant Agreement between Cellectis S.A. and Trout Capital LLC, dated March 24, 2014	F-1	333-202205	10.9	March 10, 2015

4.10†#	Change of Control Plan, effective as of September 4, 2014 (English translation)	F-1	333-202205	10.10	March 10, 2015
~~4.9†~~
4.11†#	Summary of BSA Plan	F-1	333-202205	10.11	March 10, 2015

~~4.10†~~ 4.12†#	Summary of BSPCE Plan	F-1	333-202205	10.12	March 10, 2015

~~4.11†~~ 4.13†#	2012 Free Share Plan	F-1	333-202205	10.13	March 10, 2015

~~4.12†~~ 4.14†#	2013 Free Share Plan	F-1	333-202205	10.14	March 10, 2015

~~4.13†~~ 4.15†#	2014 Free Share Plan	F-1	333-202205	10.15	March 10, 2015

~~4.14†~~ 4.16†#	2015 Free Share Plan	20-F	001-36891	4.16	March ~~21, 2016~~10, 2015


Exhibit Number	Description of Exhibit	Schedule/ Form	File Number	Exhibit	File Date

~~4.15†~~ 4.17†#	2015 Stock Option Plan	20-F	001-36891	4.17	March ~~21, 2016~~10, 2015

~~4.16†~~ 4.18†#	2016 Stock Option Plan	S-8	333-214884	99.1	December 2, 2016

~~4.17†~~ 4.19†#	2017 Stock Option Plan	S-8	333-222482	99.1	January 9, 2018

~~4.18†~~ 4.20†#	Summary of BSA Plan	S-8	333-222482	99.2	January 9, 2018

4.21†#	Free Share 2018 Plan	S-8 POS	333-222482	99.3	April 13, 2018

4.22†#	2018 Stock Option Plan	S-8	333-227717	99.1	October 5, 2018

4.23†#	Summary of BSA Plan	S-8	333-227717	99.2	October 5, 2018

4.24†#	Second Free Share 2018 Plan	S-8	333-227717	99.3	October 5, 2018

4.25*	License Agreement between Allogene Therapeutics, Inc. and Cellectis S.A. dated March 7, 2019				Filed herewith

4.26*	License, Development and Commercialization Agreement between Les Laboratoires Servier and Cellectis S.A. dated March 6, 2019				Filed herewith

4.27	Management Services Agreement between Cellectis S.A., Cellectis, Inc. and Calyxt, Inc. dated as of January 1, 2016				Filed herewith

4.28	Management Services Agreement Amendment dated July 25, 2017 between Cellectis S.A. and Calyxt, Inc.				Filed herewith

4.29	Separation Agreement dated July 25, 2017 between Cellectis S.A. and Calyxt, Inc.				Filed herewith

4.30	Stockholders Agreement dated July 25, 2017 between Cellectis S.A. and Calyxt, Inc.				Filed herewith

4.31	License Agreement dated July 25, 2017 between Cellectis S.A. and Calyxt, Inc.				Filed herewith

8.1	List of subsidiaries of the registrant				Filed ~~Herewith~~

~~200~~


~~Exhibit~~ ~~Number~~	~~Description of Exhibit~~	~~Schedule/~~ ~~Form~~	~~File Number~~	~~Exhibit~~	~~File Date~~ herewith

12.1	Certificate of Principal Executive Officer pursuant to Securities Exchange Act Rules13a-14(a) and15d-14(a) as adopted pursuant to Section 302 of the Sarbanes-Oxley Act of 2002				Filed ~~Herewith~~herewith

12.2	Certification by the Principal Financial Officer pursuant to Securities Exchange Act Rules13a-14(a) and15d-14(a) as adopted pursuant to Section 302 of the Sarbanes-Oxley Act of 2002				Filed ~~Herewith~~herewith

13.1	Certification by the Principal Executive Officer pursuant to 18 U.S.C. Section 1350, as adopted pursuant to Section 906 of the Sarbanes-Oxley Act of 2002				Filed ~~Herewith~~herewith

13.2	Certification by the Principal Financial Officer pursuant to 18 U.S.C. Section 1350, as adopted pursuant to Section 906 of the Sarbanes-Oxley Act of 2002				Filed ~~Herewith~~herewith

15.1	Consent of Ernst & Young et Autres				Filed ~~Herewith~~herewith

†	Indicates a management contract or any compensatory plan, contract or arrangement.

#	Indicates a document previously filed with the Commission.

*	Portions of this exhibit (indicated by asterisks) have been omitted pursuant to a request for confidential treatment

~~201~~

INDEX TO CONSOLIDATED FINANCIAL STATEMENTS

Financial Statements for the Years Ended December 31, ~~2015,~~ 2016, 2017 and ~~2017:~~2018:


Report of Independent Registered Public Accounting Firm ~~(2/2) ‘~~			F-2
Statements of Consolidated Financial Position as of December 31, ~~2016~~2017 and ~~2017~~2018			F-4
Statements of Consolidated Operations for the Years Ended December 31, ~~2015,~~ 2016, 2017 and ~~2017~~2018			F-5
Statements of Consolidated Comprehensive Income (Loss) for the Years Ended December 31, ~~2015,~~ 2016, 2017 and ~~2017~~2018			F-6
Statements of Consolidated Cash Flows for the Years Ended December 31, ~~2015,~~ 2016, 2017 and ~~2017~~2018			F-7
Statements of Consolidated Changes in Shareholders’ Equity for the Years Ended December 31, ~~2015,~~ 2016, 2017 and ~~2017~~2018			F-8
Notes to the Consolidated Financial Statements			F-10

~~F-1~~

REPORT OF INDEPENDENT REGISTERED PUBLIC ACCOUNTING FIRM

To the Board of Directors and Shareholders of Cellectis S.A.,

Opinion on the Consolidated Financial Statements

We have audited the accompanying statements of consolidated financial position of Cellectis S.A. (the Company) as of December 31, ~~2017~~2018 and ~~2016,~~2017, and the related statements of consolidated operations, consolidated comprehensive loss, consolidated cash flows and changes in consolidated shareholders’ equity for each of the three years in the period ended December 31, ~~2017,~~2018, and the related notes (collectively referred to as the “consolidated financial statements”). In our opinion, the consolidated financial statements present fairly, in all material respects, the consolidated financial position of the Company at December 31, ~~2017~~2018 and ~~2016,~~2017, and the consolidated results of its operations and its cash flows for each of the three years in the period ended December 31, ~~2017,~~2018, in conformity with International Financial Reporting Standards as issued by the International Accounting Standards Board and in conformity with International Financial Reporting Standards as endorsed by the European Union.

We also have audited, in accordance with the standards of the Public Company Accounting Oversight Board (United States) (PCAOB), the Company’s internal control over financial reporting as of December 31, ~~2017,~~2018, based on criteria established in the Internal Control – Integrated Framework issued by the Committee of Sponsoring Organizations of the Treadway Commission (2013 framework) and our report dated March ~~13, 2018~~11, 2019 expressed an unqualified opinion thereon.

Change in Accounting Principle

As discussed in ~~the~~ note ~~2.2 of~~2.3 to the consolidated financial statements, the Company ~~has~~ changed ~~the presentation currency of the consolidated financial statements from the euro~~its revenue recognition method on January 1, 2018, due to the ~~U.S. dollar.~~adoption of IFRS 15 “Revenue from Contracts with Customers”.

Basis for Opinion

These consolidated financial statements are the responsibility of the Company’s management. Our responsibility is to express an opinion on the Company’s consolidated financial statements based on our audits. We are a public accounting firm registered with the PCAOB and are required to be independent with respect to the Company in accordance with the U.S. federal securities laws and the applicable rules and regulations of the Securities and Exchange Commission and the PCAOB.

We conducted our audits in accordance with the standards of the PCAOB. Those standards require that we plan and perform the audit to obtain reasonable assurance about whether the consolidated financial statements are free of material misstatement, whether due to error or fraud. Our audits included performing procedures to assess the risks of material misstatement of the consolidated financial statements, whether due to error or fraud, and performing procedures that respond to those risks. Such procedures included examining, on a test basis, evidence regarding the amounts and disclosures in the consolidated financial statements. Our audits also included evaluating the accounting principles used and significant estimates made by management, as well as evaluating the overall presentation of the consolidated financial statements. We believe that our audits provide a reasonable basis for our opinion.

Ernst & Young et Autres

Ernst & Young et Autres has served as the Company’s auditor since 2012.

Paris-La Défense, March ~~13, 2018~~

~~F-2~~11, 2019

REPORT OF INDEPENDENT REGISTERED PUBLIC ACCOUNTING ~~FIRMS~~FIRM

To the Board of Directors and Shareholders of Cellectis S.A.

Opinion on Internal Control over Financial Reporting

We have audited Cellectis S.A.’s internal control over financial reporting as of December 31, ~~2017,~~2018, based on criteria established in Internal Control – Integrated Framework issued by the Committee of Sponsoring Organizations of the Treadway Commission (2013 Framework) (the COSO criteria). In our opinion, Cellectis S.A. (the Company) maintained, in all material respects, effective internal control over financial reporting as of December 31, ~~2017,~~2018, based on the COSO criteria.

We also have audited, in accordance with the standards of the Public Company Accounting Oversight Board (United States) (PCAOB), the statements of consolidated financial position of the Company as of December 31, ~~2017~~2018 and ~~2016,~~2017, and the related statements of consolidated operations, consolidated comprehensive loss, consolidated cash flows and changes in consolidated shareholders’ equity for each of the three years in the period ended December 31, ~~2017~~2018 and the related notes, and our report dated March ~~13, 2018~~11, 2019 expressed an unqualified opinion thereon.

Basis for Opinion

The Company’s management is responsible for maintaining effective internal control over financial reporting and for its assessment of the effectiveness of internal control over financial reporting included in the accompanying Report of Management on Internal Control Over Financial Reporting. Our responsibility is to express an opinion on the Company’s internal control over financial reporting based on our audit. We are a public accounting firm registered with the PCAOB and are required to be independent with respect to the Company in accordance with the U.S. federal securities laws and the applicable rules and regulations of the Securities and Exchange Commission and the PCAOB.

We conducted our audit in accordance with the standards of the PCAOB. Those standards require that we plan and perform the audit to obtain reasonable assurance about whether effective internal control over financial reporting was maintained in all material respects.

Our audit included obtaining an understanding of internal control over financial reporting, assessing the risk that a material weakness exists, testing and evaluating the design and operating effectiveness of internal control based on the assessed risk, and performing such other procedures as we considered necessary in the circumstances. We believe that our audit provides a reasonable basis for our opinion.

Definition and Limitations of Internal Control Over Financial Reporting

A company’s internal control over financial reporting is a process designed to provide reasonable assurance regarding the reliability of financial reporting and the preparation of financial statements for external purposes in accordance with generally accepted accounting principles. A company’s internal control over financial reporting includes those policies and procedures that (1) pertain to the maintenance of records that, in reasonable detail, accurately and fairly reflect the transactions and dispositions of the assets of the company; (2) provide reasonable assurance that transactions are recorded as necessary to permit preparation of financial statements in accordance with generally accepted accounting principles, and that receipts and expenditures of the company are being made only in accordance with authorizations of management and directors of the company; and (3) provide reasonable assurance regarding prevention or timely detection of unauthorized acquisition, use, or disposition of the company’s assets that could have a material effect on the financial statements.

Because of its inherent limitations, internal control over financial reporting may not prevent or detect misstatements. Also projections of any evaluation of effectiveness to future periods are subject to the risk that controls may become inadequate because of changes in conditions, or that the degree of compliance with the policies and procedures may deteriorate.

Ernst & Young et Autres

Paris-La Défense, March ~~13, 2018~~

~~F-3~~11, 2019

~~Cellectis S.A.~~

Cellectis S.A.

STATEMENTS OF CONSOLIDATED FINANCIAL POSITION

$ in thousands


			As of								As of
	Notes		December 31, 2016			December 31, 2017			Notes		December 31, 2017 as restated (*)			December 31, 2018
ASSETS
Non-current assets
Intangible assets		5		1,343			1,431			5		1,431			1,268
Property, plant, and equipment		6		16,900			7,226			6		7,226			10,041
Othernon-current financial assets				691			1,004			7.2		1,004			1,891

Totalnon-current assets				18,935			9,661					9,661			13,199
Current assets
Inventories		8		118			250			8		250			275
Trade receivables		9.1		3,627			2,753			9.1		2,753			2,971
Subsidies receivables		9.2		8,723			9,524			9.2		9,524			17,173
Other current assets		9.3		8,870			13,713			9.3		13,713			15,333
Current financial assets		10.1		36,592			40,602			10.1		40,602			388
Cash and cash equivalents		10.2		254,568			256,380			10.2		256,380			451,501

Total current assets				312,498			323,221					323,221			487,641

TOTAL ASSETS				331,432			332,882					332,882			500,840

LIABILITIES
Shareholders’ equity
Share capital		14.1		2,332			2,367			14		2,367			2,765
Premiums related to the share capital		14.1		568,185			614,037			14		614,037			828,525
Treasury share reserve		14.4		(416	)		(297	)				(297	)		—
Currency translation adjustment				(22,174	)		1,978					1,834			(16,668	)
Retained earnings (deficit)				(207,875	)		(251,927	)				(253,702	)		(326,628	)
Net income (loss)				(67,255	)		(99,368	)				(99,368	)		(78,693	)

Total shareholders’ equity—Group Share				272,795			266,791
Total shareholders’ equity - Group Share												264,872			409,301
Non-controlling interests		14.3		1,876			19,113					19,113			40,970

Total shareholders’ equity				274,671			285,904					283,985			450,272
Non-current liabilities
Non-current financial liabilities		11		30			13			11		13			1,018
Non-current provisions		17		560			3,430			17		3,430			2,681

Totalnon-current liabilities				590			3,443					3,443			3,699

Current liabilities
Current financial liabilities		11		1,730			21			11		21			333
Trade payables				9,722			9,460			11		9,460			15,883
Deferred revenues and deferred income		13		38,929			26,056
Deferred revenues and contract liabilities										13		27,975			20,754
Current provisions		17		594			1,427			17		1,427			1,530
Other current liabilities		12		5,196			6,570			12		6,570			8,369

Total current liabilities				56,171			43,534					45,453			46,869

TOTAL LIABILITIES AND SHAREHOLDERS’ EQUITY				331,432			332,882					332,882			500,840

The accompanying notes form an integral part of these Consolidated Financial Statements

~~F-4~~
(*)
2017 consolidated financial statements have been restated for the purpose of IFRS15 application. Reconciliation between the consolidated financial statements presented in previous periods and the 2018 consolidated financial statements is available in Note 2.3.

~~Cellectis S.A.~~

Cellectis S.A.

STATEMENTS OF CONSOLIDATED OPERATIONS

For the year ended December 31

$ in thousands, except per share amounts


			For the year ended December 31,											For the year ended December 31,
	Notes		2015			2016			2017			Notes		2016			2017			2018
Revenues and other income

Revenues		3.1		55,864			44,808			25,188			3.1		44,808			25,188			12,731

Other income		3.1		6,701			11,637			8,528			3.1		11,637			8,528			8,701

Total revenues and other income				62,565			56,444			33,715					56,444			33,715			21,432

Operating expenses

Royalty expenses		3.2		(2,746	)		(1,777	)		(2,620	)		3.2		(1,777	)		(2,620	)		(2,739	)
Research and development expenses		3.2		(58,154	)		(78,458	)		(79,227	)		3.2		(78,458	)		(79,227	)		(76,567	)
Selling, general and administrative expenses		3.2		(30,223	)		(43,413	)		(44,750	)		3.2		(43,413	)		(44,750	)		(47,248	)
Other operating income and expenses				(2,425	)		(99	)		232
Other operating income (expenses)															(99	)		232			31

Total operating expenses				(93,549	)		(123,746	)		(126,366	)				(123,746	)		(126,366	)		(126,523	)

Operating income (loss)				(30,984	)		(67,302	)		(92,650	)				(67,302	)		(92,650	)		(105,091	)

Financial income		3.4		10,253			7,147			7,262			3.3		7,147			7,262			20,572
Financial expenses		3.4		(1,876	)		(7,101	)		(18,294	)		3.3		(7,101	)		(18,294	)		(3,813	)

Financial gain (loss)				8,378			46			(11,032	)				46			(11,032	)		16,758

Income tax		3.5		—			—			—					—			—			—

Net income (loss)				(22,606	)		(67,255	)		(103,683	)				(67,255	)		(103,683	)		(88,333	)

Attributable to shareholders of Cellectis				(22,796	)		(67,255	)		(99,368	)				(67,255	)		(99,368	)		(78,693	)
Attributable tonon-controlling interests				190			—			(4,315	)				—			(4,315	)		(9,640	)
Basic / Diluted net income (loss) per share attributable to shareholders of Cellectis		16											16
Basic net income (loss) per share ($ /share)				(0.67	)		(1.91	)		(2.78	)				(1.91	)		(2.78	)		(1.93	)
Diluted net income (loss) per share ($ /share)				(0.67	)		(1.91	)		(2.78	)				(1.91	)		(2.78	)		(1.93	)

The accompanying notes form an integral part of these Consolidated Financial Statements

~~F-5~~

STATEMENTS OF CONSOLIDATED COMPREHENSIVE INCOME (LOSS)

For the year ended December 31

$ in thousands


	For the year ended December 31,									For the year ended December 31,
	2015			2016			2017			2016			2017			2018
Net income (loss)		(22,606	)		(67,255	)		(103,683	)		(67,255	)		(103,683	)		(88,333	)

Actuarial gains and losses		(15	)		(30	)		(515	)		(30	)		(515	)		70

Other comprehensive income (loss) that will not be reclassified subsequently to income or loss		(15	)		(30	)		(515	)		(30	)		(515	)		70

Currency translation adjustment		(6,769	)		(4,333	)		23,745			(4,278	)		23,512			(19,192	)

Other comprehensive income (loss) that will be reclassified subsequently to income or loss		(6,769	)		(4,333	)		23,745			(4,278	)		23,512			(19,192	)

Total Comprehensive income (loss)		(29,391	)		(71,618	)		(80,453	)		(71,563	)		(80,686	)		(107,455	)

Attributable to shareholders of Cellectis		(29,640	)		(71,607	)		(75,731	)		(71,551	)		(75,963	)		(97,125	)
Attributable tonon-controlling interests		249			(12	)		(4,723	)		(12	)		(4,723	)		(10,330	)

The accompanying notes form an integral part of these Consolidated Financial Statements

~~F-6~~

~~Cellectis S.A.~~

Cellectis S.A.

STATEMENTS OF CONSOLIDATED CASH FLOWS

For the year ended December 31

$ in thousands


			For the year ended December 31,											For the year ended December 31,
	Notes		2015			2016			2017			Notes		2016			2017			2018
Cash flows from operating activities
Net loss for the period				(22,606	)		(67,255	)		(103,683	)				(67,255	)		(103,683	)		(88,333	)

Reconciliation of net loss and of the cash provided by (used in) operating activities
Adjustments for
Amortization and depreciation				1,937			2,211			3,371					2,211			3,371			2,377
Net loss (income) on disposals				(11	)		65			40					65			40			20
Net financial loss (gain)				(8,378	)		(46	)		11,032					(46	)		11,032			(16,759	)
Expenses related to share-based payments				33,402			58,622			50,418					58,622			50,418			37,218
Provisions				(279	)		(365	)		2,908					(365	)		2,908			(468	)
Other non cash items				—			(1,432	)		2					(1,432	)		2			—
Interest (paid) / received				1,070			1,694			1,371					1,694			1,371			6,905

Operating cash flows before change in working capital				5,135			(6,507	)		(34,540	)				(6,507	)		(34,540	)		(59,040	)

Decrease (increase) in inventories				(25	)		50			(109	)				50			(109	)		(37	)
Decrease (increase) in trade receivables and other current assets				1,268			(997	)		(549	)				(997	)		(549	)		(3,696	)
Decrease (increase) in subsidies receivables				(679	)		(1,122	)		305					(1,122	)		305			(8,257	)
(Decrease) increase in trade payables and other current liabilities				2,961			(4,384	)		(335	)				(4,384	)		(335	)		9,374
(Decrease) increase in deferred income				(5,070	)		(19,750	)		(17,099	)
(Decrease) increase in deferred income and contract liabilities															(19,750	)		(17,099	)		(6,480	)

Change in working capital				(1,544	)		(26,203	)		(17,787	)				(26,203	)		(17,787	)		(9,096	)

Net cash flows provided by (used in) operating activities				3,591			(32,710	)		(52,327	)				(32,710	)		(52,327	)		(68,137	)

Cash flows from investment activities
Proceeds from disposal of property, plant and equipment				111			24			7,164					24			7,164			1,262
Sale (Acquisition) of subsidiaries net of cash disposed of				(3,162	)		—			—
Acquisition of intangible assets				(97	)		(337	)		(273	)				(337	)		(273	)		(171	)
Acquisition of property, plant and equipment				(4,316	)		(13,696	)		(2,383	)				(13,696	)		(2,383	)		(4,715	)
Net change innon-current financial assets				(264	)		175			(125	)				175			(125	)		221
Sale (Acquisition) of current financial assets				—			(39,302	)		(2,598	)				(39,302	)		(2,598	)		39,025

Net cash flows provided by (used in) investing activities				(7,728	)		(53,137	)		1,784					(53,137	)		1,784			35,623

Cash flows from financing activities
Increase in share capital net of transaction costs				221,142			713			2,930					713			2,930			186,382
Shares of Calyxt issued to third parties				—			—			38,257					—			38,257			49,942
Decrease in borrowings				(625	)		(91	)		(41	)				(91	)		(41	)		(127	)
Treasury shares				75			(137	)		120					(137	)		120			297

Net cash flows provided by financing activities				220,591			485			41,266					485			41,266			236,494

(Decrease) increase in cash				216,454			(85,362	)		(9,277	)				(85,362	)		(9,277	)		203,981

Cash and cash equivalents at the beginning of the year				136,400			342,111			254,568					342,111			254,568			256,380
Effect of exchange rate changes on cash				(10,743	)		(2,181	)		11,089					(2,181	)		11,089			(8,860	)

Cash and cash equivalents at the end of the period		7		342,111			254,568			256,380			10		254,568			256,380			451,501

We present our consolidated statements of cash flows using the indirect method.

The accompanying notes form an integral part of these Consolidated Financial Statements

~~F-7~~

~~Cellectis S.A.~~

Cellectis S.A.

STATEMENTS OF CHANGES IN CONSOLIDATED SHAREHOLDERS’ EQUITY

For the year ended December 31

$ in thousands, except share data

   Share Capital
Ordinary Shares       Equity
   Notes    Number of
shares    Amount    Premiums
related to
share
capital    Treasury
shares
reserve Currency
translation
adjustment Retained
earnings
(deficit) Income
(Loss) attributable to
shareholders
of Cellectis Non
controlling
interests Total
Shareholders’
Equity
As of January 1, 2015
      29,446,721    2,014    263,100    (354 ) (11,024 ) (179,962 ) 27 73,801 (1,529 ) 72,272
Net Loss
      —      —      —      —   —   —   (22,796 ) (22,796 ) 190 (22,606 )
Other comprehensive income (loss)
      —      —      —      —   (6,829 ) (15 ) —   (6,844 ) 59 (6,785 )








Total comprehensive income (loss)
      —      —      —      —   (6,829 ) (15 ) (22,796 ) (29,640 ) 249 (29,391 )
Allocation of prior period loss
      —      —      —      —   —   27 (27 ) —   —   —
Capital Increase
   14.1    5,500,000    297    209,899    —   —   (6 ) —   210,189 —   210,189
Purchase ofnon-controlling interests
   14.3    —      —      —      —   —   (5,164 ) —   (5,164 ) 1,280 (3,884 )
Treasury shares
   14.4    —      —      —      75 —   —   —   75 —   75
Exercise of share warrants and employee warrants
   14.2    231,893    13    4,325    —   —   —   —   4,338 —   4,338
Non-cash stock-based compensation expense
   15    —      —      32,614    —   —   —   —   32,614 789 33,403








As of December 31, 2015
      35,178,614    2,323    509,938    (279 ) (17,853 ) (185,120 ) (22,796 ) 286,212 789 287,002








As of January 1, 2016
      35,178,614    2,323    509,938    (279 ) (17,853 ) (185,120 ) (22,796 ) 286,212 789 287,002
Net Loss
      —      —      —      —   —   —   (67,255 ) (67,255 ) —   (67,255 )
Other comprehensive income (loss)
      —      —      —      —   (4,321 ) (30 ) —   (4,352 ) (12 ) (4,363 )








Total comprehensive income (loss)
      —      —      —      —   (4,321 ) (30 ) (67,255 ) (71,607 ) (12 ) (71,618 )
Allocation of prior period loss
      —      —      —      —   —   (22,796 ) 22,796 —   —   —
Treasury shares
      —      —      —      (137 ) —   —   —   (137 ) —   (137 )
Exercise of share warrants and employee warrants
      156,446    9    723    —   —   (6 ) —   726 —   726
Non-cash stock-based compensation expense
   12    —      —      57,524    —   —   —   —   57,524 1,098 58,622
Other movements
      —      —      —      —   —   77 —   77 —   77








As of December 31, 2016
      35,335,060    2,332    568,185    (416 ) (22,174 ) (207,875 ) (67,255 ) 272,795 1,876 274,671








~~F-8~~


	Share Capital Ordinary Shares																					Equity											Share Capital Ordinary Shares																			Equity
	Notes		Number of shares		Amount		Premiums related to share capital			Treasury shares reserve			Currency translation adjustment			Retained earnings (deficit)			Income (Loss)			attributable to shareholders of Cellectis			Non controlling interests			Total Shareholders’ Equity			Notes		Number of shares		Amount		Premiums related to share capital			Treasury shares reserve			Currency translation adjustment			Retained earnings (deficit)			Income (Loss)			attributable to shareholders of Cellectis			Non controlling interests			Total Shareholders’ Equity
As of January 1, 2017				35,335,060		2,332		568,185			(416	)		(22,174	)		(207,875	)		(67,255	)		272,795			1,876			274,671
As of January 1, 2016, as restated (*)																																		35,178,614		2,323		509,938			(279	)		(17,819	)		(185,120	)		(24,571	)		284,471			789			285,260
Net Loss																																		—		—		—			—			—			—			(67,255	)		(67,255	)		—			(67,255	)
Other comprehensive income (loss)																																		—		—		—			—			(4,266	)		(30	)		—			(4,296	)		(12	)		(4,308	)

Total comprehensive income (loss)																																		—		—		—			—			(4,266	)		(30	)		(67,255	)		(71,551	)		(12	)		(71,563	)
Allocation of prior period loss																																		—		—		—			—			—			(24,571	)		24,571			—			—			—
Treasury shares																																		—		—		—			(137	)		—			—			—			(137	)		—			(137	)
Exercise of share warrants and employee warrants																																14.1		156,446		9		723			—			—			(6	)		—			726			—			726
Share based compensation																																15		—		—		57,524			—			—			—			—			57,524			1,098			58,622
Other movements																																		—		—		—			—			—			77			—			77			—			77

As of December 31, 2016, as restated (*)																																		35,335,060		2,332		568,185			(416	)		(22,085	)		(209,651	)		(67,255	)		271,109			1,876			272,984

As of January 1, 2017, as restated (*)																																		35,335,060		2,332		568,185			(416	)		(22,085	)		(209,651	)		(67,255	)		271,109			1,876			272,984
Net Loss				—		—		—			—			—			—			(99,368	)		(99,368	)		(4,315	)		(103,683	)				—		—		—			—			—			—			(99,368	)		(99,368	)		(4,315	)		(103,683	)
Other comprehensive income (loss)				—		—		—			—			24,152			(515	)		—			23,637			(408	)		23,230					—		—		—			—			23,920			(515	)		—			23,405			(408	)		22,997

Total comprehensive income (loss)				—		—		—			—			24,152			(515	)		(99,368	)		(75,731	)		(4,723	)		(80,453	)				—		—		—			—			23,920			(515	)		(99,368	)		(75,963	)		(4,723	)		(80,686	)
Allocation of prior period loss				—		—		—			—			—			(67,255	)		67,255			—			—			—					—		—		—			—			—			(67,255	)		67,255			—			—			—
Capital Increase				466,950		26		—			—			—			(26	)		—			—			—			—			14.1		466,950		26		—			—			—			(26	)		—			—			—			—
Transaction with subsidiaries (1)				—		—		—			—			—			23,747			—			23,747			14,510			38,257					—		—		—			—			—			23,747			—			23,747			14,510			38,257
Treasury shares				—		—		—			120			—			—			—			120			—			120					—		—		—			120			—			—			—			120			—			120
Exercise of share warrants and employee warrants		11		158,052		9		2,921			—			—			—			—			2,930			—			2,930
Exercise of share warrants, employee warrants and stock options																																14.1		158,052		9		2,921			—			—			—			—			2,930			—			2,930
Non-cash stock-based compensation expense		12		—		—		42,968			—			—			—			—			42,968			7,450			50,418			15		—		—		42,968			—			—			—			—			42,968			7,450			50,418
Other movements				—		—		(37	)		—			—			(1	)		—			(38	)		—			(38	)				—		—		(37	)		—			—			(1	)		—			(38	)		—			(38	)

As of December 31, 2017				35,960,062		2,367		614,037			(297	)		1,978			(251,927	)		(99,368	)		266,791			19,113			285,904
As of December 31, 2017, as restated (*)																																		35,960,062		2,367		614,037			(297	)		1,835			(253,702	)		(99,368	)		264,873			19,113			283,986

As of January 1, 2018, as restated (*)																																		35,960,062		2,367		614,037			(297	)		1,835			(253,702	)		(99,368	)		264,873			19,113			283,986
Net Loss																																		—		—		—			—						—			(78,693	)		(78,693	)		(9,640	)		(88,333	)
Other comprehensive income (loss)																																		—		—		—			—			(18,502	)		70			—			(18,432	)		(690	)		(19,122	)

Total comprehensive income (loss)																																		—		—		—			—			(18,502	)		70			(78,693	)		(97,125	)		(10,330	)		(107,455	)
Allocation of prior period loss																																		—		—		—			—			—			(99,368	)		99,368			—			—			—
Capital Increase																																14.1		6,146,000		379		178,230			—			—			2			—			178,611			—			178,611
Transaction with subsidiaries (2)																																		—		—		—			—			—			26,454			—			26,454			23,488			49,942
Treasury shares																																		—		—		—			297			—			(58	)		—			239			—			239
Exercise of share warrants, employee warrants and stock options																																14.1		324,007		19		7,751			—			—			—			—			7,770			—			7,770
Non-cash stock-based compensation expense																																15		—		—		28,507			—			—			—			—			28,507			8,711			37,218
Other movements																																		—		—		—			—			—			(28	)		—			(28	)		(12	)		(40	)

As of December 31, 2018																																		42,430,069		2,765		828,525			—			(16,668	)		(326,628	)		(78,693	)		409,301			40,970			450,272

The accompanying notes form an integral part of these Consolidated Financial Statements

(1) Net proceeds to Calyxt from the Calyxt IPO of $58.0 million after deduction of $3.1 million of underwriting discounts and commissions and $3.3 million of other offering expenses. Equity of Calyxt attributable to~~non-controlling~~ interests of 20.3% was $11.8 million and equity of Calyxt attributable to Cellectis of 79.73% is $26.4 million (after consideration of Cellectis’ investment in shares of Calyxt issued as part of the Calyxt IPO for a purchase price of $20 million).

~~F-9~~

~~Cellectis began its IFRS 15 implementation project with a diagnostic phase. The different categories of contracts with customers of Cellectis, which have been reviewed, are:~~


Title of each class		Name of each exchange on which registered
American Depositary Shares, each representing one ordinary share, nominal value €0.05 per share		Nasdaq Global Market
Ordinary shares, nominal value €0.05 per share*		Nasdaq Global Market*


		PAGE
INTRODUCTION			i2

SPECIAL NOTE REGARDING FORWARD-LOOKING STATEMENTS			i2

PART I

Item 1.	Identity of Directors, Senior Management and Advisers		15

Item 2.	Offer Statistics and Expected Timetable		15

Item 3.	Key Information		15

Item 4.	Information on the Company		6763

Item 4A.	Unresolved Staff Comments		~~125~~109

Item 5.	Operating and Financial Review and Prospects		~~125~~109

Item 6.	Directors, Senior Management and Employees		~~144~~126

Item 7.	Major Shareholders and Related Party Transactions		~~158~~137

Item 8.	Financial Information		~~166~~144

Item 9.	The Offer and Listing		~~167~~145

Item 10.	Additional Information		~~168~~145

Item 11.	Quantitative and Qualitative Disclosures About Market Risk		~~189~~163

Item 12.	Description of Securities Other than Equity Securities		~~190~~164

PART II

Item 13.	Defaults, Dividend Arrearages and Delinquencies		~~192~~166

Item 14.	Material Modifications to the Rights of Security Holders and Use of Proceeds		~~192~~166

Item 15.	Controls and Procedures		~~192~~167

Item 15T.	Controls and Procedures		~~193~~167

Item 16.	Reserved		~~193~~167

Item 16A.	Audit Committee Financial Expert		~~193~~167

Item 16B.	Code of Ethics		~~193~~167

Item 16C.	Principal Accountant Fees and Services		~~194~~168

Item 16D.	Exemptions from the Listing Standards for Audit Committees		~~195~~168

Item 16E.	Purchases of Equity Securities by the Issuer and Affiliated Purchasers		~~195~~169

Item 16F.	Change in Registrant’s Certifying Accountant		~~195~~169

Item 16G.	Corporate Governance		~~195~~169

Item 16H.	Mine Safety Disclosure		~~198~~171

PART III

Item 17.	Financial Statements		~~198~~172

Item 18.	Financial Statements		~~198~~172

Item 19.	Exhibits		~~198~~172


	Year Ended December 31,
	2013		2014		2015		2016		2017
High		1.381		1.395		1.204		1.156		1.206
Low		1.276		1.214		1.055		1.036		1.038
Rate at end of period		1.379		1.214		1.088		1.054		1.199
Average rate per period		1.338		1.328		1.109		1.106		1.129


	September 2017		October 2017		November 2017		December 2017		January 2018		February 2018
High		1.206		1.185		1.195		1.199		1.249		1.249
Low		1.174		1.160		1.156		1.073		1.193		1.221
Rate at end of period		1.180		1.163		1.184		1.199		1.245		1.221


	2013 (4)			2014			2015			2016			2017
	($ in thousands, except share and per share data)
Revenues and other income		16,900			35,151			62,565			56,444			33,715

Operating expenses
Royalty expenses		(720	)		(4,033	)		(2,746	)		(1,777	)		(2,620	)
Research and development expenses		(23,700	)		(19,144	)		(58,154	)		(78,458	)		(79,227	)
Selling, general and administrative expenses		(25,281	)		(17,426	)		(30,223	)		(43,413	)		(44,750	)
Other operating income and expenses		(2,433	)		(1,517	)		(2,425	)		(99	)		232

Operating income (loss)		(35,234	)		(6,970	)		(30,984	)		(67,302	)		(92,650	)

Loss from discontinued operations		(39,288	)		(3,750	)		—			—			—
Financial gain (loss)		(414	)		9,428			8,378			46			(11,032	)

Net income (loss)		(74,936	)		(1,292	)		(22,606	)		(67,255	)		(103,683	)


			As of December 31,										As of December 31,
	2013 (1)		2014		2015		2016		2017		2014 (1)		2015 (1)		2016 (2)		2017 (2)		2018
	($ in thousands)										$ in thousands
Current financial assets and Cash and cash equivalents		10,425		136,400		342,111		291,159		296,982		136,400		342,111		291,159		296,982		451,889
Total assets		39,822		167,077		371,314		331,432		332,882		167,077		371,314		331,432		332,882		500,840
Total shareholders’ equity		3,471		72,272		287,002		274,671		285,904		72,272		287,002		272,985		283,985		450,272
Total non current liabilities		5,257		3,912		547		590		3,443
Totalnon-current liabilities												3,912		547		590		3,443		3,699
Total current liabilities		31,093		90,894		83,765		56,171		43,534		90,894		83,765		57,857		45,453		46,869


	For the years ended December 31,
	2014		2015			2016			2017			2018
	$ in thousands
Net Income (Loss) attributable to shareholders of Cellectis		27		(22,796	)		(67,255	)		(99,368	)		(78,693	)
Adjustment ofnon-cash stock-based compensation expense:
Research and development expenses		299		20,563			33,207			23,832			18,057
Selling, general and administrative expenses		430		12,839			25,415			26,586			19,161

Totalnon-cash stock-based compensation expense:		728		33,402			58,622			50,418			37,218

Non-cash stock-based compensation expense attributable to non controlling interests		—		—			—			(1,493	)		(2,655	)

Adjusted Net Income (Loss) attributable to shareholders of Cellectis		755		10,606			(8,633	)		(50,443	)		(44,130	)


	For the year ended December 31, 2015									For the year ended December 31, 2016									For the year ended December 31, 2017
$ in thousands	Plants			Therapeutics			Total reportable segments			Plants			Therapeutics			Total reportable segments			Plants			Therapeutics			Total reportable segments
Segment revenues and other income (1)		49			65,159			65,208			716			59,458			60,173			914			35,584			36,498
Inter-segment revenues (1)		—			(2,643	)		(2,643	)		(130	)		(3,599	)		(3,729	)		(167	)		(2,615	)		(2,782	)

External revenues and other income		49			62,516			62,565			585			55,859			56,444			747			32,969			33,715

Research and development expenses		(2,874	)		(55,280	)		(58,154	)		(4,112	)		(74,345	)		(78,458	)		(6,057	)		(73,170	)		(79,227	)
Selling, general and administrative expenses		(1,834	)		(28,390	)		(30,223	)		(4,809	)		(38,603	)		(43,413	)		(13,143	)		(31,607	)		(44,750	)
Royalties and other operating income and expenses		(272	)		(4,899	)		(5,171	)		(474	)		(1,402	)		(1,876	)		(384	)		(2,005	)		(2,389	)

Total operating expenses		(4,980	)		(88,569	)		(93,549	)		(9,395	)		(114,351	)		(123,746	)		(19,584	)		(106,782	)		(126,366	)

Operating income (loss) before tax		(4,931	)		(26,053	)		(30,984	)		(8,810	)		(58,492	)		(67,302	)		(18,837	)		(73,813	)		(92,650	)

Financial gain (loss)		259			8,119			8,378			87			(41	)		46			—			(11,032	)		(11,032	)

Net income (loss)		(4,672	)		(17,934	)		(22,606	)		(8,722	)		(58,533	)		(67,255	)		(18,837	)		(84,846	)		(103,683	)

Non controlling interests		—			(190	)		(190	)		—			—			—			4,315			—			4,315

Net income (loss) attributable to shareholders of Cellectis		(4,672	)		(18,124	)		(22,796	)		(8,722	)		(58,533	)		(67,255	)		(14,522	)		(84,846	)		(99,368	)

Adjustment of share-based compensation		789			32,613			33,402			1,098			57,524			58,622			5,957			42,968			48,925

Adjusted net income (loss) attributable to shareholders of Cellectis		(3,883	)		14,489			10,606			(7,625	)		(1,009	)		(8,633	)		(8,565	)		(41,877	)		(50,442	)

Depreciation and amortization		(99	)		(1,838	)		(1,937	)		(345	)		(1,866	)		(2,211	)		(551	)		(2,820	)		(3,371	)
Additions to tangible and intangible assets		526			3,886			4,413			10,410			4,164			14,573			792			1,849			2,642
Impairment of tangible assets		—			—			—			—			—			—			—			(798	)		(798	)


Name ⁽¹⁾	Current Position		Year of Initial Appointment		Term Expiration Year
André Choulika, Ph.D.		Chairman		2000		2018
David Sourdive, Ph.D.		Director		2000		2018
Alain Godard		Director		2007		2018
Pierre Bastid		Director		2011		2020
Laurent Arthaud		Director		2011		2020
Annick Schwebig, M.D.		Director		2011		2020
Jean-Marie Messier		Director		2015		2018
Hervé Hoppenot		Director		2017		2020
Rainer Boehm		Director		2017		2020


Period	High		Low
Annual
2015 (beginning March 24, 2015)	$	47.66	$	23.67
2016	$	33.64	$	16.40
2017	$	35.01	$	17.52
2018 (through March 12, 2018)	$	34.17	$	27.78
Quarterly
First Quarter 2016	$	30.16	$	18.77
Second Quarter 2016	$	33.64	$	25.34
Third Quarter 2016	$	28.35	$	24.07
Fourth Quarter 2016	$	23.24	$	16.40
First Quarter 2017	$	24.37	$	17.52
Second Quarter 2017	$	25.92	$	21.96
Third Quarter 2017	$	32.18	$	21.65
Fourth Quarter 2017	$	35.01	$	23.33
First Quarter 2018 (through March 12, 2018)	$	34.17	$	27.78
Monthly
September 2017	$	32.18	$	24.96
October 2017	$	35.01	$	27.51
November 2017	$	29.18	$	23.33
December 2017	$	29.15	$	23.70
January 2018	$	34.17	$	27.78
February 2018	$	32.65	$	28.53
March 2018 (through March 12, 2018)	$	34.07	$	31.57


As of December 31, 2014	Consolidated financial statement as reported (€ in thousand)			Consolidated financial statement as reported converted (a) ($ in thousand)			Adjustments (b)			Consolidated financial statement ($ in thousand)
As of December 31, 2016													Consolidated financial statement as reported (€ in thousand)			Consolidated financial statement as reported converted (a) ($ in thousand)			Adjustments (b)			Consolidated financial statement ($ in thousand)
Totalnon-current assets		5,613			6,815			—			6,815			17,963			18,935			—			18,935
Total current assets		132,001			160,262			—			160,262			296,459			312,497			—			312,497

TOTAL ASSETS		137,614			167,077			—			167,077			314,422			331,432			—			331,432

Shareholders’ equity
Share capital		1,472			1,788			226			2,014
Capital														1,767			1,862			470			2,332
Premiums, Retained earnings (deficit) and Net income (loss)		60,327			73,243			9,922			83,165			254,834			268,622			24,432			293,054
Treasury share reserve		(251	)		(305	)		(49	)		(354	)		(307	)		(324	)		(92	)		(416	)
Currency translation adjustment		(762	)		(925	)		(10,099	)		(11,024	)		2,501			2,636			(24,810	)		(22,174	)

Total shareholders’ equity—Group Share		60,786			73,801			—			73,801
Total shareholders’ equity - Group Share														258,795			272,795			—			272,795

Non-controlling interests		(1,259	)		(1,529	)		—			(1,529	)		1,779			1,875			—			1,875
Total shareholders’ equity		59,527			72,272			—			72,272			260,574			274,671			—			274,671
Totalnon-current liabilities		3,222			3,911			—			3,911			560			590			—			590
Total current liabilities		74,865			90,894			—			90,894			53,288			56,171			—			56,171

TOTAL LIABILITIES AND SHAREHOLDERS’ EQUITY		137,614			167,077			—			167,077			314,422			331,432			—			331,432


As of December 31, 2015	Consolidated financial statement as reported (€ in thousand)			Consolidated financial statement as reported converted (a) ($ in thousand)			Adjustments (b)			Consolidated financial statement ($ in thousand)
Totalnon-current assets		6,844			7,451			—			7,451
Total current assets		334,218			363,863			—			363,863

TOTAL ASSETS		341,062			371,314			—			371,314

Shareholders’ equity
Capital		1,759			1,915			408			2,323
Premiums, Retained earnings (deficit) and Net income (loss)		262,950			286,274			15,748			302,021
Treasury share reserve		(184	)		(200	)		(80	)		(279	)
Currency translation adjustment		(1,632	)		(1,776	)		(16,077	)		(17,853	)

Total shareholders’ equity—Group Share		262,894			286,213			—			286,213

Non-controlling interests		725			789			—			789
Total shareholders’ equity		263,619			287,002			—			287,002
Totalnon-current liabilities		503			548			—			548
Total current liabilities		76,940			83,765			—			83,765

TOTAL LIABILITIES AND SHAREHOLDERS’ EQUITY		341,062			371,314			—			371,314


For the full year ended December 31, 2014	Consolidated financial statement as reported (€ in thousand)			Consolidated financial statement as reported converted (a) ($ in thousand)			Adjustments		Consolidated financial statement ($ in thousand)
For the full year ended December 31, 2015												Consolidated financial statement as reported (€ in thousand)			Consolidated financial statement as reported converted (a) ($ in thousand)			Adjustments		Consolidated financial statement ($ in thousand)
Total revenues and other income		26,453			35,152			—		35,152			56,385			62,565			—		62,565
Total operating expenses		(31,698	)		(42,122	)		—		(42,122	)		(84,309	)		(93,549	)		—		(93,549	)
Operating income (loss)		(5,245	)		(6,970	)		—		(6,970	)		(27,924	)		(30,984	)		—		(30,984	)
Financial gain (loss)		7,095			9,428			—		9,428			7,550			8,378			—		8,378
Income tax		—			—			—		—			—			—			—		—
Income (loss) from continuing operations		1,850			2,458			—		2,458			(20,373	)		(22,606	)		—		(22,606	)
Loss from discontinued operations		(2,822	)		(3,750	)		—		(3,750	)		—			—			—		—
Net income (loss)		(972	)		(1,292	)		—		(1,292	)		(20,373	)		(22,606	)		—		(22,606	)


	January 1^st, 2016 as presented			IFRS 15 restatement			January 1st, 2016 as restated
	$ in thousands
Total non-current assets		7,451			—			7,451

Total current assets		363,863			—			363,863

TOTAL ASSETS		371,314			—			371,314

Shareholders’ equity								—
Share capital		2,323			—			2,323
Premiums related to the share capital		509,938			—			509,938
Treasury share reserve		(279	)		—			(279	)
Currency translation adjustment		(17,853	)		34			(17,819	)
Retained earnings (deficit)		(185,120	)		—			(185,120	)
Net income (loss)		(22,796	)		(1,775	)		(24,571	)

Total shareholders’ equity - Group Share		286,212			(1,742	)		284,471
Non-controlling interests		789			—			789

Total shareholders’ equity		287,002			(1,742	)		285,260

Totalnon-current liabilities		548			—			548

Current liabilities								—
Current financial liabilities		2,091			—			2,091
Trade payables		7,197			—			7,197
Deferred revenues and contract liabilities		59,615			1,742			61,357
Current provisions		1,038			—			1,038
Other current liabilities		13,823			—			13,823

Total current liabilities		83,765			1,742			85,506

TOTAL LIABILITIES AND SHAREHOLDERS’ EQUITY		371,314			—			371,314


	December 31, 2016 as presented			IFRS 15 restatement			December 31, 2016 as restated
				$ in thousands
Total non-current assets		18,935			—			18,935

Total current assets		312,498			—			312,498

TOTAL ASSETS		331,432			—			331,432

Shareholders’ equity								—
Share capital		2,332			—			2,332
Premiums related to the share capital		568,185			—			568,185
Treasury share reserve		(416	)		—			(416	)
Currency translation adjustment		(22,174	)		89			(22,085	)
Retained earnings (deficit)		(207,875	)		(1,775	)		(209,650	)
Net income (loss)		(67,255	)		—			(67,255	)

Total shareholders’ equity - Group Share		272,795			(1,686	)		271,109
Non-controlling interests		1,876			—			1,876

Total shareholders’ equity		274,671			(1,686	)		272,985

Totalnon-current liabilities		590			—			590

Current liabilities								—
Current financial liabilities		1,730			—			1,730
Trade payables		9,722			—			9,722
Deferred revenues and contract liabilities		38,929			1,686			40,615
Current provisions		594			—			594
Other current liabilities		5,196			—			5,196

Total current liabilities		56,171			1,686			57,857

TOTAL LIABILITIES AND SHAREHOLDERS’ EQUITY		331,432			—			331,432


	December 31, 2017 as presented			IFRS 15 restatement			December 31, 2017 as restated
	$ in thousands
Totalnon-current assets		9,661			—			9,661

Total current assets		323,221			—			323,221

TOTAL ASSETS		332,882			—			332,882

Shareholders’ equity								—
Share capital		2,367			—			2,367
Premiums related to the share capital		614,037			—			614,037
Treasury share reserve		(297	)		—			(297	)
Currency translation adjustment		1,978			(144	)		1,834
Retained earnings (deficit)		(251,927	)		(1,775	)		(253,702	)
Net income (loss)		(99,368	)		—			(99,368	)

Total shareholders’ equity - Group Share		266,791			(1,919	)		264,873
Non-controlling interests		19,113			—			19,113

Total shareholders’ equity		285,904			(1,919	)		283,985

Totalnon-current liabilities		3,443			—			3,443

Current liabilities								—
Current financial liabilities		21			—			21
Trade payables		9,460			—			9,460
Deferred revenues and contract liabilities		26,056			1,919			27,975
Current provisions		1,427			—			1,427
Other current liabilities		6,570			—			6,570

Total current liabilities		43,534			1,919			45,453

TOTAL LIABILITIES AND SHAREHOLDERS’ EQUITY		332,882			—			332,882


	As of December 31,		IFRS 15		As of December 31,
	2016, as presented		restatement		2016, as restated
	$ in thousands
Deferred revenues and contract liabilities		38,929		1,686		40,615

Total Deferred revenue and contract liabilities		38,929		1,686		40,615


	For the year ended December 31, 2015									For the year ended December 31, 2016									For the year ended December 31, 2017
$ in thousands	Plants			Therapeutics			Total reportable segments			Plants			Therapeutics			Total reportable segments			Plants			Therapeutics			Total reportable segments
Segment revenues and other income (1)		49			65,159			65,208			716			59,458			60,173			914			35,584			36,498
Inter-segment revenues (1)		—			(2,643	)		(2,643	)		(130	)		(3,599	)		(3,729	)		(167	)		(2,615	)		(2,782	)

External revenues and other income		49			62,516			62,565			585			55,859			56,444			747			32,969			33,715

Research and development expenses		(2,874	)		(55,280	)		(58,154	)		(4,112	)		(74,345	)		(78,458	)		(6,057	)		(73,170	)		(79,227	)
Selling, general and administrative expenses		(1,834	)		(28,390	)		(30,223	)		(4,809	)		(38,603	)		(43,413	)		(13,143	)		(31,607	)		(44,750	)
Royalties and other operating income and expenses		(272	)		(4,899	)		(5,171	)		(474	)		(1,402	)		(1,876	)		(384	)		(2,005	)		(2,389	)

Total operating expenses		(4,980	)		(88,569	)		(93,549	)		(9,395	)		(114,351	)		(123,746	)		(19,584	)		(106,782	)		(126,366	)

Operating income (loss) before tax		(4,931	)		(26,053	)		(30,984	)		(8,810	)		(58,492	)		(67,302	)		(18,837	)		(73,813	)		(92,650	)

Financial gain (loss)		259			8,119			8,378			87			(41	)		46			—			(11,033	)		(11,032	)

Net income (loss)		(4,672	)		(17,934	)		(22,606	)		(8,722	)		(58,533	)		(67,255	)		(18,837	)		(84,846	)		(103,683	)

Non controlling interests		—			(190	)		(190	)		—			—			—			4,315			—			4,315

Net income (loss) attributable to shareholders of Cellectis		(4,672	)		(18,124	)		(22,796	)		(8,722	)		(58,533	)		(67,255	)		(14,522	)		(84,846	)		(99,368	)

Adjustment of share-based compensation attributable to shareholders of Cellectis		789			32,613			33,402			1,098			57,524			58,622			5,957			42,968			48,925

Adjusted net income (loss) attributable to shareholders of Cellectis		(3,883	)		14,489			10,606			(7,625	)		(1,009	)		(8,633	)		(8,565	)		(41,877	)		(50,442	)

Depreciation and amortization		(99	)		(1,838	)		(1,937	)		(345	)		(1,866	)		(2,211	)		(551	)		(2,820	)		(3,371	)
Additions to tangible and intangible assets		526			3,886			4,413			10,410			4,164			14,573			792			1,849			2,642
Impairment of tangible assets		—			—			—			—			—			—			—			(798	)		(798	)


	For the year ended December 31, 2016									For the year ended December 31, 2017									For the year ended December 31, 2018
$ in thousands	Plants			Therapeutics			Total reportable segments			Plants			Therapeutics			Total reportable segments			Plants			Therapeutics			Total reportable segments
External revenues		399			44,409			44,808			508			24,680			25,188			236			12,495			12,731
External other income		186			11,450			11,637			239			8,290			8,528			178			8,523			8,701

External revenues and other income		585			55,859			56,444			747			32,969			33,715			414			21,018			21,432

Royalty expenses		(468	)		(1,309	)		(1,777	)		(390	)		(2,230	)		(2,620	)		(595	)		(2,144	)		(2,739	)
Research and development expenses		(4,112	)		(74,345	)		(78,458	)		(6,057	)		(73,170	)		(79,227	)		(8,638	)		(67,929	)		(76,567	)
Selling, general and administrative expenses		(4,809	)		(38,603	)		(43,413	)		(13,143	)		(31,607	)		(44,750	)		(21,067	)		(26,180	)		(47,248	)
Other operating income and expenses		(6	)		(93	)		(99	)		6			225			232			(50	)		81			31

Total operating expenses		(9,395	)		(114,351	)		(123,746	)		(19,584	)		(106,782	)		(126,366	)		(30,351	)		(96,172	)		(126,523	)

Operating income (loss) before tax		(8,810	)		(58,492	)		(67,302	)		(18,837	)		(73,813	)		(92,650	)		(29,937	)		(75,154	)		(105,091	)

Financial gain (loss)		87			(41	)		46			—			(11,032	)		(11,032	)		1,420			15,339			16,758

Net income (loss)		(8,722	)		(58,533	)		(67,255	)		(18,837	)		(84,846	)		(103,683	)		(28,517	)		(59,816	)		(88,333	)

Net income (loss) from discontinued operations		—			—			—			—			—			—			—			—			—
Non controlling interests		—			—			—			4,315			—			4,315			9,640			—			9,640

Net income (loss) attributable to shareholders of Cellectis		(8,722	)		(58,533	)		(67,255	)		(14,522	)		(84,846	)		(99,368	)		(18,877	)		(59,816	)		(78,693	)

R&Dnon-cash stock-based expense attributable to shareholder of Cellectis		477			32,731			33,208			967			22,623			23,590			838			16,852			17,689
SG&Anon-cash stock-based expense attributable to shareholder of Cellectis		621			24,793			25,414			4,990			20,345			25,335			5,218			11,655			16,873

Adjustment of share-based compensation attributable to shareholders of Cellectis		1,098			57,524			58,622			5,957			42,967			48,924			6,056			28,507			34,563

Adjusted net income (loss) attributable to shareholders of Cellectis		(7,625	)		(1,009	)		(8,633	)		(8,565	)		(41,877	)		(50,443	)		(12,821	)		(31,309	)		(44,130	)

Depreciation and amortization		(345	)		(1,866	)		(2,211	)		(551	)		(2,820	)		(3,371	)		(637	)		(1,740	)		(2,377	)
Additions to tangible and intangible assets		10,410			4,164			14,573			792			1,849			2,642			1,871			3,040			4,911
Impairment of tangible assets		—			—			—			—			(798	)		(798	)		—			—			—


$ in thousands	For the full year ended December 31, 2018
$ in thousands	Cellectis Consolidated financial statements Reportable segments note (IFRS)			Non-cash stock-based compensation booked in IFRS (1)		Non-cash stock-based compensation in US GAAP (1)			Intersegment transactions (2)			Reclassifications (3)			Other (4)			Calyxt Stand alone financial statements (US GAAP)
External revenues and other income		414			—		—			—			(177	)		—			236

Research and development expenses		(8,638	)		1,205		(630	)		—			(1,783	)		—			(9,846	)
Selling, general and administrative expenses		(21,067	)		7,506		(3,756	)		(3,090	)		1,245			657			(18,505	)
Royalties and other operating income and expenses		(645	)		—		—			(71	)		716			—			—

Total operating expenses		(30,350	)		8,711		(4,386	)		(3,161	)		177			657			(28,351	)

Operating income (loss) before tax		(29,937	)		8,711		(4,386	)		(3,161	)		—			657			(28,115	)

Financial gain (loss)		1,420			—		—			51			—			(1,244	)		218

Net income (loss)		(28,517	)		8,711		(4,386	)		(3,110	)		—			(587	)		(27,897	)


	Software and Patents			Assets under construction			Total
Net book value as of January 1, 2015		1,246			—			1,246
$ in thousands										Software and Patents			Assets under construction			Total
Net book value as of January 1, 2016											1,041			—			1,041

Change in scope		—			—			—			—			—			—
Additions to intangible assets		97			—			97
Depreciation expense		(174	)		—			(174	)
Translation adjustments		(127	)		—			(127	)

Net book value as of December 31, 2015		1,041			—			1,041

Gross value at end of period		2,194			—			2,194
Accumulated depreciation and impairment at end of period		(1,153	)		—			(1,153	)
Net book value as of January 1, 2016		1,041			—			1,041

Additions to intangible assets		212			439			652			212			439			652
Disposal of intangible assets		(74	)		—			(74	)		(74	)		—			(74	)
Depreciation expense		(226	)		—			(226	)		(226	)		—			(226	)
Translation adjustments		(28	)		(21	)		(49	)		(28	)		(21	)		(49	)

Net book value as of December 31, 2016		924			419			1,343			924			419			1,343

Gross value at end of period		2,256			419			2,675			2,256			419			2,675
Accumulated depreciation and impairment at end of period		(1,332	)		—			(1,332	)		(1,332	)		—			(1,332	)
Net book value as of January 1, 2017		924			419			1,343			924			419			1,343

Additions to intangible assets		6			135			141			6			135			141
Depreciation expense		(231	)		—			(231	)		(231	)		—			(231	)
Translation adjustments		112			66			178			112			66			178

Net book value as of December 31, 2017		811			619			1,431			811			619			1,431

Gross value at end of period		2,571			517			3,190			2,571			517			3,190
Accumulated depreciation and impairment at end of period		(1,759	)		—			(1,759	)		(1,759	)		—			(1,759	)
Net book value as of January 1, 2018											811			619			1,431

Additions to intangible assets											14			103			117
Disposal of intangible assets											(7	)		—			(7	)
Reclassification											6			—			6
Depreciation expense											(217	)		—			(217	)
Translation adjustments											(30	)		(31	)		(61	)

Net book value as of December 31, 2018											577			691			1,268

Gross value at end of period											2,454			691			3,146
Accumulated depreciation and impairment at end of period											(1,878	)		—			(1,878	)


~~• Buildings and other outside improvements~~		~~10-20~~ ~~years~~
~~• Leasehold improvements~~		~~5-10~~ ~~years~~
~~• Office furniture~~		~~10 years~~
~~• Laboratory equipment~~		~~3-10~~ ~~years~~
~~• Office equipment~~		~~5 years~~
~~• IT equipment~~		~~3 years~~


	Lands and Buildings			Technical equipment			Fixtures, fittings and other equipment			Assets under construction			Total
	$ in thousands
Net book value as of January 1, 2015		1,416			1,703			50			—			3,169

Additions to tangible assets		1,477			2,488			331			186			4,481
Disposal of tangible assets		—			118			—			—			118
Depreciation expense		(681	)		(1,023	)		(58	)		—			(1,761	)
Reclassification		—			(19	)		20			—			1
Translation adjustments		(139	)		(370	)		(3	)		(4	)		(517	)

Net book value as of December 31, 2015		2,072			2,897			340			182			5,490


	Lands and Buildings			Technical equipment			Fixtures, fittings and other equipment			Assets under construction			Total
	$ in thousands
Net book value as of January 1, 2016		2,072			2,897			340			182			5,490

Additions to tangible assets		11,164			1,076			562			902			13,704
Disposal of tangible assets		—			(3	)		(1	)		(183	)		(186	)
Depreciation expense		(741	)		(1,077	)		(167	)		—			(1,986	)
Reclassification		—			3			(3	)		—			—
Translation adjustments		(59	)		(38	)		(23	)		(4	)		(122	)

Net book value as of December 31, 2016		12,436			2,858			707			898			16,900

Gross value at end of period		15,085			10,634			1,104			898			27,721
Accumulated depreciation and impairment at end of period		(2,649	)		(7,775	)		(397	)		—			(10,821	)

Net book value as of January 1, 2017		12,436			2,858			707			898			16,900

Additions to tangible assets		718			701			203			878			2,501
Disposal of tangible assets		(9,243	)		(103	)		2			(109	)		(9,453	)
Reclassification		14			47			18			(79	)		—
Depreciation expense		(972	)		(1,126	)		(245	)		(798	)		(3,140	)
Translation adjustments		206			127			68			18			418

Net book value as of December 31, 2017		3,159			2,505			753			809			7,226

Gross value at end of period		6,936			12,114			1,447			1,606			22,103
Accumulated depreciation and impairment at end of period		(3,777	)		(9,609	)		(693	)		(798	)		(14,877	)

Net book value as of January 1, 2018		3,159			2,505			753			809			7,226

Additions to tangible assets		879			1,622			1,820			1,942			6,263
Disposal of tangible assets		—			(49	)		(690	)		(426	)		(1,164	)
Reclassification		39			216			793			(1,053	)		(6	)
Depreciation expense		(758	)		(854	)		(478	)		—			(2,091	)
Translation adjustments		(90	)		(46	)		(27	)		(25	)		(188	)

Net book value as of December 31, 2018		3,229			3,393			2,172			1,247			10,041

Gross value at end of period		7,604			13,297			3,215			2,045			26,160
Accumulated depreciation and impairment at end of period		(4,375	)		(9,903	)		(1,043	)		(798	)		(16,119	)


	Accounting category				Book value on the statement of financial position		Fair Value
2017	Fair value through profit and loss		Amortized cost				Fair Value
	$ in thousands
Financial assets
Non-current financial assets		—		1,004		1,004		1,004
Trade receivables		—		2,753		2,753		2,753
Subsidies receivables		—		9,524		9,524		9,524
Current financial assets		40,602		—		40,602		40,602
Cash and cash equivalents		256,380		—		256,380		256,380

Total financial assets		296,982		13,281		310,263		310,263

Financial liabilities
Non-current financial liabilities		—		13		13		13
Current financial liabilities		—		21		21		21
Trade payables		—		9,460		9,460		9,460
Other current liabilities		—		6,570		6,570		6,570

Total financial liabilities		—		16,064		16,064		16,064


	Accounting category				Book value on the statement of financial position		Fair Value
2018	Fair value through profit and loss		Amortized cost				Fair Value
	$ in thousands
Financial assets
Non-current financial assets		—		1,891		1,891		1,891
Trade receivables		—		2,971		2,971		2,971
Subsidies receivables		—		17,173		17,173		17,173
Current financial assets		7		381		388		388
Cash and cash equivalents		451,501		—		451,501		451,501

Total financial assets		451,508		22,416		473,924		473,924

Financial liabilities
Non-current financial liabilities		—		1,018		1,018		1,018
Current financial liabilities		—		333		333		333
Trade payables		—		15,883		15,883		15,883
Other current liabilities		—		8,369		8,369		8,369

Total financial liabilities		—		25,603		25,603		25,603


	Accounting category				Book value on the statement of financial position		Fair Value
2016	Fair value through profit and loss		Amortized cost		Book value on the statement of financial position		Fair Value
	$ in thousands
Financial assets
Non-current financial assets		—		691		691		691
Trade receivables		—		3,627		3,627		3,627
Subsidies receivables		—		8,723		8,723		8,723
Current financial assets		36,592		—		36,592		36,592
Cash and cash equivalents		254,568		—		254,568		254,568

Total financial assets		291,159		13,042		304,201		304,201

Financial liabilities
Non-current financial liabilities		—		30		30		30
Current financial liabilities		1,692		38		1,730		1,730
Trade payables		—		9,722		9,722		9,722
Other current liabilities		—		5,196		5,196		5,196

Total financial liabilities		1,692		14,986		16,678		16,678


	Accounting category				Book value on the statement of financial position		Fair Value
2017	Fair value through profit and loss		Amortized cost		Book value on the statement of financial position		Fair Value
	$ in thousands
Financial assets
Non-current financial assets		—		1,004		1,004		1,004
Trade receivables		—		2,753		2,753		2,753
Subsidies receivables		—		9,524		9,524		9,524
Current financial assets		40,602		—		40,602		40,602
Cash and cash equivalents		256,380		—		256,380		256,380

Total financial assets		296,982		13,281		310,263		310,263

Financial liabilities
Non-current financial liabilities		—		13		13		13
Current financial liabilities		—		21		21		21
Trade payables		—		9,460		9,460		9,460
Other current liabilities		—		6,570		6,570		6,570

Total financial liabilities		—		16,064		16,064		16,064


2016	Notional		Fair Value			Maturity
	$ in thousands
USD forward sale contracts		44,914		(1,776	)		2017 to 2018
USD forward purchase contracts		—		—			—

Total derivative financial instruments				(1,776	)

of which :
Derivative financial assets				—
Derivative financial liabilities				(1,776	)


As of December 31, 2016	Carrying amount		Unrealized Gains/(Losses)		Estimated fair value
	$ in thousands
Current financial assets		36,592		—		36,592
Cash and cash equivalents		254,567		—		254,567

Current financial assets and cash and cash equivalents		291,159		—		291,159


As of December 31, 2017	Carrying amount		Unrealized Gains/(Losses)		Estimated fair value
	$ in thousands
Current financial assets		40,602		—		40,602
Cash and cash equivalents		256,380		—		256,380

Current financial assets and cash and cash equivalents		296,982		—		296,982


As of December 31, 2017	Carrying amount		Unrealized Gains/(Losses)		Estimated fair value
			$ in thousands
Current financial assets		40,602		—		40,602
Cash and cash equivalents		256,380		—		256,380

Current financial assets and cash and cash equivalents		296,982		—		296,982


As of December 31, 2018	Carrying amount		Unrealized Gains/(Losses)		Estimated fair value
			$ in thousands
Current financial assets		388		—		388
Cash and cash equivalents		451,501		—		451,501

Current financial assets and cash and cash equivalents		451,889		—		451,889


Balance as of December 31, 2017	Gross Amount		Less than One Year		One to Five Years		More than Five Years
Balance as of December 31, 2018									Gross Amount		Less than One Year		One to Five Years		More than Five Years
	$ in thousands								$ in thousands
Finance leases		34		21		13		—		1,350		333		1,018		—
Derivative instruments		—		—		—		—		—		—		—		—

Financial liabilities		34		21		13		—		1,350		333		1,018		—

Trade payables		9,460		9,460		—		—		15,883		15,883		—		—
Other current liabilities		6,570		6,570		—				8,369		8,369		—

Total financial liabilities		16,064		16,051		13		—		25,603		24,585		1,018		—


	As of December 31,		As of December 31,		As of December 31,		As of December 31,
	2016		2017		2017		2018
	$ in thousands				$ in thousands
VAT Payables		192		9		9		291
Accruals for personnel related expenses		4,140		5,982		5,982		7,041
Other		864		579		579		1,037

Total		5,196		6,570		6,570		8,369


	As of December 31, 2017 as restated (*)		As of December 31, 2018
	$ in thousands
Deferred revenues and contract liabilities		27,975		20,454
Other		—		299

Total Deferred revenue and contract liabilities		27,975		20,754


Nature of the Transactions	Share Capital		Share premium			Number of shares		Nominal value
	$ in thousands							in $
Balance as of January 1, 2015		2,014		263,100			29,446,721		0.05
Capital increase by issuance of common shares (IPO Nasdaq)		297		209,899			5,500,000		—
Capital increase by issuance of ordinary shares (BSA, BSPCE, SO and free shares)		13		4,325			231,893		—
Non-cash stock based compensation expense		—		32,614			—		—

Balance as of December 31, 2015		2,323		509,938			35,178,614		0.05

Capital increase by issuance of ordinary shares (BSA, BSPCE, and free shares)		9		723			156,446		—
Non-cash stock based compensation expense		—		57,524			—		—

Balance as of December 31, 2016		2,332		568,185			35,335,060		0.05

Capital increase by issuance of ordinary shares (BSA, BSPCE, SO and free shares)		35		2,921			625,002		—
Share based compensation		—		42,968			—		—
Other movements		—		(37	)		—		—

Balance as of December 31, 2017		2,367		614,037			35,960,062		0.05


Nature of the Transactions	Share Capital		Share premium			Number of shares		Nominal value
	$ in thousands							in $
Balance as of January 1, 2016		2,323		509,938			35,178,614		0.05

Capital increase by issuance of ordinary shares (BSA, BSPCE and free shares)		9		723			156,446		—

Non-cash stock based compensation expense		—		57,524			—		—

Balance as of December 31, 2016		2,332		568,185			35,335,060		0.05

Capital Increase		26					466,950
Exercise of share warrants, employee warrants and stock options		9		2,921			158,052
Non-cash stock based compensation expense				42,968
Other movements				(37	)

Balance as of December 31, 2017		2,367		614,037			35,960,062		0.05

Capital Increase		379		178,230			6,146,000
Exercise of share warrants, employee warrants and stock options		19		7,751			324,007
Non-cash stock based compensation expense				28,507

Balance as of December 31, 2018		2,765		828,525			42,430,069		0.05

Large accelerated filer

Accelerated filer

Non-accelerated filer

Emerging Growth Company