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Proposal 1: Election of one (1) Class I director, to serve a term of three years;

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Proposal 2: Ratification of the appointment of Ernst & Young LLP as our independent registered public accounting firm for the fiscal year ending December 31, 2025;

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Proposal 3: Advisory approval of the compensation of the Company’s named executive officers, as disclosed in this Proxy Statement in accordance with SEC rules; and

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Proposal 4: Approval of a series of alternate amendments to the Company’s Amended and Restated Certificate of Incorporation to effect a reverse stock split of its common stock at a ratio in the range of 1-for-10 to 1-for-25, with such ratio to be determined in the discretion of our Board of Directors and with such reverse stock split to be effected at such time and date, if at all, as determined by our Board of Directors in its sole discretion

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To vote during the Annual Meeting, you must be present via live webcast and follow the instructions at www.virtualshareholdermeeting.com/LYEL2025. You will need to enter the 16-digit Control Number found on your proxy card.

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To vote prior to the Annual Meeting (until 11:59 p.m. Eastern Time on May 14, 2025), you may vote via the internet at www.proxyvote.com, by telephone or by completing and returning the enclosed proxy card, as described below.

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To vote through the internet prior to the Annual Meeting, go to www.proxyvote.com and follow the instructions to submit your vote on an electronic proxy card. You will be asked to provide the company number and Control Number from the proxy card. Your internet vote must be received by 11:59 p.m. Eastern Time on May 14, 2025 to be counted.

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To vote over the telephone, dial the number provided on the proxy card using a touch-tone phone and follow the recorded instructions. You will be asked to provide the company number and Control Number from your proxy card. Your telephone vote must be received by 11:59 p.m. Eastern Time on May 14, 2025 to be counted.

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To vote using the proxy card, simply complete, sign and date the proxy card and return it promptly in the envelope provided. If you return your signed proxy card to us before the Annual Meeting, we will vote your shares as you direct.

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You may request and submit another properly completed proxy card with a later date.

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You may grant a subsequent proxy by telephone or through the internet.

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You may send a timely written notice that you are revoking your proxy to our Corporate Secretary at

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You may attend the Annual Meeting and vote online. Simply attending the meeting will not, by itself, revoke your proxy.

(1)

This proposal is considered to be a “routine” matter under NYSE rules. Accordingly, if you hold your shares in street name and do not provide voting instructions to your broker, bank or other agent that holds your shares, your broker, bank or other agent has discretionary authority under NYSE rules to vote your shares on this proposal.

(1)

Member of the Audit Committee

(2)

Chair of the Compensation Committee and Lead Independent Director

(1)

Chair of the Nominating and Corporate Governance Committee

(2)

Member of the Audit Committee

(3)

Member of the Nominating and Corporate Governance Committee

(4)

Chair of the Audit Committee

(5)

Member of the Compensation Committee

*

Committee Chair

**

Lead Independent Director

(1)

In December 2024, upon the recommendation of the Nominating and Corporate Governance Committee, the Board of Directors appointed Dr. Ramachandra as a member of the Nominating and Corporate Governance Committee to replace Mr. Bishop.

(2)

Mr. Bishop resigned from the Board of Directors and the Nominating and Corporate Governance Committee in November 2024.

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helping our Board of Directors oversee our corporate accounting and financial reporting processes;

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managing the selection, engagement, qualifications, independence and performance of a qualified firm to serve as the independent registered public accounting firm to audit our financial statements;

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discussing the scope and results of the audit with the independent registered public accounting firm, and reviewing, with management and the independent accountants, our interim and year-end operating results;

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developing procedures for employees to submit concerns anonymously about questionable accounting, audit or other matters;

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reviewing related person transactions;

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obtaining and reviewing a report by the independent registered public accounting firm at least annually, that describes our internal quality control procedures, any material issues with such procedures and any steps taken to deal with such issues when required by applicable law;

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approving, or, as permitted, pre-approving, audit and permissible non-audit services to be performed by the independent registered public accounting firm;

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reviewing and assessing our risk management, risk assessment and major risk exposures with respect to financial, accounting, operational, environmental sustainability, competition and regulation, reviewing and discussing such risks with management and the auditors and reviewing the steps taken by our management to monitor, mitigate or otherwise control these exposures and identify future risks;

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overseeing and assessing our risks related to data privacy, technology and information security, including cybersecurity, and regularly reviewing with management related issues, including cybersecurity threats faced by us, and steps we are taking to address them; and

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reviewing with management our investment philosophy and policies, including management of investment risk and applicable policies pertinent to our investment portfolio.

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reviewing our overall compensation strategy, including base salary, incentive compensation and equity-based grants, to assure that it promotes stockholder interests, supports our strategic and tactical objectives and provides appropriate rewards and incentives for our management and employees;

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reviewing and approving the compensation of our chief executive officer, other executive officers and senior management and the corporate goals and objectives to be considered in such determination;

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reviewing and approving the compensation paid to our non-employee directors;

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administering our equity incentive plans and other benefit programs;

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reviewing, adopting, amending and terminating, incentive compensation and equity plans, severance agreements, profit sharing plans, bonus plans, change-of-control protections and any other compensatory arrangements for our executive officers and other senior management;

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periodically reviewing with management our major compensation-related risk exposures and steps taken to monitor or mitigate such exposures;

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overseeing periodic review of our workforce in terms of diversity and inclusion and trends on gender and racial/ethnic compensation equity;

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periodically reviewing, approving and overseeing the application of our clawback policy and any required recoupment and disclosure; and

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reviewing and approving the list of companies to be included in any compensation peer group used to determine pay levels based on criteria the Compensation Committee deems appropriate.

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evaluate our existing compensation strategy and practices in supporting and reinforcing our long-term strategic goals;

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assist in refining our compensation strategy and in developing and implementing an executive compensation program to execute that strategy; and

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ensure our compensation strategy adheres to best in market governance practices.

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identifying and evaluating candidates, including the nomination of incumbent directors for reelection, new directors to fill vacancies and nominees recommended by stockholders, to serve on our Board of Directors;

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considering and making recommendations to our Board of Directors regarding the composition and chairmanship of our Board of Directors and the committees of our Board of Directors;

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developing and instituting plans or programs for the continuing education of our Board of Directors and orientation of new directors, as necessary;

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developing and making recommendations to our Board of Directors regarding corporate governance guidelines and related principles;

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periodically reviewing and discussing with management our programs, policies and risks related to social responsibility, environmental and sustainability matters, including, but not limited to, material environmental, social and governance trends and related long- and short-term impacts on the company, and the steps we have taken to monitor or mitigate such exposures;

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overseeing our quality, corporate and healthcare compliance programs and periodically reviewing and discussing with management our programs, policies and risks related to such matters, including, but not limited to, material trends and related long- and short-term impacts on the Company, and the steps we have taken to monitor or mitigate such exposures;

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periodically reviewing with our Chief Executive Officer the plans for succession to the offices of our Chief Executive Officer and other key executive officers;

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overseeing periodic evaluations of the Board of Directors’ performance, including committees of the Board of Directors, and review of the committee charters; and

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annually reviewing any stockholder proposals submitted for inclusion in our proxy statement and recommending to the Board any statements in response and considering any stockholder nominees for election to the Board at our annual meeting of stockholders.

(1)

As of December 31, 2024, our non-employee directors held options to purchase shares of our common stock as set forth below:

(2)

All of the option awards were granted under our 2021 Equity Incentive Plan. The amounts shown represent the grant date fair values of option awards granted in 2024 as computed in accordance with Financial Accounting Standards Board Accounting Standards Codification, Topic 718 (“FASB ASC Topic 718”). See Note 14, Stock-based Compensation, to our audited consolidated financial statements included in Part II, Item 8 of our Annual Report on Form 10-K for the year ended December 31, 2024 for a discussion of the assumptions used in the calculation.

(3)

The amount shown corresponds to (i) for each of our directors except Dr. Ramachandra, the annual grant of an option to purchase 130,000 shares of our common stock on June 14, 2024 and (ii) for Dr. Ramachandra, an initial option grant to purchase 260,000 shares of our common stock granted on October 31, 2024, each pursuant to the then-current Non-Employee Director Compensation Policy.

(4)

Dr. Ramachandra joined the Board of Directors in October 2024 and was appointed to the Nominating and Corporate Governance Committee in December 2024, and his fees were pro-rated.

(5)

Mr. Bishop resigned from the Board of Directors in November 2024.

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an annual cash retainer of $50,000 (same as under the Prior Policy) for all non-employee directors other than the lead director/Chair of our Board of Directors;

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an annual cash retainer of $80,000 (same as under the Prior Policy) for the lead independent director of our Board of Directors;

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an annual cash retainer of $85,000 (was $80,000 under the Prior Policy) for the Chair of our Board of Directors;

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an additional annual cash retainer of $20,000, $15,000, and $10,000 (was $15,000, $12,000 and $10,000, respectively, under the Prior Policy) for service as Chair of the Audit Committee, Compensation Committee and the Nominating and Corporate Governance Committee, respectively;

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an additional annual cash retainer of $10,000, $7,500 and $5,000 (was $7,500, $6,000 and $5,000, respectively, under the Prior Policy) for service as a member of the Audit Committee, Compensation Committee and the Nominating and Corporate Governance Committee, respectively (other than for as the chair for any such committee);

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an appointment option grant, upon the date a new non-employee director is first elected or appointed to our Board of Directors, to purchase the lesser of (i) shares of common stock calculated to have a Black-Scholes value of $500,000 on the date of grant, rounded to the nearest whole number; and (ii) 260,000 shares of common stock, with such grant vesting in 36 equal monthly installments measured from the date the non-employee director is first elected or appointed to our Board of Directors, subject to the non-employee director’s continued service on each applicable vesting date; and

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an annual option grant, upon the date of each of our annual meeting of stockholders, to purchase the lesser of (i) shares of common stock calculated to have a Black-Scholes value of $300,000 on the date of grant, rounded to the nearest whole number; and (ii) 130,000 shares of common stock, all of which shares vest on the earlier of (a) the date of the next annual meeting (or the date immediately prior to such date if the non-employee director’s service as a director ends at such annual meeting due to the director’s failure to be re-elected or the director not standing for re-election) or (b) the first anniversary of the date of grant, in each case subject to the non-employee director’s continued service on each applicable vesting date.

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purchasing financial instruments, or otherwise engaging in transactions, that hedge or offset, or are designed to hedge or offset, any decrease in the market value of our common stock, such as prepaid variable forward contracts, equity swaps, collars, forward sale contracts, and exchange funds;

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purchasing our common stock on margin or holding it in a margin account at any time;

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pledging our common stock as collateral for a personal loan; and

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engaging in short sales, transactions in put options, call options or other derivative securities on an exchange or in any other organized market, or in any other inherently speculative transactions with respect to our common stock.

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effect a reverse stock split (“Reverse Stock Split”) of all issued and outstanding shares of our common stock, at a ratio ranging from 1-for-10 to 1-for-25, inclusive.

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Stock Price Requirements: We understand that many brokerage houses, institutional investors and funds have internal policies and practices that either prohibit them from investing in stocks trading below a certain price point or tend to discourage individual brokers from recommending such stocks to their customers or by restricting or limiting the ability to purchase such stocks on margin. Additionally, a Reverse Stock Split could help increase analyst and broker interest in our common stock as their internal policies might discourage them from following or recommending companies with low stock prices.

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Stock Price Volatility: Because of the trading volatility often associated with stocks trading below a certain price point, many brokerage houses and institutional investors have internal policies and practices that either prohibit them from investing in such stocks or tend to discourage individual brokers from recommending such stocks to their customers. Some of those policies and practices may also make the processing of trades in stocks trading below a certain price point economically unattractive to brokers.

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Transaction Costs: Investors may be dissuaded from purchasing stocks trading below certain prices because brokers’ commissions, as a percentage of the total transaction value, can be higher for such stocks.

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the historical trading price and trading volume of our common stock;

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the then-prevailing trading price and trading volume of our common stock and the expected impact of the Reverse Stock Split on the trading market for our common stock in the short- and long-term;

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our ability to maintain our listing on The Nasdaq Stock Market;

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which Reverse Stock Split ratio would result in the least administrative cost to us;

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prevailing general market and economic conditions; and

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whether and when our Board of Directors desires to have the additional authorized but unissued shares of common stock that will result from the implementation of a Reverse Stock Split available to provide the flexibility to use our common stock for business and/or financial purposes, as well as to accommodate the shares of our common stock to be authorized and reserved for future equity awards.

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depending on the Reverse Stock Split ratio selected by our Board of Directors, each ten to twenty-five shares of our common stock owned by a stockholder will be combined into one new share of our common stock;

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no fractional shares of common stock will be issued in connection with any Reverse Stock Split; instead, holders of common stock who would otherwise receive a fractional share of common stock pursuant to the Reverse Stock Split will receive cash in lieu of the fractional share as explained more fully below;

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the total number of authorized shares of our common stock will not be reduced proportionally to the Reverse Stock Split and, therefore, the Reverse Stock Split will result in an effective increase in the authorized number of shares of our common stock;

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based upon the Reverse Stock Split ratio selected by our Board of Directors, proportionate adjustments will be made to the per share exercise price and/or the number of shares issuable upon the exercise or vesting of all then outstanding stock options, restricted stock units and warrants (if any), which will result in a proportional decrease in the number of shares of our common stock reserved for issuance upon exercise or vesting of such stock options, restricted stock units and warrants, and, in the case of stock options and warrants, a proportional increase in the exercise price of all such stock options and warrants; and

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the number of shares then reserved for issuance under our equity compensation plans will be reduced proportionately based upon the Reverse Stock Split ratio selected by the Board.

(1)

Represents all shares reserved for issuance under our equity plans, including shares reserved for issuance pursuant to outstanding awards and shares available for future grant under such plans, as well as the milestone shares that may become issuable to the former of stockholders of ImmPACT pursuant to the Agreement and Plan of Merger between us and ImmPACT, dated October 24, 2024. Shares reserved for future issuance exclude any shares issuable pursuant to success payment obligations under our research and collaboration agreements with the Fred Hutchinson Cancer Center and The Board of Trustees of the Leland Stanford Junior University, each as more fully described under Note 4, License, Collaboration and Success Payment Agreements, to our audited consolidated financial statements included in Part II, Item 8 of our Annual Report on Form 10-K for the fiscal year ended December 31, 2024.

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an individual who is a citizen or resident of the United States or someone treated as a U.S. citizen or resident for U.S. federal income tax purposes;

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a corporation (or other entity treated as a corporation for U.S. federal income tax purposes) created or organized in or under the laws of the United States, any state thereof, or the District of Columbia;

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an estate, the income of which is subject to U.S. federal income taxation regardless of its source; or

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a trust if (i) a U.S. court can exercise primary supervision over the trust’s administration and one or more “United States persons” (within the meaning of Section 7701(a)(30) of the Code) are authorized or have the authority to control all substantial decisions of the trust or (ii) it has a valid election in effect under applicable U.S. Treasury Regulations to be treated as a United States person for U.S. federal income tax purposes.

*

Represents beneficial ownership of less than 1%.

(1)

Based on information set forth in a Schedule 13G/A filed with the SEC on February 14, 2025 by ARCH Venture Fund IX, L.P. (“AVF IX”); ARCH Venture Partners IX, L.P. (“AVP IX LP”); ARCH Venture Partners IX, LLC (“AVP IX LLC”); ARCH Venture Fund IX Overage, L.P. (“AVF IX Overage”); ARCH Venture Partners IX Overage, L.P. (“AVF IX Overage GP”) (collectively, the “Reporting Entities”); and Keith Crandell, Clinton Bybee and Robert Nelsen (collectively, the “AVP IX Managing Directors” and individually, each an “AVP IX Managing Director”). The Reporting Entities and the AVP IX Managing Directors collectively are referred to as the “Reporting Persons.” AVF IX is the record owner of 18,206,358 shares of common stock (the “AVF IX Shares”) as of December 31, 2024. AVP IX LP, as the sole general partner of AVF IX, may be deemed to beneficially own the AVF IX Shares. AVF IX Overage is the record owner of 18,206,358 shares of common stock (the “AVF IX Overage Shares”; combined with AVF IX Shares, the “Record Shares”) as of December 31, 2024. AVF IX Overage GP, as the sole general partner of AVF IX Overage, may be deemed to beneficially own the AVF IX Overage Shares. AVP IX LLC, as the sole general partner of AVP IX LP and AVF IX Overage LP, may be deemed to beneficially own the Record Shares. As managing directors of AVP IX LLC, each AVP IX Managing Director may also be deemed to share the power to direct the disposition and vote of the Record Shares. Each Reporting Person disclaims beneficial ownership of the Record Shares except for the

(2)

Based solely on information set forth in a Schedule 13G filed with the SEC on February 10, 2022 by GlaxoSmithKline plc (“GSK”). Represents 30,253,189 shares of common stock, of which GSK has sole voting and dispositive power through its indirect wholly-owned subsidiary, Glaxo Group Limited (“GGL”). The Schedule 13G filed by GSK provides information as of December 31, 2021 and, consequently, the beneficial ownership of GSK may have changed between December 31, 2021 and March 31, 2025. The mailing address of each of GSK and GGL is 980 Great West Road, Brentford, Middlesex, TW8 9GS, United Kingdom.

(3)

Based solely on information set forth in a Schedule 13G filed with the SEC on February 14, 2022 by Milky Way Investments Group Limited (“Milky Way”) and MWG Management Limited. Milky Way is controlled by MWG Management Limited, its corporate director. The Schedule 13G filed by the reporting entities provides information as of December 31, 2021 and, consequently, the beneficial ownership of the reporting entities may have changed between December 31, 2021 and March 31, 2025. The mailing address of Milky Way is c/o Trident Trust Company (B.V.I.) Limited, Trident Chambers, P.O. Box 146, Road Town, Tortola, British Virgin Islands.

(4)

Based solely on information set forth in a Schedule 13G filed with the SEC on February 14, 2025 by Foresite Capital Fund IV, L.P. (“Foresite IV”), Foresite Capital Management IV, LLC (“FCM IV”), Foresite Capital Fund V, L.P. (“Foresite V”), Foresite Capital Management V, LLC (“FCM V”), Foresite Capital Opportunity Fund V, L.P. (“Foresite Opp V”), Foresite Capital Opportunity Management V, LLC (“FCM Opp V”) and James Tananbaum (“Tananbaum”). Foresite IV is the record owner of 8,325,000 shares of common stock (the “Foresite IV Shares”) as of December 31, 2024, over which it has sole voting power and sole dispositive power, except that FCM IV, as the general partner of Foresite IV, may be deemed to have sole power to vote and dispose of these shares, and Tananbaum, the managing member of FCM IV, may be deemed to have sole power to vote and dispose of these shares. Foresite V is the record owner of 8,133,047 shares of common stock (the “Foresite V Shares”) as of December 31, 2024, over which it has sole voting power and sole dispositive power, except that FCM V, as the general partner of Foresite V, may be deemed to have sole power to vote and dispose of these shares, and Tananbaum, the managing member of FCM V, may be deemed to have sole power to vote and dispose of these shares. Foresite Opp V is the record owner of 3,020,846 shares of common stock (the “Foresite Opp V Shares”) as of December 31, 2024, over which it has sole voting power and sole dispositive power, except that FCM Opp V, as the general partner of Foresite Opp V, may be deemed to have sole power to vote and dispose of these shares, and Tananbaum, the managing member of FCM Opp V, may be deemed to have sole power to vote and dispose of these shares. The Schedule 13G filed by the reporting entities provides information as of December 31, 2024 and, consequently, the beneficial ownership of the reporting person may have changed between December 31, 2024 and March 31, 2025. The mailing address of Foresite Capital is c/o Foresite Capital Management, 900 Larkspur Landing Circle, Suite 150, Larkspur, CA 94939.

(5)

Based solely on information set forth in a Schedule 13G filed with the SEC on February 14, 2022 by Gemini Investments, L.P. (“Gemini Investments”), Gemini GP Limited (“Gemini GP”), Orland Properties Limited (“Orland Properties”) and Newton (PTC) Limited (“Newton”). Gemini GP is the general partner of Gemini Investments, and as such, may be deemed to beneficially own the shares held directly by Gemini Investments. Orland Properties wholly owns Gemini GP, and as such, may be deemed to beneficially own the shares beneficially owned by Gemini GP. Newton wholly owns Orland Properties, and as such, may be deemed to beneficially own the shares beneficially owned by Orland Properties. The Schedule 13G filed by the reporting entities provides information as of December 31, 2021 and, consequently, the beneficial ownership of the reporting entities may have changed between December 31, 2021 and March 31, 2025. The mailing address of Gemini Investments is c/o Trident Trust Company (Cayman) Limited, One Capital Place, P.O. Box 847, Grand Cayman, KY1-1103, Cayman Islands.

(6)

Consists of 175,000 shares of common stock and 4,090,000 shares of common stock issuable upon exercise of stock options held by Dr. Seely that are exercisable within 60 days of March 31, 2025.

(7)

Consists of 20,399 shares of common stock and 2,361,779 shares of common stock issuable upon exercise of stock options held by Mr. Hill that are exercisable within 60 days of March 31, 2025.

(8)

Consists of 1,187,500 shares of common stock issuable upon exercise of stock options held by Mr. Lang that are exercisable within 60 days of March 31, 2025. In March 2025, Mr. Lang resigned as our Chief Business Officer and from all other offices of the Company, effective April 14, 2025. Mr. Lang was one of our two other most highly compensated executive officers as of December 31, 2024. As a result, he is one of our “named executive officers” for this Proxy Statement.

(9)

Consists of (i) 3,811,199 shares of common stock, of which 3,125,834 shares of common stock are held by Dr. Klausner, and 245,515 shares are held by each of The Ariella Klausner Delaware Trust and The Olivia Klausner Delaware Trust, and 194,335 shares are held by The Eli Klausner Delaware Trust (collectively, the “Klausner Trusts”); and (ii) 8,354,148 shares of common stock issuable upon exercise of stock options held by Dr. Klausner that are exercisable within 60 days of March 31, 2025. Dr. Klausner is a grantor of each of the Klausner Trusts and therefore may be deemed to share the power to direct the disposition and vote of the shares held by the trusts. Dr. Klausner disclaims beneficial ownership of all shares held by the Klausner Trusts, except to any pecuniary interest therein, if any, other than for purposes of determining his obligations under Section 13 of the Exchange Act. The JTC Trust Company (Delaware) is the trustee of each of the Klausner Trusts.

(10)

Consists of 35,640 shares of common stock and 715,000 shares of common stock issuable upon exercise of stock options held by Dr. Brawley that are exercisable within 60 days of March 31, 2025.

(11)

Consists of (i) 965,000 shares of common stock issuable upon exercise of stock options held by Ms. Friedman that are exercisable within 60 days of March 31, 2025, (ii) 100,000 shares of common stock held by The Duane Irrevocable Trust 2020 (“Duane Trust”) and (iii) 236,370 shares of common stock held by the Duane Family Trust (“Duane Family Trust”). Ms. Friedman is a trustee of the Duane Trust and the Duane Family Trust and therefore may be deemed to share the power to direct the disposition and vote of the shares held by the Duane Trust and/or the Duane Family Trust. Ms. Friedman disclaims beneficial ownership of all shares held by the Duane Trust and the Duane Family Trust, except to any pecuniary interest therein, other than for purposes of determining her obligations under Section 13 of the Exchange Act.

(12)

Consists of 715,000 shares of common stock issuable upon exercise of stock options held by Dr. Nabel that are exercisable within 60 days of March 31, 2025.

(13)

Includes 315,000 shares of common stock issuable upon exercise of stock options held by Mr. Nelsen that are exercisable within 60 days of March 31, 2025. Mr. Nelsen is an AVP IX Managing Director and may be deemed to beneficially own the shares held by AVF IX and AVF IX Overage as discussed in footnote 1. Mr. Nelsen disclaims beneficial ownership of such shares except to the extent of his pecuniary interest therein, if any, other than for purposes of determining his obligations under Section 13 of the Exchange Act.

(14)

Consists of 200,000 shares of common stock held by the Sumant Ramachandra Revocable Trust DTD 01/24/12, for which Dr. Ramachandra is grantor and trustee, and 43,333 shares of common stock issuable upon exercise of stock options held by Dr. Ramachandra that are exercisable within 60 days of March 31, 2025.

(15)

Consists of 715,000 shares of common stock issuable upon exercise of stock options held by Mr. Rieflin that are exercisable within 60 days of March 31, 2025.

(16)

Consists of (i) 41,235,745 shares of common stock held by our current directors and executive officers as a group and (ii) 25,185,559 shares of common stock issuable upon exercise of stock options held by our current directors and executive officers that are exercisable within 60 days of March 31, 2025.

(1)

The amounts represent the aggregate grant date fair value, as computed in accordance with FASB ASC Topic 718, of awards of PSUs, granted to our named executive officers, which are subject to performance conditions and market conditions. The grant date fair values of the PSUs subject to performance conditions are based on the probable outcome of such conditions and given the Company determined that it was not probable at grant that any of the performance conditions applicable to the PSUs would be met, the grant date fair values reported in the stock awards column for all of the PSUs subject to performance conditions is $0. The value of the PSUs subject to performance conditions at the grant date assuming that the highest level of performance conditions would be achieved is $2,160,000 for Dr. Seely, $432,000 for Mr. Hill and $540,000 for Mr. Lang. The grant date fair values of PSUs subject to the market conditions are estimated using Monte Carlo simulations, accounting for the maximum percentage of the PSUs subject to the market conditions being earned. See Note 14, Stock-based Compensation, to our audited consolidated financial statements included in Part II, Item 8 of our Annual Report on Form 10-K for the fiscal year ended December 31, 2024 for a discussion of the assumptions used in the calculation. These amounts do not necessarily correspond to the actual value recognized or that may be recognized by the named executive officers.

(2)

Except as otherwise noted below, the amounts shown represent the sum of (a) the grant date fair values of option awards granted in 2024 and 2023, as computed in accordance with FASB ASC Topic 718 and (b) for 2023, the incremental fair values of the option awards resulting from the one-time repricing of certain stock options that had been granted under the 2021 Plan and 2018 Equity Incentive Plan (“2018 Plan”), approved by our Board of Directors in November 2023 (the “Repricing”). The Repricing impacted stock options with exercise prices greater than $2.37, and each such option was repriced to have a per share exercise price of $1.87, the closing price of our common stock on the repricing date, November 16, 2023. To be eligible to exercise the option at the new exercise price, option holders were required to remain employed with us through November 15, 2024. Any exercise of a repriced option prior to that date would have required payment of the original, higher exercise price. Additionally, as a condition of receiving the Repricing, the vesting schedule for the unvested shares underlying repriced stock options held by executives at the level of senior vice president and above, including our named executive officers, was extended for an additional year. No changes were made to the expiration dates of or number of shares underlying the repriced stock options. See Note 14, Stock-based Compensation, to our audited consolidated financial statements included in Part II, Item 8 of our Annual Report on Form 10-K for the year ended December 31, 2024 for a discussion of the assumptions used in the calculation. These amounts do not necessarily correspond to the actual value recognized or that may be recognized by the named executive officers.

(3)

The amounts shown represent the annual performance-based cash bonus earned by our named executive officers based on the achievement of certain corporate performance objectives during 2024 and 2023. For more information about the 2024 annual incentive bonus program, see the subsection below titled “Narrative to the Summary Compensation Table — Annual Performance Bonus – Non-Equity Incentive Plan Compensation.” These amounts were paid in early 2025 and 2024, respectively.

(4)

For Dr. Seely, the amount shown represents $16,586 of life insurance premiums paid by us on her behalf, which includes $5,918 for associated taxes, $4,750 in 401(k) matching contributions and $78 in other compensation.

(5)

For Mr. Hill, the amount shown represents $2,554 of life insurance premiums paid by us on his behalf, which includes $622 for associated taxes and $4,750 in 401(k) matching contributions.

(6)

Mr. Lang joined the Company in July 2023 and his 2023 salary amount was pro-rated for the period of time he was employed in 2023. In March 2025, Mr. Lang resigned as our Chief Business Officer and from all other offices of the Company, effective April 14, 2025.

(7)

The amount shown represents an advance sign-on bonus paid in 2023 pursuant to Mr. Lang’s offer letter and earned in 2024 upon his completion of one year of employment with us. See the subsection below titled “Employment Contracts and Change in Control Arrangements – Offer Letters – Matthew Lang, J.D.” for more information.

(8)

For Mr. Lang, the amount shown represents $1,959 of life insurance premiums paid by us on his behalf, which includes $699 for associated taxes, and $4,750 in 401(k) matching contributions.

(1)

The 2024 base salaries of the named executive officers became effective on March 1, 2024.

(2)

The 2023 base salaries of the named executive officers became effective on March 1, 2023 except for Mr. Lang’s 2023 base salary, which became effective on July 1, 2023, when he commenced employment with us as Chief Business Officer.

(3)

In March 2025, Mr. Lang resigned as our Chief Business Officer and from all other offices of the Company, effective April 14, 2025.

(1)

In March 2025, Mr. Lang resigned as our Chief Business Officer and from all other offices of the Company, effective April 14, 2025.

(1)

Represents the 2024 option grant made to the applicable named executive officer. Each option vests as to 12.5% of the total number of shares subject to the option six months after the vesting commencement date of February 9, 2024, and as to 1/48th of the total number of shares subject to the option each month thereafter on the same day of the month as the vesting commencement (or if there is no corresponding day, on the last day of the month), subject to the applicable named executive officer’s continued service to the Company through the applicable vesting date.

(2)

The options are subject to vesting acceleration, as described in more detail below under the section titled “Employment Contracts and Change in Control Arrangements.”

(3)

Represents the number of shares of our common stock subject to the 2024 PSU grants made to the applicable named executive officer that could be earned, assuming that the applicable performance conditions to which certain of the PSUs are subject will be achieved, and that the market conditions to which the remaining PSUs are subject will be achieved at the target level. If the PSUs subject to market conditions were achieved at the maximum level, the applicable named executive officer could earn an additional number of shares of our common stock with respect to such PSUs equal to 200,000, 40,000 and 50,000 shares of our common stock for Dr. Seely, Mr. Hill and Mr. Lang, respectively.

(4)

In March 2025, Mr. Lang resigned as our Chief Business Officer and from all other offices of the Company, effective April 14, 2025.

(1)

All of the option awards with grant dates prior to June 16, 2021 were granted under the 2018 Plan. Option awards with grant dates on or after June 16, 2021 were granted under the 2021 Plan.

(2)

Represents the number of PSUs subject to performance-based conditions and the threshold number of PSUs subject to market-based conditions that may vest upon or after, as applicable, the Compensation Committee’s certification of the attainment of the applicable performance and market conditions. The PSUs granted to the named executive officers that are subject to market conditions vest based upon the Company’s performance against two- and three-year relative total shareholder return goals (with such PSUs allocated equally between the performance periods), and the PSUs subject to performance conditions vest upon the achievement of certain clinical development milestones. With respect to PSUs subject to clinical development milestones, (a) two-thirds of such PSUs vest as follows: 50% vest upon certification of the achievement of the applicable milestone by our Compensation Committee, and the remaining 50% vest upon the earlier of (i) one year of continuous service by the applicable named executive officer from the date of certification of such achievement by our Compensation Committee and (ii) the end of the three-year performance period; and (b) the remaining one-third of such PSUs vest upon certification of the applicable clinical milestone by our Compensation Committee. The vesting of all PSU awards granted is also subject to the respective named executive officer’s continued service through each applicable vesting date.

(3)

The dollar amounts shown are determined by multiplying (x) the number of PSUs reported in the preceding column by (y) $0.64, the closing price of the Company’s common stock on December 31, 2024, the last trading day of 2024. These amounts do not necessarily correspond to the actual value recognized or that may be recognized by the named executive officers.

(4)

The option awards granted to Dr. Seely on May 20, 2021 and June 8, 2022 were in connection with her service as a member of our Board of Directors and, as such, their exercise prices were not eligible for the Repricing. The option award granted on December 15, 2022 was in connection with her commencement of employment with us as our President and Chief Executive Officer.

(5)

The option is early-exercisable, meaning that it can be exercised before it vests for shares of our common stock, subject to a repurchase right in favor of the Company that lapses in accordance with the same vesting provisions as the underlying option. Accordingly, the number of shares shown for the option in this column represents the number of shares that are early-exercisable, regardless of vesting status.

(6)

The option is early-exercisable, meaning that it can be exercised before it vests for shares of our common stock, subject to a repurchase right in favor of the Company that lapses in accordance with the same vesting provisions as the underlying option. Accordingly, there are no shares shown for the option in this column as all shares underlying the options are early-exercisable, regardless of vesting status.

(7)

The option vests as to 1/36th of the shares initially underlying the option each month until fully vested on the third anniversary of the vesting commencement date, subject to continued service to us through the applicable vesting date.

(8)

The option fully vests as to 100% of the shares initially underlying the option on the first anniversary of the vesting commencement date, subject to continued service to us through the applicable vesting date.

(9)

Pursuant to the Repricing, the exercise price of the repriced options has been modified to be $1.87 per share, the closing price of our common stock on the repricing date, November 16, 2023. To be eligible to exercise the option at the new exercise price, option holders were required to remain employed with us through November 15, 2024. All vested shares underlying the repriced options as of the repricing date remained vested. Pursuant to the Repricing, the vesting schedules for the unvested shares underlying repriced options held by the named executive

(10)

Under the original vesting schedule in effect prior to the Repricing, the options vest as to 25% of the shares initially underlying the option on the first anniversary of the vesting commencement date and as to 1/48th of the shares initially underlying the option each month until fully vested on the fourth anniversary of the vesting commencement date, subject to continued service to us through the applicable vesting date. The original vesting schedule has since been modified pursuant to the Repricing, as described under footnote 9 above.

(11)

Under the original vesting schedule in effect prior to the Repricing, the option vests as to 1/48th of the shares initially underlying the option each month until fully vested on the fourth anniversary of the vesting commencement date, subject to continued service to us through the applicable vesting date. The original vesting schedule has since been modified pursuant to the Repricing, as described under footnote 9 above.

(12)

Under the original vesting schedule in effect prior to the Repricing, the option vests as to 12.5% of the shares initially underlying the option on the date that is six (6) months after the vesting commencement date and as to 1/48th of the shares initially underlying the option each month thereafter until fully vested on the fourth anniversary of the vesting commencement date, subject to continued service to us through the applicable vesting date. The original vesting schedule has since been modified pursuant to the Repricing, as described under footnote 9 above.

(13)

The option vests as to 12.5% of the shares initially underlying the option on the date that is six (6) months after the vesting commencement date and as to 1/48th of the shares initially underlying the option each month thereafter until fully vested on the fourth anniversary of the vesting commencement date, subject to continued service to us through the applicable vesting date.

(14)

In March 2025, Mr. Lang resigned as our Chief Business Officer and from all other offices of the Company, effective April 14, 2025.

(15)

The option vests as to 25% of the shares initially underlying the option on the first anniversary of the vesting commencement date and as to 1/48th of the shares initially underlying the option each month until fully vested on the fourth anniversary of the vesting commencement date, subject to continued service to us through the applicable vesting date.