SYROS PHARMACEUTICALS, INC.

620 Memorial35 CambridgePark Drive, Suite 300

4th Floor, Cambridge, Massachusetts 0213902140

(617) 744-1340

NOTICE OF 20172022 ANNUAL MEETING OF STOCKHOLDERS

To Be Held on June 8, 2017

1, 2022

Dear Stockholders:

You are cordially invited to attend the 20172022 annual meeting of stockholders of Syros Pharmaceuticals, Inc. toThe meeting will be held virtually via the Internet at www.meetnow.global/M7TXWW9 on Thursday,Wednesday, June 8, 20171, 2022 at 1:11:00 p.m.a.m., Eastern Daylight Time, at our corporate offices located at 620 Memorial Drive, Suite 300, Cambridge, Massachusetts 02139. Time.

At the annual meeting, stockholders will consider and vote on the following matters:

As noted above, our annual meeting will be a “virtual meeting” of stockholders, which will be conducted exclusively via the Internet. There will not be a physical meeting location, and stockholders will not be able to attend the annual meeting in person. This means that you can attend the annual meeting online, vote your shares during the online meeting and submit questions for consideration prior to and during the online meeting, all by following the instructions described in the accompanying proxy statement. Stockholders of record at the close of business on April 13, 20174, 2022 will be entitled to notice of and to vote at the annual meeting or any adjournment or postponement thereof. We believe that hosting a “virtual meeting” will enable greater stockholder attendance and participation from any location around the world.

This Notice, the accompanying proxy statement and a form of proxy card are being mailed beginning on or about April 21, 2022 to all stockholders entitled to vote at the annual meeting. We have elected to provide access to our proxy materials over the Internet under the Securities and Exchange Commission’s “notice and access” rules. We believe that providing our proxy materials over the Internet expedites stockholders’ receipt of proxy materials, lowers costs and reduces the environmental impact of our annual meeting.

We encourage all stockholders to attend the annual meeting in person.online. However, whether or not you plan to attend the annual meeting in person,online, we encourage you to read this proxy statement and submit your proxy or voting instructions as soon as possible. Please review the instructions on each of your voting options described in the proxy statement.

Thank you for your ongoing support and continued interest in Syros.

By Order of the Board of Directors,

Nancy A. Simonian, M.D.

President and Chief Executive Officer

Cambridge, Massachusetts

April 25, 201721, 2022

Important Notice Regarding Internet Availability of Proxy Materials: The attached proxy statement and our 20162021 annual report to stockholders, which includes our Annual Report on Form 10-K for the fiscal year ended December 31, 2016,2021, are available atwww.envisionreports.com/SYRS. These documents are also available to any stockholder who wishes to receive a paper copy by calling (866) 641-4276, by emailing investorvote@computershare.com with “Proxy Materials Syros Pharmaceuticals, Inc.” in the subject line, or by submitting a request over the Internet at www.envisionreports.com/SYRS.

TABLE OF CONTENTS

620 Memorial35 CambridgePark Drive, Suite 300

4th Floor, Cambridge, Massachusetts 0213902140

(617) 744-1340

PROXY STATEMENT

20172022 ANNUAL MEETING OF STOCKHOLDERS

To Be Held on June 8, 20171, 2022

INFORMATION CONCERNING SOLICITATION AND VOTING

This proxy statement and the enclosed proxy card are being furnished in connection with the solicitation of proxies by the board of directors of Syros Pharmaceuticals, Inc. for use at the annual meeting of stockholders to be held on Thursday,Wednesday, June 8, 20171, 2022 at 1:11:00 p.m.a.m., Eastern Daylight Time, at our corporate offices located at 620 Memorial Drive, Suite 300, Cambridge, Massachusetts 02139, and at any adjournment thereof. The meeting will be held virtually via the Internet at www.meetnow.global/M7TXWW9. You will be able to attend and participate in the annual meeting online, vote your shares electronically and submit your questions prior to and during the meeting, all by following the instructions described in this proxy statement in the section entitled “Important Information About the Annual Meeting and Voting.” There will not be a physical meeting location, and stockholders will not be able to attend the annual meeting in person. Except where the context otherwise requires, references to “Syros Pharmaceuticals,” “the Company,” “we,” “us,” “our” and similar terms refer to Syros Pharmaceuticals, Inc.

This proxy statement summarizes information about the proposals to be considered at the meeting and other information you may find useful in determining how to vote. The proxy card is the means by whichhow you actually authorize another person to vote your shares in accordance with your instructions. We are making this proxy statement, the related proxy card and our annual report to stockholders for the fiscal year ended December 31, 20162021 available to stockholders for the first time on or about April 25, 2017.21, 2022.

A copy of our Annual Report on Form 10-K for the fiscal year ended December 31, 2016,2021, as filed with the Securities and Exchange Commission, or SEC, except for exhibits, will be furnished without charge to any stockholder upon written or oral request to Syros Pharmaceuticals, Inc., 620 Memorial35 CambridgePark Drive, Suite 300,4th Floor, Cambridge, Massachusetts 0213902140 or by calling (866) 641-4276, by emailing investorvote@computershare.com with “Proxy Materials Syros Pharmaceuticals, Inc.” in the subject line, or by submitting a request over the Internet at www.envisionreports.com/SYRS. This proxy statement and our Annual Report on Form 10-K for the fiscal year ended December 31, 20162021 are also available on the SEC’s website atwww.sec.gov.

IMPORTANT INFORMATION ABOUT THE ANNUAL MEETING AND VOTING

A.          Our board of directors has made these materials available to you on the Internet in connection with the solicitation of proxies for use at our 2017 annual meeting of stockholders to be held at our corporate offices located at 620 Memorial Drive, Suite 300, Cambridge, Massachusetts 02139 on Thursday, June 8, 2017 at 1:00 p.m.

A.

Our board of directors has made these materials available to you on the Internet in connection with the solicitation of proxies for use at our 2022 annual meeting of stockholders to be held virtually at www.meetnow.global/M7TXWW9 on Wednesday, June 1, 2022 at 11:00 a.m., Eastern Daylight Time. As a holder of common stock, you are invited to attend the annual meeting online and are requested to vote on the items of business described in this proxy statement. This proxy statement includes information that we are required to provide to you under Securities and Exchange Commission, or SEC rules and that is designed to assist you in voting your shares.

If your shares are held in “street name,” meaning they are held for your account by an intermediary, such as a broker, then you are deemed to be the beneficial owner of your shares and the broker that actually holds the shares for you is the record holder and is required to vote the shares it holds on your behalf according to your instructions. The proxy materials, as well as voting and revocation instructions, should have been forwarded to you by the broker that holds your shares. In order toTo vote your shares, you will need to follow the instructions that your broker provides you. Many brokers solicit voting instructions over the Internet or by telephone.

If you do not give instructions to your broker, your broker will still be able to vote your shares with respect to certain “discretionary” items. The ratification of the appointment of Ernst & Young LLP as our independent registered public accounting firm (Proposal 2) is considered a discretionary item. Accordingly, your broker may vote your shares in its discretion with respect to that matter even if you do not give voting instructions on Proposal 2.

However, under applicable stock exchange rules that regulate voting by registered brokerage firms, the election of our nominees to serve as Class IIII directors (Proposal 1) is, the advisory vote on the compensation paid to our named executive officers (Proposal 3), and the advisory vote to determine the frequency of future advisory votes on the compensation paid to our named executive officers (Proposal 4) are not considered to be a discretionary item.items. Accordingly, if you do not give your broker voting instructions on Proposal 1, Proposal 3 or Proposal 4, your broker may not vote your shares with respect to this matterthese matters and

Q.Can I change my vote?

A.	If your shares are registered directly in your name, you may revoke your proxy and change your vote at any time before the vote is taken at the annual meeting. To do so, you must do one of the following:

A.          If your shares are registered directly in your name, you may revoke your proxy and change your vote at any time before the vote is taken at the annual meeting. To do so, you must do one of the following:

If your shares are held in “street name,” you may submit new voting instructions by contacting your broker or other nominee. You may also vote in person atonline during the annual meeting if you obtain a legal proxy and register in advance to attend the meeting, as described inabove under “How do I register to attend the answer above.virtual annual meeting?”.

A.          A majority of our shares of common stock outstanding at the record date must be present in person or represented by proxy to hold the annual meeting. This is called a quorum. For purposes of determining whether a quorum exists, we count as present any shares that are voted over the Internet, by telephone, by completing and submitting a proxy card by mail or that are represented in person

A.

A majority of our shares of common stock outstanding at the record date must be present virtually or represented by proxy to hold the annual meeting. This is called a quorum. For purposes of determining whether a quorum exists, we count as present any shares that are voted over the Internet, by telephone, by completing and submitting a proxy card by mail or that are represented virtually at the meeting. Further, for purposes of establishing a quorum, we will count as present shares that a stockholder holds even if the stockholder votes to abstain or only votes on one of the proposals. In addition, we will count as present shares held in “street name” by brokers who indicate on their proxies that they do not have authority to vote those shares. If a quorum is not present, we expect to adjourn the annual meeting until we obtain a quorum.

A.          Proposal 1—Election of Class I

A.	Proposal 1—Election of Class III Directors

A nominee will be elected as a director at the annual meeting if the nominee receives a plurality of the votes cast “for” the applicable seat on the board of directors.

Proposal 2—Ratification of the Appointment of Independent Registered Public Accounting Firm

The affirmative vote of the holders of shares of common stock representing a majority of the votes cast on the matter is required for the ratification of the appointment of Ernst & Young LLP as our independent registered public accounting firm for the year ended December 31, 2017. 2022.

Q.How does the board of directors recommend that I vote on the proposals?

A.          Our board of directors recommends that you vote:

FOR the election of the three nominees to serve as Class IIII directors, each for a three yearthree-year term; and

FOR the ratification of the appointment of Ernst & Young LLP as our independent registered public accounting firm for the fiscal year ending December 31, 2017.2022;

Q.Are there other matters to be voted on atFOR the annual meeting?

A.          We do not know of any matters that may come before the annual meeting other than the election of our Class I directors and the ratification of the appointment of Ernst & Young LLP as our independent registered public accounting firm. If any other matters are properly presented at the annual meeting, the persons named in the accompanying proxy intend to vote, or otherwise act, in accordance with their judgment on the matter.

Q.Where can I find the voting results?

A.          We plan to announce preliminary voting results at the annual meeting and will report final voting results in a Current Report on Form 8-K filed with the SEC within four business days following the date of our annual meeting.

Q.What are the costs of soliciting these proxies?

A.          We will bear the cost of soliciting proxies. In addition to solicitation by mail, our directors, officers and employees may solicit proxies by telephone, e-mail, facsimile, and in person without additional compensation. We may reimburse brokers or persons holding stock in their names, or in the names of their nominees, for their expenses in sending proxies and proxy material to beneficial owners.

Implications of Being an “Emerging Growth Company”

We are an “emerging growth company,” as defined in the Jumpstart Our Business Startups Act of 2012, or the JOBS Act, and may remain an emerging growth company for up to five years from the date of our initial public offering, or IPO. The JOBS Act contains provisions that, among other things, reduce certain reporting requirements for an “emerging growth company.” For so long as we remain an emerging growth company, we are permitted and plan to rely on exemptions from certain disclosure requirements that are applicable to other public companies that are not emerging growth companies. These exemptions include reduced disclosure obligations regarding executive compensation. In addition, as an emerging growth company, we are not required to conduct votes seeking approval, on an advisory(non-binding) basis, of the compensation of our named executive officers orofficers; and

FOR holding, on an advisory (non-binding) basis, an annual vote on the frequency with which such votes must be conducted. We may take advantage of some or all these exemptions until such time as we are no longer an emerging growth company. We would cease to be an emerging growth company if we have more than $1 billion in annual revenue, we have more than $700 million in market valuecompensation of our stock held by non-affiliates or we issue more than $1 billion of non-convertible debt over a three-year period. We have taken advantage of certain reduced reporting obligations in this proxy statement. Accordingly, the information contained herein may be different than the information you receive from other public companies in which you hold stock.named executive officers.

Class IIII Director Nominees

Amir Nashat, Ph.D.S. Gail Eckhardt, M.D. has served on our board of directors since January 2016. He is a managing partner at Polaris Partners, a venture capital firm, where he has worked since 2002. Dr. Nashat also serves on the board of the Partners Innovation Fund and the MIT Sandbox Innovation Fund Program. Dr. Nashat serves on the boards of directors of aTyr Pharma, Inc., Fate Therapeutics, Inc. and Selecta Biosciences, Inc., all biopharmaceutical companies.  He previously served on the boards of directors of Bind Therapeutics, Inc. and Receptos, Inc., each a biopharmaceutical company.  Dr. Nashat received a Ph.D. in chemical engineering from the Massachusetts Institute of Technology, and an M.S. and B.S. in materials science and mechanical engineering from the University of California, Berkeley. We believe Dr. Nashat is qualified to serve on our board of directors because of his experience on the boards of directors of other publicly-traded companies and his experience as an investor in biotechnology and life sciences companies.

Robert T. Nelsen has served on our board of directors since our inception in November 2011 and was our lead independent director from December 2015 to January 2017. Mr. Nelsen was a co-founder of ARCH Venture Partners, a venture capital firm, and has served in various capacities for ARCH and affiliated entities since 1986. He is currently a managing director of ARCH Venture Corporation. Mr. Nelsen is a director of Agios Pharmaceuticals, Inc. and Juno Therapeutics, Inc., each a biopharmaceutical company. Mr. Nelsen previously served as a trustee of the Fred Hutchinson Cancer Research Institute, a trustee of the Institute for Systems Biology, and a director of the National Venture Capital Association.  He has also served as a director of the following biopharmaceutical companies: Bellerophon Therapeutics, Inc., Fate Therapeutics, Inc., Kythera Biopharmaceuticals, Inc. and NeurogesX, Inc. Mr. Nelsen received a B.S. with majors in biology and economics from the University of Puget Sound and an M.B.A. from the University of Chicago. We believe Mr. Nelsen is qualified to serve on our board of directors due to his extensive experience as an investor in, and director of, early stage biopharmaceutical and life sciences companies. 

Peter Wirth has served as Chair of our board of directors since January 2017.  Mr. Wirth currently serves as Chairman of FORMA Therapeutics Holdings LLC, a privately-held small molecule drug discovery company, Executive Chairman of ZappRx, Inc., a privately-held digital health company, Senior Advisor to Zai Lab Limited, a privately-held biopharmaceutical company based in Shanghai, China, and director of Aura Biosciences, Inc., a privately-held biopharmaceutical company. From November 2011 to April 2014, Mr. Wirth served as President and director of Lysosomal Therapeutics, Inc., a privately-held biopharmaceutical company. Mr. Wirth was a senior executive at Genzyme Corporation from 1996 until after its acquisition by Sanofi-Aventis in 2011, most recently serving as Executive Vice President, Legal and Corporate Development, Chief Risk Officer and Corporate Secretary. During his time at Genzyme, Mr. Wirth had senior management responsibility for the company’s legal function, corporate development function, molecular oncology division, polymer drug discovery and development division and enterprise risk management function. Mr. Wirth previously served as a director of Synageva BioPharma Corp. a biopharmaceutical company. Mr. Wirth was a 2012 Advanced Leadership Fellow at Harvard University. He received his B.A. from the University of Wisconsin-Madison and his J.D. from Harvard Law School.  We believe Mr. Wirth is qualified to serve on our board of directors due to his expertise in corporate governance and his experience in corporate strategy, product development and law in the biotechnology industry.

Class I Director Not Standing for Re-Election

Stéphane Bancel has served asbeen a member of our board of directors since November 2013. He has been president, founding chief executive officerSeptember 2020. Dr. Eckhardt is a tenured Professor, inaugural Director of the Livestrong Cancer Institutes, Chair of the Department of Oncology, and a directorAssociate Dean of Moderna Therapeutics Inc., a privately-held biotechnology company, since July 2011. Mr. Bancel was previously chief executive officer of bioMérieux SA, a publicly-traded diagnostics company, from 2006 to July 2011. Prior to his timeCancer Programs at bioMérieux, Mr. Bancel was managing director of Eli Lilly in Belgium and executive director of global manufacturing strategy and supply chain at Eli Lilly in Indianapolis, Indiana. Mr. Bancel currently serves as a supervisory director of Qiagen N.V., a molecular diagnostics company. Mr. Bancel holds an M.E. from École Central Paris, an M.S. in chemical engineering from the University of Minnesota and an M.B.A. from Harvard Business School.

Class II Directors

Vicki L. Sato, Ph.D. has served on our board of directors since August 2013.Texas at Austin’s Dell Medical School, or UT Austin. She has been a professorfaculty member at the institution since January of management practice2017. Prior to joining UT Austin, Dr. Eckhardt was at Harvard Businessthe University of Colorado School since September 2006of Medicine from 1999 to January 2017, where she had numerous roles and was a professor inresponsibilities, including Division Head of Medical Oncology, Associate Director for Translational Research at the DepartmentUniversity of MolecularColorado Comprehensive Cancer Center and Cell Biology at Harvard University from July 2005 until October 2015. Previously, sheDirector of the Phase I Program and Fellowship. Dr. Eckhardt served as presidenton the Board of Vertex Pharmaceuticals,Directors of NuGenerex Immuno-Oncology, Inc., a publicly-tradedpublic biotechnology company, which she joined in 1992. Priorfrom March 2020 to becoming presidentMay 2021, and has served on the Board of Vertex, she was the chief scientific officer and senior vice presidentDirectors of research and development. Prior to joining Vertex, Dr. Sato served as vice president of research at BiogenOncoTex Inc., a publicly-tradedprivate biotechnology company. Dr. Satocompany, since January 2022. She has also served on numerous committees and study sections, including the ASCO Molecular Oncology Task Force, the ASCO Board of Directors, the FDA Oncology Drugs Advisory Committee, and the National Cancer Institute (NCI) Cancer Centers Study Section. She is a member of the NCI Investigational Drug Steering Committee and serves on several external advisory boards of directorsNCI-designated cancer centers, was a lead mentor in ASCO’s Leadership Development Program and currently is a member of the following public companies: Bristol Myers Squibb Company, PerkinElmer Corporation and BorgWarner, Inc.Board of Directors of the Association of American Cancer Institutes. Dr. Sato receivedEckhardt earned her A.B.B.S. degree in chemistry from Radcliffe CollegeStephen F. Austin State University and her A.M. and Ph.D. degreesM.D. from Harvard University.the University of Texas Medical Branch in Galveston. She conducted her postdoctoral workinternship and residency in Internal Medicine at boththe University of Virginia Medical School, followed by a post-doctoral research fellowship in Experimental and Molecular Medicine at Scripps Research Institute in La Jolla, California, and a fellowship in Medical Oncology at the University of California Berkeley and Stanford Medical Center.San Diego. We believe that Dr. SatoEckhardt is qualified to serve on our board of directors because of her experience as a senior executive and as a director of several life sciences companies, and because of her knowledge of our industry.

Phillip A. Sharp, Ph.D. has served on our board of directors since December 2012. Dr. Sharp has been an institute professor at the Massachusetts Institute of Technology since 1999. Much of Dr. Sharp’s scientific work has been conducted at MIT’s Center for Cancer Research (now the Koch Institute), which he joined in 1974 and directed from 1985 to 1991. He subsequently led the Department of Biology from 1991 to 1999 before assuming the directorship of the McGovern Institute from 2000 to 2004. Dr. Sharp is the winner of the 1993 Nobel Prize in Physiology or Medicine. Dr. Sharp has served on the board of directors of Alnylam Pharmaceuticals, Inc., a biopharmaceutical company, since 2002. He earned his B.A. from Union College (Kentucky) in 1966 and a Ph.D. in chemistry from the University of Illinois, Champaign Urbana in 1969. He did his postdoctoral training at the California Institute of Technology. We believe Dr. Sharp is qualified to serveservice on our board of directors due to his scientificher expertise in the preclinical and his experience as a directorearly clinical development of a publicly traded company.novel agents and her extensive drug development experience.

Richard A. Young, Ph.D. has served on our board of directors since our inception in November 2011. He is also one of our scientific co-founders and a member of our scientific advisory board. He has been a member of the Whitehead Institute and professor of Biology at the Massachusetts Institute of Technology since 1984. In May 2012, he was elected into the National Academy of Sciences. Dr. Young has served as an advisor to Science magazine and the World Health Organization. Dr. Young previously served as a director of Enzon Pharmaceuticals, Inc., a biopharmaceutical company. Dr. Young received his Ph.D. in molecular biophysics and biochemistry from Yale University. We believe Dr. Young is qualified to serve on our board of directors because of his scientific expertise and his role as one of our scientific co-founders.

Class III Directors

Marsha H. Fanucci has been a member of our board of directors since October 2015. Since 2009, Ms. Fanucci has been an independent consultant. From 2004 throughto 2009, she served as senior vice president and chief financial officer of Millennium Pharmaceuticals, Inc., a biopharmaceutical company, or Millennium, that was subsequently acquired by Takeda Pharmaceuticals Company, a publicly traded biopharmaceutical company,

Genzyme Therapeutics’ Lysosomal Storage Disorder franchise. Previously, Mr. Patel held several cross functional senior leadership roles at Genzyme, including vice president, clinical research and head of the Global Clinical Research Operations Council. Prior to Genzyme, Mr. Patel held roles in clinical research and commercial development with increasing levels of responsibility at Burroughs Wellcome, a private foundation; Hoechst Marion Roussel, a privately-held life sciences company; and Fujisawa Pharmaceutical Co. Ltd., a privately-held pharmaceutical company. Mr. Patel is a member of the board of directors of BioCryst Pharmaceuticals, Inc., a biotechnology company. Mr. Patel obtained his B.Sc. with honors in biotechnology from the University of the South Bank, London. He completed his post graduate management and business degree, and doctorate level research program at Ealing College, London, and the Wellcome Foundation, respectively. We believe Mr. Patel is qualified to serve as a director based on his extensive healthcare industry leadership experience.

Nancy A. Simonian, M.D.has been our chief executive officer since July 2012. From 2001 to October 2011, Dr. Simonian was employed by Takeda Pharmaceuticals Company, a publicly-held biopharmaceutical company, and at Millennium Pharmaceuticals, Inc.Millennium. prior to its acquisition by Takeda, most recently serving as chief medical officer and senior vice president of clinical, medical and regulatory affairs. From 1995 to 2001, Dr. Simonian served at Biogen, Inc., a publicly-heldpublicly traded biotechnology company, and most recently served as vice president of clinical development. She is a member of the boardsboard of directors of Seattle Genetics, Inc., a biotechnologypublicly traded biopharmaceutical company, the Damon Runyon Cancer Research Foundation and the Biotechnology Industry Organization. She previously served as a member of the board of directors of Evelo Biosciences, Inc. from April 2018 to June 2021. Prior to joining the biopharmaceutical industry, Dr. Simonian was on the faculty of Massachusetts General Hospital and Harvard Medical School as an assistant professor of neurology. She received a B.A. in biology from Princeton University and an M.D. from the University of Pennsylvania School of Medicine. We believe Dr. Simonian is qualified to serve on our board because of her role as our chief executive officer, her experience in the biopharmaceutical industry and her other executive leadership and board of directors experience.

Class I Directors

Mark J. Alles has served on our board of directors since December 2019. Mr. Alles served as chief executive officer of Celgene Corporation, a global biopharmaceutical company, or Celgene, from March 2016 to June 2018 and as its chairman and chief executive officer from February 2018 until its acquisition by Bristol-Myers Squibb Company in November 2019. Prior to these roles, Mr. Alles served as Celgene’s president and chief operating officer from August 2014 to February 2016 and as its chief commercial officer and executive vice president, hematology & oncology from December 2012 to July 2014. Mr. Alles first joined Celgene in April 2004 and served in a number of commercial management positions of increasing responsibility at the company. Before joining Celgene, he held senior commercial management roles at Aventis Pharmaceuticals Inc. (Rhône-Poulenc Rorer) from 1993 to 2004. He is currently chairman of the board of directors of Turning Point Therapeutics, Inc., a public oncology company, and also serves on the board of directors of Antengene Corporation Limited and BioMarin Pharmaceuticals, both public biopharmaceutical companies. He is also a member of the board of directors and consulting CEO for PIKSci, Inc., a private biotechnology company. Mr. Alles received B.S. degree from Lock Haven University of Pennsylvania and served as a Captain in the United States Marine Corps. We believe that Mr. Alles is qualified for service on our board of directors due to his extensive executive experience and his track record of building successful global oncology organizations and commercializing innovative therapies.

Amir Nashat, Ph.D. has served on our board of directors since January 2016. He is a managing partner at Polaris Partners, a venture capital firm, where he has worked since 2002. Dr. Nashat also serves on the advisory board of the Partners Healthcare Innovation Fund. Dr. Nashat serves on the board of directors of Scholar Rock, Inc., a biopharmaceutical company. He previously served on the boards of directors of aTyr Pharma, Inc., Bind Therapeutics, Inc., Fate Therapeutics, Inc., Receptos, Inc. and Selecta Biosciences, Inc., each a biopharmaceutical company. Dr. Nashat received a Ph.D. in chemical engineering from MIT, and an M.S. and B.S. in materials science and mechanical engineering from the University of California, Berkeley. We believe Dr. Nashat is qualified to serve on our board of directors because of his experience on the boards of directors of other publicly traded companies and his experience as an investor in biotechnology and life sciences companies.

Board Leadership Structure

Our corporate governance guidelines provide that the nominating and corporate governance committee shall periodically assess the board of directors’ leadership structure, including whether the offices of chief executive officer and chair of the board of directors should be separate. Our guidelines provide the board of directors with flexibility to determine whether the two roles should be combined or separated based upon our needs and the board of directors’ assessment of its leadership from time to time. We currently separate the roles of chief executive officer and chair of the board of directors. OurDr. Simonian, as our president and chief executive officer, is responsible for setting the strategic direction for our company and the for providing day-to-day leadership and performance of our company, while theMr. Wirth, as chair of our board of directors, presides over meetings of the board of directors,

on the audit committee and the nominating and governance committee for part of 2016, and Stéphane Bancel and Vicki L. Sato, Ph.D., who served on the compensation committee for part of 2016, at the times of such service satisfied the applicable independence standards for such committees. In making such determinations, our board of directors considered the relationships that each such non-employee director has with our company and all other facts and circumstances our board of directors deemed relevant in determining independence, including the beneficial ownership of our capital stock by each non-employee director.

The current members of our audit committee are Mark J. Alles, Marsha H. Fanucci and Amir Nashat, Ph.D. and Sanj K. Patel. Ms. Fanucci chairs the audit committee. Our board of directors has determined that Ms. Fanucci qualifies as an “audit committee financial expert” within the meaning of applicable SEC rules. Richard A. Young served on the audit committee from January to September 2016. The audit committee held threefive meetings during fiscal 2016.2021.

Compensation Committee

Our compensation committee’s responsibilities include:

To the extent permitted by and consistent with applicable law and the provisions of a given equity-based plan, the compensation committee may delegate to one or more of our executive officers the power to grant options or other stock awards pursuant to an equity-based plan to our employees who are not directors or executive officers.

The current members of our compensation committee are Deborah Dunsire, M.D., Phillip A. Sharp, Ph.D., Robert T. Nelsen and Peter Wirth. Dr. SharpMr. Wirth chairs the compensation committee. Stéphane Bancel served on the compensation committee from January to September 2016, and Vicki L. Sato, Ph.D. served on this committee from January 2016 to January 2017. The compensation committee held eightseven meetings during fiscal 2016.2021.

The current members of our nominating and corporate governance committee are Vicki L. Sato,Srinivas Akkaraju, M.D., Ph.D., Stéphane BancelS. Gail Eckhardt, M.D., and Amir Nashat, Ph.D. Dr. SatoNashat chairs the nominating and corporate governance committee. Richard A. Young, Ph.D. served on the nominating and corporate governance committee from January to September 2016. The nominating and corporate governance committee held one meetingthree meetings during fiscal 2016.2021.

Research and Development Committee

Our research and development committee’s responsibilities include:

The current members of our research and development committee are Srinivas Akkaraju, M.D., Ph.D., Deborah Dunsire, M.D., S. Gail Eckhardt, M.D., Phillip A. Sharp, Ph.D., and Richard A. Young, Ph.D. Dr. Young chairs the research and development committee. The research and development committee held three meetings during fiscal 2021.

Director Nomination Process

The process followed by our nominating and corporate governance committee to identify and evaluate director candidates includes requests to board members and others for recommendations, meetings from time to time to evaluate biographical information and background material relating to potential candidates and interviews of selected candidates by members of the nominating and corporate governance committee and our board of directors.

Criteria and Diversity

In considering whether to recommend to our board of directors any particularspecific candidate for inclusion in our board of directors’ slate of recommended director nominees, including candidates recommended by stockholders, the nominating and corporate governance committee of our board of directors applies the criteria set forth in our corporate governance guidelines. These criteria include the candidate’s integrity, business acumen, knowledge of our business and industry, the ability to act in the interests of all stockholders, and lack of conflicts of interest.

candidates by following substantially the same process, and applying substantially the same criteria, as it follows for candidates submitted by others. If the board determines to nominate a stockholder-recommended candidate and recommends his or her election, then his or her name will be included in our proxy card for the next annual meeting.

Stockholders also have the right under our by-laws to directly nominate director candidates, without any action or recommendation on the part of the nominating and corporate governance committee or our board of directors, by following the procedures set forth under “Other Matters – Stockholder Proposals for our 20182023 Annual Meeting.”

Oversight of Risk

Our board of directors oversees our risk management processes directly and through its committees. Our management is responsible for risk management on a day-day basis. The role of our board of directors and its committees is to oversee the risk management activities of management. Our board of directors fulfills this duty by discussing with management the policies and practices utilized by management in assessing and managing risks and providing input on those policies and practices. In general, our board of directors oversees risk management activities relating to business strategy, acquisitions, capital allocation, organizational structure and certain operational risks; our audit committee oversees risk management activities related to financial controls, and legal and compliance risks, and cybersecurity risks; our compensation committee oversees risk management activities relating to our compensation policies and practices;practices and management succession planning; and our nominating and corporate governance committee oversees risk management activities relating to the composition of our board of directors and management succession planning.corporate governance. Each committee reports to the full board of directors on a regular basis, including reports with respect to the committee’s risk oversight activities as appropriate. In addition, since risk issues often overlap, committees from time to timetime-to-time request that the full board of directors discuss particular risks.

approve proposed related person transactions that arise between committee meetings, subject to ratification by the committee at its next meeting. Any related person transactions that are ongoing in nature will be reviewed annually.

A related person transaction reviewed under this policy will be considered approved or ratified if it is authorized by the audit committee in accordance with the standards set forth in the policy after full disclosure of the related person’s interests in the transaction. As appropriate for the circumstances, the policy provides that the audit committee will review and consider:

EXECUTIVE AND DIRECTOR COMPENSATION

Executive Officers

Certain information regarding our executive officers who are not also directors, as of April 1, 2017,March 31, 2022 is set forth below.

Kyle D. Kuvalanka Conley Cheehas served asbeen our chief operatingcommercial officer since September 2015.2021. He previously served as global head of portfolio management, global pipeline strategy and precision medicine at Novartis Oncology, or Novartis, a global pharmaceutical company from October 2018 to February 2021. In this role, Mr. Chee had strategic responsibility for shaping Novartis’ overall pipeline, including driving commercial planning for the company’s early-stage portfolio and diagnostics strategy for oncology. Earlier in his career at Novartis, Mr. Chee served as vice president – global head oncology lung franchise from 2015 to October 2018, as well as in multiple U.S. sales and marketing leadership roles. Prior to joining Syros,Novartis, Mr. KuvalankaChee spent five years in roles of increasing responsibility at Pfizer, Inc., a global pharmaceutical company, ultimately serving as team leader of international business development. He holds a B.Sc. Pharm from the University of Alberta and an M.B.A. from the Richard Ivey School of Business at the University of Western Ontario and completed his residency in Clinical Pharmacy at the University of British Columbia.

Jason Haas has been our chief financial officer since October 2021. He previously served as chief business officer and principal financial and accounting officerco-head of Blueprint Medicines Corporation,Americas, healthcare investment banking at Barclays from June 2016 to October 2021. Previously, he served as head of Americas, healthcare investment banking at Deutsche Bank from 2012 to June 2016. Prior to his role at Deutsche Bank, he was a biopharmaceutical company, from September 2013 to September 2015. From 2002 to September 2013,managing director on the healthcare investment banking team at Goldman Sachs & Co. Mr. Kuvalanka worked at Takeda Pharmaceutical Company and at Millennium Pharmaceuticals, Inc. prior to its acquisition by Takeda, including as vice president, corporate strategy, business development and alliance management from 2009 to September 2013. Mr. KuvalankaHaas holds a B.A. in International Relations and Economics from WesleyanColgate University and an M.B.A. in Finance from The WhartonColumbia Business School of the University of Pennsylvania.School.

Eric R. Olson, Ph.D. has been our chief scientific officer since April 2013. He previously served as research vice president for respiratory diseases at Vertex Pharmaceuticals, Inc., a biotechnology company, from 2001 to May 2013. Dr. Olson has also held positions as the director of antibacterials and molecular sciences departments at Warner-Lambert Co. as well as a research scientist focused on gene expression systems with The Upjohn Company, both of which were acquired by Pfizer Inc., a pharmaceutical company.company, or Pfizer. Dr. Olson serves on the boards of the Cystic Fibrosis Foundation and the National Brain Tumor Society. Dr. Olson received a B.S. in microbiology and immunology from the University of Minnesota and a Ph.D. in microbiology and immunology from the University of Michigan.

Gerald E. Quirk, Esq. has served as our chief legal officer since September 2016. He previously served as executive vice president, business operations and general counsel of Tokai Pharmaceuticals, Inc., a biotechnology company, from May 2015 to August 2016. Prior to that, Mr. Quirk was a partner and co-chair of the life sciences practice at Choate, Hall & Stewart LLP, a Boston law firm, from August 2012 until May 2015. Mr. Quirk previously served as vice president, corporate affairs and general counsel at Infinity Pharmaceuticals, Inc. with responsibility for the company’s legal, intellectual property, finance and corporate communications activities, from August 2006 until August 2012, and prior to that held legal and business development positions of increasing responsibility at Genzyme Corporation. Mr. Quirk holds a B.A. in political science from Swarthmore College, an Ed.M. in educational administration from Harvard University, and a J.D. from Northeastern University.

David A. Roth, M.D. has been our chief medical officer since December 2015. Previously, Dr. Roth was employed by Infinity Pharmaceuticals, Inc., a pharmaceutical company, from September 2013 until September 2015, serving most recently as its executive vice president and chief medical officer and previously as its senior vice president of clinical development and medical affairs. Prior to joining Infinity, Dr. Roth was the vice president, early development in the oncology business unit of Pfizer Inc., a pharmaceutical company, from 2009 to August 2013. Prior to joining the pharmaceutical industry, Dr. Roth’s experience included over 10ten years in research and clinical practice as an academic hematologist, and he served on the full-time faculty at Harvard Medical School and Beth Israel Deaconess Medical Center in Boston. Dr. Roth received his B.S. from the Massachusetts Institute of TechnologyMIT and his M.D. from Harvard Medical School in the Harvard MIT Division of Health Sciences and Technology, where he remains on the affiliated faculty.Technology.

executive officers, Kyle D. Kuvalanka,Jason Haas, our Chief Operating Officer, Gerald E. Quirk, Esq., our Chief LegalFinancial Officer, and David A. Roth, M.D., our Chief Medical Officer. In addition, this section provides qualitative information regarding the manner and context in which compensation is awarded to and earned by our named executive officers and is intended to place in perspective the data presented in the following tables and the corresponding narrative.

Summary Compensation Table

The following table sets forth information regarding compensation earned by our named executive officers during the years indicated.

cash bonuses to the compensation committee. The compensation committee conducts a performance evaluation of Dr. Simonian. The compensation committee consults with the board of directors as to the achievement of corporate objectives that drive contingent compensation awards.

During its annual compensation review, our compensation committee also consults with external advisors. In fiscal 2016,2021, the compensation committee engaged Compensia Inc. as its independent compensation consultant to provide comparative data on executive compensation practices in our industry and assess our executives’ compensation relative to comparable companies.

Base Salary

We use base salaries to recognize the experience, skills, knowledge and responsibilities required of all our employees, including our named executive officers. None of our named executive officers is currently party to an employment agreement or other agreement or arrangement that provides for automatic or scheduled increases in base salary.

In 2015, we paid aFebruary 2020, the compensation committee set Dr. Simonian’s annualized base salary of $425,000 to Dr. Simonian.$600,000 effective February 10, 2020. In January 2016,February 2021, the compensation committee increased Dr. Simonian’s annualized base salary wasto $618,000 effective February 22, 2021. In February 2022, the compensation committee increased to $455,000 and in September 2016, her annualDr. Simonian’s annualized base salary was further increased to $500,000 retroactive to the closing of our IPO in July 2016.  Dr. Simonian’s base salary increased to $516,000$640,000 effective February 16, 2017.

21, 2022.

Mr. KuvalankaHaas commenced employment with us in September 2015.October 2021. In 2015,2021, we paid him a base salary of $106,692$91,385 for his partial year of employment and hisbased on an annualized base salary was $380,000.  Dr. Roth commenced employment with us in December 2015.of $440,000. In 2015, we paid him a base salaryFebruary 2022, the compensation committee of $26,064 for his partial yearour board of employment, and hisdirectors increased Mr. Haas’s annualized base salary was $380,000. Base salaries for Mr. Kuvalanka and Dr. Roth did not increase in fiscal 2016, but were increased to $392,000 for both officers$470,000 effective February 16, 2017.  Mr. Quirk commenced employment with us in September 2016.  21, 2022.

In 2016, we paid him a base salary of $98,269 for his partial year of employment, and hisFebruary 2020, the compensation committee set Dr. Roth’s annualized base salary was $365,000.  Mr. Quirk’sto $470,000 effective February 10, 2020. In February 2021, the compensation committee increased Dr. Roth’s annualized base salary was not adjusted for 2017.to $484,000 effective February 22, 2021. In February 2022, the compensation committee of our board of directors increased Dr. Roth’s annualized base salary to $500,000 effective February 21, 2022.

Cash Incentives

During 2016, weWe have established a framework under which the compensation committee would, in its discretion, award annual performance-based cash bonuses to our executive officers for up to a specific percentage of his or her salary as a vehicle to reward achievement of value driving milestones and recognize individual performance. Dr. Simonian iswas eligible for a performance-based cash bonus of up to 50%a percentage of her annual base salary, subject to achievement of corporate goals as determined by the compensation committee. Her bonus target was 55% of her annual base salary in the year ended December 31, 2020, or fiscal 2020, and fiscal 2021. Our other named executive officers are eligible for a performance-based cash bonus of up to 35%a percentage of such named executive

In 2015, we madethe equity incentive awards that it grants in order to ensure the long-term sustainability of our equity incentive plan. As a result of the ongoing volatility in the equity markets for development-stage companies in our industry and the imperative to continue to motivate and retain our executive officers including Dr. Simonian, Mr. Kuvalanka and Dr. Roth.through the completion of the potential value-creating clinical milestones in the future, we expect that the number of shares subject to the equity incentive awards granted by the compensation committee during 2022 will increase.

In February 2020, Dr. Simonian was granted three optionsan option to purchase up to 100,368, up to 43,048 and up to 43,047294,000 shares of our common stock respectively. The first and second grantsDr. Roth was granted an option to purchase 94,000 shares of our common stock. These options vested as to 25% of the shares on February 5, 2016 and June 9, 2016, respectively,28, 2021, with the remaining shares vesting in equal monthly installments thereafter through February 5, 2019 and June 9, 2019, respectively. Dr. Simonian’s third grant vests upon29, 2024, subject to the achievement of performance‑based criteria, with 50% ofapplicable officer’s continued service through the shares having vested upon the closing of our IPO in July 2016 and the remaining 50% of the sharesapplicable vesting upon the earlier to occur of clinical proof‑of‑concept studies for a company drug candidate and a significant development or partnership deal, in each case as determined by our board of directors, and in any event will vest in full on June 9, 2021.

date.

In March 2016,February 2021, Dr. Simonian was granted an option to purchase up to 93,333304,000 shares of our common stock whichand Dr. Roth was granted an option to purchase 119,000 shares of our common stock. These options vested as to 25% of the shares on March 31, 2017,February 28, 2022, with the remaining shares vesting in equal monthly installments thereafter through March 31, 2020.  February 28, 2025, subject to the applicable officer’s continued service through the applicable vesting date.

In September 2016, Dr. SimonianOctober 2021, Mr. Haas was granted an option to purchase an additional 75,000750,000 shares of our common stock whichin connection with his appointment as our Chief Financial Officer as an inducement material to Mr. Haas’ acceptance of employment with the company in accordance with Nasdaq Listing Rule 5635(c)(4). This option will vestvests as to 25% of the shares subject to the option on September 16, 2017,October 12, 2022, with the remaining shares vesting in equal monthly installments thereafter through September 16, 2020.

In September 2015,October 12, 2025, subject to Mr. KuvalankaHaas’s continued service through each applicable vesting date. The size of this equity award was driven largely by the value of new-hire equity incentive awards granted two options to purchase upchief financial officers at comparable companies as well as the compensation committee’s desire to 189,252 and up to 68,819 shares of common stock, respectively. The first grant vested as to 25%make Mr. Haas whole of the shares on September 21, 2016, with the remaining shares vesting in equal monthly installments thereafter through September 21, 2019. Mr. Kuvalanka’s second grant vests upon the achievement of performance‑based criteria, with 40%value of the shares having vesteddeferred compensation he would be forfeiting upon the closing of our IPO in July 2016, 40% vesting upon the execution by us of a collaboration agreement and 20% of the shares vesting upon clinical proof‑of‑concept for a company drug candidate, in each case as determined by our board of directors, and in any event will vest in full on September 21, 2021. Mr. Kuvalanka was not granted any stock options during 2016.leaving his prior employer.

Outstanding Equity Awards at Fiscal Year End 2016

2021

The following table sets forth information regarding outstanding equity awards which consist entirely of stock options, held by our named executive officers as of December 31, 2016.2021.