Document:

EX-4.1

 Exhibit 4.1 

EXECUTION VERSION 
 LYRA
THERAPEUTICS, INC. 
 EIGHTH AMENDED AND RESTATED INVESTOR RIGHTS AGREEMENT 

This Agreement dated as of January 10, 2020 is entered into by and among Lyra Therapeutics, Inc., a Delaware corporation (the
“Company”), Carmichael Roberts, George Whitesides and Robert Langer (individually, a “Founder” and collectively, the “Founders”) and the individuals and entities listed as preferred stockholders on
Schedule A attached hereto (individually, an “Investor” and collectively, the “Investors”). 

Recitals 
 WHEREAS, the
Company was a party with the stockholders of the Company named therein to a Seventh Amended and Restated Investor Rights Agreement dated as of June 5, 2018, by and among the Company, the Investors and the other parties thereto (as amended, the
“Prior IRA”). 
 WHEREAS, certain parties, concurrently herewith are purchasing shares of the Company’s Series
C Convertible Preferred Stock pursuant to that certain Series C Convertible Preferred Stock and Warrant Purchase Agreement of even date herewith (the “Series C Purchase Agreement”). 

WHEREAS, the Investors’ obligations in the Series C Purchase Agreement are conditioned upon the Company’s execution and delivery of
this Agreement. 
 WHEREAS, the signatories to this Agreement hold the requisite number of shares to effect the amendment of the Prior IRA
and desire to amend and restate the Prior IRA in its entirety. 
 WHEREAS, the Company and the Investors desire to provide for certain
arrangements with respect to (i) the registration of shares of capital stock of the Company under the Securities Act of 1933, as amended and (ii) the Investors’ right of first refusal with respect to certain issuances of securities of
the Company. 
 NOW, THEREFORE, in consideration of the mutual promises and covenants contained in this Agreement and for other good and
valuable consideration, the receipt and sufficiency of which are hereby acknowledged, the parties hereto agree as follows: 
 1. Certain
Definitions.     
 As used in this Agreement, the following terms shall have the following respective meanings:

 “Available Undersubscription Amount” shall have the meaning set forth in Section 3.1(b). 

“Board of Directors” means the Company’s board of directors. 

“Charter” means the Company’s Amended and Restated Certificate of Incorporation. 

 “Code” means the Internal Revenue Code of 1986, as amended. 

“Commission” means the Securities and Exchange Commission, or any other federal agency at the time administering the
Securities Act. 
 “Common Stock” means the common stock, $0.001 par value per share, of the Company. 

“Company” shall have the meaning set forth in the Preamble. 

“Company Sale” means: (a) a merger or consolidation in which (i) the Company is a constituent party, or (ii) a
subsidiary of the Company is a constituent party and the Company issues shares of its capital stock pursuant to such merger or consolidation, except in the case of either clause (i) or (ii), any such merger or consolidation involving the
Company or any of its subsidiaries in which the shares of capital stock of the Company outstanding immediately prior to such merger or consolidation continue to represent, or are converted into or exchanged for equity interests which represent,
immediately following such merger or consolidation, more than fifty percent (50%) by voting power of the equity interests of (A) the surviving or resulting entity or (B) if the surviving or resulting entity is a wholly owned subsidiary of
another entity immediately following such merger or consolidation, the entity that is the parent entity of such surviving or resulting entity, in each case in substantially the same proportions as such stockholders held the outstanding stock of the
Company immediately prior thereto; (b) the sale, lease, transfer, exclusive license or other disposition, in a single transaction or series of related transactions, by the Company or any of its subsidiaries, of all or substantially all the
assets of the company and its subsidiaries taken as a whole (except where such sale, lease, transfer, exclusive license or other disposition is to a wholly owned subsidiary of the Company); or (c) the sale or transfer, in a single transaction
or series of related transactions, by the stockholders of the Company of more than fifty percent (50%) by voting power of the then-outstanding capital stock of the Company to any person or entity or group of affiliated entities. 

“Exchange Act” means the Securities Exchange Act of 1934, as amended, or any successor federal statute, and the rules and
regulations of the Commission issued under such Act, as they each may, from time to time, be in effect. 
 “Founder” shall
have the meaning set forth in the Preamble. 
 “Founders” shall have the meaning set forth in the Preamble. 

“Founder Registrable Shares” shall have the meaning set forth in Section 2.2(a). 

“Indemnified Party” shall have the meaning set forth in Section 2.5(c). 

“Indemnifying Party” shall have the meaning set forth in Section 2.5(c). 

“Initiating Holders” means the Stockholders initiating a request for registration pursuant to Section 2.1(a) or 2.1(b),
as the case may be. 
 “Initial Public Offering” means the initial underwritten public offering of shares of Common Stock
pursuant to an effective Registration Statement. 

  
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 “Investors” shall have the meaning set forth in the Preamble. 

“Other Holders” shall have the meaning set forth in Section 2.1(d). 

“Notice of Acceptance” shall have the meaning set forth in Section 3.1(b). 

“Preferred Directors” shall have the meaning assigned to that term in the Charter. 

“Preferred Stock” means the Series A-1 Preferred (as defined below), Series A-2 Preferred (as defined below), Series A-3 Preferred (as defined below), Series A-4 Preferred (as defined below), Series B Preferred
(as defined below) and Series C Preferred (as defined below). 
 “Prospectus” means the prospectus included in any
Registration Statement, as amended or supplemented by an amendment or prospectus supplement, including post-effective amendments, and all material incorporated by reference or deemed to be incorporated by reference in such Prospectus. 

“Qualifying Holder” means any Stockholder that is an “accredited investor” within the meaning of Rule 501(a) under
the Securities Act. 
 “Refused Securities” shall have the meaning set forth in Section 3.1(c). 

“Registration Statement” means a registration statement filed by the Company with the Commission for a public offering and
sale of securities of the Company (other than a registration statement on Form S-8 or Form S-4, or their successors, or any other form for a similar limited purpose, or
any registration statement covering only securities proposed to be issued in exchange for securities or assets of another corporation).     

“Registration Expenses” means the expenses described in Section 2.4. 

“Registrable Shares” means (i) the shares of Common Stock issued or issuable to an Investor or other Stockholder upon
conversion of Preferred Stock, (ii) any shares of Common Stock held by an Investor or other Stockholder, and any shares of Common Stock issued or issuable upon the conversion or exercise of any other securities, in each case that are acquired
by such Investor or other Stockholder prior to the Initial Public Offering, and (iii) any other shares of Common Stock issued in respect of the shares described in clauses (i) and (ii) (because of stock dividends, splits,
reclassifications, recapitalizations, or similar events occurring after the date of this Agreement); provided, however, that shares of Common Stock which are Registrable Shares shall cease to be Registrable Shares upon (i) any
sale pursuant to a Registration Statement or Rule 144 under the Securities Act or (ii) any sale in any manner to a person or entity which, by virtue of Section 4 of this Agreement, is not entitled to the rights provided by this Agreement.
Wherever reference is made in this Agreement to a request or consent of holders of a certain percentage of Registrable Shares, the determination of such percentage shall include shares of Common Stock issuable upon conversion of the Preferred Stock
even if such conversion has not been effected. 

  
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 “Securities Act” means the Securities Act of 1933, as amended, or any
successor federal statute, and the rules and regulations of the Commission issued under such Act, as they each may, from time to time, be in effect. 

“Selling Stockholder” means any Stockholder owning Registrable Shares included in a Registration Statement. 

“Series A-1 Preferred” means the Series A-1/A
Convertible Preferred Stock of the Company, $0.001 par value per share, the Series A-1/B Convertible Preferred Stock of the Company, $0.001 par value per share and the Series
A-1/C Convertible Preferred Stock of the Company, $0.001 par value per share. 
 “Series A-2 Preferred” means the Series A-2 Convertible Preferred Stock of the Company, $0.001 par value per share. 

“Series A-3 Preferred” means the Series A-3
Convertible Preferred Stock of the Company, $0.001 par value per share. 
 “Series A-4
Preferred” means the Series A-4 Convertible Preferred Stock of the Company, $0.001 par value per share. 

“Series B Director” shall have the meaning assigned to that term in the Charter. 

“Series B Preferred” means the Series B Convertible Preferred Stock of the Company, $0.001 par value per share. 

“Series C Director” shall have the meaning assigned to that term in the Charter. 

“Series C Original Issue Date” shall mean the date on which a share of Series C Preferred was first issued. 

“Series C Preferred” means the Series C Convertible Preferred Stock of the Company, $0.001 par value per share. 

“Significant Stockholder” shall mean each Stockholder who holds not less than seven million (7,000,000) shares of Preferred
Stock (subject to appropriate adjustment in the event of any stock dividend, stock split, combination or other similar recapitalization occurring after the date of this Agreement and affecting the number of issued and outstanding shares of Preferred
Stock), including, for this purpose, shares of Common Stock issued upon conversion of Preferred Stock so long as such Common Stock is held by a current or former holder of Preferred Stock, but excluding any person that is a competitor of the Company
as determined by the Board of Directors, including the Preferred Directors, in good faith). 
 “Stockholder” means any
Investor and any person or entity to whom the rights granted under this Agreement are transferred by any holder of the Preferred Stock, its successors or assigns pursuant to Section 4 hereof. 

  
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 2. Registration Rights 

2.1. Required Registrations. 

(a) At any time after the earlier of (i) the fifth anniversary of the Series C Original Issue Date or (ii) six months after the
closing of the Initial Public Offering, a Stockholder or Stockholders holding in the aggregate at least thirty percent (30%) of the Registrable Shares then outstanding (which in the event that an Initial Public Offering has not yet occurred, must
include holders of at least fifty percent (50%) of the Registrable Shares issued or issuable upon conversion of the Series B Preferred and Series C Preferred then outstanding) may request, in writing, that the Company effect the registration on Form
S-1 (or any successor form) of Registrable Shares owned by such Stockholder or Stockholders having an aggregate value of at least Five Million Dollars ($5,000,000) (based on the market price or fair value on
the date of such request). 
 (b) At any time after the Company becomes eligible to file a Registration Statement on Form S-3 (or any successor form relating to secondary offerings), a Stockholder or Stockholders may request, in writing, that the Company effect the registration on Form S-3 (or
such successor form) (if Form S-3 (or such successor form) is available for such offering), of Registrable Shares having an aggregate value of at least Two Million Dollars ($2,000,000) (based on the public
market price on the date of such request). 
 (c) Upon receipt of any request for registration pursuant to this Section 2.1,
the Company shall promptly give written notice of such proposed registration to all other Stockholders. Such Stockholders shall have the right, by giving written notice to the Company within thirty (30) days after the Company provides its
notice, to elect to have included in such registration such of their Registrable Shares as such Stockholders may request in such notice of election, subject in the case of an underwritten offering to the approval of the managing underwriter as
provided in Section 2.1(d) below. Thereupon, the Company shall use its reasonable best efforts to effect, as expeditiously as possible, but no later than sixty (60) days following the date of the registration request under
Section 2.1(a) or (b) as applicable, the registration on an appropriate registration form of all Registrable Shares which the Company has been requested to so register (provided, however, that in the case of a
registration requested under Section 2.1(b), the Company will only be obligated to effect such registration on Form S-3 (or any successor form)). 

(d) If the Initiating Holders intend to distribute the Registrable Shares covered by their request by means of an underwriting, they shall so
advise the Company as a part of their request made pursuant to Section 2.1(a) or (b), as the case may be, and the Company shall include such information in its written notice referred to in Section 2.1(c). The right of
any other Stockholder to include its Registrable Shares in such registration pursuant to Section 2.1(a) or (b), as the case may be, shall be conditioned upon such other Stockholder’s participation in such underwriting on the
terms set forth herein. 

  
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 If the Company desires that any securities of the Company held by officers and directors of
the Company be included in any registration for an underwritten offering requested pursuant to Sections 2.1(a) or (b) or if other holders of securities of the Company who are entitled, by contract with the Company, to have
securities included in such a registration (the “Other Holders”) request such inclusion, the Company may include the securities of such officers, directors and Other Holders in such registration and underwriting on the terms set
forth herein. The Company shall (together with all Stockholders, officers, directors and Other Holders proposing to distribute their securities through such underwriting) enter into an underwriting agreement in customary form (including, without
limitation, customary indemnification and contribution provisions on the part of the Company) with the managing underwriter. Notwithstanding any other provision of this Section 2.1(d), if the managing underwriter advises the Company that the
inclusion of all shares requested to be registered would adversely affect the offering, the securities of the Company held by officers or directors of the Company (other than Registrable Shares) and the securities held by Other Holders (other than
Registrable Shares) shall be excluded from such registration and underwriting to the extent deemed advisable by the managing underwriter, and if a further limitation of the number of shares is required, the number of shares that may be included in
such registration and underwriting shall be allocated among all holders of Registrable Shares requesting registration in proportion, as nearly as practicable, to the respective number of Registrable Shares held by them at the time of the request for
registration made by the Initiating Holders pursuant to Section 2.1(a) or (b), as the case may be. If any holder of Registrable Shares, officer, director or Other Holder who has requested inclusion in such registration as provided
above disapproves of the terms of the underwriting, such person may elect to withdraw therefrom by written notice to the Company, and the securities so withdrawn shall also be withdrawn from registration. If the managing underwriter has not limited
the number of Registrable Shares or other securities to be underwritten, the Company may include securities for its own account in such registration if the managing underwriter so agrees and if the number of Registrable Shares and other securities
which would otherwise have been included in such registration and underwriting will not thereby be limited. 
 (e) The Initiating Holders
shall have the right to select the managing underwriter(s) for any underwritten offering requested pursuant to Section 2.1(a) or (b), subject to the approval of the Company, which approval will not be unreasonably withheld or
delayed. 
 (f) The Company shall not be required to effect more than two (2) registrations pursuant to Section 2.1(a) or
more than two (2) registrations in any 12-month period pursuant to Section 2.1(b). In addition, the Company shall not be required to effect any registration (other than on Form S-3 or any successor form relating to secondary offerings) within six months after the effective date of any other Registration Statement of the Company. For purposes of this Section 2.1(f), a
Registration Statement shall not be counted until such time as such Registration Statement has been declared effective by the Commission (unless the Initiating Holders withdraw their request for such registration, other than as a result of
information concerning the business or financial condition of the Company which is made known to the Stockholders after the date on which such registration was requested, and elect not to pay the Registration Expenses therefor pursuant to
Section 2.4). 
 (g) If at the time of any request to register Registrable Shares by Initiating Holders pursuant to this
Section 2.1, the Company is engaged or has plans to engage in a registered public offering or is engaged in any other activity which, in the good faith determination of the Company’s Board of Directors, would be adversely affected by the
requested registration, then the Company may at its option direct that such request be delayed for a period not in excess of ninety (90) days from the date of such request, such right to delay a request to be exercised by the Company not more
than once in any 12-month period. 

  
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 2.2. Incidental Registration. 

(a) Whenever the Company proposes to file a Registration Statement (other than in connection with the Initial Public Offering but in the case
of Stockholders, other than a Registration Statement filed pursuant to Section 2.1, but in the case of the Founders, including a Registration Statement filed pursuant to Section 2.1), at any time and from time to time, it
will, prior to such filing, give written notice to all Stockholders and Founders of its intention to do so. Upon the written request of a Stockholder or Stockholders, or Founder or Founders, given within twenty (20) days after the Company
provides such notice (which request shall state the intended method of disposition of the Registrable Shares or, in the case of Founders, Common Stock, requested to be registered), the Company shall use its reasonable best efforts to cause all
Registrable Shares which the Company has been requested by such Stockholder or Stockholders to register, and all of the shares of Common Stock that Founder or Founders have requested that the Company register (together with any other shares of
Common Stock issued in respect of the shares requested to be included by the Founders (because of stock dividends, splits, reclassifications, recapitalizations or similar events occurring after the date of this Agreement) the “Founder
Registrable Shares”) to be registered under the Securities Act to the extent necessary to permit their sale or other disposition in accordance with the intended methods of distribution specified in such request; provided that the
Company shall have the right to postpone or withdraw any registration effected pursuant to this Section 2.2 without obligation to any Stockholder or Founder. 

(b) If the registration for which the Company gives notice pursuant to Section 2.2(a) is a registered public offering involving
an underwriting, the Company shall so advise the Stockholders and Founders as a part of the written notice given pursuant to Section 2.2(a). In such event, the right of any Stockholder to include its Registrable Shares or Founder to
include its Founder Registrable Shares in such registration pursuant to Section 2.2 shall be conditioned upon such Stockholder’s or Founder’s participation in such underwriting on the terms set forth herein. All Stockholders
and Founders proposing to distribute their securities through such underwriting shall (together with the Company, Other Holders, and any officers or directors distributing their securities through such underwriting) enter into an underwriting
agreement in customary form with the underwriter or underwriters selected for the underwriting by the Company. Notwithstanding any other provision of this Section 2.2, if the managing underwriter determines that the inclusion of all
shares requested to be registered would adversely affect the offering, the Company may limit (to zero) the number of Registrable Shares and Founder Registrable Shares to be included in the registration and underwriting. The Company shall so advise
all holders of Registrable Shares and Founder Registrable Shares requesting registration, and the number of shares that are entitled to be included in the registration and underwriting shall be allocated in the following manner: the securities of
the Company held by officers and directors of the Company (other than Registrable Shares and Founder Registrable Shares) shall be excluded from such registration and underwriting to the extent deemed advisable by the managing underwriter, and, if a
further limitation on the number of shares is required the other securities held by officers and directors of the Company (including Registrable Shares and Founder Registrable Shares), shall be excluded from such registration and underwriting to the
extent deemed advisable 

  
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by the managing underwriter, and if a further limitation on the number of shares is required, the number of shares that may be included in such registration and underwriting (other than shares to
be sold by the Company) shall be allocated among all Stockholders, Founders and Other Holders requesting registration in proportion, as nearly as practicable, to the respective number of shares of Common Stock (on an
as-converted basis) which they held at the time the Company gave the notice specified in Section 2.2(a) (excluding from calculation any securities excluded from such registrations as set forth
above in this sentence, provided that, if any shares are to be sold in such offering other than on behalf of the Company, then the total number of Registrable Shares permitted to be included therein shall in any event be at least fifty
percent (50%) of the securities included therein (based on aggregate market values). If any Stockholder, Founders or Other Holder would thus be entitled to include more securities than such holder requested to be registered, the excess shall be
allocated among other requesting Stockholders, Founders and Other Holders pro rata in the manner described in the preceding sentence. If any holder of Registrable Shares or Founder Registrable Shares or any officer, director or Other Holder
disapproves of the terms of any such underwriting, such person may elect to withdraw therefrom by written notice to the Company, and any Registrable Shares or Founder Registrable Shares or other securities excluded or withdrawn from such
underwriting shall be withdrawn from such registration. For the avoidance of doubt, no shares to be offered on behalf of the Company shall be required to be excluded from such Registration Statement and underwriting unless all shares held by holders
of securities of the Company (including Stockholders, Founders and Other Holders) have been excluded from such Registration Statement and underwriting. 

(c) The Company shall have the right to select the managing underwriter for any underwritten offering requested pursuant to Section 2.2,
subject to the approval of the holders of a majority of the Registrable Shares, which approval will not be unreasonably withheld or delayed. 

2.3. Registration Procedures.     

(a) If and whenever the Company is required by the provisions of this Agreement to use its reasonable best efforts to effect the registration
of any Registrable Shares under the Securities Act, the Company shall: 
 (i) file with the Commission a Registration Statement with respect
to such Registrable Shares and use its reasonable best efforts to cause that Registration Statement to become and remain effective for one hundred eighty (180) days from the effective date or such lesser period until all such Registrable Shares
are sold; 
 (ii) as expeditiously as possible prepare and file with the Commission any amendments and supplements to the Registration
Statement and the Prospectus as may be necessary to comply with the provisions of the Securities Act (including the anti-fraud provisions thereof) and to keep the Registration Statement effective for one hundred eighty (180) days from the
effective date or such lesser period until all such Registrable Shares are sold; 
 (iii) as expeditiously as possible furnish to each
Selling Stockholder such reasonable number of copies of the Prospectus, including any preliminary prospectus, in conformity with the requirements of the Securities Act, and such other documents as such Selling Stockholder may reasonably request in
order to facilitate the public sale or other disposition of the Registrable Shares owned by such Selling Stockholder; 

  
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 (iv) use its reasonable best efforts to, as expeditiously as possible, register or qualify
the Registrable Shares covered by the Registration Statement under the securities or Blue Sky laws of such states as the Selling Stockholders shall reasonably request, and do any and all other acts and things that may be necessary or desirable to
enable the Selling Stockholders to consummate the public sale or other disposition in such states of the Registrable Shares owned by the Selling Stockholder; provided, however, that the Company shall not be required in connection with
this paragraph (iv) to qualify as a foreign corporation or execute a general consent to service of process in any jurisdiction; 
 (v)
as expeditiously as possible, cause all such Registrable Shares to be listed on each securities exchange or automated quotation system on which similar securities issued by the Company are then listed; 

(vi) promptly provide a transfer agent and registrar for all such Registrable Shares not later than the effective date of such registration
statement; 
 (vii) promptly make available for inspection by the Selling Stockholders, any managing underwriter participating in any
disposition pursuant to such Registration Statement, and any attorney or accountant or other agent retained by any such underwriter or selected by the Selling Stockholders, all financial and other records, pertinent corporate documents and
properties of the Company and cause the Company’s officers, directors, employees and independent accountants to supply all information reasonably requested by any such seller, underwriter, attorney, accountant or agent in connection with such
Registration Statement; 
 (viii) as expeditiously as possible, notify each Selling Stockholder of the time when such Registration
Statement has become effective or a supplement to any Prospectus forming a part of such Registration Statement has been filed; and 
 (ix)
as expeditiously as possible following the effectiveness of such Registration Statement, notify each Selling Stockholder of such Registrable Shares of any request by the Commission for the amending or supplementing of such Registration Statement or
Prospectus. 
 (b) If the Company has delivered a Prospectus to the Selling Stockholders, and after having done so the Prospectus is
amended to comply with the requirements of the Securities Act, the Company shall promptly notify the Selling Stockholders of such amendment and, if necessary, request that the Selling Stockholders immediately cease making offers of Registrable
Shares and return all Prospectuses to the Company. The Company shall promptly provide the Selling Stockholders with revised Prospectuses and, following receipt of the revised Prospectuses, the Selling Stockholders shall be free to resume making
offers of the Registrable Shares. 

  
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 (c) In the event that, in the judgment of the Company, it is advisable to suspend use of a
Prospectus included in a Registration Statement due to pending material developments or other events that have not yet been publicly disclosed and as to which the Company believes public disclosure would be detrimental to the Company, the Company
shall notify all Selling Stockholders to such effect, and, upon receipt of such notice, each such Selling Stockholder shall immediately discontinue any sales of Registrable Shares pursuant to such Registration Statement until such Selling
Stockholder has received copies of a supplemented or amended Prospectus or until such Selling Stockholder is advised in writing by the Company that the then current Prospectus may be used and has received copies of any additional or supplemental
filings that are incorporated or deemed incorporated by reference in such Prospectus. Notwithstanding anything to the contrary herein, the Company shall not exercise its rights under this Section 2.3(c) to suspend sales of Registrable Shares
for a period in excess of ninety (90) days in any three hundred sixty-five-day (365-day) period. 

2.4. Allocation of Expenses. The Company will pay all Registration Expenses for all registrations under this Agreement;
provided, however, that if a registration under Section 2.1 is withdrawn at the request of the Initiating Holders (other than as a result of material information concerning the business or financial condition of the Company which
is made known to the Stockholders after the date on which such registration was requested) and if the Initiating Holders elect not to have such registration counted as a registration requested under Section 2.1, the requesting Stockholders
shall pay the Registration Expenses of such registration pro rata in accordance with the number of their Registrable Shares included in such registration. For purposes of this Section, the term “Registration Expenses” shall mean all
expenses incurred by the Company in complying with this Agreement, including, without limitation, all registration and filing fees, exchange listing fees, printing expenses, fees and expenses of counsel for the Company and the reasonable fees and
expenses (not to exceed $75,000) of one counsel selected by the Selling Stockholders to represent the Selling Stockholders (and, in connection with the Initial Public Offering, the Company shall also pay the fees and expenses of one special counsel
to represent the Significant Stockholders), state Blue Sky fees and expenses, and the expense of any special audits incident to or required by any such registration, but excluding underwriting discounts, selling commissions and the fees and expenses
of Selling Stockholders’ own counsel (other than the counsel selected to represent all Selling Stockholders). 
 2.5.
Indemnification and Contribution.     
 (a) In the event of any registration of any of the Registrable Shares
or Founder Registrable Shares under the Securities Act pursuant to this Agreement, the Company will indemnify and hold harmless the seller of such Registrable Shares or Founder Registrable Shares, each of its directors, partners, and officers, each
underwriter of such Registrable Shares or Founder Registrable Shares, and each other person, if any, who controls such seller or underwriter within the meaning of the Securities Act or the Exchange Act against any losses, claims, damages or
liabilities, joint or several, to which such seller, underwriter or controlling person may become subject under the Securities Act, the Exchange Act, state securities or Blue Sky laws or otherwise, insofar as such losses, claims, damages or
liabilities (or actions 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 Registrable Shares or Founder Registrable
Shares were registered under the Securities Act, any preliminary prospectus or final prospectus contained in the Registration Statement, or any amendment or supplement to such Registration Statement, or arise out of or are based upon the omission or
alleged omission to state a material fact required to 

  
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be stated therein or necessary to make the statements therein not misleading; and the Company will reimburse such seller, underwriter and each such controlling person for any legal or any other
expenses reasonably incurred by such seller, underwriter or controlling person in connection with investigating or defending any such loss, claim, damage, liability or action; provided, however, that the Company will not be liable in
any such case to the extent that any such loss, claim, damage or liability arises out of or is based upon any untrue statement or omission of a material fact made in such Registration Statement, preliminary prospectus or final prospectus, or any
such amendment or supplement, in reliance upon and in conformity with information furnished to the Company, in writing, by or on behalf of such seller, underwriter or controlling person specifically for use in the preparation thereof or to the
extent that such loss, claim, damage or liability arises out of such seller’s failure to deliver a copy of the preliminary or final prospectus or any amendment or supplement thereto. 

(b) In the event of any registration of any of the Registrable Shares or Founder Registrable Shares under the Securities Act pursuant to this
Agreement, each seller of Registrable Shares or Founder Registrable Shares, severally and not jointly, will indemnify and hold harmless the Company, each of its directors and officers and each underwriter (if any) and each person, if any, who
controls the Company or any such underwriter within the meaning of the Securities Act or the Exchange Act, against any losses, claims, damages or liabilities, joint or several, to which the Company, such directors and officers, underwriter or
controlling person may become subject under the Securities Act, Exchange Act, state securities or Blue Sky laws or otherwise, insofar as such losses, claims, damages or liabilities (or actions in respect thereof) arise out of or are based upon any
untrue statement or alleged untrue statement of a material fact contained in any Registration Statement under which such Registrable Shares or Founder Registrable Shares were registered under the Securities Act, any preliminary prospectus or final
prospectus contained in the Registration Statement, or any amendment or supplement to the Registration Statement, or arise out of or are based upon any omission or alleged omission to state a material fact required to be stated therein or necessary
to make the statements therein not misleading, if the statement or omission was made in reliance upon and in conformity with information relating to such seller furnished in writing to the Company by or on behalf of such seller specifically for use
in connection with the preparation of such Registration Statement, preliminary prospectus or final prospectus, amendment or supplement; provided, however, that the obligations of a seller of Registrable Shares or Founder Registrable
Shares hereunder shall be limited to an amount equal to the net proceeds to such seller of Registrable Shares or Founder Registrable Shares sold in connection with such registration. 

(c) Each party entitled to indemnification under this Section (the “Indemnified Party”) shall give notice to the party
required to provide indemnification (the “Indemnifying Party”) promptly after such Indemnified Party has actual knowledge of any claim as to which indemnity may be sought, and shall permit the Indemnifying Party to assume the
defense of any such claim or any litigation resulting therefrom; provided that counsel for the Indemnifying Party, who shall conduct the defense of such claim or litigation, shall be approved by the Indemnified Party (whose approval shall not
be unreasonably withheld); and provided further that the failure of any Indemnified Party to give notice as provided herein shall not relieve the Indemnifying Party of its obligations under this Section except to the extent that the
Indemnifying Party is adversely affected by such failure. The Indemnified Party may participate in such defense at such party’s expense; provided, however, that the Indemnifying Party shall pay 

  
 11 

 such expense if representation of such Indemnified Party by the counsel retained by the Indemnifying Party
would be inappropriate due to actual or potential differing interests between the Indemnified Party and any other party represented by such counsel in such proceeding; provided further that in no event shall the Indemnifying Party be
required to pay the expenses of more than one law firm per jurisdiction as counsel for the Indemnified Party. The Indemnifying Party also shall be responsible for the expenses of such defense if the Indemnifying Party does not elect to assume such
defense. No Indemnifying Party, in the defense of any such claim or litigation shall, except with the consent of each Indemnified Party, consent to entry of any judgment or enter into any settlement 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 of such claim or litigation, and no Indemnified Party shall consent to entry of any judgment or settle such claim or litigation
without the prior written consent of the Indemnifying Party, which consent shall not be unreasonably withheld. 
 (d) In order to provide
for just and equitable contribution in circumstances in which the indemnification provided for in this Section 2.5 is due in accordance with its terms but for any reason is held to be unavailable to an Indemnified Party in respect to any
losses, claims, damages and liabilities referred to herein, then the Indemnifying Party shall, in lieu of indemnifying such Indemnified Party, contribute to the amount paid or payable by such Indemnified Party as a result of such losses, claims,
damages or liabilities to which such party may be subject in such proportion as is appropriate to reflect the relative fault of the Indemnifying Party on the one hand and Indemnified Party on the other in connection with the statements or omissions
which resulted in such losses, claims, damages or liabilities, as well as any other relevant equitable considerations. The relative fault of a party shall be determined by reference to, among other things, whether the untrue or alleged untrue
statement of material fact related to information supplied by such party and the parties’ relative intent, knowledge, access to information and opportunity to correct or prevent such statement or omission. The Company and the sellers of
Registrable Shares or Founder Registrable Shares agree that it would not be just and equitable if contribution pursuant to this Section 2.5 were determined by pro rata allocation or by any other method of allocation which does not take
account of the equitable considerations referred to above. Notwithstanding the provisions of this paragraph of Section 2.5, in no case shall any one seller of Registrable Shares or Founder Registrable Shares be liable or responsible for
any amount in excess of the net proceeds received by such seller of Registrable Shares or Founder Registrable Shares from the offering of Registrable Shares or Founder Registrable Shares; provided, however, that 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 party entitled to contribution will,
promptly after receipt of notice of commencement of any action, suit or proceeding against such party in respect of which a claim for contribution may be made against another party or parties under this Section 2.5, notify such party or
parties from whom contribution may be sought, but the omission so to notify such party or parties from whom contribution may be sought shall not relieve such party from any other obligation it or they may have thereunder or otherwise under this
Section. No party shall be liable for contribution with respect to any action, suit, proceeding or claim settled without its prior written consent, which consent shall not be unreasonably withheld. 

  
 12 

 2.6. Other Matters with Respect to Underwritten Offerings. In the event that
Registrable Shares are sold pursuant to a Registration Statement in an underwritten offering pursuant to Section 2.1, the Company agrees to (a) enter into an underwriting agreement containing customary representations and warranties
with respect to the business and operations of the Company and customary covenants and agreements to be performed by the Company, including without limitation customary provisions with respect to indemnification by the Company of the underwriters of
such offering; (b) use its reasonable best efforts to cause its legal counsel to render customary opinions to the underwriters with respect to the Registration Statement; and (c) use its reasonable best efforts to cause its independent
public accounting firm to issue customary “cold comfort letters” to the underwriters with respect to the Registration Statement. 

2.7. Information by Holder. As a condition to the Company’s obligation to register the Registrable Shares of any holder, such
holder shall furnish to the Company such information regarding such holder and the distribution proposed by such holder as the Company may reasonably request in writing and as shall be required in connection with any registration, qualification or
compliance referred to in this Agreement. 
 2.8. “Stand-Off” Agreement;
Confidentiality of Notices. Each Stockholder agrees not to sell or otherwise transfer or dispose of any Registrable Shares or other securities of the Company held by such Stockholder prior to the initial public offering of the Company’s
Common Stock (and for the avoidance of doubt, not including any securities acquired by it in connection with such offering or at any time thereafter), for a period of one hundred eighty (180) days following the effective date of a Registration
Statement subject to a potential extension (not to exceed fifteen (15) days) to prevent a prohibition on publications by research analysts pursuant to NASD Rule 2711; provided that: 

(a) such agreement shall only apply to the initial public offering of Common Stock of the Company to be sold by or on behalf of the Company in
an underwritten offering; and 
 (b) all stockholders of the Company then holding at least one percent (1%) of the outstanding Common Stock
(on an as converted basis) and all Founders, officers and directors of the Company enter into similar agreements. 
 The Company may impose
stop-transfer instructions with respect to the Registrable Shares or other securities subject to the foregoing restriction until the end of such one hundred eighty-day
(180-day) period. 
 Each Holder further agrees to execute such agreements as may be reasonably
requested by the underwriters in connection with such registration that are consistent with this Section 2.8 or that are necessary to give further effect thereto. 

Any Stockholder receiving any written notice from the Company regarding the Company’s plans to file a Registration Statement shall treat
such notice confidentially and shall not disclose such information to any person other than as necessary to exercise its rights under this Agreement. 

2.9. Limitations on Subsequent Registration Rights. The Company shall not, without the prior written consent of Stockholders holding at
least a majority of the Registrable Shares then held by all Stockholders and a majority of Series B Preferred and Series C Preferred (voting together as a single class on an as-converted basis), enter into any
agreement (other than 

  
 13 

 this Agreement) with any holder or prospective holder of any securities of the Company which grant such
holder or prospective holder rights to include securities of the Company in any Registration Statement, unless (a) such rights to include securities in a registration initiated by the Company or by Stockholders are junior in all respects to the
rights granted to Other Holders under Sections 2.1 and 2.2 of this Agreement, and (b) any such rights to initiate a registration provide that Stockholders are entitled to include Registrable Shares in priority to such holders. 

2.10. Rule 144 Requirements. After the earliest of (i) the closing of the sale of securities of the Company pursuant to a
Registration Statement, (ii) the registration by the Company of a class of securities under Section 12 of the Exchange Act, or (iii) the issuance by the Company of an offering circular pursuant to Regulation A under the Securities
Act, the Company agrees to, so long as any Stockholder is a holder of Registrable Shares: 
 (a) make and keep current public information
about the Company available, as those terms are understood and defined in Rule 144; 
 (b) use its reasonable best efforts to file with the
Commission in a timely manner all reports and other documents required of the Company under the Securities Act and the Exchange Act (at any time after it has become subject to such reporting requirements); and 

(c) furnish to any holder of Registrable Shares upon request (i) a written statement by the Company as to its compliance with the
reporting requirements of Rule 144 and of the Securities Act and the Exchange Act (at any time after it has become subject to such reporting requirements), (ii) a copy of the most recent annual or quarterly report of the Company, and (iii) such
other reports and documents of the Company as such holder may reasonably request to avail itself of any similar rule or regulation of the Commission allowing it to sell any such securities without registration. 

2.11. Termination. All of the Company’s obligations to register Registrable Shares under Sections 2.1 and 2.2 of this Agreement
shall terminate upon the earliest of (i) five (5) years after the closing of the Initial Public Offering, (ii) the date on which no Stockholder holds any Registrable Shares, (iii) a Company Sale or (iv) at such time as SEC Rule
144 or another similar exemption under the Securities Act is available for the sale of all of a holder’s or Founder’s shares without limitation during a three (3) month period without registration. 

3. Right Of Preferred Stockholders. 

(a) The Company shall not issue, sell or exchange, agree to issue, sell or exchange, or reserve or set aside for issuance, sale or exchange,
(A) any shares of its Common Stock, (B) any other equity securities of the Company, including, without limitation, shares of preferred stock, (C) any option, warrant or other right to subscribe for, purchase or otherwise acquire any
equity securities of the Company, or (D) any debt securities convertible into capital stock of the Company (collectively, the “Offered Securities”), unless in each such case the Company shall have first complied with this
Section 3.1. 

  
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 (i) The Company shall deliver to each Stockholder that is a Qualifying Holder a written
notice of any proposed or intended issuance, sale or exchange of Offered Securities (the “Offer”), which Offer shall (w) identify and describe the Offered Securities, (x) describe the price and other terms upon which they are
to be issued, sold or exchanged, and the number or amount of the Offered Securities to be issued, sold or exchanged, (y) identify the persons or entities (if known) to which or with which the Offered Securities are to be offered, issued, sold
or exchanged and (z) offer to issue and sell to or exchange with such Qualifying Holder (A) such portion of the Offered Securities as is equal to (1) one hundred percent (100%) of the Offered Securities multiplied by (2) a
fraction the numerator of which is the aggregate number of shares of Common Stock issued and issuable upon conversion of the Preferred Stock held by such Qualifying Holder and the denominator of which is the total number of shares of Common Stock
issued and issuable upon conversion of all the outstanding shares of Preferred Stock then held by all Qualifying Holders (the “Basic Amount”), and (B) any additional portion of the Offered Securities attributable to the
Basic Amount of other Qualifying Holders as such Qualifying Holder shall indicate it will purchase or acquire should any other Qualifying Holder subscribe for less than its Basic Amount (the “Undersubscription Amount”). 

(ii) If more than one Qualifying Holder indicates that it wishes to purchase the Undersubscription Amount, or a portion thereof, and the
Undersubscription Amount is not sufficient to allow all such Qualifying Holders to purchase the amounts indicated, the Undersubscription Amount shall be allocated among the Qualifying Holders who indicated an interest therein, pro rata based on such
Qualifying Holders’ relative Basic Amounts. 
 (iii) Each Qualifying Holder shall have the right, for a period of thirty
(30) days following the delivery, to purchase or acquire, at the price and upon the other terms specified in the Offer, the number and amount of Offered Securities described above. For the purpose of determining both the numbers of shares each
Qualifying Holder owns and the number of shares each Qualifying Holder is entitled to purchase, each Qualifying Holder shall be treated as one holder together with its affiliates (regardless of whether such affiliates currently own securities of the
Company). 
 (b) To accept an Offer, in whole or in part, a Qualifying Holder must deliver a written notice to the Company prior to the end
of the thirty-day (30-day) period of the Offer, setting forth the portion of the Qualifying Holder’s Basic Amount that such Qualifying Holder elects to purchase
and, if such Qualifying Holder shall elect to purchase all of its Basic Amount, the Undersubscription Amount (if any) that such Qualifying Holder elects to purchase (the “Notice of Acceptance”). If the Basic Amounts subscribed for
by all Qualifying Holders are less than the total of all of the Basic Amounts available for purchase, then each Qualifying Holder who has set forth an Undersubscription Amount in its Notice of Acceptance shall be entitled to purchase, in addition to
the Basic Amount it subscribed for, the Undersubscription Amount it has subscribed for; provided, however, that if the Undersubscription Amount subscribed for exceeds the difference between the total of all of the Basic Amounts
available for purchase and the Basic Amounts subscribed for (the “Available Undersubscription Amount”), each Qualifying Holder who has subscribed for any Undersubscription Amount shall be entitled to purchase only that portion of
the Available Undersubscription Amount as the Undersubscription Amount subscribed for by such Qualifying Holder bears to the total Undersubscription Amounts subscribed for by all Qualifying Holders, subject to rounding by the Board of Directors to
the extent it deems reasonably necessary. 

  
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 (c) The Company shall have ninety (90) days from the expiration of the period set
forth in Section 3.1(b) above to issue, sell or exchange all or any part of such Offered Securities as to which a Notice of Acceptance has not been given by the Qualifying Holders (the “Refused Securities”), but only to the
offerees or Qualifying Holders described in the Offer (if so described therein) and only upon terms and conditions (including, without limitation, unit prices and interest rates) which are not more favorable, in the aggregate, to the acquiring
person or persons or less favorable to the Company than those set forth in the Offer. 
 (d) In the event the Company shall propose to sell
less than all the Refused Securities (any such sale to be in the manner and on the terms specified in Section 3.1(c) above), then each Qualifying Holder may, at its sole option and in its sole discretion, reduce the number or amount of the
Offered Securities specified in its Notice of Acceptance to an amount that shall be not less than the number or amount of the Offered Securities that the Qualifying Holder elected to purchase pursuant to Section 3.1(b) above multiplied by a
fraction, (i) the numerator of which shall be the number or amount of Offered Securities the Company actually proposes to issue, sell or exchange (including Offered Securities to be issued or sold to Qualifying Holders pursuant to
Section 3.1(b) above prior to such reduction) and (ii) the denominator of which shall be the original amount of the Offered Securities. In the event that any Qualifying Holder so elects to reduce the number or amount of Offered Securities
specified in its Notice of Acceptance, the Company may not issue, sell or exchange more than the reduced number or amount of the Offered Securities unless and until such securities have again been offered to the Qualifying Holders in accordance with
Section 3.1(a) above. 
 (e) Upon the closing of the issuance, sale or exchange of all or less than all of the Refused Securities, the
Qualifying Holders shall acquire from the Company, and the Company shall issue to the Qualifying Holders, the number or amount of Offered Securities specified in the Notices of Acceptance, as reduced pursuant to Section 3.1(d) above if the
Qualifying Holders have so elected, upon the terms and conditions specified in the Offer. The purchase by the Qualifying Holders of any Offered Securities is subject in all cases to the preparation, execution and delivery by the Company and the
Qualifying Holders of a purchase agreement relating to such Offered Securities reasonably satisfactory in form and substance to the Qualifying Holders and the Company. 

(f) Any Offered Securities not acquired by the Qualifying Holders or other persons in accordance with Section 3.1(c) above may not be
issued, sold or exchanged until they are again offered to the Qualifying Holders under the procedures specified in this Agreement. 
 (g)
The rights of the Qualifying Holders under this Section 3 shall not apply to: 
 (i) Common Stock issued as a stock dividend to holders
of Common Stock or Preferred Stock or upon any subdivision or combination of shares of Common Stock or Preferred Stock; 
 (ii) the
issuance of any shares of Common Stock upon conversion of shares of convertible preferred stock; 

  
 16 

 (iii) the issuance of (A) shares of Common Stock, or the grant of options therefor
subject in either case to appropriate adjustment in the event of any stock dividend, stock split, combination or other similar recapitalization occurring after the date of this Agreement affecting such shares, issued, or deemed issued to directors,
consultants and employees of the Company or any subsidiary, pursuant to the Company’s 2005 Equity Incentive Plan, as amended (the “2005 Plan”), (B) shares of Common Stock, or the grant of options therefor subject in either case
to appropriate adjustment in the event of any stock dividend, stock split, combination or other similar recapitalization occurring after the date of this Agreement affecting such shares, issued, or deemed issued to directors, consultants and
employees of the Company or any subsidiary, pursuant to the Company’s 2016 Equity Incentive Plan, as amended (the “2016 Plan”, and together with the 2005 Plan, the “Plans”) or (C) shares of Common Stock,
or the grant of options therefore, pursuant to any other plan, agreement or arrangement approved by a majority of the Board of Directors (which majority must include a majority of the Preferred Directors) of the Company; 

(iv) securities issued solely in consideration for the acquisition (whether by merger or otherwise) by the Company or any of its subsidiaries
of all or substantially all of the stock or assets of any other entity; provided that such transaction, and the issuance of shares in connection therewith, has been approved by a majority of the Board of Directors (which majority must include
a majority of the Preferred Directors); 
 (v) securities sold by the Company in an underwritten public offering pursuant to an effective
registration statement under the Securities Act; 
 (vi) securities issued or issuable to financial institutions or lessors in connection
with commercial credit agreements, equipment financings or similar transactions; provided that such transaction, and the issuance of shares in connection therewith, has been approved by a majority of the Board of Directors, (which majority
must include a majority of the Preferred Directors); 
 (vii) securities issued or issuable in connection with strategic transactions
involving the Company and other entities, including (i) joint ventures, manufacturing, marketing or distribution arrangements and technology transfer or development arrangements; provided that such strategic transactions, and the
issuance of shares in connection therewith, has been approved by a majority of the Board of Directors (which majority must include a majority of the Preferred Directors); or 

(viii) securities issued or issuable pursuant to the Series C Purchase Agreement, as such Series C Purchase Agreement may be amended from
time to time in accordance with its terms. 
 3.2. Termination. This Section 3 shall terminate upon the earlier of the following
events: 
  

	 	(a)	 The closing of a Company Sale; or 

 

	 	(b)	 The closing of an Initial Public Offering. 

  
 17 

 4. Transfers of Rights. 

(a) This Agreement, and the rights and obligations of each Stockholder hereunder, may be assigned by such Stockholder to (i) any
affiliate, partner or stockholder of such Stockholder to which such Stockholder transfers at least ten percent (10%) of the aggregate shares of Preferred Stock (or underlying Common Stock) held by such Stockholder (or one hundred percent (100%) of
the Shares purchased by such Stockholder if fewer than five hundred thousand (500,000) (subject to appropriate adjustment in the event of any stock dividend, stock split, combination or other similar recapitalization occurring after the date of this
Agreement and affecting the number of issued and outstanding shares of Preferred Stock)), and such transferee shall be deemed a “Stockholder” for purposes of this Agreement, provided that no transfer shall be made to any entity that
is a competitor of the Company as determined in good faith by the Board of Directors of the Company (including the Preferred Directors); provided further that the transferee provides prior written notice of such assignment to the
Company and agrees in writing to be bound hereby as a “Stockholder”. 
 (b) In connection with any transfer of Founder
Registrable Shares permitted under an applicable restricted stock purchase agreement and made in compliance with the Eighth Amended and Restated Right of First Refusal and Co-Sale Agreement of even date
therewith, the rights and obligations of each Founder under Section 2.2 hereunder may be assigned by such Founder to (i) any person or entity to which at least five hundred thousand (500,000) Founder Registrable Shares (subject to
appropriate adjustment in the event of any stock dividend, stock split, combination or other similar recapitalization occurring after the date of this Agreement and affecting the number of issued and outstanding shares of Common Stock) are
transferred by such Founder (or one hundred percent (100%) of the Founder Registrable Shares held by such Founder if less than five hundred thousand (500,000) (subject to appropriate adjustment in the event of any stock dividend, stock split,
combination or other similar recapitalization occurring after the date of this Agreement and affecting the number of issued and outstanding shares of Common Stock) or (ii) to his spouse or child or a trust or similar entity established for the
benefit of his spouse, his child or himself, and such transferee shall be deemed a Founder for purposes of Section 2.2 of this Agreement; provided that the transferee provides written notice of such assignment to the Company and agrees
in writing to be bound hereby as a “Founder”. 
 (c) Each Institutional Investor (as defined in Section 6(j) below) may
assign this Agreement and its rights and obligations hereunder to any entity that may be considered an affiliate (as defined in Rule 12b-2 promulgated under the Exchange Act) of such Institutional Investor and
such affiliate of such Institutional Investor shall be deemed a “Stockholder” for purposes of this Agreement, and all such affiliates of such Institutional Investor may from time to time reassign their rights to other affiliates of such
Institutional Investor; provided that the transferee provides written notice of such assignment to the Company and agrees to be bound by the terms and conditions set forth herein as a “Stockholder”. 

  
 18 

 5. Covenants 

5.1. Affirmative Covenants. The Company covenants and agrees that, in addition to any other rights provided by law, so long as at least
20,000,000 shares of Preferred Stock (such number to be proportionately adjusted in the event of any stock splits, stock dividends, recapitalizations or similar events occurring after the date of this Agreement affecting the number of issued and
outstanding shares of Preferred Stock) are outstanding, it will perform and observe the following covenants and provisions, unless waived by the prior written consent or affirmative vote of the holders of not less than a majority of the then
outstanding shares of Preferred Stock, voting together as a separate class on an as-converted basis: 

(a) pay and discharge all taxes, assessments and governmental charges or levies imposed upon it or upon its income or profits or business, or
upon any properties belonging to it, prior to the date on which penalties attach thereto, and all lawful claims, which, if unpaid, might become a lien or charge upon any properties of the Company, other than those which are being contested in good
faith by appropriate proceedings; 
 (b) preserve and maintain its corporate existence, rights, franchises and privileges in the
jurisdiction of its incorporation, and qualify and remain qualified as a foreign corporation in each jurisdiction in which such qualification is required, unless the failure to so qualify does not and will not have a material and adverse effect on
the business, operations or financial condition of the Company and preserve and maintain all material licenses and other rights to use patents, processes, licenses, trademarks, trade names, inventions, intellectual property rights or copyrights
owned or possessed by it as are reasonably necessary or advisable for it to conduct its business; and 
 (c) maintain commercially
reasonable and adequate general liability, casualty and product liability insurance, and directors and officers liability insurance in amounts as requested by holders of the a majority of the Preferred Stock, voting together as a separate class on
an as-converted basis and as approved by the Board of Directors. 
 5.2. Inspection. The
Company shall permit each Significant Stockholder or any authorized representative thereof, to visit and inspect the properties of the Company, including its corporate and financial records, and to discuss its business and finances with officers of
the Company, during normal business hours and without disruption of the Company’s business following reasonable notice and as often as may be reasonably requested; provided, however, that the Company shall not be obligated
pursuant to this Section 5.2 to provide access to any information which it reasonably considers to be a trade secret. 
 5.3.
Observer Rights. 
 (a) As long as Perceptive Life Sciences Master Fund Ltd (“Perceptive”) owns not less than fifty
percent (50%) of the shares of the Series B Preferred Stock it holds as of the date hereof (or an equivalent amount of Common Stock issued upon conversion thereof), the Company shall invite a representative of Perceptive to attend all meetings of
its Board of Directors in a nonvoting observer capacity, who shall initially be Keyvan Mirsaeedi-Farahani and, in this respect, shall give such representative copies of all notices, minutes, consents, and other materials that it provides to its
directors at the same time and in the same manner as provided to such directors; provided, however, that such representative shall agree to hold in confidence and trust and to act in a fiduciary manner with respect to all information
so provided; and provided further, that the Company reserves the right to withhold any information and to exclude such representative from any meeting or portion thereof if access to such information or attendance at such meeting could
adversely affect the attorney-client privilege between the Company and its counsel or result in disclosure of trade secrets or a conflict of interest, or if such Investor or its representative is a competitor of the Company. 

  
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 (b) As long as RA Capital Healthcare Fund, L.P. (“RA Capital”) owns not
less than fifty percent (50%) of the shares of the Series B Preferred Stock it holds as of the date hereof (or an equivalent amount of Common Stock issued upon conversion thereof), the Company shall invite a representative of RA Capital to attend
all meetings of its Board of Directors in a nonvoting observer capacity, who shall initially be Neta Batscha, and, in this respect, shall give such representative copies of all notices, minutes, consents, and other materials that it provides to its
directors at the same time and in the same manner as provided to such directors; provided, however, that such representative shall agree to hold in confidence and trust and to act in a fiduciary manner with respect to all information
so provided; and provided further, that the Company reserves the right to withhold any information and to exclude such representative from any meeting or portion thereof if access to such information or attendance at such meeting could
adversely affect the attorney-client privilege between the Company and its counsel or result in disclosure of trade secrets or a conflict of interest, or if such Investor or its representative is a competitor of the Company. 

5.4. Financial Statements and Other Information. The Company shall deliver to each Significant Stockholder: 

(a) within one hundred eighty (180) days after the end of each fiscal year of the Company, an audited balance sheet of the Company as at
the end of such year and audited statements of income and of cash flows of the Company for such year, certified by certified public accountants of recognized regional standing selected by the Company, and prepared in accordance with GAAP; 

(b) within forty-five (45) days after the end of each fiscal quarter of the Company (other than the last quarter in each fiscal year),
an unaudited balance sheet of the Company as at the end of such quarter, and unaudited statements of income and of cash flows of the Company for such fiscal quarter and for the current fiscal year to the end of such fiscal quarter; 

(c) on or before the thirtieth (30th) day following the first day of each fiscal year, a
budget, consisting of a business plan and projected financial statements for such fiscal year, which budget has been approved by the Board of Directors; 

(d) such other notices, information and data with respect to the Company as the Company delivers to the holders of its capital stock
generally at the same time it delivers such items to such holders; and 
 (e) with reasonable promptness, such other information and data
as such Significant Stockholder may from time to time reasonably request. 
 The foregoing financial statements shall be prepared on a consolidated basis if
the Company then has any subsidiaries. The financial statements delivered pursuant to clause (b) above shall be accompanied by a certificate of the chief financial officer of the Company stating that such statements have been prepared in
accordance with GAAP consistently applied (except as noted) and fairly present the financial condition and results of operations of the Company at the date thereof and for the periods covered thereby. 

  
 20 

 Notwithstanding anything else in this Subsection 5.4 to the contrary, the Company may cease providing
the information set forth in this Subsection 5.4 during the period starting with the date 60 days before the Company’s good-faith estimate of the date of filing of a registration statement if it reasonably concludes it must
do so to comply with the Commission rules applicable to such registration statement and related offering; provided that the Company’s covenants under this Subsection 5.4 shall be reinstated at such time as the Company is no longer
actively employing its commercially reasonable efforts to cause such registration statement to become effective. 
 5.5. Material Changes
and Litigation. The Company shall promptly notify the Board of Directors of (a) any material adverse change in the business, prospects, assets or condition, financial or otherwise, of the Company and (b) any litigation or governmental
proceeding or investigation brought or, to the Company’s knowledge, threatened against the Company, or against any Founder, officer, director, key employee or principal stockholder of the Company which, if adversely determined, would have a
material adverse effect on the Company. 
 5.6. Director and Officer and Key Man Insurance. The Company shall maintain term life
insurance on the life of the Chief Executive Officer, currently Maria Palasis, in the amount of One Million Dollars ($1,000,000), with the proceeds payable to the Company. The Company shall maintain, or within ninety (90) days after the date
hereof the Company shall obtain and thereafter maintain, workers’ compensation insurance and insurance with respect to the properties and business of the Company of the kinds and in the amounts not less than is customarily obtained by
corporations in the same or similar business, similarly situated and in similar size, including, without limitation, insurance against loss, damage, fire, theft, public liability and other risks. 

5.7. Agreements with Employees; Options. 

(a) The Company shall require (i) all persons hereafter employed by, or consulting with the Company, and each current employee or person
consulting who has not already done so, to enter into Nondisclosure and Developments Agreements substantially in the Company’s standard form, which form shall be reasonably acceptable to Perceptive; (ii) all persons hereafter employed by
the Company as officers or designated by the Chief Executive Officer or the Board of Directors of the Company as key contributors to the Company to enter into Noncompetition, Nonsolicitation, Nondisclosure and Developments Agreements substantially
in the Company’s standard form which form shall be reasonably acceptable to Perceptive, for a period ending no less than one (1) year following the termination of employment of such key contributor; and (iii) each employee who is
designated by the Chief Executive Officer or the Board of Directors of the Company as a key contributor and has not already done so to enter into Noncompetition, Nonsolicitation, Nondisclosure and Developments Agreements substantially in the
Company’s standard form, or such other form as may be approved by the Board of Directors of the Company, and in either case, which form shall be reasonably acceptable to Perceptive, for a period ending no less than one (1) year following
the termination of employment of such key contributor. For the avoidance of doubt, the form agreements provided to Perceptive in conjunction with the Series C Purchase Agreement are deemed to be reasonably acceptable to Perceptive. 

  
 21 

 (b) Other than options or restricted stock granted to the Founders (which shall be governed
by the terms of the applicable restricted stock agreement), unless otherwise agreed by a majority of the members of the Board of Directors who are not employees of the Company or a subsidiary of the Company (which majority shall include a majority
of the Preferred Directors), all options or restricted stock granted or issued under the Plans shall become exercisable at the rate of twenty-five percent (25%) on the one-year anniversary of the date of
issuance with the remainder vesting in equal monthly installments (2.0833%) over the subsequent three (3) years so long as the holder continues to be an employee or consultant of the Company. 

5.8. Directors. 
 (a)
The Company shall promptly reimburse in full each director of the Company who is not an employee of the Company and who was elected as a director solely or in part by the holders of Preferred Stock for all of his reasonable out-of-pocket expenses incurred in attending each meeting of the Board of Directors of the Company or any committee thereof. 

(b) The Board of Directors shall meet at least quarterly unless otherwise agreed by all of the members of the Board of Directors. 

5.9. Reservation of Common Stock. The Company shall reserve and maintain a sufficient number of shares of Common Stock for issuance
upon conversion of all of the outstanding Shares. 
 5.10. Related Party Transactions. Except for transactions contemplated by this
Agreement, the Series C Purchase Agreement, the Ninth Amended and Restated Stockholders’ Voting Agreement, or the Eighth Amended and Restated Right of First Refusal and Co-Sale Agreement, each entered
into on or about the date hereof, or transactions on customary terms related to such person’s employment, the Company shall not enter into any agreement with any stockholder, officer or director of the Company, or, to the extent known to the
Company, any “affiliate” or “associate” of such persons (as such terms are defined in the rules and regulations promulgated under the Securities Act), including without limitation any agreement or other arrangement providing for
the furnishing of services by, rental of real or personal property from, or otherwise requiring payments to, any such person or entity, without the consent of at least a majority of the members of the Company’s Board of Directors having no
interest in such agreement or arrangement. 
 5.11. Matters Requiring Director Approval. The Company hereby covenants and agrees with
each of the Investors that it shall not, without approval of the Board of Directors (which approval must include either the Series B Director or the Series C Director): 

(a) Incur any new indebtedness for borrowed money that, together with any other indebtedness for borrowed money incurred since the Series C
Original Issue Date, exceeds Two Hundred Fifty Thousand Dollars ($250,000) in the aggregate; 
 (b) Pay or declare any dividend or
distribution on any shares of its capital stock, or apply any of its assets to the redemption, retirement, purchase or acquisition, directly or indirectly, through subsidiaries or otherwise, of any shares of its capital stock, except for
(i) the repurchase by the Company of capital stock held by an employee, director or consultant 

  
 22 

 
of the Company at the original purchase price upon termination of their employment or services with the Company, or (ii) as contemplated by the Charter or by the Eighth Amended and Restated
Right of First Refusal and Co-Sale Agreement, dated as the date hereof (the “ROFR Agreement”), as such ROFR Agreement may be amended from time to time in accordance with its terms. 

(c) guarantee, directly or indirectly, or permit any subsidiary to guarantee, directly or indirectly, any indebtedness except for trade
accounts of the Company or any subsidiary arising in the ordinary course of business; 
 (d) make any investment inconsistent with any
investment policy approved by the Board of Directors; 
 (e) change the principal business of the Company, enter new lines of business, or
exit the current line of business; 
 (f) sell, assign, license, pledge, or encumber material technology or intellectual property, other
than licenses granted in the ordinary course of business; or 
 (g) enter into any corporate strategic relationship involving the payment,
contribution, or assignment by the Company or to the Company of money or assets greater than $250,000. 
 5.12. Termination of
Covenants. All covenants of the Company contained in this Section 5 shall terminate upon the earlier of the closing of a Company Sale or the closing of an Initial Public Offering. 

6. General. 
 (a)
Severability. The invalidity or unenforceability of any provision of this Agreement shall not affect the validity or enforceability of any other provision of this Agreement. 

(b) Specific Performance. In addition to any and all other remedies that may be available at law in the event of any breach of this
Agreement, each Stockholder shall be entitled to specific performance of the agreements and obligations of the Company hereunder and to such other injunctive or other equitable relief as may be granted by a court of competent jurisdiction. 

(c) Governing Law; Dispute Resolution. This Agreement shall be governed by and construed in accordance with the internal laws of the
State of Delaware (without reference to the conflicts of law provisions thereof). The parties (a) hereby irrevocably and unconditionally submit to the jurisdiction of the state courts of Delaware and to the jurisdiction of the United States
District Court for the District of Delaware for the purpose of any suit, action or other proceeding arising out of or based upon this Agreement, (b) agree not to commence any suit, action or other proceeding arising out of or based upon this
Agreement except in the state courts of Delaware or the United States District Court for the District of Delaware, and (c) hereby waive, and agree not to assert, by way of motion, as a defense, or otherwise, in any such suit, action or
proceeding, any claim that it is not subject personally to the jurisdiction of the above-named courts, 

  
 23 

 
that its property is exempt or immune from attachment or execution, that the suit, action or proceeding is brought in an inconvenient forum, that the venue of the suit, action or proceeding is
improper or that this Agreement or the subject matter hereof may not be enforced in or by such court. WAIVER OF JURY TRIAL: EACH PARTY HEREBY WAIVES ITS RIGHTS TO A JURY TRIAL OF ANY CLAIM OR CAUSE OF ACTION BASED UPON OR ARISING OUT OF THIS
AGREEMENT, THE OTHER TRANSACTION DOCUMENTS, THE SECURITIES OR THE SUBJECT MATTER HEREOF OR THEREOF. THE SCOPE OF THIS WAIVER IS INTENDED TO BE ALL-ENCOMPASSING OF ANY AND ALL DISPUTES THAT MAY BE FILED IN ANY
COURT AND THAT RELATE TO THE SUBJECT MATTER OF THIS TRANSACTION, INCLUDING, WITHOUT LIMITATION, CONTRACT CLAIMS, TORT CLAIMS (INCLUDING NEGLIGENCE), BREACH OF DUTY CLAIMS, AND ALL OTHER COMMON LAW AND STATUTORY CLAIMS. THIS SECTION HAS BEEN FULLY
DISCUSSED BY EACH OF THE PARTIES HERETO AND THESE PROVISIONS WILL NOT BE SUBJECT TO ANY EXCEPTIONS. EACH PARTY HERETO HEREBY FURTHER WARRANTS AND REPRESENTS THAT SUCH PARTY HAS REVIEWED THIS WAIVER WITH ITS LEGAL COUNSEL, AND THAT SUCH PARTY
KNOWINGLY AND VOLUNTARILY WAIVES ITS JURY TRIAL RIGHTS FOLLOWING CONSULTATION WITH LEGAL COUNSEL. 
 (d) Notices. All notices,
requests, consents, and other communications under this Agreement shall be in writing and shall be deemed delivered (i) two business days after being sent by registered or certified mail, return receipt requested, postage prepaid or
(ii) one business day after being sent via a reputable nationwide overnight courier service guaranteeing next business day delivery, in each case to the intended recipient as set forth below: 

If to the Company, Lyra Therapeutics, Inc., 480 Arsenal Street, Watertown, MA 02472, Attn: Chief Executive Officer, or at such other address or
addresses as may have been furnished in writing by the Company to the Preferred Stockholders, with a copy to Latham & Watkins LLP, 200 Clarendon Street, 27th Floor, Boston, Massachusetts
02116, Attn: Johan V. Brigham, Esq.; 
 If to any Founder, at his address set forth on the signature page hereto, or at such other address
or addresses as may have been furnished to the Company and the Investors in writing by such Founder; or 
 If to an Investor, at his or its
address set forth on Schedule A, or at such other address or addresses as may have been furnished to the Company in writing by such an Investor, with a copy to: Cooley LLP, 500 Boylston Street, 14th Floor, Boston, MA 02116-3736, Attention:
Marc Recht. 
 Any party may give any notice, request, consent or other communication under this Agreement using any other means (including,
without limitation, personal delivery, messenger service, telecopy, first class mail or electronic mail), but no such notice, request, consent or other communication shall be deemed to have been duly given unless and until it is actually received by
the party for whom it is intended. Any party may change the address to which notices, requests, consents or other communications hereunder are to be delivered by giving the other parties notice in the manner set forth in this Section. 

  
 24 

 (e) Complete Agreement. This Agreement constitutes the entire agreement and
understanding of the parties hereto with respect to the subject matter hereof and supersedes all prior agreements and understandings relating to such subject matter. 

(f) Amendments and Waivers. Any term of this Agreement may be amended or terminated and the observance of any term of this Agreement
may be waived (either generally or in a particular instance and either retroactively or prospectively), with the written consent of the Company and the holders of at least a majority of the Registrable Shares held by all of the Stockholders;
provided that any amendment, termination or waiver of any term of Section 2.2 of this Agreement which has a disproportionately negative effect on the Founders shall also require the written consent of the holders of at least a
majority of the Founder Registrable Shares; and provided further that this Agreement may be amended with the consent of the holders of less than all Registrable Shares only in a manner which affects all such holders in the same fashion
and no amendment shall be effected to any provision hereof which will have the effect of reducing the approval threshold for any matter herein unless such amendment is approved by holders holding such threshold of votes; and provided
further that no consent shall be required for an amendment pursuant to Section 6(k). Any such amendment, termination or waiver effected in accordance with this Section 6(f) shall be binding on all parties hereto, even
if they do not execute such consent. No waivers of or exceptions to any term, condition or provision of this Agreement, in any one or more instances, shall be deemed to be, or construed as, a further or continuing waiver of any such term, condition
or provision. 
 (g) Pronouns. Whenever the context may require, any pronouns used in this Agreement shall include the corresponding
masculine, feminine or neuter forms, and the singular form of nouns and pronouns shall include the plural, and vice versa. 
 (h)
Counterparts; Facsimile Signatures. This Agreement may be executed and delivered by facsimile signature and in any number of counterparts, each of which shall be deemed to be an original, and all of which together shall constitute one and the
same document. 
 (i) Section Headings. The section headings are for the convenience of the parties and in no way alter, modify,
amend, limit or restrict the contractual obligations of the parties. 
 (j) One Holder. For purposes of determining the number of
Shares held by each Stockholder under this Agreement, (i) all affiliates of North Bridge Venture Partners (collectively, “North Bridge”) shall be deemed to be one holder, (ii) all affiliates of Polaris Venture Partners
(collectively, “Polaris”) shall be deemed to be one holder and (iii) all affiliates of Perceptive Life Science Master Fund, Ltd. (collectively, “Perceptive”) shall be deemed to be one holder (with North Bridge,
Polaris, Perceptive and Intersouth Partners VII, L.P. collectively the “Institutional Investors” and each an “Institutional Investor”). 

(k) Addition of Investors. Notwithstanding anything to the contrary contained herein, if the Company issues additional shares of
Series C Preferred after the date hereof, any purchaser of such shares of Series C Preferred shall become a party to this Agreement by executing and delivering an additional counterpart signature page to this Agreement, and

  
 25 

 
thereafter shall be deemed an “Investor” for all purposes hereunder. No action or consent by the Investors shall be required for such joinder to this Agreement by such additional
Investor so long as such additional Investor has agreed in writing to be bound by all of the obligations as an “Investor” hereunder. 

(l) Prior IRA. The Prior IRA is hereby amended and restated in its entirety and restated herein. Such amendment and restatement is
effective upon execution of this Agreement by the Company, the Founders and the Investors constituting the requisite votes necessary to amend such Prior IRA. Upon such execution, all provisions of, rights granted and covenants made in the Prior IRA
are hereby waived, released and superseded in their entirety and shall have no further force or effect. 
 [Remainder of page intentionally
left blank.] 

  
 26 

 IN WITNESS WHEREOF, the parties have executed this Eighth Amended and Restated Investor
Rights Agreement as of the date first written above. 
  

			
	LYRA THERAPEUTICS, INC.

 
			
		
	By:	 	/s/ Maria Palasis

 
			
	Name:	 	Maria Palasis
	Title:	 	President and Chief Executive Officer

 [Signature Page to Eighth Amended and Restated Investor Rights Agreement] 

 IN WITNESS WHEREOF, the parties have executed this Eighth Amended and Restated Investor
Rights Agreement as of the date first written above. 
  

			
	INVESTORS:
	
	ARROWMARK FUNDAMENTAL OPPORTUNITY FUND, L.P.
		
	By:	 	 ArrowMark Partners GP, LLC
 Its General
Partner

 
			
		
	By:	 	 /s/ David Corkins

	Name:	 	David Corkins
	Title:	 	Managing Member

 [Signature Page to Eighth Amended and Restated Investor Rights Agreement] 

 IN WITNESS WHEREOF, the parties have executed this Eighth Amended and Restated Investor
Rights Agreement as of the date first written above. 
  

			
	INVESTORS:
	
	INTERSOUTH PARTNERS VII, L.P.
		
	By:	 	 Intersouth Associates VII, LLC
 Its General
Partner

		
	By:	 	/s/ Dennis Dougherty
	Name:	 	Dennis Dougherty
	Title:	 	 Member Manager

 [Signature Page to Eighth Amended and Restated Investor Rights Agreement] 

 IN WITNESS WHEREOF, the parties have executed this Eighth Amended and Restated Investor
Rights Agreement as of the date first written above. 
  

			
	INVESTORS:
	
	MERIDIAN SMALL CAP GROWTH FUND
		
	By:	 	 ArrowMark Colorado Holdings, LLC
 Its
Investment Advisor

		
	By:	 	/s/ David Corkins
	Name:	 	David Corkins
	Title:	 	 Managing Member

 [Signature Page to Eighth Amended and Restated Investor Rights Agreement] 

 IN WITNESS WHEREOF, the parties have executed this Eighth Amended and Restated Investor
Rights Agreement as of the date first written above. 
  

			
	INVESTORS:
	
	NORTH BRIDGE VENTURE PARTNERS V-A, L.P.
		
	By:	 	 North Bridge Venture Management V, L.P.
 Its
General Partner

		
	By:	 	 NBVM GP, LLC
 Its General
Partner

		
	By:	 	/s/ Edward T. Anderson
	Name:	 	Edward T. Anderson
	Title:	 	 Manager

  

			
	NORTH BRIDGE VENTURE PARTNERS V-B, L.P.
		
	By:	 	 North Bridge Venture Management V, L.P.
 Its
General Partner

		
	By:	 	 NBVM GP, LLC
 Its General
Partner

		
	By:	 	/s/ Edward T. Anderson
	Name:	 	Edward T. Anderson
	Title:	 	 Manager

  

			
	NORTH BRIDGE VENTURE PARTNERS VI, L.P.
		
	By:	 	 North Bridge Venture Management VI, L.P.
 Its
General Partner

		
	By:	 	 NBVM GP, LLC
 Its General
Partner

		
	By:	 	/s/ Edward T. Anderson
	Name:	 	Edward T. Anderson
	Title:	 	 Manager

 [Signature Page to Eighth Amended and Restated Investor Rights Agreement] 

 IN WITNESS WHEREOF, the parties have executed this Eighth Amended and Restated Investor
Rights Agreement as of the date first written above. 
  

			
	INVESTORS:
	
	PERCEPTIVE LIFE SCIENCES MASTER FUND, LTD.
		
	By:	 	/s/ Joseph Edelman
	Name:	 	Joseph Edelman
	Title:	 	 Investment Manager

  

			
	PERCEPTIVE LS (A), LLC
		
	By:	 	 Perceptive LS GP, LLC
 Its
Manager

		
	By:	 	/s/ Joseph Edelman
	Name:	 	Joseph Edelman
	Title:	 	 Investment Manager

 [Signature Page to Eighth Amended and Restated Investor Rights Agreement] 

 IN WITNESS WHEREOF, the parties have executed this Eighth Amended and Restated Investor
Rights Agreement as of the date first written above. 
  

			
	INVESTORS:
	
	POLARIS VENTURE PARTNERS V, L.P.
		
	By:	 	 Polaris Venture Management Co. V, L.L.C.
 Its
General Partner

		
	By:	 	/s/ Lauren Crockett
	Name:	 	Lauren Crockett
	Title:	 	
Attorney-in-fact

  

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

		
	By:	 	/s/ Lauren Crockett
	Name:	 	Lauren Crockett
	Title:	 	
Attorney-in-fact

  

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

		
	By:	 	/s/ Lauren Crockett
	Name:	 	Lauren Crockett
	Title:	 	
Attorney-in-fact

  

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

		
	By:	 	/s/ Lauren Crockett
	Name:	 	Lauren Crockett
	Title:	 	
Attorney-in-fact

 [Signature Page to Eighth Amended and Restated Investor Rights Agreement] 

 IN WITNESS WHEREOF, the parties have executed this Eighth Amended and Restated Investor
Rights Agreement as of the date first written above. 
  

			
	INVESTORS:
	
	POLARIS VENTURE PARTNERS IV, L.P.
		
	By:	 	 Polaris Venture Management Co. IV, L.L.C.

Its General Partner

		
	By:	 	/s/ Lauren Crockett
	Name:	 	Lauren Crockett
	Title:	 	
Attorney-in-Fact

  

			
	POLARIS VENTURE PARTNERS ENTREPRENEURS’ FUND IV, L.P.
		
	By:	 	 Polaris Venture Management Co. IV, L.L.C.

Its General Partner

		
	By:	 	/s/ Lauren Crockett
	Name:	 	Lauren Crockett
	Title:	 	
Attorney-in-Fact

 [Signature Page to Eighth Amended and Restated Investor Rights Agreement] 

 IN WITNESS WHEREOF, the parties have executed this Eighth Amended and Restated Investor
Rights Agreement as of the date first written above. 
  

			
	INVESTORS:
	
	RA CAPITAL HEALTHCARE FUND, L.P.
		
	By:	 	 RA Capital Healthcare Fund GP, LLC
 Its
General Partner

		
	By:	 	 /s/ Rajeev Shah
  

	Name:	 	Rajeev Shah
	Title:	 	 Manager

 [Signature Page to Eighth Amended and Restated Investor Rights Agreement] 

 IN WITNESS WHEREOF, the parties have executed this Eighth Amended and Restated Investor
Rights Agreement as of the date first written above. 
  

	
	INVESTORS AND FOUNDERS:
	
	 /s/ Carmichael S. Roberts

 

	 Carmichael S. Roberts

 [Signature Page to Eighth Amended and Restated Investor Rights Agreement] 

 IN WITNESS WHEREOF, the parties have executed this Eighth Amended and Restated Investor
Rights Agreement as of the date first written above. 
  

	
	INVESTORS AND FOUNDERS:
	
	 /s/ George M. Whitesides

	George M. Whitesides

 [Signature Page to Eighth Amended and Restated Investor Rights Agreement] 

 IN WITNESS WHEREOF, the parties have executed this Eighth Amended and Restated Investor
Rights Agreement as of the date first written above. 
  

	
	INVESTORS AND FOUNDERS:
	
	 /s/ Robert Langer

	Robert Langer

 [Signature Page to Eighth Amended and Restated Investor Rights Agreement] 

 Lyra Therapeutics, Inc. 

Signature Page 
 to 

Series C Convertible Preferred Stock and Warrant Purchase Agreement, 

Eighth Amended and Restated Investor Rights Agreement, 

Eighth Amended and Restated Right of First Refusal and Co-Sale Agreement, and 

Ninth Amended and Restated Stockholders’ Voting Agreement 

By executing and delivering this signature page, the undersigned hereby (i) agrees to become a party to and to be bound by the terms and
conditions of that certain Series C Convertible Preferred Stock and Warrant Purchase Agreement by and among Lyra Therapeutics, Inc., a Delaware corporation (the “Company”), and the other parties thereto, dated as of January 10,
2020 (the “Purchase Agreement”), as a “Purchaser” thereunder, (ii) acknowledges having read the representations in Section 4 of the Purchase Agreement entitled “Representations of Purchasers” and
(iii) represents and warrants that the statements contained in such Section 4 are complete and accurate with respect to the undersigned as a Purchaser. 

The undersigned further hereby joins in, becomes a party to and agrees to be bound by the terms and conditions of: 

(a)    that certain Eighth Amended and Restated Investor Rights Agreement, dated as of January 10, 2020, by and among
the Company and the other parties thereto (the “Investors’ Rights Agreement”) as an “Investor” thereunder; 

(b)    that certain Eighth Amended and Restated Right of First Refusal and Co-Sale
Agreement, dated as of January 10, 2020, by and among the Company and the other parties named therein (the “Co-Sale Agreement”) as an “Investor” thereunder; and 

(c)    that certain Ninth Amended and Restated Stockholders’ Voting Agreement, dated as of January 10, 2020, by
and among the Company and the other parties thereto (the “Voting Agreement”) as an “Investor” and “Stockholder” thereunder. 

This Joinder Agreement is contingent upon the Company’s receipt of funds from the undersigned in the amount of the Aggregate Purchase
Price (as set forth below), which the undersigned hereby agrees it shall provide to the Company by a date no later than January 28, 2020. The undersigned hereby authorizes this signature page to be attached to the Purchase Agreement, the
Investors’ Rights Agreement, the Co-Sale Agreement and the Voting Agreement or counterparts thereof. 
 Number
of Shares of 
 Series C Preferred Stock: 5,153,178 
 Number of
Warrant Shares 
 (Common Stock): 1,545,953 
 Aggregate
Purchase Price: $1,999,999.92 

  
 [Signature Page Follows]

 In witness Whereof, the undersigned has executed this Signature Page to Series C Convertible
Preferred Stock and Warrant Purchase Agreement, Eighth Amended and Restated Investors’ Rights Agreement, Eighth Amended and Restated Right of First Refusal and Co-Sale Agreement, and Ninth Amended and
Restated Stockholders’ Voting Agreement as of the 28th day of January, 2020. 

Clifton Capital L.P. 
  

			
	By:	 	 /s/ Tracy Lange

	Name:	 	Tracy Lange
	Title:	 	 Authorised Representative
 For Clifton Capital
LP

	
	AGREED TO AND ACCEPTED:
	
	LYRA THERAPEUTICS, INC.
	
	 /s/ Maria Palasis

	Maria Palasis
	President and Chief Executive Officer

 Lyra Therapeutics, Inc. 

Signature Page 
 to 

Series C Convertible Preferred Stock and Warrant Purchase Agreement, 

Eighth Amended and Restated Investor Rights Agreement, 

Eighth Amended and Restated Right of First Refusal and Co-Sale Agreement, and 

Ninth Amended and Restated Stockholders’ Voting Agreement 

By executing and delivering this signature page, the undersigned hereby (i) agrees to become a party to and to be bound by the terms and
conditions of that certain Series C Convertible Preferred Stock and Warrant Purchase Agreement by and among Lyra Therapeutics, Inc., a Delaware corporation (the “Company’’), and the other parties thereto, dated as of
January 10, 2020 (the “Purchase Agreement”), as a “Purchaser” thereunder, (ii) acknowledges having read the representations in Section 4 of the Purchase Agreement entitled “Representations of
Purchasers” and (iii) represents and warrants that the statements contained in such Section 4 are complete and accurate with respect to the undersigned as a Purchaser. 

The undersigned further hereby joins in, becomes a party to and agrees to be bound by the terms and conditions of: 

(a)    that certain Eighth Amended and Restated Investor Rights Agreement, dated as of January 10, 2020, by and among
the Company and the other parties thereto (the “Investors’ Rights Agreement”) as an “Investor” thereunder; 

(b)    that certain Eighth Amended and Restated Right of First Refusal and Co-Sale
Agreement, dated as of January 10, 2020, by and among the Company and the other parties named therein (the “Co-Sale Agreement”) as an “Investor” thereunder; and 

(c)    that certain Ninth Amended and Restated Stockholders’ Voting Agreement, dated as of January 10, 2020, by
and among the Company and the other parties thereto (the “Voting Agreement”) as an “Investor” and “Stockholder” thereunder. 

This Joinder Agreement is contingent upon the Company’s receipt of funds from the undersigned in the amount of the Aggregate Purchase
Price (as set forth below), which the undersigned hereby agrees it shall provide to the Company by a date no later than January 27, 2020. The undersigned hereby authorizes this signature page to be attached to the Purchase Agreement, the
Investors’ Rights Agreement, the Co-Sale Agreement and the Voting Agreement or counterparts thereof. 
 Number
of Shares of 
 Series C Preferred Stock: 3,607,224 
 Aggregate
Purchase Price: $1,399,999.71 

  
 [Signature Page Follows]

 In witness Whereof, the undersigned has executed this Signature Page to Series C Convertible
Preferred Stock and Warrant Purchase Agreement, Eighth Amended and Restated Investors’ Rights Agreement, Eighth Amended and Restated Right of First Refusal and Co-Sale Agreement, and Ninth Amended and
Restated Stockholders’ Voting Agreement as of the 27 day of January, 2020. 
  

					
	Soleus Private Equity Fund I, L.P.
	
	By: Soleus Private Equity GP I, LLC, as general partner

 
					
			
		 	By:	 	 /s/ Steven Musumeci

		 	Name:	 	Steven Musumeci
		 	Title:	 	Chief Operating Officer

	
	AGREED TO AND ACCEPTED:
	
	LYRA THERAPEUTICS, INC.
	
	 /s/ Maria Palasis

	Maria Palasis
	President and Chief Executive Officer

 Lyra Therapeutics, Inc. 

Signature Page 
 to 

Series C Convertible Preferred Stock and Warrant Purchase Agreement, 

Eighth Amended and Restated Investor Rights Agreement, 

Eighth Amended and Restated Right of First Refusal and Co-Sale Agreement, and 

Ninth Amended and Restated Stockholders’ Voting Agreement 

By executing and delivering this signature page, the undersigned hereby (i) agrees to become a party to and to be bound by the terms and
conditions of that certain Series C Convertible Preferred Stock and Warrant Purchase Agreement by and among Lyra Therapeutics, Inc., a Delaware corporation (the “Company”), and the other parties thereto, dated as of January 10,
2020 (the “Purchase Agreement”), as a “Purchaser” thereunder, (ii) acknowledges having read the representations in Section 4 of the Purchase Agreement entitled “Representations of Purchasers” and
(iii) represents and warrants that the statements contained in such Section 4 are complete and accurate with respect to the undersigned as a Purchaser. 

The undersigned further hereby joins in, becomes a party to and agrees to be bound by the terms and conditions of: 

(a)    that certain Eighth Amended and Restated Investor Rights Agreement, dated as of January 10, 2020, by and among
the Company and the other parties thereto (the “Investors’ Rights Agreement”) as an “Investor” thereunder; 

(b)    that certain Eighth Amended and Restated Right of First Refusal and Co-Sale
Agreement, dated as of January 10, 2020, by and among the Company and the other parties named therein (the “Co-Sale Agreement”) as an “Investor” thereunder; and 

(c)    that certain Ninth Amended and Restated Stockholders’ Voting Agreement, dated as of January 10, 2020, by
and among the Company and the other parties thereto (the “Voting Agreement”) as an “Investor” and “Stockholder” thereunder. 

This Joinder Agreement is contingent upon the Company’s receipt of funds from the undersigned in the amount of the Aggregate Purchase
Price (as set forth below), which the undersigned hereby agrees it shall provide to the Company by a date no later than 31, 2020. The undersigned hereby authorizes this signature page to be attached to the Purchase Agreement, the Investors’
Rights Agreement, the Co-Sale Agreement and the Voting Agreement or counterparts thereof. 
 Number of Shares of

 Series C Preferred Stock: 235,965 
 Number of Warrant Shares

 (Common Stock): 70,789 
 Aggregate Purchase Price:
$91,580.38 

  
 [Signature Page Follows]

 In witness Whereof, the undersigned has executed this Signature Page to Series C Convertible
Preferred Stock and Warrant Purchase Agreement, Eighth Amended and Restated Investors’ Rights Agreement, Eighth Amended and Restated Right of First Refusal and Co-Sale Agreement, and Ninth Amended and
Restated, Stockholders’ Voting Agreement as of the 31st day of January, 2020. 
  

			
	Emmant, LLC
		
	By:	 	 /s/ Guido Neels

	Name:	 	Guido Neels
	Title:	 	Manager

	
	AGREED TO AND ACCEPTED:
	
	LYRA THERAPEUTICS, INC.
	
	 /s/ Maria Palasis

	Maria Palasis
	President and Chief Executive Officer

 Lyra Therapeutics, Inc. 

Signature Page 
 to 

Series C Convertible Preferred Stock and Warrant Purchase Agreement, 

Eighth Amended and Restated Investor Rights Agreement, 

Eighth Amended and Restated Right of First Refusal and Co-Sale Agreement, and 

Ninth Amended and Restated Stockholders’ Voting Agreement 

By executing and delivering this signature page, the undersigned hereby (i) agrees to become a party to and to be bound by the terms and
conditions of that certain Series C Convertible Preferred Stock and Warrant Purchase Agreement by and among Lyra Therapeutics, Inc., a Delaware corporation (the “Company”), and the other parties thereto, dated as of January 10,
2020 (the “Purchase Agreement”), as a “Purchaser” thereunder, (ii) acknowledges having read the representations in Section 4 of the Purchase Agreement entitled “Representations of Purchasers” and
(iii) represents and warrants that the statements contained in such Section 4 are complete and accurate with respect to the undersigned as a Purchaser. 

The undersigned further hereby joins in, becomes a party to and agrees to be bound by the terms and conditions of: 

(a)    that certain Eighth Amended and Restated Investor Rights Agreement, dated as of January 10, 2020, by and among
the Company and the other parties thereto (the “Investors’ Rights Agreement”) as an “Investor” thereunder; 

(b)    that certain Eighth Amended and Restated Right of First Refusal and Co-Sale
Agreement, dated as of January 10, 2020, by and among the Company and the other parties named therein (the “Co-Sale Agreement”) as an “Investor” thereunder; and 

(c)    that certain Ninth Amended and Restated Stockholders’ Voting Agreement, dated as of January 10, 2020, by
and among the Company and the other parties thereto (the “Voting Agreement”) as an “Investor” and “Stockholder” thereunder. 

This Joinder Agreement is contingent upon the Company’s receipt of funds from the undersigned in the amount of the Aggregate Purchase
Price (as set forth below), which the undersigned hereby agrees it shall provide to the Company by a date no later than January 31, 2020. The undersigned hereby authorizes this signature page to be attached to the Purchase Agreement, the
Investors’ Rights Agreement, the Co-Sale Agreement and the Voting Agreement or counterparts thereof. 
 Number
of Shares of 
 Series C Preferred Stock: 644,147 
 Number of
Warrant Shares 
 (Common Stock): 193,244 
 Aggregate Purchase
Price: $249,999.90 

  
 [Signature Page Follows]

 In witness Whereof, the undersigned has executed this Signature Page to Series C Convertible
Preferred Stock and Warrant Purchase Agreement, Eighth Amended and Restated Investors’ Rights Agreement, Eighth Amended and Restated Right of First Refusal and Co-Sale Agreement, and Ninth Amended and
Restated Stockholders’ Voting Agreement as of the 29th day of January, 2020. 
  

			
	RFA Holding LLC
		
	By:	 	 /s/ Giles Carmichael

	Name:	 	Giles Carmichael
	Title:	 	President

	
	AGREED TO AND ACCEPTED:
	
	LYRA THERAPEUTICS, INC.
	
	 /s/ Maria Palasis

	Maria Palasis
	President and Chief Executive Officer

 SCHEDULE A 

Investors 
  

			
	 Intersouth Partners VII, L.P.
 406 Blackwell
Street
 Suite 200
 Durham, NC 27701

 
 North Bridge Venture Partners V-A, L.P.

950 Winter Street, Suite 4600
 Waltham, MA 02451

 
 North Bridge Venture Partners V-B, L.P.

950 Winter Street, Suite 4600
 Waltham, MA 02451

 
 North Bridge Venture Partners VI, L.P.

950 Winter Street, Suite 4600
 Waltham, MA 02451

 
 Polaris Venture Partners IV, L.P.

1 Marina Park Drive
 10th Floor

Boston, MA 02210
  

Polaris Venture Partners Entrepreneurs’ Fund IV, L.P.
 1
Marina Park Drive
 10th Floor
 Boston, MA 02210

 
 Polaris Venture Partners V, L.P.

1 Marina Park Drive
 10th Floor

Boston, MA 02210
  

Polaris Venture Partners Entrepreneurs’ Fund V, L.P.
 1
Marina Park Drive
 10th Floor
 Boston, MA 02210
	  	 Polaris Venture Partners Founders’ Fund V, L.P.

1 Marina Park Drive
 10th Floor

Boston, MA 02210
  

Polaris Venture Partners Special Founders’ Fund V, L.P.
 1
Marina Park Drive
 10th Floor
 Boston, MA 02210

 
 Citigroup Global Markets, Inc., as Custodian

FBO Robert Duane Dickson IRA
 50 Deepwood Road

Darien, CT 06820
  

Duane Dickson
 [xxx]

[xxx]
  

Carmichael Roberts
 [xxx]

[xxx]
  

Robert Langer
 [xxx]

[xxx]
  

Michael Loberg
 [xxx]

[xxx]
  

George M. Whitesides
 [xxx]

[xxx]
  

Mountainview Ventures, LLC
 20 River Oaks Circle

Pikesville, MD 21208

			
	 Shaw Warren
 [xxx]

[xxx]
  

Paramjith Anand
 [xxx]

[xxx]
  

Milan Mrksich
 [xxx]

[xxx]
  

MS Pace LP
 8500 Normandale Lake Blvd

Suite 1070
 Bloomington, MN 55437

 
 Broadview Ventures I LLC

265 Franklin Street, Suite 1902
 Boston, MA 02110

 
 Emmant, LLC

c/o Guido Neels
 7916 Dean Road

Indianapolis, IN 46240
  

Endoluminal Therapeutics, Inc.
 c/o Dr. Marvin Slepian

2540 N Yellow Flower Trail
 Tucson, AZ 85715

 
 RA Capital Healthcare Fund, L.P.

c/o RA Capital Management, L.P.
 200 Berkeley Street

18th Floor
 Boston, MA 02116

Attn: General Counsel
  

Soleus Private Equity Fund I, L.P.
 104 Field Point Road

Second Floor
 Greenwich, CT 06830

 
 ArrowMark Fundamental Opportunity Fund, L.P.

100 Fillmore Street
 Denver, CO 80206
	  	 Meridian Small Cap Growth Fund
 100 Fillmore
Street
 Denver, CO 80206
  

Perceptive Life Sciences Master Fund, Ltd
 51 Astor Place, 10th Floor
 New York, NY 10003
  

Perceptive LS (A), LLC
 51 Astor Place, 10th Floor
 New York, NY 10003
  

RFA Holding LLC
 401 Park Avenue South

New York, NY 10016
  

Clifton Capital LP
 c/o MJM Europe Limited

Corporate House
 10 Oratory Street

Naxxar
 NXR 2504

Malta
  

Giselle Weiss
 Ochsengasse 14

CH-4123 Allschwil
 Switzerland

 
 Tracie Smart

76 Benton Road
 Somerville, MA 02143

 
 Milan Mrksich

560 N. Lincoln Street
 Hinsdale, IL 60521

 
 Karen Shannon

552 N. Vine St
 Hinsdale IL 60521EX-4.3

 Exhibit 4.3 

Schedule of Holders of 

Warrants to Purchase Common Stock 
  

											
	 Holder
	  	Warrant
No.	  	Number of Shares	 	  	Issuance Date	  	Expiration Date /
Void After
	 ArrowMark Fundamental Opportunity Fund, L.P.
	  	1-A	  	 	386,488	 	  	1/10/2020	  	1/10/2030
	 Meridian Small Cap Growth Fund
	  	1-B	  	 	772,976	 	  	1/10/2020	  	1/10/2030
	 Intersouth Partners VII, L.P.
	  	2	  	 	309,190	 	  	1/10/2020	  	1/10/2030
	 North Bridge Venture Partners V-A, L.P.
	  	3-A	  	 	726,219	 	  	1/17/2020	  	1/17/2030
	 North Bridge Venture Partners V-B, L.P.
	  	3-B	  	 	355,949	 	  	1/17/2020	  	1/17/2030
	 North Bridge Venture Partners VI, L.P
	  	3-C	  	 	463,785	 	  	1/17/2020	  	1/17/2030
	 Polaris Venture Partners V, L.P.
	  	4-A	  	 	596,697	 	  	1/10/2020	  	1/10/2030
	 Polaris Venture Partners Entrepreneurs’ Fund V, L.P.
	  	4-B	  	 	11,630	 	  	1/10/2020	  	1/10/2030
	 Polaris Venture Partners Founders’ Fund V, L.P.
	  	4-C	  	 	4,087	 	  	1/10/2020	  	1/10/2030
	 Polaris Venture Partners Special Founders’ Fund V, L.P.
	  	4-D	  	 	5,967	 	  	1/10/2020	  	1/10/2030
	 Perceptive Life Sciences Master Fund, Ltd
	  	5-A	  	 	4,869,753	 	  	1/17/2020	  	1/17/2030
	 Perceptive LS (A), LLC
	  	5-B	  	 	11,710,597	 	  	1/17/2020	  	1/17/2030
	 RA Capital Healthcare Fund, L.P.
	  	6	  	 	386,488	 	  	1/17/2020	  	1/17/2030
	 Soleus Private Equity Fund I, L.P.
	  	7	  	 	1,082,167	 	  	1/27/2020	  	1/27/2030
	 Emmant, LLC
	  	8	  	 	70,789	 	  	1/31/2020	  	1/31/2030
	 Clifton Capital LP
	  	9	  	 	1,545,953	 	  	1/29/2020	  	1/29/2030
	 RFA Holding LLC
	  	11	  	 	193,244	 	  	1/29/2020	  	1/29/2030

 THIS WARRANT AND THE UNDERLYING SECURITIES HAVE BEEN ACQUIRED FOR INVESTMENT AND HAS NOT BEEN REGISTERED
UNDER THE SECURITIES ACT OF 1933, AS AMENDED. THIS WARRANT AND THE UNDERLYING SECURITIES MAY NOT BE SOLD OR TRANSFERRED WITHOUT (I) AN EFFECTIVE REGISTRATION STATEMENT RELATED THERETO, (II) AN OPINION OF COUNSEL FOR THE HOLDER,
SATISFACTORY TO THE COMPANY, THAT SUCH REGISTRATION IS NOT REQUIRED UNDER THE ACT OR (III) RECEIPT OF A NO ACTION LETTER FROM THE SECURITIES AND EXCHANGE COMMISSION. COPIES OF THE AGREEMENT COVERING THE ACQUISITION OF THIS WARRANT AND
RESTRICTING ITS TRANSFER MAY BE OBTAINED AT NO COST BY WRITTEN REQUEST MADE BY THE HOLDER OF RECORD OF THIS WARRANT TO THE SECRETARY OF THE COMPANY AT THE PRINCIPAL EXECUTIVE OFFICES OF THE COMPANY. 

Void after _________________ 

LYRA THERAPEUTICS, INC. 

COMMON STOCK WARRANT 

Warrant No. __ 
 THIS
CERTIFIES THAT, for value received, ___________ or its registered assigns (hereinafter called the “Holder”) is entitled to purchase from Lyra Therapeutics, Inc., a Delaware corporation, with its principal place of business at
480 Arsenal Way, Watertown, MA 02472 (the “Company”), at any time after the date specified in Section 1 hereof and ending at 5:00 p.m. Eastern Time on the Expiration Date, as such term is defined in Section 1 hereof,
up to ______ shares of the Company’s common stock, par value $0.001 per share (the “Common Stock”), subject to adjustment as set forth herein. 

This Warrant (the “Warrant”) is being issued pursuant to the terms of that certain Series C Convertible Preferred Stock and
Warrant Purchase Agreement, dated as of January 10, 2020, by and among, the Company and the purchasers set forth on Schedule A therein (the “Purchase Agreement”). This Warrant may be exercised in whole or in part, at any time
through to the Expiration Date (as defined below) at the option of the Holder. 
 1. Definitions. As used herein, the
following terms shall have the following respective meanings. Any capitalized terms not defined herein shall have the meaning given to them in the Purchase Agreement. 

(a) “Exercise Price” shall mean the fair market value of the Company’s Common Stock following the Initial Closing (as
determined by the Board of Directors, in good faith, based on the most recent independent third party valuation of the Company available following the Initial Closing performed pursuant to Section 409A of the Code, and taking into account any
changes to the Company’s business between the date of such third party valuation and the Initial Closing), subject to adjustments pursuant to Section 5 below. 

 (b) Subject to Section 2, “Expiration Date” shall
mean the period ending on the earlier of (i) _________________, (ii) immediately prior to the closing of the initial public offering of Common Stock (“IPO”), (iii) immediately prior to the closing of a Liquidation Transaction, or
(iv) unless terminated earlier as provided below. 
 (c) “Liquidation Transaction” shall be deemed to be occasioned
by, or to include, any Deemed Liquidation Event (as defined in the Amended and Restated Certificate of Incorporation of the Company, as in effect as of the date hereof). 

(d) “Warrant Shares” shall mean the shares of the Common Stock issuable upon exercise of this Warrant, subject to adjustments
pursuant to the terms herein, including but not limited to adjustment pursuant to Section 5 below. 
 2.
Term and Exercise Schedule. This Warrant shall be exercisable, in whole or in part, at any time, through the Expiration Date. If the Expiration Date is occasioned by either the closing of the IPO or a Liquidation Transaction and to the
extent that the fair market value of one Warrant Share (as determined in accordance with Section 3.3(b)) would be greater than the Exercise Price in effect on such date immediately prior to such IPO or Liquidation
Transaction, and the Holder has not exercised this Warrant pursuant to Sections 3.1 and 3.2 below as to all Warrant Shares, then this Warrant shall automatically be deemed to be Net Exercised pursuant to
Section 3.3(a) as to all Warrant Shares for which it shall not previously have been exercised, effective immediately prior to and contingent upon the closing of the IPO or Liquidation Transaction. In the event of an IPO or
Liquidation Transaction where the fair market value of one Warrant Share as determined in accordance with Section 3.3(b) would be less than the Exercise Price in effect immediately prior to such IPO or Liquidation
Transaction, then this Warrant will expire immediately prior to the closing of such IPO or Liquidation Transaction. 
 3. Method of
Exercise; Payment; Issuance of New Warrant. The purchase right represented by this Warrant may be exercised by the Holder, in whole or in part, by: 

3.1. the surrender of this Warrant (with an executed notice of exercise in the form attached hereto as Attachment A
and an duly executed Investment Representation Statement in the form attached hereto as Attachment B) by delivery to the Company at its address set forth above (or such other address as it may designate by notice in writing to the Holder);
and 
 3.2. the payment to the Company, by check, wire transfer, forgiveness of indebtedness, or any combination of the foregoing, of an
amount equal to the then applicable Exercise Price per share multiplied by the number of Warrant Shares then being purchased. 
 If this
Warrant should be exercised, the Company shall, upon surrender of this Warrant, execute and deliver a certificate representing the Common Stock issued to the Holder upon such exercise, and if this Warrant should be exercised in part only, a new
Warrant of the same tenor evidencing the rights of the Holder thereof to purchase the balance of the Warrant 

  
 2 

 
Shares purchasable hereunder. Upon receipt by the Company of this Warrant and such notice of exercise, together with, if applicable, the aggregate Exercise Price, at such office, or by the stock
transfer agent or warrant agent of the Company at its office, the Holder shall be deemed to be the holder of record of the applicable Warrant Shares, notwithstanding that the stock transfer books of the Company shall then be closed. The Company
shall pay any and all documentary stamp or similar issue or transfer taxes payable in respect of the issue or delivery of the Warrant Shares. 

3.3. Net Exercise. 

(a) Conversion Right. In addition to and without limiting the rights of the Holder under the terms of this Warrant, the Holder
may elect to convert this Warrant or any portion thereof (the “Conversion Right”) into Warrant Shares, the aggregate value of such Warrant Shares shall be equal to the value of this Warrant or the portion thereof being converted.
The Conversion Right may be exercised automatically as provided herein and by the Holder by surrender of this Warrant at the principal office of the Company together with notice of the Holder’s intention to exercise the Conversion Right, in
which event the Company shall issue to the Holder a number of Warrant Shares computed using the following formula: 
  
 

 
 Where: 

X - The number of Warrant Shares to be issued to the Holder upon exercise of the
Conversion Right. 
 Y - The number of Warrant Shares issuable upon exercise of this
Warrant (or such lesser number as are being exercised). 
 A - The fair market value
of one Warrant Share, as determined pursuant to Section 3.3(b) hereof, as of the time the Conversion Right is exercised pursuant to this Section 3. 

B - Exercise Price for one Warrant Share under this Warrant (as adjusted to the date
of such calculations). 
 Notwithstanding the foregoing, this Warrant shall be deemed to have converted into Warrant Shares pursuant to this
Section 3.3(a) upon the Expiration Date if not previously exercised or converted before such date. 
 (b)
Fair Market Value. For purposes of Section 3.3(a), the fair market value of one Warrant Share shall be determined by the Board of Directors of the Company, acting in good faith; provided, however, that:

 (i) where a public market exists for the Common Stock at the time of such exercise, then: 

(A) the fair market value of one Warrant Share shall be the last closing price per share of the Common Stock on the principal national
securities exchange on which the Common Stock is listed or admitted to trading; or 

  
 3 

 (B) the average of the bid and asked price per share as reported in the “pink
sheets” published by the National Quotation Bureau, Inc. if the Common Stock is not listed or traded on any exchange; or 
 (C) if
such quotations are not available, the fair market value of that number of shares of the Common Stock into which one Warrant Share is convertible on the date such notice was received by the Company, as determined in good faith by the Board of
Directors of the Company. 
 (ii) if the Warrant is exercised in connection with the IPO, the fair market value of one Warrant Share shall
be the per share offering price of Common Stock to the public in the IPO; and 
 (iii) all such determinations to be appropriately adjusted
for stock dividend, stock split, stock combination or other similar transactions during the applicable calculation period. 
 4. Stock
Fully Paid; Reservation of Warrant Shares. All shares of stock which may be issued upon the exercise of the rights represented by this Warrant will, upon issuance, be duly authorized, validly issued, fully paid and nonassessable, and free
from all taxes, liens, encumbrances and charges with respect to the issue thereof, except for the restrictions on transfer provided herein or under applicable federal and state securities laws and any liens or encumbrances created by or imposed by
the Holder. During the period within which the rights represented by this Warrant may be exercised, the Company covenants that it will at all times cause to be reserved and kept available out of its authorized and unissued capital stock a sufficient
number of shares of its Common Stock to provide for the exercise of the rights represented by this Warrant into Common Stock. 
 5.
Adjustment of Exercise Price and Number of Warrant Shares. The number and kind of Warrant Shares purchasable upon the exercise of this Warrant and the Exercise Price shall be subject to adjustment from time to time upon the occurrence
of certain events, as follows: 
 5.1. Reclassification. Upon any event whereby all of the outstanding shares of Common
Stock are reclassified, exchanged, combined, substituted or replaced for, into, with or by Company securities of a different class and/or series, then from and after the consummation of such event, this Warrant will be exercisable for the number,
class and series of Company securities that the Holder would have had, if the Warrant Shares been outstanding on and as of the consummation of such event, provided that the aggregate purchase price shall remain the same and subject to further
adjustment thereafter from time to time in accordance with the provisions of this Warrant. The provisions of this Section 5.1 shall similarly apply to successive reclassifications, exchanges, combinations, substitutions,
replacements or similar events. 

  
 4 

 5.2 Subdivision or Combination of Warrant Shares. If the Company at any time
while this Warrant remains outstanding and unexpired shall subdivide the outstanding Common Stock into a greater number of shares, the number of Warrant Shares purchasable hereunder shall be proportionally increased and the Exercise Price shall be
proportionately decreased, provided that the aggregate purchase price shall remain the same. If the Company at any time while this Warrant remains outstanding and unexpired shall combine or consolidate the outstanding Common Stock, into a lesser
number of shares, then the number of Warrant Shares purchasable hereunder shall be proportionately decreased and the Exercise Price shall be proportionately increased, provided that the aggregate purchase price shall remain the same. 

5.3. Stock Dividends. If the Company at any time while this Warrant is outstanding and unexpired declares or pays a dividend or
distribution (other than as set forth in Sections 5.1 and 5.2) on the outstanding shares of Common Stock payable in Common Stock or other securities or property (other than cash), then upon exercise of this Warrant, for each Warrant
Share acquired, Holder shall receive, without additional cost to Holder, the total number and kind of securities and property which Holder would have received had Holder owned the Warrant Shares of record as of the date the dividend or distribution
occurred. 
 5.4. Adjustment of Number of Warrant Shares. Upon each adjustment in the Exercise Price, the number of shares of
stock purchasable hereunder shall be adjusted, to the nearest whole share, to the product obtained by multiplying the number of Warrant Shares purchasable immediately prior to such adjustment in the Exercise Price by a fraction, the numerator of
which shall be the Exercise Price immediately prior to such adjustment and the denominator of which shall be the Exercise Price immediately thereafter. 

5.5 Notice/Certificate of Adjustment. Upon each adjustment of the Exercise Price, class and/or number of Warrant Shares, the
Company, at the Company’s expense, shall notify the Holder in writing within a reasonable time setting forth the adjustments to the Exercise Price, class and/or number of Warrant Shares and facts upon which any such adjustment is based. The
Company shall, upon written request from the Holder, furnish the Holder with a certificate of its Chief Executive Officer or Chief Financial Officer, including computations of such adjustment and the Exercise Price, class and number of shares of
Common Stock in effect upon the date of such adjustment. 
 6. Fractional Warrant Shares. No fractional Warrant Shares will be
issued in connection with any exercise hereunder, but in lieu of such fractional shares the Company shall make a cash payment therefor upon the basis of the Exercise Price then in effect. 

7. Representations of the Company. The Company represents and warrants to the Holder that
(i) Section 2.2(j) of the Schedule of Exceptions sets forth a true and correct pro forma capitalization table of the Company reflecting the Company’s fully diluted equity capitalization immediately
following the Initial Closing (including all shares, options (including shares reserved under the Stock Plan), warrants, and other rights to acquire capital stock) and (ii) such capitalization table includes the names of the Company’s
securityholders, the number of shares of capital stock issued or issuable to such securityholder, and the fully diluted percentage interest of each such securityholder. 

  
 5 

 8. Compliance with Securities Act;
Non-transferability of Warrant; Disposition of Shares of Stock. 
 8.1. Compliance
with Securities Act. The Holder, by acceptance hereof, agrees that this Warrant and the Warrant Shares are being acquired for investment and that he, she or it will not offer, sell or otherwise dispose of this Warrant or any Warrant Shares
except under circumstances which will not result in a violation of the Securities Act of 1933, as amended (the “Act”). Upon exercise of this Warrant, the Holder hereof shall confirm in writing, in a form attached hereto as
Attachment B, that the Warrant Shares so purchased are being acquired for investment and not with a view toward distribution or resale. In addition, the Holder shall provide such additional information regarding such Holder’s financial
and investment background, as the Company may reasonably request, as is relevant for purposes of determining the Holder’s suitability with respect to a purchase of the Warrant Shares. All Warrant Shares (unless registered under the Act) shall
be stamped or imprinted with a legend in substantially the following form (in addition to any legend required under applicable state securities laws): 

THIS SECURITY HAS NOT BEEN REGISTERED UNDER THE SECURITIES ACT OF 1933, AS AMENDED. THIS SECURITY MAY NOT BE SOLD OR TRANSFERRED WITHOUT
(I) AN EFFECTIVE REGISTRATION STATEMENT RELATED THERETO, (II) AN OPINION OF COUNSEL FOR THE HOLDER, SATISFACTORY TO THE COMPANY, THAT SUCH REGISTRATION IS NOT REQUIRED UNDER THE ACT OR (III) RECEIPT OF A NO ACTION LETTER FROM THE
SECURITIES AND EXCHANGE COMMISSION. COPIES OF THE WARRANT AGREEMENT COVERING THE ACQUISITION OF THIS SECURITY AND RESTRICTING ITS TRANSFER MAY BE OBTAINED AT NO COST BY WRITTEN REQUEST MADE BY THE HOLDER OF RECORD OF THIS SECURITY TO THE SECRETARY
OF THE COMPANY AT THE PRINCIPAL EXECUTIVE OFFICES OF THE COMPANY. 
 8.2. Transferability of Warrant. This Warrant may not be
transferred or assigned in whole or in part without (i) an effective registration statement related thereto, (ii) an opinion of counsel for the Holder, satisfactory to the Company, that such registration is not required under the Act or
(iii) receipt of a no action letter from the Securities and Exchange Commission (together, “Securities Law Compliance Guarantees”); provided, however, that the Warrant may be transferred in whole or in part
without Securities Law Compliance Guarantees upon any of the following provided that the transferee agrees in writing, by way of documentation reasonably acceptable to the Company, to be subject to the terms hereof to the same extent as if he/she
were an original Holder hereunder: 
 (a) A transfer of the Warrant by a Holder who is a natural person during such Holder’s lifetime
or on death by will or intestacy to such Holder’s immediate family or to any custodian or trustee for the account of such Holder or such Holder’s immediate family, or a transfer of the Warrant by a Holder that is a trust to a natural
person who is the beneficiary of such trust, to such beneficiary’s immediate family or to any custodian or trustee for the account of such beneficiary or such beneficiary’s immediate family. “Immediate family” as used
herein shall mean spouse, lineal descendant, father, mother, brother, or sister of the Holder; 

  
 6 

 (b) A transfer of the Warrant to the Company; 

(c) A transfer of the Warrant to a parent, subsidiary or affiliate of a Holder ; or 

(d) A transfer of the Warrant by a Holder which is a limited or general partnership to any of its partners or former partners (with, a, b and
c, a “Permitted Transfer”). 
 9. Rights of Stockholders. No Holder of this Warrant shall be entitled to vote
or receive dividends or be deemed the holder of stock or any other securities of the Company which may at any time be issuable on the exercise hereof for any purpose, nor shall anything contained herein be construed to confer upon the Holder of this
Warrant, as such, any of the rights of a stockholder of the Company or any right to vote for the election of directors or upon any matter submitted to stockholders at any meeting thereof, or to give or withhold consent to any corporate action
(whether upon any recapitalization, issuance of stock, reclassification of stock, change of par value or change of stock to no par value, consolidation, merger, conveyance, or otherwise) or to receive notice of meetings, or to receive dividends or
subscription rights or otherwise until this Warrant has been exercised and the Warrant Shares shall have become deliverable, as provided herein. 

10. Governing Law. This Warrant and all acts and transactions pursuant hereto and the rights and obligations of the parties
hereto shall be governed, construed and interpreted in accordance with the laws of the state of Delaware, without giving effect to principles of conflicts of law. 

11. Miscellaneous. The headings in this Warrant are for purposes of convenience and reference only, and shall not be deemed to
constitute a part hereof. All notices and other communications shall be delivered by hand or mailed by first-class registered or certified mail, postage prepaid, to the respective addresses provided in the Purchase Agreement, or to such other
address as the Company or Holder may designate to the other parties hereto. 
 12. Purchase Agreement. This Warrant is a
Warrant referred to in the Purchase Agreement and is entitled to all the benefits provided therein. 
 13. Loss, Theft or Destruction
of Warrant. Upon receipt by the Company of evidence reasonably satisfactory to it of the loss, theft, mutilation or destruction of this Warrant and of indemnity or security reasonably satisfactory to it, the Company will make and deliver an
affidavit of lost warrant which shall carry the same rights carried by this Warrant, stating that such affidavit of lost warrant is issued in replacement of this Warrant, making reference to the original date of issuance of this Warrant (and any
successors hereto) and dated as of such cancellation, in lieu of this Warrant. 

  
 7 

 14. Amendment and Waiver. Any term of this Warrant may be amended or waived
only with the written consent of the Company and the holders of at least a majority of the Warrant Shares then issuable upon exercise of the Warrants then outstanding that were issued pursuant to the Purchase Agreement. Any amendment or waiver
effected in accordance with this Section 14 shall be binding upon the Holder and each transferee of the Warrant, each future holder of all such Warrants, and the Company. 

15. Counterparts: Facsimile/Electronic Signature. This Warrant may be executed in counterparts, all of which together shall
constitute one and the same agreement. Any signature page delivered electronically or by facsimile shall be binding to the same extent as an original signature page with regards to any agreement subject to the terms hereof or any amendment thereto.

 (Remainder of Page Intentionally Left Blank) 

  
 8 

 IN WITNESS WHEREOF, the Company has caused this Warrant to be duly executed by its officers,
thereunto duly authorized this ___ day of _______, 2020. 
  

			
	LYRA THERAPEUTICS, INC.

 
			
		
	By:	 	 

 
			
	Name:	 	Maria Palasis
	Title:	 	Chief Executive Officer

 [Signature Pages to Common Stock Warrant] 

 ATTACHMENT A TO WARRANT 

NOTICE OF EXERCISE 
 TO: Lyra Therapeutics,
Inc. 
 1. The undersigned hereby elects to purchase ____________ shares of Common Stock of Lyra Therapeutics, Inc. as defined in that
certain Series C Convertible Preferred Stock and Warrant Purchase Agreement, dated as of January 10, 2020, and pursuant to the terms of the attached Warrant, and tenders herewith payment of the purchase price of such shares in full, together
with all applicable transfer taxes, if any. 
 2. The undersigned hereby elects to convert the attached Warrant into Warrant Shares in the
manner specified in Section 3.3 of the Warrant. This conversion is exercised with respect to _______________________ of the Shares covered by the Warrant. 

[Strike paragraph above that does not apply.] 

3. Please issue a certificate or certificates representing said shares of stock in the name of the undersigned or in such other name as is
specified below: 
  

							
		  	Name:	  	 	  	

							
				
		  	Address:	  	 	  	

							
				
		  	              	  	 	  	
				
		  		  	 	  	

 4. The undersigned represents that the aforesaid shares of stock are being acquired for the account of the
undersigned for investment and not with a view to, or for resale in connection with, the distribution thereof and that the undersigned has no present intention of distributing or reselling such shares. In support thereof, the undersigned has
executed an Investment Representation Statement attached hereto as Attachment B. 
  

			
	[HOLDER]

 
			
		
	By:	 	 

 
			
	Title:	 	 

 
			
	Date:	 	 

 ATTACHMENT B TO WARRANT 

INVESTMENT REPRESENTATION STATEMENT 
  

							
	PURCHASER	 	:	 	  
	  	
	COMPANY	 	:	 	Lyra Therapeutics, Inc.	  	
	SECURITY	 	:	 	  
	  	
	AMOUNT	 	:	 	  
	  	
	DATE	 	:	 	  
	  	

 In connection with the purchase of the above-listed securities and underlying stock (the
“Securities”), the undersigned represents to the Company the following: 
 (a) We are purchasing these Securities for our
own account for investment purposes only and not with a view to, or for the resale in connection with, any “distribution” thereof for purposes of the Securities Act of 1933, as amended (the “Act”). 

(b) We understand that the Securities have not been registered under the Act in reliance upon a specific exemption therefrom, which exemption
depends upon, among other things, the bona fide nature of our investment intent as expressed herein. In this connection, we understand that, in the view of the Securities and Exchange Commission (the “SEC”), the statutory basis for
such exemption may be unavailable if our representation was predicated solely upon a present intention to hold these Securities for the minimum capital gains period specified under tax statutes, for a deferred sale, for or until an increase or
decrease in the market price of the Securities, or for a period of one year or any other fixed period in the future. 
 (c) We further
understand that the Securities must be held indefinitely unless subsequently registered under the Act or unless an exemption from registration is otherwise available. Moreover, we understand that the Company is under no obligation to register the
Securities. In addition, we understand that the certificate evidencing the Securities will be imprinted with a legend which prohibits the transfer of the Securities unless they are registered or such registration is not required in the opinion of
counsel for the Company. 
 (d) We are aware of the provisions of Rule 144, promulgated under the Act, which, in substance, permits limited
public resale of “restricted securities” acquired, directly or indirectly, from the issuer thereof (or from an affiliate of such issuer), in a non-public offering subject to the satisfaction of
certain conditions. 
 (e) We further understand that at the time we wish to sell the Securities there may be no public market upon which to
make such a sale. 
  

			
	[HOLDER]
		
		 	 
		 	(signature)
		
		 	 
		 	(title)

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