DEF 14A of Spero Therapeutics, Inc., Apr 27, 2026

DEF 14A of Spero Therapeutics, Inc.

Filed Apr 27, 2026 · FY20250001193125-26-182201

Open on EDGAR
  1. NOTICE OF 2026 ANNUAL MEETING OF STOCKHOLDERS

    | 5. | To approve the Spero Therapeutics, Inc. 2026 Stock Incentive Plan. |

  2. How does the Board of Directors recommend that I vote on the proposals?

    | • | “FOR” the approval of the Spero Therapeutics, Inc. 2026 Stock Incentive Plan. |

  3. Information Regarding Directors

    Information Regarding Directors Set forth below are the names of the persons nominated as directors and directors whose terms do not expire this year, their ages, their offices in the Company, if any, their principal occupations or employment for at least the past five years, the length of their tenure as directors and the names of other public companies in which such persons hold or have held directorships during the past five years. Additionally, information about the specific experience, qualifications, attributes or skills that our director nominees and directors contribute to the Board of Directors’ experience and effectiveness, as concluded by our Board of Directors at the time of filing of this proxy statement is set forth below:

  4. Information Regarding Directors

    | Name | Age | Position with the Company | | Milind Deshpande, Ph.D. | 69 | Director | | Scott Jackson | 61 | Director | | John C. Pottage, Jr., M.D. | 73 | Director | | Esther Rajavelu | 47 | Director, President, Chief Executive Officer, Chief Financial Officer, Chief Business Officer and Treasurer | | Cynthia Smith | 57 | Director | | Frank E. Thomas | 56 | Chairman | | Kathleen Tregoning | 55 | Director | | Patrick Vink, M.D. | 62 | Director |

  5. Information Regarding Directors

    Our Board of Directors has reviewed the materiality of any relationship that each of our directors has with Spero Therapeutics, Inc., either directly or indirectly. Based upon this review, our Board of Directors has determined that the following members of the Board of Directors are “independent directors” as defined by rules of The Nasdaq Stock Market (“Nasdaq”): Milind Deshpande, Ph.D., Scott Jackson, Cynthia Smith, Frank E. Thomas, Kathleen Tregoning and Patrick Vink, M.D. Ms. Rajavelu is not an independent director under applicable rules because she is our President, Chief Executive Officer, Chief Financial Officer, Chief Business Officer and Treasurer. Dr. Pottage is not an independent director under applicable rules because of his consulting relationship with us. Milind Deshpande, Ph.D. has served on our Board of Directors since January 2014 and previously served as Chairman of our Board of Directors from January 2014 to August 2023. Dr. Deshpande is the current chairman of the board of directors of Avilar Therapeutics, Inc., a biopharmaceutical company, where he has served since May 2021, and a Venture Partner at RA Capital Management, LP, an investment manager, where he has served since October 2018. Dr. Deshpande previously served as the acting President and Chief Executive Officer, and as a member of the board of directors of Hyku Biosciences Inc., a biotechnology company, from January 2022 to March 2024. Prior to Hyku Biosciences, Dr. Deshpande served as the President and Chief Executive Officer at Nayan Therapeutics, Inc., a biotechnology company, from February 2019 to February 2022. Prior to his tenure at Nayan Therapeutics, Dr. Deshpande served as President and Chief Executive Officer of Avilar Therapeutics from January 2020 to June 2021. Additionally, Dr. Deshpande served as President and Chief Executive Officer, and as a member of the board of directors of Achillion Pharmaceuticals, Inc., a biopharmaceutical company (“Achillion”), from May 2013 until May 2018. He joined Achillion in September 2001 as Vice President of Chemistry, was named Head of Drug Discovery in April 2002, Senior Vice President of Drug Discovery in December 2002, Senior Vice President and Chief Scientific Officer in December 2004, Executive Vice President of Research and Chief Scientific Officer in June 2007 and President of Research and Development in October 2010. Prior to joining Achillion, from 1991 to 2001, Dr. Deshpande was Associate Director of Lead Discovery and Early Discovery Chemistry at the Pharmaceutical Research Institute at Bristol-Myers Squibb Co., where he managed the identification of new clinical candidates to treat infectious and neurological diseases. From 1988 to 1991, he held a faculty position at Boston University Medical School. Dr. Deshpande received his Ph.D. in Organic Chemistry from Ohio University, following his undergraduate education in India. We believe that Dr. Deshpande is qualified to serve on our Board of Directors due to his extensive executive leadership experience and his expertise in research and development in the life sciences industry. Scott Jackson has served on our Board of Directors since April 2020. Mr. Jackson served as Chief Executive Officer and as a member of the board of directors of Celator Pharmaceuticals, Inc., a biopharmaceutical company, from April 2008 until July 2016, when the company was acquired by Jazz Pharmaceuticals plc. Mr. Jackson has more than thirty years of corporate leadership experience in the pharmaceutical and biotechnology industry and has held positions of increasing responsibility in sales, marketing and commercial development at Eli Lilly & Company, SmithKline Beecham plc, ImClone Systems Incorporated, Centocor Inc., a division of Johnson & Johnson, Eximias Pharmaceutical Corporation and YM BioSciences Inc. Mr. Jackson presently serves on the board of MacroGenics, Inc. Mr. Jackson previously served as chairman of the board of Mural Oncology plc from November 2023 until December 2025 and served on the board of GlycoMimetics, Inc. from November 2018 until February 2025. Mr. Jackson also serves on the board of directors at Philabundance, a non-profit organization that addresses food insecurity in the Philadelphia area. Mr. Jackson holds a B.S. in pharmacy from the Philadelphia College of Pharmacy and Science and an M.B.A. from the University of Notre Dame. We believe that Mr. Jackson’s extensive executive leadership experience in the pharmaceutical industry and his experience as a member of the board of directors of other publicly traded biotechnology companies, as well as his broad life sciences industry knowledge qualifies him to serve on our Board of Directors. John C. Pottage, Jr., M.D. has served on our Board of Directors since September 2018. Dr. Pottage is currently a lead scientific consultant at Intrepid Alliance Inc., a consortium of innovative biopharmaceutical companies, a position he has held since January 2023. Previously, Dr. Pottage served as Senior Vice President and Chief Scientific and Medical Officer of ViiV Healthcare Company, a pharmaceutical company, from November 2009 to October 2019. From September 2008 to November 2009, Dr. Pottage served as Senior Vice President, Head of Infectious Disease Medicine Development Center at GlaxoSmithKline LLC, a global biopharma company, and, from June 2007 to September 2008, he served as Vice President, Global Clinical Development of Antivirals also at GlaxoSmithKline. Prior to joining GlaxoSmithKline, from May 2002 to May 2007, Dr. Pottage served as Chief Medical Officer and Senior Vice President of Drug Development of Achillion. From July 1998 to May 2002, Dr. Pottage served as Medical Director of Vertex Pharmaceuticals Incorporated. Dr. Pottage served on the board of directors of Pardes Biosciences, Inc. from September 2022 to August 2023. He received his A.B. in biology from Colgate University and his M.D. from Saint Louis University School of Medicine. We believe that Dr. Pottage’s medical background coupled with his broad expertise within the biopharmaceutical sector and his knowledge of the life sciences industry qualifies him to serve on our Board of Directors.

  6. Information Regarding Directors

    Esther Rajavelu has served on our Board of Directors since June 2025. Ms. Rajavelu has served as our President and Chief Executive Officer since May 2025 and as our Chief Financial Officer, Chief Business Officer and Treasurer since November 2023. From January 2025 to May 2025, Ms. Rajavelu served as our Interim President and Chief Executive Officer. Prior to employment with us, Ms. Rajavelu served as Chief Financial Officer of Fulcrum Therapeutics, Inc., a biopharmaceutical company, from January 2022 to April 2023. Prior to that, Ms. Rajavelu served as a senior equities research analyst at UBS Securities from July 2020 to December 2021;as a senior equities research analyst at Oppenheimer & Co. Inc. from June 2018 to July 2020; as an equities research analyst at Deutsche Bank AG from June 2014 to June 2018; as vice president, life sciences mergers and acquisitions and capital and debt advisory at Ernst & Young Capital Advisors, LLC from 2011 to 2014; and in the healthcare investment banking group at Bank of America Merrill Lynch from 2006 to 2011 where she focused on financing and strategic transactions in the biotechnology and medical devices sectors. Ms. Rajavelu received an M.B.A. from The Wharton School of the University of Pennsylvania and a B.A. in Economics and International Relations from Wesleyan University. We believe that Ms. Rajavelu is qualified to serve on our Board of Directors because of the leadership qualities she has demonstrated during her service in various executive leadership roles at our company and other biotechnology companies, as well as her extensive life sciences advisory experience in capital markets, mergers and acquisitions, and corporate strategy during her prior employment as a senior equities research analyst, investment banker and strategy consultant in the biopharmaceutical sector. Cynthia Smith has served on our Board of Directors since March 2019. Since 2017, she has consulted as a strategic advisor for biotechnology companies. Previously, Ms. Smith served as Chief Commercial Officer of ZS Pharma, Inc., a biopharmaceutical company, from June 2013 to December 2016. In December 2015, ZS Pharma, Inc. was acquired and became a subsidiary of AstraZeneca. Prior to joining ZS Pharma, Inc., from October 2008 to March 2013, Ms. Smith served as Vice President, Market Access and Commercial Development at Affymax, Inc. (“Affymax”), a biotechnology company focused on the development and commercialization of novel renal therapies. Prior to Affymax, from 2000 to 2008, she held various senior leadership positions in market access, corporate strategy, government relations and external affairs at Merck & Co, Inc. (“Merck”). Before joining the pharmaceutical industry, Ms. Smith served in the White House Office of Management and Budget (OMB) in the Clinton Administration. Ms. Smith serves on the board of directors of Agios Pharmaceuticals, Inc., Akebia Therapeutics, Inc., Protara Therapeutics, Inc. and Tvardi Therapeutics, Inc. Ms. Smith earned an M.B.A. from the Wharton School of the University of Pennsylvania, an M.S. in public policy from the Eagleton Institute of Politics at Rutgers University, and a B.A. from the University of North Carolina at Chapel Hill. We believe that Ms. Smith is qualified to serve on our Board of Directors due to her broad experience as a member of the board of directors of other publicly traded biotechnology companies, her more than two decades serving in senior leadership roles in the healthcare industry, as well as her extensive life sciences industry knowledge. Frank E. Thomas has served on our Board of Directors since July 2017. Mr. Thomas is currently President and Chief Operating Officer of Orchard Therapeutics plc (“Orchard”), a development-stage biotechnology company headquartered in the United Kingdom, where he served as Chief Financial Officer and Chief Business Officer from January 2018 to March 2020. Prior to Orchard, Mr. Thomas served as the President and Chief Operating Officer of AMAG Pharmaceuticals, Inc. (“AMAG”), a commercial-stage pharmaceutical company, which was a publicly traded company that was subsequently acquired by Covis Pharmaceuticals, Inc., from April 2015 to April 2017, as AMAG’s Executive Vice President and Chief Operating Officer from May 2012 through April 2015 and as Executive Vice President, Chief Financial Officer and Treasurer from August 2011 through May 2012. Prior to AMAG, he served as Senior Vice President, Chief Operating Officer and Chief Financial Officer for Molecular Biometrics, Inc., a commercial-stage medical diagnostics company, from October 2008 to July 2011. Prior to Molecular Biometrics, Mr. Thomas spent four years at Critical Therapeutics, Inc., which was a publicly traded company that subsequently merged with Cornerstone Therapeutics Inc., from April 2004 to March 2008, where he was promoted to President in June 2006 and Chief Executive Officer in December 2006 from the position of Senior Vice President and Chief Financial Officer. He also served on the board of directors of Critical Therapeutics from 2006 to 2008. Prior to that, Mr. Thomas served as the Chief Financial Officer and Vice President of Finance and Investor Relations at Esperion Therapeutics, Inc., a public biopharmaceutical company, from February 2000 to April 2004. Mr. Thomas has served as a member of the board of directors of Larimar Therapeutics Inc. since May 2020. Mr. Thomas holds a B.B.A. from the University of Michigan, Ann Arbor. We believe that Mr. Thomas is qualified to serve on our Board of Directors because of his extensive commercial and operational management experience at biopharmaceutical companies, as well as his extensive expertise with capital markets and other financial matters. Kathleen Tregoning has served on our Board of Directors since October 2021. Ms. Tregoning previously served as Chief Corporate Affairs Officer, from July 2020, and Head of Commercial Strategy, from September 2023, of Cerevel Therapeutics Holdings, Inc. (“Cerevel”), a biopharmaceutical, until its acquisition by Abbvie Inc. in August 2024. Prior to joining Cerevel, from February 2017 to March 2020, Ms. Tregoning served as Executive Vice President for External Affairs at Sanofi S.A. (“Sanofi”), a French multinational pharmaceutical company, where she was responsible for leading an integrated organization that brought together market access, communications, public policy, government affairs, patient advocacy and corporate social responsibility. Prior to joining Sanofi, Ms. Tregoning spent more than a decade at Biogen Inc., a multinational biotechnology company, first as Vice President, Public Policy and Government Affairs, from 2006 to 2015, and then as Senior Vice President, Corporate Affairs, from December 2015 to February 2017. Previously, Ms. Tregoning served as a professional staff member in the United States Congress, where she held health policy roles with the Senate Budget Committee, the House Energy and Commerce Committee, and the House Ways and Means Committee. Ms. Tregoning began her career with Andersen Consulting, where she developed business strategies and processes for clients in a range of industries, and later served as an Assistant Deputy Mayor for Policy and Budget in the office of the Mayor of Los Angeles. Ms. Tregoning graduated from Stanford University with a B.A. in International Relations and holds an M.A. in Public Policy from the Kennedy School of Government at Harvard University. We believe that Ms. Tregoning is qualified to serve on our Board of Directors because of her senior and executive leadership experience in several biopharmaceutical companies, including her knowledge of healthcare public policy and knowledge of the life sciences industry.

  7. Information Regarding Directors

    Patrick Vink, M.D. has served on our Board of Directors since September 2015. Dr. Vink has been an advisor to the pharmaceutical industry since 2015, including his current role as a senior advisor at Athyrium Capital Management, LP, a financial services company, a role that he has held since 2019. Previously, Dr. Vink was employed at Cubist Pharmaceuticals, Inc. (“Cubist”), a biopharmaceutical company, from September 2012 to April 2015, most recently serving as Executive Vice-President and Chief Operating Officer, overseeing all worldwide commercial and technical operations as well as global alliance management, from January 2015 to April 2015. Prior to joining Cubist, Dr. Vink served as Senior Vice President, Global Head of Hospital Business and Global Head of Biologics for Mylan Inc. (“Mylan”), a pharmaceutical company, from September 2007 to September 2012. In this role, Dr. Vink managed Mylan’s global hospital business. Before joining Mylan, Dr. Vink held several leadership positions across the life sciences industry, including Head of Global Business Franchise Biopharmaceuticals for Sandoz Group AG, a division of Novartis AG; Vice President International Business for Biogen Inc.; and Head of Worldwide Marketing, Cardiovascular and Thrombosis for Sanofi-Synthélabo SA. From 2013 to 2015, Dr. Vink served as a member of the Executive Committee of the European Federation of Pharmaceutical Industries and Associations. Dr. Vink is chairman of the board of directors of two privately held companies and previously served on the boards of directors of Santhera Pharmaceuticals AG and Amryt Pharma PLC until June 2023 and April 2023, respectively. Dr. Vink graduated as a medical doctor from the University of Leiden, Netherlands in 1988 and obtained his M.B.A. in 1992 from the University of Rochester. We believe that Dr. Vink is qualified to serve on our Board of Directors because of his extensive operational business experience, significant knowledge of the activities of our company, and diverse background serving on the board of directors of various public and private life science companies.

  8. Committees of the Board of Directors and Meetings

    | • | exercising all rights, authority and functions of the Board of Directors under all of our stock option, stock incentive, employee stock purchase and other equity-based plans; |

  9. Committees of the Board of Directors and Meetings

    | • | reviewing and making recommendations to the Board of Directors with respect to director compensation; |

  10. Committees of the Board of Directors and Meetings

    | • | appointing, compensating and overseeing the work of any compensation consultant, legal counsel or other advisor retained by the Compensation Committee; |

  11. Committees of the Board of Directors and Meetings

    | • | conducting the independence assessment outlined in Nasdaq rules with respect to any compensation consultant, legal counsel or other advisor retained by the Compensation Committee; |

  12. EXECUTIVE OFFICER AND DIRECTOR COMPENSATION

    EXECUTIVE OFFICER AND DIRECTOR COMPENSATION

  13. Summary Compensation Table

    Summary Compensation Table The following table shows the total compensation paid to or accrued by our named executive officers during the last two fiscal years ended December 31, 2024 and 2025.

  14. Summary Compensation Table

    | Name and Principal Position | Year | Salary ($) | Bonus ($)(1) | Stock Awards ($)(2) | Non-Equity Incentive Plan Compensation ($)(3) | All other Compensation ($)(4) | Total ($) | | Esther Rajavelu | 2025 | 577,179 | 687,479 | 413,705 | 372,000 | 11,134 | 2,061,496 | | President and Chief Executive Officer, Chief Financial Officer, Chief Business Officer and Treasurer(5) | 2024 | 483,313 | — | 188,190 | 173,993 | 10,984 | 856,479 | | Tim Keutzer | 2025 | 484,566 | 498,818 | 213,710 | 232,592 | 11,134 | 1,440,820 | | Chief Operating Officer | 2024 | 475,065 | — | 376,378 | 171,023 | 10,984 | 1,033,450 | | Satyavrat Shukla | 2025 | 244,153 | 452,813 | 603,380 | 307,912 | 416,919 | 2,025,178 | | Former President and Chief Executive Officer(6) | 2024 | 603,750 | — | 1,066,817 | 271,686 | 10,984 | 1,953,237 |

  15. Summary Compensation Table

    | (1) Consists of retention bonuses paid to Ms. Rajavelu, Mr. Keutzer and Mr. Shukla, during the year ended December 31, 2025. See “Retention Cash Bonus” below for additional information regarding these retention bonuses. |

  16. Summary Compensation Table

    | (2) These amounts represent the aggregate grant date fair value for restricted stock unit (“RSU”) awards computed in accordance with the rules of Financial Accounting Standards Board Accounting Standard Codification Topic 718, Compensation—Stock Compensation (“ASC Topic 718”). A discussion of the assumptions used in determining grant date fair value may be found in Note 8 to our consolidated financial statements for the year ended December 31, 2025 included in our Annual Report on Form 10-K for the fiscal year ended December 31, 2025 filed with the SEC on March 26, 2026. |

  17. Summary Compensation Table

    | (3) Amounts represent annual cash bonuses earned for the applicable fiscal year. The annual cash bonuses are paid in the first quarter of the calendar year following the year to which the cash bonus relates. |

  18. Summary Compensation Table

    | (4) Amounts in this column include for the year ended December 31, 2025 (i) in the case of Ms. Rajavelu, $634 consists of the dollar value of life insurance premiums we paid with respect to term life insurance, and $10,500 in a matching contribution under our 401(k) plan, (ii) in the case of Mr. Keutzer, $634 consists of the dollar value of life insurance premiums we paid with respect to term life insurance and $10,500 in a matching contribution under our 401(k) plan, and (iii) in the case of Mr. Shukla, $410,550 in severance, $211 consists of the dollar value of life insurance premiums we paid with respect to term life insurance and $6,158 in a matching contribution under our 401(k) plan |

  19. Summary Compensation Table

    | (5) Ms. Rajavelu was appointed President and Chief Executive Officer, effective as of May 2, 2025, and was elected as a member of the Board of Directors at our 2025 annual meeting of stockholders in June 2025. Ms. Rajavelu was previously serving as our Interim President and Chief Executive Officer, since January 2025. Ms. Rajavelu continues to serve as our Chief Financial Officer, Treasurer, and Chief Business Officer since November 2023. |

  20. Summary Compensation Table

    | (6) Mr. Shukla separated from the Company and resigned from the Board of Directors effective as of May 2, 2025. |

  21. Outstanding Equity Awards at 2025 Fiscal Year-End

    Outstanding Equity Awards at 2025 Fiscal Year-End The following table shows grants of stock options and awards outstanding on the last day of the fiscal year ended December 31, 2025 to each of our named executive officers.

  22. Outstanding Equity Awards at 2025 Fiscal Year-End

    | | 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) | | Esther Rajavelu | — | — | — | — | 150,000(2) | 349,500 | | | — | — | — | — | 89,899(3) | 209,465 | | | — | — | — | — | 366,139(4) | 853,104 | | | — | — | — | — | 185,000(5) | 431,050 | | Timothy Keutzer | 55,943(6) | — | 5.90 | 7/5/2027 | — | — | | | 25,016(6) | — | 11.63 | 12/12/2027 | — | — | | | 25,000(6) | — | 6.26 | 1/1/2029 | — | — | | | 55,000(6) | — | 9.34 | 2/2/2030 | — | — | | | 57,432(6) | — | 19.18 | 1/31/2031 | — | — | | | 50,974(7) | 2,216(7) | 11.18 | 1/31/2032 | — | — | | | — | — | — | — | 9,503(8) | 22,142 | | | — | — | — | — | 132,978(3) | 309,839 | | | — | — | — | — | 179,798(4) | 418,929 | | | — | — | — | — | 266,139(5) | 620,104 |

  23. Outstanding Equity Awards at 2025 Fiscal Year-End

    | (1) The market value of the stock awards is based on the closing price of our common stock of $2.33 per share on December 31, 2025. |

  24. Outstanding Equity Awards at 2025 Fiscal Year-End

    | (2) Consists of RSUs. Each RSU represents the right to receive one share of common stock upon vesting. The RSUs vest in four equal annual installments beginning on November 13, 2023, subject to the individual’s continued service through the applicable vesting date. |

  25. Outstanding Equity Awards at 2025 Fiscal Year-End

    | (3) Consists of RSUs. Each RSU represents the right to receive one share of common stock upon vesting. The RSUs vest in four equal annual installments beginning on February 1, 2024 subject to the individual’s continued service through the applicable vesting date. |

  26. Outstanding Equity Awards at 2025 Fiscal Year-End

    | (4) Consists of RSUs. Each RSU represents the right to receive one share of common stock upon vesting. The RSUs vest in four equal annual installments beginning on February 3, 2025, subject to the individual’s continued service through the applicable vesting date. |

  27. Outstanding Equity Awards at 2025 Fiscal Year-End

    | (5) Consists of RSUs. Each RSU represents the right to receive one share of common stock upon vesting. The RSUs vest in four equal annual installments beginning on May 2, 2025, subject to the individual’s continued service through the applicable vesting date. |

  28. Outstanding Equity Awards at 2025 Fiscal Year-End

    | (6) These options are fully vested. |

  29. Outstanding Equity Awards at 2025 Fiscal Year-End

    | (7) 25% of the options vested on December 2, 2023 and an additional 1/36th of the remaining shares vest monthly until the option is fully vested. In addition, in the event of a termination in connection with a change of control, the vesting of these options will accelerate in accordance with the terms of the option and Mr. Keutzer’s employment agreement. |

  30. Outstanding Equity Awards at 2025 Fiscal Year-End

    | (8) Consists of RSUs. Each RSU represents the right to receive one share of common stock upon vesting. The RSUs vest in four equal annual installments beginning on February 1, 2023 subject to the individual’s continued service through the applicable vesting date. |

  31. Outstanding Equity Awards at 2025 Fiscal Year-End

    During the year ended December 31, 2025, our named executive officers did not exercise any stock options.

  32. Director Compensation

    Director Compensation The following table shows the total compensation paid or accrued during the fiscal year ended December 31, 2025 to each of our non-employee directors. Directors who are employed by us are not compensated for their service on our Board of Directors.

  33. Director Compensation

    | Name | Fees Earned or Paid in Cash ($) | Stock Awards ($)(1) | All Other Compensation ($)(2) | Total ($) | | Milind Deshpande, Ph.D. | 62,500 | 57,800 | — | 120,300 | | Scott Jackson | 57,500 | 57,800 | — | 115,300 | | Ankit Mahadevia, M.D.(3) | 45,000 | 57,800 | — | 102,800 | | John C. Pottage, Jr., M.D. | 50,000 | 57,800 | 225,000 | 332,800 | | Cynthia Smith | 60,000 | 57,800 | — | 117,800 | | Frank E. Thomas | 130,000 | 57,800 | — | 187,800 | | Kathleen Tregoning | 50,000 | 57,800 | — | 107,800 | | Patrick Vink, M.D. | 83,750 | 57,800 | — | 141,550 |

  34. Director Compensation

    | (1) These amounts represent the aggregate grant date fair value for RSU awards computed in accordance with ASC 718. A discussion of the assumptions used in determining grant date fair value may be found in Note 8 to our consolidated financial statements, included in our Annual Report on Form 10-K for the fiscal year ended December 31, 2025, filed with the SEC on March 26, 2026. |

  35. Director Compensation

    | (2) For Dr. Pottage, this amount consists of compensation earned during the year ended December 31, 2025 pursuant to the Pottage Consulting Agreement (as defined below). |

  36. Director Compensation

    | (3) Dr. Mahadevia resigned from our Board of Directors effective January 30, 2026. |

  37. Director Compensation

    As of December 31, 2025, the aggregate number of options held by each of our directors was as follows (representing both exercisable and unexercisable option awards, none of which have been exercised):

  38. Director Compensation

    | Name | Number of Shares Underlying Outstanding Stock Options | | Milind Deshpande, Ph.D. | 83,664 | | Scott Jackson | 30,000 | | Ankit Mahadevia, M.D. | 1,338,937 | | John C. Pottage, Jr., M.D. | 33,219 | | Cynthia Smith | 35,848 | | Frank E. Thomas | 63,893 | | Kathleen Tregoning | 85,974 | | Patrick Vink, M.D. | 82,067 |

  39. Director Compensation Policy

    Director Compensation Policy Under our Non-Employee Director Compensation Policy, as amended (the “Non-Employee Director Compensation Policy”), each non-employee director is eligible to receive compensation for his or her service consisting of annual cash retainers and equity awards. Our non-employee directors received the following annual retainers for their service as of December 31, 2025:

  40. Director Compensation Policy

    | Position | Retainer | | Board Member | $40,000 | | Board Chairperson (additional retainer) | 30,000 | | Lead Director, if any (additional retainer) | 18,750 | | Audit Committee Chair | 20,000 | | Compensation Committee Chair | 20,000 | | Nominating and Corporate Governance Committee Chair | 15,000 | | Development Committee Chair | 10,000 | | Audit Committee Member | 10,000 | | Compensation Committee Member | 10,000 | | Nominating and Corporate Governance Committee Member | 7,500 | | Development Committee Member | 5,000 |

  41. Director Compensation Policy

    Our Non-Employee Director Compensation Policy provides the following with respect to equity awards to non-employee directors: (i) the initial equity award consisting of a non-qualified stock option to purchase shares of our common stock upon first appointment to our Board of Directors and vesting in equal monthly installments until the third anniversary of the grant date subject to the non-employee director’s continued service in the amount of 15,000 shares, and (ii) annual equity awards consisting of a grant of 20,000 RSUs (each RSU relating to one share of our common stock) on the date of our annual meeting of stockholders. The RSUs vest on the first anniversary of the grant date, subject to the director’s continued service, and are issued under, and are subject to the terms of, the 2017 Plan. The policy also provides that, prior to the beginning of each calendar year, a non-employee director may elect to receive all or a portion of his or her base annual fee for service on our Board of Directors in the form of a non-qualified stock option to purchase a number of shares of our common stock based on the Black-Scholes value of such option, which option will be granted on the first business day of the calendar year. These options vest in four quarterly installments on the last day of each calendar quarter during the calendar year, subject to the continued service of the non-employee director. Directors may be reimbursed for travel, food, lodging and other expenses directly related to their service as directors. Directors are also entitled to the protection provided by their indemnification agreements and the indemnification provisions in our Amended and Restated Certificate of Incorporation, as amended, and Amended and Restated Bylaws.

  42. Pay Versus Performance

    Pay Versus Performance As required by Item 402(v) of Regulation S-K, we are providing the following information about the relationship between “compensation actually paid” to our principal executive officers (“PEOs”) and the average of our other named executive officers (“NEOs”) and certain financial metrics of our Company. The following table also provides information regarding company performance over the same periods, as well as the relationship of “compensation actually paid” to our PEOs and NEOs to company performance.

  43. Pay Versus Performance

    | | | | | | | | | | Value of Initial Fixed $100 Investment Based On: | | | Year | Summary Compensation Table Total for Esther Rajavelu ($)(1) | Summary Compensation Table Total for Satyavrat Shukla ($)(2) | Summary Compensation Table Total for Ankit Mahadevia ($)(3) | Compensation Actually Paid to Esther Rajavelu ($)(4)(5) | Compensation Actually Paid to Satyavrat Shukla ($)(3)(5) | Compensation Actually Paid to Ankit Mahadevia ($)(4)(5) | Average Summary Compensation Table Total Compensation for Other NEOs ($)(6) | Average Compensation Actually Paid to Other NEOs ($)(6)(7) | Company TSR ($)(8) | Net Income (loss) ($ in thousands) (9) | | 2025 | 2,061,496 | 2,025,178 | — | 3,344,368 | 982,033 | — | 1,440,820 | 2,268,982 | 12 | 8,572 | | 2024 | — | 1,953,238 | — | | 2,245,682 | — | 944,965 | 1,721,025 | 5 | (68,566) | | 2023 | — | 2,024,647 | 2,385,905 | | 2,020,658 | 2,622,030 | 1,446,848 | 1,487,365 | 8 | 22,806 |

  44. Pay Versus Performance

    | (1) Ms. Rajavelu was one of our PEOs during 2025. |

  45. Pay Versus Performance

    | (2) Calculated in accordance with Item 402(v)(2) of Regulation S-K. The following adjustments were made to Mr. Shukla’s total compensation as reported in the Summary Compensation Table to determine “compensation actually paid.” |

  46. Pay Versus Performance

    | Year | Summary Compensation Table Total ($) | Deductions for Reported Grant Date Fair Value of Stock Awards ($)(a) | Deductions for Reported Grant Date Fair Value of Option Awards ($)(a) | Additions for Pay Versus Performance Equity Adjustments ($)(b) | Compensation Actually Paid ($) | | 2025 | 2,025,178 | (603,380) | — | (439,765) | 982,033 | | 2024 | 1,953,238 | (1,066,817) | — | 1,359,261 | 2,245,682 |

  47. Pay Versus Performance

    | (a) Reflects the amounts reported in the Stock Awards column of the Summary Compensation Table in the relevant years. |

  48. Pay Versus Performance

    | (b) The pay versus performance equity adjustments reflect the aggregated sum of the following values for the respective years. |

  49. Pay Versus Performance

    | Year | Year-End Fair Value of Outstanding and Unvested Equity Awards Granted in the Covered Year ($) | Year Over Year Change in Fair Value of Outstanding and Unvested Equity Awards Granted in Prior Years ($) | Year Over Year Change in Fair Value of Equity Awards Granted in Prior Years That Vested in the Covered Year ($) | Total Pay Versus Performance Equity Adjustments ($) | | 2025 | — | (404,790) | (34,975) | (439,765) | | 2024 | 858,445 | 494,488 | 6,328 | 1,359,261 |

  50. Pay Versus Performance

    | (3) Dr. Mahadevia was one of our PEOs during 2023. Calculated in accordance with Item 402(v)(2) of Regulation S-K. The following adjustments were made to Dr. Mahadevia’s total compensation as reported in the Summary Compensation Table to determine “compensation actually paid.” |

  51. Pay Versus Performance

    | Year | Summary Compensation Table Total ($) | Deductions for Reported Grant Date Fair Value of Stock Awards ($)(a) | Deductions for Reported Grant Date Fair Value of Option Awards ($)(a) | Additions for Pay Versus Performance Equity Adjustments ($)(b) | Compensation Actually Paid ($) | | 2023 | 2,385,905 | (999,998) | — | 1,236,123 | 2,622,030 |

  52. Pay Versus Performance

    | (a) Reflects the amounts reported in the Stock Awards columns of the Summary Compensation Table in the relevant year. |

  53. Pay Versus Performance

    | (b) The pay versus performance equity adjustments reflect the aggregated sum of the following values for the respective years. |

  54. Pay Versus Performance

    | (4) Calculated in accordance with Item 402(v)(2) of Regulation S-K. The following adjustments were made to Ms. Rajavelu’s total compensation as reported in the Summary Compensation Table to determine “compensation actually paid.” |

  55. Pay Versus Performance

    | Year | Summary Compensation Table Total ($) | Deductions for Reported Grant Date Fair Value of Stock Awards ($)(a) | Deductions for Reported Grant Date Fair Value of Option Awards ($) | Additions for Pay Versus Performance Equity Adjustments ($)(b) | Compensation Actually Paid ($) | | 2025 | 2,061,496 | (413,705) | — | 1,696,577 | 3,344,368 |

  56. Pay Versus Performance

    | (a) Reflects the amounts reported in the Stock Awards column of the Summary Compensation Table in the relevant years. |

  57. Pay Versus Performance

    | (b) The pay versus performance equity adjustments reflect the aggregated sum of the following values in the relevant year. |

  58. Pay Versus Performance

    | Year | Year-End Fair Value of Outstanding and Unvested Equity Awards Granted in the Covered Year ($) | Year Over Year Change in Fair Value of Outstanding and Unvested Equity Awards Granted in Prior Years ($) | Year Over Year Change in Fair Value of Equity Awards Granted in Prior Years That Vested in the Covered Year ($) | Total Pay Versus Performance Equity Adjustments ($) | | 2025 | 1,284,154 | 311,869 | 100,554 | 1,696,577 |

  59. Pay Versus Performance

    | (5) Measurement date equity fair values are calculated with assumptions derived on a basis consistent with those used for grant date fair value purposes. RSUs are valued based on the closing stock price on the relevant measurement date. Stock options are valued using a Black-Scholes model as at the relevant measurement dates. |

  60. Pay Versus Performance

    | (6) Reflects compensation for the following non-PEO NEOs: |

  61. Pay Versus Performance

    2025: Mr. Keutzer 2024: Ms. Rajavelu and Mr. Keutzer 2023: Ms. Tamara Joseph, our former Chief Legal Officer and Corporate Secretary and Mr. Keutzer

  62. Pay Versus Performance

    | (7) Average “compensation actually paid” for the non-PEO NEOs has been calculated in accordance with Item 402(v)(2) of Regulation S-K. The following adjustments were made to average Summary Compensation Table total compensation for each year to determine average “compensation actually paid.” |

  63. Pay Versus Performance

    | Year | Summary Compensation Table Total | Deductions for Reported Grant Date Fair Value of Stock Awards(a) | Deductions for Reported Grant Date Fair Value of Option Awards(a) | Additions for Pay Versus Performance Equity Adjustments(b) | Compensation Actually Paid | | 2025 | 1,440,820 | (213,710) | — | 1,041,872 | 2,268,982 | | 2024 | 944,965 | (282,284) | — | 1,058,344 | 1,721,025 | | 2023 | 1,446,848 | (499,999) | — | 540,516 | 1,487,365 |

  64. Pay Versus Performance

    | (a) Reflects the average amounts reported in the Stock Awards and Option Awards columns of the Summary Compensation Table in the relevant years. |

  65. Pay Versus Performance

    | (b) The pay versus performance equity adjustments reflect the aggregated sum of the following values for the respective years. |

  66. Pay Versus Performance

    | Year | Average Year-End Fair Value of Outstanding and Unvested Equity Awards Granted in the Covered Year ($) | Average Year Over Year Change in Fair Value of Outstanding and Unvested Equity Awards Granted in Prior Years ($) | Average Year Over Year Change in Fair Value of Equity Awards Granted in Prior Years That Vested in the Covered Year ($) | Total Pay Versus Performance Equity Adjustments ($) | | 2025 | 620,104 | 427,938 | (6,170) | 1,041,872 | | 2024 | 908,191 | 153,664 | (3,511) | 1,058,344 | | 2023 | 147,945 | 371,929 | 20,642 | 540,516 |

  67. Pay Versus Performance

    | (8) As required under Item 201(e) of Regulation S-K, total shareholder return (“TSR”) measures the cumulative value of $100 invested on the last trading day before the earliest fiscal year in the table, or December 31, 2022, through and including the end of the applicable fiscal year for which total shareholder return is calculated, or December 31, 2023, December 31, 2024, and December 31, 2025, respectively. |

  68. Pay Versus Performance

    | (9) The dollar amounts are our net income (loss) amounts reflected in our audited financial statements for the applicable year. |

  69. Pay Versus Performance

    The following graph visually describes the relationship between “compensation actually paid” to PEOs for each applicable year and the average “compensation actually paid” to our other Non-PEO NEOs, to the cumulative total stockholder return of our company. The following graph visually describes the relationship between “compensation actually paid” to our PEOs for each applicable year and the average “compensation actually paid” to our other Non-PEO NEOs, to net income (loss).

  70. PROPOSAL 1: ELECTION OF DIRECTORS

    PROPOSAL 1: ELECTION OF DIRECTORS The Board of Directors has nominated Milind Deshpande, Ph.D. and Kathleen Tregoning for re-election at the Annual Meeting. The Board of Directors currently consists of eight members, classified into three classes as follows: (i) Milind Deshpande, Ph.D. and Kathleen Tregoning constitute our Class III directors with a term ending at the Annual Meeting, (ii) Scott Jackson, John C. Pottage, Jr., M.D. and Cynthia Smith constitute our Class I directors with a term ending at the 2027 annual meeting of stockholders, and (iii) Esther Rajavelu, Frank E. Thomas and Patrick Vink, M.D. constitute our Class II directors with a term ending at the 2028 annual meeting of stockholders. At each annual meeting of stockholders, directors are elected for a full term of three years to succeed those directors whose terms are expiring. The Board of Directors has voted to nominate Milind Deshpande, Ph.D. and Kathleen Tregoning for election at the Annual Meeting as Class III directors for a term of three years to serve until the 2029 annual meeting of stockholders, and until their respective successors are elected and qualified. Unless authority to vote for any of these nominees is withheld, the shares represented by the enclosed proxy will be voted FOR the election as directors of Milind Deshpande, Ph.D. and Kathleen Tregoning. In the event that either nominee becomes unable or unwilling to serve, the shares represented by the enclosed proxy will be voted for the election of such other person as the Board of Directors may recommend in that nominee’s place. We have no reason to believe that any nominee will be unable or unwilling to serve as a director. THE BOARD OF DIRECTORS RECOMMENDS THE ELECTION OF MILIND DESHPANDE, PH.D. AND KATHLEEN TREGONING AS DIRECTORS, AND PROXIES SOLICITED BY THE BOARD OF DIRECTORS WILL BE VOTED IN FAVOR THEREOF UNLESS A STOCKHOLDER HAS INDICATED OTHERWISE ON THE PROXY.

  71. Highlights of the 2026 Plan

    | • | Limit on Non-Employee Director Compensation. The 2026 Plan generally provides that the maximum aggregate amount of cash and value of stock rights (calculated based on grant date fair value for financial reporting purposes) granted in any calendar year to any non-employer director for services as a director may not exceed $750,000. |