DEF 14A of aTYR Pharma Inc, Mar 26, 2026

DEF 14A of aTYR Pharma Inc

Filed Mar 26, 2026 · FY20250001339970-26-000021

Open on EDGAR

  1. ELECTION OF DIRECTORS

    ELECTION OF DIRECTORS

  2. General

    General Our Amended and Restated Certificate of Incorporation, as amended, provides for a board of directors that is divided into three classes, each with a staggered, three-year term. The term of the Class II directors will expire at the Annual Meeting, and three of our Class II directors are nominated for re-election at the Annual Meeting. Our Board of Directors is currently comprised of seven members. If the Class II director nominees are re-elected at the Annual Meeting, the composition of our Board of Directors will be as follows: Class I— Dr. Paul Schimmel and Dr. Sara L. Zaknoen; Class II— Mr. Timothy P. Coughlin, Dr. Jane A. Gross and Mr. Eric Benevich; and Class III— Dr. Svetlana Lucas and Dr. Sanjay S. Shukla. On March 24, 2026, Dr. Paul Schimmel informed the Company that he is retiring from the Board of Directors effective as of May 29, 2026. Dr. Schimmel has served on our Board of Directors since founding our company in 2005, and we thank him for his years of service and scientific leadership. Dr. Schimmel’s decision to retire was not the result of any disagreement with the Company on any matter related to the Company’s operations, policies or practices. The Board of Directors intends to reduce the size of the Board of Directors and rebalance class designations, effective upon Dr. Schimmel’s retirement. In the absence of instructions to the contrary, the persons named as proxy holders in the accompanying proxy intend to vote in favor of the election of the Class II director nominees designated below to serve until the 2029 annual meeting of stockholders and until their successors shall have been duly elected and qualified. Each nominee is currently a director. Our Board of Directors expects that each nominee will be available to serve as a director, but if any such nominee should become unavailable or unwilling to stand for election, it is intended that the shares represented by the proxy will be voted for such substitute nominee as may be designated by the Board of Directors, or alternatively, the Board of Directors may leave a vacancy on the Board of Directors or reduce the size of the Board of Directors. The biographies of our directors and their ages as of March 1, 2026 are set forth below.

  3. General

    | Name | Age | Position | | Sanjay S. Shukla, M.D., M.S. | 54 | President, Chief Executive Officer and Director | | Timothy P. Coughlin (1) | 59 | Chairman of the Board | | Eric Benevich (1)(2) | 60 | Director | | Jane A. Gross, Ph.D. (2)(3) | 69 | Director | | Svetlana Lucas, Ph.D. (1)(3) | 54 | Director | | Paul Schimmel, Ph.D. (4) | 85 | Director | | Sara L. Zaknoen, M.D. (2)(3) | 67 | Director |

  4. General

    (1) Member of the Audit Committee. (2) Member of the Compensation Committee. (3) Member of the Nominating and Corporate Governance Committee. (4) Dr. Schimmel is retiring from the Board of Directors on May 29, 2026.

  5. Nominees for Director

    Nominees for Director Class II: The three individuals listed below are nominated for election as Class II directors to serve a three-year term expiring at the 2029 annual meeting of stockholders and until their respective successors are elected and qualified. Mr. Coughlin and Dr. Gross were previously elected by the stockholders and Mr. Benevich was appointed by the Board of Directors to serve on our Board of Directors in December 2024. Eric Benevich has served as a director since December 2024. Mr. Benevich has served as Chief Commercial Officer of Neurocrine Biosciences, Inc. (“Neurocrine”) since May 2015 and is responsible for all aspects of commercial development, marketing and sales of the Neurocrine product portfolio, including INGREZZA® and CRENESSITYTM. Mr. Benevich has over 30 years of commercial experience in the pharmaceutical industry and previously served in various positions of increasing responsibility at AstraZeneca, Amgen, Peninsula Pharmaceuticals and Avanir Pharmaceuticals in the sales and marketing of drugs such as Prilosec®, Epogen®, Enbrel® and Neudexta®. Mr. Benevich has a Bachelor of Business Administration in International Business from Washington State University. Our Board of Directors believes that Mr. Benevich is qualified to serve on our Board of Directors due to his experience in commercial development in the life science industry. Timothy P. Coughlin has served as a director since April 2017, and as Chairman of our Board of Directors since March 2024. Mr. Coughlin is the former Chief Financial Officer of Neurocrine, a biopharmaceutical company that has received U.S. Food and Drug Administration approval for INGREZZA® and ORILISSA®, both of which were discovered, developed and commercialized during his tenure at Neurocrine from 2002 to 2018. Prior to joining Neurocrine, he was with Catholic Health Initiatives, a nationwide integrated healthcare delivery system, where he served as Vice President, Financial Services. Mr. Coughlin also served as a Senior Manager in the Health Sciences practice of Ernst & Young LLP and its predecessors from 1989 to 1999. Mr. Coughlin serves on the board of directors of Travere Therapeutics, Inc. and ADC Therapeutics SA, both biotechnology companies. He also served on the board of directors of Peloton Therapeutics prior to its sale to Merck & Co in 2019 and Fate Therapeutics, Inc. from 2013 to 2025. Mr. Coughlin holds a master’s degree in international business from San Diego State University and a bachelor’s degree in accounting from Temple University. Mr. Coughlin is a certified public accountant in both California and Pennsylvania. Our Board of Directors believes that Mr. Coughlin is qualified to serve on our Board of Directors due to his extensive background in financial and accounting matters for public companies, his experience overseeing information technology functions, his experience in the life science industry and his years of business and leadership experience. Jane A. Gross, Ph.D. has served as a director since June 2019. Dr. Gross served as the Chief Scientific Officer and Senior Vice President of Research and Development at Aptevo Therapeutics Inc. (“Aptevo”) from September 2016 to September 2021. She currently serves as a consultant to, and on the scientific advisory boards of several biotechnology companies developing drugs for autoimmune, inflammatory and oncology therapeutics. At Aptevo, Dr. Gross led the discovery of novel protein therapeutics based on the ADAPTIR™ and ADAPTIR-FLEX™ platform technologies focusing on development of therapeutics based on immuno-oncology, leading research efforts in molecular biology and protein engineering, immunology, protein and cell sciences, pharmacology, and translational research. Prior to joining Aptevo, Dr. Gross served as Vice President, Applied Research and Non-Clinical Development at Emergent BioSolutions Inc. and Vice President, Immunology Research at ZymoGenetics, Inc., where she led efforts in discovery and development of therapeutics from novel genes. Dr. Gross serves on the board of directors of BriaCell Therapeutics Corp., a biotechnology company. Dr. Gross holds a Ph.D. in Immunology from the University of California, Berkeley under Jim Allison (2018 recipient of the Nobel Prize in Physiology and Medicine) and a Post-Doctoral Fellowship from the University of Washington in Immunology. Our Board of Directors believes that Dr. Gross is qualified to serve on our Board of Directors due to her extensive experience in the biotherapeutics industry and her expertise in immunology and oncology. Continuing Directors:

  6. Class III: Currently Serving Until the 2027 Annual Meeting

    Class III: Currently Serving Until the 2027 Annual Meeting Svetlana Lucas, Ph.D. has served as a director since June 2019. Dr. Lucas currently serves as Chief Business Officer at Scribe Therapeutics, a private genetic medicines company, where she has implemented multiple strategic collaborations with pharmaceutical companies, including Sanofi and Prevail Therapeutics, a subsidiary of Eli Lilly and Company. Prior to her current role, she served as Senior Vice President, Business Development at Tizona Therapeutics, Inc. (“Tizona”), a clinical stage immunotherapy company, where she was responsible for the company’s business development strategy and transactions, including a global strategic collaboration with AbbVie Inc. (“AbbVie”). Before joining Tizona, Dr. Lucas was Head of Oncology and Inflammation External R&D at Amgen Inc. (“Amgen”), where she oversaw business development activities, including Amgen’s strategic cancer immunotherapy research collaboration and licensing agreement with Kite Pharma, and collaborated with Amgen Ventures on several investments in oncology and inflammation. Dr. Lucas joined Amgen following the acquisition of Onyx Pharmaceuticals, Inc. (“Onyx”), where she spearheaded the company’s oncology partnering strategy and due diligence of new opportunities. Prior to Onyx, she held positions of increasing responsibility in strategy, business development and strategic marketing at Amgen, PDL BioPharma/Facet Biotech (acquired by AbbVie), and XOMA Corporation. She began her career as a strategy consultant in the Life Sciences practice of McKinsey & Company, Inc. Dr. Lucas serves on the board of directors of Jasper Therapeutics, Inc., a biotechnology company. Dr. Lucas received her Ph.D. in Molecular Biology and Biochemistry from California Institute of Technology, and an undergraduate degree in Biology from Moscow State University. Our Board of Directors believes that Dr. Lucas is qualified to serve on our Board of Directors due to her extensive business development experience in the biotherapeutics industry. Sanjay S. Shukla, M.D., M.S. has served as our President and Chief Executive Officer and as a director since November 2017. Dr. Shukla served as our Chief Medical Officer from March 2016 to November 2017. From April 2015 to March 2016, Dr. Shukla worked in an advisory capacity for a number of companies, including as a consultant to our company from January 2016 to March 2016. Prior to that, from October 2012 to April 2015, Dr. Shukla served as Vice President and Global Head of Integrated Medical Services for Novartis, a biopharmaceutical company, where he led global medical affairs operations, with oversight for all pharma general medicines therapies, both inline and in development. Dr. Shukla served as Chief Executive Officer of RXMD, a clinical development consultancy that assisted in advancing proof of concept for early stage drug candidates, from April 2009 to September 2012. Prior to that, Dr. Shukla served in a variety of clinical development, data analytics and drug safety roles at Vifor Pharma, a biopharmaceutical company, and Aspreva Pharmaceuticals (acquired by Vifor Pharma). Dr. Shukla received his M.D. from Howard University College of Medicine and his Bachelors of Science in microbiology and Master of Science in epidemiology and biostatistics from the University of Maryland. Our Board of Directors believes that Dr. Shukla is qualified to serve on our Board of Directors due to his experience as our Chief Executive Officer and previously as our Chief Medical Officer, as well as his medical background, experience in the life science industry and his leadership experience.

  7. Class I: Currently Serving Until the 2028 Annual Meeting*

    Class I: Currently Serving Until the 2028 Annual Meeting* Paul Schimmel, Ph.D. has served as a director since September 2005. Dr. Schimmel is currently a director of several private companies. He was a cofounder of Repligen Corporation, Alkermes, Inc., Cubist Pharmaceuticals, Sirtris Pharmaceuticals, and Alnylam Pharmaceuticals, Inc., and a founding Director of Momenta, Inc. Dr. Schimmel is a Professor of Molecular Medicine and of Chemistry at Scripps Research. He was formerly the John D. and Catherine T. MacArthur Professor of Biochemistry and Biophysics in the Department of Biology at the Massachusetts Institute of Technology. Dr. Schimmel holds a B.A. from Ohio Wesleyan University and a Ph.D. in biochemistry and biophysics from the Massachusetts Institute of Technology. He is an elected member of the National Academy of Sciences, the National Academy of Medicine, the National Academy of Inventors, the American Philosophical Society and the American Academy of Arts and Sciences. Our Board of Directors believes Dr. Schimmel is qualified to serve on our Board of Directors due to his role as one of our scientific founders and his discoveries and scientific leadership in the field of tRNA synthetase biology and other areas important to the development of therapeutics. Sara L. Zaknoen, M.D. has served as a director since May 2021. Since June 2014, through her company, Zed Strategic Consulting, Dr. Zaknoen has worked as a clinical drug development consultant with large pharmaceutical and biotechnology companies across multiple disease indications. Previously, Dr. Zaknoen held Chief Medical Officer positions at several biotechnology companies, including Ignyta, Inc., Polynoma LLC, Tragara Pharmaceuticals, Inc. and Cabrellis Pharmaceuticals Corporation. Prior to that, Dr. Zaknoen served as Executive Director of Phase 2/3 Clinical Oncology Research at Novartis Pharmaceutical Corporation, where she provided oversight for a number of important marketed therapies, such as Gleevec®, Tasigna® and Exiade®. This included supervising the execution of clinical studies, including registrational trials, and involvement with new drug applications and label expansion activities. As Director of Clinical Oncology Research at Schering-Plough (now Merck) she was the lead physician on the Temodar® program, supporting its approval and launch. Additional professional experience includes: Assistant Professor of Medicine at the University of Cincinnati Medical Center; Director of Experimental Therapeutics at the Western Pennsylvania Hospital, Western Pennsylvania Cancer Institute; and Medical Staff Fellow at the National Cancer Institute. Dr. Zaknoen completed her residency, internship and fellowship in hematology/oncology at the University of Minnesota. She received her M.D. from Indiana University School of Medicine and her B.S. in chemistry and biology from Valparaiso University. Our Board of Directors believes that Dr. Zaknoen is qualified to serve on our Board of Directors due to her extensive experience in clinical research, her medical background and her experience in the biotherapeutics industry. * Dr. Schimmel is retiring from the Board of Directors on May 29, 2026, and will not serve until the 2028 Annual Meeting.

  8. Executive Summary

    Executive Summary Corporate Performance Highlights In 2025, we executed on our plans to advance the clinical development of efzofitimod, complete key manufacturing milestones for efzofitimod, advance our pipeline, enhance sustainability through meeting financial objectives, and complete critical pre-commercial launch readiness activities for efzofitimod. The Compensation Committee considered the achievements described below in determining achievement of our corporate goals. Certain elements of executive compensation are tied to the level of corporate goal achievement as described further below. Highlights of our performance in 2025 include: • Completed the global pivotal Phase 3 EFZO-FIT study. In September 2025, we announced top-line data from the EFZO-FIT study. The EFZO-FIT study was the largest interventional study ever conducted in pulmonary sarcoidosis. The study did not meet its primary endpoint of change from baseline in mean daily OCS dose at week 48. However, the study demonstrated a clinically meaningful improvement across multiple endpoints, which we believe indicate drug activity for efzofitimod. We have scheduled a Type C meeting with the FDA in mid-April 2026 to review the results and determine the path forward for efzofitimod in pulmonary sarcoidosis. • Completed key manufacturing milestones related to efzofitimod During 2025, our contracted development and manufacturing organizations (CDMOs) completed three process performance qualification manufacturing batches that would be required as part of a potential BLA submission for efzofitimod. We also demonstrated that the drug substance manufactured by our CDMO is comparable in quality, safety and potency to the drug substance manufactured by our previous CDMO, which is currently being used in the EFZO-CONNECT study. • Progressed Enrollment of the Phase 2 EFZO-CONNECT study and announced interim data from the study. We believe efzofitimod has potential applications in the treatment of other ILDs, such as CHP and CTD-ILD, including SSc-ILD and rheumatoid arthritis-associated ILD. As such, we designed the EFZO-CONNECT study in patients with SSc-ILD. In June 2025, we announced interim data from the study showing three out of four efzofitimod-treated diffuse SSc-ILD patients showed clinically important improvement based on the modified Rodnan Skin Score (“mRSS”) assessment at 12 weeks and that efzofitimod was generally well-tolerated at all doses. We expect to complete enrollment of the study in the first half of 2026. • Advanced our diverse pipeline of biologics product candidates based on our understanding of extracellular tRNA synthetase biology. We have advanced two additional tRNA synthetase programs, ATYR0101 and ATYR0750, into preclinical development. We plan to further elucidate the therapeutic potential of these candidates through mechanistic investigations, including in vitro and in vivo preclinical studies. In 2026, we presented a poster related to ATYR0101 at the Keystone Symposia on Fibrosis. The poster demonstrated that subcutaneous delivery of ATYR0101 yielded a favorable pharmacokinetic and immunogenicity profile compared to standard delivery methods while reducing lung inflammation. The findings presented in the poster suggest the ability of ATYR0101 to potentially resolve the cycle of chronic inflammation and fibrosis utilizing a novel mechanism and further support its therapeutic profile for patients suffering from fibrosis. • Enhanced sustainability through meeting financial and operational objectives. We ended 2025 with $80.9 million in cash, cash equivalents, restricted cash and available-for-sale investments. In April 2022, we entered into an Open Market Sale AgreementSM with Jefferies implementing an “at-the-market” offering program (the “ATM Offering Program”), pursuant to which we may offer and sell, from time to time and at our option, shares of our common stock through Jefferies, acting as sales agent. Jefferies is entitled to a fixed commission rate of up to 3.0% of the gross sales proceeds of shares sold under the ATM Offering Program. During 2025, we sold an aggregate of 13,887,177 shares of common stock at a weighted-average price of $4.94 per share for net proceeds of approximately $66.4 million under the ATM Offering Program. Completed critical pre-commercial launch readiness activities prior to top-line data announcement from EFZO-FIT study. In March 2025, we appointed Dalia R. Rayes as Head of Commercial, Global Efzofitimod Franchise. Ms. Rayes oversees our global commercial strategy and operations for the efzofitimod program. Ms. Rayes brings over 25 years of experience building and leading commercial organizations at biotechnology and pharmaceutical companies, including leading rare disease product launches. Under Ms. Rayes, we completed several pre-commercialization efforts in marketing, commercial operations and commercial supply during 2025. 2025 Compensation Actions We structured a significant proportion of target compensation for our Named Executive Officers as variable or at-risk pay, consisting of annual performance bonus and equity awards in the form of stock options. “Target compensation” consists of base salary, target performance bonus opportunity and equity awards granted in 2025. • The Compensation Committee and the Board of Directors determined that we had achieved 75% level of performance in 2025. However, given the results of the EFZO-FIT study, the Compensation Committee lowered the performance-based bonus payout for our executive management team to 60%. • In January 2025, we granted equity awards in the form of stock options to our Named Executive Officers to incentivize and reward for stockholder value creation as part of our annual performance evaluation. Executive Compensation Practices Our Compensation Committee conducts oversight of our compensation program and policies. Our executive compensation philosophy is based on the following objectives: • Aligning executive interests and stockholder interests through long-term incentives linked to Company performance; • Attracting, retaining and motivating superior executive talent who exemplify and enhance the Company culture; and • Providing incentives that reward the achievement of performance goals that directly correlate to the enhancement of stockholder value, as well as to facilitate executive retention. Our executive compensation program generally consists of the following three principal components: base salary, annual performance-based bonuses and long-term incentive compensation. We also provide our executive officers with severance and change-in-control payments and benefits, as well as other benefits generally available to all our employees, including retirement benefits under our tax-qualified retirement plan (“401(k) Plan”) and participation in employee benefit plans. In evaluating our executive compensation program and policies, as well as the short- and long-term value of our executive compensation plans and arrangements, the Compensation Committee focuses on providing a competitive compensation package that provides significant short- and long-term incentives for the achievement of measurable corporate objectives and individual contribution towards our corporate performance. We believe that this approach provides an appropriate blend of short- and long-term incentives to maximize stockholder value.

  9. Executive Summary

    (1) Acelyrin, Inc. merged with Alumis Inc. in February 2025. (2) Longboard Pharmaceuticals, Inc. was acquired by H. Lundbeck A/S in December 2024, and is no longer part of our peer group as of the date of this Proxy Statement. Factors Used in Determining Executive Compensation The Compensation Committee sets, or recommends to the Board of Directors, the compensation of our executive officers at levels the Compensation Committee determines to be competitive and appropriate for each executive officer, using the Compensation Committee’s professional experience and judgment. Compensation decisions are not made by use of a formulaic approach; the Compensation Committee believes that these decisions require consideration of a multitude of relevant factors that may vary from year to year. In making executive compensation decisions, the Compensation Committee generally takes into consideration the following factors: • our corporate performance and business needs; • each executive officer’s individual performance, experience, job function, change in position or responsibilities, and expected future contributions to our company; • internal pay equity among our executive officers and positions; • the need to attract new talent to our executive team and retain existing talent in a highly competitive industry; • a range of market data reference points; • the total compensation cost and stockholder dilution from executive compensation actions; • trends and compensation paid to similarly situated executives within our market; • its compensation consultant’s recommendations, if applicable; • a review of each executive officer’s total targeted and historical compensation and equity ownership; our Chief Executive Officer’s recommendations, based on the Chief Executive Officer’s direct knowledge of the performance of each Named Executive Officer and review of competitive market data, including peer group data provided by a compensation consultant.

  10. SUMMARY COMPENSATION TABLE

    SUMMARY COMPENSATION TABLE The following table presents all of the compensation earned by or paid to our Named Executive Officers during the fiscal years ending December 31, 2025 and 2024.

  11. SUMMARY COMPENSATION TABLE

    | Name and Principal Position | Year | Salary ($) | Option Awards ($) (1) | Nonequity Incentive Plan Compensation ($) (2) | All Other Compensation ($) (3) | Total ($) | | Sanjay S. Shukla, M.D., M.S. | 2025 | 638,977 | 2,595,200 | 210,862 | 11,214 | 3,456,253 | | President and CEO | 2024 | 580,887 | 641,760 | 246,877 | 10,998 | 1,480,522 | | Jill M. Broadfoot | 2025 | 471,790 | 713,680 | 113,229 | 11,214 | 1,309,913 | | Chief Financial Officer | 2024 | 428,899 | 160,440 | 145,826 | 10,998 | 746,163 | | Nancy E. Denyes | 2025 | 451,570 | 713,680 | 108,377 | 11,214 | 1,284,841 | | General Counsel | 2024 | 410,517 | 160,440 | 139,576 | 18,772 | 729,305 |

  12. SUMMARY COMPENSATION TABLE

    (1) Amounts shown reflect aggregate full grant date fair value of equity-based awards granted during the year in accordance with Financial Accounting Standards Board Accounting Standards Codification Topic 718 (“FASB ASC Topic 718”). For additional information on the valuation assumptions underlying the value of these equity awards, see Part II, Item 8 “Financial Statements and Supplementary Data” of our Annual Report in the Notes to Consolidated Financial Statements, Note 7, “Stockholders’ Equity”. These amounts do not necessarily correspond to the actual value recognized or that may be recognized by the Named Executive Officers. Our Named Executive Officers will only realize compensation from grants of stock options to the extent the trading price of our common stock is greater than the exercise price of such stock options. (2) The amounts reported reflect the cash bonus determined by our Compensation Committee (for the Named Executive Officers other than Dr. Shukla), and by our Board of Directors upon recommendation of our Compensation Committee (for Dr. Shukla), based on certain performance goals and achievement of certain developmental, clinical or regulatory milestones as specified by our Board of Directors upon recommendation of our Compensation Committee, as discussed more fully in the section below titled “Annual Performance-Based Bonuses”. (3) The amounts reported in 2025 in this column include: (i) 401(k) Plan employer match of $10,500 and life insurance premium of $714 to Dr. Shukla; (ii) 401(k) Plan employer match of $10,500 and life insurance premium of $714 to Ms. Broadfoot; and (iii) 401(k) Plan employer match of $10,500 and life insurance premium of $714 to Ms. Denyes.

  13. Outstanding Equity Awards at Fiscal Year-End

    Outstanding Equity Awards at Fiscal Year-End The following table sets forth certain information with respect to outstanding equity awards held by each of our Named Executive Officers as of December 31, 2025:

  14. Outstanding Equity Awards at Fiscal Year-End

    | | | Option Awards | | | | | Stock Awards | | | Name | Vesting Commencement Date | Number of Securities Underlying Unexercised Options (#) Exercisable | Number of Securities Underlying Unexercised Options (#) Unexercisable | Equity Incentive Plan Awards: Number of Securities Underlying Unexercised Unearned Options (#) | Option Exercise Price ($) | Option Expiration Date | Number Of Shares Or Units Of Stock That Have Not Vested (#) | Market Value Of Shares Or Units Of Stock That Have Not Vested ($) (5) | | Sanjay S. Shukla, | 3/30/2016 | 11,642(1) | — | — | 68.04 | 3/30/2026 | — | — | | M.D., M.S. | 9/13/2016 | 1,964(2) | — | — | 42.84 | 9/13/2026 | — | — | | | 2/7/2017 | 4,642(2) | — | — | 46.20 | 2/7/2027 | — | — | | | 11/1/2017 | 32,142(2) | — | — | 56.00 | 11/1/2027 | — | — | | | 2/6/2018 | 21,428(2) | — | — | 46.20 | 2/6/2028 | — | — | | | 2/6/2019 | 14,285(2) | — | — | 7.24 | 2/6/2029 | — | — | | | 2/5/2020 | 39,327(2) | — | — | 4.51 | 2/5/2030 | — | — | | | 5/26/2020 | 36,571(2) | — | — | 4.06 | 5/26/2030 | — | — | | | 2/4/2021 | 112,469(2) | — | — | 4.52 | 2/4/2031 | — | — | | | 5/21/2021 | 207,000(2) | — | — | 4.45 | 5/21/2031 | — | — | | | 2/3/2022 | 79,542(2) | 3,458(2) | — | 5.52 | 2/3/2032 | — | — | | | 2/3/2022 | — | — | — | — | — | 10,375(4) | 8,125 | | | 5/24/2022 | 462,501(2) | 53,779(2) | — | 2.81 | 5/24/2032 | — | — | | | 2/16/2023 | 238,067(2) | 98,028(2) | — | 2.20 | 2/16/2033 | — | — | | | 1/5/2024 | 287,500(2) | 312,500(2) | — | 1.50 | 1/5/2034 | — | — | | | 1/8/2025 | 229,166(2) | 770,834(2) | — | 3.74 | 1/8/2035 | — | — | | Jill M. Broadfoot | 7/30/2018 | 14,285(1) | — | — | 11.41 | 7/30/2028 | — | — | | | 2/6/2019 | 3,571(2) | — | — | 7.24 | 2/6/2029 | — | — | | | 2/5/2020 | 23,359(2) | — | — | 4.51 | 2/5/2030 | — | — | | | 5/26/2020 | 15,000(2) | — | — | 4.06 | 5/26/2030 | — | — | | | 2/4/2021 | 50,000(2) | — | — | 4.52 | 2/4/2031 | — | — | | | 5/21/2021 | 54,000(2) | — | — | 4.45 | 5/21/2031 | — | — | | | 2/3/2022 | 28,750(2) | 1,250(2) | — | 5.52 | 2/3/2032 | — | — | | | 2/3/2022 | — | — | — | — | — | 3,750(4) | 2,937 | | | 5/24/2022 | 67,498(2) | 7,849(2) | — | 2.81 | 5/24/2032 | — | — | | | 2/16/2023 | 59,516(2) | 24,507(2) | — | 2.20 | 2/16/2033 | — | — | | | 1/5/2024 | 71,875(2) | 78,125(2) | — | 1.50 | 1/5/2034 | — | — | | | 1/8/2025 | 63,020(2) | 211,980(2) | — | 3.74 | 1/8/2035 | — | — | | Nancy E. Denyes | 1/27/2016 | 1,428(2) | — | — | 85.96 | 1/27/2026 | — | — | | | 9/13/2016 | 2,607(2) | — | — | 42.84 | 9/13/2026 | — | — | | | 2/7/2017 | 2,500(2) | — | — | 46.20 | 2/7/2027 | — | — | | | 2/6/2018 | 3,571(2) | — | — | 46.20 | 2/6/2028 | — | — | | | 5/16/2018 | 3,571(3) | — | — | 11.90 | 5/16/2028 | — | — | | | 2/6/2019 | 8,928(2) | — | — | 7.24 | 2/6/2029 | — | — | | | 2/5/2020 | 13,778(2) | — | — | 4.51 | 2/5/2030 | — | — | | | 5/26/2020 | 15,000(2) | — | — | 4.06 | 5/26/2030 | — | — | | | 2/4/2021 | 45,000(2) | — | — | 4.52 | 2/4/2031 | — | — | | | 5/21/2021 | 59,000(2) | — | — | 4.45 | 5/21/2031 | — | — | | | 2/3/2022 | 20,604(2) | 896(2) | — | 5.52 | 2/3/2032 | — | — | | | 2/3/2022 | — | — | — | — | — | 2,687(4) | 2,104 | | | 5/24/2022 | 78,920(2) | 9,177(2) | — | 2.81 | 5/24/2032 | — | — | | | 2/16/2023 | 59,516(2) | 24,507(2) | — | 2.20 | 2/16/2033 | — | — | | | 1/5/2024 | 71,875(2) | 78,125(2) | — | 1.50 | 1/5/2034 | — | — | | | 1/8/2025 | 63,020(2) | 211,980(2) | — | 3.74 | 1/8/2035 | — | — |

  15. Outstanding Equity Awards at Fiscal Year-End

    1/4th of the shares subject to the option vest one year from Vesting Commencement Date and the remaining shares subject to the option vest in 36 equal monthly installments following the one-year anniversary of the Vesting Commencement Date. The option is subject to full acceleration in the event the employee is terminated by us without Cause or resigns for Good Reason within the period commencing two months prior to and ending 12 months following a Change in Control or Sale Event (as such capitalized terms are defined in the 2015 Stock Plan or the respective stock option agreements evidencing such stock option, as applicable). 1/48th of the total shares subject to the option vest monthly from the Vesting Commencement Date set forth in the table above. The option is subject to full acceleration in the event the employee is terminated by us without Cause or resigns for Good Reason within the period commencing two months prior to and ending 12 months following a Change in Control or Sale Event (as such capitalized terms are defined in the 2015 Stock Plan or the respective stock option agreements evidencing such stock option, as applicable). (3) 1/36th of the total shares subject to the option vest monthly from the Vesting Commencement Date set forth in the table above. The option is subject to full acceleration in the event the employee is terminated by our company without Cause or resigns for Good Reason within the period commencing two months prior to and ending 12 months following a Change in Control or Sale Event (as such capitalized terms are defined in the 2015 Stock Plan, as applicable or the respective stock option agreements evidencing such stock option). (4) Vested in four equal annual installments from Vesting Commencement Date set forth in the table above. (5) Value is based on our closing stock price on December 31, 2025 of $0.78, as reported on the Nasdaq Capital Market.

  16. PAY VERSUS PERFORMANCE

    PAY VERSUS PERFORMANCE We are providing the following information about the relationship between executive compensation actually paid and certain financial performance of our company as required by Section 953(a) of the Dodd-Frank Wall Street Reform and Consumer Protection Act and Item 402(v) of Regulation S-K. Use of the term “compensation actually paid” is required by the SEC’s rules and as a result of the calculation methodology required by the SEC, such amounts differ from compensation actually received by the individuals and the compensation decisions described in the “Executive Compensation” section above.

  17. PAY VERSUS PERFORMANCE

    | Year | Summary Compensation Table Total for Principal Executive Officer (“PEO”) (1) | Compensation Actually Paid to PEO (2) | Average Summary Compensation Table Total for Non-PEO Named Executive Officers (“NEOs”) (3) | Average Compensation Actually Paid to Non-PEO NEOs (2) | Value of Initial Fixed $100 Investment Based On Total Stockholder Return (“TSR”) (4) | Company Net Income (Loss) (thousands)(5) | | (a) | (b) | (c) | (d) | (e) | (f) | (g) | | 2025 | $3,456,253 | $575,965 | $1,297,377 | $506,659 | $35.76 | $(74,118) | | 2024 | $1,480,522 | $3,137,857 | $737,734 | $1,682,268 | $165.30 | $(64,023) | | 2023 | $1,300,525 | $596,670 | $664,560 | $473,280 | $64.38 | $(50,389) |

  18. PAY VERSUS PERFORMANCE

    (1) The dollar amounts reported in column (b) are the amounts of total compensation reported for Dr. Shukla (our Chief Executive Officer and PEO) for each corresponding year in the “Total” column of the Summary Compensation Table. Refer to “Executive Compensation—Summary Compensation Table”. The dollar amounts reported in column (c) represent the amount of “compensation actually paid” to Dr. Shukla and the amounts reported in column (e) represent average “compensation actually paid” to the NEOs as a group (excluding Dr. Shukla), as computed in accordance with Item 402(v) of Regulation S-K. The dollar amounts do not reflect the actual amount of compensation earned by or paid to our NEOs during the applicable year. In accordance with the requirements of Item 402(v) of Regulation S-K, the following adjustments were made to the NEO’s total compensation for each year to determine the compensation actually paid:

  19. PAY VERSUS PERFORMANCE

    | Year | NEOs | Summary Compensation Table (“SCT”) Total Compensation | Deduct: Grant Date Fair Value of the “Stock Awards” and “Option Awards” Columns in the SCT for Applicable Year* | Add: Fair Value at Applicable Year End of Awards Granted during Applicable Year that Remain Unvested as of Applicable Year End* | Add: Change in Fair Value from the end of the Prior Year to the end of the Applicable Year of Awards Granted during Prior Year that were Outstanding and Unvested as of Applicable Year End* | Add: Vesting Date Fair Value of Awards Granted during the Applicable Year that Vested During the Applicable Year* | Add: Change in Fair Value as of the Vesting Date of Awards Granted in Prior Year that Vested in the Applicable Year | Compensation Actually Paid | | 2025 | PEO | $3,456,253 | $2,595,200 | $367,278 | $(1,048,998) | $501,491 | $(104,859) | $575,965 | | | Average Non-PEO NEOs | $1,297,377 | $713,680 | $202,002 | $(510,468) | $275,820 | $(44,392) | $506,659 |

  20. PAY VERSUS PERFORMANCE

    * The valuation assumptions used to calculate fair values did not materially differ from those disclosed at the time of grant. The dollar amounts reported in column (d) represent the average of the amounts reported for Ms. Broadfoot and Ms. Denyes (our non-PEO NEOs for each year presented) as a group in the “Total” column of the Summary Compensation Table in each applicable year. Refer to “Executive Compensation—Summary Compensation Table”. (4) Cumulative TSR is calculated by dividing the sum of the cumulative amount of dividends for the measurement period, assuming dividend reinvestment, and the difference between our company’s share price at the end and the beginning of the measurement period by our company’s share price at the beginning of the measurement period. This amount assumes an initial investment of $100 on December 31, 2022. No dividends were paid on stock or option awards in 2023, 2024 or 2025. (5) The dollar amounts reported represent the amount of net income (loss) reflected in our consolidated audited financial statements for the applicable year. Due to the fact that we are not a commercial-stage company, we did not have any revenue during the periods presented, other than revenue associated with a milestone payment and collaboration revenue under our license and collaboration agreement with Kyorin. Consequently, the Company did not use net income (loss) as a performance measure in its executive compensation program.

  21. Analysis of the Information Presented in the Pay versus Performance Table

    Analysis of the Information Presented in the Pay versus Performance Table We generally seek to incentivize long-term performance, and therefore do not specifically align our performance measures with “compensation actually paid” (as computed in accordance with Item 402(v) of Regulation S-K) for a particular year. In accordance with Item 402(v) of Regulation S-K, we are providing the following descriptions of the relationships between information presented in the Pay Versus Performance table. Compensation Actually Paid and Net Income (Loss) Our company has not historically looked to net income (loss) as a performance measure for our executive compensation program. Our net loss was $(64.0) million in 2024 and $(74.1) million in 2025. As a result, we do not believe there is any meaningful relationship between our net loss and compensation actually paid to our NEOs during the periods presented. The graph below shows the relationship between compensation actually paid to our PEO and the average amount of compensation actually paid to our non-PEO NEOs, on one hand, to the Company’s net loss over the three years presented, on the other hand. Compensation Actually Paid and Cumulative TSR The graph below shows the relationship between compensation actually paid to our PEO and the average amount of compensation actually paid to our non-PEO NEOs, on one hand, to the Company’s cumulative TSR over the three years presented, on the other hand. All information provided above under the “Pay Versus Performance” heading will not be deemed to be incorporated by reference in any filing of our company under the Securities Act of 1933, as amended, or the Securities Exchange Act of 1934, as amended, whether made before or after the date hereof and irrespective of any general incorporation language in any such filing, except to the extent the Company specifically incorporates such information by reference.

  22. Director Compensation

    Director Compensation Our Board of Directors adopted a non-employee director compensation policy in May 2015 that 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. In January 2016, February 2017, February 2022, February 2024, and December 2024 our Board of Directors adopted amendments to the policy with respect to the cash and equity component of compensation to our non-employee directors. Under this policy, as amended to date, all non-employee directors are paid cash compensation for service on the Board of Directors and committees of the Board of Directors as set forth below, prorated based on days of service during a calendar year. Under the policy in place for 2025, non-employee directors were eligible to receive cash and equity compensation as follows:

  23. Director Compensation

    | Board of Directors | Annual Retainer | | All non-employee members | $40,000 | | Additional retainer for Chairperson | $30,000 | | Audit Committee: | | | Chairperson | $20,000 | | Non-Chairperson members | $9,000 | | Compensation Committee: | | | Chairperson | $15,000 | | Non-Chairperson members | $7,000 | | Nominating and Corporate Governance Committee: | | | Chairperson | $10,000 | | Non-Chairperson members | $5,000 |

  24. Director Compensation

    In addition, under the policy, each newly appointed or elected non-employee director will receive an option grant to purchase up to 100,000 shares of common stock, which will vest in equal monthly installments during the 36 months following the grant date, subject to the director’s continued service on our Board of Directors. Thereafter, on the date of each annual meeting of stockholders, each continuing non-employee director will be eligible to receive an annual option grant to purchase up to 50,000 shares of common stock, which will vest in full upon the earlier of the first anniversary of the date of grant or the date of the following annual meeting of stockholders, subject to the director’s continued service on our Board of Directors. All of the foregoing options will be granted with an exercise price equal to the fair market value of our common stock on the date of grant. We have agreed to reimburse all reasonable out-of-pocket expenses incurred by non-employee directors in attending Board and committee meetings.

  25. Director Compensation Table—2025

    Director Compensation Table—2025 The following table sets forth information with respect to the compensation earned by our non-employee directors during the fiscal year ended December 31, 2025. During 2025, Dr. Shukla did not receive compensation for his service on the Board of Directors. The compensation paid to Dr. Shukla as an employee of our company is set forth under the heading “Executive Compensation—Summary Compensation Table”.

  26. Director Compensation Table—2025

    | Name | Fees Earned or Paid in Cash | Option Awards(1) | Total | | Timothy P. Coughlin (Chairman) (2) | $82,200 | $112,085 | $194,285 | | Eric Benevich (3) | $51,764 | $112,085 | $163,849 | | John K. Clarke (4) | $19,775 | $— | $19,775 | | Jane A. Gross, Ph.D. (5) | $58,338 | $112,085 | $170,423 | | Svetlana Lucas, Ph.D. (6) | $63,044 | $112,085 | $175,129 | | Paul Schimmel, Ph.D. (7) | $40,000 | $112,085 | $152,085 | | Sara L. Zaknoen, M.D. (8) | $55,337 | $112,085 | $167,422 |

  27. Director Compensation Table—2025

    In accordance with SEC rules, this column reflects the aggregate grant date fair value of the stock options granted during 2025 computed in accordance with FASB ASC Topic 718. For additional information on the valuation assumptions underlying the value of these stock options, see Part II, Item 8 “Financial Statements and Supplementary Data” of our Annual Report in the Notes to Consolidated Financial Statements, Note 7, “Stockholders Equity”. These amounts do not reflect the actual economic value that may be realized by the directors upon the vesting of the stock options, the exercise of the stock options, or the sale of the common stock underlying such stock options. Our non-employee directors will only realize compensation to the extent the trading price of our common stock is greater than the exercise price of such stock options. Mr. Coughlin held stock options to purchase an aggregate of 95,425 shares of common stock as of December 31, 2025. (3) Mr. Benevich held stock options to purchase an aggregate of 150,000 shares of common stock as of December 31, 2025. (4) Mr. Clarke retired from the Board of Directors as of our 2025 annual meeting of stockholders. Mr. Clarke held stock options to purchase an aggregate of 45,812 shares of common stock as of December 31, 2025. (5) Dr. Gross held stock options to purchase an aggregate of 91,141 shares of common stock as of December 31, 2025. (6) Dr. Lucas held stock options to purchase an aggregate of 91,141 shares of common stock as of December 31, 2025. (7) Dr. Schimmel held stock options to purchase an aggregate of 95,812 shares of common stock as of December 31, 2025. Dr. Schimmel is retiring from the Board of Directors on May 29, 2026. Dr. Zaknoen held stock options to purchase an aggregate of 88,285 shares of common stock as of December 31, 2025.

  28. Executive Officer and Director Compensation

    Executive Officer and Director Compensation

  29. (e) Trading Policy Restrictions. Option exercises and other Awards under the Plan shall be subject to the Company’s insider trading policies and procedures, as in effect from time to time.

    (e) Trading Policy Restrictions. Option exercises and other Awards under the Plan shall be subject to the Company’s insider trading policies and procedures, as in effect from time to time.