EX-4.31



 Exhibit 4.31 

LICENSE AGREEMENT 

This LICENSE AGREEMENT (this “Agreement”), dated as of July 25, 2017 (the “Effective Date”), is
entered into by and between Cellectis S.A., a corporation existing and registered under the laws of France, located at 8 rue de la Croix Jarry, 75013 Paris, France (“Cellectis”), and Calyxt, Inc., a corporation existing
and registered under the laws of Delaware, located at 600 County Road D West, Suite 8, New Brighton, MN 55112, USA (“Calyxt”) (each a “Party” and collectively, the “Parties”). 

W I T N E S S E T H: 

WHEREAS, Cellectis owns or otherwise controls certain Intellectual Property Rights and desires to grant to Calyxt, and Calyxt desires to
receive from Cellectis, a license to use and otherwise exploit such Intellectual Property Rights, in each case upon the terms and conditions set forth herein. 

NOW, THEREFORE, in consideration of the mutual covenants and agreements hereinafter set forth and for other good and valuable consideration,
the receipt and sufficiency of which are hereby acknowledged, the Parties hereby agree as follows: 
 ARTICLE 1 

DEFINITIONS 

Section 1.01. Definitions. (a) For purposes of this Agreement, the following terms shall have the following
meanings: 
 “Affiliate” means, with respect to any Person, any other Person directly or indirectly
controlling, controlled by, or under common control with such other Person, whether now or in the future. For purposes of this definition, (i) “control” when used with respect to any Person means the power to direct the
management and policies of such Person, directly or indirectly, whether through the ownership of voting securities, by contract or otherwise, and the terms “controlling” and “controlled” have correlative meanings
and (ii) neither Cellectis nor any of its Subsidiaries shall be considered to be an Affiliate of Calyxt or any of its Subsidiaries (and vice versa). 

“Applicable Law” means, with respect to any Person, any transnational, domestic or foreign federal, state or local law
(statutory, common or otherwise), constitution, treaty, convention, ordinance, code, rule, regulation, order, injunction, judgment, decree, ruling or other similar requirement enacted, adopted, promulgated or applied by a Governmental Authority that
is binding upon or applicable to such Person, as amended unless expressly specified otherwise. 
 “Bare
Sublicense” means any sublicense granted by Calyxt to any third party of rights to some or all of the Licensed Cellectis Patents pursuant to Section 2.03, without any Calyxt Licensed Product developed by or in collaboration with Calyxt.


  
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 “Bare Sublicense Revenue” means any and all consideration, payments and
revenue (including the fair market value of any non-cash consideration) received by Calyxt pursuant to any Bare Sublicense.  

“Business Day” means a day, other than Saturday, Sunday or other day on which commercial banks in Paris, France are
authorized or required by Applicable Law to close. 
 “Calyxt Field” means the field of researching,
developing and commercializing agricultural and food products, including, but not limited to traits, seeds, proteins, oils, carbohydrates, food, and food and animal feed ingredients, excluding any application in connection with animals and animal
cells. 
 “Calyxt Improvement” means any improvements, modifications, refinements to, enhancements, derivatives or
combinations of, any Licensed Cellectis IP made by Calyxt or any of its Affiliates after the Effective Date and all Intellectual Property Rights in any of the foregoing. 

“Calyxt Improvement Patents” means any Patents owned or controlled by Calyxt or any of its Affiliates Covering any Calyxt
Improvements. 
 “Calyxt Licensed Products” means any and all products (i) the creation, generation, development,
making or use of which is, in whole or in part, Covered by a Licensed Cellectis Patent, or (ii) which is created, generated, bred or made by use of a process Covered by a Licensed Cellectis Patent. For sake of clarity, any plant or seed which
contains one or more modifications made using a process Covered by any of the Licensed Cellectis Patents, as well as any progeny of such plant or seed, any part of such plant or seed, and any product derived from such plant or seed (such as, for
example, meal and oil derived from any soybean), is a Calyxt Licensed Product. 
 “Cellectis Improvement” means any
improvements, modifications or refinements to, or enhancements or derivatives of any Licensed Cellectis IP made by Cellectis or any of its Affiliates after the Effective Date and all Intellectual Property Rights in any of the foregoing. 

“Confidential Information” means any and all non-public, proprietary or other confidential information disclosed by a
Party (“disclosing party”) to the other Party (“receiving party”) and includes all information licensed hereunder without the need for any further notice or marking, excluding any information that: (i) the
receiving party independently develops without reference to the disclosed information; (ii) the receiving party independently receives on a non-confidential and authorized basis from a source other than the disclosing party; (iii) becomes
public knowledge through no fault of the receiving party; or (iv) is in the public domain at the time the receiving party receives the disclosed information. 

“Cover” means, with respect to any product, service or process, and any Intellectual Property Right, that the
manufacture, use, offer for sale, sale, distribution, importation, development or other commercialization of such product, service or process would, but for any ownership of or license under such Intellectual Property Right, constitute an
infringement, misappropriation or other violation of any of such Intellectual Property Right. “Covered” and “Covering” have correlative meanings. 

  
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 “Exclusively Licensed Cellectis Patents” means any and all Licensed
Cellectis Patents exclusively related to the Calyxt Field for which Calyxt is granted exclusive rights under the Calyxt License. 

“Governmental Authority” means any transnational, domestic or foreign federal, state or local governmental, regulatory
or administrative authority, department, court, agency or official, including any political subdivision thereof. 

“Intellectual Property Rights” means any and all intellectual property rights or similar proprietary rights throughout
the world, including all (i) national and multinational statutory invention registrations and similar statutory rights, patents and patent applications, including all provisionals, non-provisionals, divisionals, continuations,
continuations-in-part, reissues, renewals, reexaminations, extensions, supplemental protection certificates and the equivalents of any of the foregoing in any jurisdiction, and all inventions disclosed in any of the foregoing
(“Patents”); (ii) trademarks, service marks, certification marks, logos, trade names, trade dress, domain names and other indications of origin, including all registrations and applications for registration of, and all goodwill
associated with, any of the foregoing (“Trademarks”); (iii) copyrights and registrations and applications for registration thereof, including all derivative works, moral rights, renewals, extensions, reversions or restorations
associated with such copyrights, regardless of the medium of fixation or means of expression; (iv) trade secrets, know-how and other confidential or proprietary information (including processes, techniques and research and development
information); and (v) mask works, industrial designs (whether or not registered), database rights, publicity rights and privacy rights. 

“Licensable” means, with respect to any Intellectual Property Right, that a Person has the power and authority to
grant a license (or sublicense, as the case may be), on the applicable terms and conditions of this Agreement, to such Intellectual Property Right without any of the following: (i) the consent of any third party (unless such consent can be
obtained without providing any additional consideration to such third party); (ii) impairing such Person’s existing rights in respect of such Intellectual Property Right (it being understood that the grant of any license hereunder, in and
of itself, shall not be construed as an impairment of any of such Person’s rights); (iii) imposing any additional obligations on such Person under any preexisting agreement relating to such Intellectual Property Right; and/or (iv) the
payment of royalties or other consideration on or after the Effective Date by such Person to any third party under any preexisting agreement relating to such Intellectual Property Right (other than to the University of Minnesota pursuant to the
UMinn License). For the avoidance of doubt, in no event shall any Intellectual Property Right be “Licensable” if any of the foregoing conditions in clauses (i)-(iv) apply. 

“Licensed Cellectis IP” means the (i) Licensed Cellectis Patents; and (ii) Other Licensed Intellectual
Property Rights. 

  
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 “Licensed Cellectis Patents” means any and all Patents that are:
(i) related to the Calyxt Field; (ii) necessary for Calyxt to operate in the Calyxt Field; and (iii) Licensable by Cellectis and existing as of the Effective Date. 

“Licensed TALEN Mark” means the trademark “TALEN” and all registrations and applications for registration
thereof, in each case as owned by Cellectis as of the Effective Date, including the registration set forth on Schedule A. 

“Net Sales” means, with respect to any Calyxt Licensed Product, the gross amount invoiced by Calyxt or any of its
sublicensees for any such Calyxt Licensed Product, in each case less (i) all trade, quantity, and cash discounts actually allowed; (ii) all credits and allowances actually granted due to rejections, returns, billing errors, and retroactive
price reductions; (iii) applicable duties; (iv) all credits or allowances given or made for uncollectible amounts and for which a provision is made in Calyxt’s financial statements; (v) the commodity price for seed and grain; and
(vi) applicable excise, sale and use taxes. Net Sales shall also include the fair market value of all other consideration received as consideration for the sale or disposition of any Calyxt Licensed Product, whether such consideration is in
cash, payment in kind, exchange or another form. Net Sales shall be determined by using generally accepted accounting principles consistently applied. 

“Non-Exclusive Field” means the field of researching, developing and commercializing a modified or mutated I-CreI homing
endonuclease that is a homodimer, heterodimer or single chain endonuclease, but solely to the extent the foregoing falls within the Calyxt Field. 

“Other Licensed Intellectual Property Rights” means any and all know-how and other Intellectual Property Rights (excluding
any Patents and Trademarks) that are (i) related to the Calyxt Field; (ii) necessary for Calyxt to operate in the Calyxt Field; and (iii) Licensable by Cellectis and existing as of the Effective Date.  

“Patent-Related Expenses” means costs and expenses (including out-of-pocket attorneys’ fees, patent agent fees and
governmental filing fees) that Cellectis or any of its Affiliates incurs in prosecuting and maintaining the Licensed Cellectis Patents which are exclusively and solely related to the Calyxt Field.  

“Person” means an individual, corporation, partnership, limited liability company, association, trust or other entity
or organization, including a Governmental Authority. 
 “Royalty Term” means, with respect to any Calyxt
Licensed Product in any jurisdiction, the period commencing on the Effective Date and ending upon the expiration of the last-to-expire Valid Claim Covering such Calyxt Licensed Product in such jurisdiction. 

“Sublicense Revenue Term” means the period commencing on the Effective Date and ending upon the expiration of the
last-to-expire Valid Claim. 

  
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 “Subsidiary” means, with respect to any Person, any entity of which
securities or other ownership interests having ordinary voting power to elect a majority of the board of directors or other Persons performing similar functions are at the time directly or indirectly owned by such Person. 

“UMinn License” means the Exclusive Patent License Agreement between the University of Minnesota and Cellectis dated
as of January 10, 2011 (as amended, including by the First Amendment to the Exclusive Patent License Agreement, dated as of May 24, 2012, Second Amendment to the Exclusive Patent License Agreement, dated as of April 1, 2014, and Third
Amendment to the Exclusive Patent License Agreement, dated as of December 16, 2015). 
 “University of
Minnesota” means the Regents of the University of Minnesota. 
 “Valid Claim” means any
(i) claim in any unexpired and issued Patent included in the Licensed Cellectis Patents that has not been (A) disclaimed, revoked or held invalid or unenforceable by a decision of a court or other Governmental Authority of competent
jurisdiction from which no appeal (other than an appeal to the highest appellate court of such jurisdiction) can be taken, or with respect to which an appeal is not taken within the time allowed for appeal, or (B) irretrievably abandoned,
disclaimed or admitted to be invalid or unenforceable by Cellectis through reissue, disclaimer or otherwise, or (ii) pending claim in a pending Patent application included in the Licensed Cellectis Patents that has not been abandoned or finally
rejected without the possibility of appeal or refiling. 
 (b) Each of the following terms is defined in the Section set forth
opposite such term: 
  

			
	 Term
	  	Section
	 Agreement
	  	Preamble
	 Calyxt
	  	Preamble
	 Calyxt License
	  	2.01
	 Calyxt TM License
	  	2.02
	 Cellectis
	  	Preamble
	 Controlling Party
	  	9.02
	 Effective Date
	  	Preamble
	 Indemnified Party
	  	8.03
	 Indemnifying Party
	  	8.03
	 Infringement
	  	9.02
	 Losses
	  	8.01
	 Necessary Third Party License
	  	5.02
	 Negotiation Period
	  	2.06
	 Non-Controlling Party
	  	9.02
	 Option Period
	  	2.06
	 Parties
	  	Preamble
	 Party
	  	Preamble
	 Remainder
	  	9.02
	 Third Party Claim
	  	8.03
	 UMinn IP
	  	2.04


  
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 Section 1.02. Other Definitional and Interpretative Provisions. The words
“hereof”, “herein” and “hereunder” and words of like import used in this Agreement shall refer to this Agreement as a whole and not to any particular provision of this Agreement. The captions herein are included for
convenience of reference only and shall be ignored in the construction or interpretation hereof. References to Articles, Sections and Schedules are to Articles, Sections and Schedules of this Agreement unless otherwise specified. All Schedules
annexed hereto or referred to herein are hereby incorporated in and made a part of this Agreement as if set forth in full herein. Any capitalized terms used in any Schedule but not otherwise defined therein, shall have the meaning as defined in this
Agreement. Any singular term in this Agreement shall be deemed to include the plural, and any plural term the singular. Whenever the words “include”, “includes” or “including” are used in this Agreement, they shall be
deemed to be followed by the words “without limitation”, whether or not they are in fact followed by those words or words of like import. “Writing”, “written” and comparable terms refer to printing, typing and other
means of reproducing words (including electronic media) in a visible form. References to any agreement or contract are to that agreement or contract as amended, modified or supplemented from time to time in accordance with the terms hereof and
thereof. References to any Person include the successors and permitted assigns of that Person. References from or through any date mean, unless otherwise specified, from and including or through and including, respectively. References to
“law”, “laws” or to a particular statute or law shall be deemed also to include any and all Applicable Law. 

ARTICLE 2 
 GRANT
OF LICENSE 
 Section 2.01. Calyxt License. Subject to the terms and conditions of this Agreement,
Cellectis hereby grants to Calyxt an exclusive (except as otherwise provided herein and subject to existing licenses granted by Cellectis to third parties prior to the Effective Date) worldwide, perpetual license, with the right to sublicense (in
accordance with Section 2.03) under the Licensed Cellectis IP to use, have used, make, have made, sell, have sold, offer for sale, export, import and otherwise exploit any and all Calyxt Licensed Products within the Calyxt Field (the “Calyxt
License”). Notwithstanding the foregoing, the Calyxt License shall be non-exclusive solely in the Non-Exclusive Field, such that the rights as set forth above in this Section 2.01 are granted to Calyxt on a non-exclusive basis in the
Non-Exclusive Field under the Calyxt License. 
 Section 2.02. Trademark License Grant. Subject to the terms and conditions of
this Agreement, Cellectis hereby grants to Calyxt a worldwide, non-exclusive, sublicensable (in accordance with Section 2.03), royalty-free and fully paid-up, non-transferable (except as set forth in Section 11.01) license under the Licensed TALEN
Mark to use, make, have made, sell, offer for sale, import and otherwise exploit any and all Calyxt Licensed Products within the Calyxt Field (the “Calyxt TM License”). 

  
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 Section 2.03. Sublicense Rights. The Calyxt License and the Calyxt TM License
include the right of Calyxt to grant sublicenses to any Person; provided that (a) any such sublicense shall be in writing and automatically terminate upon any termination of this Agreement (it being understood that any such sublicense
shall include express terms and conditions to effect such automatic termination); (b) sublicensees of the Calyxt License shall not be permitted to grant further sublicenses thereunder with respect to any UMinn IP (as defined below);
(c) Calyxt shall cause each of its sublicensees to abide by all applicable terms and conditions of this Agreement, enforce such terms and conditions and the provisions of any sublicense against each such sublicensee; and (d) Calyxt shall
remain responsible and liable to Cellectis for the performance of each such sublicensee’s obligations and for all acts or omissions of such sublicensee as if they were acts of Calyxt under this Agreement. 

Section 2.04. UMinn License. The Parties acknowledge and agree that Calyxt has received a copy of the UMinn License
and certain Licensed Cellectis IP is owned by the University of Minnesota (“UMinn IP”) and the Calyxt License with respect to the UMinn IP is granted as a sublicense under, and subject to the terms and conditions
of, the UMinn License. Accordingly, in exercising its rights under the Calyxt License, Calyxt shall comply with any and all terms and conditions of the UMinn License as they would apply to Calyxt as a sublicensee with respect to any UMinn IP.
Without limiting the generality of the foregoing, promptly following receipt of written notice thereof, Calyxt shall reimburse Cellectis for any and all payments made by Cellectis to the University of Minnesota pursuant to Sections 11.6.4 (Milestone
Payments), 11.11 (Annual Fee), and 11.12 (Commercialization Fee) of the UMinn License, but solely to the extent that any such payments are required as a result of the applicable activities of Calyxt hereunder or thereunder. Calyxt shall not be
liable to the University of Minnesota for any other payments other than those as specifically set forth in the previous sentence. Without the prior written consent of Calyxt, Cellectis shall not (A) terminate the UMinn License, or
(B) amend or waive any rights under the UMinn License in any manner that would reasonably be expected to have a material adverse effect on any of Calyxt’s rights under this Agreement. In addition, Calyxt shall provide, and shall cause its
sublicensees to provide, to Cellectis all reports, information and other assistance in connection with Calyxt’s and its sublicensees’ activities pursuant to this Agreement that are reasonably required to enable Calyxt to comply with its
obligations under the UMinn License.  
 Section 2.05. Calyxt Improvements. (a) As between the Parties, any
and all Calyxt Improvements shall be solely and exclusively owned by Calyxt. 
 (b) Subject to the terms and conditions of this
Agreement, Calyxt, on behalf of itself and its Affiliates, hereby grants to Cellectis and its Affiliates an exclusive, perpetual, worldwide, sublicensable, non-transferable (except as set forth in Section 11.01), royalty-free and fully paid-up
license to use and otherwise exploit Calyxt Improvements for any purpose outside of the Calyxt Field. On a continuing basis during the term of this Agreement, Calyxt shall promptly make available to Cellectis all Calyxt Improvements then in
existence that are necessary or reasonably useful for the commercialization of such Calyxt Improvements by Cellectis outside of the Calyxt Field, and provide Cellectis with all reasonable assistance to enable Cellectis to understand and use such
Calyxt Improvement. Calyxt shall use commercially reasonable efforts to identify and disclose all Calyxt Improvements based on facts known to Calyxt, as well as in response to specific requests made by Cellectis. 

  
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 Section 2.06. Calyxt Option on Cellectis Improvements. Solely during the
period in which Cellectis and its Affiliates own, in the aggregate, a number of Calyxt common shares equal to at least fifty percent (50%) of the then outstanding common shares of Calyxt, Cellectis shall promptly disclose to Calyxt all
Cellectis Improvements then in existence that are necessary or reasonably useful for the commercialization of such Cellectis Improvements by Calyxt in the Calyxt Field, and provide Calyxt an option to obtain a license within the Calyxt Field. Calyxt
may exercise its option by sending a written notice to Cellectis within thirty (30) days after it has the knowledge of such Cellectis Improvements (the “Option Period”). If Calyxt has exercised its option within
the Option Period to obtain such a license, then for a period of thirty (30) days after Cellectis receives such notice from Calyxt (the “Negotiation Period”), the Parties shall, in good faith, negotiate the terms
and conditions of a definitive agreement pursuant to which Cellectis would grant Calyxt a royalty-bearing license with respect to such Cellectis Improvement; provided that if Calyxt does not exercise its option within the Option Period, or if
the Parties do not agree on the terms and conditions of such a definitive agreement for such license within the Negotiation Period, Cellectis shall be free to grant any third party any license or other rights with respect to such Cellectis
Improvement. 
 Section 2.07. No Other Licenses. Except as expressly provided in this Agreement, no other licenses
are granted to either Party under this Agreement. Each Party acknowledges and agrees that (a) any use by Calyxt or any of its sublicensees of the Licensed Cellectis IP outside the scope of the Calyxt License or the Licensed TALEN Mark outside
the scope of the Calyxt TM License is expressly prohibited and (b) any use by Cellectis or any of its sublicensees of the Calyxt Improvements outside the scope of the licenses granted to Cellectis or any of its sublicensees pursuant to Section
2.05 is expressly prohibited. 
 ARTICLE 3 

LICENSED TALEN MARK; QUALITY CONTROL 

Section 3.01. Quality Control. Cellectis reserves the right to practice reasonable quality control with regard to the use
of the Licensed TALEN Mark by Calyxt or any of its sublicensees and Calyxt shall, and shall cause its sublicensees to, adhere to such quality, appearance, reputational, distinctiveness and other standards with respect to the use of the Licensed
TALEN Mark and with respect to the goods and services sold or rendered under the Licensed TALEN Mark as Cellectis may require from time to time. Calyxt hereby acknowledges the validity of the Licensed TALEN Mark and Cellectis’ exclusive right,
title, and interest in and to the Licensed TALEN Mark, subject to the license granted hereunder. Calyxt shall, and shall cause its sublicensees to, (a) not take any action or make any statement which would reasonably be expected to damage the
reputation or goodwill associated with Cellectis, any of its Affiliates, or the Licensed TALEN Mark, or to prejudice, infringe or impair the rights of Cellectis with respect to the Licensed TALEN Mark and (b) comply with all Applicable Law
governing the use of the Licensed TALEN Mark, including all services performed under the Licensed TALEN Mark and all goods to which the Licensed TALEN Mark is applied. At Calyxt’s sole expense, Calyxt shall supply, and shall cause its
sublicensees to supply, Cellectis with specimens of all uses of the Licensed TALEN Mark upon the reasonable request of Cellectis. 

  
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 Section 3.02. Reservation of Rights; Ownership. (a) Calyxt, on behalf of itself
and its sublicensees, acknowledges and agrees that (i) Cellectis is the sole and exclusive owner of all right, title and interest in and to the Licensed TALEN Mark; (ii) Cellectis shall have the sole and exclusive right to prosecute and
maintain the Licensed TALEN Mark; and (iii) neither Calyxt nor any of its sublicensees has acquired, and shall not acquire, any right, title or interest in or to the Licensed TALEN Mark other than the rights expressly set forth in this
Agreement. Calyxt shall cooperate with Cellectis in taking all appropriate measures for the protection of the Licensed TALEN Mark. 
 (b)
Calyxt shall not, and shall cause its sublicensees not to, (i) challenge the validity, enforceability or ownership of the Licensed TALEN Mark or claim adversely or assist in any claim adverse to Cellectis concerning any right, title or interest
in the Licensed TALEN Mark; (ii) do or permit any act which may directly or indirectly impair or prejudice Cellectis’ title to the Licensed TALEN Mark or be detrimental to the reputation and goodwill of Cellectis or any of its Affiliates,
including any act which might assist or give rise to any application to remove or de-register any of the Licensed TALEN Mark; or (iii) register or attempt to register any trademarks for any words, names, graphics, or other source identifiers
that are identical or confusingly similar to the Licensed TALEN Mark. All use of the Licensed TALEN Mark by Calyxt and any of its sublicensees, and all goodwill associated with such use, shall inure to the benefit of Cellectis. 

Section 3.03. Trademark Notice. In connection with the use of the Licensed TALEN Mark, Calyxt and its sublicensees shall
mark each use with the registered trademark symbol, “®,” or such other trademark notice symbol as designated by Cellectis. 

ARTICLE 4 
 COMMERCIALIZATION 

Section 4.01 Commercialization and Performance Milestones. Calyxt shall use its commercially reasonable efforts, and shall
require all sublicensees to use commercially reasonable efforts, in each case consistent with sound and reasonable business practices and judgment, to commercialize the Licensed Cellectis Patents in the Calyxt Field and to manufacture, offer to sell
and sell Calyxt Licensed Products as soon as practicable and to maximize sales thereof.  
 Section 4.02 Covenants
Regarding the Manufacture of Licensed Products. Calyxt hereby covenants and agrees, and shall require all sublicensees to covenant and agree, that (a) the manufacture, use, sale, or transfer of Calyxt Licensed Products shall comply with all
Applicable Laws, including all federal export laws and regulations; and (b) it will make commercially reasonable efforts such that the Calyxt Licensed Products shall not be defective in design or manufacture. Calyxt hereby further covenants and
agrees that, pursuant to 35 United States Code Section 204, it shall, and it shall cause each sublicensee, to substantially manufacture in the United States of America all products embodying or produced through the use of an invention
that is subject to the rights of the federal government of the United States of America. 

  
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 Section 4.03 Export and Regulatory Compliance. Calyxt understands that the
Arms Export Control Act (AECA), including its implementing International Traffic In Arms Regulations (ITAR) and the Export Administration Act (EAA), including its Export Administration Regulations (EAR), are some (but not all) of the laws and
regulations that comprise the U.S. export laws and regulations. Calyxt further understands that the U.S. export laws and regulations include (but are not limited to): (i) ITAR and EAR product/service/data-specific requirements; (ii) ITAR
and EAR ultimate destination-specific requirements; (iii) LIAR and EAR end user-specific requirements; (iv) Foreign Corrupt Practices Act; and (v) antiboycott laws and regulations. Calyxt shall comply with all then-current applicable
export laws and regulations of the U.S. Government (and other applicable U.S. laws and regulations) pertaining to the Calyxt Licensed Products (including any associated products, items, articles, computer software, media, services, technical data,
and other information). Calyxt certifies that it shall not, directly or indirectly, export (including any deemed export), nor re-export (including any deemed re-export) the Calyxt Licensed Products (including any associated products, items,
articles, computer software, media, services, technical data, and other information) in violation of U.S. export laws and regulations or other applicable U.S. laws and regulations. Calyxt shall include an appropriate provision in its agreements with
its authorized sublicensees to assure that these parties comply with all then-current applicable U.S. export laws and regulations and other applicable U.S. laws and regulations. 

Section 4.04 Commercialization Reports. As requested by Cellectis in writing, no more than once per year, Calyxt shall
deliver to Cellectis written reports of Calyxt’s and its sublicensees’ efforts and plans to commercialize the Licensed Cellectis Patents in the Calyxt Field and to manufacture, offer to sell, or sell Calyxt Licensed Products. 

Section 4.05 Use of Cellectis and University of Minnesota Names and Trademarks. Except for the Licensed TALEN Mark, no
provision of this Agreement grants Calyxt or any of its sublicensees any right or license to use the name, logo, or any marks owned by or associated with Cellectis, the University of Minnesota or the names, or identities of any member of the
faculty, staff, or student body of the University of Minnesota. Calyxt shall not use and shall not permit any of its sublicensees to use any such logos, marks, names, or identities without the prior written approval of Cellectis or the University of
Minnesota, as applicable. 
 Section 4.06 Governmental Markings. 

(a) Calyxt and its sublicensees may mark all Calyxt Licensed Products in a manner consistent with their current patent marking practices for
their own products and Applicable Laws. Where marking is to be performed but the Calyxt Licensed Product cannot be marked, the patent notice shall be placed on associated tags, labels, packaging, or accompanying documentation (either electronic or
paper) as appropriate. 

  
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 (b) Calyxt and its sublicensees are solely responsible for obtaining all necessary approvals from
Governmental Authorities for the development, production, distribution, sale, and use of any Calyxt Licensed Product, at Calyxt’s expense, including, without limitation, any safety studies. Calyxt is solely responsible for including with the
Calyxt Licensed Product any warning labels, packaging and instructions as to the use and the quality control for such Calyxt Licensed Product. 

(c) Calyxt agrees to register this Agreement with any foreign Governmental Authority that requires such registration, and Calyxt shall pay all
costs and legal fees in connection with such registration. Calyxt shall comply with all foreign laws affecting this Agreement or the sale of Calyxt Licensed Products. 

ARTICLE 5 

ROYALTIES AND BARE SUBLICENSE REVENUE 

Section 5.01. Royalties and Bare Sublicense Revenue. As consideration for the Calyxt License, Calyxt shall pay to Cellectis the
following amounts during the following periods: 
 (a) With respect to each Calyxt Licensed Product in any jurisdiction, three percent
(3%) of all Net Sales of such Calyxt Licensed Product in such jurisdiction during the Royalty Term for such Calyxt Licensed Product; and 

(b) thirty percent (30%) of all Bare Sublicense Revenue during the Sublicense Revenue Term. 

Section 5.02. Royalty Stacking for Third Party Rights. (a) If Calyxt is presently required, or in the future is
required, to secure a royalty-bearing or fee-bearing license under any patent to use, make, have made, sell, offer for sale or import any Calyxt Licensed Product in the Calyxt Field in any jurisdiction (a “Necessary Third Party
License”), then, during the period in which Calyxt is required to make royalty payments to the licensor under such Necessary Third Party License, Calyxt shall have the right to reduce the royalty rate contemplated in Section 5.01(a)
with respect to the Net Sales of such Calyxt Licensed Product in such jurisdiction by an amount equal to one quarter (1/4) of the royalty rate payable to such licensor pursuant to such Necessary Third Party License; provided that, in no
event shall the royalty rate contemplated in Section 5.01(a) with respect to any Net Sales of any Calyxt Licensed Product be reduced to less than two percent (2%). If such Necessary Third Party License includes a royalty stacking provision of like
intent to this Section 5.02, the royalty rate reduction provided for in this Section 5.02 will be calculated as if such provision in such Necessary Third Party License were absent. For the avoidance of doubt, and notwithstanding anything in this
Agreement to the contrary, in no event shall any agreement to which Calyxt is a party as of or prior to the Effective Date be deemed a Necessary Third Party License under this Agreement.  

  
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 (b) Without limiting Section 5.02(a), and by way of example only: 

(i) If, after the Effective Date, Calyxt is required to enter into a Necessary Third Party License, pursuant to which Calyxt is
required to pay the licensor thereunder a royalty of two percent (2%) of the net sales of a Calyxt Licensed Product in a jurisdiction, then, during the period in which such royalty is required to be paid, the royalty rate contemplated in
Section 5.01(a) with respect to the Net Sales of such Calyxt Licensed Product in such jurisdiction would equal two and one half percent (2.5%). 

(ii) If, after the Effective Date, Calyxt is required to enter into a Necessary Third Party License, pursuant to which Calyxt
is required to pay the licensor thereunder a royalty of eight percent (8%) of the net sales of a Calyxt Licensed Product in a jurisdiction, then, during the period in which such royalty is required to be paid, the royalty rate contemplated in
Section 5.01(a) with respect to the Net Sales of such Calyxt Licensed Product in such jurisdiction would equal two percent (2%). 

Section 5.03. Reimbursement of Patent-Related Expenses. Commencing on the Effective Date, Calyxt shall pay all invoices
issued by Cellectis or any of its Affiliates for any Patent-Related Expenses under this Agreement within thirty (30) days of its receipt of each such invoice.  

Section 5.04. Reporting; Audit Rights. Calyxt shall render to Cellectis, on a calendar quarterly basis, commencing with the first
calendar quarter after the Effective Date, a detailed written report of the royalties and Bare Sublicense Revenue due to Cellectis. Such report shall be accompanied by a remittance of such royalties and Bare Sublicense Revenue as shown to be due
hereunder. Each report shall be rendered within thirty (30) days following the end of each calendar quarterly period. Calyxt shall keep books and records in sufficient detail to enable the royalty payments and Bare Sublicense Revenue due
hereunder to be adequately determined. Once per calendar year, upon reasonable written notice, Cellectis or any third party owner of Patent rights included in the Licensed Cellectis IP shall have the right at its sole cost and expense to cause a
nationally recognized independent certified public accountant reasonably acceptable to Calyxt to examine and inspect such books and records during Calyxt’s normal business hours, but only to the extent necessary to verify the computation of
royalties and Bare Sublicense Revenue payable hereunder. Such books and records shall be deemed Confidential Information of Calyxt hereunder, and such nationally recognized independent certified public accountant shall disclose to Cellectis or such
third party only the royalties and Bare Sublicense Revenue payable and the percentage under/overpayment by Calyxt. In the event that such examination determines that Calyxt has underpaid royalties and Bare Sublicense Revenue by more than three
percent (3%), Calyxt shall reimburse Cellectis for its reasonable costs in conducting such examination. At Calyxt’s expense, Calyxt shall also provide Cellectis with all reasonably requested cooperation in connection with complying with any
audit regarding the activities of Calyxt hereunder that is conducted by or on behalf of the University of Minnesota pursuant to the UMinn License. 

Section 5.05. Method of Payment. Each payment by Calyxt hereunder shall be made by electronic transfer in immediately
available funds, at Cellectis’ election, via either a bank wire transfer or any other means of electronic funds transfer to a bank account specified in writing by Cellectis to Calyxt. Cellectis may change such account by written notice at least
five (5) Business Days before any payment is due. All royalties and Bare Sublicense Revenue of Calyxt  

  
 12 








 
shall be computed and paid in U.S. dollars. For the purposes of determining the amount of any royalties or Bare Sublicense Revenue due for any relevant calendar quarter, the amount of Net Sales
or Bare Sublicense Revenue in any foreign currency shall be converted into U.S. dollars in a manner consistent with Calyxt’s customary practices used to prepare its audited financial reports. No more than once per calendar year, upon written
request of Cellectis, Calyxt shall provide Cellectis with a written explanation of such customary practices of Calyxt. 

Section 5.06. Withholding Taxes. To the extent either Party is required by Applicable Law to withhold or deduct any amounts
from any payments to be made under this Agreement, such Party shall be entitled to withhold or deduct such amounts and such amounts shall be treated for all purposes of this Agreement as having been paid to the Party in respect of which such
deduction and withholding were made. Each Party shall (in consultation and cooperation with the other) use commercially reasonable efforts to attempt to lawfully mitigate, reduce or avoid such withholdings or deductions. Promptly after the execution
of this Agreement, Cellectis shall provide to Calyxt a valid Form W-8BEN-E establishing Cellectis’ right to a zero percent rate of withholding tax with respect to the amounts payable by Calyxt under this Article 5 under Article 12 of the United
States – France income tax treaty. 
 ARTICLE 6 

REPRESENTATIONS AND WARRANTIES; 

DISCLAIMERS; LIMITATION OF LIABILITY 

Section 6.01. Mutual Representations and Warranties. As of the Effective Date, each Party hereby represents and warrants to
the other Party that (a) the execution, delivery and performance by such Party of this Agreement are within such Party’s corporate powers and have been duly authorized by all necessary corporate action on the part of such Party and
(b) this Agreement constitutes a valid and binding agreement of such Party enforceable against such Party in accordance with its terms (subject to applicable bankruptcy, insolvency, fraudulent transfer, reorganization, moratorium and other laws
affecting creditors’ rights generally and general principles of equity). 
 Section 6.02. Disclaimers and Limitation of
Liability. EXCEPT FOR THE REPRESENTATIONS AND WARRANTIES IN SECTION 6.01, ALL LICENSES AND RIGHTS GRANTED HEREIN ARE MADE ON AN “AS IS” BASIS, AND THE PARTIES EACH HEREBY DISCLAIM ANY EXPRESS OR IMPLIED REPRESENTATIONS OR WARRANTIES OF
ANY KIND, INCLUDING WITHOUT LIMITATION, THOSE REGARDING MERCHANTABILITY, FITNESS FOR A PARTICULAR PURPOSE, OR OF NON-INFRINGEMENT. NOTWITHSTANDING ANYTHING IN THIS AGREEMENT TO THE CONTRARY, CALYXT ACKNOWLEDGES AND AGREES THAT ALL RIGHTS GRANTED TO
CALYXT UNDER THIS AGREEMENT ARE SUBJECT IN ALL RESPECTS TO ANY AND ALL LICENSES OR OTHER RIGHTS GRANTED BY CELLECTIS OR ANY OF ITS AFFILIATES TO ANY THIRD PARTIES WITH RESPECT TO ANY LICENSED CELLECTIS IP AS OF OR PRIOR TO THE EFFECTIVE DATE. TO THE
EXTENT PERMITTED BY APPLICABLE LAW, NEITHER PARTY SHALL BE LIABLE UNDER 

  
 13 








 
ANY LEGAL OR EQUITABLE THEORY FOR ANY INDIRECT, SPECIAL, INCIDENTAL, OR CONSEQUENTIAL DAMAGES OF ANY KIND ARISING OUT OF OR OTHERWISE RELATED TO THIS AGREEMENT, EVEN IF SUCH PARTY HAS BEEN
ADVISED OF THE POSSIBILITY OF SUCH DAMAGES. 
 Section 6.03. University of Minnesota Disclaimer; Limitation of Liability; and
Release. 
 (a) Calyxt acknowledges and agrees that the disclaimer, remedy limitation and damages cap provisions applicable to and
exclusion of representations and warranties by the University Minnesota, set forth in Sections 10.2, 11.1 and 11.2 of the UMinn License shall apply with respect to Calyxt’s rights and remedies under this Agreement and such Sections are hereby
incorporated by reference herein, mutatis mutandis. 
 (b) Calyxt, on behalf of itself and its Affiliates and its and their respective
employees, hereby releases the University of Minnesota, Cellectis and their respective regents, employees and agents forever from any and all suits, actions, claims, liabilities, demands, damages, losses, and expenses (including reasonable
attorneys’ and investigative expenses) relating to or arising out of the manufacture, use, lease, sale, or other disposition of any Calyxt Licensed Product. 

ARTICLE 7 

CONFIDENTIALITY 

Section 7.01. Confidentiality. (a) The receiving party shall keep confidential the disclosing party’s
Confidential Information, and shall not use any of the disclosing party’s Confidential Information for any purpose other than the exercise of the receiving party’s rights, or as otherwise permitted, under this Agreement. The receiving
party shall preserve the confidentiality of the disclosing party’s Confidential Information as it would customarily take to preserve the confidentiality of its own similar type of confidential information and shall not disclose the disclosing
party’s Confidential Information to any third party without the prior written consent of the disclosing party, except as expressly permitted hereunder. The receiving party may disclose the Confidential Information to (i) any of its
employees, agents, independent contractors and sublicensees who need it in connection with this Agreement and are bound in writing by restrictions regarding disclosure and use of the Confidential Information comparable to and no less restrictive
than those set forth herein or (ii) the extent it is in response to a valid order of a court or other Governmental Authority or to otherwise comply with Applicable Law; provided that, in the case of clause (ii), the receiving party shall
first provide written notice to the disclosing party and reasonably cooperate with the disclosing party to obtain a protective order or other measures preserving the confidential treatment of such Confidential Information and requiring that the
information or documents so disclosed be used only for the purposes for which the order was issued or is otherwise required by Applicable Law. 

(b) The terms and conditions of this Agreement shall be deemed Confidential Information for the purposes of this Agreement;
provided that each Party may disclose the terms and conditions of this Agreement: (i) in confidence, to its accountants, banks and present and prospective financing sources and their advisors; (ii) in connection with the enforcement
of this Agreement or rights under  

  
 14 








 
this Agreement; (iii) in confidence, in connection with an actual or proposed merger, acquisition or similar transaction involving such Party; (iv) in confidence, to its Affiliates;
(v) in confidence, to its third party independent contractors who have a need to know, solely in connection with their provision of services to such Party; (vi) as required by applicable securities laws or the rules of any stock exchange
on which securities of such Party are traded or any other Applicable Law; or (vii) as mutually agreed upon by the Parties in writing. 

ARTICLE 8 

INDEMNIFICATION 

Section 8.01. Indemnification by Calyxt. Calyxt shall defend Cellectis and its Affiliates and their respective officers,
directors, employees, contractors, customers and agents against any action, suit, proceeding or other claim, and indemnify and hold each of them harmless from any and all damages, liabilities, expenses, and other losses (including reasonable
attorneys’ fees and court costs) (“Losses”) to the extent arising from any (a) breach of this Agreement by Calyxt; (b) breach of the UMinn License caused by any activities of Calyxt or its sublicensees;
(c) use, making, having made, sale, offer for sale, importation or any other exploitation of any Calyxt Licensed Products or any exploitation of the Licensed Cellectis IP by Calyxt or any of its sublicensees; (d) gross negligence or
willful misconduct by Calyxt; and/or (e) violation of Applicable Law by Calyxt. 
 Section 8.02. Indemnification
by Cellectis. Cellectis shall defend Calyxt and its Affiliates and their respective officers, directors, employees, and agents against any action, suit, proceeding or other claim, and indemnify and hold each of them harmless from any and all
Losses to the extent arising from any (a) breach of this Agreement by Cellectis; (b) gross negligence or willful misconduct by Cellectis; and/or (c) violation of Applicable Law by Cellectis. 

Section 8.03. Third Party Claim Procedures. (a) The Party seeking indemnification under Section 8.01 or 8.02 (the
“Indemnified Party”) shall give prompt notice in writing to the Party against whom indemnity is to be sought (the “Indemnifying Party”) of the assertion of any claim or the commencement
of any action, suit, proceeding or other claim by any third party (“Third Party Claim”) in respect of which indemnity may be sought under such Section. Such notice shall set forth in reasonable detail such Third Party
Claim and the basis for indemnification (taking into account the information then available to the Indemnified Party). The failure to so notify the Indemnifying Party shall not relieve the Indemnifying Party of its obligations hereunder, except to
the extent such failure shall have actually prejudiced the Indemnifying Party.  
 (b) The Indemnifying Party shall be entitled
to participate in the defense of any Third Party Claim and, subject to the limitations set forth in this Section, shall be entitled to control and appoint lead counsel for such defense, in each case at its own expense; provided that prior to
assuming control of such defense, the Indemnifying Party must acknowledge that it would have an indemnity obligation for the Losses resulting from such Third Party Claim as provided under this Article 8. 

  
 15 








 (c) The Indemnifying Party shall not be entitled to assume or maintain control of the defense of
any Third Party Claim and shall pay the fees and expenses of counsel retained by the Indemnified Party if (i) the Indemnifying Party does not deliver the acknowledgment referred to in Section 8.03(b) within thirty (30) days of receipt of
notice of the Third Party Claim pursuant to Section 8.03(a); (ii) the Third Party Claim relates to or arises in connection with any criminal action, proceeding or claim; (iii) the Third Party Claim seeks an injunction or equitable relief
against the Indemnified Party or any of its Affiliates; or (iv) the Indemnifying Party has failed or is failing to prosecute or defend vigorously the Third Party Claim. 

(d) If the Indemnifying Party shall assume the control of the defense of any Third Party Claim in accordance with the provisions of this
Section 8.03, the Indemnifying Party shall obtain the prior written consent of the Indemnified Party before entering into any settlement of such Third Party Claim. 

(e) In circumstances where the Indemnifying Party is controlling the defense of a Third Party Claim in accordance with paragraphs
(b) and (c) above, the Indemnified Party shall be entitled to participate in the defense of any Third Party Claim and to employ separate counsel of its choice for such purpose, in which case the fees and expenses of such separate counsel
shall be borne by the Indemnified Party; provided that in such event the Indemnifying Party shall pay the fees and expenses of such separate counsel (i) incurred by the Indemnified Party prior to the date the Indemnifying Party assumes
control of the defense of the Third Party Claim or (ii) if representation of both the Indemnifying Party and the Indemnified Party by the same counsel would create a conflict of interest. 

(f) Each Party shall cooperate, and cause their respective Affiliates to cooperate, in the defense or prosecution of any Third Party Claim and
shall furnish or cause to be furnished such records, information and testimony, and attend such conferences, discovery proceedings, hearings, trials or appeals, as may be reasonably requested in connection therewith. 

Section 8.04. Direct Claim Procedures. In the event that an Indemnified Party has a claim for indemnity under Section 8.01
or 8.02 against an Indemnifying Party that does not involve a Third Party Claim, the Indemnified Party shall give prompt notice in writing of such claim to the Indemnifying Party. Such notice shall set forth in reasonable detail such claim and the
basis for indemnification (taking into account the information then available to the Indemnified Party). The failure to so notify the Indemnifying Party shall not relieve the Indemnifying Party of its obligations hereunder, except to the extent such
failure shall have actually prejudiced the Indemnifying Party. 
 ARTICLE 9 

PROSECUTION, MAINTENANCE; LITIGATION 

Section 9.01. Prosecution and Maintenance. (a) As between the Parties, Cellectis shall have the sole and exclusive right to
prosecute and maintain the Licensed Cellectis IP and Licensed TALEN Mark (it being understood that, and Calyxt acknowledges and agrees that, pursuant to the UMinn License, the University of Minnesota has the sole and exclusive right to


  
 16 








 
prosecute and maintain the UMinn IP). Subject to any rights granted to any third parties prior to the date hereof with respect to any Exclusively Licensed Cellectis Patents, Cellectis shall
(i) keep Calyxt reasonably informed of all steps to be taken in connection with the prosecution and maintenance of the Exclusively Licensed Cellectis Patents, and (ii) consider in good faith (or, in the case of any Exclusively Licensed
Cellectis Patents being prosecuted by the University of Minnesota, use commercially reasonable efforts to cause the University of Minnesota to consider in good faith) all reasonable comments and suggestions by Calyxt regarding such matters,
including in respect of any actions, decisions, applications, amendments, submissions or correspondence related thereto. Notwithstanding the foregoing, subject to any rights granted to any third parties prior to the date hereof with respect to any
Exclusively Licensed Cellectis Patents, in the event that Cellectis elects to abandon or otherwise cease prosecuting and maintaining any Exclusively Licensed Cellectis Patent (excluding the Patents licensed to Cellectis under the UMinn License),
prior to any such abandonment, Calyxt shall have the option to acquire at no cost any such Exclusively Licensed Cellectis Patent and assume the responsibility for the prosecution and maintenance of such Exclusively Licensed Cellectis Patent (it
being understood that, in the event that Calyxt exercises such option to acquire such Exclusively Licensed Cellectis Patent, (A) Cellectis shall execute and deliver any documents and perform any other acts, in each case as may be reasonably
necessary to effect the foregoing and (B) effective as of Calyxt acquiring ownership of such Exclusively Licensed Cellectis Patent, such Exclusively Licensed Cellectis Patent shall thereafter be automatically deemed to be licensed to Cellectis
under the licenses granted to Cellectis pursuant to Section 2.05). 
 (b) As between the Parties, Calyxt shall have the sole and exclusive
right to prosecute and maintain all Intellectual Property Rights owned or otherwise controlled by Calyxt or any of its Affiliates, including all Intellectual Property Rights in or to any Calyxt Improvements. To the extent that any Calyxt Improvement
Patents relate to any subject matter outside of the Calyxt Field, Calyxt shall (i) keep Cellectis reasonably informed of all steps to be taken in connection with the prosecution and maintenance of such Calyxt Improvement Patents, and
(ii) consider in good faith all reasonable comments and suggestions by Cellectis regarding such matters, including in respect of any actions, decisions, applications, amendments, submissions or correspondence related thereto. Notwithstanding
the foregoing, in the event that Calyxt elects to abandon or otherwise cease prosecuting or maintaining any such Calyxt Improvement Patent related to any subject matter outside of the Calyxt Field, prior to any such abandonment, Cellectis shall have
the option to acquire at no cost any such Calyxt Improvement Patent and assume the responsibility for the prosecution and maintenance of such Calyxt Improvement Patent (it being understood that, in the event that Cellectis exercises such option to
acquire such Calyxt Improvement Patent, (x) Calyxt shall execute and deliver any documents and perform any other acts, in each case as may be reasonably necessary to effect the foregoing and (y) effective as of Cellectis acquiring
ownership of such Calyxt Improvement Patent, such Calyxt Improvement Patent shall thereafter be automatically deemed to be licensed to Calyxt under the Calyxt License). 

Section 9.02. Litigation. (a) If either Party becomes aware of any actual or threatened infringement or other violation by
any third party of any Licensed Cellectis Patent or Calyxt Improvement Patent, or any challenge to any Licensed Cellectis Patent or Calyxt Improvement Patent by any third party, then such Party shall promptly notify the other Party in writing
thereof. 

  
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 (b) Cellectis shall have the first right, but not the obligation, at its expense and using
counsel of its choice, to enforce any Licensed Cellectis Patent against any Person or defend any challenge with respect to any such Licensed Cellectis Patent. Cellectis shall have sole and exclusive control of any decisions or other aspects of any
such enforcement or defense; provided that if Cellectis elects to not (i) enforce any Exclusively Licensed Cellectis Patent against any infringement or other violation of the exclusive rights granted to Calyxt under the Calyxt License or
(ii) defend any such Exclusively Licensed Cellectis Patent from any challenge that would be reasonably expected to have a material adverse effect on Calyxt’s exclusive rights under the Calyxt License, then in either case (but only to the
extent that prior to the date hereof Cellectis has not granted any third party any right to enforce or defend any such Exclusively Licensed Cellectis Patent), Cellectis shall promptly provide Calyxt with written notice of such election and,
following receipt of such notice, Calyxt may, at its sole option and expense, enforce its rights under or defend any challenge to such Exclusively Licensed Cellectis Patent, as applicable. 

(c) Calyxt shall have the first right, but not the obligation, at its expense and using counsel of its choice, to enforce any Calyxt
Improvement Patent against any Person or defend any challenge with respect to any such Calyxt Improvement Patent. Calyxt shall have sole and exclusive control of any decisions or other aspects of any such enforcement or defense; provided that
if Calyxt elects to not (i) enforce any such Calyxt Improvement Patent against any infringement or other violation thereof outside of the Calyxt Field or (ii) defend any such Calyxt Improvement Patent from any challenge that would be
reasonably expected to have a material adverse effect on Cellectis’ rights under the licenses granted to Cellectis pursuant to Section 2.05, then in either case, Calyxt shall promptly provide Cellectis with written notice of such election
and, following receipt of such notice, Cellectis may, at its sole option and expense, enforce its rights under or defend any challenge to such Calyxt Improvement Patent, as applicable. 

(d) The Party controlling any enforcement or defense under Section 9.02(b) or 9.02(c) (the “Controlling
Party”) shall keep the other Party (the “Non-Controlling Party”) reasonably and regularly informed of the status and progress of such enforcement or defense efforts, and shall reasonably consider the
Non-Controlling Party’s comments on any such efforts. The Non-Controlling Party shall provide the Controlling Party with all reasonable assistance in the enforcement or defense of any Exclusively Licensed Cellectis Patents or Calyxt Improvement
Patents, as applicable, as the Controlling Party may reasonably request, including by signing or executing any necessary documents and consenting to it being named a party to any applicable proceedings. The Non-Controlling Party shall have the right
to be represented in any enforcement or defense of any Exclusively Licensed Cellectis Patents or Calyxt Improvement Patents, as applicable, by counsel of its choice and at its own expense. Neither Party shall settle any action, suit, proceeding or
other claim involving any Exclusively Licensed Cellectis Patent or Calyxt Improvement Patent in any manner without the prior written consent of the other Party, such consent not to be unreasonably withheld. 

  
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 (e) Any recoveries resulting from an action, suit, proceeding or other claim brought by a
Party under Section 9.02(b) or 9.02(c) shall be first applied against payment of each Party’s costs and expenses in connection therewith. Any such recoveries in excess of such costs and expenses (the “Remainder”)
shall be retained by the Controlling Party; provided that if Calyxt is the Controlling Party, the Remainder with respect to any enforcement of the Exclusively Licensed Cellectis Patents shall be included in Net Sales and Bare Sublicense
Revenue, as applicable, for purposes of calculating royalties and payments owed to Cellectis hereunder. 
 (f) For the avoidance of
doubt, as between the Parties, (i) Cellectis shall have the sole and exclusive right, but not the obligation, to bring and control any legal action in connection with any actual, alleged, or threatened infringement of any Licensed Cellectis
Patents (A) outside of the Calyxt Field and (B) within the Non-Exclusive Field, in each case at Cellectis’ own expense as it reasonably determines appropriate and (ii) Calyxt shall have the sole and exclusive right, but not the
obligation, to bring and control any legal action in connection with any actual, alleged, or threatened infringement of any Calyxt Improvement Patents within the Calyxt Field at its own expense as it reasonably determines appropriate. 

ARTICLE 10 
 TERM
AND TERMINATION 
 Section 10.01. Term. This Agreement shall remain in full force and effect in
perpetuity unless earlier terminated, in whole or in part, pursuant to Section 10.02 or 10.03. 
 Section 10.02. Mutual
Agreement. This Agreement may be terminated in its entirety at any time upon the mutual written agreement of the Parties. 

Section 10.03. For Cause. Either Party may, by written notice to the other Party, immediately terminate this Agreement
(a) if such other Party is in material breach of any provision of this Agreement (it being understood that if such breach is capable of being cured, such other Party shall have the right to cure such breach within sixty (60) days of
receiving written notice thereof) and (b) upon the bankruptcy, dissolution or winding up of such other Party, or the making or seeking to make or arrange an assignment for the benefit of creditors of such other Party, or the initiation of
proceedings in voluntary or involuntary bankruptcy, or the appointment of a receiver or trustee of such other Party’s property that is not discharged within ninety (90) days. 

Section 10.04. Survival. Notwithstanding anything in this Agreement to the contrary, Sections 2.05(a), 2.07, 3.02, 5.04,
6.02, 6.03, 9.02(f) and 10.04, and Articles 7, 8 and 11 shall survive any expiration or termination of this Agreement. 

  
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 ARTICLE 11 

MISCELLANEOUS 

Section 11.01. Assignment. Neither this Agreement nor any rights or obligations hereunder may be assigned or otherwise
transferred by either Party, in whole or in part, whether voluntarily or by operation of Applicable Law, without the prior written consent of the other Party; provided that either Party may, without the consent of the other Party, assign or
otherwise transfer this Agreement to (a) any of its Affiliates or (b) any successor to all or substantially all of the assets or business of such Party to which this Agreement relates. Any attempted assignment or transfer in contravention
of this Section 11.01 shall be void ab initio. 
 Section 11.02. Notices. All notices, requests and other
communications to either Party hereunder shall be in writing (including facsimile transmission and electronic mail transmission, so long as a receipt of such electronic mail is requested and received) and shall be given, 

if to Cellectis, to: 
 Cellectis
S.A. 
 8 rue de la Croix Jarry 

75013 Paris, France 
 Attention:
Marie-Bleuenn Terrier 
 E-mail: marie-bleuenn.terrier@cellectis.com 

if to Calyxt, to: 
 Calyxt, Inc.

 600 County Road D West, Suite 8 

New Brighton, MN 55112 

Attention: Federico A. Tripodi 

E-mail: Federico.tripodi@calyxt.com 
 or such
other address or facsimile number as such Party may hereafter specify for the purpose by notice to the other Party. All such notices, requests and other communications shall be deemed received on the date of receipt by the recipient thereof if
received prior to 5:00 p.m. in the place of receipt and such day is a Business Day in the place of receipt. Otherwise, any such notice, request or communication shall be deemed not to have been received until the next succeeding Business Day in the
place of receipt. 
 Section 11.03. Amendments and Waivers. (a) Any provision of this Agreement may be amended or
waived if, but only if, such amendment or waiver is in writing and is signed, in the case of an amendment, by each Party, or in the case of a waiver, by the Party against whom the waiver is to be effective. 

(b) No failure or delay by any Party in exercising any right, power or privilege hereunder shall operate as a waiver thereof nor shall any
single or partial exercise thereof preclude any other or further exercise thereof or the exercise of any other right, power or privilege. The rights and remedies herein provided shall be cumulative and not exclusive of any rights or remedies
provided by Applicable Law. 

  
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 Section 11.04. Expenses. Except as otherwise provided herein, all costs and
expenses incurred in connection with this Agreement shall be paid by the Party incurring such cost or expense. 

Section 11.05. Governing Law. This Agreement shall be governed by and construed in accordance with the laws of France

 Section 11.06. Jurisdiction. The Parties agree that any suit, action or proceeding seeking to enforce any
provision of, or based on any matter arising out of or in connection with, this Agreement or the transactions contemplated hereby shall be brought in a court of competent jurisdiction sitting in Paris, France and each of the Parties hereby
irrevocably consents to the exclusive jurisdiction of such court (and of the appropriate appellate courts therefrom) in any such suit, action or proceeding and irrevocably waives, to the fullest extent permitted by law, any objection that it may now
or hereafter have to the laying of the venue of any such suit, action or proceeding in any such court or that any such suit, action or proceeding brought in any such court has been brought in an inconvenient forum. Process in any such suit, action
or proceeding may be served on any Party anywhere in the world, whether within or without the jurisdiction of any such court. Without limiting the foregoing, each Party agrees that service of process on such Party as provided in Section 11.02 shall
be deemed effective service of process on such Party. 
 Section 11.07. WAIVER OF JURY TRIAL. EACH OF THE PARTIES
HEREBY IRREVOCABLY WAIVES ANY AND ALL RIGHT TO TRIAL BY JURY IN ANY LEGAL PROCEEDING ARISING OUT OF OR RELATED TO THIS AGREEMENT OR THE TRANSACTIONS CONTEMPLATED HEREBY. 

Section 11.08. Counterparts; Effectiveness; Third Party Beneficiaries. This Agreement may be signed in any number of
counterparts, each of which shall be an original, with the same effect as if the signatures thereto and hereto were upon the same instrument. This Agreement shall become effective when each Party shall have received a counterpart hereof signed by
the other Parties. Until and unless each Party has received a counterpart hereof signed by the other Party, this Agreement shall have no effect and neither Party shall have any right or obligation hereunder (whether by virtue of any other oral or
written agreement or other communication). No provision of this Agreement is intended to confer any rights, benefits, remedies, obligations, or liabilities hereunder upon any Person other than the Parties and their respective successors and assigns;
provided that the University of Minnesota shall be a third party beneficiary of Section 6.03. 
 Section 11.09.
Entire Agreement. This Agreement constitutes the entire agreement between the Parties with respect to the subject matter hereof and thereof and supersedes all prior agreements and understandings, both oral and written, between the Parties
with respect to the subject matter hereof and thereof. 

  
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 Section 11.10. Relationship of the Parties. Nothing contained in this Agreement is
intended or shall be deemed to make either Party the agent, employee, partner or joint venturer of the other Party or be deemed to provide such Party with the power or authority to act on behalf of the other Party or to bind the other Party to any
contract, agreement or arrangement with any other individual or entity. 
 Section 11.11. Severability. If any term,
provision, covenant or restriction of this Agreement is held by a court of competent jurisdiction or other Governmental Authority to be invalid, void or unenforceable, the remainder of the terms, provisions, covenants and restrictions of this
Agreement shall remain in full force and effect and shall in no way be affected, impaired or invalidated so long as the economic or legal substance of the transactions contemplated hereby is not affected in any manner materially adverse to either
Party. Upon such a determination, the Parties shall negotiate in good faith to modify this Agreement so as to effect the original intent of the Parties as closely as possible in an acceptable manner in order that the transactions contemplated hereby
be consummated as originally contemplated to the fullest extent possible. 
 Section 11.12. Specific Performance.
The Parties agree that irreparable damage would occur if any provision of this Agreement were not performed in accordance with the terms hereof and that the Parties shall be entitled to an injunction or injunctions to prevent breaches of this
Agreement or to enforce specifically the performance of the terms and provisions hereof in any court set forth in Section 11.06, in addition to any other remedy to which they are entitled at law or in equity. 

[Remainder of page intentionally left blank] 

  
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 IN WITNESS WHEREOF, the Parties have caused this Agreement to be duly executed as of the date
first written above. 
  

			
	CELLECTIS S.A.
		
	By:	 	 /s/ André Choulika

		 	Name: André Choulika
		 	Title: Chief Executive Officer


  

			
	CALYXT, INC.
		
	By:	 	 /s/ Federico Tripodi

		 	Name: Federico Tripodi
		 	Title: Chief Executive Officer



  
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 Schedule A 

Licensed TALEN Mark 
  

											
	 Mark
	  	 Serial No.
	  	 Registration No.
	  	 Filing Date
	  	 Registration Date
	  	 Country

	 TALEN
	  	79/107519	  	4,729,507	  	October 27, 2011	  	May 5, 2015	  	U.S.


  
 A-1