Document:

EX-10.11

 Exhibit 10.11 

EXECUTION VERSION 

REGISTRATION AND VOTING RIGHTS AGREEMENT 

This Registration and Voting Rights Agreement (the “Agreement”) is made and entered into as of March 31, 2015
among aTyr Pharma, Inc., a Delaware corporation (the “Corporation”), and the Holders named in Schedule A hereto. 

R E C I T A L S 

WHEREAS, the Corporation, the holders of Series A Preferred Stock, the holders of Series B Preferred Stock, the holders of Series B-2
Preferred Stock, the holders of Series C Preferred Stock and the holders of Series D Preferred Stock are parties to that certain Fourth Amended and Restated Registration Rights Agreement, dated as of April 8, 2013 (the “Prior
Registration Rights Agreement”); and  
 WHEREAS, concurrently herewith, the Corporation and the Holders (as herein
defined) are entering into a Stock Purchase Agreement (as amended from time to time and including all exhibits, attachments and appendices thereto, the “Series E Purchase Agreement”). The execution and delivery of this Agreement is
a condition precedent to the Holders’ obligations under the Series E Purchase Agreement. 
 NOW, THEREFORE, the parties to this
Agreement hereby agree as follows: 
 ARTICLE I 

DEFINITIONS 
 Unless
otherwise defined herein, capitalized terms used herein and not defined shall have the same meaning as provided in the Purchase Agreement. 

In addition, the following terms shall have the meanings set forth in this Article I: 

“Affiliate” means, with respect to any person, any other person who or which, directly or indirectly, controls, is controlled
by, or is under common control with such specified person, including, without limitation, any direct or indirect subsidiary of such person that is at least 50% controlled by such person, general partner, officer, director or manager of such person
and any venture capital or other investment fund now or hereafter existing that is controlled by one or more general partners or managing members of, or is under common investment management with, such person and any mutual fund, pension fund,
pooled investment vehicle or institutional separate account advised by the same or affiliated registered investment advisor. 

“Agreement” has the meaning given such term in the Recitals. 

“Baker” means Baker Brothers Life Sciences, L.P. and 667, L.P. 

“Capital Securities” means, as to any Person that is a corporation, the authorized shares of such Person’s capital
stock, including all classes of common, preferred, voting and nonvoting capital stock, and, as to any Person that is not a corporation or an individual, the ownership 

 
interests in such Person, including, without limitation, the right to share in profits and losses, the right to receive distributions of cash and property, and the right to receive allocations of
items of income, gain, loss, deduction and credit and similar items from such Person, whether or not such interests include voting or similar rights entitling the holder thereof to exercise control over such Person. 

“Commission” means the U.S. Securities and Exchange Commission or any successor governmental agency that administers the
Securities Act and the Exchange Act. 
 “Commission Form S-3” has the meaning specified in Section 2.1(b). 

“Common Stock” means the Common Stock, par value $.001 per share, of the Corporation, as constituted on the date hereof,
any shares of the Corporation’s capital stock into which such Common Stock shall be changed, and any shares of the Corporation’s capital stock resulting from any reclassification of such Common Stock or recapitalization of the Corporation.

 “Common Stock Equivalents” means any options, warrants or other securities or rights convertible into or exercisable or
exchangeable for, whether directly or following conversion into or exercise or exchange for other options, warrants or other securities or rights, shares of Common Stock. 

“Conversion Stock” means Common Stock issued or issuable upon the conversion of shares of the Corporation’s Preferred
Stock. 
 “Exchange Act” means the Securities Exchange Act of 1934, as amended, or any successor statute thereto, and
the rules and regulations of the Commission promulgated from time to time thereunder, all as the same shall be in effect at the time. 

“Holders” means, collectively, (i) the Persons named in Schedule A hereto, (ii) any other Person
holding Registrable Securities to whom any such Person assigns the registration rights contemplated hereby pursuant to Article VII of this Agreement and in the case of (i) or (ii) provided such Person signs a counterpart to this Agreement.

 “Incidental Registration” has the meaning specified in Section 2.2. 

“Incidental Registration Cutback” has the meaning specified in Section 2.2(b) of this Agreement. 

“Indemnified Parties” has the meaning specified in Section 5.1 of this Agreement. 

“Indemnifying Party” has the meaning specified in Section 5.1 of this Agreement. 

“Liquidation Event” means (i) any liquidation, dissolution or winding up of the Corporation, whether voluntary or
involuntary, (ii) a merger or consolidation of the Corporation into or with any other Person or Persons in a single transaction or a series of related transactions in which the stockholders of the Corporation immediately prior to such merger,
consolidation, transaction or first of such related series of transaction possess less than fifty percent (50%) of the surviving entity’s issued and outstanding voting Capital Securities immediately after such 

  
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merger, consolidation, transaction or related series of such transactions, (iii) a transaction or series of related transactions in which a Person, entity or group unrelated to the
Corporation acquires Capital Securities representing more than 50% of the outstanding voting power of the Corporation, or (iv) a sale of all or substantially all of the Corporation’s assets to any Person (including indirectly by the grant
of an exclusive license or licenses to all or substantially all of the Corporation’s intellectual property). Notwithstanding the foregoing, the Corporation’s initial public offering of equity securities or any conversion of Preferred
Stock to Common Stock pursuant to the Corporation’s Amended and Restated Certificate of Incorporation as then in effect, after which the stockholders of the Corporation as of immediately prior to such event continue to maintain an ownership
interest in the Corporation as of immediately after such event, or the sale by the Corporation of Capital Securities for bona fide capital raising purposes, shall not be a “Liquidation Event”. 

“Participating Investor” means Sofinnova and Baker. 

“Person” or “person” shall mean an individual, partnership, corporation, limited liability company,
association, trust, joint venture, unincorporated organization and any government, governmental department or agency or political subdivision thereof. 

“Preferred Stock” means, collectively, shares of the Series A Preferred Stock, Series B Preferred Stock, Series B-2
Preferred Stock, Series C Preferred Stock, Series D Preferred Stock and Series E Preferred Stock. 
 “Registrable
Securities” means the following (in each case as adjusted for stock splits, recapitalizations and other similar events): (i) the Conversion Stock; (ii) any Common Stock or other securities issued or issuable on conversion of or
otherwise with respect to the Conversion Stock (including pursuant to any stock split, stock dividend, recapitalization, or similar event); (iii) any shares of Common Stock acquired by any Holder after the date hereof including upon the
exercise of warrants, and (iv) securities issued in replacement or exchange of any Conversion Stock or securities issued in clauses (i), (ii) or (iii) above; provided, however, that any and all shares described in clauses
(i)-(iv) above shall cease to be Registrable Securities upon their sale pursuant to a registration statement under the Securities Act. 

“Registration Expenses” means all expenses incident to the Corporation’s performance of or compliance with this
Agreement in connection with each Requested Registration or Incidental Registration, including, without limitation, all registration, filing, listing and Financial Industry Regulatory Authority (“FINRA”) fees, all fees and expenses
of complying with securities or blue sky laws, all word processing, duplicating and printing expenses, all messenger and delivery expenses, any transfer taxes, the fees and expenses of the Corporation’s legal counsel and independent public
accountants, the reasonable fees and disbursements up to a maximum of $75,000 of one counsel for all Holders participating in each such registration, and any fees and disbursements of underwriters customarily paid by issuers or sellers of
securities; provided, however, that Registration Expenses shall not include underwriting discounts and commissions. 

“Requested Registration” has the meaning specified in Section 2.1(b) of this Agreement. 

  
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 “Requested Registration Cutback” has the meaning specified in
Section 2.1(c) of this Agreement. 
 “Required Amount” means, as to a Participating Investor, at least 50% of the
shares of Series E Preferred Stock purchased by such Participating Investor in connection with the Initial Issuance (as adjusted for stock splits, recapitalizations and other similar events), or at least 50% of the shares of Common Stock issued upon
conversion of such shares of Series E Preferred Stock (as adjusted for stock splits, recapitalizations and other similar events). 

“Requisite Stockholders” means the holders of a majority of the issued and outstanding shares of Series B-2 Preferred Stock,
Series C Preferred Stock, Series D Preferred Stock, Series E Preferred Stock and any Common Stock issued upon conversion of such shares of Preferred Stock, voting together as a single class on an as-if-converted to Common Stock basis. 

“S-1 Registration” has the meaning specified in Section 2.1(a) of this Agreement. 

“S-1 Registration Notice” has the meaning specified in Section 2.1(a) of this Agreement. 

“S-1 Registration Request” has the meaning specified in Section 2.1(a) of this Agreement. 

“S-3 Registration” has the meaning specified in Section 2.1(b) of this Agreement. 

“S-3 Registration Notice” has the meaning specified in Section 2.1(b) of this Agreement. 

“S-3 Registration Request” has the meaning specified in Section 2.1(b) of this Agreement. 

“Securities Act” means the Securities Act of 1933, as amended, or any successor statute thereto, and the rules and
regulations of the Commission promulgated from time to time thereunder, all as the same shall be in effect at the time. 
 “Series A
Preferred Stock” shall mean the Series A Preferred Stock of the Corporation, par value $.001 per share. 
 “Series B
Preferred Stock” shall mean the Series B Preferred Stock of the Corporation, par value $.001 per share. 
 “Series B-2
Preferred Stock” shall mean the Series B-2 Preferred Stock of the Corporation, par value $.001 per share. 
 “Series C
Preferred Stock” shall mean the Series C Preferred Stock of the Corporation, par value $.001 per share. 
 “Series D
Preferred Stock” shall mean the Series D Preferred Stock of the Corporation, par value $.001 per share. 

  
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 “Series E Preferred Stock” shall mean the Series E Preferred Stock of the
Corporation, par value $.001 per share. 
 “Sofinnova” means Sofinnova Venture Partners IX, L.P. 

“Subscription Agreements” has the meaning specified in the Recitals hereto. 

“Underwriter’s Maximum Number” has the meaning specified in Section 2.1(c) of this Agreement. 

ARTICLE II 

REGISTRATIONS 

SECTION 2.1. Requested Registrations. 

(a) Registrations on Form S-1. 

(i) Request for S-1 Registration. Subject to Section 2.1(a)(ii), if at any time beginning on the earlier of (i) March 31,
2019 and (ii) 180 days following the effective date of the Corporation’s initial public offering of equity securities, the Corporation shall receive a written request from the Requisite Stockholders (an “S-1 Registration
Request”) that the Corporation effect the registration under the Securities Act of all or any portion of the Registrable Securities (an “S-1 Registration”), then the Corporation shall (x) promptly, and in any event
within 10 days, give written notice of the proposed registration to all other Holders (“S-1 Registration Notice”), and (y) use its best efforts to effect the registration under the Securities Act of the Registrable Securities
that the Corporation has been so requested to register on behalf of the Holder(s) and any Holder joining in such request (as is specified in a written request by each such Holder received by the Corporation within 20 days after delivery of the S-1
Registration Notice) in accordance herewith within sixty (60) days after the receipt of the S-1 Registration Request. Subject to Section 2.1(c), the Corporation may include in such S-1 Registration other securities of the Corporation for
sale, for the Corporation’s account or for the account of any other person. 
 (ii) Limitations on S-1 Registrations. 

(1) Offering Price Limitation. The Corporation shall not be obligated to effect an S-1 Registration pursuant to this
Section 2.1(a) unless the anticipated aggregate offering price of the Registrable Securities to be sold pursuant thereto is at least $10,000,000; 

(2) Limitation on the Number of S-1 Registrations. The Corporation shall not be obligated to effect more than two (2) S-1
Registrations hereunder provided each such registration has been declared or ordered and kept effective for the time period indicated in Article III(a)(iii) below and provided, however, that if the Corporation is not entitled to use
Commission Form S-3 due to the Corporation’s failure to comply with its filing obligations under the Exchange Act, the Holders shall be entitled to additional S-1 Registrations under Section 2.1(a) notwithstanding the foregoing limitation.

  
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 (3) Alternative S-3 Registration. The Corporation shall, if permitted by law, effect any
registration requested under Section 2.1(a) by the filing of a registration statement on Commission Form S-3 pursuant to Section 2.1(b), provided, however, that any such registration conducted pursuant to Form S-3 shall not count towards
the limit on registration requests provided in Section 2.1(a)(ii)(2) above. 
 (4) Recent Registration Limitation. If the
Corporation has effected a Requested Registration within the preceding 180 days and such registration has been declared or ordered effective, the Corporation shall have the right to defer such requested registration for a period of not more than
ninety (90) days after receipt of the request of the Holders, provided that such right to delay a requested registration shall be exercised by the Corporation not more than once in any twelve (12)-month period. 

(5) Delay Limitation. If the Corporation shall furnish to Holders requesting the S-1 Registration, a certificate signed by the
Corporation’s Chief Executive Officer or Chairman of the Board of Directors stating that in the good faith judgment of the Board of Directors of the Corporation such registration at the time requested would be materially detrimental to the
Corporation and its stockholders for such S-1 Registration to be effected at such time in which event the Corporation shall have the right to defer such requested registration for a period of not more than ninety (90) days after receipt of the
request of the Holders, provided that such right to delay a request shall be exercised by the Corporation not more than once in any twelve (12)-month period. 

(6) Simultaneous Corporation Registration Limitation. During the period starting with the date of the filing of, and ending on a date
one hundred eighty (180) days following the effective date of, a registration on Form S-1 pertaining to the initial public offering of securities of the Corporation, the Corporation shall not be obligated to effect a registration under this
Section 2.1 unless otherwise consented to by the underwriter of such offering and only if the Corporation is actively employing in good faith all reasonable efforts to cause such registration statement to become and remain effective. 

(7) Termination. The right to request an S-1 Registration shall terminate on the seventh anniversary of the Corporation’s initial
public offering of its securities. 
 (b) Registrations on Form S-3. 

(i) Request for S-3 Registration. Subject to Section 2.2(b)(ii), if at any time after the Corporation is a registrant entitled to
file a registration statement on Form S-3 or any successor or similar short-form registration statement promulgated by the Commission (collectively, “Commission Form S-3”), the Corporation shall receive a written request from
Holders of at least ten percent (10%) of the Registrable Securities then outstanding (an “S-3 Registration Request”) that the Corporation effect the registration under the Securities Act of all or part of the Registrable
Securities (an “S-3 Registration”, and together with S-1 Registration, a “Requested Registration”), then the Corporation shall (x) promptly, and in any event within 10 days, give written notice of the proposed
registration to all other Holders (an “S-3 Registration Notice”), and (y) use its best efforts to effect the registration under the Securities Act of the Registrable Securities that the Corporation has been so requested to
register on behalf 

  
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of the requesting Holder(s) and any Holder joining in such request (as is specified in a written request by each such Holder received by the Corporation within 15 days after delivery of the S-3
Registration Notice) in accordance herewith within thirty (30) days after receipt of the S-3 Registration Request. Subject to Section 2.1(c), the Corporation may include in such S-3 Registration other securities of the Corporation for
sale, for the Corporation’s account or for the account of any other person. 
 (ii) Limitations on S-3 Registrations. 

(1) Offering Price Limitation. The Corporation shall not be obligated to effect an S-3 Registration pursuant to this
Section 2.1(b) unless the anticipated aggregate offering price of the Registrable Securities to be sold pursuant thereto is at least $1,000,000; provided, however, that if the aggregate number of Registrable Securities held by the Holders have
a fair market value of less than $1,000,000, the Holders of such Registrable Securities may request registration of such Registrable Securities so long as all Registrable Securities are requested to be registered. 

(2) No Limitation on the Number of S-3 Registrations. Subject to the other limitations of this Section 2.1(b)(ii), there shall be
no limitation on the number of S-3 registrations that the Corporation may be required to effect under this Section 2.1(b). 
 (3)
Multiple Simultaneous S-3 Limitation. The Corporation shall not be obligated to keep effective at any one time more than three Commission Form S-3 registration statements in accordance with this Section 2.1(b), and if the Corporation is
requested to effect an additional S-3 Registration at a time when it is keeping three such registration statements effective, it may delay effecting such S-3 Registration until it is no longer required in accordance with Article III(a)(iii) to keep
effective one (or more) of the then effective Commission Form S-3 registration statements. 
 (4) Recent Registration Limitation. The
Corporation shall not be obligated to effect an S-3 Registration pursuant to this Section 2.1(b) if the Corporation has effected a Requested Registration within the preceding 180 days, and such registration has been declared or ordered
effective. 
 (5) Delay Limitation. If the Corporation shall furnish to Holders requesting the S-3 Registration, a certificate signed
by the Corporation’s Chief Executive Officer or Chairman of the Board stating that in the good faith judgment of the Board of Directors of the Corporation such registration at the time requested would be materially detrimental to the
Corporation and its stockholders for such S-3 Registration to be effected at such time, in which event the Corporation shall have the right to defer such filing for a period of not more than ninety (90) days after receipt of the request of the
Holders, provided that such right to delay a request shall be exercised by the Corporation not more than once in any twelve (12)-month period. 

(6) Termination. The rights to request an S-3 Registration shall terminate on the seventh anniversary of the Corporation’s initial
public offering of its securities. 
 (c) Priority in Registration. If a Requested Registration is an underwritten
offering, and the managing underwriters shall give written advice to the Holders and the Corporation that, 

  
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in their opinion, market conditions dictate that no more than a specified maximum number of securities (the “Underwriter’s Maximum Number”) could successfully be
included in such registration within a price range acceptable to the Holders initiating the Requested Registration, then the Corporation shall be required only to include in such registration only such number of securities as is equal to the
Underwriter’s Maximum Number (“Requested Registration Cutback”) and the Corporation and the Holders will participate in such offering in the following order of priority: 

(i) First, there shall be included in such registration that number of Registrable Securities that the Holders shall have requested to be
included in such offering up to the Underwriter’s Maximum Number; and 
 (ii) Second, the Corporation shall be entitled to include in
such registration that number of securities that it proposes to offer and sell for its own account to the full extent of any remaining portion of the Underwriter’s Maximum Number. 

In the event that a Requested Registration Cutback results in less than 50% of the Registrable Securities of Holders that were requested to be included in
such registration actually being included in such registration, then (i) each requesting Holder will be entitled to include his, her or its pro rata share, calculated as the Underwriter’s Maximum Number, multiplied by a fraction, the
numerator of which is the number of shares that such Holder requested to be included in such offering and the denominator of which is the total number of shares requested by Holders to be included in such registration and (ii) such registration
shall not be counted for purposes of the limitations on requested registrations in Section 2.1(a)(ii)(2) above. 
 SECTION 2.2.
Incidental Registrations. 
 (a) Incidental Registration. If the Corporation for itself or any of its
security holders shall (except for registrations under Section 2.1(a)(i), which shall not be deemed registrations for the purposes of this Section 2.2) at any time or times after the date hereof undertake to register (including a Requested
Registration pursuant to Section 2.1(b)) under the Securities Act any shares of its capital stock or other securities (other than (i) the registration of an offer, sale or other disposition of securities solely to employees of, or other
persons providing services to, the Corporation, or any subsidiary pursuant to an employee or similar benefit plan or (ii) relating to a merger, acquisition or other transaction of the type described in Rule 145 under the Securities Act or a
comparable or successor rule, registered on Form S-4 or similar or successor forms promulgated by the Commission), on each such occasion the Corporation will notify each Holder of such determination or request at least thirty (30) days prior to
the filing of such registration statement, and upon the request of any Holder given in writing within twenty (20) days after the receipt of such notice, subject to Section 2.2(b), the Corporation shall use its best efforts as soon as
practicable thereafter to cause any of the Registrable Securities specified by any such Holder to be included in such registration statement to the extent such registration is permissible under the Securities Act and subject to the conditions of the
Securities Act (an “Incidental Registration”). If a Holder decides not to include all of its Registrable Securities in any Incidental Registration filed by the Corporation, such Holder shall nevertheless continue to have the right
to include any Registrable Securities in any subsequent Incidental Registration as may be filed by the Corporation with respect to offerings of its securities, all upon the terms and 

  
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conditions set forth herein. The Corporation shall have the right to terminate or withdraw any Incidental Registration initiated by it under this Section 2.2 prior to the effectiveness of
such registration whether or not any Holder has elected to include securities in such registration. The Registration Expenses of such withdrawn registration shall be borne by the Corporation in accordance with Section 2.3. 

(b) Priority in Registration. If an Incidental Registration is an underwritten offering, and the managing underwriters
shall give written advice to the Holders and the Corporation that, in their opinion, market conditions dictate that no more than the Underwriter’s Maximum Number could successfully be included in such registration, then the Corporation shall be
required only to include in such registration only such number of securities as is equal to the Underwriter’s Maximum Number (“Incidental Registration Cutback”) and the Corporation and the Holders will participate in such
offering in the following order of priority: 
 (i) First, subject to Section 2.2(b)(ii) below, the Corporation shall be entitled
to include in such registration that number of securities that the Corporation proposes to offer and sell for its own account in such registration and that does not exceed the Underwriter’s Maximum Number; and 

(ii) Second, the Corporation will be obligated and required to include in such registration that number of Registrable Securities that the
Holders shall have requested to be included in such offering to the full extent of the remaining portion of the Underwriter’s Maximum Number; provided further, that the Corporation shall make at least thirty percent (30%) of the
Underwriter’s Maximum Number available to Holders that have requested to include Registrable Securities. 
 In the event that an Incidental
Registration Cutback results in less than all of the Registrable Securities of Holders that were requested to be included in such registration actually being included in such registration, then each requesting Holder will be entitled to include his,
her or its pro rata share, calculated as the portion of the Underwriter’s Maximum Number available to Holders who requested to include shares in such offering, multiplied by a fraction, the numerator of which is the number of shares that such
Holder requested to be included in such offering and the denominator of which is the total number of shares requested by Holders to be included in such registration. 

SECTION 2.3. Expenses. The Corporation shall pay all Registration Expenses incurred in connection with all Incidental
Registrations and all Requested Registrations effected in accordance with this Article II. 
 SECTION 2.4.
Effective Registration Statement. A Requested Registration or an Incidental Registration effected pursuant to Section 2.1 or Section 2.2, respectively, shall not be deemed to have been effected unless the
registration statement filed with respect thereto in accordance with the Securities Act has become effective with the Commission and kept effective in accordance with the provisions of Article III(a)(iii) below. Notwithstanding the foregoing, a
registration statement will not be deemed to have become effective if (a) after it has become effective with the Commission, such registration is made subject to any stop order, injunction, or other order or requirement of the Commission or
other governmental agency or any court  

  
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proceeding for any reason other than a misrepresentation or omission by any Holder, or (b) the conditions to closing specified in the purchase agreement or underwriting agreement entered
into in connection with such registration are not satisfied, other than solely by reason of some act or omission by any Holder. 

SECTION 2.5. Jurisdictional Limitations. Notwithstanding anything in this Agreement to the contrary, the Corporation
shall not be obligated to take any action to effect registration, qualification or compliance with respect to its Registrable Securities: 

(a) In any particular jurisdiction in which the Corporation would be required to execute a general consent to service of process unless the
Corporation is already subject to service in such jurisdiction and except as required by the Securities Act; 
 (b) That would require it to
qualify generally to do business in any jurisdiction in which it is not already so qualified or obligated to qualify; or 
 (c) That would
subject it to taxation in a jurisdiction in which it is not already subject generally to taxation. 
 ARTICLE III 

REGISTRATION PROCEDURES 

(a) Corporation Obligations. If and whenever the Corporation is required to use its best efforts to effect the registration of any
Registrable Securities under the Securities Act as provided in Article II, the Corporation, as expeditiously as possible and subject to the terms and conditions of Article II, will do the following: 

(i) Prepare and file with the Commission the appropriate registration statement in the form requested by the required percentage of Holders, in
the case of a Requested Registration, to effect such registration and use its diligent efforts to cause such registration statement to become and remain effective for the period set forth in Article III(a)(iii) below; 

(ii) Permit any Holder who, in the reasonable judgment of the Corporation’s counsel, might be deemed to be an underwriter or a
controlling person of the Corporation, to participate in the preparation of such registration statement (including making available for inspection by any such Person and any attorney, accountant or other agent retained by such Person, all financial
and other records, pertinent corporate documents and all other information reasonably requested in connection therewith) and give to the Holders of Registrable Securities to be sold under such registration statement, the underwriters, if any, and
their respective counsel and accountants, advance draft copies of such registration statement, each prospectus included therein or filed with the Commission at least five (5) business days prior to the filing thereof with the Commission, and
any amendments and supplements thereto promptly as they become available, and will give each of them such access to its books and records and such opportunities to discuss the business of the Corporation with its officers and the independent public
accountants who have certified its financial statements as shall be necessary, in the opinion of such Holders and such underwriters’ respective counsel, to conduct a reasonable investigation within the meaning of the Securities Act; 

  
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 (iii) Prepare and file with the Commission such amendments and supplements to such registration
statement and the prospectus used in connection therewith as may be necessary to keep such registration statement effective and to comply with the provisions of the Securities Act with respect to the disposition of all securities covered by such
registration statement until the earlier of such time as all of such securities have been disposed of in accordance with the intended methods of disposition by the seller or sellers thereof set forth in such registration statement or the expiration
of 180 days after such registration statement becomes effective (such period of 180 days to be extended one day for each day or portion thereof during such period that such registration statement shall be subject to any stop order suspending the
effectiveness of the registration statement, or of any order suspending or preventing the use of any related prospectus or suspending the qualification of any Registrable Securities included in such registration statement for sale in any
jurisdiction); 
 (iv) Furnish to the Holders participating in such registration without charge to the Holders, such number of conformed
copies of such registration statement and of each such amendment and supplement thereto (in each case including all exhibits), such number of copies of the prospectus contained in such registration statement (including each preliminary prospectus
and any summary prospectus) and any other prospectus filed under Rule 424 under the Securities Act, in conformity with the requirements of the Securities Act, and such other documents, as the purchaser or any Holder of Registrable Securities to be
sold under such registration statement may reasonably request; 
 (v) Use its best efforts to register or qualify all Registrable Securities
covered by such registration statement under such other United States state securities or blue sky laws of such jurisdictions as any Holder of Registrable Securities to be sold under such registration statement shall reasonably request, to keep such
registration or qualification in effect for the time period set forth in Article III(a)(iii) hereof, and take any other action that may be reasonably necessary or advisable to enable the Holders who are participating in such registration to sell
Registrable Securities in such jurisdictions; 
 (vi) Use its best efforts to cause all Registrable Securities covered by such registration
statement to be registered with or approved by such other United States state governmental agencies or authorities as may be necessary to enable the Holders who are participating in such registration to sell Registrable Securities as intended by
such registration statement; 
 (vii) In the event of the issuance of any stop order suspending the effectiveness of the registration
statement, or of any order suspending or preventing the use of any related prospectus or suspending the qualification of any Registrable Securities included in such registration statement for sale in any jurisdiction, the Corporation shall use its
best efforts promptly to obtain the withdrawal of such order; 
 (viii) Use its best efforts to furnish to the Holders registering
Registrable Securities under such registration statement: 
 (1) An opinion, dated the effective date of the registration statement, of the
independent counsel representing the Corporation for the purposes of such registration, 

  
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addressed to the underwriters, if any, and to the Holders making such request, stating that such registration statement has become effective under the Securities Act and that: 

(A) To the knowledge of such counsel, no stop order suspending the effectiveness thereof has been issued and no proceedings for that purpose
have been instituted or are pending or contemplated under the Securities Act; 
 (B) The registration statement, the related prospectus,
and each amendment or supplement thereto, comply as to form in all material respects with the requirements of the Securities Act and the applicable rules and regulations of the Commission thereunder (except that such counsel need express no opinion
as to financial statements and related schedules contained therein); 
 (C) To the knowledge of such counsel, as of the effective date,
neither the registration statement, the prospectus, nor any amendment or supplement thereto (other than the financial statements and related schedules therein), contains any untrue statement of a material fact or omits a material fact necessary to
make the statements therein, in light of the circumstances under which they were made, not misleading; 
 (D) The descriptions in the
registration statement or the prospectus, or any amendment or supplement thereto, of the securities to be registered, insofar as such description purports to constitute a summary of the terms of the securities to be registered, and the description
of the underwriting, insofar as such description purports to describe the provisions of the laws and documents, which have been provided to counsel, directly pertaining to the underwriting are accurate and fairly present the information required to
be shown; and 
 (E) Except as disclosed in the registration statement or other public filing made by the Corporation with the Commission,
such counsel does not know of any pending legal or governmental proceedings to which the Corporation is a party or of which any property of the Corporation is the subject that, if determined adversely to the Corporation, would individually or in the
aggregate have a material adverse effect on the then-correct or future consolidated financial position, stockholders’ equity or results of operation of the Corporation, nor of any contracts or documents or instruments of a character required to
be described in the registration statement or prospectus, or any amendment or supplement thereto or to be filed as exhibits to the registration statement that are not described and filed as required (such opinion of counsel shall additionally cover
such legal matters with respect to the registration in respect of which such opinion is being given as a majority in interest of Holders participating in such registration may reasonably request and may contain such qualifications and limitations as
are customarily included in opinions of such sort); and 
 (2) A letter, dated the effective date of the registration statement, from the
independent certified public accountants of the Corporation, addressed to the underwriters, if any, and to the Holders making such request, stating that they are independent certified public accountants within the meaning of the Securities Act and
that in the opinion of such accountants, the financial statements and other financial data of the Corporation included in the registration statement or the prospectus, or any amendment or supplement thereto, comply as to form in all material
respects with the applicable accounting requirements of the Securities Act (such letter 

  
 12 

 
from the independent certified public accountants shall additionally cover such other financial matters (including information as to the period ending not more than five business days prior to
the date of such letter) with respect to the registration in respect of which such letter is being given as the Holders may reasonably request); 

(ix) Immediately notify the Holders of Registrable Securities included in such registration statement at any time when a prospectus relating
thereto is required to be delivered under the Securities Act, of its becoming aware of any event as result of which the prospectus included in such registration statement, as then in effect, includes an untrue statement of material fact or omits to
state any material fact required to be stated therein or necessary to make the statements therein not misleading in the light of the circumstances under which they were made, and at the request of the Holders promptly prepare and furnish to the
Holders a reasonable number of copies of a supplement to or an amendment of such prospectus as may be necessary so that, as thereafter delivered to the purchasers of such securities, such prospectus shall not include an untrue statement of a
material fact or omit to state a material fact required to be stated therein or necessary to make the statements therein not misleading in the light of the circumstances under which they were made; 

(x) Otherwise use its best efforts to comply with all applicable rules and regulations of the Commission, and make available to its security
holders, as soon as reasonably practicable, an earnings statement covering the period of at least twelve months, but not more than eighteen months, beginning with the first full calendar month after the effective date of such registration statement,
which earnings statement shall satisfy the provisions of Section 11(a) of the Securities Act and Rule 158 thereunder; 
 (xi) Provide a
transfer agent and registrar for all Registrable Securities covered by such registration statement not later than the effective date of such registration statement; and 

(xii) Use its best efforts to list all Registrable Securities covered by such registration statement on any securities exchange on which the
same class of securities issued by the Corporation are then listed. or, if no such equity securities are then listed, apply for listing or quotation of the Registrable Securities on an exchange or quotation system selected by a majority in interest
of Holders participating in such registration. 
 (b) Holder Obligations. 

(i) The Corporation may require each Holder of Registrable Securities to be sold under such registration statement to furnish the Corporation
with such information as it may reasonably request in writing (1) regarding such Holder’s proposed distribution of such securities and (2) as required in connection with any registration (including an amendment to a registration
statement or prospectus), qualification or compliance referred to in this Article III. The Corporation agrees not to file or make any amendment to any registration statement with respect to any Registrable Securities, or any amendment of or
supplement to the prospectus used in connection therewith, which refers to any seller of any Registrable Securities covered thereby by name, or otherwise identifies such seller as the holder of any Registrable Securities, without the prior written
consent of such seller, such consent not to be unreasonably withheld, unless such disclosure is required by law, in which case the Corporation will notify such Holder of its intent 

  
 13 

 
to make such amendment or supplement as soon as possible, but in any event in advance of effecting such amendment or supplement. 

(ii) Each Holder, by execution of this Agreement, agrees (1) that upon receipt of any notice from the Corporation, or upon such
Holder’s otherwise becoming aware, of the happening of any event of the kind described in subdivision (a)(ix) of this Article III, such Holder will forthwith discontinue its disposition of Registrable Securities pursuant to the registration
statement relating to such Registrable Securities until the receipt by such Holder of the copies of the supplemented or amended prospectus contemplated by subdivision (a)(ix) of this Article III and, if so directed by the Corporation, will deliver
to the Corporation all copies other than permanent file copies, then in possession of the Holders of the prospectus relating to such Registrable Securities current at the time of receipt of such notice and (2) that it will immediately notify
the Corporation, at any time when a prospectus relating to the registration of such Registrable Securities is required to be delivered under the Securities Act, of the happening of any event as a result of which information previously furnished in
writing by such Holder to the Corporation specifically for inclusion in such prospectus contains an untrue statement of a material fact or omits to state any material fact required to be stated therein or necessary to make the statements therein not
misleading in the light of the circumstances under which they were made. In the event the Corporation or any such Holder shall give any such notice, the period referred to in subdivision (a)(iii) of this Article III shall be extended by a number of
days equal to the number of days during the period from and including the date of the giving of notice pursuant to subdivision (a)(ix) of this Article III to and including the date when such Holder shall have received the copies of the supplemented
or amended prospectus contemplated by subdivision (a)(ix) of this Article III. 
 ARTICLE IV 

UNDERWRITTEN OFFERINGS 

SECTION 4.1. Underwritten Offerings. 

(a) Underwritten Offering. In connection with any underwritten offering pursuant to a registration requested under
Section 2.1, the Corporation will enter into an underwriting agreement (and any other customary agreements) with the underwriters for such offering, such agreement to be in form and substance reasonably satisfactory to the Holders of a majority
of the Registrable Securities to be included in such offering and to such underwriters in their reasonable judgment and to contain such representations and warranties by the Corporation and such other terms as are customarily contained in agreements
of that type, including, without limitation, indemnities to the effect and to the extent provided in Section 5.1. The Corporation will also take all such other actions as the participating Holders or the underwriters reasonably request in order
to expedite or facilitate the disposition of Registrable Securities (including effecting a stock split or combination of shares and the participation of senior management in so-called “road shows” and similar events). Each Holder
participating in such underwritten offering shall be a party to such underwriting agreement and may, at such Holder’s option, require that any or all of the representations and warranties by, and the other agreements on the part of, the
Corporation to and for the benefit of such underwriters shall also be made to and for the benefit of each such Holder and that any or all of the conditions precedent to the obligations of such 

  
 14 

 
underwriters under such underwriting agreement be conditions precedent to the obligations of such Holder. No Holder participating in any such underwritten offering shall be required by the
provisions hereof to make any representations or warranties to or agreements with the Corporation or the underwriters other than representations, warranties or agreements regarding such Holder and its intended method of distribution and any other
representation required by law. 
 (b) Selection of Underwriters. Whenever a registration requested pursuant to
Section 2.1 is for an underwritten offering, the Requisite Stockholders will have the right, but not the obligation, to select the managing underwriter to administer the offering. 

SECTION 4.2. Holdback Agreements. 

(a) Each Holder hereby agrees in connection with the Corporation’s initial public offering of equity securities not to effect (except as
part of such underwritten registration in accordance with the provisions hereof or pursuant to a transaction exempt from registration (other than under Rule 144 or Rule 145 of the Securities Act)) any sale, distribution, short sale, loan, grant of
options for the purchase of, or otherwise dispose of, any Registrable Securities held by such Holder prior to the effective date of such registration for such period as such managing underwriter requests, such period in no event to commence earlier
than seven (7) days prior to, or to end more than 180 days after, the effective date of such registration. In addition, each holder of Registrable Securities agrees to execute and deliver to any managing underwriter (or, in the case of any
offering that is not underwritten, an investment banker) in connection with such registration any lock-up letter requested of such Holder and in form and substance reasonably satisfactory to the Holder by such managing underwriter. Each Holder
further agrees that the Corporation may instruct its transfer agent to place stop transfer notations in its records to enforce the provisions of this Section 4.2(a). The foregoing restrictions shall be conditioned on each officer, director of
the Corporation and holder of one percent or more of the Corporation’s Common Stock or securities convertible or exchangeable for one percent or more of its Common Stock (determined in all instances on a fully diluted basis) being bound by
substantially the same restrictions as are set forth above; provided that the lock-up letters entered into by Sofinnova and Baker may reflect modifications to certain transfer restrictions in connection with required filings under Section 13 or
Section 16(a) of the Exchange Act. 
 (b) After receipt of notice of a Requested Registration pursuant to Section 2.1, the
Corporation shall not initiate, without the consent of a majority in interest of Holders participating in such Requested Registration, a registration of any of its securities for its own account until 90 days after such registration has become
effective or such registration has been terminated. 
 (c) Any discretionary waiver or termination of any of the restrictions included in
Section 4.2(a) by the Corporation or the underwriters shall apply pro rata to all Holders subject to such agreements, based on the number of shares subject to such agreements. 

ARTICLE V 

INDEMNIFICATION AND CONTRIBUTION 

  
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 SECTION 5.1. Indemnification. 

(a) Indemnification by the Corporation. In the event of any registration under the Securities Act pursuant to Article II
of any Registrable Securities covered by such registration, the Corporation will, to the extent permitted by law, and hereby does, indemnify and hold harmless each Holder of Registrable Securities to be sold under such registration statement, the
partners, members, officers, directors, and stockholders of each such Holder, each such Holder’s legal counsel and independent accountants, each other person who participates as an underwriter in the offering or sale of such securities (if so
required by such underwriter as a condition to including the Registrable Securities of the Holders in such registration) and each other person, if any, who controls any such Holder or any such underwriter within the meaning of the Securities Act or
the Exchange Act (each an “Indemnified Party” and collectively, the “Indemnified Parties”), against any losses, claims, damages or liabilities, joint or several, to which any Indemnified Party may become subject
under the Securities Act, the Exchange Act, any state securities laws or otherwise, insofar as such losses, claims, damages or liabilities (or actions or proceedings, whether commenced or threatened, in respect thereof) arise out of or are based
upon any untrue statement or alleged untrue statement of any material fact contained in any registration statement under which such securities were registered under the Securities Act, any preliminary prospectus, final prospectus or summary
prospectus contained therein or any document incorporated therein by reference, or any amendment or supplement thereto, or any omission or alleged omission to state therein a material fact required to be stated therein or necessary to make the
statements therein in light of the circumstances in which they were made not misleading, or arise out of any violation by the Corporation of any rule or regulation promulgated under the Securities Act or state securities law applicable to the
Corporation and relating to action or inaction required of the Corporation in connection with any such registration, and the Corporation will reimburse the Indemnified Parties for any legal or any other expenses reasonably incurred by them in
connection with investigating or defending any such loss, claim, liability, action or proceeding as such expenses are incurred; provided, however, that the indemnity agreement contained in this Section 5.1(a) shall not apply to amounts paid in
settlement of any such loss, claim, damage, liability or action if such settlement is effected without the consent of the Corporation (which consent shall not be unreasonably withheld; and provided, further, however that the Corporation shall
not be liable to an Indemnified Party in any such case to the extent that any such loss, claim, damage, liability (or action or proceeding in respect thereof) or expense arises out of or is based upon any untrue statement or alleged untrue statement
or omission or alleged omission made in such registration statement, any such preliminary prospectus, final prospectus, summary prospectus, amendment or supplement in reliance upon and in conformity with information furnished to the Corporation in
writing by such Indemnified Party specifically for use therein. 
 (b) Indemnification by the Holders. As a condition
to including any Registrable Securities of any person or entity in any registration statement filed pursuant to Article II, each Holder of Registrable Securities, to the extent permitted by law, hereby agrees to indemnify and hold harmless (in the
same manner and to the same extent as set forth in subdivision (a) of this Section 5.1 the Corporation), each director of the Corporation, each officer of the Corporation, each other person, if any, who controls the Corporation within the
meaning of the Securities Act, any other Holder selling securities in such registration statement, and any controlling Person of any such other Holder, with respect to any statement or alleged statement in or omission or

  
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alleged omission from such registration statement, any preliminary prospectus, final prospectus or summary prospectus contained therein, or any amendment or supplement thereto, if, and only
if, such statement or alleged statement or omission or alleged omission was made in reliance upon and in conformity with information furnished in writing to the Corporation directly by such Holder specifically for use therein; provided,
however, that the indemnity agreement contained in this Section 5.1(b) shall not apply to amounts paid in settlement of any losses, claims, damages, liabilities or actions if such settlement is effected without the consent of the Holder; and
provided, further, however, that the obligation of any Holder hereunder shall be limited to an amount equal to the net proceeds (after deduction of all underwriters discounts and commissions paid by such Holder) received by such Holder upon
the sale of Registrable Securities sold in the offering covered by such registration, unless such liability arises out of or is based upon such Holder’s willful misconduct. 

(c) Notices of Claims, etc. Promptly after receipt by an Indemnified Party of notice of the commencement of any action
or proceeding involving a claim referred to in the preceding subdivisions of this Section 5.1, such Indemnified Party will, if a claim in respect thereof is to be made against a party required to provide indemnification (an
“Indemnifying Party”), give written notice to the latter of the commencement of such action, provided, however, that the failure of any Indemnified Party to give notice as provided herein shall not relieve the Indemnifying
Party of its obligation under the preceding subdivisions of this Section 5.1, except to the extent that the Indemnifying Party is actually prejudiced by such failure to give notice. In case any such action is brought against an Indemnified
Party, unless in such Indemnified Party’s reasonable judgment a conflict of interest between such Indemnified and indemnifying parties may exist in respect of such claim, the Indemnifying Party shall be entitled to participate in and to assume
the defense thereof, jointly with any other Indemnifying Party similarly notified to the extent that it may wish, with counsel reasonably satisfactory to such Indemnified Party, and after notice from the Indemnifying Party to such Indemnified Party
of its election so to assume the defense thereof, the Indemnifying Party shall not be liable to such Indemnified Party for any legal or other expenses subsequently incurred by the latter in connection with the defense thereof other than reasonable
costs of investigation. No Indemnifying Party shall consent to entry of any judgment or enter into any settlement without the consent of the Indemnified Party which does not include as an unconditional term thereof the giving by the claimant or
plaintiff to such Indemnified Party of a release from all liability in respect to such claim or litigation. 
 (d)
Other Indemnification. Indemnification similar to that specified in the preceding subdivisions of this Section 5.1 (with appropriate modifications) shall be given by the Corporation and each holder of Registrable Securities included
in any registration statement to each other and any underwriter, as applicable, with respect to any required registration or other qualification of securities under any Federal or state law or regulation of any governmental authority, other than the
Securities Act. 
 (e) Indemnification Payment. The indemnification required by this Section 5.1 shall be made by periodic
payments of the amount thereof during the course of the investigation or defense, as and when bills are received or expense, loss, damage or liability is incurred. 

  
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 (f) Survival of Obligations. The obligations of the Corporation and of the Holders under
this Section 5.1 and Section 5.2 shall survive the completion of any offering of Registrable Securities under this Agreement. 

SECTION 5.2. Contribution. If the indemnification provided for in Section 5.1 is unavailable or insufficient to hold
harmless an Indemnified Party, then each Indemnifying Party shall contribute to the amount paid or payable to such Indemnified Party as a result of the losses, claims, damages or liabilities referred to in Section 5.1 an amount or additional
amount, as the case may be, in such proportion as is appropriate to reflect the relative fault of the Indemnifying Party or parties, on the one hand, and the Indemnified Party, on the other, in connection with the statements or omissions which
resulted in such losses, claims, demands or liabilities as well as any other relevant equitable considerations. The relative fault shall be determined by reference to, among other things, whether the untrue or alleged untrue statement of a material
fact or the omission or alleged omission to state a material fact relates to information supplied by the Indemnifying Party or parties, on the one hand, or the Indemnified Party, on the other, and the parties’ relative, intent, knowledge,
access to information and opportunity to correct or prevent such untrue statement or omission. The amount paid to an Indemnified Party as a result of the losses, claims, damages or liabilities referred to in the first sentence of this
Section 5.2 shall be deemed to include any legal or other expenses reasonably incurred by such Indemnified Party in connection with investigating or defending any action or claim which is the subject of this Article V. No person guilty of
fraudulent misrepresentation within the meaning of Section 11(f) of the Securities Act shall be entitled to contribution from any Person who was not guilty of such fraudulent misrepresentation. Any amounts paid or payable pursuant to this
Section 5.2 shall be limited to an amount equal to the net proceeds (after deduction of all underwriters discounts and commissions paid by such Person) received by such Person upon the sale of Registrable Securities sold in the offering covered
by the applicable registration. 
 ARTICLE VI 

CORPORATION COVENANTS 

SECTION 6.1. Covenants Relating to Rule 144; Reports Under The Exchange Act. With a view to
(a) making available the benefits of certain rules and regulations of the Commission which may at any time permit the sale of securities of the Corporation to the public without registration after such time as a public market exists for the
Common Stock of the Corporation or (b) causing the Corporation to be and remain eligible to file a registration on Commission Form S-3, the Corporation agrees to do the following: 

(i) To make and keep public information available in accordance with Rule 144 under the Securities Act at all times after the effective date of
the first registration under the Securities Act filed by the Corporation for an offering of its securities to the general public; 
 (ii) To
take such action, including the voluntary registration of its Common Stock under Section 12 of the Exchange Act, as is necessary to enable the Holders to utilize Commission Form S-3 for the sale of their Registrable Securities, such action to
be taken as soon as practicable after the end of the fiscal year in which the first registration statement filed by the Corporation for the offering of its securities to the general public is declared effective; 

  
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 (iii) To use its best efforts to then file with the Commission in a timely manner all reports and
other documents required of the Corporation under the Securities Act and the Exchange Act, as amended (at any time after it has become subject to such reporting requirements); 

(b) So long as a Holder owns any Registrable Securities, to furnish to such Holder forthwith upon request a written statement by the
Corporation as to its compliance with the reporting requirements of said Rule 144 (at any time after 90 days after the effective date of the first registration statement filed by the Corporation for an offering of its securities to the general
public), and of the Securities Act and the Exchange Act (at any time after it has become subject to such reporting requirements) and a copy of the most recent annual or quarterly report of the Corporation, and such other reports and documents of the
Corporation as an Holder may reasonably request in availing itself of any rule or regulation of the Commission allowing an Holder to sell any such securities without registration; and 

(i) The Corporation shall use its best efforts to take any action necessary to maintain its eligibility to utilize Commission Form S-3 to
permit resales as requested by the Holders with respect to “Transactions Involving Secondary Offerings” as described in General Instruction I.B.3 of Commission Form S-3. 

SECTION 6.2. Other Registration Rights. Except with the written consent of the Requisite Stockholders, the Corporation
shall not grant to any Person any registration rights so long as any of the registration rights under this Agreement remain in effect. 

SECTION 6.3. Right of First Offer.  

(a) Right of First Offer. Subject to the terms and conditions of this Section 6.3 and applicable securities laws, if, after the
one year anniversary of the date of the first issuance of shares of Series E Preferred Stock by the Corporation (the “Initial Issuance”), the Corporation proposes to offer or sell any new shares of Common Stock or any Common Stock
Equivalents pursuant to a public offering by the Corporation registered under the Securities Act, including an initial public offering, for bona fide capital-raising purposes (the “New Securities”), the Corporation shall give
written notice (the “Offer Notice”) to the Participating Investors stating (i) its bona fide intention to offer such New Securities, (ii) the number of such New Securities to be offered, and (iii) the structure of the
proposed offering or sale. By written notification to the Corporation within five (5) business days after the date of the Offer Notice (the “Exercise Period”), each Participating Investor may elect to purchase, upon the same
terms and conditions as other purchasers in the offering or sale of the New Securities, up to that number of the New Securities as shall be equal to the number of New Securities multiplied by a percentage determined by dividing (x) the number
of shares of Common Stock represented by the Common Stock and Common Stock Equivalents then owned by such Participating Investor by (y) the aggregate number of shares of Common Stock and Common Stock Equivalents then outstanding. Each
Participating Investor shall be entitled to apportion the right of first offer hereby granted to it in such proportions as it deems appropriate, among itself and its Affiliates that are “accredited investors” within the meaning of Rule
501(a) under the Securities Act. At the Corporation’s reasonable request from time to time, each Participating Investor shall confirm in writing the number of securities it owns in the Corporation within three (3) business days of such
request. 

  
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 (b) Limitations. Notwithstanding the foregoing, the right of first offer in this
Section 6.3 shall not be applicable to: 
 (i) securities offered pursuant to resale registration statements; 

(ii) securities issued or issuable in exchange and as consideration in connection with a bona fide collaboration,
partnering, joint venture, or licensing transaction involving the Corporation or any of its Affiliates; 
 (iii) securities
issued or issuable in exchange and as consideration for the bona fide acquisition of another corporation or entity by the Corporation by consolidation, merger, purchase of all or substantially all of the assets, or
other bona fide reorganization in which the Corporation acquires, in a single transaction or series of related transactions, all or substantially all of the assets of such other corporation or entity or fifty percent (50%) or
more of the voting power of such other corporation or entity or fifty percent (50%) or more of the equity ownership of such other entity; 

(iv) securities issuable upon conversion of or with respect to any then-previously-issued or outstanding securities; 

(v) securities issued or issuable to the Corporation by any of the Corporation’s Affiliates; 

(vi) securities issued or issuable in connection with financings by banks, financial institutions, venture debt financing entities or similar
entities in the business of providing debt financing, or in connection with lines of credit, or in connection with royalty or other payment stream monetization transactions; 

(vii) shares of Common Stock and/or Common Stock Equivalents issued or issuable for compensatory purposes to employees, officers, directors,
contractors, vendors, advisors or consultants of the Corporation or any of its subsidiaries (whether or not issued pursuant to a Corporation equity incentive plan); 

(viii) securities issued as a dividend, stock split or distribution on the Common Stock; and 

(ix) any right, option or warrant to acquire any securities set forth in Section 6.2(b)(i)-(viii) above. 

(c) Expiration. 
 (i) In
the event the Participating Investor fails to exercise its right of first refusal within the Exercise Period for the entire amount of New Securities offered pursuant to Section 6.3(a) above, the Corporation shall have sixty (60) days
thereafter to sell or enter into an agreement (pursuant to which the sale of New Securities covered thereby shall be closed, if at all, within thirty (30) days from the date of such agreement) to sell the New Securities respecting which the
Investor’s right of first refusal set forth in Section 6.3(a) was not exercised, upon terms no more favorable to the purchasers thereof than specified in the Offer Notice. In the event the 

  
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Corporation has not sold within such sixty (60) day period or entered into an agreement to sell the New Securities in accordance with the foregoing within such thirty (30) days, the
Corporation shall not thereafter issue or sell any New Securities without first again offering such securities to the Participating Investors in the manner provided in this Section 6.3. 

(ii) The right of a Participating Investor to purchase New Securities pursuant to this Section 6.3 shall terminate upon the earliest of
(a) two years after the closing of the Corporation’s initial public offering, (b) three years after the Initial Issuance, (c) the time when such Participating Investor no longer holds its Required Amount, and (d) a
Liquidation Event. 
 SECTION 6.4. Designation Rights. 

(a) On or after the date of the closing of the Corporation’s initial public offering, at any time during which Baker does not have
a designated representative then serving on the Board of Directors of the Corporation, Baker may, upon written notice to the Corporation (which notice shall indicate the number of securities it then owns in the Corporation), elect to cause, and the
Corporation shall cause, its Board of Directors promptly to appoint one individual selected by Baker (the “Designated Director”) to the Board of Directors (and in the event the Corporation maintains a classified Board of Directors,
the Designated Director shall be appointed to the director class that has the latest date of expiration of the applicable term as of the date of such appointment, subject to applicable law), and to nominate for election at each meeting of the
stockholders of the Corporation at which members of the Corporation’s Board of Directors (or members of the applicable class of the Corporation’s Board of Directors, as the case may be) are elected and included within the slate of
directors contained in the Corporation’s proxy statement, provided that such Designated Director meets the criteria that are reasonably acceptable to the nominating committee (or equivalent committee or the full Board, as applicable) of the
Board of Directors of the Corporation, including the criteria as set forth in Section 6.4(e) hereof. The Corporation shall create a vacancy on the Board of Directors, if needed, to cause the Designated Director to be appointed or elected, as
the case may be, to the Board of Directors pursuant to this Section 6.4(a). 
 (b) The Corporation shall have the right to block a
Designated Director from serving on the Board of Directors or require that such Designated Director resign from the Board of Directors, if such Designated Director holds, or is nominated to hold, a management position or board seat at a company that
the Board of Directors of the Corporation reasonably and in good faith determines directly competes with the Corporation. 
 (c) In the
event the Designated Director is nominated for election at a meeting of the stockholders of the Corporation but is not elected by the stockholders, the Corporation’s obligations pursuant to this Section 6.4 shall terminate and the
Corporation shall have no further obligations under this Section 6.4. 
 (d) During the time that Baker has a right to designate a
director pursuant to this Section 6.4, in the event a vacancy is created by the death, disability, retirement, resignation or removal of a Designated Director (other than pursuant to Section 6.4(c)), Baker may upon written notice to the
Corporation (which notice shall indicate the number of securities it then owns in the Corporation) appoint a new Designated Director to fill the resulting vacancy and the 

  
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Corporation shall cause such person to be promptly appointed to the Board of Directors (and in the event the Corporation maintains a classified Board of Directors, the new Designated Director
shall be appointed to the director class that has the latest date of expiration of the applicable term as of the date of such appointment, subject to applicable law), and nominated for election at each meeting of the stockholders of the Corporation
at which members of the Corporation’s Board of Directors (or members of the applicable class of the Corporation’s Board of Directors, as the case may be) are elected and included within the slate of directors contained in the
Corporation’s proxy statement, provided that such new Designated Director meets the criteria that are reasonably acceptable to the nominating committee (or equivalent committee or the full Board, as applicable) of the Board of Directors of the
Corporation, including the criteria as set forth in Section 6.4(e) hereof. 
 (e) Notwithstanding the foregoing, the rights of Baker to
designate a director shall at all times be subject to applicable rules and published guidance of The NASDAQ Stock Market LLC, including, but not limited to, listing rule 5640 (or any successor rule), and any Corporation policies with respect to
directors. The Designated Director must at all times be considered “independent” as determined in accordance with the rules of the NASDAQ Stock Market, LLC and the Commission. 

(f) The right of Baker to designate a director pursuant to this Section 6.4 shall terminate upon the earliest of (i) two
years after the earlier of the closing of the Corporation’s initial public offering, (ii) the time when Baker no longer holds its Required Amount, and (iii) a Liquidation Event (the “Termination Date”). For purposes
of clarity, in the event the Designated Director’s term ends following the Termination Date, the Corporation shall be under no obligation to re-designate or otherwise nominate the Designated Director or any other director pursuant to this
Section 6.4 following the Termination Date.  
 SECTION 6.5. Section 16 Matters. For so long as any
Participating Investor has a representative on the Corporation’s Board of Directors, the Corporation shall take such reasonable and customary actions as may be required, in accordance with the procedures set forth in Rule 16b-3 promulgated
under the Exchange Act, to cause any acquisition or deemed acquisition or disposition or deemed disposition of securities by such Participating Investor and its Affiliates pursuant to the transactions contemplated by the Series E Purchase Agreement,
including as a result of any adjustments to the Conversion Price (as defined in the Corporation’s certificate of incorporation, as amended and/or restated on or after the date hereof), to be exempt from the provisions of Section 16(b) of
the Exchange Act pursuant to Rule 16b-3 promulgated thereunder, provided that such Participating Investor acknowledges the “deputization” of its director representative on the Corporation’s Board of Directors.  

ARTICLE VII 

ASSIGNABILITY 
 This
Agreement and all of the provisions hereof may be assigned, without the consent of the Corporation, by any Holder to, and shall inure to the benefit of, any purchaser, transferee or assignee of any shares of Registrable Securities (as adjusted for
stock splits, recapitalizations, and other similar events) unless (i) the Holder is transferring less than five percent (5%) of the 

  
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Corporation’s capital stock held by such Holder, unless the transferee is a stockholder, partner or member (or former partner or member) of such Holder; (ii) the purchaser, transferee
or assignee is a direct or indirect competitor of the Corporation as reasonably determined by a majority of the disinterested directors of the Corporation; or (iii) the Holder specifies otherwise in connection with particular transfers of
Registrable Securities, and any such purchaser, transferee or assignee shall take shares of Registrable Securities subject to, and shall be bound by, the terms of this Agreement; provided in each instance that the transferee or assignee of such
rights assumes in writing the obligations of such Holder under this Agreement. However, the Corporation shall not be required to recognize any such purchaser, transferee or assignee as an “Holder” under this Agreement unless and until
either (i) such person becomes the holder of record of Registrable Securities or (ii) the Corporation receives written notice of such purchase, transfer or assignment and (iii) such person executes and delivers to the Corporation a
counter-part signature page to this Agreement. 
 ARTICLE VIII 

MISCELLANEOUS 
 SECTION
8.1. Waivers and Amendments. The rights and obligations of the Corporation and all other parties hereto under this Agreement may be waived (either generally or in a particular instance, either retroactively or prospectively,
and either for a specified period of time or indefinitely) or amended if and only if such waiver or amendment is consented to in writing by the Corporation and by the Requisite Stockholders; provided, however, that this Agreement may not be
amended and the observance of any term hereof may not be waived in a manner that would adversely affect the rights of one or more Holders (the “Adversely Affected Holder”) without the written consent of each Adversely Affected
Holder, unless such amendment, termination, or waiver applies to all Holders in the same fashion; provided, however that no amendment, termination or waiver of Section 6.3 (or any of the defined terms as used therein) with respect to Baker
shall be effective without the written consent of Baker and no amendment, termination or waiver of Section 6.3 (or any of the defined terms as used therein) with respect to Sofinnova shall be effective without the written consent of Sofinnova,
and provided, further, that no amendment, termination or waiver of Section 6.4 (or any of the defined terms as used therein) will be effective without the written consent of Baker. Each Holder shall be bound by any amendment or waiver effected
in accordance with this Section, whether or not such Holder has consented to such amendment or waiver. Upon the effectuation of each such waiver or amendment, the Corporation shall promptly give written notice thereof to the Holders who have not
previously consented thereto in writing. 
 SECTION 8.2. Successors and Assigns. Except as otherwise
expressly provided herein, the provisions hereof shall inure to the benefit of, and be binding upon, the successors, assigns, heirs, executors and administrators of the parties hereto. 

SECTION 8.3. Entire Agreement. This Agreement constitutes the full and entire understanding and agreement of the
parties with regard to the subjects hereof and amends, restates and supersedes in their entirety all other or prior agreements, whether oral or written, with respect thereto (including, without limitation, the Prior Registration Rights Agreement).
This Agreement constitutes consent of the Corporation and the Holders of at least sixty percent 

  
 23 

 
(60%) of the issued and outstanding Series B-2 Preferred Stock and Series C Preferred Stock to the amendment and restatement of the Prior Registration Rights Agreement. 

SECTION 8.4. Notices. All demands, notices, requests, consents and other communications required or permitted under this
Agreement shall be in writing and shall be personally delivered or sent by facsimile machine (with a confirmation copy sent by one of the other methods authorized in this Section), reputable commercial overnight delivery service (including Federal
Express and U.S. Postal Service overnight delivery service) or, deposited with the U.S. Postal Service mailed first class, registered or certified mail, postage prepaid, as set forth below: 

If to the Corporation, addressed to: 

aTyr Pharma, Inc. 
 3545 John
Hopkins Court, Suite 250 
 San Diego, CA 92121 

Attn: Chief Executive Officer 

Fax: (858) 731-8394 
 with a
copy to: 
 Goodwin Procter LLP 

53 State Street 
 Boston, MA 02109

 Attn: Kingsley L. Taft, Esq. 

Fax: (617) 523-1231 
 If to
any Holder, to it at its address specified on Schedule A, with a copy (which shall not constitute notice) to counsel, if any, specified on Schedule A. 

Notices shall be deemed given upon the earlier to occur of (i) receipt by the party to whom such notice is directed; (ii) if sent by
facsimile machine, on the day (other than a Saturday, Sunday or legal holiday in the jurisdiction to which such notice is directed) such notice is sent if sent (as evidenced by the facsimile confirmed receipt) prior to 5:00 p.m. Eastern Time
and, if sent after 5:00 p.m. Eastern Time, on the day (other than a Saturday, Sunday or legal holiday in the jurisdiction to which such notice is directed) after which such notice is sent; (iii) on the first business day (other than a
Saturday, Sunday or legal holiday in the jurisdiction to which such notice is directed) following the day the same is deposited with the commercial courier if sent by commercial overnight delivery service; or (iv) the fifth day (other than a
Saturday, Sunday or legal holiday in the jurisdiction to which such notice is directed) following deposit thereof with the U.S. Postal Service as aforesaid. Each party, by notice duly given in accordance therewith may specify a different address for
the giving of any notice hereunder. 
 SECTION 8.5. Governing Law. This Agreement shall be construed and enforced in
accordance with and governed by the laws of the State of Delaware (without giving effect to any conflicts or choice of laws provisions thereof that would cause the application of the domestic substantive laws of any other jurisdiction). 

  
 24 

 SECTION 8.6. Consent To Jurisdiction. 

(a) EACH OF THE PARTIES HERETO HEREBY CONSENTS TO THE EXCLUSIVE JURISDICTION OF ALL STATE AND FEDERAL COURTS LOCATED IN DELAWARE,
STATE, AS WELL AS TO THE JURISDICTION OF ALL COURTS TO WHICH AN APPEAL MAY BE TAKEN FROM SUCH COURTS, FOR THE PURPOSE OF ANY SUIT, ACTION OR OTHER PROCEEDING ARISING OUT OF, OR IN CONNECTION WITH, THIS AGREEMENT OR ANY OF THE TRANSACTIONS
CONTEMPLATED HEREBY. EACH PARTY HEREBY EXPRESSLY WAIVES ANY AND ALL RIGHTS TO BRING ANY SUIT, ACTION OR OTHER PROCEEDING IN OR BEFORE ANY COURT OR TRIBUNAL OTHER THAN THE COURTS DESCRIBED ABOVE AND COVENANTS THAT IT SHALL NOT SEEK IN ANY MANNER TO
RESOLVE ANY DISPUTE OTHER THAN AS SET FORTH IN THIS SECTION 8.6 OR TO CHALLENGE OR SET ASIDE ANY DECISION, AWARD OR JUDGMENT OBTAINED IN ACCORDANCE WITH THE PROVISIONS HEREOF. 

(b) EACH OF THE PARTIES HERETO HEREBY EXPRESSLY WAIVES ANY AND ALL OBJECTIONS IT MAY HAVE TO VENUE, INCLUDING, WITHOUT LIMITATION,
THE INCONVENIENCE OF SUCH FORUM, IN ANY OF SUCH COURTS. IN ADDITION, EACH OF THE PARTIES CONSENTS TO THE SERVICE OF PROCESS BY PERSONAL SERVICE OR ANY MANNER IN WHICH NOTICES MAY BE DELIVERED HEREUNDER IN ACCORDANCE WITH SECTION 8.4 OF THIS
AGREEMENT. 
 SECTION 8.7. Equitable Remedies. The parties hereto agree that irreparable harm would occur in the
event that any of the agreements and provisions this Agreement were not performed fully by the parties hereto in accordance with their specific terms or conditions or were otherwise breached, and that money damages are an inadequate remedy for
breach of this Agreement because of the difficulty of ascertaining and quantifying the amount of damage that will be suffered by the parties hereto in the event that this Agreement is not performed in accordance with its terms or conditions or is
otherwise breached. It is accordingly hereby agreed that the parties hereto shall be entitled to an injunction or injunctions to restrain, enjoin and prevent breaches of this Agreement by the other parties and to enforce specifically such terms and
provisions of this Agreement, such remedy being in addition to and not in lieu of, any other rights and remedies to which the other parties are entitled to at law or in equity. 

SECTION 8.8. WAIVER OF JURY TRIAL. EACH OF THE PARTIES HERETO HEREBY VOLUNTARILY AND IRREVOCABLY WAIVES TRIAL BY JURY IN ANY ACTION OR
OTHER PROCEEDING BROUGHT IN CONNECTION WITH THIS AGREEMENT, ANY OF THE RELATED AGREEMENTS, DOCUMENTS OR ANY OF THE TRANSACTIONS CONTEMPLATED HEREBY OR THEREBY. 

SECTION 8.9. No Third Party Beneficiary. Except for Indemnified Parties that are not parties hereto, there are no third
party beneficiaries of this Agreement. 
 SECTION 8.10. Expenses. In addition to the payment of the Registration
Expenses set forth in Section 2.3, the Corporation hereby agrees to pay on demand all reasonable documented 

  
 25 

 
out-of-pocket fees, costs and expenses, including reasonable attorneys’ fees incurred by the Holder(s) in connection with the following: (a) the interpretation, proposed amendment,
modification or enforcement of this Agreement, (provided, that the Corporation shall have no obligation to reimburse the Holder(s) for (i) expenses specifically excluded from the definition of “Registrable Securities” and
(ii) expenses incurred in any enforcement action in which the Holder(s) are not the prevailing parties other than expenses payable pursuant to Section 5.2), and (b) any approvals, consents or waivers with respect to this Agreement.

 SECTION 8.11. Addition of Holders. If the Corporation shall issue additional shares of Series E Preferred Stock, or
if any holder of shares of Preferred Stock transfers shares of Preferred Stock, any such purchaser or transferee of such shares of Series E Preferred Stock or any other Preferred Stock may become a party to this Agreement (in the case of any
transferee, subject to Article VII) by executing and delivering an additional counterpart signature page to this Agreement and shall be deemed a “Holder” hereunder. 

SECTION 8.12. Severability; Titles and Subtitles; Gender; Singular and Plural; Counterparts; Facsimile. 

(a) In case any provision of this Agreement shall be invalid, illegal or unenforceable, the validity, legality and enforceability of the
remaining provisions of this Agreement shall not in any way be affected or impaired thereby. 
 (b) The titles of the sections and
subsections of this Agreement are for convenience of reference only and are not to be considered in construing this Agreement. 
 (c) The
use of any gender in this Agreement shall be deemed to include the other genders, and the use of the singular in this Agreement shall be deemed to include the plural (and vice versa), wherever appropriate. 

(d) This Agreement may be executed in any number of counterparts, each of which shall be an original, but all of which together constitute one
instrument. 
 (e) Counterparts of this Agreement (or applicable signature pages hereof) that are manually signed and delivered by facsimile
transmission shall be deemed to constitute signed original counterparts hereof and shall bind the parties signing and delivering in such manner. 

SECTION 8.13. Massachusetts Business Trust. 

A copy of the Agreement and Declaration of Trust of Fidelity or any affiliate thereof is on file with the Secretary of State of the
Commonwealth of Massachusetts and notice is hereby given that this Agreement is executed on behalf of the trustees of Fidelity or any affiliate thereof as trustees and not individually and that the obligations of this Agreement are not binding on
any of the trustees, officers or stockholders of Fidelity or any affiliate thereof individually but are binding only upon Fidelity or any affiliate thereof and its assets and property. 

  
 26 

 SECTION 8.14. Aggregation of Stock. All shares of capital stock of the
Corporation held or acquired by Affiliates shall be aggregated together for the purpose of determining the availability of any rights under this Agreement and such Affiliated persons may apportion such rights as among themselves in any manner they
deem appropriate. 
 [signature page follows] 

  
 27 

 IN WITNESS WHEREOF, the parties hereto have executed this Registration and Voting Rights
Agreement as of the day and year first above written. 
  

			
	CORPORATION
	
	aTYR PHARMA, INC.
		
	By:		/s/ John D. Mendlein
	Name:		John D. Mendlein, Ph.D.
	Title:		Executive Chairman and
Chief Executive Officer

 Signature Page To Registration and Voting Rights Agreement 

 IN WITNESS WHEREOF, the parties hereto have executed this Registration and Voting Rights
Agreement as of the day and year first above written. 
  

			
	HOLDERS:
	
	DOMAIN PARTNERS VIII, L.P.
	
	 By: One Palmer Square Associates VIII, L.L.C.,

Its General Partner

		
	By:		/s/ Lisa A. Kraeutler
			Name: Lisa A. Kraeutler
			Title:  Attorney-in-fact

  

			
	DP VIII ASSOCIATES, L.P.
	
	 By: One Palmer Square Associates VIII, L.L.C.,

Its General Partner

		
	By:		/s/ Lisa A. Kraeutler
			Name: Lisa A. Kraeutler
			Title:  Attorney-in-fact

 Signature Page To Registration and Voting Rights Agreement 

 IN WITNESS WHEREOF, the parties hereto have executed this Registration and Voting Rights
Agreement as of the day and year first above written. 
  

					
	HOLDERS:
	
	ROCK SPRINGS CAPITAL MASTER FUND LP
	
	 By: Rock Springs GP LLC
 Its:
General Partner

		
	By:		/s/ Graham McPhail
			Name:		Graham McPhail
			Title:		Managing Director
					Rock Springs Capital
					650 S. Exeter St., Suite 1070
					Baltimore, MD 21202

 Signature Page To Registration and Voting Rights Agreement 

 IN WITNESS WHEREOF, the parties hereto have executed this Registration and Voting Rights
Agreement as of the day and year first above written. 
  

			
	HOLDERS:
	
	667, L.P.
	
	By: BAKER BROS. ADVISORS LP, management company and investment adviser to 667, L.P., pursuant to authority granted to it by Baker Biotech Capital, L.P., general partner to 667, L.P., and not as the general
partner.
		
	By:		/s/ Scott L. Lessing
			Name: Scott L. Lessing
			Title:  President

  

			
	BAKER BROTHERS LIFE SCIENCES, L.P.
	
	By: BAKER BROS. ADVISORS LP, management company and investment adviser to Baker Brothers Life Sciences, L.P., pursuant to authority granted to it by Baker Brothers Life Sciences Capital, L.P., general partner to Baker
Brothers Life Sciences, L.P., and not as the general partner.
		
	By:		/s/ Scott L. Lessing
			Name: Scott L. Lessing
			Title:  President

 Signature Page To Registration and Voting Rights Agreement 

 IN WITNESS WHEREOF, the parties hereto have executed this Registration and Voting Rights
Agreement as of the day and year first above written. 
  

			
	HOLDERS:
	
	MARSHFIELD ADVISERS, LLC
		
	By:		/s/ Scott Carman
			Name: Scott Carman
			Title: Head of Private Equity

 Signature Page To Registration and Voting Rights Agreement 

 IN WITNESS WHEREOF, the parties hereto have executed this Registration and Voting Rights
Agreement as of the day and year first above written. 
  

			
	HOLDERS:
	
	ALTA PARTNERS VIII, L.P.
	
	By: Alta Partners Management VIII, LLC, Its General Partner
		
	By:		/s/ Larry Randall
			Name: Larry Randall
			Title:   CFO

 Signature Page To Registration and Voting Rights Agreement 

 IN WITNESS WHEREOF, the parties hereto have executed this Registration and Voting Rights
Agreement as of the day and year first above written. 
  

									
			HOLDERS:
			
	POLARIS VENTURE PARTNERS V, L.P.				POLARIS VENTURE PARTNERS FOUNDERS’ FUND V, L.P.
					
	By:		Polaris Venture Management Co. V, L.L.C.,				By:		Polaris Venture Management Co. V, L.L.C.,
	Its General Partner				Its General Partner

  

									
	By:		/s/ William E. Bilodeau				By:		/s/ William E. Bilodeau
			Name:William E. Bilodeau						Name: William E. Bilodeau
			Title: Attorney-in-fact						Title: Attorney-in-fact

  

									
	 POLARIS VENTURE PARTNERS

ENTREPRENEURS’ FUND V, L.P.
				 POLARIS VENTURE PARTNERS SPECIAL

FOUNDERS’ FUND V, L.P.

					
	By:		Polaris Venture Management Co. V, L.L.C.,				By:		Polaris Venture Management Co. V, L.L.C.,
	Its General Partner				Its General Partner

  

									
	By:		/s/ William E. Bilodeau				By:		/s/ William E. Bilodeau
			Name: William E. Bilodeau						Name: William E. Bilodeau
			Title: Attorney-in-fact						Title: Attorney-in-fact

 Signature Page To Registration and Voting Rights Agreement 

 IN WITNESS WHEREOF, the parties hereto have executed this Registration and Voting Rights
Agreement as of the day and year first above written. 
  

			
	HOLDERS:
	
	DLA PIPER VENTURE FUND 2013, LLC
		
	By:		DLA Piper LLP (US), Managing Member

  

			
	By:		/s/ Randy Socol
			 Name: Randy Socol
 Title:
Partner

  

			
	DLA PIPER VENTURE FUND 2008, L.L.C.
		
	By:		DLA Piper LLP (US), Managing Member

  

			
	By:		/s/ Randy Socol
			 Name: Randy Socol
 Title:
Partner

  

			
	DLA PIPER VENTURE FUND 2011, LLC
		
	By:		DLA Piper LLP (US), Managing Member

  

			
	By:		/s/ Randy Socol
			 Name: Randy Socol
 Title:
Partner

 Signature Page To Registration and Voting Rights Agreement 

 IN WITNESS WHEREOF, the parties hereto have executed this Registration and Voting Rights
Agreement as of the day and year first above written. 
  

			
	HOLDERS:
	
	ECOR1 CAPITAL FUND, L.P.
		
	By:		/s/ Oleg Nodelman
			Name: Oleg Nodelman
			Title: Managing Director, EcoR1 Capital, LLC

  

			
	ECOR1 CAPITAL FUND QUALIFIED, L.P.
		
	By:		/s/ Oleg Nodelman
			 Name: Oleg Nodelman
 Title: Managing
Director, EcoR1 Capital, LLC

 Signature Page To Registration and Voting Rights Agreement 

 IN WITNESS WHEREOF, the parties hereto have executed this Registration and Voting Rights
Agreement as of the day and year first above written. 
  

			
	HOLDERS:
	
	SOFINNOVA VENTURE PARTNERS IX, L.P.
		
	By:		Sofinnova Management IX, L.L.C.,
	Its General Partner

  

			
	By:		/s/ Srinivas Akkaraju
			Name: Srinivas Akkaraju, M.D., Ph.D.
			Title: Managing Member

 Signature Page To Registration and Voting Rights Agreement 

 IN WITNESS WHEREOF, the parties hereto have executed this Registration and Voting Rights
Agreement as of the day and year first above written. 
  

			
	HOLDERS:
	
	DEERFIELD PRIVATE DESIGN FUND III, L.P.
		
	By:		 Deerfield Mgmt III, L.P.
 General
Partner

			 By: J.E. Flynn Capital III, LLC

General Partner

  

			
	By:		/s/ David J. Clark
			 Name: David J. Clark
 Title:
Authorized Signatory

  

			
	DEERFIELD SPECIAL SITUATIONS FUND, L.P.
		
	By:		 Deerfield Mgmt, L.P.
 General
Partner

			 By: J.E. Flynn Capital, LLC

General Partner

  

			
	By:		/s/ David J. Clark
			 Name: David J. Clark
 Title: Authorized
Signatory

 Signature Page To Registration and Voting Rights Agreement 

 IN WITNESS WHEREOF, the parties hereto have executed this Registration and Voting Rights
Agreement as of the day and year first above written. 
  

			
	HOLDERS:
	
	CHP II, L.P.
		
	By:		 CHP II Management, LLC
 Its General
Partner

  

			
	By:		/s/ John J. Park
			 Name: John J. Park
 Title: Managing
Member

 Signature Page To Registration and Voting Rights Agreement 

 IN WITNESS WHEREOF, the parties hereto have executed this Registration and Voting Rights
Agreement as of the day and year first above written. 
  

			
	HOLDERS:
	
	HOWARD & DENISE SCHWARTZ TR DTD 1/12/89
		
	By:		/s/ Howard Schwartz
			 Name: Howard Schwartz
 Title:
Trustee

 Signature Page To Registration and Voting Rights Agreement 

 IN WITNESS WHEREOF, the parties hereto have executed this Registration and Voting Rights
Agreement as of the day and year first above written. 
  

			
	HOLDERS:
	
	THE SOCOL FAMILY TRUST U/T/D 7/6/04
		
	By:		/s/ Randy Socol
			 Name: Randy Socol
 Title:
Trustee

 Signature Page To Registration and Voting Rights Agreement 

 IN WITNESS WHEREOF, the parties hereto have executed this Registration and Voting Rights
Agreement as of the day and year first above written. 
  

			
	HOLDERS:
	
	LONGFELLOW VENTURE PARTNERS
		
	By:		/s/ George H. Conrades
			 Name: George H. Conrades
 Title: Managing
Member

 Signature Page To Registration and Voting Rights Agreement 

 IN WITNESS WHEREOF, the parties hereto have executed this Registration and Voting Rights
Agreement as of the day and year first above written. 
  

			
	HOLDERS:
	
	PAUL R. SCHIMMEL PROTOTYPE PSP
		
	By:		/s/ Paul Schimmel
			 Name: Paul R. Schimmel
 Title:
Trustee

 Signature Page To Registration and Voting Rights Agreement 

 IN WITNESS WHEREOF, the parties hereto have executed this Registration and Voting Rights
Agreement as of the day and year first above written. 
  

			
	HOLDERS:
	
	SCHIMMEL REVOCABLE TRUST U/A DTD 9/6/2000
		
	By:		/s/ Paul Schimmel
			Name: Paul R. Schimmel
			Title: Trustee

 Signature Page To Registration and Voting Rights Agreement 

 IN WITNESS WHEREOF, the parties hereto have executed this Registration and Voting Rights
Agreement as of the day and year first above written. 
  

			
	HOLDERS:
	
	PAUL R. SCHIMMEL
	
	/s/ Paul Schimmel
	(Signature)

 Signature Page To Registration and Voting Rights Agreement 

 IN WITNESS WHEREOF, the parties hereto have executed this Registration and Voting Rights
Agreement as of the day and year first above written. 
  

	
	HOLDERS:
	
	JOHN K. CLARKE
	
	/s/ John K. Clarke
	(Signature)

 Signature Page To Registration and Voting Rights Agreement 

 IN WITNESS WHEREOF, the parties hereto have executed this Registration and Voting Rights
Agreement as of the day and year first above written. 
  

	
	HOLDERS:
	
	JOHN D. MENDLEIN, PH.D.
	
	/s/ John D. Mendlein
	(Signature)

 Signature Page To Registration and Voting Rights Agreement 

 IN WITNESS WHEREOF, the parties hereto have executed this Registration and Voting Rights
Agreement as of the day and year first above written. 
 HOLDERS: 

 

									
	 FIDELITY SELECT PORTFOLIOS:

BIOTECHNOLOGY PORTFOLIO
				 FIDELITY ADVISOR SERIES VII:

FIDELITY ADVISOR BIOTECHNOLOGY
 FUND

					
	By:		/s/ Stacie M. Smith				By:		/s/ Stacie M. Smith
			Name: Stacie M. Smith						Name: Stacie M. Smith
			Title: Authorized Signatory						Title: Authorized Signatory

  

									
	 FIDELITY MT. VERNON STREET TRUST:

FIDELITY GROWTH COMPANY FUND
				 FIDELITY ADVISOR SERIES I:
 FIDELITY
ADVISOR GROWTH OPPORTUNITIES
 FUND

					
	By:		/s/ Stacie M. Smith				By:		/s/ Stacie M. Smith
			Name: Stacie M. Smith						Name: Stacie M. Smith
			Title: Authorized Signatory						Title: Authorized Signatory

  

			
	VARIABLE INSURANCE PRODUCTS FUND III: GROWTH OPPORTUNITIES PORTFOLIO
		
	By:		/s/ Stacie M. Smith
			Name: Stacie M. Smith
			Title: Authorized Signatory

 Signature Page To Registration and Voting Rights Agreement 

 IN WITNESS WHEREOF, the parties hereto have executed this Registration and Voting Rights
Agreement as of the day and year first above written. 
  

			
	HOLDERS:
	
	SPHERA GLOBAL HEALTHCARE MASTER FUND
		
	By:		/s/ Doron Breen
			Name: Doron Breen
			Title: Director

 Signature Page To Registration and Voting Rights Agreement 

 IN WITNESS WHEREOF, the parties hereto have executed this Registration and Voting Rights
Agreement as of the day and year first above written. 
  

			
	HOLDERS:
	
	FEDERATED KAUFMANN SMALL CAP FUND, A PORTFOLIO OF FEDERATED EQUITY FUNDS
		
	By:		/s/ Lawrence Auriana
			 Name: Lawrence Auriana
 Title: Vice
President, Federated Global Investment Management, as attorney-in-fact for Federated Kaufmann Small Cap Fund, a portfolio of Federated Equity Funds

 
			
	FEDERATED KAUFMANN FUND, A PORTFOLIO OF FEDERATED EQUITY FUNDS
		
	By:		/s/ Lawrence Auriana
			 Name: Lawrence Auriana
 Title: Vice
President, Federated Global Investment Management, as attorney-in-fact for Federated Kaufmann Small Cap Fund, a portfolio of Federated Equity Funds

  

			
	FEDERATED KAUFMANN FUND II, A PORTFOLIO OF FEDERATED INSURANCE SERIES
		
	By:		/s/ Lawrence Auriana
			Name: Lawrence Auriana
			Title: Vice President, Federated Global Investment Management, as attorney-in-fact for Federated Kaufmann Small Cap Fund, a portfolio of Federated Equity Funds

 IN WITNESS WHEREOF, the parties hereto have executed this Registration and Voting Rights
Agreement as of the day and year first above written. 
  

			
	HOLDERS:
	
	SERIES VIII, A SERIES OF ASTRUM PARTNERSLLC
		
	By:		Magnetar Financial LLC
	Its:		Manager
		
	By:		/s/ Anthony Fox
			Name: Anthony Fox
			 Title: Chief Financial Officer – Funds

          Magnetar Financial LLC

 Signature Page To Registration and Voting Rights Agreement 

 IN WITNESS WHEREOF, the parties hereto have executed this Registration and Voting Rights
Agreement as of the day and year first above written. 
  

			
	HOLDERS:
	
	 T. Rowe Price Health Sciences Fund, Inc.

TD Mutual Funds – TD Health Sciences Fund
 VALIC Company I
– Health Sciences Fund
 T. Rowe Price Health Sciences Portfolio

John Hancock Variable Insurance Trust –Health Sciences Trust

John Hancock Funds II – Health Sciences Fund
 Each fund,
several and not jointly

 
			
		
	By:		T. Rowe Price Associates Inc., Investment Adviser
		
	By:		/s/ Adam Poussard
		
	Name:		Adam Poussard
		
	Title:		Vice President

 
			
	
	 T. Rowe Price New Horizons Fund, Inc.

T. Rowe Price New Horizon Trust
 T. Rowe Price U.S. Equities
Trust
 Each fund, severally and not jointly

 
			
		
	By:		T. Rowe Price Associates Inc., Investment Adviser
		
	By:		/s/ Henry Ellenbogen
		
	Name:		Henry Ellenbogen
		
	Title:		Vice President
	
	Address:
	
	 T. Rowe Price Associates, Inc.
 100
East Pratt Street
 Baltimore, MD 21202
 Attn: Andrew Baek, Vice
President
 Phone: 410-345-2090
 Email:
Andrew_baek@troweprice.com

 Signature Page To Registration and Voting Rights Agreement 

 Schedule A 

List of Holders 
 Holder Names and
Residence or Principal Place of Business 
 Fidelity Select Portfolios: Biotechnology Portfolio 

Brown Brothers Harriman & Co. 

525 Washington Blvd. 
 Jersey
City, NJ 07310 
 Attn: Michael Lerman, 15th Floor 

Corporate Actions 
 with a copy
to: 
 Andrew Boyd 
 Fidelity
Investments 
 82 Devonshire Street, V13H 

Boston, MA 02109 
 Tel:
617-563-5144 
 Fax: 617-385-2818 

Email: andrew.boyd@fmr.com 

Fidelity Advisor Series VII: 

Fidelity Advisor Biotechnology Fund 

State Street Bank & Trust 

P.O. Box 5756 
 Boston, MA 02206

 Attn: Fidelity Advisor Series VII: 

Fidelity Advisor Biotechnology Fund 

with a copy to: 
 Andrew Boyd

 Fidelity Investments 
 82
Devonshire Street, V13H 
 Boston, MA 02109 

Tel: 617-563-5144 
 Fax:
617-385-2818 
 Email: andrew.boyd@fmr.com 

 Holder Names and Residence or Principal Place of Business 

Fidelity Mt. Vernon Street Trust: Fidelity Growth Company Fund 

Ball & Co. 
 c/o Citibank
N.A./Custody 
 IC&D Lock Box 

P.O. Box 7247-7057 
 Philadelphia,
PA 19170-7057 
 Account #: 206681 

with a copy to: 
 Andrew Boyd

 Fidelity Investments 
 82
Devonshire Street, V13H 
 Boston, MA 02109 

Tel: 617-563-5144 
 Fax:
617-385-2818 
 Email: andrew.boyd@fmr.com 

Fidelity Advisor Series I: 

Fidelity Advisor Growth Opportunities Fund 

BNY Mellon 
 Attn: Stacey Wolfe

 525 William Penn Place, Rm 0400 

Pittsburgh, PA 15259 
 with a
copy to: 
 Andrew Boyd 

Fidelity Investments 
 82
Devonshire Street, V13H 
 Boston, MA 02109 

Tel: 617-563-5144 
 Fax:
617-385-2818 
 Email: andrew.boyd@fmr.com 

Variable Insurance Products Fund III: 

Growth Opportunities Portfolio 

BNY Mellon 
 Attn: Stacey Wolfe

 525 William Penn Place, Rm 0400 

Pittsburgh, PA 15259 
 with a
copy to: 
 Andrew Boyd 

Fidelity Investments 
 82
Devonshire Street, V13H 

 Holder Names and Residence or Principal Place of Business 

Boston, MA 02109 
 Tel:
617-563-5144 
 Fax: 617-385-2818 

Email: andrew.boyd@fmr.com 
 CHP
II, L.P. 
 230 Nassau Street 

Princeton, NJ 08542 
 Fax:
(609) 683-0174 
 Attention: John Clarke 

Paul Schimmel Prototype PSP 

Schimmel Revocable Trust U/A Dtd 9/6/2000 

TSRI 
 10550 N. Torrey Pines Road

 Mail Stop BCC 379 
 La Jolla,
CA 92037 
 Alta Partners VIII, L.P. 

c/o Alta Partners Management VIII, LLC 

One Embarcadero Center 
 37th Floor 
 San Francisco, CA 94111 

Fax: (415) 362-6178 

Attention: Finance 
 Polaris
Venture Partners V, L.P. 
 Polaris Venture Partners Entrepreneurs’ Fund V, L.P. 

Polaris Venture Partners Founders’ Fund V, L.P. 

Polaris Venture Partners Special Founders’ Fund V, L.P. 

1000 Winter Street 
 Suite 3350

 Waltham, MA 02451 
 Fax:
(781) 290-0880 
 Attention: Amir Nashat 

Domain Partners VIII, L.P. 
 DP
VIII Associates, L.P. 
 One Palmer Square, Suite 515 

Princeton, NJ 08542 
 Attention:
Lisa Kraeutler 

 Holder Names and Residence or Principal Place of Business 

Imagene Co., Ltd. 
 13F GyeongGi
Bio Center, 
 864-1, iui-dong, Yeongtong-gu, Suwon-city, 

GyeongGi-do 443-270, Korea 
 Fax:
82-31-888-6735 
 Attention: Ms. Myeong-Hee Jo 

Alexandria Equities, LLC 
 385 E.
Colorado Boulevard 
 Suite 299 

Pasadena, CA 91101 
 Fax:
(626) 578-0770 
 Attention: Amanda Cashin or Silvia Chung 

Franklin C. Salisbury, Jr. 

Alexander Rich 
 Longfellow
Venture Partners 
 Pelmea, LP 

c/o George H. Conrades & Meredith Clark Shachoy 

P.O. Box 380199 
 Cambridge, MA
02238 
 Fax: (617) 945-5009 

Timothy J. Rink 
 John Mendlein,
Ph.D. 

 Holder Names and Residence or Principal Place of Business 

Howard & Denise Schwartz Tr dtd 1/12/89 

The Socol Family Trust u/t/d 7/6/04 

c/o Randy Socol 
 DLA Piper
Venture Fund 2011, LLC 
 Attn: L. Burch or F. Hensley 

6225 Smith Ave. 
 Baltimore, MD
21209-3600 
 Telephone: Laura Burch (410) 580-4159 or 

            Fred Hensley (410) 580-4013 

Facsimile: (410) 580-3001 

E-mail: accounting-gcinvests@dlapiper.com 

DLA Piper Venture Fund 2008, L.L.C. 

Attn: L. Burch or F. Hensley 

6225 Smith Ave. 
 Baltimore, MD
21209-3600 
 Telephone: Laura Burch (410) 580-4159 or 

            Fred Hensley (410) 580-4013 

Facsimile: (410) 580-3001 

E-mail: accounting-gcinvests@dlapiper.com 

Cyrus E. Rich 
 Zachary C. Rich

 Josiah D. Rich FBO Nicholas J. Rich 

 Holder Names and Residence or Principal Place of Business 

Josiah D. Rich FBO Nola J. Rich 
 Jessica J. Rich
FBO Abraham W. Sturley 
 Jessica J. Rich FBO Rachel E.G. Sturley 

Jessica J. Rich FBO Abigail Sturley 
 Katherine Schimmel 

TSRI 
 10550 N. Torrey Pines Road 

Mail Stop BCC 379 
 La Jolla, CA 92037 

K. Leyla Schimmel 
 TSRI 

10550 N. Torrey Pines Road 
 Mail Stop BCC 379 

La Jolla, CA 92037 
 Sofinnova Venture Partners IX, L.P. 

3000 Sand Hill Road, Bldg 4, Suite 250 
 Menlo Park, CA 94025 

Baker Brothers Life Sciences, L.P. 
 c/o Baker Brothers
Investments 
 667 Madison Ave., 21st Floor 

New York, NY 10065 
 667, L.P. 

c/o Baker Brothers Investments 
 667 Madison Ave., 21st Floor 
 New York, NY 10065 

Deerfield Private Design Fund III, L.P. 
 787 Third Ave., 37th Floor 
 New York, NY 10017 

 Deerfield Special Situations Fund, L.P. 

787 Third Ave., 37th Floor 

New York, NY 10017 
 T. Rowe Price Health Sciences Fund, Inc.

 c/o T. Rowe Price Associates, Inc. 
 100 E. Pratt St. 

Baltimore, MD 21202 
 Attn: Matthew Dow, Vice President 

Phone: 410-345-3468 
 E-mail: andrew_baek@troweprice.com 

TD Mutual Funds – TD Health Sciences Fund 
 c/o T. Rowe
Price Associates, Inc. 
 100 E. Pratt St. 
 Baltimore, MD 21202

 Attn: Matthew Dow, Vice President 
 Phone: 410-345-3468 

E-mail: andrew_baek@troweprice.com 
 VALIC Company I –
Health Sciences Fund 
 c/o T. Rowe Price Associates, Inc. 
 100
E. Pratt St. 
 Baltimore, MD 21202 
 Attn: Matthew Dow, Vice
President 
 Phone: 410-345-3468 
 E-mail:
andrew_baek@troweprice.com 
 T. Rowe Price Health Sciences Portfolio 

c/o T. Rowe Price Associates, Inc. 
 100 E. Pratt St. 

Baltimore, MD 21202 
 Attn: Matthew Dow, Vice President 

Phone: 410-345-3468 
 E-mail: andrew_baek@troweprice.com 

John Hancock Variable Insurance Trust – Health Sciences Trust 

c/o T. Rowe Price Associates, Inc. 
 100 E. Pratt St. 

Baltimore, MD 21202 
 Attn: Matthew Dow, Vice President 

Phone: 410-345-3468 
 E-mail: andrew_baek@troweprice.com 

John Hancock Funds II – Health Sciences Fund 
 c/o T. Rowe
Price Associates, Inc. 
 100 E. Pratt St. 
 Baltimore, MD 21202

 Attn: Matthew Dow, Vice President 
 Phone: 410-345-3468 

E-mail: andrew_baek@troweprice.com 

 T. Rowe Price New Horizons Fund, Inc. 

c/o T. Rowe Price Associates, Inc. 
 100 E. Pratt St. 

Baltimore, MD 21202 
 Attn: Matthew Dow, Vice President 

Phone: 410-345-3468 
 E-mail: andrew_baek@troweprice.com 

T. Rowe Price New Horizons Trust 
 c/o T. Rowe Price Associates,
Inc. 
 100 E. Pratt St. 
 Baltimore, MD 21202 

Attn: Matthew Dow, Vice President 
 Phone: 410-345-3468 

E-mail: andrew_baek@troweprice.com 
 T. Rowe Price U.S. Equities
Trust 
 c/o T. Rowe Price Associates, Inc. 
 100 E. Pratt St.

 Baltimore, MD 21202 
 Attn: Matthew Dow, Vice President 

Phone: 410-345-3468 
 E-mail: andrew_baek@troweprice.com 

Marshfield Advisers, LLC 
 60 East South Temple Street, Suite 400

 Salt Lake City, UT 84111 
 Attention: Scott Carman 

Federated Kaufmann Fund 
 c/o Federated Equity Funds 

5800 Corporate Drive 
 Pittsburgh, PA 15237 

Federated Kaufmann Small Cap Fund 
 c/o Federated Equity Funds

 5800 Corporate Drive 
 Pittsburgh, PA 15237 

Federated Kaufmann Fund II 
 c/o Federated Equity Funds 

5800 Corporate Drive 
 Pittsburgh, PA 15237 

eCor1 Capital Fund, L.P. 
 c/o EcoR1 Capital, LLC 

409 Illinois St. 
 San Francisco, CA 94158 

 eCor Capital Fund Qualified, L.P. 

c/o EcoR1 Capital, LLC 
 409 Illinois St. 

San Francisco, CA 94158 
 Series VIII, a series of Astrum
Partners LLC 
 c/o Magnetar Capital LLC 
 1603 Orrington 

Evanston, IL 60201 
 Rock Springs Capital Master Fund LP 

c/o Rock Springs Capital 
 650 S. Exeter St., Suite 1070 

Baltimore, MD 21202 
 Sphera Global Healthcare Master Fund 

c/o Sphera Funds Management Ltd. 
 21 Ha’arba’ah St.

 Tel Aviv, Israel 64739 
 John Clarke 

DLA Piper Venture Fund 2013, LLC 
 Attn: L. Burch or F. Hensley

 6225 Smith Ave. 
 Baltimore, MD 21209-3600 

Telephone: Laura Burch (410) 580-4159 or 

        Fred Hensley (410) 580-4013 

Facsimile: (410) 580-3001 
 E-mail:
accounting-gcinvests@dlapiper.comEX-10.12

 Exhibit 10.12 

[AMENDED AND RESTATED] 

INDEMNIFICATION AGREEMENT 

This [Amended and Restated] Indemnification Agreement (“Agreement”) is made as of
[                  ] [      ], 201[    ] by and between aTyr Pharma, Inc., a Delaware
corporation (the “Company”), and [                  ] (“Indemnitee”). 

RECITALS 
 WHEREAS, the Company
desires to attract and retain the services of highly qualified individuals, such as Indemnitee, to serve the Company; 
 WHEREAS, in order
to induce Indemnitee to provide or continue to provide services to the Company, the Company wishes to provide for the indemnification of, and advancement of expenses to, Indemnitee to the maximum extent permitted by law; 

WHEREAS, the Bylaws (the “Bylaws”) of the Company require indemnification of the officers and directors of the Company, and
Indemnitee may also be entitled to indemnification pursuant to the General Corporation Law of the State of Delaware (the “DGCL”); 

WHEREAS, the Bylaws and the DGCL expressly provide that the indemnification provisions set forth therein are not exclusive, and thereby
contemplate that contracts may be entered into between the Company and members of the Board of Directors of the Company (the “Board”), officers and other persons with respect to indemnification; 

WHEREAS, the Board has determined that the increased difficulty in attracting and retaining highly qualified persons such as Indemnitee is
detrimental to the best interests of the Company’s stockholders; 
 WHEREAS, it is reasonable and prudent for the Company contractually
to obligate itself to indemnify, and to advance expenses on behalf of, such persons to the fullest extent permitted by applicable law, regardless of any amendment or revocation of the Company’s Certificate of Incorporation (the
“Charter”) or the Bylaws, so that they will serve or continue to serve the Company free from undue concern that they will not be so indemnified; 

WHEREAS, this Agreement is a supplement to and in furtherance of the indemnification provided in the Charter, the Bylaws and any resolutions
adopted pursuant thereto, and shall not be deemed a substitute therefor, nor to diminish or abrogate any rights of Indemnitee thereunder; and 

[WHEREAS, Indemnitee has certain rights to indemnification and/or insurance provided by [Name of Fund/Sponsor] which Indemnitee and [Name of
Fund/Sponsor] intend to be secondary to the primary obligation of the Company to indemnify Indemnitee as provided in this Agreement, with the Company’s acknowledgment and agreement to the foregoing being a material condition to
Indemnitee’s willingness to serve or continue to serve on the Board.] 

  
 1 

 NOW, THEREFORE, in consideration of the premises and the covenants contained herein, the Company
and Indemnitee do hereby covenant and agree as follows: 
 Section 1.         Services
to the Company. Indemnitee agrees to serve as a director of the Company. Indemnitee may at any time and for any reason resign from such position (subject to any other contractual obligation or any obligation imposed by law), in which event the
Company shall have no obligation under this Agreement to continue Indemnitee in such position. This Agreement shall not be deemed an employment contract between the Company (or any of its subsidiaries or any Enterprise) and Indemnitee. 

Section 2.         Definitions. 

As used in this Agreement: 

(a)       “Change in Control” shall mean: 

(i)        the date any “person,” as such term is used in
Sections 13(d) and 14(d) of the Securities Exchange Act of 1934, as amended (the “Act”) (other than the Company, any of its subsidiaries, or any trustee, fiduciary or other person or entity holding securities under any employee benefit
plan or trust of the Company or any of its subsidiaries), together with all “affiliates” and “associates” (as such terms are defined in Rule 12b-2 under the Act) of such person, becomes the “beneficial owner” (as such
term is defined in Rule 13d-3 under the Act), directly or indirectly, of securities of the Company representing fifty percent (50%) or more of the combined voting power of the Company’s then outstanding securities having the right to vote
in an election of the Board (“Voting Securities”) (in such case other than as a result of an acquisition of securities directly from the Company); or 

(ii)       the date a majority of the members of the Board is replaced
during any 12-month period by directors whose appointment or election is not endorsed by a majority of the members of the Board before the date of the appointment or election; or 

(iii)      the date of consummation of (A) any consolidation or merger of
the Company where the stockholders of the Company, immediately prior to the consolidation or merger, would not, immediately after the consolidation or merger, beneficially own (as such term is defined in Rule 13d-3 under the Act), directly or
indirectly, shares representing in the aggregate more than fifty percent (50%) of the voting shares of the resulting or successor entity in the consolidation or merger (or of its ultimate parent entity, if any), or (B) any sale or other
transfer (in one transaction or a series of transactions contemplated or arranged by any party as a single plan) of all or substantially all of the assets of the Company on a consolidated basis to a person or entity not affiliated with the Company.

 Notwithstanding the foregoing, a “Change in Control” will not be deemed to have

  
 2 

 
occurred for purposes of the foregoing clause (i) solely as the result of an acquisition of securities by the Company which, by reducing the number of shares of Voting Securities
outstanding, increases the proportionate number of Voting Securities beneficially owned by any person to fifty percent (50%) or more of the combined voting power of all of the then outstanding Voting Securities; provided, however, that if any
person referred to in this sentence will thereafter become the beneficial owner of any additional shares of Voting Securities (other than pursuant to a stock split, stock dividend, or similar transaction or as a result of an acquisition of
securities directly from the Company) and immediately thereafter beneficially owns fifty percent (50%) or more of the combined voting power of all of the then outstanding Voting Securities, then a “Change in Control” will be deemed to
have occurred for purposes of the foregoing clause (i). 
 (b)        “Corporate
Status” describes the status of a person as a current or former director of the Company or current or former director, manager, partner, officer, employee, agent or trustee of any other Enterprise which such person is or was serving at the
request of the Company. 
 (c)        “Enforcement Expenses” shall include all
reasonable attorneys’ fees, court costs, transcript costs, fees of experts, travel expenses, duplicating costs, printing and binding costs, telephone charges, postage, delivery service fees, and all other out-of-pocket disbursements or expenses
of the types customarily incurred in connection with an action to enforce indemnification or advancement rights, or an appeal from such action. Expenses, however, shall not include fees, salaries, wages or benefits owed to Indemnitee. 

(d)        “Enterprise” shall mean any corporation (other than the Company),
partnership, joint venture, trust, employee benefit plan, limited liability company, or other legal entity of which Indemnitee is or was serving at the request of the Company as a director, manager, partner, officer, employee, agent or trustee. 

(e)        “Expenses” shall include all reasonable attorneys’ fees, court
costs, transcript costs, fees of experts, travel expenses, duplicating costs, printing and binding costs, telephone charges, postage, delivery service fees, and all other out-of-pocket disbursements or expenses of the types customarily incurred in
connection with prosecuting, defending, preparing to prosecute or defend, investigating, being or preparing to be a witness in, or otherwise participating in, a Proceeding or an appeal resulting from a Proceeding. Expenses, however, shall not
include amounts paid in settlement by Indemnitee, the amount of judgments or fines against Indemnitee or fees, salaries, wages or benefits owed to Indemnitee. 

(f)         “Independent Counsel” means a law firm, or a partner (or, if
applicable, member or shareholder) of such a law firm, that is experienced in matters of Delaware corporation law and neither presently is, nor in the past five (5) years has been, retained to represent: (i) the Company, any subsidiary of
the Company, any Enterprise or Indemnitee in any matter material to any such party; or (ii) any other party to the Proceeding giving rise to a claim for indemnification hereunder. Notwithstanding the foregoing, the term “Independent
Counsel” shall not include any person who, under the applicable standards of professional conduct then prevailing, would have a conflict of interest in representing either the Company or Indemnitee in an action to determine Indemnitee’s
rights under this Agreement. The Company agrees to pay 

  
 3 

 
the reasonable fees and expenses of the Independent Counsel referred to above and to fully indemnify such counsel against any and all expenses, claims, liabilities and damages arising out of or
relating to this Agreement or its engagement pursuant hereto. 
 (g)      The term
“Proceeding” shall include any threatened, pending or completed action, suit, arbitration, alternate dispute resolution mechanism, investigation, inquiry, administrative hearing or any other actual, threatened or completed
proceeding, whether brought in the right of the Company or otherwise and whether of a civil, criminal, administrative, regulatory or investigative nature, and whether formal or informal, in which Indemnitee was, is or will be involved as a party or
otherwise by reason of the fact that Indemnitee is or was a director of the Company or is or was serving at the request of the Company as a director, manager, partner, officer, employee, agent or trustee of any Enterprise or by reason of any action
taken by Indemnitee or of any action taken on his or her part while acting as a director of the Company or while serving at the request of the Company as a director, manager, partner, officer, employee, agent or trustee of any Enterprise, in each
case whether or not serving in such capacity at the time any liability or expense is incurred for which indemnification, reimbursement or advancement of expenses can be provided under this Agreement; provided, however, that the term
“Proceeding” shall not include any action, suit or arbitration, or part thereof, initiated by Indemnitee to enforce Indemnitee’s rights under this Agreement as provided for in Section 12(a) of this Agreement. 

Section 3.        Indemnity in Third-Party Proceedings. The Company shall indemnify
Indemnitee to the extent set forth in this Section 3 if Indemnitee is, or is threatened to be made, a party to or a participant in any Proceeding, other than a Proceeding by or in the right of the Company to procure a judgment in its favor.
Pursuant to this Section 3, Indemnitee shall be indemnified against all Expenses, judgments, fines, penalties, excise taxes, and amounts paid in settlement actually and reasonably incurred by Indemnitee or on his or her behalf in connection
with such Proceeding or any claim, issue or matter therein, if Indemnitee acted in good faith and in a manner he or she reasonably believed to be in or not opposed to the best interests of the Company and, in the case of a criminal proceeding, had
no reasonable cause to believe that his or her conduct was unlawful. 

Section 4.        Indemnity in Proceedings by or in the Right of the Company. The Company
shall indemnify Indemnitee to the extent set forth in this Section 4 if Indemnitee is, or is threatened to be made, a party to or a participant in any Proceeding by or in the right of the Company to procure a judgment in its favor. Pursuant to
this Section 4, Indemnitee shall be indemnified against all Expenses actually and reasonably incurred by Indemnitee or on his or her behalf in connection with such Proceeding or any claim, issue or matter therein, if Indemnitee acted in good
faith and in a manner he or she reasonably believed to be in or not opposed to the best interests of the Company. No indemnification for Expenses shall be made under this Section 4 in respect of any claim, issue or matter as to which Indemnitee
shall have been finally adjudged by a court to be liable to the Company, unless and only to the extent that the Delaware Court of Chancery (the “Delaware Court”) shall determine upon application that, despite the adjudication of
liability but in view of all the circumstances of the case, Indemnitee is fairly and reasonably entitled to indemnification for such expenses as the Delaware Court shall deem proper. 

  
 4 

 Section 5.        Indemnification for Expenses of
a Party Who is Wholly or Partly Successful. Notwithstanding any other provisions of this Agreement and except as provided in Section 7, to the extent that Indemnitee is a party to or a participant in any Proceeding and is successful in such
Proceeding or in defense of any claim, issue or matter therein, the Company shall indemnify Indemnitee against all Expenses actually and reasonably incurred by him or her in connection therewith. If Indemnitee is not wholly successful in such
Proceeding but is successful as to one or more but less than all claims, issues or matters in such Proceeding, the Company shall indemnify Indemnitee against all Expenses actually and reasonably incurred by Indemnitee or on his or her behalf in
connection with each successfully resolved claim, issue or matter. For purposes of this Section and without limitation, the termination of any claim, issue or matter in such a Proceeding by dismissal, with or without prejudice, shall be deemed to be
a successful result as to such claim, issue or matter. 
 Section 6.        Reimbursement
for Expenses of a Witness or in Response to a Subpoena. Notwithstanding any other provision of this Agreement, to the extent that Indemnitee, by reason of his or her Corporate Status, (i) is a witness in any Proceeding to which Indemnitee
is not a party and is not threatened to be made a party or (ii) receives a subpoena with respect to any Proceeding to which Indemnitee is not a party and is not threatened to be made a party, the Company shall reimburse Indemnitee for all
Expenses actually and reasonably incurred by him or her or on his or her behalf in connection therewith. 

Section 7.        Exclusions. Notwithstanding any provision in this Agreement to the
contrary, the Company shall not be obligated under this Agreement: 
 (a)      to indemnify for amounts
otherwise indemnifiable hereunder (or for which advancement is provided hereunder) if and to the extent that Indemnitee has otherwise actually received such amounts under any insurance policy, contract, agreement or otherwise[; provided that
the foregoing shall not affect the rights of Indemnitee or the Fund Indemnitors as set forth in Section 13(c)]; 

(b)      to indemnify for an accounting of profits made from the purchase and sale (or sale and purchase) by
Indemnitee of securities of the Company within the meaning of Section 16(b) of the Securities Exchange Act of 1934, as amended, or similar provisions of state statutory law or common law; 

(c)      to indemnify with respect to any Proceeding, or part thereof, brought by Indemnitee against the
Company, any legal entity which it controls, any director or officer thereof or any third party, unless (i) the Board has consented to the initiation of such Proceeding or part thereof and (ii) the Company provides the indemnification, in
its sole discretion, pursuant to the powers vested in the Company under applicable law; provided, however, that this Section 7(d) shall not apply to (A) counterclaims or affirmative defenses asserted by Indemnitee in an
action brought against Indemnitee or (B) any action brought by Indemnitee for indemnification or advancement from the Company under this Agreement or under any directors’ and officers’ liability insurance policies maintained by the
Company in the suit for which indemnification or advancement is being sought as described in Section 12; or 

  
 5 

 (e)      to provide any indemnification or advancement of expenses
that is prohibited by applicable law (as such law exists at the time payment would otherwise be required pursuant to this Agreement). 

Section 8.        Advancement of Expenses. Subject to Section 9(b), the Company shall
advance, to the extent not prohibited by law, the Expenses incurred by Indemnitee in connection with any Proceeding, and such advancement shall be made within thirty (30) days after the receipt by the Company of a statement or statements
requesting such advances (including any invoices received by Indemnitee, which such invoices may be redacted as necessary to avoid the waiver of any privilege accorded by applicable law) from time to time, whether prior to or after final disposition
of any Proceeding. Advances shall be unsecured and interest free. Advances shall be made without regard to Indemnitee’s ability to repay the expenses and without regard to Indemnitee’s ultimate entitlement to indemnification under the
other provisions of this Agreement. Indemnitee shall qualify for advances upon the execution and delivery to the Company of this Agreement which shall constitute an undertaking providing that Indemnitee undertakes to the fullest extent required by
law to repay the advance if and to the extent that it is ultimately determined by a court of competent jurisdiction in a final judgment, not subject to appeal, that Indemnitee is not entitled to be indemnified by the Company. The right to advances
under this paragraph shall in all events continue until final disposition of any Proceeding, including any appeal therein. Nothing in this Section 8 shall limit Indemnitee’s right to advancement pursuant to Section 12(e) of this
Agreement. 
 Section 9.        Procedure for Notification and Defense of Claim. 

(a)      To obtain indemnification under this Agreement, Indemnitee shall submit to the Company a written
request therefor specifying the basis for the claim, the amounts for which Indemnitee is seeking payment under this Agreement, and all documentation related thereto as reasonably requested by the Company. 

(b)      In the event that the Company shall be obligated hereunder to provide indemnification for or make any
advancement of Expenses with respect to any Proceeding, the Company shall be entitled to assume the defense of such Proceeding, or any claim, issue or matter therein, with counsel approved by Indemnitee (which approval shall not be unreasonably
withheld or delayed) upon the delivery to Indemnitee of written notice of the Company’s election to do so. After delivery of such notice, approval of such counsel by Indemnitee and the retention of such counsel by the Company, the Company will
not be liable to Indemnitee under this Agreement for any fees or expenses of separate counsel subsequently employed by or on behalf of Indemnitee with respect to the same Proceeding; provided that (i) Indemnitee shall have the right to
employ separate counsel in any such Proceeding at Indemnitee’s expense and (ii) if (A) the employment of separate counsel by Indemnitee has been previously authorized by the Company, (B) Indemnitee shall have reasonably concluded
that there may be a conflict of interest between the Company and Indemnitee in the conduct of such defense, or (C) the Company shall not continue to retain such counsel to defend such Proceeding, then the fees and expenses actually and
reasonably incurred by Indemnitee with respect to his or her separate counsel shall be Expenses hereunder. 

  
 6 

 (c)       In the event that the Company does not assume the
defense in a Proceeding pursuant to paragraph (b) above, then the Company will be entitled to participate in the Proceeding at its own expense. 

(d)       The Company shall not be liable to indemnify Indemnitee under this Agreement for any amounts
paid in settlement of any Proceeding effected without its prior written consent (which consent shall not be unreasonably withheld or delayed). The Company shall not, without the prior written consent of Indemnitee (which consent shall not be
unreasonably withheld or delayed), enter into any settlement which (i) includes an admission of fault of Indemnitee, any non-monetary remedy imposed on Indemnitee or any monetary damages for which Indemnitee is not wholly and actually
indemnified hereunder or (ii) with respect to any Proceeding with respect to which Indemnitee may be or is made a party or may be otherwise entitled to seek indemnification hereunder, does not include the full release of Indemnitee from all
liability in respect of such Proceeding. 
 Section 10.        Procedure Upon Application
for Indemnification. 
 (a)        Upon written request by Indemnitee for indemnification
pursuant to Section 9(a), a determination, if such determination is required by applicable law, with respect to Indemnitee’s entitlement to indemnification hereunder shall be made in the specific case by one of the following methods:
(x) if a Change in Control shall have occurred, by Independent Counsel in a written opinion to the Board; or (y) if a Change in Control shall not have occurred: (i) by a majority vote of the disinterested directors, even though less
than a quorum; (ii) by a committee of disinterested directors designated by a majority vote of the disinterested directors, even though less than a quorum; or (iii) if there are no disinterested directors or if the disinterested directors
so direct, by Independent Counsel in a written opinion to the Board. For purposes hereof, disinterested directors are those members of the Board who are not parties to the action, suit or proceeding in respect of which indemnification is sought. In
the case that such determination is made by Independent Counsel, a copy of Independent Counsel’s written opinion shall be delivered to Indemnitee and, if it is so determined that Indemnitee is entitled to indemnification, payment to Indemnitee
shall be made within thirty (30) days after such determination. Indemnitee shall cooperate with the Independent Counsel or the Company, as applicable, in making such determination with respect to Indemnitee’s entitlement to
indemnification, including providing to such counsel or the Company, upon reasonable advance request, any documentation or information which is not privileged or otherwise protected from disclosure and which is reasonably available to Indemnitee and
reasonably necessary to such determination. Any out-of-pocket costs or expenses (including reasonable attorneys’ fees and disbursements) actually and reasonably incurred by Indemnitee in so cooperating with the Independent Counsel or the
Company shall be borne by the Company (irrespective of the determination as to Indemnitee’s entitlement to indemnification) and the Company hereby indemnifies and agrees to hold Indemnitee harmless therefrom. 

(b)        If the determination of entitlement to indemnification is to be made by Independent
Counsel pursuant to Section 10(a), the Independent Counsel shall be selected by the Board if a Change in Control shall not have occurred or, if a Change in Control shall have occurred, by Indemnitee. Indemnitee or the Company, as the case may
be, may, within ten (10) days after written notice of such selection, deliver to the Company or Indemnitee, as the case 

  
 7 

 
may be, a written objection to such selection; provided, however, that such objection may be asserted only on the ground that the Independent Counsel so selected does not meet the
requirements of “Independent Counsel” as defined in Section 2 of this Agreement, and the objection shall set forth with particularity the factual basis of such assertion. Absent a proper and timely objection, the person so selected
shall act as Independent Counsel. If such written objection is so made and substantiated, the Independent Counsel so selected may not serve as Independent Counsel unless and until such objection is withdrawn or the Delaware Court has determined that
such objection is without merit. If, within twenty (20) days after the later of (i) submission by Indemnitee of a written request for indemnification pursuant to Section 9(a), and (ii) the final disposition of the Proceeding,
including any appeal therein, no Independent Counsel shall have been selected without objection, either Indemnitee or the Company may petition the Delaware Court for resolution of any objection which shall have been made by Indemnitee or the Company
to the selection of Independent Counsel and/or for the appointment as Independent Counsel of a person selected by the court or by such other person as the court shall designate. The person with respect to whom all objections are so resolved or the
person so appointed shall act as Independent Counsel under Section 10(a) hereof. Upon the due commencement of any judicial proceeding or arbitration pursuant to Section 12(a) of this Agreement, Independent Counsel shall be discharged and
relieved of any further responsibility in such capacity (subject to the applicable standards of professional conduct then prevailing). 

Section 11.        Presumptions and Effect of Certain Proceedings. 

(a)        To the extent permitted by applicable law, in making a determination with respect to
entitlement to indemnification hereunder, it shall be presumed that Indemnitee is entitled to indemnification under this Agreement if Indemnitee has submitted a request for indemnification in accordance with Section 9(a) of this Agreement, and
the Company shall have the burden of proof to overcome that presumption in connection with the making of any determination contrary to that presumption. Neither (i) the failure of the Company or of Independent Counsel to have made a
determination prior to the commencement of any action pursuant to this Agreement that indemnification is proper in the circumstances because Indemnitee has met the applicable standard of conduct, nor (ii) an actual determination by the Company
or by Independent Counsel that Indemnitee has not met such applicable standard of conduct, shall be a defense to the action or create a presumption that Indemnitee has not met the applicable standard of conduct. 

(b)        The termination of any Proceeding or of any claim, issue or matter therein, by judgment,
order, settlement or conviction, or upon a plea of guilty, nolo contendere or its equivalent, shall not (except as otherwise expressly provided in this Agreement) of itself adversely affect the right of Indemnitee to indemnification or
create a presumption that Indemnitee did not act in good faith and in a manner which he or she reasonably believed to be in or not opposed to the best interests of the Company or, with respect to any criminal Proceeding, that Indemnitee had
reasonable cause to believe that his or her conduct was unlawful. 
 (c)        The knowledge
and/or actions, or failure to act, of any director, manager, partner, officer, employee, agent or trustee of the Company, any subsidiary of the Company, or 

  
 8 

 
any Enterprise shall not be imputed to Indemnitee for purposes of determining the right to indemnification under this Agreement. 

Section 12.        Remedies of Indemnitee. 

(a)        Subject to Section 12(f), in the event that (i) a determination is made pursuant
to Section 10 of this Agreement that Indemnitee is not entitled to indemnification under this Agreement, (ii) advancement of Expenses is not timely made pursuant to Section 8 of this Agreement, (iii) no determination of
entitlement to indemnification shall have been made pursuant to Section 10(a) of this Agreement within sixty (60) days after receipt by the Company of the request for indemnification for which a determination is to be made other than by
Independent Counsel, (iv) payment of indemnification or reimbursement of expenses is not made pursuant to Section 5 or 6 or the last sentence of Section 10(a) of this Agreement within thirty (30) days after receipt by the Company
of a written request therefor (including any invoices received by Indemnitee, which such invoices may be redacted as necessary to avoid the waiver of any privilege accorded by applicable law) or (v) payment of indemnification pursuant to
Section 3 or 4 of this Agreement is not made within thirty (30) days after a determination has been made that Indemnitee is entitled to indemnification, Indemnitee shall be entitled to an adjudication by the Delaware Court of his or her
entitlement to such indemnification or advancement. Alternatively, Indemnitee, at his or her option, may seek an award in arbitration to be conducted by a single arbitrator pursuant to the Commercial Arbitration Rules of the American Arbitration
Association. Indemnitee shall commence such proceeding seeking an adjudication or an award in arbitration within 180 days following the date on which Indemnitee first has the right to commence such proceeding pursuant to this Section 12(a);
provided, however, that the foregoing time limitation shall not apply in respect of a proceeding brought by Indemnitee to enforce his or her rights under Section 5 of this Agreement. The Company shall not oppose Indemnitee’s
right to seek any such adjudication or award in arbitration. 
 (b)        In the event that a
determination shall have been made pursuant to Section 10(a) of this Agreement that Indemnitee is not entitled to indemnification, any judicial proceeding or arbitration commenced pursuant to this Section 12 shall be conducted in all
respects as a de novo trial, or arbitration, on the merits and Indemnitee shall not be prejudiced by reason of that adverse determination. In any judicial proceeding or arbitration commenced pursuant to this Section 12, the Company shall have
the burden of proving Indemnitee is not entitled to indemnification or advancement, as the case may be. 

(c)        If a determination shall have been made pursuant to Section 10(a) of this Agreement
that Indemnitee is entitled to indemnification, the Company shall be bound by such determination in any judicial proceeding or arbitration commenced pursuant to this Section 12, absent (i) a misstatement by Indemnitee of a material fact,
or an omission of a material fact necessary to make Indemnitee’s statement not materially misleading, in connection with the request for indemnification, or (ii) a prohibition of such indemnification under applicable law. 

(d)        The Company shall be precluded from asserting in any judicial proceeding or arbitration
commenced pursuant to this Section 12 that the procedures and presumptions of this Agreement are not valid, binding and enforceable and shall stipulate in any such court or before any such arbitrator that the Company is bound by all the
provisions of this Agreement. 

  
 9 

 (e)        The Company shall indemnify Indemnitee to the
fullest extent permitted by law against any and all Enforcement Expenses and, if requested by Indemnitee, shall (within thirty (30) days after receipt by the Company of a written request therefor) advance, to the extent not prohibited by law,
such Enforcement Expenses to Indemnitee, which are incurred by Indemnitee in connection with any action brought by Indemnitee for indemnification or advancement from the Company under this Agreement or under any directors’ and officers’
liability insurance policies maintained by the Company in the suit for which indemnification or advancement is being sought. Such written request for advancement shall include invoices received by Indemnitee in connection with such Enforcement
Expenses but, in the case of invoices in connection with legal services, any references to legal work performed or to expenditures made that would cause Indemnitee to waive any privilege accorded by applicable law need not be included with the
invoice. 
 (f)        Notwithstanding anything in this Agreement to the contrary, no determination
as to entitlement to indemnification under this Agreement shall be required to be made prior to the final disposition of the Proceeding, including any appeal therein. 

Section 13.        Non-exclusivity; Survival of Rights; Insurance; [Primacy of
Indemnification;] Subrogation. 
 (a)        The rights of indemnification and to receive
advancement as provided by this Agreement shall not be deemed exclusive of any other rights to which Indemnitee may at any time be entitled under applicable law, the Charter, the Bylaws, any agreement, a vote of stockholders or a resolution of
directors, or otherwise. No amendment, alteration or repeal of this Agreement or of any provision hereof shall limit or restrict any right of Indemnitee under this Agreement in respect of any action taken or omitted by such Indemnitee in his or her
Corporate Status prior to such amendment, alteration or repeal. To the extent that a change in Delaware law, whether by statute or judicial decision, permits greater indemnification or advancement than would be afforded currently under the Charter,
Bylaws and this Agreement, it is the intent of the parties hereto that Indemnitee shall enjoy by this Agreement the greater benefits so afforded by such change. No right or remedy herein conferred is intended to be exclusive of any other right or
remedy, and every other right and remedy shall be cumulative and in addition to every other right and remedy given hereunder or now or hereafter existing at law or in equity or otherwise. The assertion or employment of any right or remedy hereunder,
or otherwise, shall not prevent the concurrent assertion or employment of any other right or remedy. 

(b)        To the extent that the Company maintains an insurance policy or policies providing
liability insurance for directors, managers, partners, officers, employees, agents or trustees of the Company or of any other Enterprise, Indemnitee shall be covered by such policy or policies in accordance with its or their terms to the maximum
extent of the coverage available for any such director, manager, partner, officer, employee, agent or trustee under such policy or policies. If, at the time of the receipt of a notice of a claim pursuant to the terms hereof, the Company has director
and officer liability insurance in effect, the Company shall give prompt notice of the commencement of such proceeding to the insurers in accordance with the procedures set forth in the respective policies. 

  
 10 

 (c)        [The Company hereby acknowledges that
Indemnitee has certain rights to indemnification, advancement of expenses and/or insurance provided by [Name of Fund/Sponsor] and certain of [its][their] affiliates (collectively, the “Fund Indemnitors”). The Company hereby agrees
(i) that it is the indemnitor of first resort (i.e., its obligations to Indemnitee are primary and any obligation of the Fund Indemnitors to advance expenses or to provide indemnification for the same expenses or liabilities incurred by
Indemnitee are secondary), (ii) that notwithstanding anything to the contrary in the Charter or Bylaws, the Company shall be required to advance the full amount of expenses incurred by Indemnitee and shall be liable for the full amount of all
Expenses, judgments, penalties, fines and amounts paid in settlement to the extent legally permitted and as required by the terms of this Agreement and the Charter and/or Bylaws (or any other agreement between the Company and Indemnitee), without
regard to any rights Indemnitee may have against the Fund Indemnitors, and (iii) that it irrevocably waives, relinquishes and releases the Fund Indemnitors from any and all claims against the Fund Indemnitors for contribution, subrogation or
any other recovery of any kind in respect thereof. The Company further agrees that no advancement or payment by the Fund Indemnitors on behalf of Indemnitee with respect to any claim for which Indemnitee has sought indemnification from the Company
shall affect the foregoing and the Fund Indemnitors shall have a right of contribution and/or be subrogated to the extent of such advancement or payment to all of the rights of recovery of Indemnitee against the Company. The Company and Indemnitee
agree that the Fund Indemnitors are express third party beneficiaries of the terms of this Section 13(c).] 

(d)        [Except as provided in paragraph (c) above,] [I/i]n the event of any payment under
this Agreement, the Company shall be subrogated to the extent of such payment to all of the rights of recovery of Indemnitee [(other than against the Fund Indemnitors)], who shall execute all papers required and take all action necessary to secure
such rights, including execution of such documents as are necessary to enable the Company to bring suit to enforce such rights. 

(e)        [Except as provided in paragraph (c) above,] [T/t]he Company’s obligation to
provide indemnification or advancement hereunder to Indemnitee who is or was serving at the request of the Company as a director, manager, partner, officer, employee, agent or trustee of any other Enterprise shall be reduced by any amount Indemnitee
has actually received as indemnification or advancement from such other Enterprise. 

Section 14.        Duration of Agreement. This Agreement shall continue until and
terminate upon the later of: (a) ten (10) years after the date that Indemnitee shall have ceased to serve as a director of the Company or as a director, manager, partner, officer, employee, agent or trustee of any other Enterprise for
which Indemnitee is or was serving at the request of the Company in the above-described capacity or (b) one (1) year after the final termination of any Proceeding, including any appeal, then pending in respect of which Indemnitee is
granted rights of indemnification or advancement hereunder and of any proceeding commenced by Indemnitee pursuant to Section 12 of this Agreement relating thereto. This Agreement shall be binding upon the Company and its successors and assigns
and shall inure to the benefit of Indemnitee and his or her heirs, executors and administrators. The Company shall require and cause any successor (whether direct or indirect by purchase, merger, consolidation or otherwise) to all, substantially all
or a substantial part, of the business and/or assets of the Company, by written agreement in form and substance satisfactory to Indemnitee, expressly to assume and agree to perform this 

  
 11 

 
Agreement in the same manner and to the same extent that the Company would be required to perform if no such succession had taken place. 

Section 15.        Severability. If any provision or provisions of this Agreement shall be
held to be invalid, illegal or unenforceable for any reason whatsoever: (a) the validity, legality and enforceability of the remaining provisions of this Agreement (including, without limitation, each portion of any section of this Agreement
containing any such provision held to be invalid, illegal or unenforceable, that is not itself invalid, illegal or unenforceable) shall not in any way be affected or impaired thereby and shall remain enforceable to the fullest extent permitted by
law; (b) such provision or provisions shall be deemed reformed to the extent necessary to conform to applicable law and to give the maximum effect to the intent of the parties hereto; and (c) to the fullest extent possible, the provisions
of this Agreement (including, without limitation, each portion of any section of this Agreement containing any such provision held to be invalid, illegal or unenforceable, that is not itself invalid, illegal or unenforceable) shall be construed so
as to give effect to the intent manifested thereby. 
 Section 16.        Enforcement.

 (a)        The Company expressly confirms and agrees that it has entered into this Agreement and
assumed the obligations imposed on it hereby in order to induce Indemnitee to serve or continue to serve as a director of the Company, and the Company acknowledges that Indemnitee is relying upon this Agreement in serving as a director of the
Company. 
 (b)        This Agreement constitutes the entire agreement between the parties hereto
with respect to the subject matter hereof and supersedes all prior agreements and understandings, oral, written and implied, between the parties hereto with respect to the subject matter hereof; provided, however, that this Agreement
is a supplement to and in furtherance of the Charter, the Bylaws and applicable law, and shall not be deemed a substitute therefor, nor to diminish or abrogate any rights of Indemnitee thereunder. 

Section 17.        Modification and Waiver. No supplement, modification or amendment, or
waiver of any provision, of this Agreement shall be binding unless executed in writing by the parties thereto. No waiver of any of the provisions of this Agreement shall be deemed or shall constitute a waiver of any other provisions of this
Agreement nor shall any waiver constitute a continuing waiver. No supplement, modification or amendment of this Agreement or of any provision hereof shall limit or restrict any right of Indemnitee under this Agreement in respect of any action taken
or omitted by such Indemnitee prior to such supplement, modification or amendment. 

Section 18.        Notice by Indemnitee. Indemnitee agrees promptly to notify the Company
in writing upon being served with any summons, citation, subpoena, complaint, indictment, information or other document relating to any Proceeding or matter which may be subject to indemnification, reimbursement or advancement as provided hereunder.
The failure of Indemnitee to so notify the Company shall not relieve the Company of any obligation which it may have to Indemnitee under this Agreement or otherwise. 

  
 12 

 Section 19.        Notices. All notices,
requests, demands and other communications under this Agreement shall be in writing and shall be deemed to have been duly given if (i) delivered by hand and receipted for by the party to whom said notice or other communication shall have been
directed, (ii) mailed by certified or registered mail with postage prepaid, on the third business day after the date on which it is so mailed, (iii) mailed by reputable overnight courier and receipted for by the party to whom said notice
or other communication shall have been directed or (iv) sent by facsimile transmission, with receipt of oral confirmation that such transmission has been received: 

(a)        If to Indemnitee, at such address as Indemnitee shall provide to the Company. 

(b)        If to the Company to: 

aTyr Pharma, Inc. 
 3545 John
Hopkins Court, Suite #250 
 San Diego, CA 92121 

Attention: Secretary 
 or to any other address
as may have been furnished to Indemnitee by the Company. 

Section 20.        Contribution. To the fullest extent permissible under applicable law,
if the indemnification provided for in this Agreement is unavailable to Indemnitee for any reason whatsoever, the Company, in lieu of indemnifying Indemnitee, shall contribute to the amount incurred by Indemnitee, whether for judgments, fines,
penalties, excise taxes, amounts paid or to be paid in settlement and/or for Expenses, in connection with any Proceeding in such proportion as is deemed fair and reasonable in light of all of the circumstances in order to reflect (i) the
relative benefits received by the Company and Indemnitee in connection with the event(s) and/or transaction(s) giving rise to such Proceeding; and/or (ii) the relative fault of the Company (and its directors, officers, employees and agents) and
Indemnitee in connection with such event(s) and/or transactions. 

Section 21.        Internal Revenue Code Section 409A. The Company intends for this
Agreement to comply with the Indemnification exception under Section 1.409A-1(b)(10) of the regulations promulgated under the Internal Revenue Code of 1986, as amended (the “Code”), which provides that indemnification of, or
the purchase of an insurance policy providing for payments of, all or part of the expenses incurred or damages paid or payable by Indemnitee with respect to a bona fide claim against Indemnitee or the Company do not provide for a deferral of
compensation, subject to Section 409A of the Code, where such claim is based on actions or failures to act by Indemnitee in his or her capacity as a service provider of the Company. The parties intend that this Agreement be interpreted and
construed with such intent. 
 Section 22.        Applicable Law and Consent to
Jurisdiction. This Agreement and the legal relations among the parties shall be governed by, and construed and enforced in accordance with, the laws of the State of Delaware, without regard to its conflict of laws rules. Except with respect to
any arbitration commenced by Indemnitee pursuant to Section 12(a) of this Agreement, the Company and Indemnitee hereby irrevocably and unconditionally (i) agree that any action or proceeding arising out of or in connection with this
Agreement shall be brought 

  
 13 

 
only in the Delaware Court, and not in any other state or federal court in the United States of America or any court in any other country, (ii) consent to submit to the exclusive
jurisdiction of the Delaware Court for purposes of any action or proceeding arising out of or in connection with this Agreement, (iii) consent to service of process at the address set forth in Section 19 of this Agreement with the same
legal force and validity as if served upon such party personally within the State of Delaware, (iv) waive any objection to the laying of venue of any such action or proceeding in the Delaware Court, and (v) waive, and agree not to plead or
to make, any claim that any such action or proceeding brought in the Delaware Court has been brought in an improper or inconvenient forum. 

Section 23.        Headings. The headings of the paragraphs of this Agreement are inserted
for convenience only and shall not be deemed to constitute part of this Agreement or to affect the construction thereof. 

Section 24.        Identical Counterparts. This Agreement may be executed in one or more
counterparts, each of which shall for all purposes be deemed to be an original but all of which together shall constitute one and the same Agreement. Only one such counterpart signed by the party against whom enforceability is sought needs to be
produced to evidence the existence of this Agreement. 
 [Remainder of Page Intentionally Left Blank] 

  
 14 

 IN WITNESS WHEREOF, the parties have caused this [Amended and Restated] Indemnification Agreement
to be signed as of the day and year first above written. 
  

							
					aTYR PHARMA, INC.
				
					By:		  

							Name:
							Title:
				
							  

							[Name of Indemnitee]

 SIGNATURE PAGE TO 

[AMENDED AND RESTATED] INDEMNIFICATION AGREEMENT

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