DEF 14A of Voyager Therapeutics, Inc., Apr 28, 2026

DEF 14A of Voyager Therapeutics, Inc.

Filed Apr 28, 2026 · FY20250001104659-26-050297

Open on EDGAR

  1. ELECTION OF DIRECTORS

    ELECTION OF DIRECTORS Our Board of Directors is authorized to have, and currently consists of, ten members. The Board of Directors is divided into three classes, with one class of directors standing for election each year, for a term of three years. Directors for each class are elected at the annual meeting of stockholders held in the year in which the term for their class expires and hold office until their successors are duly elected and qualified or their earlier resignation or removal. In accordance with our Certificate of Incorporation and Bylaws, vacancies on our Board of Directors shall be filled solely and exclusively by the affirmative vote of a majority of the remaining directors, even if less than a quorum is present, and not by stockholders. The Board of Directors currently consists of the following: • Class II directors: James A. Geraghty, Steven Hyman, M.D., and Alfred Sandrock, M.D., Ph.D., whose terms expire at the Annual Meeting; • Class III directors: Michael Higgins, Jude Onyia, Ph.D., and Nancy Vitale, whose terms expire at the 2027 annual meeting of stockholders; and • Class I directors: Grace E. Colón, Ph.D., Catherine J. Mackey, Ph.D., Glenn Pierce, M.D., Ph.D., and George Scangos, Ph.D., whose terms expire at the at the 2028 annual meeting of stockholders. The Board of Directors, on the recommendation of our Nominating and Corporate Governance Committee, nominated James A. Geraghty, Steven Hyman, M.D., and Alfred Sandrock, M.D., Ph.D., for election by the stockholders as Class II directors at the upcoming Annual Meeting. If elected, each nominee will serve as a director until the 2029 annual meeting of stockholders and until such nominee’s successor is duly elected and qualified, or until such nominee’s earlier death, resignation, or removal. The names of and certain information about the directors in each of the three classes are set forth below. There are no family relationships among any of our directors or executive officers. If no contrary indication is made, proxies will be voted for James A. Geraghty, Steven Hyman, M.D., and Alfred Sandrock, M.D., Ph.D. or, in the event that any of these nominees is not a candidate or is unable to serve as a director at the time of election (which is not currently expected), for any substitute nominee who is designated by our Board of Directors. The proxy cannot be voted for a greater number of persons than the number of nominees named in this proxy statement.

  2. Nominees for Class II Directors

    Nominees for Class II Directors The name of the nominees for Class II directors and certain information about each nominee as of March 31, 2026, which has been furnished to us by each nominee, as applicable, are set forth below.

  3. Nominees for Class II Directors

    | Name | Positions and Offices Held with Voyager | Director Since | Age | | James A. Geraghty | Director | 2014 | 71 | | Steven Hyman, M.D. | Director | 2015 | 73 | | Alfred Sandrock, M.D., Ph.D. | Director, President and Chief Executive Officer | 2022 | 68 |

  4. Nominees for Class II Directors

    Set forth below are the biographies of each director, as well as a discussion of the particular experience, qualifications, attributes, and skills that led the Board of Directors to conclude that each person nominated to serve or currently serving on the Board of Directors should serve as a director. In addition to the information presented below, we believe that each director nominee meets the minimum qualifications established by the Nominating and Corporate Governance Committee. The process of the Board of Directors and its committees to identify and evaluate candidates for director nominees, as well as the applicable minimum qualifications for director nominees, is described below under the heading “Corporate Governance — Board and Committee Matters.”

  5. Director Nominees for Election for a Three-year Term Ending at the 2029 Annual Meeting of Stockholders

    Director Nominees for Election for a Three-year Term Ending at the 2029 Annual Meeting of Stockholders James A. Geraghty. Mr. Geraghty has been a member of the Board of Directors since January 2014. He has served as a member of the boards of directors of publicly-traded biotechnology companies Fulcrum Therapeutics since July 2016 and CanBridge Pharma since May 2020. Mr. Geraghty previously served as Chairman of the board of directors of privately-held biotechnology company OMass Therapeutics from May 2023 to January 2026, and of publicly-traded biotechnology companies Pieris Pharmaceuticals from May 2017 to December 2024, and Orchard Therapeutics from May 2020 to January 2024. He also served as a member of the board of directors of Aceragen Inc. (formerly Idera Pharmaceuticals) from May 2013 to March 2023. Mr. Geraghty was an Entrepreneur-in-Residence at Third Rock Ventures from May 2013 to October 2016. He served as Senior Vice President, North America Strategy and Business Development at Sanofi from February 2011 to October 2013, and held many roles at Genzyme Corporation from 1992 to 2011, including Senior Vice President of International Development and President of Genzyme Europe. He also served as Chairman, President and CEO of GTC Biotherapeutics. Mr. Geraghty received a B.A. from Georgetown University, an M.S. from the University of Pennsylvania, and a J.D. from Yale Law School. We believe Mr. Geraghty’s experience as a senior executive and service on the Boards of Directors of other life sciences companies qualifies him to serve on the Board of Directors. Steven Hyman, M.D. Dr. Hyman has been a member of the Board of Directors since September 2015. He has served as founding director of the Program in Brain Health at the Broad Institute of Harvard and MIT and as a Core Faculty Member of the Broad Institute since March 2012. Dr. Hyman has served as a Harvard University Distinguished Service Professor of Stem Cell and Regenerative Biology since July 2011. He has served as a member of the board of directors of publicly-traded biotechnology company Cyclerion Therapeutics, Inc. since July 2022. From December 2001 to June 2011, he served as Provost of Harvard University, the University’s chief academic officer. From 1996 to 2001, he served as Director of the U.S. National Institute of Mental Health, where he emphasized investment in neuroscience and emerging genetic technologies. He was elected to the Institute of Medicine in 2000, renamed the National Academy of Medicine, where he completed his second term on the Governing Council from 2012 to 2018, and has chaired the Forum on Neuroscience and Nervous Systems Disorders, which brings together government, industry, patient groups, and academia (2012 to 2018). He also served on the governing board of the National Research Council, the operating arm of the U.S. National Academies, from 2016 to 2018. He is a fellow of the American Academy of Arts and Sciences, a fellow of the American Association for the Advancement of Science, former president of the Society for Neuroscience, former president of the American College of Neuropsychopharmacology, and a Distinguished Life Fellow of the American Psychiatric Association. Dr. Hyman has served as Board Chair of the Charles A. Dana Foundation (NY) since April 2018. Dr. Hyman received a B.A. from Yale College, an M.A. from the University of Cambridge, which he attended as a Mellon fellow, and an M.D. from Harvard Medical School. We believe that Dr. Hyman’s extensive knowledge of neuroscience combined with his leadership skills qualifies him to serve as a member of the Board of Directors. Alfred Sandrock, M.D., Ph.D. Dr. Sandrock has been a member of the Board of Directors since February 2022 and has served as our President and Chief Executive Officer since March 2022. From February 1998 to December 2021, Dr. Sandrock served in positions of increasing responsibility at Biogen Inc., culminating in his service as Executive Vice President, Research and Development from October 2019 to December 2021. Dr. Sandrock also served in various Chief Medical Officer roles from 2012 to 2020, including as Executive Vice President, Chief Medical Officer for Biogen from October 2015 to January 2020, and served on Biogen’s Executive Committee from June 2013 to December 2021. Dr. Sandrock held other senior executive positions at Biogen during his tenure, including Group Senior Vice President and Chief Medical Officer, Chief Medical Officer and Senior Vice President of Development Sciences, Senior Vice President of Neurology Research and Development, and Vice President of Clinical Development, Neurology. Dr. Sandrock has served on the boards of directors of privately-held biotechnology companies Transition Bio, Inc. since February 2022, Neurimmune, Inc. since February 2022, and Neutargeton Therapeutics since February 2025. Dr. Sandrock served on the boards of directors of publicly-traded biotechnology company Neurocrine Biosciences, Inc. (“Neurocrine”) from September 2015 to May 2020 and privately-held biotechnology companies Praxis Precision Medicines, Inc. from May 2017 to February 2020, Disarm Therapeutics, Inc. from April 2017 to October 2020, Atalanta Therapeutics, Inc. from March 2022 to September 2024, and Verge Genomics Inc. from February 2022 to March 2026. Dr. Sandrock earned a B.A. in human biology from Stanford University, an M.D. from Harvard Medical School, and a Ph.D. in neurobiology from Harvard University. We believe Dr. Sandrock’s experience as a senior executive and service on the boards of directors of other life sciences companies qualifies him to serve on the Board of Directors. The Board of Directors unanimously recommends that stockholders vote FOR the election of the Class II director nominees listed above.

  6. Continuing Directors

    Continuing Directors

  7. Class III Directors Continuing in Office Until the 2027 Annual Meeting of Stockholders

    Class III Directors Continuing in Office Until the 2027 Annual Meeting of Stockholders

  8. Class III Directors Continuing in Office Until the 2027 Annual Meeting of Stockholders

    | Name | Positions and Offices Held with Voyager | Director Since | Age | | Michael Higgins | Director | 2015 | 63 | | Jude Onyia, Ph.D. | Director | 2023 | 62 | | Nancy Vitale | Director | 2020 | 55 |

  9. Class III Directors Continuing in Office Until the 2027 Annual Meeting of Stockholders

    Michael Higgins. Mr. Higgins was appointed Chairman of the Board in June 2019. He has been a member of the Board of Directors since July 2015 and also served as our Interim President and Chief Executive Officer from June 2021 to March 2022. Mr. Higgins has served as chairman of the board of directors of Pulmatrix, Inc., a publicly-traded biopharmaceutical company, since April 2020, and Nocion Therapeutics, Inc., a privately-held biopharmaceutical company, since September 2020. Mr. Higgins has also served as a member of the board of directors of Cyclerion Therapeutics, Inc., a publicly-traded biopharmaceutical company, since November 2023; Camp4 Therapeutics Corporation, a publicly-traded biopharmaceutical company, since October 2017; Kindex Pharmaceuticals from 2015 to 2021; and Sea Pharmaceuticals, LLC, a privately-held pharmaceutical company, since October 2016. Mr. Higgins previously served as a member of the board of directors of Genocea Biosciences Inc., a publicly-traded immuno-oncology company, from February 2015 to May 2022. Mr. Higgins is a serial entrepreneur who has helped launch and build numerous companies during his career. He served as Entrepreneur-in-Residence and Advisor at Polaris Partners, an investment company, from 2015 to 2021. From 2003 to 2014 he served as Senior Vice President, Chief Operating Officer at Ironwood Pharmaceuticals Inc, a biopharmaceutical company. Prior to 2003, Mr. Higgins held a variety of senior business positions at Genzyme Corporation, including Vice President of Corporate Finance and Vice President of Business Development. Mr. Higgins earned a B.S. from Cornell University and an M.B.A. from the Amos Tuck School of Business at Dartmouth College. We believe that Mr. Higgins’ financial and business expertise, including his diversified background as an executive officer in public pharmaceutical companies and service on the boards of directors of other life sciences companies, qualifies him to serve as a member of the Board of Directors. Jude Onyia, Ph.D. Dr. Onyia has been a member of the Board of Directors since February 2023. Dr. Onyia has served as the Chief Scientific Officer of Neurocrine since November 2021 and leads the drug discovery and non-clinical development teams. Dr. Onyia previously served as the Chief Scientific Officer of Capsida Biotherapeutics, Inc., a gene therapy platform company, from June 2021 to November 2021. From January 1995 to June 2021, Dr. Onyia served in various roles of increasing responsibility with Eli Lilly and Company, a pharmaceutical company, including as Vice President of Biotechnology Discovery Research from September 2014 to June 2021. Dr. Onyia holds a B.S. in forest biology from the State University of New York (SUNY) College of Environmental Science and Forestry, and a Ph.D. in cell and molecular biology from the SUNY Health Science Center at Syracuse. We believe Dr. Onyia’s more than 25 years of experience in leading research and development in large and small biopharmaceutical and biotechnology companies qualifies him to serve on the Board of Directors. Nancy Vitale. Ms. Vitale has been a member of the Board of Directors since September 2020. Ms. Vitale has served as Chief People Officer at Omada Health, Inc., since January 2022. She has also served as a Co-Founder at Partners for Wellbeing, LLC, a boutique human resources consulting firm, since July 2019, and served as Managing Partner from July 2019 January 2022. Previously, Ms. Vitale served in various human resources leadership roles at biotechnology company Genentech, Inc., a member of the Roche Group, from September 2006 to July 2019, including Chief Human Resources Officer and Senior Vice President. Prior to her time at Genentech, Ms. Vitale served as Head of Human Resources, Gillette North America Commercial Operations at Procter & Gamble, a consumer goods company, from April 2004 to August 2006 and as Vice President of Human Resources at Cigna Corporation, a health service company, from April 2001 to April 2004. Ms. Vitale served on the Board of Directors for the Make-A-Wish Foundation from January 2017 to July 2021. Ms. Vitale received a B.B.A. from the University of Michigan and an M.B.A. from the Goizueta Business School at Emory University. We believe Ms. Vitale’s business and human resources expertise, including her experience as a human resources executive in the biotechnology industry, qualifies her to serve as a member of the Board of Directors.

  10. Class I Directors Continuing in Office Until the 2029 Annual Meeting of Stockholders

    Class I Directors Continuing in Office Until the 2029 Annual Meeting of Stockholders

  11. Class I Directors Continuing in Office Until the 2029 Annual Meeting of Stockholders

    | Name | Positions and Offices Held with Voyager | Director Since | Age | | Grace E. Colón, Ph.D. | Director | 2023 | 59 | | Catherine J. Mackey, Ph.D. | Director | 2022 | 70 | | Glenn Pierce, M.D., Ph.D. | Director | 2017 | 70 | | George Scangos, Ph.D. | Director | 2023 | 78 |

  12. Class I Directors Continuing in Office Until the 2029 Annual Meeting of Stockholders

    Grace E. Colón, Ph.D. Dr. Colón has been a member of the Board of Directors since January 2023. She has been CEO, President and member of the board of directors of privately-held OmniPulse Biosciences since January 2026. She has also served as Chair of the boards of directors of privately-held biotechnology company Aronora, Inc., since August 2025, privately-held biotechnology company Bloom Science, Inc. since May 2023, and privately-held consumer health platform company Emm Technology Ltd. since March 2023, and a member of the board of directors and cofounder of privately-held biotechnology company Inaya Therapeutics, Inc. since July 2023. From March 2013 to September 2022, Dr. Colón served as President, Chief Executive Officer and Director of InCarda Therapeutics, Inc., a privately-held biopharmaceutical company, where she oversaw advancement of the lead cardiovascular program from pre-clinical stage to Phase 3. She previously served as the Executive Chair and a member of the board of directors of ProterixBio, Inc. (formerly Bioscale, Inc.), a privately-held diagnostics company, from November 2016 to December 2023, and President and Chief Executive Officer of ProterixBio from February 2015 to October 2016. Dr. Colón also served in various capacities with New Science Ventures, a venture capital firm, including as Partner from January 2014 to June 2016 and Senior Advisor from July 2016 through June 2019. Prior to her time at New Science Ventures, her experiences include roles such as Senior Vice President and the founding President of the Industrial Products Division at Precigen, Inc. (formerly Intrexon Corporation), a publicly-traded biopharmaceutical company, from October 2010 to May 2012; in various capacities of increasing responsibility at Gilead Sciences, Inc., a publicly-traded biopharmaceutical company, from February 2005 to August 2010, most recently as head of Global Clinical Operations; and as Vice President of Corporate Planning at Affymetrix, Inc., a publicly-traded genomics company (acquired by Thermo Fisher Scientific, Inc.), from August 2000 to January 2005. Previously, Dr. Colón served as a member of the boards of directors of CareDx, Inc., a publicly-traded molecular diagnostics company, from July 2019 to June 2024; Cocoon Biotech, Inc., a privately-held biotechnology company, from October 2016 to March 2020; InterLink AI (formerly PerceptiMed, Inc.), a privately-held healthcare technology company, from January 2015 to June 2019; and Paradigm Diagnostics, Inc., a privately-held molecular diagnostics company (acquired by Exact Sciences Corp.), from February 2017 to June 2019. Dr. Colón has served as a term member of the Massachusetts Institute of Technology Corporation (MIT Board of Trustees) since July 2021, and a member of the board of directors of the Biotechnology Innovation Organization (BIO) since June 2021. On the BIO board of directors, she also served as Vice Chair of the Emerging Companies Section and a member of the Executive Committee from June 2023 through June 2025. In March 2024, Dr. Colón was elected as a Fellow of the American Institute for Medical and Biological Engineering (AIMBE). Dr. Colón received her Ph.D. in chemical engineering from the Massachusetts Institute of Technology, where she was a National Science Foundation Fellow, and earned a B.S. in chemical engineering from the University of Pennsylvania, where she was a Benjamin Franklin Scholar. We believe Dr. Colón’s more than 25 years of experience in leading biopharma, genomics, healthcare and industrial biotechnology companies, serving as a venture capital partner, and service on the boards of directors of other life sciences companies qualifies her to serve on the Board of Directors. Catherine J. Mackey, Ph.D. Dr. Mackey has been a member of the Board of Directors since August 2022. Dr. Mackey has served on the board of directors of publicly-traded biotechnology company IDEAYA Biosciences, Inc. since April 2022. She previously served as the Chair of the board of directors of Cour Pharmaceuticals Development Co., Inc., a privately-held, clinical-stage company focused on immunomodulation, from May 2014 to December 2022. She also previously served as a member of the board of directors of publicly-traded biotechnology company Avid Bioservices, Inc. from July 2019 to February 2025, when Avid was acquired by GHO Capital Partners LLP and Ampersand Capital Partners; publicly traded biotechnology company Trillium Therapeutics Inc. from June 2021 to November 2021, when Trillium was acquired by Pfizer Inc.; publicly-traded biopharmaceutical company Poseida Therapeutics, Inc. from January 2019 to June 2021; and publicly-traded biopharmaceutical company GW Pharmaceuticals plc from December 2017 to May 2021, when GW Pharmaceuticals was acquired by Jazz Pharmaceuticals. Dr. Mackey previously served as Senior Vice President, Pfizer Global Research and Development, and Director, La Jolla Laboratories, of Pfizer, Inc. from 2001 to 2010. Dr. Mackey earned her B.S. and Ph.D. in microbiology from Cornell University. We believe Dr. Mackey’s more than 30 years of experience in leading biotechnology research, development, and operations in large life sciences companies and service on the boards of directors of other life sciences companies qualifies her to serve on the Board of Directors. Glenn Pierce, M.D., Ph.D. Dr. Pierce has been a member of the Board of Directors since January 2017 and also served as our Interim Chief Scientific Officer from June 2021 to June 2022 and a consultant from June 2022 to June 2023. Dr. Pierce served as consultant at Ambys Medicines, a biotechnology company, from August 2020 to March 2022, and previously served as Chief Medical Officer of Ambys from August 2018 to August 2020. Since August 2014, Dr. Pierce has served as a consultant to several biotechnology companies. He has served as an Entrepreneur-in-Residence at Third Rock Ventures, LLC, a life sciences venture capital firm focused on the formation, development and strategy of new companies, since January 2016. He retired from Biogen Idec, Inc., a biotechnology company, in May 2014, where he had worked since March 2009 and most recently served as Senior Vice President leading the Hematology, Cell and Gene Therapies division. Prior to Biogen, he served in a variety of biotechnology and biopharmaceutical companies, including Bayer AG, Inspiration Pharma Ltd., Avigen, Inc., Selective Genetics, Inc., and Amgen, Inc. in the areas of tissue regeneration and hematology research, development, or both. Dr. Pierce is the co-author of more than 170 scientific papers and an inventor on more than 15 patents, and he has contributed to the development of a number of novel marketed proteins. He served on the Medical and Scientific Advisory Council, the board of directors and was President of the board of directors of the National Hemophilia Foundation during a span of two decades. Dr. Pierce also served on the Blood Products Advisory Committee at the U.S. Food and Drug Administration and the Committee on Blood Safety and Availability at the U.S. Department of Health and Human Services. He has served on the board of directors of the World Federation of Hemophilia since 2015 and as the organization’s Vice President, Medical since 2018. Dr. Pierce also served on the board of directors of publicly-traded biopharmaceutical company Global Blood Therapeutics, Inc. (acquired by Pfizer, Inc.) from February 2016 to October 2022. Dr. Pierce received a B.A. in Biology, an M.D., and a Ph.D. in immunology, all from Case Western Reserve University in Cleveland, and completed his postgraduate training in pathology and hematology research at Washington University in St. Louis. We believe Dr. Pierce’s more than 35 years of experience in leading biotechnology research and development in small and large, public and private biotechnology and biopharmaceutical companies qualifies him to serve on the Board of Directors.

  13. Class I Directors Continuing in Office Until the 2029 Annual Meeting of Stockholders

    George Scangos, Ph.D. Dr. Scangos has been a member of the Board of Directors since May 2023. Dr. Scangos served as President and Chief Executive Officer of Vir Biotechnology, Inc. from January 2017 until April 2023. He was a member of the board of directors of Vir Biotechnology, Inc., from January 2017 until May 2024. From July 2010 to December 2016, Dr. Scangos served as Chief Executive Officer and as a member of the board of directors of Biogen Inc., a biotechnology company. While at Biogen, Dr. Scangos was named a “Business Person of the Year” by Fortune Magazine and one of “The Best Performing CEOs in the World” by Harvard Business Review for each of the years 2014, 2015, and 2016. Dr. Scangos previously served as President and Chief Executive Officer of Exelixis, Inc., a drug discovery and development company, from October 1996 to July 2010 and as a member of its board of directors from October 1996 to May 2020. From 1987 to 1996, Dr. Scangos served in positions of increasing responsibility at Bayer Corporation, a pharmaceutical company, culminating with his service as President of Biotechnology from 1993 to 1996. Before joining Bayer, Dr. Scangos was a Professor of Biology at Johns Hopkins University. Dr. Scangos also currently serves on the boards of directors of Agilent Technologies, Inc., a publicly traded life sciences, diagnostics and applied chemical analysis company, where he has been a director since 2014; Octave Bioscience, Inc., a privately-held company developing precision medicine approaches for neurodegenerative diseases, where he has been a director since 2017; and Rezo Therapeutics, Inc., a privately-held biotechnology company elucidating the networks involved in a variety of diseases, where he has been a director since 2021. Dr. Scangos served as Chair of Pharmaceutical Research and Manufacturers of America in 2016, and as the Chair of the California Healthcare Institute in 2010. He was a member of the board of directors of the Global Alliance for TB Drug Development from 2006 until 2010. Dr. Scangos currently serves on the board of directors of the Bay Area chapter of Life Science Cares, an organization devoted to using the resources of the biotechnology industry to help individuals from disadvantaged backgrounds. Dr. Scangos received his B.A. in Biology from Cornell University and a Ph.D. in Microbiology from the University of Massachusetts. We believe Dr. Scangos’ experience as a senior executive and service on the boards of directors of other life sciences companies qualifies him to serve on the Board of Directors.

  14. Arrangements between Officers, Directors and Director Nominees

    Arrangements between Officers, Directors and Director Nominees In connection with the 2023 Neurocrine Collaboration Agreement, we and Neurocrine amended and restated our existing investor agreement on January 8, 2023 (the “Amended and Restated Investor Agreement”), providing for standstill and lock-up restrictions and a voting agreement with respect to shares of the Company owned by Neurocrine, and the right for Neurocrine to nominate a director for election to the Board of Directors. Pursuant to the Amended and Restated Investor Agreement, the Board of Directors appointed Jude Onyia, Ph.D., Chief Scientific Officer of Neurocrine, to serve as a Class III director effective February 23, 2023. Under the Amended and Restated Investor Agreement, Neurocrine’s right to designate an individual to serve on the Board of Directors and our agreement to nominate such individual for election is subject to specified conditions and shall terminate upon the earliest of (a) Neurocrine holding less than 10% of our outstanding common stock; (b) a change of control of our company or Neurocrine; (c) a liquidation or dissolution of our Company; and (d) the date that is ten years from the closing date of the Stock Purchase Agreement by and between Neurocrine and us, dated as of January 8, 2023. Other than as disclosed above, there is no arrangement or understanding between any of our executive officers, directors or director nominees and any other person, pursuant to which such person was selected to serve as an executive officer or director, as applicable.

  15. DIRECTOR COMPENSATION

    DIRECTOR COMPENSATION The following table sets forth a summary of the compensation we paid to our non-employee directors for service during 2025. Other than as set forth in the table and described more fully below, we did not pay any compensation to, reimburse any expense of, make any equity awards or non-equity awards to, or pay any other compensation to any of the other non-employee members of the Board of Directors in 2025. We reimburse non-employee directors for reasonable travel expenses. The compensation received by Dr. Sandrock, our President and Chief Executive Officer, is presented under the heading “Summary Compensation Table” and the stock option, restricted stock unit and performance-based restricted stock unit (“PRSU”) grants to Dr. Sandrock are presented in the “Outstanding Equity Awards at Fiscal Year-End Table — 2025”.

  16. DIRECTOR COMPENSATION

    | Name | Fees Earned or Paid in Cash ($) | Option Awards ($)(1) | All Other Compensation ($) | Total ($) | | Grace E. Colón, Ph.D.(2) | 54,000 | 53,760 | — | 107,760 | | James A. Geraghty(3) | 53,500 | 53,760 | — | 107,260 | | Michael Higgins(4) | 92,500 | 53,760 | — | 146,260 | | Steven Hyman, M.D.(5) | 54,000 | 53,760 | — | 107,760 | | Catherine J. Mackey, Ph.D.(6) | 53,000 | 53,760 | — | 106,760 | | Jude Onyia, Ph.D.(7) | 45,000 | 53,760 | — | 98,760 | | Glenn Pierce, M.D., Ph.D.(8) | 51,000 | 53,760 | — | 104,760 | | George Scangos, Ph.D.(9) | 50,000 | 53,760 | — | 103,760 | | Nancy Vitale(10) | 55,000 | 53,760 | — | 108,760 |

  17. DIRECTOR COMPENSATION

    (1) Amounts represent the aggregate grant date fair value of stock option awards granted to our directors in 2025 computed in accordance with Financial Accounting Standards Board Accounting Standards Codification Topic 718 (“FASB ASC Topic 718”). The assumptions used in the valuation of these awards are consistent with the valuation methodologies specified in the notes to our consolidated financial statements included in our 2025 Annual Report on Form 10-K. The amounts above reflect our aggregate accounting expense for these awards and do not necessarily correspond to the actual value that will be recognized by the directors upon the vesting of the awards or any sale of the underlying securities. (2) Fees earned or paid in cash include $40,000 for service as a non-employee director, $6,000 for service on the Compensation Committee, and $8,000 for service on the Audit Committee. Dr. Colón held options to purchase an aggregate of 114,000 shares of our common stock as of December 31, 2025. (3) Fees earned or paid in cash include $40,000 for service as a non-employee director, $6,000 for service as the Chair of the Audit Committee and $7,500 for service as the Chair of the Nominating and Corporate Governance Committee. The Audit Committee fees reflect proration based on service through June 30, 2025, and service as the Chair of the Audit Committee through March 31, 2025. The Nominating and Corporate Governance Committee fees reflect proration based on service as Chair through June 30, 2025, although Mr. Geraghty remained a member. Mr. Geraghty held options to purchase an aggregate of 182,000 shares of our common stock as of December 31, 2025. (4) Fees earned or paid in cash include $40,000 for service as a non-employee director, $32,500 for service as the Chair of the Board of Directors, $12,000 for service as the Chair of the Audit Committee, $3,000 for service on the Compensation Committee, and $5,000 for service on the Nominating and Corporate Governance Committee. The Audit Committee fees reflect proration based on service as a member and Chair beginning April 1, 2025. The Compensation Committee fees reflect proration based on service through June 30, 2025. Mr. Higgins held options to purchase an aggregate of 443,000 shares of our common stock as of December 31, 2025. Fees earned or paid in cash include $40,000 for service as a non-employee director, $4,000 for service on the Audit Committee and $10,000 for service as the Chair of the Science and Technology Committee. The Audit Committee fees reflect proration based on service through June 30, 2025. Dr. Hyman held options to purchase an aggregate of 182,000 shares of our common stock as of December 31, 2025. Fees earned or paid in cash include $40,000 for service as a non-employee director, $8,000 for service on the Audit Committee, and $5,000 for service on the Science and Technology Committee. Dr. Mackey held options to purchase an aggregate of 114,000 shares of our common stock as of December 31, 2025. (7) Fees earned or paid in cash include $40,000 for service as a non-employee director and $5,000 for service on the Science and Technology Committee. Dr. Onyia held options to purchase an aggregate of 114,000 shares of our common stock as of December 31, 2025. (8) Fees earned or paid in cash include $40,000 for service as a non-employee director, $6,000 for service on the Compensation Committee, and $5,000 for service on the Science and Technology Committee. Dr. Pierce held options to purchase an aggregate of 324,000 shares of our common stock as of December 31, 2025. (9) Fees earned or paid in cash include $40,000 for service as a non-employee director, $2,500 for service on the Science and Technology Committee, $7,500 for service as the Chair on the Nominating and Corporate Governance Committee. The Science and Technology Committee fees reflect proration based on a services through June 30, 2025. The Nominating and Corporate Governance Committee fees reflect proration based on a service as Chair beginning on July 1, 2025. Dr. Scangos held options to purchase an aggregate of 114,000 shares of our common stock as of December 31, 2025. (10) Fees earned or paid in cash include $40,000 for service as a non-employee director, $12,500 for service as the Chair of the Compensation Committee and $2,500 for service on the Nominating and Corporate Governance Committee. The Nominating and Corporate Governance Committee fees reflect proration based on a service through June 30, 2025. Ms. Vitale held options to purchase an aggregate of 137,000 shares of our common stock as of December 31, 2025. Our non-employee director compensation policy is designed to provide a total compensation package that enables us to attract and retain, on a long-term basis, high caliber non-employee directors. Under the current policy, adopted in July 2018, and most recently amended in March 2026, all non-employee directors will be paid cash compensation as set forth in the tables below. None of the annual cash retainer amounts were amended during March 2026.

  18. DIRECTOR COMPENSATION

    | | Annual Retainer | | Board of Directors: | | | All non-employee members | $40,000 | | Additional retainer for Chair | $32,500 | | Audit Committee: | | | Chair | $16,000 | | Non-Chair members | $8,000 | | Compensation Committee: | | | Chair | $12,500 | | Non-Chair members | $6,000 | | Science and Technology Committee: | | | Chair | $10,000 | | Non-Chair members | $5,000 | | Nominating and Corporate Governance Committee: | | | Chair | $10,000 | | Non-Chair members | $5,000 |

  19. DIRECTOR COMPENSATION

    From June 2024 to March 2026, under our non-employee director compensation policy then in effect, each person who was initially appointed or elected to the Board of Directors was eligible for an option grant to purchase up to 48,000 shares of our common stock under our stock option plan on the date he or she first became a non-employee director, which vested in equal monthly installments over a period of 36 months, subject to the non-employee director’s continued service as a member the Board of Directors. In addition, on the date of the Annual Meeting of stockholders, each continuing non-employee director who had served on the Board of Directors was eligible to receive an annual option grant to purchase up to 24,000 shares of our common stock, which option would vest in full upon the earlier of (i) the first anniversary of the date of grant or (ii) the next annual meeting of stockholders, in each case, subject to the non-employee director’s continued service as a member of the Board of Directors. All the foregoing options were granted at fair market value on the date of grant. Since March 2026, under our non-employee director compensation policy, as amended, each person who is initially appointed or elected to the Board of Directors is eligible for an option grant to purchase up to 60,000 shares of our common stock under our stock option plan on the date he or she first becomes a non-employee director, which vests in equal monthly installments over 36 months, subject to the non-employee director’s continued service as a member the Board of Directors. In addition, on the date of the annual meeting of stockholders, each continuing non-employee director who has served on the Board of Directors is eligible to receive an annual option grant to purchase up to 30,000 shares of our common stock, which option will vest in full upon the earlier of (i) the first anniversary of the date of grant or (ii) the next annual meeting of stockholders, in each case, subject to the non-employee director’s continued service as a member of the Board of Directors. All the foregoing options will be granted at fair market value on the date of grant.

  20. Role of the Compensation Committee

    Role of the Compensation Committee Our Compensation Committee is primarily responsible for developing and implementing our compensation policies. The Compensation Committee oversees our compensation and benefit plans and policies, administers our equity incentive plans, makes recommendations to the independent members of the Board of Directors on compensation for the Chief Executive Officer, and reviews and approves annually all compensation decisions relating to our other officers, including our executive officers. The Compensation Committee considers recommendations from our Chief Executive Officer regarding the compensation of our executive officers other than himself. Our Compensation Committee has the authority under its charter to engage the services of a consulting firm or other outside advisors to assist it in designing our compensation programs and in making compensation decisions. The Compensation Committee engaged Aon Human Capital Solutions, a division of Aon plc (“Aon”), as its compensation consultant until March 2025, at which time the Compensation Committee engaged Alpine Rewards, LLC (“Alpine”). Alpine is the Compensation Committee’s current outside compensation consultant. Historically, our Compensation Committee reviews all compensation components, including base salary, bonus, benefits, equity incentives, and other perquisites, as well as severance arrangements, change-in-control benefits and other forms of executive officer compensation, provides a recommendation on the compensation of our Chief Executive Officer to the Board of Directors, and approves the compensation of our other executive officers. In addition, the Compensation Committee also considers matters related to individual compensation, such as compensation for new executive hires, as well as high-level strategic issues, such as the efficacy of our compensation strategy, potential modifications to that strategy, and new trends, plans, or approaches to compensation, at various meetings throughout the year. The Compensation Committee also makes recommendations to the Board of Directors regarding the compensation of non-employee directors and has the authority to administer our equity-based plans, both directly and through delegated authority, to the Chief Executive Officer.

  21. Compensation Consultant

    Compensation Consultant Until March 2025, the Compensation Committee engaged Aon as its compensation consultant. In March 2024, the Compensation Committee reviewed information regarding the independence and potential conflicts of interest of Aon, considering, among other things, the relationship between Aon, the members of the Board of Directors, and our executive officers and the factors set forth in the Nasdaq listing rules. Based on such review, the Compensation Committee concluded that the engagement of Aon did not raise any conflict of interest. In connection with its engagement, Aon assisted the Committee in conducting competitive compensation assessments for our executive officers for their compensation levels for the fiscal year ended December 31, 2025. In March 2025, the Compensation Committee engaged Alpine as its compensation consultant. In March 2025 and again in March 2026, the Compensation Committee reviewed information regarding the independence and potential conflicts of interest of Alpine, considering, among other things, the relationship between Alpine, the members of the Board of Directors, and our executive officers and the factors set forth in the Nasdaq listing rules. Based on such review, the Compensation Committee concluded that the engagement of Alpine did not raise any conflict of interest. In connection with its engagement, Alpine assisted the Compensation Committee in conducting competitive compensation assessments for our executive officers for their compensation levels for the fiscal year ending December 31, 2026.

  22. Competitive Assessment of Compensation — Peer Group and Market Data

    Competitive Assessment of Compensation — Peer Group and Market Data In evaluating the total compensation of our executive officers, the Compensation Committee, with the assistance of Aon, established a peer group of 18 publicly-traded companies in the biopharmaceutical industry that was comprised of companies whose market capitalization, number of employees, maturity of product development pipeline or area of therapeutic focus are similar to Voyager. Our 2025 peer group includes the following companies:

  23. Competitive Assessment of Compensation — Peer Group and Market Data

    | Voyager 2025 Compensation Peer Group | | | | 4D Molecular Therapeutics, Inc. | MeiraGTx Holdings plc | Solid Biosciences, Inc. | | Adverum Biotechnologies, Inc. | Monte Rosa Therapeutics, Inc. | Stoke Therapeutics, Inc. | | Editas Medicine, Inc. | Neurogene, Inc. | Tenaya Therapeutics, Inc. | | Entrada Therapeutics, Inc. | PepGen, Inc. | Verve Therapeutics, Inc. | | Foghorn Therapeutics, Inc. | Prelude Therapeutics Inc. | Vigil Neuroscience, Inc. | | Lexeo Therapeutics, Inc. | Prime Medicine, Inc. | Wave Life Sciences Ltd. |

  24. Competitive Assessment of Compensation — Peer Group and Market Data

    For the full compensation analysis, Aon supplemented the peer group information with published survey data, which provided a broader market representation of companies and deeper position reporting.

  25. Summary Compensation Table

    Summary Compensation Table The following table presents information regarding the total compensation awarded to, earned by, and paid during the fiscal years ended December 31, 2025, and 2024, to any individual who served as our chief executive officer during 2025, the two most highly compensated executive officers (other than those individuals who served as principal executive officer) who were serving as executive officers at the end of the fiscal year ended December 31, 2025, and up to two individuals who would have been among the most highly compensated executive officers but were not serving as executive officers at the end of the fiscal year ended December 31, 2025. All of these individuals are our named executive officers for 2025:

  26. Summary Compensation Table

    | Name and Principal Position | Year | Salary ($) | Bonus ($) | Stock Awards ($)(1) | Option Awards ($)(2) | Non-equity Incentive Plan Compensation ($)(3) | All Other Compensation ($) | Total ($) | | Alfred Sandrock, M.D., Ph.D. Chief Executive Officer, President, and Director | 2025 | 674,903 | — | 611,900(4) | 843,900(5) | 361,917 | 15,750(6) | 2,508,370 | | | 2024 | 652,080 | — | 988,000(7) | 1,372,800(8) | 351,471 | 15,525(6)(9) | 3,379,876 | | Jacquelyn Fahey Sandell Former Chief Legal Officer | 2025 | 297,842 | — | 175,130(10) | 241,530(11) | — | 760,484(12) | 1,474,986 | | | 2024 | 468,758 | — | 235,600(13) | 327,360(14) | 185,066 | 15,525(6) | 1,232,309 | | Robin Swartz Chief Business Officer & Chief Operating Officer | 2025 | 520,190 | — | 300,675(15) | 414,675(16) | 205,475 | 13,285(6) | 1,454,300 | | | 2024 | 490,745 | — | 380,000(17) | 528,000(18) | 199,534 | 15,525(6) | 1,613,804 | | Toby Ferguson, M.D., Ph.D. Former Chief Medical Officer | 2025 | 513,369 | — | 265,860(19) | 366,660(20) | 202,781 | 15,750(6) | 1,364,420 | | | 2024 | 363,750(9) | — | 1,106,700(21) | 1,333,500(22) | 148,159 | 14,688(6) | 2,966,797 |

  27. Summary Compensation Table

    Amounts represent the aggregate grant date fair value of restricted stock unit awards granted to our named executive officers in 2025 and 2024 computed in accordance with FASB ASC Topic 718. The amounts above reflect our aggregate accounting expense for these awards and do not necessarily correspond to the actual value that will be recognized by the named executive officers upon the vesting and settlement of the awards or any sale of the underlying securities. Amounts represent the aggregate grant date fair value of option awards granted to our named executive officers in 2025 and 2024 computed in accordance with FASB ASC Topic 718. The assumptions used in the valuation of these awards are consistent with the valuation methodologies specified in the notes to our consolidated financial statements included in our 2025 Annual Report on Form 10-K. The amounts above reflect our aggregate accounting expense for these awards and do not necessarily correspond to the actual value that will be recognized by the named executive officers upon the vesting of the awards or any sale of the underlying securities. (3) Amounts represent each executive’s 2025 performance-based variable cash bonus, which was earned for the 2025 performance year and paid in February 2026, and each executive’s 2024 performance-based variable cash bonus, which was earned for the 2024 performance year and paid in February 2025, as applicable. (4) Dr. Sandrock was granted a restricted stock unit award on February 21, 2025, in connection with his ongoing service as our President and Chief Executive Officer. The shares underlying this restricted stock unit award vest and become settleable, subject to Dr. Sandrock’s continued service as an employee on each applicable vesting date, with one-third of the award vesting on each of the first, second and third anniversaries of the grant date. Dr. Sandrock additionally was granted a PRSU award on February 21, 2025, and the shares underlying this award vest and become settleable, subject to Dr. Sandrock’s continued service as an employee on each applicable vesting date, upon the achievement of specified performance milestones over multiple years. As the probability of achieving these performance milestones was not considered “probable” in accordance with FASB ASC Topic 718, the grant date fair value of Dr. Sandrock’s PRSUs was zero. (5) Dr. Sandrock was granted a stock option award on February 21, 2025, in connection with his ongoing service as our President and Chief Executive Officer. The shares underlying this option vest and become exercisable in equal monthly installments over 48 months thereafter, subject to Dr. Sandrock’s continued service as an employee on each applicable vesting date. (6) Amounts represent the company 401(k) matching contributions. (7) Dr. Sandrock was granted a restricted stock unit award on February 9, 2024, in connection with his ongoing service as our President and Chief Executive Officer. The restricted stock unit award vests and becomes settleable, subject to Dr. Sandrock’s continued service as an employee on each applicable vesting date with one-third of the award vesting on each of the first, second and third anniversaries of the grant date. (8) Dr. Sandrock was granted a stock option award on February 9, 2024, in connection with his ongoing service as our President and Chief Executive Officer. The shares underlying this option vest and become exercisable in equal monthly installments over 48 months thereafter, subject to Dr. Sandrock’s continued service as an employee on each applicable vesting date. (9) Due to clerical error in the prior year’s proxy statement, this amount has been corrected in this proxy statement. (10) Ms. Fahey Sandell was granted a restricted stock unit award on February 21, 2025, in connection with her service as our Chief Legal Officer. The restricted stock unit award would have vested and become settled, subject to Ms. Fahey Sandell’s continued service as an employee on each applicable vesting date with one-third of the award vesting on each of the first, second and third anniversaries of the grant date. Ms. Fahey Sandell additionally was granted a PRSU award on February 21, 2025, that would have vested and become settleable, subject to Ms. Fahey Sandell’s continued service as an employee on each applicable vesting date, upon the achievement of specified performance milestones over multiple years. As the probability of achieving these performance milestones was not considered “probable” in accordance with FASB ASC Topic 718, the grant date fair value of Ms. Fahey Sandell’s PRSUs was zero. Ms. Fahey Sandell was granted a stock option award on February 21, 2025, in connection with her service as our Chief Legal Officer. The shares underlying this option vest and become exercisable in equal monthly installments over 48 months thereafter, subject to Ms. Fahey Sandell’s continued service as an employee on each applicable vesting date. All Other Compensation for Ms. Fahey Sandell in 2025 is comprised of (a) 401(k) matching contributions, (b) payments under the Fahey Sandell Consulting Agreement (defined below), (c) payments under the Fahey Sandell Separation Agreement (defined below), including salary continuation, prorated bonus, outplacement services, and COBRA premium contribution, and (d) the fair value of restricted stock unit award and stock option award modifications pursuant to the Fahey Sandell Separation Agreement. (13) Ms. Fahey Sandell was granted a restricted stock unit award on February 9, 2024, in connection with her service as former Chief Legal Officer. The restricted stock unit award vested with respect to one-third of the underlying shares on the first anniversary of the grant date, and would have vested and become settled, subject to Ms. Fahey Sandell’s continued service as an employee on each applicable vesting date thereafter with one-third of the award vesting on each of the second and third anniversaries of the grant date. (14) Ms. Fahey Sandell was granted a stock option award on February 9, 2024, in connection with her service as our Chief Legal Officer. The shares underlying this option would have vested and become exercisable in equal monthly installments over 48 months thereafter, subject to Ms. Fahey Sandell’s continued service as an employee on each applicable vesting date. (15) Ms. Swartz was granted a restricted stock unit award on February 21, 2025, in connection with her ongoing service as our Chief Business Officer and Chief Operating Officer. The restricted stock unit award vests and becomes settleable, subject to Ms. Swartz’s continued service as an employee on each applicable vesting date, with one-third of the award vesting on each of the first, second and third anniversaries of the grant date. Ms. Swartz was additionally granted a PRSU award on February 21, 2025, that vests and becomes settleable, subject to Ms. Swartz’s continued service as an employee on each applicable vesting date, upon the achievement of specified performance milestones over multiple years. As the probability of achieving these performance milestones was not considered “probable” in accordance with FASB ASC Topic 718, the grant date fair value of Ms. Swartz’s PRSUs was zero. (16) Ms. Swartz was granted a stock option award on February 21, 2025, in connection with her ongoing service as our Chief Business Officer and Chief Operating Officer. The shares underlying this option vest and become exercisable in equal monthly installments over 48 months thereafter, subject to Ms. Swartz’s continued service as an employee on each applicable vesting date. (17)

  28. Summary Compensation Table

    Ms. Swartz was granted a restricted stock unit award on February 9, 2024, in connection with her ongoing service as our Chief Business Officer and Chief Operating Officer. The restricted stock unit award vests and becomes settleable, subject to Ms. Swartz’s continued service as an employee on each applicable vesting date, with one-third of the award vesting on each of the first, second and third anniversaries of the grant date. (18) Ms. Swartz was granted a stock option award on February 9, 2024, in connection with her ongoing service as our Chief Business Officer and Chief Operating Officer. The shares underlying this option vest and become exercisable in equal monthly installments over 48 months thereafter, subject to Ms. Swartz’s continued service as an employee on each applicable vesting date. (19) Dr. Ferguson was granted a restricted stock unit award on February 21, 2025, in connection with his ongoing service as our Chief Medical Officer. The restricted stock unit award would have vested and become settleable, subject to Dr. Ferguson’s continued service as an employee on each applicable vesting date with one-third of the award vesting on each of the first, second and third anniversaries of the grant date. Dr. Ferguson was additionally granted a PRSU award on February 21, 2025, that would have vested and become settleable, subject to Dr. Ferguson’s continued service as an employee on each applicable vesting date, upon the achievement of specified performance milestones over multiple years. As the probability of achieving these performance milestones was not considered “probable” in accordance with FASB ASC Topic 718, the grant date fair value of Dr. Ferguson’s PRSUs was zero. (20) Dr. Ferguson was granted a stock option award on February 21, 2025, in connection with his ongoing service as our Chief Medical officer. The shares underlying this option would have vested and become exercisable in equal monthly installments over 48 months thereafter, subject to Dr. Ferguson’s continued service as an employee on each applicable vesting date. Dr. Ferguson was granted a restricted stock unit award on April 1, 2024, in connection with the commencement of his service as our Chief Medical Officer. The restricted stock unit award vested and became settleable with respect to one-third of the underlying shares on the first anniversary of the grant date, and would have vested and become settled, subject to Dr. Ferguson’s continued service as an employee on each applicable vesting date thereafter with one-third of the award vesting on the second and third anniversaries of the grant date. Dr. Ferguson was granted a stock option award on March 25, 2024, in connection with the commencement of his service as our Chief Medical Officer. The shares underlying this option would have vested and become exercisable, subject to Dr. Ferguson’s continued service as an employee of the Company on each applicable vesting date, as follows: 25% of the shares vested on the first anniversary of the grant date, and an additional 2.0833% of the shares will vest on a monthly basis at the end of each one-month period following the first anniversary of the grant date until the four-year anniversary of the grant date.

  29. Potential payments provided upon termination of named executive officers currently employed by the Company

    Potential payments provided upon termination of named executive officers currently employed by the Company We have entered into employment agreements with each of Dr. Sandrock, Ms. Swartz, Ms. Fahey Sandell, and Dr. Ferguson, the material terms of each of which were approved by the Compensation Committee or the Board of Directors. Ms. Fahey Sandell and Dr. Ferguson ceased employment with the Company on August 1, 2025, and February 20, 2026, respectively. These employment agreements reaffirm the executive’s position with the Company during the period of the executive’s employment and provide for at will employment. Pursuant to the applicable employment agreement, each of Dr. Sandrock, Ms. Swartz, Ms. Fahey Sandell, and Dr. Ferguson was or will be eligible, as applicable, to receive certain payments and benefits in the event that the executive’s employment is terminated by us without “cause” (as defined in the applicable employment agreement), or in the event that the executive terminates his or her employment with “good reason” (as defined in the applicable employment agreement), as summarized below. In the event that Dr. Sandrock, Ms. Swartz, Ms. Fahey Sandell, or Dr. Ferguson has terminated or terminates his or her employment with “good reason” or was or is terminated without “cause,” the applicable executive was or would be eligible, as applicable, to receive 12 months of base salary continuation, a pro rata portion of the executive’s target performance-based cash compensation for that fiscal year based on the number of days worked in that fiscal year at the time of termination, and 12 months of COBRA continuation medical benefits subsidized by us provided that the executive executes and does not revoke a separation agreement and release of claims in favor of us and our affiliates. In the event that Dr. Sandrock, Ms. Swartz, Ms. Fahey Sandell, or Dr. Ferguson has terminated or terminates his or her employment with “good reason” or was or is terminated without “cause” within the period ending 12 months following the consummation of a “sale event” (as defined in the applicable employment agreement), the applicable executive was or would be eligible, as applicable, to receive 12 months of base salary continuation, a pro rata portion of the executive’s target performance-based cash compensation for that fiscal year based on the number of days worked in that fiscal year at the time of termination, 12 months of COBRA continuation medical benefits subsidized by us, and all of the executive’s equity awards shall immediately accelerate and become fully exercisable or non-forfeitable as of the date of termination, provided that the executive executes and does not revoke a separation agreement and release of claims in favor of us and our affiliates.

  30. Outstanding Equity Awards at Fiscal Year-End Table — 2025

    Outstanding Equity Awards at Fiscal Year-End Table — 2025 The following table summarizes, for each of the named executive officers, the number of shares of common stock underlying outstanding stock options and number of shares of restricted stock awards held as of December 31, 2025.

  31. Outstanding Equity Awards at Fiscal Year-End Table — 2025

    | | Option Awards | | | | Stock Awards | | | Name | Number of Securities Underlying Unexercised Options (#) Exercisable | Number of Securities Underlying Unexercised Options (#) Unexercisable | Option Exercise Price ($) | Option Expiration Date | Number of Shares or Units of Stock That Have Not Vested (#) | Market Value of Shares or Units of Stock that Have Not Vested ($)(1) | | Alfred Sandrock, M.D., Ph.D. | 41,250 | 2,750(2) | 3.32 | 2/7/2032 | | | | | 881,812 | 58,788(3) | 8.55 | 3/22/2032 | | | | | | | | | 25,000(4) | 98,250 | | | 170,000 | 70,000(5) | 7.43 | 2/17/2033 | | | | | | | | | 40,000(6) | 157,200 | | | 114,400 | 145,600(7) | 7.60 | 2/9/2034 | | | | | | | | | 86,667(8) | 340,601 | | | 58,000 | 232,000(9) | 4.22 | 2/21/2035 | | | | | | | | | 145,000(10) | 569,850 | | | | | | | 231,000(11) | 907,830 | | Jacquline Fahey Sandell(12) | 73,708 | —(13) | 10.18 | 6/30/2026 | | | | | 27,280 | —(14) | 7.60 | 6/30/2026 | | | | | 16,600 | —(15) | 4.22 | 6/30/2026 | | |

  32. Outstanding Equity Awards at Fiscal Year-End Table — 2025

    | | Option Awards | | | | Stock Awards | | | Name | Number of Securities Underlying Unexercised Options (#) Exercisable | Number of Securities Underlying Unexercised Options (#) Unexercisable | Option Exercise Price ($) | Option Expiration Date | Number of Shares or Units of Stock That Have Not Vested (#) | Market Value of Shares or Units of Stock that Have Not Vested ($)(1) | | Robin Swartz | 76,500 | —(16) | 8.43 | 1/11/2031 | | | | | 117,500 | 2,500(17) | 2.85 | 1/13/2032 | | | | | 59,800 | 5,200(18) | 3.32 | 2/7/2032 | | | | | 56,666 | 23,334(19) | 7.43 | 2/17/2033 | | | | | | | | | 13,333(20) | 52,399 | | | 44,000 | 56,000(21) | 7.60 | 2/9/2034 | | | | | | | | | 33,334(22) | 131,003 | | | 28,500 | 114,000(23) | 4.22 | 2/21/2035 | | | | | | | | | 71,250(24) | 280,013 | | | | | | | 78,000(25) | 306,540 | | Toby Ferguson, M.D., Ph.D.(31) | 91,875 | 118,125(26) | 9.26 | 5/20/2026 | | | | | | | | | 70,000(27) | 275,100 | | | 25,200 | 100,800(28) | 4.22 | 5/20/2026 | | | | | | | | | 63,000(29) | 247,590 | | | | | | | 78,000(30) | 306,540 |

  33. Outstanding Equity Awards at Fiscal Year-End Table — 2025

    The value of restricted stock unit awards is based on the closing price of our common stock on the Nasdaq Global Select Market on December 31, 2025, which was $3.93 per share. (2) Dr. Sandrock’s stock option award to purchase 44,000 shares of our common stock was granted on February 7, 2022. The shares underlying this option award vest and become exercisable in equal quarterly installments over four years thereafter, subject to Dr. Sandrock’s continued service as a director on each applicable vesting date. (3) Dr. Sandrock’s stock option award to purchase 940,600 shares of our common stock was granted on March 22, 2022. The shares underlying this option vest and become exercisable, subject to Dr. Sandrock’s continued service as an employee on each applicable vesting date, as follows: 25% of the shares vested on the first anniversary of the grant date, and an additional 2.0833% of the shares will vest on a monthly basis at the end of each one-month period following the first anniversary of the grant date until the four-year anniversary of the grant date. (4) Dr. Sandrock was granted a restricted stock unit award of 100,000 shares of common stock on March 22, 2022. The shares underlying this restricted stock unit award vest and become settleable, subject to Dr. Sandrock’s continued service as an employee on each applicable vesting date, over a four-year period, as follows, commencing on April 1, 2022: 25% of the restricted stock unit award vested on the first anniversary of the vesting commencement date and an additional 25% of the shares underlying this restricted stock unit award vested and will vest at the end of each one-year period following the first anniversary of the vesting commencement date until the four year anniversary of the vesting commencement date. (5) Dr. Sandrock’s stock option award to purchase 240,000 shares of our common stock was granted on February 17, 2023. The shares underlying this option vest and become exercisable in equal monthly installments over 48 months thereafter, subject to Dr. Sandrock’s continued service as an employee on each applicable vesting date. Dr. Sandrock was granted a restricted stock unit award of 120,000 shares of common stock on February 17, 2023. The shares underlying this restricted stock unit award vest and become settleable, subject to Dr. Sandrock’s continued service as an employee on each applicable vesting date, over a three-year period with one-third of the award vesting on each of the first, second and third anniversaries of the grant date. Dr. Sandrock’s stock option award to purchase 260,000 shares of our common stock was granted on February 9, 2024. The shares underlying this option vest and become exercisable in equal monthly installments over 48 months thereafter, subject to Dr. Sandrock’s continued service as an employee on each applicable vesting date. (8) Dr. Sandrock was granted a restricted stock unit award of 130,000 shares of common stock on February 9, 2024. The shares underlying this restricted stock unit award vest and become settleable, subject to Dr. Sandrock’s continued service as an employee on each applicable vesting date, over a three-year period with one-third of the award vesting on each of the first, second and third anniversaries of the grant date. (9) Dr. Sandrock’s stock option award to purchase 290,000 shares of our common stock was granted on February 21, 2025. The shares underlying this option vest and become exercisable in equal monthly installments over 48 months thereafter, subject to Dr. Sandrock’s continued service as an employee on each applicable vesting date. (10) Dr. Sandrock was granted a restricted stock unit award of 145,000 shares of common stock on February 21, 2025. The shares underlying this restricted stock unit award vest and become settleable, subject to Dr. Sandrock’s continued service as an employee on each applicable vesting date, over a three-year period with one-third of the award vesting on each of the first, second and third anniversaries of the grant date. (11) Dr. Sandrock was granted a PRSU award of 231,000 shares of common stock on February 21, 2025. The shares underlying this PRSU award vest and become settleable, subject to Dr. Sandrock’s continued service as an employee on each applicable vesting date, upon the achievement of specified performance milestones over multiple years. (12) On December 31, 2025, upon the cessation of services provided by Ms. Fahey Sandell pursuant to the Fahey Sandell Consulting Agreement, the unvested 160,501 restricted stock unit awards and PRSUs, and stock option awards to purchase 149,412 shares of our common stock were forfeited. (13) Ms. Fahey Sandell’s stock option award to purchase 122,000 shares of our common stock was granted on July 10, 2023. The shares underlying this option vest and become exercisable, subject to Ms. Fahey Sandell’s continued service as an employee on each applicable vesting date, as follows: 25% of the shares vested on the first anniversary of the grant date, and an additional 2.0833% of the shares will vest on a monthly basis at the end of each one-month period following the first anniversary of the grant date until the four-year anniversary of the grant date. In connection with the Fahey Sandell Separation Agreement, the vesting of shares underlying this award ceased on December 31, 2025, upon the cessation of services provided by Ms. Fahey Sandell pursuant to the Fahey Sandell Consulting Agreement. At such time, any unvested portion of this award expired. In accordance with the Fahey Sandell Separation Agreement, any portion of this award that had vested as of such cessation of services shall remain exercisable until June 30, 2026. (14) Ms. Fahey Sandell’s stock option award to purchase 62,000 shares of our common stock was granted on February 9, 2024. The shares underlying this option vest and become exercisable in equal monthly installments over 48 months thereafter, subject to Ms. Fahey Sandell’s continued service as an employee on each applicable vesting date. In connection with the Fahey Sandell Separation Agreement, the vesting of shares underlying this award ceased on December 31, 2025, upon the cessation of services provided by Ms. Fahey Sandell pursuant to the Fahey Sandell Consulting Agreement. At such time, any unvested portion of this award expired. In accordance with the Fahey Sandell Separation Agreement, any portion of this award that had vested as of such cessation of services shall remain exercisable until June 30, 2026. (15) Ms. Fahey Sandell’s stock option award to purchase 83,000 shares of our common stock was granted on February 21, 2025. The shares underlying this option vest and become exercisable in equal monthly installments over 48 months thereafter, subject to Ms. Fahey Sandell’s continued service as an employee on each applicable vesting date. In connection with the Fahey Sandell Separation Agreement, the vesting of shares underlying this award ceased on December 31, 2025, upon the cessation of services provided by Ms. Fahey Sandell pursuant to the Fahey Sandell Consulting Agreement. At such time, any unvested portion of this award expired. In accordance with the Fahey Sandell Separation Agreement, any portion of this award that had vested as of such cessation of services shall remain exercisable until June 30, 2026. Ms. Swartz’s stock option award to purchase 76,500 shares of our common stock was granted on January 11, 2021. The shares underlying this option vest and become exercisable, subject to Ms. Swartz’s continued service as an employee on each applicable vesting date, as follows: 25% of the shares vested on the first anniversary of the grant date, and an additional 2.0833% of the shares will vest on a monthly basis at the end of each one-month period following the first anniversary of the grant date until the four-year anniversary of the grant date.

  34. Outstanding Equity Awards at Fiscal Year-End Table — 2025

    Ms. Swartz’s stock option award to purchase 120,000 shares of our common stock was granted on January 13, 2022. The shares underlying this option vest and become exercisable in equal monthly installments over 48 months thereafter, subject to Ms. Swartz’s continued service as an employee on each applicable vesting date. (18) Ms. Swartz’s stock option award to purchase 65,000 shares of our common stock was granted on February 7, 2022. The shares underlying this option vest and become exercisable in equal monthly installments over 48 months thereafter, subject to Ms. Swartz’s continued service as an employee on each applicable vesting date. (19) Ms. Swartz’s stock option award to purchase 80,000 shares of our common stock was granted on February 17, 2023. The shares underlying this option vest and become exercisable in equal monthly installments over 48 months thereafter, subject to Ms. Swartz’s continued service as an employee on each applicable vesting date. (20) Ms. Swartz was granted a restricted stock unit award of 40,000 shares of common stock on February 17, 2023. The shares underlying this restricted stock unit award vest and become settleable, subject to Ms. Swartz’s continued service as an employee on each applicable vesting date, over a three-year period with one-third of the award vesting on each of the first, second and third anniversaries of the grant date. (21) Ms. Swartz’s stock option award to purchase 100,000 shares of our common stock was granted on February 9, 2024. The shares underlying this option vest and become exercisable in equal monthly installments over 48 months thereafter, subject to Ms. Swartz’s continued service as an employee. (22) Ms. Swartz was granted a restricted stock unit award of 50,000 shares of common stock on February 9, 2024. The shares underlying this restricted stock unit award vest and become settleable, subject to Ms. Swartz’s continued service as an employee on each applicable vesting date, over a three-year period with one-third of the award vesting on each of the first, second and third anniversaries of the grant date. (23) Ms. Swartz’s stock option award to purchase 142,500 shares of our common stock was granted on February 21, 2025. The shares underlying this option vest and become exercisable in equal monthly installments over 48 months thereafter, subject to Ms. Swartz’s continued service as an employee. (24) Ms. Swartz was granted a restricted stock unit award of 71,250 shares of common stock on February 21, 2025. The shares underlying this restricted stock unit award vest and become settleable, subject to Ms. Swartz’s continued service as an employee on each applicable vesting date, over a three-year period with one-third of the award vesting on each of the first, second and third anniversaries of the grant date. (25) Ms. Swartz was granted a PRSU award of 78,000 shares of common stock on February 21, 2025. The shares underlying this PRSU award vest and become settleable, subject to Ms. Swartz’s continued service as an employee on each applicable vesting date, upon the achievement of specified performance milestones over multiple years. (26) Dr. Ferguson’s stock option award to purchase 210,000 shares of our common stock was granted on March 25, 2024. The shares underlying this option vest and become exercisable, subject to Dr. Ferguson’s continued service as an employee on each applicable vesting date, as follows: 25% of the shares underlying this restricted stock unit award vested on the first anniversary of the grant date, and an additional 2.0833% of the shares will vest on a monthly basis at the end of each one-month period following the first anniversary of the grant date until the four-year anniversary of the grant date. In connection with Dr. Ferguson’s departure, vesting of this award ceased as of February 20, 2026, and the unvested portion of this award was forfeited. Dr. Ferguson was granted a restricted stock unit award of 105,000 shares of common stock on April 1, 2024. The shares underlying this restricted stock unit award vest and become settleable subject to Dr. Ferguson’s continued service as an employee of the Company on each applicable vesting date, over a three-year period with one-third of the award vesting on each of the first, second and third anniversaries of the grant date. In connection with Dr. Ferguson’s departure, vesting of this award ceased as of February 20, 2026, and the unvested portion of this award was forfeited. Dr. Ferguson’s stock option award to purchase 126,000 shares of our common stock was granted on February 21, 2025. The shares underlying this option vest and become exercisable in equal monthly installments over 48 months thereafter, subject to Dr. Ferguson’s continued service as an employee. In connection with Dr. Ferguson’s departure, vesting of this award ceased as of February 20, 2026, and the unvested portion of this award was forfeited. (29) Dr. Ferguson was granted a restricted stock unit award of 63,000 shares of common stock on February 21, 2025. The shares underlying this restricted stock unit award vest and become settleable, subject to Dr. Ferguson’s continued service as an employee on each applicable vesting date, over a three-year period with one-third of the award vesting on each of the first, second and third anniversaries of the grant date. In connection with Dr. Ferguson’s departure, vesting of this award ceased as of February 20, 2026, and the unvested portion of this award was forfeited. (30) Dr. Ferguson was granted a PRSU award of 78,000 shares of common stock on February 21, 2025. The shares underlying this PRSU award vest and become settleable, subject to Dr. Ferguson’s continued service as an employee on each applicable vesting date, upon the achievement of specified performance milestones over multiple years. In connection with Dr. Ferguson’s departure, vesting of this award ceased as of February 20, 2026, and the unvested portion of this award was forfeited. (31) On February 20, 2026, upon the termination of Dr. Ferguson’s employment, the unvested 256,000 restricted stock unit awards and PRSUs, and stock option awards to purchase 422,030 shares of our common stock were forfeited.

  35. Pay Versus Performance

    Pay Versus Performance The following tables and related disclosures provide information regarding (i) the “total compensation” of our principal executive officer (the “PEO”), and our other named executive officers (the “Non-PEO NEOs”), in each case as presented under the caption “Summary Compensation Table” in this proxy statement or our prior proxy statements, which we refer to as the SCT Amounts; (ii) the “compensation actually paid” to our PEO and our Non-PEO NEOs, as calculated pursuant to the SEC’s pay-versus-performance rules, which amounts we refer to as our CAP Amounts; (iii) certain financial performance measures; and (iv) the relationship between the CAP Amounts to those financial performance measures.

  36. Pay Versus Performance

    | Year | Summary Compensation Table Total for PEO(1) ($) | Compensation Actually Paid to PEO(2)(3) ($) | Average Summary Compensation Table Total for Non-PEO NEOs(1) ($) | Average Compensation Actually Paid to Non-PEO NEOs(2)(3) ($) | Value of Initial Fixed $100 Investment Based On Total Shareholder Return(4) ($) | Net (Loss) Income (in thousands)(5) ($) | | 2025 | 2,508,370 | 2,050,563 | 1,431,235 | 1,152,236 | 64.43 | (119,721) | | 2024 | 3,379,876 | 891,406 | 2,419,582 | 1,713,457 | 92.95 | (65,002) | | 2023 | 3,172,433 | 5,137,629 | 1,529,285 | 1,728,086 | 138.36 | 132,330 |

  37. Pay Versus Performance

    (1) For 2025, our PEO was Dr. Sandrock, and our Non-PEO NEOs were Ms. Fahey Sandell, Ms. Swartz, and Dr. Ferguson. For 2024, our PEO was Dr. Sandrock, and our Non-PEO NEOs were Dr. Ferguson and Dr. Jorgensen. For 2023, our PEO was Dr. Sandrock and our Non-PEO NEOs were Ms. Fahey Sandell and Ms. Swartz. (2) The CAP Amounts have been calculated in accordance with Item 402(v) of Regulation S-K and do not reflect compensation actually realized or received by our PEO and Non-PEO NEOs. These amounts reflect the SCT Amounts for each year, adjusted as described in footnote 3 below. (3) The CAP Amounts reflect the exclusions and inclusions for the PEO and the Non-PEO NEOs set forth below. Equity values are calculated in accordance with FASB ASC Topic 718. (4) Total shareholder return (“TSR”) assumes an investment of $100 in our common stock on the last trading day before the earliest fiscal year in the above table through and including the end of the fiscal year for which the TSR is calculated. The closing stock price of one share of our common stock on December 31, 2022, was $6.10. (5) Net (loss) income as reported in our Annual Report on Form 10-K for the relevant fiscal year.

  38. Pay Versus Performance

    | Name and Principal Position | Year | Summary Compensation Table Total for PEO ($) | Exclusion of Stock Awards and Option Awards from Summary Compensation Table for PEO ($) | Inclusion of Year-End Fair Value of Equity Awards Granted During Year That Remained Outstanding and Unvested as of Last Day of Year for PEO ($) | Inclusion of Change In Fair Value from Last Day of Prior Year to Last Day of Year of Outstanding and Unvested Equity Awards Granted in Any Prior Year for PEO ($) | Inclusion of Vesting Date Fair Value of Awards Granted During Year that Vested During Year for PEO ($) | Inclusion of Change in Fair Value from Last Day of Prior Year to Vesting Date of Unvested Equity Awards Granted in any Prior Year that Vested During Year for PEO ($) | Exclusion of Fair Value at Last Day of Prior Year of Equity Awards Forfeited During Year for PEO ($) | Compensation Actually Paid to PEO ($) | | Alfred Sandrock, M.D., Ph.D. Chief Executive Officer, President, and Director | 2025 | 2,508,370 | (1,455,800) | 1,151,773 | (254,595) | 138,073 | (37,258) | — | 2,050,563 | | | 2024 | 3,379,876 | (2,360,800) | 1,506,700 | (1,478,112) | 257,348 | (413,606) | — | 891,406 | | | 2023 | 3,172,433 | (2,157,863) | 2,199,100 | 1,076,411 | 299,050 | 548,498 | — | 5,137,629 |

  39. Pay Versus Performance

    | Year | Average Summary Compensation Table Total for Non-PEO NEOs ($) | Exclusion of Average Stock Awards and Option Awards from Summary Compensation Table for Non-PEO NEOs ($) | Inclusion of Average Year-End Fair Value of Equity Awards Granted During Year That Remained Outstanding and Unvested as of Last Day or Year for Non-PEO NEOs ($) | Inclusion of Average Change In Fair Value from Last Day of Prior Year to Last Day of Year of Outstanding and Unvested Equity Awards Granted in Any Prior Year for Non-PEO NEOs ($) | Inclusion of Average Vesting Date Fair Value of Awards Granted During Year that Vested During Year for Non-PEO NEOs ($) | Inclusion of Average Change in Fair Value from Last Day of Prior Year to Vesting Date of Unvested Equity Awards Granted in any Prior Year that Vested During Year for Non-PEO NEOs ($) | Exclusion of Average Fair Value at Last Day or Prior Year of Equity Awards Forfeited During Year for Non-PEO NEOs ($) | Average Compensation Actually Paid to Other Non-PEO NEOs ($) | | 2025 | 1,431,235 | (588,177) | 355,461 | (101,570) | 55,784 | (54,218) | 53,721 | 1,152,236 | | 2024 | 2,419,582 | (1,989,100) | 1,282,975 | — | — | — | — | 1,713,457 | | 2023 | 1,529,285 | (1,043,962) | 1,000,309 | 116,252 | 49,839 | 76,363 | — | 1,728,086 |

  40. Description of Relationship Between NEO Compensation Actually Paid and Company Total Shareholder Return

    Description of Relationship Between NEO Compensation Actually Paid and Company Total Shareholder Return The following chart sets forth the relationship between the CAP Amount to our PEO, the average of the CAP Amount to our Non-PEO NEOs, and the Company’s cumulative TSR over the fiscal three-year period from 2023 through 2025. We utilize several performance measures to align executive compensation with our performance, but those are not financial performance measures, such as total shareholder return. For example, as described in more detail in the section titled “Executive Compensation,” part of the compensation our NEOs are eligible to receive consists of bonuses that are designed to provide appropriate incentives to achieve defined annual corporate goals and reward our NEOs for individual achievement toward those goals. Additionally, we view equity awards, which are an integral part of our executive compensation program, as related to Company performance because the value of each award increases only if the value of our common stock appreciates from the grant date and if the NEO continues employment over the vesting period. We believe equity awards strongly align our NEOs’ interests with those of our stockholders by providing a continuing financial incentive to maximize long-term value for our stockholders and encouraging our NEOs to continue in our employment for the long-term.

  41. Relationship Between NEO Compensation Actually Paid and Company TSR

    Relationship Between NEO Compensation Actually Paid and Company TSR

  42. Description of Relationship Between NEO Compensation Actually Paid and Net (Loss) Income

    Description of Relationship Between NEO Compensation Actually Paid and Net (Loss) Income The following chart sets forth the relationship between the CAP Amount to our PEO, the average of the CAP Amount to our Non-PEO NEOs, and our net loss over the fiscal three-year period from 2023 through 2025. As described in greater detail in the section above titled “Executive Compensation,” the Company’s executive compensation program reflects a variable pay-for-performance philosophy. The metrics that the Company uses for both our long-term and short-term incentive awards are selected based on an objective of incentivizing our NEOs to increase the value of our enterprise for our stockholders. We are not a commercial-stage company, and any revenues recognized during the periods presented relates to license and collaboration activities with our collaboration partners. Consequently, the Company does not currently use any financial performance measures, such as net (loss) income, to link executive compensation actually paid to our performance for our executive compensation program.

  43. Relationship Between NEO Compensation Actually Paid and Net (Loss) Income

    Relationship Between NEO Compensation Actually Paid and Net (Loss) Income

  44. Executive Officer and Director Compensation

    Executive Officer and Director Compensation See “Director Compensation” and “Executive Compensation” for information regarding compensation of directors and executive officers.