Document:

Exhibit 10.1

 

Certain information identified by [***] have
been excluded from this exhibit because it is both (i) not material and (ii) competitively harmful if publicly disclosed.

 

EXECUTION VERSION

 

RESEARCH AND COLLABORATION AGREEMENT

 

by and among

 

Eli
Lilly and Company

 

and

 

ProQR
Therapeutics N.V.

 

and

 

ProQR
Therapeutics VIII B.V.

 

    

     

    

 

CONFIDENTIAL

 

RESEARCH AND COLLABORATION AGREEMENT

 

This Research
and Collaboration Agreement (“Agreement”) is entered into as of September 3rd, 2021
(the “Effective Date”), by and among ProQR Therapeutics N.V.,
a company organized and existing under the laws of the Netherlands, having its principal place of business at Zernikedreef 9, 2333 CK
Leiden, The Netherlands, and ProQR Therapeutics VIII B.V., a company organized and
existing under the laws of the Netherlands, having its principal place of business at Zernikedreef 9, 2333 CK Leiden, the Netherlands,
(such entities, collectively, “ProQR”), and Eli Lilly and Company,
a corporation organized and existing under the laws of Indiana, with its principal business office located at Lilly Corporate Center, Indianapolis, Indiana
46285, U.S.A. (“Lilly”). Lilly and ProQR are each hereafter referred to individually as a “Party”
and together as the “Parties.”

 

Whereas,
ProQR is a biotechnology company that has developed a proprietary platform named AXIOMER® relating to the design of antisense oligonucleotides
to edit RNA in cells, and controls certain intellectual property rights with respect to using such platform to discover, generate, optimize,
identify and select antisense oligonucleotide sequences;

 

Whereas,
Lilly is a pharmaceutical company engaged in the research, development, manufacturing, marketing and distribution of pharmaceutical products,
including therapeutic products;

 

Whereas,
ProQR and Lilly desire to collaborate to use the ProQR Platform (as defined below) to discover, generate, optimize, identify and select
antisense oligonucleotide sequences which are Directed To (as defined below) Project Targets (as defined below); and

 

Whereas,
Lilly desires to obtain from ProQR, and ProQR desires to grant to Lilly, certain exclusive and non-exclusive license rights to develop,
manufacture, and commercialize Compounds or Products containing such antisense oligonucleotide sequence(s) Directed To Project Targets,
subject to the terms and conditions of this Agreement.

 

Now,
Therefore, in consideration of the mutual covenants contained herein, and for other good and valuable consideration, the receipt
and adequacy of which are hereby acknowledged, the Parties hereby agree as follows:

 

Article 1

 

DEFINITIONS

 

Capitalized terms used in
this Agreement and the Exhibits hereto shall have the following meanings (or as defined elsewhere in this Agreement):

 

1.1           “Acquirer”
has the meaning set forth in the definition of “Change of Control.”

 

1.2           “Additional
Project” has the meaning set forth in Section 3.4.

 

    

    
	 	 
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1.3           “Additional
Target” has the meaning set forth in Section 3.4.

 

1.4           “Additional
Workplan” has the meaning set forth in Section 4.4.2.

 

1.5           “Affiliate”
means, with respect to any Person, any entity that, at the relevant time (whether as of the Effective Date or thereafter), directly or
indirectly through one or more intermediaries, controls, is controlled by, or is under common control with such Person, for so long as
such control exists. As used in this Section 1.5, “control” means: (a) to possess, directly or indirectly, the
power to direct or cause the direction of the management or policies of an entity, whether through ownership of voting securities or
by contract relating to voting rights or corporate governance; or (b) direct or indirect ownership of fifty percent (50%) (or such
lesser percentage that is the maximum allowed to be owned by a foreign entity in a particular jurisdiction) or more of the voting share
capital or other equity interest in such entity.

 

1.6           “Agreement”
has the meaning set forth in the Preamble.

 

1.7           “Alliance
Manager” has the meaning set forth in Section 2.1.

 

1.8           “Applicable
Laws” means the applicable provisions of any and all federal, national, supranational, regional, state and local laws,
treaties, statutes, rules, regulations, guidelines or requirements, administrative codes, guidance, ordinances, judgments, decrees, directives,
injunctions, orders, or permits of or from any court, arbitrator, Regulatory Authority, Governmental Authority, taxing authority, national
securities exchange or exchange listing organization having jurisdiction over or related to the relevant subject item that may be in
effect from time to time during the Term, including data protection and privacy laws.

 

1.9           “Background
IP” means Lilly Background IP or ProQR Background IP, as applicable.

 

1.10         “BLA/NDA”
means a Biologic License Application or New Drug Application (as more fully described in the FD&C Act and any applicable regulations
promulgated thereunder by the FDA), or any analogous application or submission with any Regulatory Authority outside of the United States.

 

1.11         “Business
Day” means any day, other than any Saturday, Sunday, or any day that banks are authorized or required to be closed in Indianapolis, Indiana
or Amsterdam, the Netherlands.

 

1.12         “Calendar
Quarter” means each respective period of three (3) consecutive months ending on March 31, June 30, September 30,
and December 31 of any Calendar Year.

 

1.13         “Calendar
Year” means each respective period of twelve (12) consecutive months commencing on January 1 and ending on December 31.

 

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1.14         “Candidate
Success CSFs” means with respect to a Project Target, the Critical Success Factors for such Project Target established
by the JSC necessary for a Hit Directed To such Project Target to be considered a Successful Candidate.

 

1.15         “Change
of Control” means:

 

(a)         with
respect to either Party: (i) the acquisition by a Third Party, in one transaction or a series of related transactions, of direct
or indirect beneficial ownership of more than [***] of the outstanding voting equity securities of such Party; (ii) a merger or
consolidation involving such Party, as a result of which a Third Party acquires direct or indirect beneficial ownership of more than
[***] of the voting power of the surviving entity immediately after such merger, reorganization or consolidation; or (iii) a sale
of all or substantially all of the assets of such Party in one transaction or a series of related transactions to a Third Party. The
acquiring or combining Third Party in any of (i), (ii) or (iii), and any of such Third Party’s Affiliates (whether in existence
as of or at any time following the applicable transaction, but other than the acquired Party and its Affiliates as in existence prior
to the applicable transaction) are referred to collectively herein as the “Acquirer”; or

 

(b)         [***].

 

1.16         “Claim”
has the meaning set forth in Section 11.1.1.

 

1.17         “Clinical
Trial” means a human clinical trial, including a Phase I Clinical Trial, Phase II Clinical Trial or Phase III Clinical
Trial, or any post-Regulatory Approval human clinical trial, as applicable.

 

1.18         “Combination
Product” has the meaning set forth in the definition of “Net Sales.”

 

1.19         “Commercial
Milestone Event” has the meaning set forth in Section 8.2.

 

1.20         “Commercialization”
means any and all activities directed to the offering for sale of a Compound, Product or other compound, product or therapy including:
(a) activities directed to storing, marketing, promoting, detailing, distributing, importing, exporting, selling and offering to
sell that Compound, Product, or other compound, product or therapy; (b) conducting Clinical Trials after Marketing Approval of a
Compound, Product, or other compound, product or therapy with respect to such Compound, Product, or other compound, product or therapy;
(c) interacting with Regulatory Authorities regarding the foregoing; and (d) seeking Regulatory Approvals (as applicable) for
and registration of that Compound, Product, or other compound, product or therapy beyond Marketing Approval (which is addressed within
 “Development”). When used as a verb, “to Commercialize” and “Commercializing”
means to engage in Commercialization and “Commercialized” has a corresponding meaning. For clarity, “Commercialization”
shall not include any Development activities prior to Marketing Approval.

 

1.21         “Commercially
Reasonable Efforts” of a Party means [***].

 

1.22         “Competing
Program” has the meaning set forth in Section 7.2.1.

 

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1.23         “Compound”
means any antisense oligonucleotide sequence designed to target, or which does target, a specific adenosine in an RNA expressed from
a Target for deamination by an endogenous ADAR, which sequence is selected, discovered, generated, optimized or identified under the
Research Program and Directed To a Project Target.

 

1.24         “Confidential
Proprietary Information” has the meaning set forth in Section 12.1.1.

 

1.25         “Confidentiality
Agreement” means that certain Non-Disclosure Agreement entered into between ProQR and Lilly as of [***], and as it may
be amended.

 

1.26         “Control”
or “Controlled” means, with respect to any Know-How, Patents, or other Technology or Intellectual Property
Rights, that a Party has the legal authority or right (whether by ownership, license, or otherwise) to grant to the other Party a license,
covenant not to sue, sublicense, access, or right to use (as applicable) under such Know-How, Patents, or other Technology or Intellectual
Property Rights, on the terms and conditions set forth herein, in each case without violating any obligations of the granting Party owed
to a Third Party, breaching the terms of any agreement with a Third Party or subjecting the granting Party to any additional fee or charge;
provided that: (i) [***]; (ii) [***]; and (iii) [***].

 

1.27         “Cover”
means, with respect to a claim of a Patent and a relevant Product, that such claim would be infringed, absent a license, by the Exploitation
of such Product (considering claims of patent applications to be issued as then pending).

 

1.28         “Critical
Success Factors” or “CSFs” means, with respect to a Project Target, (a) the Hit CSFs,
or (b) the criteria that must be met in order for a Hit to be designated as a Successful Candidate.

 

1.29         “Development”
or “Develop” means any and all activities directed to the non-clinical and clinical drug development activities
that are necessary or useful to obtain Marketing Approval for a Compound, Product, or other compound, product or therapy, including design
and conduct of Clinical Trials and the preparation and filing of Regulatory Filings and all regulatory affairs related to the foregoing.
When used as a verb, “Developing” means to engage in Development and “Developed”
has a corresponding meaning. For clarity, “Development” shall not include any Commercialization activities.

 

1.30         “Development
Milestone Event” has the meaning set forth in Section 8.2.

 

1.31         “Development
Milestone Payment” has the meaning set forth in Section 8.2.

 

1.32         “Directed
To” means, with respect to (a) a compound, product or therapy, and (b) a Target, that the molecule contained
in such compound, product or therapy is directed to, binds to, modulates or otherwise utilizes such Target.

 

1.33         “Disclosing
Party” has the meaning set forth in Section 12.1.2.

 

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1.34         “Discontinued
Target” means: (a) a Target that was previously a Lilly Target or Project Target but has been replaced by a Replacement
Project pursuant to Section 3.5; (b) a Project Target for which Lilly discontinues Research pursuant to Section 3.6 or
(c) a Project Target for which Lilly’s rights and licenses hereunder are terminated in accordance with this Agreement, including
Article 13.

 

1.35         “Dispute”
has the meaning set forth in Section 14.2.

 

1.36         “Divestiture”
has the meaning set forth in Section 7.2.1.

 

1.37         “Dollar”
means a U.S. dollar, and “$” is to be interpreted accordingly.

 

1.38         “Effective
Date” has the meaning set forth in the preamble to this Agreement.

 

1.39         “Eli
Lilly and Company Animal Care and Use Requirement for Animal Researchers and Suppliers” has the meaning set forth in Section 4.7.

 

1.40         “Eli
Lilly and Company Good Research Practices” has the meaning set forth in Section 4.7.

 

1.41         “Enabling
Technology” has the meaning set forth in Section 9.5.

 

1.42         “Excluded
Technology” means any Patents Covering or Know-How directed to Manufacturing processes, drug delivery technology, or devices.

 

1.43         “Exclusivity
Period” has the meaning set forth in Section 7.1.3.

 

1.44         “Executive
Officers” means (a) with respect to ProQR, [***], and (b) with respect to Lilly, [***]; or any other person that
the relevant party designates from time to time.

 

1.45         “Exploitation”
means, individually or collectively, the Research, Development, registration, Manufacture, making, having made, use, storage, keeping,
importation, exportation, transportation, promotion, marketing, distribution, offering for sale, sale or other exploitation or Commercialization
of a Compound or Product or other compound, product or therapy. When used as a verb, “to Exploit” and “Exploiting”
mean to engage in Exploitation, and “Exploited” has a corresponding meaning.

 

1.46         “FD&C
Act” means the United States Federal Food, Drug and Cosmetic Act, as amended.

 

1.47         “FDA”
means the United States Food and Drug Administration or any successor agency thereto.

 

1.48         “Field”
means any and all purposes and uses, including diagnostic, prophylactic, and therapeutic treatment of any disease or medical condition
in humans and animals.

 

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1.49         “Firewall
Event” has the meaning set forth in Section 15.8.5.

 

1.50         “Firewall
Period” means, [***].

 

1.51         “Firewalls”
means effective walls and screens established between ProQR, on the one hand, and on the other hand, an Acquirer of ProQR which has a
Competing Program, to ensure that no non-public information, materials (such as lab notebooks, document management systems or other documented
or memorialized Know-How) or non-personnel resources directly relating to any Project Targets, Lilly Targets, Compounds, Products, or
the Research Program, or any non-public information, materials or non-personnel resources relating to Patents provided, or made accessible,
to ProQR by Lilly are accessible by personnel of the Acquirer working on the Competing Program during the Firewall Period. For purposes
of this definition, “Firewalls” shall include, during the Firewall Period, as necessary to satisfy this definition: (a) walls
and screens (whether technical or physical) between (i) on the one hand, personnel of ProQR performing Research or Development activities
under, or otherwise working on or involved with, the Research Program or having access to any ProQR Technology, non-public materials
(such as lab notebooks, document management systems or other forms in which such Know-How may be memorialized) or non-personnel resources,
in each case, directly relating to the Research Program (all of the foregoing, collectively, “Collaboration Personnel”)
and (ii) on the other hand, personnel of an Acquirer working on the Competing Program, or having access to any non-public materials
(such as lab notebooks, document management systems or other forms in which such Know-How may be memorialized) or non-personnel resources,
in each case, directly relating to the Competing Program (all of the foregoing, collectively, “Competing Program Personnel”);
and (b) processes ensuring that (i) Collaboration Personnel do not perform any Research or Development activities under the
Competing Program or have access to any non-public materials (such as lab notebooks, document management systems or other forms in which
such Know-How may be memorialized) or non-personnel resources, in each case, directly relating to the Competing Program and (ii) Competing
Program Personnel do not perform Research or Development activities or any other work under the Research Program or have access to any
ProQR Technology, non-public materials (such as lab notebooks, document management systems or other forms in which such Know-How may
be memorialized) or non-personnel resources, in each case, directly relating to the Research Program. Notwithstanding the foregoing,
 “Firewalls” shall not (i) require activities specific to the Research Program to be performed in a separate facility
than activities that are specific to the Competing Program, provided that the activities specific to the Research Program are performed
in a different location within such facility than the activities specific to the Competing Program, (ii) restrict Collaboration
Personnel from working together with Competing Program Personnel on Exploitation of any compound or product that is neither the subject
of the Research Program nor the subject of the Competing Program, or (iii) restrict executive officers or members of the board of
directors of ProQR or its Affiliates, including the Acquirer, from accessing or receiving disclosure of information solely as necessary
to enable executive officers and the board of directors to comply with (x) their fiduciary obligations to ProQR or its Affiliates
or (y) Applicable Laws; provided, however, that such executive officers or members of the board of directors are prohibited from
using any information pertaining to the Research Program or any other activities covered under this Agreement to inform or make decisions
regarding or relating to any Competing Programs.

 

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1.52         “First
Commercial Sale” means [***].

 

1.53         “Generic
Equivalent” means, with reference to a Product, any biologic or pharmaceutical product (including any competing equivalent
product using endogenous ADAR recruitment which effects the same Target as a Compound) that is sold by a Third Party (other than a licensee
of Lilly or any of its Affiliates) and that is approved for marketing and/or sale by a Regulatory Authority in reliance on or using data
from the Regulatory Filings for the Product that were submitted by Lilly, its Affiliates, or their licensees, and that meets the equivalency
determination by the applicable Regulatory Authority in such country as is necessary to permit substitution of such product for the Product
under applicable law in such country.

 

1.54         “GLP
Toxicology Study” means, with respect to a Compound or Product, an in vivo toxicology study that is conducted in
compliance with then-current Good Laboratory Practices.

 

1.55         “Good
Clinical Practices” or “cGCP” means all applicable current Good Clinical Practice standards for
the design, conduct, performance, monitoring, auditing, recording, analysis and reporting of Clinical Trials, including, as applicable:
(a) as set forth in the International Conference on Harmonisation of Technical Requirements for Registration of Pharmaceuticals
for Human Use (“ICH”) E6 and any other guidelines for good clinical practice for trials on medicinal products
in the Territory; (b) the Declaration of Helsinki (2004) as last amended at the 52nd World Medical Association in October 2000
and any further amendments or clarifications thereto; (c) U.S. Code of Federal Regulations Title 21, Parts 50, 54, 56, 312 and 314,
as may be amended from time to time; and (d) the equivalent Applicable Laws in any relevant country, each as may be amended and
applicable from time to time, and in each case (of (a)-(d)), that provide for, among other things, assurance that the clinical data and
reported results are credible and accurate and protect the rights, integrity, and confidentiality of trial subjects.

 

1.56         “Good
Laboratory Practices” or “GLPs” means the then-current standards for laboratory activities for
pharmaceuticals, as set forth in the FDA’s Good Laboratory Practice regulations as defined in 21 C.F.R. Part 58, the Council
Directive 87/18/EEC, as amended, the principles for Good Laboratory Practice and/or the Good Laboratory Practice principles of the Organization
for Economic Co-Operation and Development (“OECD”), and such standards of good laboratory practice as are required
by the European Union and other organizations and governmental agencies in countries in which a Product is intended to be sold, to the
extent such standards are not less stringent than United States Good Laboratory Practice.

 

1.57         “Good
Research Practices” or “GRP” means research practices consistent with: (a) the research
quality standards defining how Lilly’s research laboratories conduct good science for non-regulated work as set forth in Exhibit 4.7
Part A of this Agreement; and (b) the Research Quality Association (RQA), 2014 Quality in Research Guidelines for Working
in Non-Regulated Research; (c) the WHO Quality Practices in Basic Biomedical Research Guidelines; and (d) the equivalent Applicable
Laws if any, in any relevant country, each as may be amended and applicable from time to time.

 

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1.58         “Government
Official” has the meaning set forth in Section 10.5.7.

 

1.59         “Governmental
Authority” means any national, international, federal, state, provincial or local government, or political subdivision
thereof, or any multinational organization or any authority, agency or commission entitled to exercise any administrative, executive,
judicial, legislative, police, regulatory or taxing authority or power, and any court or tribunal (or any department, bureau or division
thereof, or any governmental arbitrator or arbitral body).

 

1.60         “Hit”
means a Compound(s) that: (a) is discovered or identified by ProQR; and (b) meets the applicable Hit CSFs for such Project
Target.

 

1.61         “Hit
CSFs” means, with respect to a Project Target, the CSFs for such Project Target established by the JSC necessary for a
Compound(s) Directed To such Project Target to achieve a “Hit.”

 

1.62         “Improvement”
means any: (a) modification, enhancement or change to the applicable Patents and/or Know-How of a Party created, conceived of or
reduced to practice or acquired in the course of performing a Project; or (b) Patent right claiming any Know-How described in the
foregoing subclause (a).

 

1.63         “IND”
means an investigational new drug application filed with the FDA or any similar application filed with a Regulatory Authority in a country
outside the U.S. required to commence Clinical Trials of a pharmaceutical product.

 

1.64         “Indemnitee”
has the meaning set forth in Section 11.1.3.

 

1.65         “Indemnitor”
has the meaning set forth in Section 11.1.3.

 

1.66         “Infringement”
has the meaning set forth in Section 8.3.7.

 

1.67         “Initial
Project” has the meaning set forth in Section 3.2.

 

1.68         “Initial
Target” means the Target referred to as IDUA (UniProt identification number P35475).

 

1.69         “Initial
Workplan” has the meaning set forth in Section 4.4.2.

 

1.70         “Initiation”
means, with respect to a Clinical Trial, the first dosing in the first human subject in such Clinical Trial.

 

1.71         “Intellectual
Property Rights” means any and all proprietary rights provided under: (a) patent law, including any Patents; (b) copyright
law; (c) trademark law, or (d) any other applicable statutory provision or common law principle, including trade secret law,
that may provide a right in ideas, formulae, algorithms, concepts, inventions (whether or not patentable), or Know-How, or the expression
or use thereof.

 

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1.72         “Inventions”
means all Know-How and inventions, whether or not patentable, including all rights, title and interest in and to the Intellectual Property
Rights in all of the foregoing.

 

1.73         “[***]
Target” means an [***] Target in the central or peripheral nervous system, but excluding any Targets selected for
the purpose of developing Compounds to treat ocular diseases or conditions. For clarity, [***] Targets may be Additional Targets, Replacement
Projects, or Reserved Targets.

 

1.74         “Joint
Patents” has the meaning set forth in Section 9.7.4.

 

1.75         “Joint
Program IP” has the meaning set forth in Section 9.2.2(d).

 

1.76         “JPC”
has the meaning set forth in Section 2.4.

 

1.77         “JSC”
has the meaning set forth in Section 2.3.

 

1.78         “JSC
Co-Chairpersons” has the meaning set forth in Section 2.3.

 

1.79         “Know-How”
means any proprietary scientific or technical information, inventions, discoveries, results and data of any type whatsoever, in any tangible
or intangible form, including inventions, discoveries, databases, safety information, practices, methods, instructions, techniques, processes,
drawings, documentation, specifications, formulations, formulae, knowledge, know-how, trade secrets, materials, skill, experience, test
data and other information and technology applicable to formulations, compositions or products or to their manufacture, development,
registration, use, marketing or sale or to methods of assaying or testing them, including pharmacological, pharmaceutical, medicinal
chemistry, biological, chemical, biochemical, toxicological and clinical test data, physical and analytical, safety, quality control
data, manufacturing, and stability data, studies and procedures, and manufacturing process and development information, results and data.

 

1.80         “Lilly”
has the meaning set forth in the Preamble.

 

1.81         “Lilly
Background IP” means any and all Patents and Know-How that Lilly or its Affiliates: (a) owns or Controls as of the
Effective Date; or (b) arises outside the scope of the Research Program after the Effective Date and is Controlled by Lilly.

 

1.82         “Lilly
Competitor” means a company that: (a) [***]; and (b) either (i) [***], or (ii) [***].

 

1.83         “Lilly
Indemnitee” has the meaning set forth in Section 11.1.1.

 

1.84         “Lilly
Patent” means any Patent constituting or claiming any Lilly Background IP or Lilly Program IP.

 

1.85         “Lilly
Program IP” has the meaning set forth in Section 9.2.2(b).

 

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1.86         “Lilly
Targets” means, individually or collectively, each Target that is a Project Target or of which a Project Target is a specific
adenosine, element or mutated form. For avoidance of doubt, (i) there may be multiple Project Targets associated with a Lilly Target
and a Lilly Target shall remain a Lilly Target so long (and only so long) as there is at least [***] Project Target associated with it
and (ii) when there is no Project Target associated with a Lilly Target (because, for example, the Project Target has become a Discontinued
Target), then such Lilly Target shall cease to be a Lilly Target.

 

1.87         “Lilly
Technology” means, individually or collectively, the Lilly Background IP, Lilly Program IP, and Lilly’s interest
in Joint Program IP.

 

1.88         “Loss
of Market Exclusivity” means, [***].

 

1.89         “Losses”
has the meaning set forth in Section 11.1.1.

 

1.90         “Manufacture”
and “Manufacturing” means any and all activities related to the production, manufacture, formulation, finishing,
packaging, labeling, shipping and holding of any Compound or Product, or other compound, product or therapy, or any component, intermediary
or precursor thereof (including, for clarity, expression vectors, cell lines, culture media and feeds), and including process development,
process qualification and validation, scale-up, pre-clinical, clinical and commercial manufacture, quality assurance and quality control
(including testing).

 

1.91         “Marketing
Approval” means all approvals, licenses, permits, notifications, registrations, clearances, authorizations, or waivers
of any Regulatory Authority, that is or are necessary to initiate marketing and selling a Product in the Field in a particular jurisdiction,
including a BLA/NDA.

 

1.92         “Milestone
Events” has the meaning set forth in Section 8.2.

 

1.93         “Milestone
Payments” has the meaning set forth in Section 8.2.

 

1.94         “Net
Sales” means, with respect to a particular Product, the gross amount invoiced by Lilly or a Lilly Affiliate or any Sublicensee
thereof to unrelated Third Parties, excluding any sublicensee, for the Product in the Territory, less the following items consistent
with U.S. Generally Accepted Accounting Principles (U.S. GAAP), consistently applied: [***]

 

Such amounts shall be determined
from the books and records of Lilly or its Affiliate or Sublicensee, maintained in accordance with U. S. GAAP or, in the case of Sublicensees,
such similar accounting principles, consistently applied. Lilly further agrees in determining such amounts, it will use Lilly’s
then current standard procedures and methodology, including Lilly’s then current standard exchange rate methodology for the translation
of foreign currency sales into U.S. Dollars or, in the case of Sublicensees, such similar methodology, consistently applied.

 

In the event that the Product
is sold as part of a Combination Product (where “Combination Product” means any pharmaceutical product which
comprises the Product and other active compound(s), ingredient(s) and/or device(s)), the Net Sales of the Product, for the purposes
of determining royalty and commercial milestone payments, shall be determined [***].

 

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In the event that the weighted
average sale price of the Product can be determined but the weighted average sale price of the other active compound(s), ingredient(s) and/or
device(s) cannot be determined, Net Sales for purposes of determining royalty payments shall be calculated by [***].

 

In the event that the weighted
average sale price of the other active compound(s), ingredient(s) and/or device(s) can be determined but the weighted average
sale price of the Product cannot be determined, Net Sales for purposes of determining royalty payments shall be calculated by [***].

 

In the event that the weighted
average sale price of both the Product and the other active compound(s), ingredient(s) and/or device(s) in the Combination
Product cannot be determined, the Net Sales of the Product shall be deemed to be equal to the mutually agreed (by the Parties) percentage
of the Net Sales of the Combination Product, based on the relative value and/or cost of the Product and other active compound(s), ingredient(s) and/or
device(s) in such Combination Product; provided, however, that in the event the Parties cannot, in spite of good faith efforts,
mutually agree to such a percentage, then such percentage shall be equal to [***] of the Net Sales of the Combination Product.

 

The weighted average sale
price for a Product, other active compound(s), ingredient(s) and/or device(s), or Combination Product shall be calculated [***]
and such price shall be used during all applicable royalty reporting periods for [***]. When determining the weighted average sale price
of a Product, other active compound(s), ingredient(s) and/or device(s), or Combination Product, the weighted average sale price
shall be calculated by [***]. In the initial Calendar Year, a forecasted weighted average sale price will be used for the Product, other
active compound(s), ingredient(s) and/or device(s), or Combination Product. Any over or under payment due to a difference between
forecasted and actual weighted average sale prices will be paid or credited in the first royalty payment of the following [***].

 

1.95         “Orphan
Indication” means an indication for use of a drug to treat a rare disease or condition where the number of people affected
by the disease or condition is less than 200,000 persons in the U.S. or where the indication for use otherwise meets the criteria for
orphan drug designation under section 526(a) of the FD&C Act and 21 C.F.R. 316.21.

 

1.96         “Party”
and “Parties” has the meaning set forth in the Preamble.

 

1.97         “Patents”
means: (a) patents and patent applications; (b) any and all divisionals, continuations, continuations-in-part, reissues, renewals,
substitutions, registrations, re-examinations, revalidations, extensions, supplementary protection certificates and the like of any such
patents and patent applications; and (c) any and all foreign equivalents of the foregoing.

 

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1.98         “Person”
means any corporation, limited or general partnership, limited liability company, joint venture, trust, unincorporated association, governmental
body, authority, bureau or agency, any other entity or body, or an individual.

 

1.99         “Personal
Information” means, in addition to any definition for any similar term (e.g., “personal data” or “personally
identifiable information” or “PII”) provided by Applicable Laws, or by either Party in any of its own privacy policies,
notices or contracts, all information that identifies, could be used to identify or is otherwise associated with an individual person,
whether or not such information is directly associated with an identified individual person.

 

1.100       “Phase
I Clinical Trial” means a clinical trial of a Product generally consistent with 21 C.F.R. § 312.21(a) (or
the non-United States equivalent thereof).

 

1.101       “Phase
II Clinical Trial” means a clinical trial of a Product generally consistent with 21 C.F.R. § 312.21(b) (or
the non-United States equivalent thereof).

 

1.102       “Phase
III Clinical Trial” means a clinical trial of a Product generally consistent with 21 C.F.R. § 312.21(c) (or
the non-United States equivalent thereof).

 

1.103       “Pricing
and Reimbursement Approval” means, with respect to a Product, the approval, agreement, determination or decision of any
Regulatory Authority establishing the price or level of reimbursement for such Product, as required in a given country or jurisdiction
prior to sale of such Product in such country or jurisdiction.

 

1.104       “Product”
means any pharmaceutical composition, preparation, formulation or product containing or comprising a Compound for which the applicable
Candidate Success CSFs were achieved during the Research Term.

 

1.105       “Product-Specific
Patent” means a Patent filed during or after the Research Term, that (a) claims (i) a Compound, Product or Project
Target or (ii) a method of making or using a Compound or Product, wherein in either case (clause (i) or (ii)), such claim of
such Patent includes Specific Identification of the relevant Compound, Product or Project Target or (b) contains a claim that includes
a genus that identifies a Compound, Product or Project Target, or a method of making or using a Compound, Product or Project Target.

 

1.106       “Program
IP” has the meaning set forth in Section 9.2.2.

 

1.107       “Project”
has the meaning set forth in Section 3.2.

 

1.108       “Project
Leader” has the meaning set forth in Section 2.2.

 

1.109       “Project
Target” means a specific adenosine in a Target, or in an element of a Target, or in a mutated form of a Target, where such
adenosine is designated as a “Project Target” by Lilly pursuant to Article 3. As used in this Agreement, “Project
Target” includes the Initial Target.

 

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1.110       “ProQR”
has the meaning set forth in the Preamble.

 

1.111       “ProQR
Background IP” means any and all Patents and Know-How that ProQR or its Affiliates: (a) Controls as of the Effective
Date; or (b) arises outside the scope of the Research Program after the Effective Date and is Controlled by ProQR.

 

1.112       “ProQR
Patent” means any Patent included in the ProQR Technology.

 

1.113       “ProQR
Platform” means AXIOMER® technology which relates to adenosine to inosine editing by adenosine deaminase acting on
RNA (ADAR) performed at the specific location targeted by the editing oligonucleotide which is covered by ProQR Patents or is proprietary
to ProQR. For the avoidance of doubt, the ProQR Platform does not include specific Compounds or Products.

 

1.114       “ProQR
Platform IP” means ProQR Background IP that relates to all or part of the ProQR Platform, and all Improvements thereto.

 

1.115       “ProQR
Platform Patent” means all Patents that claim the ProQR Platform IP. The ProQR Platform Patents are listed in Exhibit 1.115.

 

1.116       “ProQR
Program IP” has the meaning set forth in Section 9.2.2(c).

 

1.117       “ProQR
Supplemental IP” means any ProQR Background IP (other than ProQR Platform IP) that the JSC determines is useful (a) to
apply in the execution of a Project, or (b) to Exploit Compounds or Products in the Field in the Territory. For avoidance of doubt,
ProQR Supplemental IP shall not include (1) technological approaches to modulating Targets other than as included within the ProQR
Platform but may include complementary technologies such as formulation, delivery or targeting technologies, or (2) ProQR Platform
IP, which shall be included in the licenses granted to Lilly under this Agreement as of the Effective Date and shall not be subject to
JSC determination of usefulness.

 

1.118       “ProQR
Technology” means, individually or collectively, the ProQR Background IP, the ProQR Program IP, or ProQR’s interest
in any Joint Program IP, in each case, that is (i) related to the ProQR Platform, (ii) specific to the deamination of a Project
Target with a Compound or (iii) ProQR Supplemental IP that the JSC determines in accordance with Section 2.6(g) to be
useful to the Exploitation of Compounds or Products in the Field in the Territory. For avoidance of doubt, the ProQR Technology includes
the ProQR Platform IP.

 

1.119       “Prosecute
and Maintain” or “Prosecution and Maintenance” with respect to a particular Patent, means all
activities associated with the preparation, filing, prosecution and maintenance of such Patent, together with the conduct of interferences,
derivation proceedings, inter partes review and post-grant review, the defense of oppositions and other similar proceedings with
respect to that Patent, including any activities associated with claims, including as a counterclaim or declaratory judgment action,
of unpatentability, invalidity or unenforceability of such Patent that are brought by a Third Party in connection with an Infringement
under Section 9.8.

 

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1.120       “Prosecuting
Party” has the meaning set forth in Section 9.7.4.

 

1.121       “Receiving
Party” has the meaning set forth in Section 12.1.2.

 

1.122       “Regulatory
Approvals” means, collectively, any and all approvals (including supplements, amendments, pre- and post-approvals, Pricing
and Reimbursement Approvals), licenses, registrations, permits, notifications, and authorizations (including marketing and labeling authorizations)
or waivers of any Regulatory Authority that are necessary for the testing or Exploitation of a pharmaceutical product (including any
Product) in any country or jurisdiction, including Pricing and Reimbursement Approval, as applicable.

 

1.123       “Regulatory
Authority” means any Governmental Authority that has responsibility in its applicable jurisdiction over the testing or
Exploitation of pharmaceutical products (including any Product) in any country or jurisdiction. For countries or jurisdictions where
governmental approval is required for pricing or reimbursement for a pharmaceutical product (including any Product) to be reimbursed
by national health insurance (or its local equivalent), Regulatory Authority includes any Governmental Authority whose review or approval
of pricing or reimbursement of such product is required.

 

1.124       “Regulatory
Filings” means, collectively, any and all applications, filings, submissions, approvals (including supplements, amendments,
pre- and post-approvals, Pricing and Reimbursement Approvals), licenses, registrations, permits, notifications, and authorizations (including
marketing and labeling authorizations), non-clinical and clinical study authorization applications or notifications (including all supporting
files, writings, data, studies and reports) or waivers with respect to the testing or Exploitation of a Compound or Product made to or
received from any Regulatory Authority in a given country or jurisdiction, including INDs and BLAs.

 

1.125       “Replacement
Project” has the meaning set forth in Section 3.5.

 

1.126       “Replacement
Target” has the meaning set forth in Section 3.5.

 

1.127       “Research”
means any and all activities directed to the discovery, identification, screening, testing, assessment and optimization of Project Targets,
or Compounds or Products, or other compound, product or therapy, including, with respect to ProQR, such activities directed to the generation
of Hits and Successful Candidates with respect to a Project Target. “Research” refers to those activities prior to Development.
When used as a verb, “Researching” means to engage in Research and “Researched” has
a corresponding meaning.

 

1.128       “Research
Continuance Scenario” has the meaning set forth in Section 15.8.2(b).

 

1.129       “Research
Program” has the meaning set forth in Section 3.1.

 

1.130       “Research
Term” has the meaning set forth in Section 4.3.

 

1.131       “Research
Transfer Scenario” has the meaning set forth in Section 15.8.2(a).

 

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1.132       “Reserved
Target” has the meaning set forth in Section 3.3.1.

 

1.133       “Residuals”
has the meaning set forth in Section 12.1.5.

 

1.134       “Royalty”
has the meaning set forth in Section 8.3.2.

 

1.135       “Royalty
Term” has the meaning set forth in Section 8.3.1.

 

1.136       “Specific
Identification” or “Specifically Identify” means, with respect to the identification of a Compound,
Project Target or Product in a Patent, that any claim of such Patent recites such Compound, Project Target or Product, as applicable.

 

1.137       “Sublicensee”
means a Third Party that is granted a license or sublicense to Exploit Compounds or Products in the Field in the Territory, beyond the
mere right to purchase Products from Lilly and its Affiliates or resell such Products, and excludes Lilly’s Affiliates or Third
Party subcontractors that act solely for the benefit of Lilly or its Affiliates in the supply chain or that perform discrete services
(as opposed to being granted broad rights or responsibilities) on behalf of Lilly or its Affiliates.

 

1.138       “Successful
Candidate” means a Hit which has met the Project-specific Candidate Success CSFs.

 

1.139       “Target”
means an RNA sequence expressed from a specific gene or gene sequence that can be identified by a UniProt identification number (https://www.uniprot.org/)
(including pre-mRNA and mRNA).

 

1.140       “Technology”
means all technology and information, including all Inventions, discoveries, data, assays, protocols, databases, results, information,
trade secrets, ideas, concepts, formulas, techniques, and methods.

 

1.141       “Term”
has the meaning set forth in Section 13.1.

 

1.142       “Terminated
Product” has the meaning set forth in Section 13.5.

 

1.143       “Territory”
means worldwide.

 

1.144       “Third
Party” means any Person other than Lilly or ProQR or an Affiliate of ProQR or Lilly.

 

1.145       “U.S.”
means the United States of America and its territories and possessions.

 

1.146       “Unavailable
Target” means a Target that, at the time Lilly seeks to add such Target as a Reserved Target, or proposes such Target (or
specific portion or mutation thereof) for inclusion in the Research Program: (a) is a Target that is the subject of a final agreed
upon term sheet or letter of intent between ProQR and any Third Party; (b) on which ProQR is actively working either as an internal
ProQR program or with another collaborator; or (c) as to which ProQR has granted rights for such Target that would conflict with
or prevent the grant of rights to Lilly under this Agreement for such Target.

 

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1.147       “Valid
Claim” means a claim for a composition of matter, or approved labelled method of use (such that alone or with other Valid
Claims all such methods of use are covered) contained in: (a) an issued, unexpired and granted Patent owned or Controlled by ProQR
or its Affiliates, which claim has not been held unenforceable, unpatentable or invalid by a decision of a court or other Governmental
Authority of competent jurisdiction; or (b) a pending Patent owned or Controlled by ProQR or its Affiliates that: (i) has been
asserted and continues to be prosecuted in good faith, (ii) has not been abandoned or finally rejected without the possibility of
appeal or refiling, and (iii) has not been pending for more than [***] from the earliest priority date. Further, a claim shall only
be considered a Valid Claim if contained in an application (or a patent issuing from an application) that is or claims priority to an
application filed before or during the Research Term which contains support for such claim.

 

1.148       “Working
Group” has the meaning set forth in Section 2.5.

 

1.149       “Workplan”
has the meaning set forth in Section 4.4.1.

 

Article 2

 

GOVERNANCE AND JOINT STEERING COMMITTEE

 

2.1           Alliance
Managers. Within [***] following the Effective Date, each Party shall appoint [***] to act as the Alliance Manager for such Party
(each, an “Alliance Manager”). Without limiting the responsibilities and authorities of the Project Leaders
and the JSC (as expressly set forth herein), the Alliance Managers shall each be the primary point of contact for the Parties regarding
the collaboration and related activities contemplated by this Agreement and shall help facilitate all such activities hereunder. For
avoidance of doubt, the individual appointed by a Party to act as an Alliance Manager may, but need not, be the same individual appointed
by such Party as a Project Leader, but an Alliance Manager may not be appointed to serve simultaneously as a JSC member. Either Party
may change its Alliance Manager at any time upon prior written notice to the other Party.

 

2.2           Project
Leaders. Lilly and ProQR shall each assign [***] to serve as the primary point of contact between the Parties with respect to each
Project under the Research Program (each, a “Project Leader”). The Project Leaders shall regularly communicate
with each other to address Project-related issues, needs and updates and facilitate communications and organization of Working Groups
(as needed and if established by the JSC) associated with the Workplan for such Project. Either Party may change its Project Leaders
at any time upon reasonable prior written notice to the other Party. For clarity, the same employee may, but need not, be the Project
Leader for multiple Projects.

 

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2.3           Joint
Steering Committee. Within [***] after the Effective Date, the Parties shall establish a cross-functional, joint steering committee
(the “JSC”) composed of up to [***] senior representatives from each Party (provided each Party has an equal
number of representatives on the JSC at all times) that will oversee and coordinate the Parties’ activities related to the Research
Program. The JSC may, from time to time, establish subcommittees and Working Groups as it deems necessary to further the purposes of
this Agreement and the Research Program. Each Party shall appoint its respective representatives to the JSC from time to time, and may
change its representatives, in its sole discretion, effective upon reasonable prior written notice to the other Party designating such
change. The representatives from each Party shall have appropriate technical credentials, experience and knowledge pertaining to and
ongoing familiarity with the Research Program and the activities conducted thereunder. Each Party shall designate [***] of its representatives
on the JSC to serve as a JSC co-chairperson (collectively, “JSC Co-Chairpersons”), who will be jointly responsible
for calling meetings of the JSC, circulating agendas and performing administrative tasks required to assure efficient operation of the
JSC but shall not have any extra or additional votes or authority. The JSC Co-Chairpersons or their designees shall alternate responsibility
for circulating agendas at least [***] prior to each meeting and distributing minutes of the meetings pursuant to Section 2.7.

 

2.4           Formation
of JPC; JPC Functions and Powers. The JSC will establish a joint patent subcommittee (“JPC”), consisting
of [***] subject matter experts (at least one of which must be a patent attorney admitted to practice and in good standing with the USPTO
or the European Patent Office) from each Party or such other number as the JSC may agree upon (with an equal number of experts from each
of Lilly and ProQR), within [***] after the Effective Date. The JPC will be responsible for evaluating technology arising during the
Research Term under this Agreement, including (a) making initial determination of inventorship, (b) determining whether such
technology is Program IP, ProQR Platform IP, or Joint Program IP (including whether certain technologies can be separately categorized
and separately patentable and subject to separate assignment and license obligations hereunder), (c) coordination of the Parties
with respect to managing the preparation, filing, prosecution, maintenance, enforcement and defense of Joint Program IP (and determining
whether any such Joint Program IP should be maintained as trade secret in lieu of patenting), and (d) determine timing of submission
of, and evaluating, drafts for public disclosure by way of publication in a scientific journal, by presentation at scientific conferences,
or by any other means as provided in Section 12.3.

 

2.5           Working
Groups. The Parties may establish working groups consisting of members from both ProQR and Lilly (each, a “Working Group”)
to oversee aspects of the activities of the Research Program or each individual Project. From time to time, the Parties may establish
additional Working Groups as needed to oversee particular activities and/or projects. Each Working Group shall undertake the activities
specified under this Agreement for such Working Group or otherwise delegated to it by the JSC. During the process of establishing each
Working Group, such Working Group and the JSC shall agree regarding which matters such Working Group may resolve on its own (with discretion
to refer any matter to the JSC for a final decision) and which matters such Working Group will advise the JSC and/or the Project Leaders
regarding (and with respect to which such advice-specific matters the JSC must resolve).

 

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2.6           Function
and Powers of the JSC. The JSC will:

 

(a)           review,
discuss, and approve any amendments to the Workplans that may be necessary or desired, and prepare, discuss, and approve any Additional
Workplans, in accordance with Section 4.4; provided that, unless ProQR otherwise consents, each amendment to a Workplan and Additional
Workplan shall only include obligations for ProQR that are substantially similar in nature and extent to those included in the Initial
Workplan;

 

(b)           oversee
the implementation of the Workplans, including the activities, timing and deliverables thereunder, and coordination of such activities
and timing across Research Programs;

 

(c)           approve
(in the case of Additional Projects and Replacement Projects), and refine as necessary (for all Projects), the Critical Success Factors
for each Project, and determine whether the applicable Critical Success Factors have been satisfied;

 

(d)           determine
whether a Project with respect to a Project Target should be discontinued;

 

(e)           without
limiting Lilly’s rights pursuant to Section 3.5 to select a Replacement Target, and subject to Lilly’s decision-making
authority in Section 2.9, discuss Lilly’s selection of Replacement Targets;

 

(f)           discuss
the progress of the Research Program and Projects generally, and the validation and development of Compounds Directed To the Project
Targets;

 

(g)           [***];

 

(h)           provide
a forum for the Parties to share and discuss information relating to the research and validation of Compounds Directed To the Project
Targets, including the results of the activities being carried out under the Workplans;

 

(i)           address
issues arising in the performance of the Workplans;

 

(j)           establish
Working Groups, and direct and oversee any operating Working Groups on all significant issues, and resolve disputed matters that may
arise at the Working Groups;

 

(k)           facilitate
the exchange of Know-How or any materials as required hereunder (including pursuant to Section 4.9);

 

(l)           oversee
winding down of the Research Program or a Project with respect to discontinued Project Targets; and

 

(m)          perform
any and all tasks and responsibilities that are expressly attributed to the JSC under this Agreement or as otherwise agreed by the Parties
in writing.

 

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2.7           Meetings.
The JSC will meet at least [***] for so long as the JSC remains in effect. The JSC may conduct such meetings by telephone, videoconference,
or in person. Each Party may call special meetings of the JSC with at least [***] prior written notice, or a shorter time period in exigent
circumstances, to resolve particular matters requested by such Party that are within the purview of the JSC. Meetings of the JSC are
effective only if at least one (1) representative of the JSC for each Party participates in such meeting. Each Alliance Manager
shall be permitted to attend meetings of the JSC, and any Working Group, as a non-voting observer. Each Party may invite a reasonable
number of other participants, in addition to its representatives, to attend JSC meetings in a non-voting capacity; provided that if either
Party intends to have any Third Party (including any consultant) attend such a meeting, such Party shall provide prior written notice
to the other Party. Such Party shall ensure that such Third Party is bound by confidentiality and non-use obligations consistent with
the terms of this Agreement. Each Party is responsible for its own expenses incurred in connection with participating in and attending
all such meetings. The JSC Co-Chairpersons or their designees shall keep minutes of each JSC meeting that record in writing all decisions
made, action items assigned or completed and other appropriate matters. The JSC Co-Chairpersons or their designees shall send meeting
minutes to all members of the JSC promptly after a meeting for review. Each JSC member shall have [***] from receipt in which to comment
on and to approve the minutes (such approval not to be unreasonably withheld, conditioned or delayed). If a JSC member, within such time
period, does not notify the JSC Co-Chairpersons or their designees that s/he does not approve of the minutes, the minutes shall be deemed
to have been approved by such member. The Parties acknowledge and agree that, notwithstanding the requirements of this Section 2.7
for the JSC to meet [***], the Parties shall communicate and meet (as appropriate, including via the Project Leaders) on a more informal
basis as needed to discuss the progress of the Projects or the Research Program.

 

2.8           Decisions.
The JSC will endeavor to make decisions by consensus, with the representatives of each Party having, collectively, [***] on behalf of
that Party. If the JSC cannot reach consensus or a dispute arises that cannot be resolved within the JSC, either Party may refer such
dispute to the Executive Officers or their delegates for resolution. If consensus cannot be reached with respect to a decision within
[***] after attempted resolution by the Executive Officers, then (a) [***] has the final decision-making authority with respect
to how [***], and (b) [***] has the final decision-making authority with respect to all other matters within the purview of the
JSC; provided, however, [***]: (a) excuses, reduces, or delays [***]; (b) negates any consent right or other rights
specifically granted or allocated to [***] under this Agreement; (c) amends, modifies, or waives compliance with the terms of this
Agreement; or (d) materially increases [***] as a result, except to the extent [***]. In addition, [***].

 

2.9           Authority.
The Alliance Managers, Project Leaders, JSC (including the JSC Co-Chairpersons), the JPC, and each Working Group have only the powers
assigned expressly to them in this Article 2 and elsewhere in this Agreement (or in the case of Working Groups, as expressly assigned
to them by the JSC). Each Party retains the rights, powers, and discretion granted to it under this Agreement and neither Party may delegate
or vest such rights, powers, or discretion in the Alliance Manager, a Project Leader, the JSC, the JSC Co-Chairpersons, the JPC, or any
Working Group, unless expressly provided for in this Agreement or the Parties expressly so agree in writing. The Alliance Managers, Project
Leaders, the JSC, the JSC Co-Chairpersons, the JPC, and any Working Groups shall not have the power to amend, waive or modify any term
of this Agreement, and no decision of the JSC shall be in contravention of any terms and conditions of this Agreement. It is understood
and agreed that issues to be formally decided by the JSC are limited to those specific issues that are expressly provided in this Agreement
to be decided by the JSC.

 

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2.10           Discontinuation
of JSC. The JSC will automatically disband with immediate effect at the end of the Research Term.

 

Article 3

 

TARGETS AND PROJECTS

 

3.1           Overview
and Purpose. During the Research Term, Lilly and ProQR will collaborate in a research and pre-clinical development program with the
intended purpose of: (a) selecting, discovering, generating, or identifying Compounds that are Directed To Project Targets; (b) performing
Research and other activities to optimize such Compounds as Hits; and (c) performing Research and other activities to optimize such
Hits as Successful Candidates (the “Research Program”). The Research Program will consist of Projects for each
Project Target selected as set forth in this Article 3 and will be conducted pursuant to Workplans, as further set forth below in
Article 4.

 

3.2           Projects.
The Research Program will consist of projects directed to Project Targets, with the goal of identifying and developing a Successful Candidate
for each such Project Target (each, a “Project”). As of the Effective Date, the Research Program consists of
the initial Project directed to the Initial Target (the “Initial Project”). The Research Program will consist
of the Initial Project and up to four (4) Additional Projects (as may be added pursuant to Section 3.4), plus any Replacement
Projects (as may replace Project Targets pursuant to Section 3.5). For avoidance of doubt, there shall be up to but no more than
five (5) active Projects at any given time during the Research Term.

 

3.3           Reserved
Targets.

 

3.3.1           As
of the Effective Date, the Parties have identified the Targets set forth on Exhibit 3.3 as Targets to be reserved under this
Agreement (the “Reserved Targets”) for potential inclusion in the Research Program through the selection by
Lilly of one or more such Reserved Targets as either Additional Targets or Replacement Targets. Lilly may update the Reserved Target
list at any time by notifying the JSC of the identity of a Target to be added as a Reserved Target or to replace a different Reserved
Target; provided that: (a) such Target to be added as a Reserved Target is not, at such time, an Unavailable Target (unless Lilly
has consented to the non-exclusive restrictions on such Target pursuant to this Section 3.3.1); and (b) in no event shall the
aggregate number of Reserved Targets exceed [***] at any given time. Lilly may also notify the JSC at any time of any Reserved Target
to be removed from the Reserved Target list. To the extent a Target is an Unavailable Target solely because ProQR has granted non-exclusive
rights for such Target that would conflict with or prevent the grant of rights to Lilly under this Agreement for such Target, then upon
Lilly’s request to add such Target as a Reserved Target or Project Target, ProQR shall promptly provide Lilly with written notice
providing a reasonably detailed description of such non-exclusive rights, and, upon written notice by Lilly consenting to such non-exclusive
restrictions, such Target shall not be deemed an Unavailable Target and Lilly shall be permitted to request to add such Target as a Reserved
Target or Project Target (whether as an Additional Target or Replacement Target); provided however, that the Parties will, as a precondition
to the addition of such a non-exclusively licensed Target as a Reserved Target or Project Target, enter into an amendment to this Agreement
as [***] to permit ProQR to comply with its obligations to the relevant Third Party licensee without such continued compliance constituting
a breach of ProQR’s obligations under this Agreement, including by amending the rights and licenses granted hereunder to Lilly
with respect to such Target and Compounds and Products Directed to such Target so that such rights and licenses are subject to the prior
license grant, and limiting ProQR’s obligations under Article 7 as they pertain to such Target and Compounds and Products
Directed To such Target to permit compliance with the prior agreement.

 

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3.3.2           During
the Research Term, only for so long as a Target is a Reserved Target in accordance with this Section 3.3, and for the entire Research
Term with respect to all Targets that are [***] Targets: (a) ProQR shall not grant rights to a Third Party for such Target that
would conflict with or prevent the grant of rights to Lilly under this Agreement in the event that such Target were to become a Project
Target; and (b) such Reserved Target or [***] Target shall be considered a Lilly Target solely for purposes of, and shall be subject
to, the exclusivity obligations set forth in Section 7.1 (subject to Section 7.2).

 

3.3.3           During
the Research Term, for so long as a Target is a Reserved Target in accordance with this Section 3.3, if (a) ProQR or its Affiliates
intend to initiate an internal program with respect to such Reserved Target, or (b) ProQR or its Affiliates have been notified by
a Third Party that such Third Party has substantive intentions to collaborate with ProQR or its Affiliates on such Reserved Target; then
ProQR shall promptly notify Lilly of the identity of such Reserved Target. Lilly may, no later than [***] after receipt of such notification,
propose an Additional Project or Replacement Project (as applicable) directed to such Reserved Target (or any specific adenosine, element
or mutated form thereof) (in which case the Reserved Target, specific element thereof or mutated form thereof shall become a Project
Target). During such notification period, such Target shall not become an Unavailable Target. If, following such notification period,
Lilly elects not to propose an Additional Project or Replacement Project (as applicable) directed to such Reserved Target (or any specific
adenosine, element or mutated form thereof), ProQR may move forward with such internal program or collaboration, as applicable and the
Target shall no longer be considered a Reserved Target.

 

3.4           Additional
Projects. Lilly shall have the right, at any time during the period of [***] following the Effective Date, to designate up to (but
no more than) four (4) additional Targets (or specific adenosines, elements or mutated forms thereof) (“Additional Targets”)
as Project Targets under this Agreement, by notifying the JSC in writing of the identity of the proposed Additional Target. If the proposed
Additional Target is (or is a specific adenosine, element of or mutated form of) a Reserved Target or [***] Target, is not an Unavailable
Target (unless Lilly has consented to the non-exclusive restrictions on such Target pursuant to Section 3.3.1 and subject to the
terms set forth in Section 3.3.1), or is otherwise consented to by ProQR, the proposed Additional Target shall be deemed a Project
Target, and in which case a Project associated with such Additional Target shall be added to the Research Program as an “Additional
Project,” and the Parties (through the JSC) shall prepare the mutually agreed upon Workplan for the Additional Project
(in accordance with Section 4.4.2).

 

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3.5           Replacement
Projects. If the JSC elects to discontinue a Project with respect to a Project Target, upon discontinuation of that Project: (a) ProQR
shall have no further obligations under the Workplan with respect to such Project Target; and (b) such Project Target shall thereafter
no longer be considered a Project Target and shall be deemed a Discontinued Target. In addition, Lilly shall have the right, no later
than [***] before the end of the Research Term, to select a Target (or specific adenosine, element or mutated form thereof) to replace
such Project Target in the Research Program as a Replacement Target by notifying the JSC of such election (each, a “Replacement
Target”). If the proposed Replacement Target is a Reserved Target or [***] Target, is not an Unavailable Target (unless
Lilly has consented to the non-exclusive restrictions on such Target pursuant to Section 3.3.1), or is otherwise consented to by
ProQR, the proposed Replacement Target shall be deemed a Project Target, and in which case a project associated with such Replacement
Target shall replace the discontinued Project as a “Replacement Project,” and the Parties (through the JSC)
shall prepare the mutually agreed upon Workplan for the Replacement Project (in accordance with Section 4.4.2). For avoidance of
doubt, Replacement Targets may, but need not be, Reserved Targets or [***] Targets, and Replacement Projects may replace either an Initial
Project or an Additional Project, and may itself be eligible to be replaced by a subsequent Replacement Project. In no event, however,
shall there be more than [***] replacements of Project Targets, in the aggregate, pursuant to this Section 3.5.

 

3.6           Research
Discontinuance. Without limiting Sections 13.3.1 and 13.3.2, Lilly may discontinue the Research with respect to a Project Target
at any time upon notice to ProQR. Upon receipt of such notice, such Target shall cease to be a Project Target, as applicable, and shall
become a Discontinued Target.

 

3.7           Discontinued
Targets. Once a Project Target becomes a Discontinued Target: (a) all rights and licenses granted under this Agreement will
terminate with respect to such Discontinued Target, and (b) neither Party will be restricted under this Agreement from developing
and commercializing compounds and products that are Directed To such Discontinued Target at its own cost and expense. For avoidance of
doubt, the exclusivity obligations set forth in Section 7.1 shall continue to apply for the duration of such obligations set forth
therein with respect to compounds and products Directed To a Target that remains a Lilly Target.

 

Article 4

 

RESEARCH PROGRAM

 

4.1           Responsibilities.
An outline of certain anticipated activities and responsibilities of the Parties during each of the phases (from Hit identification through
achievement of Candidate Success CSFs) of a typical Project under the Research Program is attached hereto as Exhibit 4.1
(the “Research Program Outline”). As may be further set forth in the Workplans, during the Research Term, [***].

 

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4.2           Diligence
Efforts. The Parties shall use Commercially Reasonable Efforts to execute and to perform, or cause to be performed, in a good scientific
manner and in compliance with Applicable Law, the Research activities assigned to them, respectively, in each Workplan.

 

4.3           Research
Term. The Research Program shall be conducted for a period commencing on the Effective Date and continuing for [***] after the Effective
Date, or as may be extended by Lilly pursuant to this Section 4.3 (as may be extended pursuant to this Section 4.3, the “Research
Term”). To ensure the Parties have sufficient time to perform Research activities, Lilly shall have a one-time option (exercisable
by Lilly at its sole discretion) to extend the Research Term for an additional period of up to [***] by notifying ProQR via the JSC of
such election no later than [***] prior to the end of the Research Term.

 

4.4           Workplans.

 

4.4.1           Content.
The Parties shall conduct the Research Program pursuant to mutually agreed upon, comprehensive written plans for each Project (each,
a “Workplan”) which Workplan shall set forth, for such Project: (a) the objective of the applicable Workplan
and the Research activities to be conducted by each of the Parties; (b) the expected resources to be allocated to performing such
Research; (c) the anticipated timeline and milestones of achieving such activities; and (d) the Critical Success Factors.

 

4.4.2           Preparation
of Workplans. The Workplan for the Initial Project has been prepared and mutually agreed upon by the Parties and is attached hereto
as Exhibit 4.4.2 (the “Initial Workplan”). The Workplan for each Additional Project and each Replacement
Project (each an “Additional Workplan”), as applicable, shall be mutually prepared by the Parties within [***]
following the addition of the relevant Project Target to the Research Program (pursuant to Section 3.4 or Section 3.5, respectively),
which shall substantially follow, in form and substance, the form of the Initial Workplan, except to the extent the Parties agree to
any deviations from such form with respect to any particular Project Target. Each Additional Workplan shall only become effective once
it has been approved by the JSC and subject to the decision making in accordance with Section 2.8. The JSC shall use reasonable
efforts to approve each Additional Workplan within [***] following the submission of such Additional Workplan to the JSC.

 

4.4.3           Approval
and Amendments. The JSC shall regularly review the Workplans (including the coordination of the activities across the Research Program
and to account for the number of active Projects and Workplans at any given time) and the progress of activities being conducted under
the Workplans, in no event less frequently than [***] times per Calendar Year. Either Party may propose amendments to the Workplan for
a particular Project from time to time as appropriate, to take into account completion, commencement, or cessation of activities contemplated
in the then-current Workplan for such Project or any newly available information related to such Project. Such amendments shall be effective
upon JSC approval and subject to the decision making in accordance with Section 2.8.

 

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4.5           Records;
Reports.

 

4.5.1           Records.
ProQR shall maintain, or cause to be maintained, during the Research Term and for a reasonable period of time thereafter that is consistent
with industry standards and applicable regulations, complete and accurate records (paper and/or electronic) of its Research data and
results (including raw data) for each Project in sufficient detail and in a good scientific manner appropriate for scientific, patent,
and regulatory purposes, which records will reasonably reflect all work performed by or on behalf of ProQR under the Workplan for each
Project. Lilly may request a copy of any such records of ProQR.

 

4.5.2           Reports
and Data Package. ProQR shall regularly report to Lilly through the JSC (or the designated Project Leader or Working Group) its results
in conducting Research under the Workplan for each Project. For each Project, ProQR shall provide the JSC with: (a) the deliverables
set forth in the Workplan for such Project, including a written report summarizing the data and information generated under each Project,
within [***] after the completion of ProQR’s Research for such Project; and (b) on a [***] basis during the Research Term,
all data and results (including raw data) generated by or on behalf of ProQR in performance of the Research for such Project under this
Agreement, including such data and results as are relevant to a determination of whether Hit CSFs or Candidate Success CSFs for each
Project have been achieved.

 

4.6           Research
Program Funding. Each Party shall bear any and all costs and expenses it incurs associated with its activities conducted under the
Research Program.

 

4.7           Certain
Standards Applicable to Work. All Research conducted by either Party for non-regulated work under this Agreement will be conducted
in accordance with the Workplans, Eli Lilly and Company Good Research Practices, Eli Lilly and Company Animal Care and Use Requirement
for Animal Researchers and Suppliers and all Applicable Laws, including those regarding data privacy and data security laws and regulations.
For purposes of this Agreement, “Eli Lilly and Company Good Research Practices” means the compiled set of shared
research quality standards defining how Lilly’s research laboratories conduct good science for non-regulated work as set forth
in Exhibit 4.7 Part A. For purposes of this Agreement, “Eli Lilly and Company Animal Care and Use Requirement
for Animal Researchers and Suppliers” means the guidelines relating to animal care and use for research done on behalf
of Lilly as set forth in Exhibit 4.7 Part B. If Lilly reasonably requests, ProQR will complete a self-assessment examination
form based on such quality standards. If it has not done so prior to the Effective Date, a duly authorized representative of Lilly may
make an on-site visit to ProQR for the purpose of conducting a quality assessment or quality audit for non-regulated work, which shall
be performed at Lilly’s cost and expense. Additionally, Lilly may conduct compliance audits of ProQR and/or ProQR’s Affiliates
and Third Party subcontractors engaged in work related to this Agreement, during normal business hours, no more than once annually, except
in the case of audits for cause to ensure compliance with applicable cGCP, GLP and GRP requirements, provided Lilly has requested such
audit with written notice of at least [***] and such audit does not unreasonably interfere with the audited entity’s operations.
All such audits shall be done at Lilly’s cost and expense.

 

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4.8           Subcontracting.
Each Party may engage its Affiliates or Third Party subcontractors (including contract research organizations and contract manufacturing
organizations) to perform such portions of its research obligations under the Research Program that it customarily engages for its other
similar research activities. The activities of any such Third Party subcontractors will be considered activities of such subcontracting
Party under this Agreement. The subcontracting Party shall ensure compliance by such Third Party subcontractors with the terms of this
Agreement, including any applicable Workplans. The subcontracting Party shall ensure, prior to engaging any Third Party subcontractor,
that such Third Party subcontractor is subject to written agreements containing terms and conditions that: (a) protect the rights
of the Parties under this Agreement, including by imposing obligations of confidentiality on each such Third Party subcontractor that
are no less than the obligations of confidentiality on each Party under this Agreement and obligations consistent with the intellectual
property provisions of Article 9; (b) do not under any circumstance impose any payment obligations or liability on the non-subcontracting
Party; and (c) are otherwise consistent with the terms of this Agreement. If ProQR or any of its Affiliates intends to engage a
Third Party subcontractor to perform any of its obligations under the Research Program, ProQR shall provide Lilly written notice no less
than [***] prior to engaging such Third Party subcontractor, which notice shall include the identity of such subcontractor and a high
level description of the services such subcontractor will perform related to the Research Program.

 

4.9           Lilly
Materials. In the event that it is necessary to enable execution of the Workplan, Lilly may need to transfer certain Lilly materials
to ProQR that are not otherwise delivered under a supply or other separate agreement between the Parties or their Affiliates. In each
such case, the Parties will mutually agree on the terms of such material transfer, which in any case shall be subject to the terms of
Article 9 of this Agreement. Any such materials provided to ProQR shall be accompanied by a materials transfer record substantially
in the form of Exhibit 4.9 (each a “Materials Transfer Record”). In the event of such transfer,
unless otherwise mutually agreed, Lilly shall be responsible for obtaining all necessary approvals and/or filings as required under Applicable
Laws for the exportation of any such materials to ProQR and ProQR shall be responsible for obtaining all necessary approvals and/or filings
as required under Applicable Laws for their importation and use by ProQR.

 

4.10         Technical
Transfer Assistance. Following the conclusion of each Project, ProQR shall reasonably cooperate with Lilly and provide Lilly with
such assistance as is reasonably requested by Lilly in connection with the preparation and submission by Lilly of the applicable IND
or BLA/NDA, including [***]. For avoidance of doubt, ProQR shall not be required to generate any new or additional data or information
under this Section 4.10.

 

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Article 5

 

DEVELOPMENT AND COMMERCIALIZATION

 

5.1           Lilly
Sole Right and Responsibility. Except with respect to the Research activities to be conducted by ProQR pursuant to a Workplan for
the Projects, Lilly shall be solely responsible for, and shall have the exclusive right to, Exploit Compounds and Products (including
with respect to the conduct of GLP Toxicology studies) in the Field in the Territory. Subject to the terms of this Agreement, all decisions
concerning the Exploitation of Compounds and Products following Research, including the clinical and regulatory strategy of Compounds
and Products, the Development, Manufacturing, marketing and sales of Products, and the design, price, promotion, and other Commercialization
of Products, is within the sole discretion of Lilly. Upon the achievement of Candidate Success CSFs for a given Project, the applicable
Project shall be deemed concluded and Lilly will have the sole and exclusive right (but not obligation) to pursue further Exploitation
of the Product resulting from such Project, subject to the terms and conditions set forth herein, including this Section 5.1. Lilly
shall use Commercially Reasonable Efforts to achieve a First Commercial Sale in [***] for [***] Product for each Successful Candidate.

 

5.2           Development
Reports. Lilly shall keep ProQR reasonably informed as to the progress and results of its and its Affiliates’ and Sublicensees’
Development activities under this Agreement, and shall provide ProQR with a high-level written report summarizing its Development activities
and the results thereof on at least [***] basis, indicating estimated timeframes with respect to Research and Development milestones
for the then-current Calendar Year and subsequent Calendar Year. Lilly’s obligations under this Section 5.2 shall cease with
respect to a Product at such time when Marketing Approval for such Product is obtained. Following the First Commercial Sale of a Product
with respect to a Project Target, Lilly shall provide ProQR with reports as provided in Section 8.3.7, but Lilly shall have no additional
reporting obligations with respect to its Commercialization activities, unless otherwise expressly specified in this Agreement.

 

5.3           Lilly
Regulatory Control. Except as provided under a Workplan, as between the Parties, Lilly shall have sole responsibility for and control
of the preparation, submission, and maintenance of all Regulatory Filings and obtaining Regulatory Approvals (including the preparation
and submission of the IND, BLA/NDA (and any other documents needed for clinical submissions in the European Union or otherwise), or marketing
authorization application (“MAA”) filing and for seeking IND, BLA/NDA, or MAA approval) with respect to Products,
and shall have sole control over all interactions with the applicable Regulatory Authority. ProQR shall reasonably cooperate with Lilly,
at Lilly’s reasonable request and expense, with respect to any regulatory matters related to Products. Lilly will own all right,
title and interest in and to any and all Regulatory Filings and Regulatory Approvals for Products and, as between the Parties, all such
Regulatory Filings and Regulatory Approvals will be held in the name of Lilly. ProQR shall execute all documents and take all actions
as are necessary or reasonably requested by Lilly to vest such title in Lilly.

 

5.4           Adverse
Event Reporting. Lilly shall establish, hold, and maintain the global safety database for Products with respect to information on
adverse events concerning the Products, as and to the extent required by Applicable Law. [***]

 

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Article 6

 

LICENSE RIGHTS

 

6.1           Exclusive
License Grant to Lilly. ProQR (on behalf of itself and its Affiliates) hereby grants to Lilly an exclusive (even as to ProQR and
its Affiliates, subject to the remainder of this Section 6.1), royalty-bearing (as set forth in Section 8.3), license, with
the right to grant sublicenses (through multiple tiers, as provided in Section 6.4), under the ProQR Platform IP and ProQR’s
interest in any Joint Program IP to Exploit Compounds and Products in the Field in the Territory, subject to a right retained by ProQR
to perform (i) activities to be conducted by ProQR as contemplated under this Agreement and (ii) any internal research and
development activities conducted by ProQR to improve the ProQR Platform. [***]

 

6.2           Non-Exclusive
License Grant to Lilly. ProQR (on behalf of itself and its Affiliates) hereby grants to Lilly a non-exclusive license, with the right
to grant sublicenses (through multiple tiers, as provided in Section 6.4), under any ProQR Supplemental IP to Exploit Compounds
and Products Directed to Project Targets in the Field in the Territory [***].

 

6.3           Non-Exclusive
License Grant to ProQR. Subject to the terms and conditions of this Agreement and any exclusivity granted to Lilly herein, Lilly
hereby grants to ProQR a worldwide, fully paid, royalty-free, non-sublicensable (except to Third Party subcontractors acting on its behalf,
as permitted by Section 4.8, and subject to Section 6.4), non-exclusive license under the Lilly Background IP (in each case,
excluding Lilly Program IP and any Excluded Technology), solely as and to the extent necessary (a) for ProQR or its Affiliates (or
Third Party subcontractors) to perform its obligations, including its Research activities with respect to the Project Targets, under
the Workplans during the Research Term and (b) to improve the ProQR Platform.

 

6.4           Third
Party Sublicenses. Lilly and ProQR may grant one or more sublicenses under the rights and licenses granted to it under Sections 6.1
and 6.2 (in the case of Lilly) or Section 6.3 (in the case of ProQR), in full or in part, to Third Parties (with the right to sublicense
through multiple tiers); provided, that: (a) any such permitted sublicense is consistent with and subject to the terms and conditions
of this Agreement, including the confidentiality provisions of Article 12 and the intellectual property provisions of Article 9
and the audit provisions of Section 8.5; and (b) the Party granting such sublicense shall remain responsible for performance
of such Party’s obligations under this Agreement and shall be responsible for all actions of each such sublicensee as if such sublicensee
were the Party hereunder.

 

6.5           No
Implied Rights. Except as expressly set forth in this Agreement, neither Party shall be granted, by implication or otherwise, any
license or right to or under any other Intellectual Property Right, including any trademarks, Know-How, or Patents, of the other Party.

 

6.6           Safe
Harbor Research. Except to the extent ProQR has granted exclusive rights to Lilly under this Agreement, neither Party, by entering
into this Agreement, is forfeiting any rights that such Party may have to perform research activities in compliance with 35 U.S.C. § 271(e)(1) or
any experimental or research use exemption that may apply under Applicable Law or in any country.

 

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Article 7

 

EXCLUSIVITY

 

7.1           ProQR
Exclusivity Obligations.

 

7.1.1           Activities
With Respect to Lilly Targets. Neither ProQR nor any of its Affiliates shall (by themselves, or through or with any Third Party):
(a) during the Research Term, conduct a program or efforts to discover or generate, or otherwise intentionally discover or generate,
any compounds Directed To (i) a Lilly Target or (ii) any other [***] Target; (b) thereafter through the end of the Exclusivity
Period for a Lilly Target, conduct a program or efforts to discover or generate, or otherwise intentionally discover or generate, any
compound or product Directed To that Lilly Target (including any Lilly Target that is an [***] Target); or (c) during the relevant
period identified in the foregoing clause (b), provide any license or authorization to any Third Party to permit the Third Party, or
otherwise permit or enable any Third Party, to do any of the foregoing. Solely for purposes of this Section 7.1.1, and without otherwise
modifying the definition of “Lilly Target” for any other purpose, “Lilly Target” shall include the Target of
which such Lilly Target is a specific adenosine, the gene coding for such Target, and any protein product translated from such Target.
If ProQR or any of its Affiliates determines that it has unintentionally discovered or generated a compound Directed To a Lilly Target
as described above, during the relevant period of restriction indicated above, ProQR and its Affiliate shall (1) cease any and all
development activities with respect to such Compound, (2) not permit or enable any Third Party to conduct further development or
commercialization activities with respect to such Compound and (3) promptly notify Lilly in writing of the identity of such Compound,
and (if such unintentional discovery or generation occurs during the Research Term and the applicable Intellectual Property Rights are
Controlled by ProQR) Lilly shall have the option to include the Compound in the Research Program (and to Exploit such Compound); provided,
however, that such option right shall not include rights to any Compounds that are intended to treat ocular diseases or conditions.

 

7.1.2           ProQR
Exploitation Activities With Respect to Certain Compounds. Neither ProQR nor any of its Affiliates shall, themselves, nor shall any
of them, enable any Third Party to, during the Research Term or thereafter during the Exclusivity Period, directly or indirectly Exploit
any Compound that is the subject of an active Project, any Hit or any Successful Candidate. If ProQR or any of its Affiliates determines
that it has unintentionally Developed a compound Directed to a Lilly Target as described above, during the relevant period of restriction
indicated above, ProQR and its Affiliate shall cease any and all Development activities with respect to such Compound and shall not permit
or enable any Third Party to conduct further Development activities with respect to such Compound.

 

7.1.3           The
 “Exclusivity Period” with respect to a Lilly Target shall mean a period ending on the earlier of (i) the
end of the Term and (ii) the end of the period during which Lilly is using Commercially Reasonable Efforts to Research, Develop,
Manufacture or Commercialize Compounds or Products Directed To that Lilly Target (which includes at least the duration of the Research
Term) and a period of [***] thereafter. For avoidance of doubt, the “Exclusivity Period” (a) [***] and
(b) [***].

 

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7.2           Transactions
Involving Competing Programs.

 

7.2.1           Acquisition
of Existing Competing Program. [***]

 

7.2.2           Existing
Competing Program of a ProQR Acquirer. [***]

 

Article 8

 

FEES, ROYALTIES, & PAYMENTS

 

8.1           Upfront
Payment. As partial consideration for the rights granted by ProQR to Lilly pursuant to the terms of this Agreement, Lilly shall pay
to ProQR a one-time, non-refundable, non-creditable payment equal to Twenty Million Dollars ($20,000,000) within [***] following the
Effective Date.

 

8.2           Milestone
Events and Payments.

 

8.2.1           Milestone
Events and Milestone Payments. On a Project Target-by-Project Target basis, Lilly shall pay to ProQR certain milestone payments,
as follows: (a) within [***] following any Compound Directed To a given Project Target achieving a discovery and development milestone
event set forth in Table 8.2 below (each, a “Development Milestone Event”), Lilly shall pay to ProQR the corresponding
Milestone Payment indicated in Table 8.2 (each such Milestone Payment, a “Development Milestone Payment”);
and (b) within [***] following the end of the Calendar Quarter in which any commercial milestone event set forth in Table 8.2 (each,
a “Commercial Milestone Event”) is achieved, Lilly shall pay to ProQR the corresponding Milestone Payment indicated
in Table 8.2 (each such Milestone Payment, a “Commercial Milestone Payment”). The Development Milestone Events
and Commercial Milestone Events may be referred to individually or collectively as “Milestone Events,” and
Development Milestone Payments and Commercial Milestone Payments may be referred to individually or collectively as “Milestone
Payments.” In the event that a given Development Milestone Event or Commercial Milestone Event is achieved in respect
of a Project Target before payment by Lilly of all earlier Development Milestone Payments or Commercial Milestone Payments (as applicable)
in respect of such Project Target, all such earlier payments shall also become due. In the event that First Commercial Sale is achieved
in respect of a Project Target before payment by Lilly of all Development Milestone Payments in respect of such Project Target, all Development
Milestone Payments shall also become due in respect of such Project Target.

 

8.2.2           Limitations
per Project Target. Each Milestone Payment shall be payable only once per Project Target (with respect to the Development Milestone
Events, for the first Hit, Successful Candidate or Product Directed To such Project Target to achieve such Milestone Event), and no Milestone
Payment shall be payable for subsequent or repeated achievements of the same Milestone Event with respect to the same Project Target.

 

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8.2.3           Step-Down
of Thresholds for Orphan Indications. If a Product is approved solely for an Orphan Indication (and no other indication) then the
Net Sales thresholds in the Commercial Milestone portion of Table 8.2 will be deemed reduced by [***] percent ([***]%) for such Product
for so long as the Product is solely approved for an Orphan Indication. So, for example, if a Product is only approved for an Orphan
Indication and achieves Calendar Year Net Sales of [***] (and no other Product Directed To the same Project Target has achieved Calendar
Year Net Sales of [***]) then the first Commercial Milestone Event will be deemed achieved by such Product. However, if, prior to achieving
the next Commercial Milestone Event, such Product is subsequently approved for an indication that is not an Orphan Indication then it
will not be deemed to have achieved the next Commercial Milestone Event until it has Calendar Year Net Sales of at least [***].

 

Table 8.2–
Milestone Payments

 

	Development
    Milestone Event	Milestone
    Payment
	[***]	[***]
	[***]	[***]
	[***]	[***]
	[***]	[***]
	[***]	[***]
	[***]	[***]
	[***]	[***]
	[***]	[***]
	Commercial
    Milestone Event	Milestone
    Payment
	[***]	[***]
	[***]	[***]
	[***]	[***]
	[***]	[***]
	[***]	[***]

 

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8.3           Royalties
on Products.

 

8.3.1           Royalty
Term. Lilly shall pay ProQR royalties as set forth in this Section 8.3 on a country-by-country and Product-by-Product basis
in the Territory during the period of time beginning on the date of the First Commercial Sale of such Product in such country and continuing
until the latest to occur of: (a) the expiration or abandonment of the last-to-expire Valid Claim in such country Covering such
Product; (b) the expiration of all data and regulatory exclusivity periods for such Product in such country; and (c) [***]
years after the First Commercial Sale of such Product in such country (the “Royalty Term”). Upon the expiration
of the Royalty Term for a Product in a particular country, the license granted by ProQR to Lilly under Sections 6.1 and 6.2 with
respect to such Product and such country shall survive and become perpetual, fully-paid, and royalty-free, and shall (with respect to
the license under Section 6.1) remain exclusive (even as to ProQR and its Affiliates).

 

8.3.2           Royalty
Rates. On a Product-by-Product and country-by-country basis, during the Royalty Term, Lilly shall pay to ProQR a tiered royalty equal
to the percentages of annual Net Sales of such Product in such countries, as set forth in Table 8.3 below (the “Royalty”),
calculated by [***]. For clarity, the Royalty rates set forth below are intended to be tiered and incremental, and the higher incremental
rate will only apply to that portion of the Net Sales of royalty-bearing Products that fall within the indicated range of sales.

 

Notwithstanding the Royalty
rates set forth in Table 8.3 below, all Royalties payable pursuant to this Section 8.3 are subject to reduction as further
described in Sections 8.3.3 - 8.3.5 below or as expressly stated elsewhere in this Agreement.

 

Table 8.3–
Royalty Rates

 

	Annual Net Sales of the
    Applicable Product	Royalty
    Rate
	[***]	[***]
	[***]	[***]
	[***]	[***]
	[***]	[***]

 

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8.3.3           Third
Party Payments. On a Product-by-Product and country-by-country basis, Lilly may deduct from any Royalty payments to ProQR under this
Section 8.3 with respect to the sale of a given Product in a given country, an amount equal to [***] percent ([***] %) of any payments
made by Lilly to a Third Party (including a Third Party collaborating with Lilly on complementary technology) in consideration for a
right or license under such Third Party’s Intellectual Property Rights which is necessary or Materially Useful to Exploit such
Product; provided, that in no event will the Royalty payments payable to ProQR under this Section 8.3 for each Product be
reduced, solely as a result of this Section 8.3.3, by more than [***] percent ([***] %). For purposes of this Section 8.3.1,
 “Materially Useful” means that applying a Third Party Patent will result in a material advantage to the Product,
including an advantage to the safety or efficacy of the Product or reduction in the cost of Developing or Manufacturing the Product.

 

8.3.4           Valid
Claim. In any [***] during the Royalty Term for a Product for which there is no longer a Valid Claim that Covers such Product in
a country, the Royalty rates provided in Table 8.3 above for the Product will be reduced in such country by [***] percent ([***] %) for
such [***] (in addition to any reductions in Section 8.3.3 and Section 8.3.5) and thereafter for the remainder of the Royalty
Term.

 

8.3.5           Generic
Equivalents. On a country-by-country and Product-by-Product basis, commencing on the first [***] in which there is Loss of Market
Exclusivity, the Royalty rates provided in Table 8.3 above for the Product will be permanently reduced in such country by [***] percent
([***]%).

 

8.3.6           Cumulative
Royalty Reductions and Limitations. Each of the potential Royalty reductions in the foregoing Sections 8.3.3, 8.3.4 and 8.3.5 may
be taken in addition to, and not in lieu of, the potential reductions in the other such Section; provided, that in no circumstances will
the Royalties payable to ProQR under this Section 8.3 in any [***] for a Product in a given country be reduced, as a result of Sections
8.3.3, 8.3.4 and 8.3.5 in the aggregate, below [***] percent ([***]%) of the royalties otherwise payable under this Section 8.3.

 

8.3.7           Payment;
Reports. Royalty payments due by Lilly to ProQR under this Section 8.3 will be calculated and reported for each [***]. All Royalty
payments due under this Section 8.3 shall be paid within [***] after the end of each [***] and shall be accompanied by a report
setting forth, with respect to each [***], on a Product-by-Product and country-by-country basis: (a) Net Sales of the Product by
Lilly and its Affiliates and Sublicensees in such country, (b) a calculation of the Royalties due on such Net Sales, and (c) a
statement of the amount of Third Party payments deducted under Section 8.3.3.

 

8.4           Method
of Payment; Currency Conversion. Unless otherwise agreed by the Parties, all payments due under this Agreement shall be paid in Dollars
by wire transfer or electronic funds transfer of immediately available funds to an account designated by the payee; provided, however,
that Lilly shall only be required to disburse funds to the payee’s jurisdiction of incorporation or to a jurisdiction in which
the payee has a significant business presence. When conversion of payments from any currency other than Dollars is required, Lilly’s
then-current standard exchange rate methodology will be employed for the translation of foreign currency sales into Dollars; provided,
that this methodology is used by Lilly in the translation of its foreign currency operating results, is consistent with U.S. GAAP, is
audited by Lilly’s independent certified public accountants in connection with the audit of the consolidated financial statements
of Lilly, and is used for external reporting of foreign currency operating results.

 

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8.5           Records
and Audits. Lilly (including its Affiliates and Sublicensees) shall keep complete and accurate books and records which may be necessary
to ascertain properly and to verify the royalty payments due hereunder. Such records shall be kept for such period of time required by
Applicable Laws, but no less than [***] following the end of the Calendar Year to which they pertain. Within the Term, ProQR shall not
more than [***] each Calendar Year have the right to have a Big 4 independent, certified public accountant (e.g., Deloitte, KPMG,
PricewaterhouseCoopers, Ernst & Young) inspect Lilly’s records for the purpose of determining the accuracy of royalty
payments. No period will be audited more than once. The independent, certified public accountant shall keep confidential any information
obtained during such inspection and shall report to ProQR and Lilly only the amounts of Net Sales and royalties due and payable. Such
audits may be exercised during normal business hours upon reasonable prior written notice to Lilly. ProQR shall bear the full cost of
such audit unless such audit discloses an underpayment by Lilly of more than [***] percent ([***]%) of the amount of royalties or other
payments due under this Agreement for the audited period, and which underpayment is also at least [***] ($[***]), in which case, Lilly
shall bear the cost of such audit. Lilly shall remit to ProQR the amount of any underpayment, plus interest (at the rate of prime, consistent
with Section 8.6), within [***] of the date the auditor’s written report is received. Any overpayment by Lilly revealed by
an audit shall be refunded by ProQR at the request of Lilly within [***] of the receipt of the request, with interest (at the rate of
prime, consistent with Section 8.6).

 

8.6           Late
Payments. If any payment properly due under this Agreement and not subject to a good faith dispute is not paid when due in accordance
with the applicable provisions of this Agreement, the payment shall accrue interest from the date due at the rate [***] (as reported
in The Wall Street Journal (Eastern U.S. edition)) or the maximum rate allowable by Applicable Law, whichever is less. The payment
of such interest shall not limit the Party entitled to receive payment from exercising any other rights it may have as a consequence
of the lateness of any payment.

 

8.7           Taxes.

 

8.7.1           Cooperation
and Coordination. The Parties acknowledge and agree that it is their mutual objective and intent to minimize, to the extent feasible
and in compliance with Applicable Laws, taxes payable with respect to their collaborative efforts under this Agreement and that they
shall use reasonable efforts to cooperate and coordinate with each other to achieve such objective, including by completing and filing
documents required or permitted under the provisions of any Applicable Laws in connection with a claim of exemption from, or entitlement
to a reduced rate of, withholding taxes or in connection with any claim to a refund of or credit for any payment of such taxes. Notwithstanding
the foregoing, for clarity, it is ProQR’s sole responsibility to prepare and file required documents necessary to claim an exemption
from withholding tax or to claim a reduced rate of withholding tax, at ProQR’s sole expense.

 

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8.7.2           Payment
of Tax. The upfront, milestones, royalties and other amounts payable by Lilly to ProQR to this Agreement (each, a “Payment”)
shall be paid free and clear of any and all taxes, except for any withholding taxes required by Applicable Law. Except as provided in
this Section 8.7, ProQR shall be solely responsible for paying any and all taxes (other than withholding taxes required by Applicable
Law to be deducted from Payments and remitted by Lilly) levied on account of, or measured in whole or in part by reference to, any Payments
it receives. Lilly shall deduct or withhold from the Payments any taxes that it is required by Applicable Law to deduct or withhold.
Notwithstanding the foregoing, if ProQR is entitled under any applicable tax treaty to a reduction in the rate of, or the elimination
of, any applicable withholding tax, it may deliver to Lilly or the appropriate Governmental Authority (with the assistance of Lilly to
the extent that this is reasonably required and is expressly requested in writing) the prescribed forms necessary to reduce the applicable
rate of withholding or to relieve Lilly of its obligation to withhold such tax and Lilly shall apply the reduced rate of withholding
or dispense with withholding as the case may be; provided that Lilly has received ProQR’s delivery of all applicable forms in a
form satisfactory to Lilly (and, if necessary, evidence, in a form satisfactory to Lilly, of ProQR’s receipt of appropriate governmental
authorization) at least [***] prior to the time Payments are due. If in accordance with the foregoing, Lilly withholds any amounts of
tax, it shall pay to ProQR the balance when due, make timely payment to the proper tax authority of the withheld amount and send to ProQR
proof of such payment and applicable tax withholding certificates within [***] following such payments.

 

Article 9

 

INTELLECTUAL PROPERTY

 

9.1           Inventorship.
Inventorship as between the Parties will be determined in accordance with U.S. patent laws. All such determinations shall be documented
to ensure that the Patent claims in any divisional or continuation patent applications reflect appropriate inventorship.

 

9.2           Ownership
of Intellectual Property Rights.

 

9.2.1           Background
IP. As between the Parties, and subject to the licenses granted under this Agreement (a) Lilly shall solely own (or retain ownership
of) all rights, title and interests in and to the Lilly Background IP, and (b) ProQR shall solely own (or retain ownership of) all
rights, title and interests in and to the ProQR Background IP. If any Third Party becomes an Acquirer of ProQR after the Effective Date
pursuant to a Change of Control, any Intellectual Property Rights Controlled by the Acquirer before the relevant Change of Control transaction
or thereafter during the Term will not be considered part of the ProQR Background IP; provided, however, that any Intellectual Property
Rights that would otherwise constitute ProQR Background IP and are discovered, created or acquired by or on behalf of the Acquirer after
the relevant Change of Control transaction by using any ProQR Technology relating to the Research Program will be considered part of
the ProQR Background IP.

 

9.2.2           Program
IP. Ownership of Inventions arising, discovered, created, acquired, conceived or reduced to practice, by or on behalf of either Party
(or any of their Affiliates) in the course of the Research Program or otherwise in the course of performing activities under this Agreement
(“Program IP”) shall be as follows:

 

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(a)           Except
as otherwise provided in Section 9.2.2(b)–(d), all Inventions arising, discovered, created, acquired, conceived or reduced
to practice, by or on behalf of either Party (or any of their Affiliates) in the course of the Research Program, or otherwise in the
course of performing activities under this Agreement, shall be owned by the inventors as determined in accordance with inventorship rules under
U.S. patent law.

 

(b)           Lilly
shall solely own (or retain ownership of) all Program IP that is either (i) invented solely be Lilly (or any of its Affiliates),
or (ii) solely constituting an Improvement to the Lilly Background IP, in each case, except to the extent constituting Joint Program
IP (collectively, “Lilly Program IP”).

 

(c)           ProQR
shall solely own (or retain ownership of) all Program IP that is either (i) invented solely be ProQR (or any of its Affiliates),
or (ii) solely constituting an Improvement to the ProQR Background IP (which for purposes of this Agreement shall include any Inventions
related to targeted RNA editing the use of which is not confined to a particular Target), in each case, except to the extent constituting
Joint Program IP (collectively, “ProQR Program IP”).

 

(d)           Lilly
and ProQR shall jointly own all Program IP that (i) constitutes an Improvement to both the Lilly Background IP and the ProQR Background
IP, (ii) Covers a Compound, and (iii) is neither Lilly Program IP nor ProQR Program IP (collectively, “Joint Program
IP”).

 

9.3           Assignments
of Intellectual Property Rights.

 

9.3.1           Inventor
Assignment Obligation. Each Party shall cause all of its Affiliates, employees, agents, independent contractors, consultants, and
others who perform activities for such Party under this Agreement to be under an obligation to assign (or, if such Party is unable to
cause such person or entity to agree to such assignment obligation despite such Party using reasonable efforts to negotiate such assignment
obligation, provide a license, preferably exclusive, under) to such Party their rights in and to any Inventions created, conceived of,
reduced to practice, or acquired in the course or scope of the Research Program or otherwise related to this Agreement and all Intellectual
Property Rights therein, except where Applicable Law requires otherwise and except in the case of governmental, not-for-profit and public
institutions that have standard policies against such an assignment (in which case a Party shall obtain a suitable license, preferably
exclusive, or right to obtain such a license). Each Party shall use reasonable efforts to promptly disclose to the other Party in writing
all Inventions arising in the course or scope of the Research Program or otherwise under this Agreement, including any invention disclosures,
or other similar documents, submitted to it by its employees, agents or independent contractors describing such Inventions, and all information
relating to such Inventions to the extent necessary or useful for the preparation, filing and maintenance of any Patent with respect
to such Invention.

 

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9.3.2           Lilly
Assignment of ProQR Program IP. Lilly shall and hereby does assign to ProQR all of Lilly’s and its Affiliates’ right,
title, and interest in and to all ProQR Program IP made by or on behalf of Lilly; provided, that Lilly shall retain, and ProQR hereby
grants to Lilly, the right to use such ProQR Program IP solely for Lilly’s (or any of its Affiliates’) internal Research
and Development purposes. Lilly shall take (and cause its employees, agents, contractors and Sublicensees (if applicable) to take) such
further actions reasonably requested by ProQR to evidence such assignment and to support ProQR’s efforts to Patent or obtain other
Intellectual Property Rights protection for such ProQR Program IP.

 

9.3.3           ProQR
Assignment of Lilly Program IP. ProQR shall and hereby does assign to Lilly all of ProQR’s and its Affiliates’ right,
title, and interest in and to all Lilly Program IP made by or on behalf of ProQR. ProQR shall take (and cause its employees, agents,
contractors and Sublicensees (if applicable) to take) such further actions reasonably requested by Lilly to evidence such assignment
and to support Lilly’s efforts to Patent or obtain other Intellectual Property Rights protection for such Lilly Program IP.

 

9.3.4           Assignment
of Joint Program IP. (a) ProQR (on behalf of itself and its Affiliates) shall and does hereby assign to Lilly and (b) Lilly
(on behalf of itself and its Affiliates) shall and does hereby assign to ProQR, an equal, undivided interest in and to the Joint Program
IP. Subject to the licenses and obligations of exclusivity granted under this Agreement, each Party shall have full rights to Exploit
and license Joint Program IP (and any Patent rights therein), without any obligation or requirement of an accounting to the other Party;
provided that, in Exploiting or licensing such Joint Program IP, neither Party shall have any right or license to the underlying Background
IP of the other Party, in each case, not otherwise expressly granted elsewhere in this Agreement.

 

9.4           Independent
Development. Subject to the licenses and obligations of exclusivity granted hereunder, nothing in this Agreement shall be construed
as limiting either Lilly’s or ProQR’s right to research, develop, improve and in-license technology related to the Lilly
Background IP (in the case of Lilly) or ProQR Background IP (in the case of ProQR) outside the scope of this Agreement in its ordinary
course of business.

 

9.5           Enabling
Technology. [***].

 

9.6           Contribution
of Licensed ProQR Technology. ProQR shall inform Lilly in writing, prior to contributing to any Research to be conducted under any
Workplan any portion of the ProQR Technology that is in-licensed from a Third Party, the contribution of which would prevent or conflict
with the ownership and use rights with respect to Patents and Know-How contemplated by this Agreement.

 

9.7           Patent
Prosecution and Maintenance.

 

9.7.1           Rights
to Prosecute and Maintain Generally. Subject to Sections 9.7.2, 9.7.3 and 9.7.4, each Party shall control the Prosecution and Maintenance
of Patents claiming Inventions that such Party Controls (other than Control obtained pursuant to a grant of rights under this Agreement).

 

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9.7.2           Product-Specific
Patents. As between the Parties, Lilly shall have the first right, but not the obligation, to Prosecute and Maintain any Product-Specific
Patents at Lilly’s sole cost and expense. Lilly shall keep ProQR reasonably informed of the status of all Product-Specific Patents
and shall promptly provide ProQR with all material correspondence received from any patent authority in connection therewith. In addition,
Lilly shall promptly provide ProQR with drafts of all proposed material filings and correspondence to any patent authority with respect
to any Product-Specific Patents for ProQR’s review and comment prior to the submission of such proposed filings and correspondences,
and Lilly shall consider ProQR’s reasonable comments in good faith. Lilly shall notify ProQR of its intention to suspend or cease
any Prosecution and Maintenance of any Product-Specific Patent. Lilly shall provide such notice at least [***] prior to any filing or
payment due date, or any other due date that requires action, in connection with such Product-Specific Patent. In such event, Lilly shall
permit ProQR, at ProQR’s discretion and at its sole expense, to continue Prosecution and Maintenance of such Product-Specific Patent,
subject to the foregoing information sharing obligation and review and comment rights applied mutatis mutandis; provided that
ProQR shall not continue such Prosecution and Maintenance if Lilly objects to such Prosecution or Maintenance consistent with Lilly’s
strategic decision-making rights. The Parties shall in good faith cooperate through the JPC (or as otherwise agreed to by the Parties)
to communicate and determine (and, in any event, ProQR shall notify Lilly, including through the JPC) which future ProQR Patents may
reasonably become Product-Specific Patents, and any such ProQR Patents shall be subject to the foregoing information sharing obligation
and review and comment rights applied mutatis mutandis, as well as Section 9.7.5. Lilly shall have no obligation to defend
against any patent oppositions existing or filed as of the Effective Date, including those listed on Exhibit 10.2, regardless
of whether or not any of the patents involved in the oppositions could be considered Product-Specific Patents. Further, Lilly’s
election not to defend against patent oppositions existing as of the Effective Date shall not prevent Lilly from exercising any of the
remaining rights to Prosecute and Maintain any Product-Specific Patent that it wishes to control in accordance with this Agreement.

 

9.7.3           ProQR
Platform Patents. As between the Parties, subject to Section 9.7.2, ProQR shall have the sole responsibility for and control
over the Prosecution and Maintenance of any ProQR Platform Patents and ProQR Improvements IP, at ProQR’s sole cost and expense.

 

9.7.4           Joint
Program Patents. The JPC shall determine which Party has the first right, but not the obligation, to Prosecute and Maintain the Patents
(other than Product-Specific Patents) in the Joint Program IP (“Joint Patents”). The Party handling the Prosecution
and Maintenance of a given Joint Patent (the “Prosecuting Party”) shall keep the other Party reasonably informed
of the status of such Joint Patents and shall promptly provide such other Party with all material correspondence received from any patent
authority in connection therewith. In addition, the Prosecuting Party shall promptly provide the other Party with drafts of all proposed
material filings and correspondence to any patent authority for such other Party’s review and comment prior to the submission of
such proposed filings and correspondences, and the Prosecuting Party shall consider the other Party’s reasonable comments in good
faith. The Prosecuting Party shall notify the other Party of its intention to suspend or cease any Prosecution and Maintenance of any
Joint Patent at least [***] prior to any filing or payment due date, or any other due date that requires action, in connection with such
Joint Patent. In such event, the Prosecuting Party shall permit the other Party, at its discretion and at its sole expense, to continue
Prosecution and Maintenance of such Joint Patent.

 

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9.7.5           Separation
of Patent Claims. If ProQR desires to file an application for a Patent or to maintain a Patent (including such Patents filed as continuations
or divisionals after the Effective Date from Patents listed in Exhibit 1.115), in accordance with the rights granted to ProQR
under this Article 9 and such Patent would include the Specific Identification of a Project Target, Compound or Product (i.e.,
for purposes of pursuing Product-Specific Patent protection), the Parties agree that, to the extent practicable, such application or
Patent shall be divided into two (2) or more Patent applications or Patents, so that only one such application or Patent Specifically
Identifies a Project Target, Compound or Product (i.e., a Product-Specific Patent) and the other such application or Patent falls
generally within the ProQR Patents, and does not Specifically Identify a Project Target, Compound or Product. If such division or continuation
is not practicable, such Patent application, or Patent, shall be deemed to be within the Product-Specific Patents. If such division or
continuation is practicable, then the Parties shall, in good faith, coordinate filings as appropriate on the same day.

 

9.7.6           Cooperation
of the Parties. Each Party shall cooperate fully with the other Party, through the JPC, in the Prosecution and Maintenance of Patents
under this Section 9.7 at its own cost (except as expressly set forth otherwise in this Article 9), including by: (a) executing
all papers and instruments, or requiring its employees or contractors, to execute such papers and instruments, to enable the other Party
to apply for and to Prosecute and Maintain such Patents in any country as permitted by this Section 9.7; and (b) promptly informing
the other Party of any matters coming to such Party’s attention that may affect the Prosecution and Maintenance of any such Patents.
Each Party will use reasonable efforts via good faith consultation to avoid creating potential issues in Prosecution and Maintenance
of Patents under this Section 9.7.

 

9.8           Infringement
or Misappropriation by Third Parties.

 

9.8.1           Notice.
Each Party shall notify the other within [***] of becoming aware of any alleged or threatened infringement by a Third Party of any of
the ProQR Patents, Lilly Patents or Program IP, which infringing activity involves the using, making, importing, offering for sale or
selling a Compound or Product, in each case in the Field in the Territory, and any related declaratory judgment, opposition or similar
action alleging the invalidity, unenforceability or non-infringement of any of the ProQR Patents, Lilly Patents or Program IP (collectively
 “Infringement”).

 

9.8.2           Generally.
Subject to Sections 9.8.3, 9.8.4 and 9.8.5, each Party shall have the sole right to bring and control any legal action in connection
with any Infringement of Patents claiming Inventions that such Party Controls (other than Control obtained pursuant to a grant of rights
under this Agreement).

 

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9.8.3           Product-Specific
Patents. As between the Parties, Lilly shall have the first right to bring and control any legal action in connection with any Infringement
of any Product-Specific Patents at its own expense. Lilly shall keep ProQR reasonably informed of the status of such enforcement efforts
for such Product-Specific Patents and shall consider in good faith ProQR’s comments thereon. ProQR may, at its own expense, be
represented in any such action by counsel of its own choice. If Lilly does not bring such legal action within [***] after the notice
provided pursuant to Section 9.8.1, ProQR may bring and control any legal action in connection with such Infringement of such Product-Specific
Patent at its own expense as it reasonably determines appropriate so long as Lilly does not reasonably object to such action consistent
with Lilly’s strategic decision-making rights.

 

9.8.4           ProQR
Platform Patents. As between the Parties, ProQR shall have the sole right to bring and control any legal action in connection with
any Infringement of the ProQR Platform Patents at its own expense. ProQR shall keep Lilly reasonably informed of the status of the applicable
ProQR Platform Patents.

 

9.8.5           Joint
Patents. Subject to Section 9.8.3 and Section 9.8.4, the JPC shall determine which Party has the first right to bring and
control any legal action in connection with any Infringement of any Joint Patents at its own expense as it reasonably determines appropriate.
The enforcing Party shall keep the other Party reasonably informed of the status of such enforcement efforts for the Joint Patents and
shall consider in good faith such other Party’s comments thereon. The enforcing Party shall provide the other Party with drafts
of all material papers to be filed with the court and shall in good faith incorporate all reasonable comments thereto by such other Party
before filing such papers. The other Party may, at its own expense, be represented in any such action by counsel of its own choice. If
the enforcing Party does not bring such legal action within [***] after the notice provided pursuant to Section 9.8.1, the other
Party may bring and control any legal action in connection with such Infringement of any Joint Patents at its own expense as it reasonably
determines appropriate so long as the enforcing Party does not reasonably object to such action.

 

9.8.6           Allocation
of Recoveries. Any recoveries resulting from an enforcement action relating to a claim of Infringement shall be first applied against
payment of each Party’s costs and expenses in connection therewith. The enforcing Party will retain any such recoveries in excess
of such costs and expenses; provided, that if Lilly is the enforcing Party with respect to patents within the ProQR Technology that cover
a Compound or Product, then such excess recoveries shall be [***].

 

9.8.7           Cooperation.
At the request and expense of the Party bringing an action under this Section 9.8, the other Party shall provide reasonable assistance
in connection therewith, including by executing reasonably appropriate documents, cooperating in discovery and joining as a party to
the action if required by Applicable Law to pursue such action. In connection with any such enforcement action, the Party bringing the
action shall not enter into any settlement admitting the invalidity or non-infringement of, or otherwise impairing the other Party’s
rights in the applicable Patents without the prior written consent of the other Party.

 

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9.9           Defense
and Settlement of Third Party Claims. Each Party shall promptly notify the other in writing of: (a) any allegation by a Third
Party that the activity of either of the Parties pursuant to this Agreement infringes or may infringe the Intellectual Property Rights
of such Third Party; or (b) any declaratory judgment action that is brought naming either Party as a defendant and alleging invalidity
of any of the Lilly Patents, Joint Patents or ProQR Patents. ProQR has the sole right to control any defense of any such claim described
in (a) involving alleged infringement of Third Party rights by ProQR’s activities at its own expense and by counsel of its
own choice, and Lilly may, at its own expense, be represented in any such action by counsel of its own choice. Lilly has the sole right
to control any defense of any such claim described in (a) involving alleged infringement of Third Party rights by Lilly’s
activities or related to the Joint Patents at its own expense and by counsel of its own choice, and ProQR may, at its own expense, be
represented in any such action by counsel of its own choice. Neither Party may settle any patent infringement litigation under this Section 9.9
in a manner that admits the invalidity or unenforceability of the other Party’s Patents or imposes on the other Party restrictions
or obligations or other liabilities, without the written consent of such other Party, which consent shall not be unreasonably withheld,
conditioned, or delayed. Nothing in this Section 9.9 will limit any indemnification rights or obligations of a Party under Article 11.

 

9.10         Patent
Extension. The Parties shall cooperate in determining which Patent claiming or Covering a Product should be extended, and thereafter
the Parties shall cooperate in obtaining patent term restorations, supplemental protection certificates or their equivalents, and other
forms of patent term extensions for a given Product with respect to any applicable ProQR Patent or Lilly Patent in any country or region
where applicable. [***] shall have final decision-making authority with respect to decisions regarding patent term extensions with respect
to any Product-Specific Patent and any Joint Patent during the Term.

 

9.11         CREATE
Act. It is the Parties’ intention that this Agreement is a “joint research agreement” as that phrase is defined
in 35 U.S.C. § 102(c) as amended by the Cooperative Research and Technology Enhancement (CREATE) Act, including the provisions
of 35 U.S.C. § 102(b)(2)(c). The Parties agree to cooperate and to take reasonable actions to maximize the protections available
for the Compounds and Products under such safe harbor provisions.

 

9.12         Trademarks.
Lilly shall have the right to select, and will be free, in its sole discretion, to use and to register in any trademark office in the
Territory, any trademark for use with a Product (the “Product Trademarks”). As between the Parties, Lilly shall
own all right, title and interest in and to any such Product Trademarks adopted by Lilly for use with Product, and is responsible for
the registration, filing, maintenance and enforcement thereof.

 

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Article 10

 

REPRESENTATIONS, WARRANTIES AND COVENANTS

 

10.1         Mutual
Representations and Warranties. Each of Lilly and ProQR represent and warrant, as of the Effective Date, that:

 

10.1.1           it
is duly organized and validly existing under the Applicable Laws of the jurisdiction of its incorporation or formation, as applicable,
has full corporate, limited liability company or other power and authority, as applicable, to enter into this Agreement and to carry
out the provisions hereof, and has sufficient facilities, experienced personnel or other capabilities (including via Affiliates and/or
Third Parties) to enable it to perform its obligations under this Agreement;

 

10.1.2           it
is duly authorized to execute and deliver this Agreement and to perform its obligations hereunder, and the individual executing this
Agreement on its behalf has been duly authorized to do so by all requisite corporate, limited liability company or other action, as applicable;
and

 

10.1.3           this
Agreement is legally binding upon it and enforceable in accordance with its terms (except as the enforceability thereof may be limited
by bankruptcy, bank moratorium or similar laws affecting creditors’ rights generally and laws restricting the availability of equitable
remedies and may be subject to general principles of equity whether or not such enforceability is considered in a proceeding at law or
in equity) and the execution, delivery and performance of this Agreement by it have been duly authorized by all necessary corporate action
and do not and will not: (a) conflict with, or constitute a default or result in a breach under, any agreement, instrument or understanding,
oral or written, to which it is a party or by which it may be bound, or violate any Applicable Law; or (b) require any consent or
approval of its stockholders.

 

10.2         ProQR
Representations and Warranties. ProQR represents, warrants and covenants to Lilly that, as of the Effective Date:

 

10.2.1           No
Unavailable Targets. As of the Effective Date, no Reserved Target is an Unavailable Target, and the Initial Target is not subject
to an executed agreement between ProQR and a Third Party (or ProQR’s commitment to negotiate an agreement with a Third Party) that
would prevent, or conflict with, the inclusion of the Target as a Project Target (and the associated Target being a Lilly Target) under
this Agreement on an exclusive basis as set forth in Section 6.1 and Section 7.1.

 

10.2.2           No
Grants that Conflict with this Agreement. ProQR and its Affiliates have not granted, and will not grant during the Term, any rights
(or other encumbrances) to any Third Party under ProQR Technology that prevent or conflict with the rights granted to Lilly hereunder.

 

10.2.3           Control
over Know-How and Patents. ProQR has Control over all Know-How and Patent rights owned by it or its Affiliates that are necessary
or reasonably useful for the Exploitation of Compounds or Products in the Field, as known to be contemplated by this Agreement.

 

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10.2.4           Existing
Patents.

 

(a)           All
Patent rights contained in the ProQR Technology existing as of the Effective Date that are issued or subject to a pending application
for issuance are listed on Exhibit 10.2.4 (the “Existing Patents”).

 

(b)           All
Existing Patents: (i) to the extent issued and subsisting are, to ProQR’s knowledge, not invalid or unenforceable, in whole
or in part, and confer a valid right to claim priority thereto; (ii) are solely and exclusively owned by, or exclusively licensed
to ProQR, free of any encumbrance, lien or claim of ownership by any Third Party; (iii) are, to the extent subject to a pending
application for issuance, being diligently prosecuted in good faith in the respective patent offices in which such applications have
been filed in accordance with Applicable Law and, to ProQR’s knowledge, all material references, documents and information have
been presented to the relevant patent office in respect of such Existing Patents to the extent required by such patent office; and (iv) were
filed and are being maintained in accordance with applicable Patent office rules, and all applicable fees applicable thereto have been
paid on or before any final due date for payment.

 

(c)           Neither
ProQR nor any of its Affiliates have taken any action that would render unpatentable (including by means of the “on-sale bar”
doctrine or prior publication) any invention claimed in the Existing Patents.

 

(d)           The
Existing Patents represent all ProQR Patents that relate to the ProQR Platform or the exploitation thereof contemplated under this Agreement.

 

(e)           To
ProQR’s knowledge, other than the rights granted under this Agreement, no rights or licenses are required under any Third Party
Patent rights not Controlled by ProQR to practice the ProQR Technology as contemplated in the Workplan as of the Effective Date, or to
Exploit the Products as contemplated herein solely by reason of the incorporation of ProQR Technology in such Products.

 

10.2.5           No
Third Party Agreements. Except as set forth on Exhibit 10.2, there are no license or other agreements with Third Parties
regarding the exploitation of any ProQR Technology or other materials contemplated to be provided by ProQR to Lilly hereunder, to which
ProQR or its Affiliate is a party that is inconsistent with or diminishes or would conflict with or prevent the rights and licenses granted
to Lilly under this Agreement.

 

10.2.6           Litigation
and Actions Relating to Intellectual Property. Except as set forth on Exhibit 10.2, ProQR: (a) has not received
any written notice of any threatened claims or litigation seeking to invalidate or otherwise challenge the ProQR Technology, including
the ProQR Patents, or ProQR’s or its Affiliates’ rights therein; and (b) is not aware of any pending or threatened action,
suit, proceeding or claim by a Third Party asserting that ProQR or any of its Affiliates is infringing or has misappropriated or otherwise
is violating any Patent right, trade secret or other proprietary right of any Third Party as would reasonably be expected to impair the
ability of ProQR to fulfill any of its obligations under this Agreement.

 

10.2.7           Other
Material Claims and Actions. There are no claims, actions, or proceedings pending or, to ProQR’s knowledge, threatened by any
Third Party; and to ProQR’s knowledge, there are no formal inquiries initiated or written notices received that may lead to the
institution of any such legal proceedings; in each case (or in aggregate) against ProQR or its properties, assets or business, which
if adversely decided, would, individually or in the aggregate, have a material adverse effect on, or prevent ProQR’s ability to
conduct the Research or to grant the licenses or rights granted to Lilly under this Agreement.

 

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10.2.8           Intercompany
Arrangements. Both entities within the definition of “ProQR” have in place present licenses and services agreements with
all of their Affiliates sufficient to ensure compliance with ProQR’s obligations hereunder. Without limiting the generality of
the foregoing, any such Affiliate that owns any right, title or interest in or to ProQR Technology has granted a present license under
such ProQR Technology to ProQR (with appropriate scope of exclusivity) so that such technology is included in the licenses granted hereunder
from the Effective Date.

 

10.2.9           Assignment
by Employees, Agents and Consultants. ProQR has obtained from each of its current employees, consultants and contractors, in each
case who perform research or development activities pursuant to this Agreement, written agreements containing obligations of confidentiality
and non-use and an assignment to ProQR of all inventions (and all of such Person’s rights thereto) for which ProQR or Lilly is
intended to have ownership or license rights under this Agreement such that no such employee, contractor or consultant shall retain any
rights to such inventions that would prevent or conflict with Lilly’s rights of ownership or use of such inventions contemplated
by this Agreement.

 

10.2.10         No
Government Funding. The inventions claimed or covered by the ProQR Patents: (a) were not conceived, discovered, developed or
otherwise made in connection with any research activities funded, in whole or in part, by, or otherwise using the resources of, any Governmental
Authority (whether of the U.S., the United Kingdom, or otherwise); (b) are not a “subject invention” as that term is
described in 35 U.S.C. Section 201(e) and (c) are not otherwise subject to the provisions of the Patent and Trademark
Law Amendments Act of 1980, as amended, codified at 35 U.S.C. §§ 200-212, as amended, as well as any regulations promulgated
pursuant thereto, including in 37 C.F.R. Part 401 (the “Bayh-Dole Act”). ProQR and its Affiliates have
complied with the applicable provisions of the Bayh-Dole Act, in a manner that protects and preserves ProQR’s right, title and
interest in such inventions to the maximum extent permitted by law.

 

10.2.11         Regulatory
Documentation. No Regulatory Documentation exists or has been generated relating to ProQR’s contemplated activities and obligations
under this Agreement. “Regulatory Documentation” means all: (a) applications (including all INDs, or equivalent,
BLA/NDAs, and applications for Regulatory Approval), registrations, licenses, authorizations and approvals (including Regulatory Approvals);
(b) correspondence and reports submitted to or received from Regulatory Authorities (including minutes and official contact reports
relating to any communications with any Regulatory Authority) and all supporting documents with respect thereto, including all adverse
event files and complaint files; (c) supplements or changes to any of the foregoing following Regulatory Approval; and (d) clinical
and other data, including Clinical Trial data, contained or relied upon in any of the foregoing; in each case ((a)–(d)) relating
to a Lilly Target and Products Directed To a Lilly Target.

 

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10.3         Mutual
Covenants.

 

10.3.1           Debarment.
Each Party represents, warrants and covenants to the other Party that neither it nor its officers, employees, agents, consultants or
any other person used by such Party in the performance of the respective research and development activities under this Agreement is:
(a) debarred or disqualified under the FD&C Act; (b) listed by any government or regulatory agencies as ineligible to participate
in any government healthcare programs or government procurement or non-procurement programs (as that term is defined in 42 U.S.C. § 1320a-7b(f)),
or excluded, debarred, suspended or otherwise made ineligible to participate in any such program; or (c) convicted of a criminal
offense related to the provision of healthcare items or services, or is subject to any such pending action. Each Party will not during
the Term knowingly, employ or use, directly or indirectly, including through Affiliates the services of any such person. In the event
that either Party becomes aware of the debarment or disqualification or threatened debarment or disqualification of any person providing
services to such Party, directly or indirectly, including through Affiliates or, in the case of Lilly, Sublicensees, which directly or
indirectly relate to activities contemplated by this Agreement, such Party shall promptly notify the other Party in writing and such
Party shall cease employing, contracting with, or retaining any such person to perform any such services.

 

10.3.2           Protection
of Information. Each Party agrees that during the Term of this Agreement, and without limiting its obligations hereunder, each Party
shall implement technical and organizational measures to protect all information under the Agreement that are appropriate and that provide
no less protection than both (i) good industry practice (i.e., in accordance with ISO 27001 and/or similar industry standards)
and (ii) such Party’s measures to protect its own information of a similar nature or importance.

 

10.4         Maintenance
of Intercompany Arrangements. ProQR shall maintain the agreements referenced in Section 10.2.8 in full force and effect throughout
the Term and shall ensure that such Affiliates remain compliant with the terms of such agreements. Should any such entity cease to be
an Affiliate of ProQR, ProQR shall ensure that such cessation has no adverse impact on ProQR’s performance hereunder or Lilly’s
rights hereunder.

 

10.5         Compliance.

 

10.5.1           Compliance
with this Agreement. Each of the Parties shall, and shall cause their respective Affiliates to, comply in all material respects with
the terms of this Agreement.

 

10.5.2           Compliance
with Applicable Laws. Each Party covenants to the other that in the performance of its obligations under this Agreement, such Party
shall comply, and shall cause its Affiliates and its and its Affiliates’ employees and contractors to comply, with all Applicable
Laws. No Party shall, or shall be required to, undertake any activity under or in connection with this Agreement which violates, or which
it believes, in good faith, may violate, any Applicable Laws.

 

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10.5.3           Compliance
with Party-Specific Regulations. In carrying out their respective obligations under this Agreement, the Parties agree to cooperate
with each other as may reasonably be required to help ensure that each is able to fully meet its obligations with respect to all judgments,
decrees, orders or similar decisions issued by any Governmental Authority specific to a Party, and all consent decrees, corporate integrity
agreements, or other agreements or undertakings of any kind by a Party with any Governmental Authority, in each case as the same may
be in effect from time to time and applicable to a Party’s activities contemplated by this Agreement (the “Party-Specific
Regulations”). Each Party shall be responsible for providing the other Party with any Party-Specific Regulations applicable
to the other Party, including any updates to such Party-Specific Regulations, and the covenant in the preceding sentence shall only apply
to the extent such Party-Specific Regulations and any updates thereto have been provided to the other Party. Neither Party shall be obligated
to pursue any course of conduct that would result in such Party being in material breach of any Party-Specific Regulation applicable
to it; provided that in the event that a Party refuses to fulfill its obligations under this Agreement in any material respect on such
basis, the other Party shall have the right to terminate this Agreement in accordance with Section 13.2; however, under such circumstances,
such termination, including the applicable effects of such termination set forth in Sections 13.5 and 13.6, shall be the sole remedy
for such terminating Party and such terminating Party shall not be entitled to any other remedy under law or equity. All Party-Specific
Regulations are binding only in accordance with their terms and only upon the Party to which they relate.

 

10.5.4           Compliance
with Internal Compliance Codes. All Internal Compliance Codes shall apply only to the Party to which they relate. The Parties agree
to cooperate with each other to help ensure that each Party is able to comply with the substance of its respective Internal Compliance
Codes and, to the extent practicable, each Party shall operate in a manner consistent with its Internal Compliance Codes applicable to
its performance under this Agreement. “Internal Compliance Codes,” as used in this Section 10.5.4, means
a Party’s internal policies and procedures intended to ensure that a Party complies with Applicable Laws, Party-Specific Regulations,
and such Party’s internal ethical, medical and similar standards.

 

10.5.5           Compliance
with Anti-Corruption Laws. In connection with this Agreement, the Parties shall comply with all applicable local, national, and international
laws, regulations, and industry codes dealing with government procurement, conflicts of interest, corruption or bribery, including, if
applicable, the U.S. Foreign Corrupt Practices Act of 1977, as amended, and any laws enacted to implement the Organisation of Economic
Cooperation and Development Convention on Combating Bribery of Foreign Officials in International Business Transactions.

 

10.5.6           Compliance
with Privacy Laws. In connection with this Agreement, Lilly and ProQR (and its and their Affiliates), and any Person acting for or
on its or their behalf, will comply with all Applicable Laws with respect to data protection and privacy laws with respect to the receipt,
collection, compilation, use, storage, processing, sharing, safeguarding, security (technical, physical and administrative), disposal,
destruction, disclosure, or transfer (including cross-border) of Personal Information, including providing any notice, obtaining any
consent or prior authorization, and conducting any assessment required under Applicable Laws.

 

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10.5.7           Prohibited
Conduct. Without limiting the other obligations of the Parties set forth in this Section 10.4, each Party covenants to the other
that, as of the Effective Date and in the performance of its obligations under this Agreement through the expiration and termination
of this Agreement, such Party and, to its knowledge, its Affiliates and its and its Affiliates’ employees and contractors, in connection
with the performance of their respective obligations under this Agreement, have not made, offered, given, promised to give, or authorized,
and will not make, offer, give, promise to give, or authorize, any bribe, kickback, payment or transfer of anything of value, directly
or indirectly through Third Parties, to any Government Official for the purpose of: (a) improperly influencing any act or decision
of the Person or Government Official; (b) inducing the Person or Government Official to do or omit to do an act in violation of
a lawful or otherwise required duty; (c) securing any improper advantage; or (d) inducing the Person or Government Official
to improperly influence the act or decision of any organization, including any government or government instrumentality, to assist any
Party in obtaining or retaining business. For the purpose of this Section 10.5.7 “Government Official”
means: (x) any officer, employee (including physicians, hospital administrators, or other healthcare professionals), agent, representative,
department, agency, de facto official, representative, corporate entity, instrumentality or subdivision of any government, military or
international organization, including any ministry or department of health or any state-owned or affiliated company or hospital; (y) any
candidate for political office, any political party or any official of a political party, in each case for the purpose of obtaining or
retaining business for or with, or directing business to, any Person, including either Party; or (z) any Person acting in an official
capacity on behalf of any of the foregoing.

 

10.6         Disclaimer.
EXCEPT AS OTHERWISE EXPRESSLY SET FORTH IN THIS Article 10, NEITHER PARTY MAKES ANY REPRESENTATIONS OR EXTENDS ANY WARRANTIES OF
ANY KIND, EITHER EXPRESS OR IMPLIED, INCLUDING WARRANTIES OF MERCHANTABILITY, QUALITY, FITNESS FOR A PARTICULAR PURPOSE, NONINFRINGEMENT,
OR VALIDITY OF PATENT CLAIMS. NOTHING IN THIS AGREEMENT SHALL BE CONSTRUED AS A REPRESENTATION MADE OR WARRANTY GIVEN BY EITHER PARTY
THAT EITHER PARTY WILL BE SUCCESSFUL IN OBTAINING ANY PATENTS OR THAT ANY PATENTS WILL ISSUE BASED ON A PENDING APPLICATION. WITHOUT
LIMITING THE RESPECTIVE RIGHTS AND OBLIGATIONS OF THE PARTIES EXPRESSLY SET FORTH HEREIN, EACH PARTY SPECIFICALLY DISCLAIMS ANY GUARANTEE
THAT ANY PROGRAM OR PRODUCTS WILL BE SUCCESSFUL, IN WHOLE OR IN PART.

 

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Article 11

 

INDEMNIFICATION

 

11.1         Indemnity.

 

11.1.1           By
ProQR. Subject to Section 11.1.3, ProQR shall defend, indemnify and hold harmless Lilly and its Affiliates, and their respective
directors, officers, employees, and agents (each, a “Lilly Indemnitee”) from and against any and all costs,
fees, expenses, losses, liabilities, and damages, including reasonable legal expenses and attorneys’ fees (collectively, “Losses”)
to which any Lilly Indemnitee may become subject as a result of (a) any claim, demand, action or other proceeding by any Third Party
(a “Claim”) to the extent such Claim and Losses arise out of: (i) the gross negligence, fraud or willful
misconduct of ProQR or its Affiliates in connection with performance of its activities under this Agreement; (ii) the breach of
this Agreement or the representations, warranties, and covenants made hereunder by ProQR; or (iii) ProQR’s, its Affiliates’
or their respective licensees’ or sublicensees’ conduct of the Research Program during the Research Term (but only to the
extent such activities directly cause such Losses and specifically excluding any product liability-related Claims and Losses); or (b) [***];
except, in each case, to the extent such Losses result from matters described in clause (a), (b), (c) or (d) of Section 11.1.2.

 

11.1.2           By
Lilly. Subject to Section 11.1.3, Lilly shall defend, indemnify and hold harmless ProQR, its Affiliates, and their respective
directors, officers, employees and agents (each, a “ProQR Indemnitee”) from and against any and all Losses
to which any ProQR Indemnitee may become subject as a result of any Claim to the extent such Claim and Losses arise out of: (a) the
gross negligence, fraud or willful misconduct of Lilly, its Affiliates, or their respective Sublicensees in connection with performance
of its or their activities under this Agreement; (b) the breach of this Agreement or the representations, warranties and covenants
made hereunder by Lilly; (c) Lilly’s, its Affiliates’ or their respective Sublicensees’ (excluding ProQR and its
Affiliates and its and their licensees and contractors) conduct of the Research Program during the Research Term, other than where such
activity is performed by ProQR or its Affiliates on behalf of Lilly, its Affiliates or their respective Sublicensees, or (d) the
Exploitation of any Product by or on behalf of Lilly, its Affiliates, or their respective Sublicensees; except, in each case, to the
extent such Losses result from matters subject to clause (a) or (b) of Section 11.1.1.

 

11.1.3           Procedure.
A Party that intends to claim indemnification under this Article 11 (the “Indemnitee”) shall promptly
notify the Indemnitor (the “Indemnitor”) in writing of any Claim in respect of which the Indemnitee intends
to claim such indemnification. The failure to deliver written notice to the Indemnitor within a reasonable time after the commencement
of any action with respect to a Claim shall only relieve the Indemnitor of its indemnification obligations under this Article 11
if and to the extent the Indemnitor is actually and materially prejudiced thereby. The Indemnitor has sole control of the defense or
settlement thereof. The Indemnitee shall cooperate fully with the Indemnitor and its legal representatives in the investigation of any
action with respect to a Claim covered by this indemnification. The Indemnitee may participate at its expense in the Indemnitor’s
defense of and settlement negotiations for any Claim with counsel of the Indemnitee’s own selection. The Indemnitor shall not settle
any Claim without the prior written consent of the Indemnitee, not to be unreasonably withheld, conditioned or delayed. So long as the
Indemnitor is actively defending the Claim in good faith, the Indemnitee shall not settle or compromise any such Claim without the prior
written consent of the Indemnitor. If the Indemnitor does not assume and conduct the defense of the Claim as provided above: (a) the
Indemnitee may defend against, consent to the entry of any judgment, or enter into any settlement with respect to such Claim in any manner
the Indemnitee may deem reasonably appropriate (and the Indemnitee need not consult with, or obtain any consent from, the Indemnitor
in connection therewith); and (b) the Indemnitor shall remain responsible to indemnify the Indemnitee as provided in this Article 11.

 

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11.2         Insurance.
During the Term, each Party shall maintain such types and amounts of liability insurance (including self-insurance) as is normal and
customary in the industry generally for similarly situated parties and adequate to cover its obligations under this Agreement, and [***]
will upon request provide [***] with a certificate of insurance in that regard.

 

Article 12

 

CONFIDENTIALITY

 

12.1         Confidential
Proprietary Information.

 

12.1.1           Confidential
Proprietary Information. In connection with this Agreement, each Party may disclose technical, business or other confidential information
in connection with this Agreement, whether prior to, on, or after the Effective Date, including: (a) any unpublished Patents; and
(b) any information regarding the scientific, regulatory or business affairs or other activities of either Party (such confidential
information, “Confidential Proprietary Information”). Without limiting the foregoing, the terms of this Agreement
are the Confidential Proprietary Information of both Parties and shall be treated confidentially by each of the Parties, subject to the
exceptions set forth in Section 12.1.6. Information exchanged by the Parties pursuant to the Confidentiality Agreement shall be
treated as Confidential Proprietary Information under this Agreement and governed by the terms of this Agreement. Without limiting the
foregoing, until such time as the applicable information has become available to the public in accordance with this Agreement, ProQR
agrees not to disclose the sequence of any Compound except pursuant to Section 12.1.4.

 

12.1.2           Restrictions.
A Party (the “Receiving Party”) that receives Confidential Proprietary Information from the other Party (the
 “Disclosing Party”) shall keep all the Disclosing Party’s Confidential Proprietary Information in confidence
with the same degree of care with which the Receiving Party holds its own confidential information (but in no event less than a commercially
reasonable degree of care). A Receiving Party shall not use the Disclosing Party’s Confidential Proprietary Information except
in connection with the performance of its obligations and exercise of its rights under this Agreement.

 

12.1.3           Exceptions.
The obligations of confidentiality and restriction on use of Confidential Proprietary Information under Section 12.1.2 do not apply
to any information that the Receiving Party can prove by competent written evidence: (a) is now, or hereafter becomes, through no
act or failure to act on the part of the Receiving Party, generally known or available to the public; (b) is known by the Receiving
Party at the time of receiving such information, other than by previous disclosure of the Disclosing Party, or its Affiliates, employees,
agents, consultants, or contractors; (c) is hereafter furnished to the Receiving Party without restriction by a Third Party who
has no obligation of confidentiality or limitations on use with respect thereto, as a matter of right; or (d) is independently discovered
or developed by the Receiving Party without the use of or reference to Confidential Proprietary Information belonging to the Disclosing
Party. Specific information shall not be deemed to be within any of the foregoing exclusions merely because it is embraced by more general
information falling within those exclusions.

 

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12.1.4           Permitted
Disclosures. The Receiving Party may disclose Confidential Proprietary Information belonging to the Disclosing Party as expressly
permitted by this Agreement or if and to the extent such disclosure is reasonably necessary in the following instances:

 

(a)           made
by or on behalf of the Receiving Party to a Patent authority as may be reasonably necessary or useful for purposes of Prosecution and
Maintenance of Patents as permitted by this Agreement; provided, that neither Party shall file a patent application that discloses Program
IP that is solely owned by the other Party pursuant to this Agreement without the prior written consent of the owning Party (such consent
not to be unreasonably withheld, conditioned or delayed);

 

(b)           made
by or on behalf of the Receiving Party to Regulatory Authorities as required in connection with any Regulatory Filings for a product
that such Party has a license or right to develop in a given country or jurisdiction;

 

(c)           made
by or on behalf of the Receiving Party as may be reasonably necessary for prosecuting or defending litigation as permitted by this Agreement;

 

(d)           made
by or on behalf of the Receiving Party for the purpose of complying with a valid order of a court of competent jurisdiction or other
Governmental Authority of competent jurisdiction or, if in the opinion of the Receiving Party’s legal counsel, such disclosure
is otherwise required by Applicable Law;

 

(e)           made
by or on behalf of the Receiving Party where such disclosure is required by a Regulatory Authority (including in filings with the Securities
and Exchange Commission or other agency) of certain material developments or material information generated under this Agreement;

 

(f)           made
by or on behalf of the Receiving Party as of the Effective Date in response to a valid request by a U.S., state, foreign, provincial,
or local tax authority, in which case either Party may disclose a copy of this Agreement (including any Exhibits, Appendices, ancillary
agreements, and amendments hereto);

 

(g)           made
by the Receiving Party to its and its Affiliates’ employees, consultants, contractors and agents, and to Sublicensees (in the case
of Lilly), in each case on a need-to-know basis (as reasonably determined by the Receiving Party) in connection with the Exploitation
of Products or Terminated Products (if applicable) in the Field in the Territory, in each case under written obligations of confidentiality
and non-use at least as stringent as those herein; and

 

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(h)           made
by the Receiving Party to potential and actual investors, acquirers, licensees and other financial or commercial partners solely for
the purpose of evaluating or carrying out an actual or potential investment, acquisition, or collaboration, in each case under written
obligations of confidentiality and non-use at least as stringent as those herein; provided, however, that with respect to disclosure
to actual or bona fide potential investors, such disclosure is under a written obligation of confidentiality that is consistent with
market terms, including a shorter period of time during which such information must be held confidential.

 

Notwithstanding the foregoing, if a Party is
required to make a disclosure of the other Party’s Confidential Proprietary Information pursuant to Section 12.1.4(c) or
Section 12.1.4(d), it shall, except where impracticable, give reasonable advance notice to the other Party of such disclosure and
use efforts to secure confidential treatment of such Confidential Proprietary Information at least as diligent as such Party would use
to protect its own Confidential Proprietary Information, but in no event less than reasonable efforts. Any information disclosed pursuant
to this Section 12.1.4 remains Confidential Proprietary Information and subject to the restrictions set forth in this Agreement,
including the foregoing provisions of this Article 12.

 

12.1.5           Public
Domain Information and Residual Knowledge. Nothing in this Agreement shall prevent a Party from using any information that is in
the public domain. A Party shall also not be restricted under, and shall not be in breach of, this Agreement from using, within or outside
this Agreement and for any purpose, any general knowledge, skill, and expertise acquired by its employees (or its Affiliates’ employees)
in their performance of this Agreement (“Residuals”) solely to the extent such Residuals shall have been retained
in the unaided memory (without intentional memorization) of such employees in intangible form and without use by the Party or such employees
of tangible copies of any Confidential Proprietary Information of the other Party; provided that this provision will not be deemed in
any event to provide any right to infringe, or to grant any license to or under, the Patent rights of the other Party or of Third Parties
that have licensed or provided materials to the other Party; provided, further, that a Party’s use of such Residuals is on an “as
is, where is” basis, with all faults and all representations and warranties disclaimed and at such Party’s sole risk.

 

12.1.6           Disclosure
of Agreement. Notwithstanding the foregoing, either Party or its Affiliates may disclose the relevant terms of this Agreement: (a) to
the extent required or advisable to comply with the rules and regulations promulgated by the U.S. Securities and Exchange Commission
or any equivalent governmental agency in any country in the Territory, provided that such Party shall submit a confidential treatment
request in connection with such disclosure and shall submit with such confidential treatment request only such redacted form of this
Agreement, which redacted form of this Agreement shall be provided to the other Party for review and comment and which comments shall
be considered in good faith by the disclosing Party; (b) upon request from a Governmental Authority (such as a tax authority), provided
the disclosing Party uses reasonable efforts to ensure the Governmental Authority maintains such terms as confidential; (c) to applicable
licensors, to the extent necessary to comply with the terms of any Third Party license agreement, the rights under which are sublicensed
to the other Party under this Agreement; and (d) to the extent necessary to perform obligations or exercise rights under this Agreement,
to any sublicensee, collaborator or potential sublicensee or potential collaborator of such Party, provided that any sublicensee, collaborator
or potential sublicensee or collaborator agree in writing to be bound by obligations of confidentiality and non-use no less protective
of the Disclosing Party than those set forth in this Agreement.

 

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12.1.7           Survival.
Each Party’s obligations under this Section 12.1 (other than Section 12.1.5) shall apply during the Term and continue
for [***] thereafter with respect to Confidential Proprietary Information, except for information which is a “trade secret,”
for which each Party’s obligations under this Section 12.1 shall remain in place as long as the applicable Confidential Proprietary
Information retains its status as a trade secret. Section 12.1.5 shall apply during the Term and shall survive any expiration or
termination of this Agreement.

 

12.2         Publicity.
Promptly following the Effective Date, ProQR may issue a press release in a mutually agreed upon form attached hereto as Exhibit 12.2.
Thereafter, either Party may make subsequent public disclosure of the contents of such press release and, except as permitted under Section 12.1.4
and this Section 12.2, ProQR shall not issue any subsequent press release or public statement disclosing information relating to
this Agreement or the transactions contemplated hereby or the terms hereof without the prior written consent of Lilly, not to be unreasonably
withheld, conditioned, or delayed; provided however, that ProQR will not be prevented from complying with any duty of disclosure it may
have pursuant to Applicable Laws or pursuant to the rules of any recognized stock exchange or quotation system subject to the restrictions
set forth in Sections 12.1.4 and 12.1.6. If a Party desires to issue a press release disclosing information relating to this Agreement
or the transactions contemplated hereby or the terms hereof, it will provide the other Party with a copy of the proposed press release.
The disclosing Party shall specify with each such proposed press release or public statement, taking into account the urgency of the
matter being disclosed, a reasonable period of time within which the other Party may provide any comments on such proposed press release
or public statement. If such other Party provides any comments, the Parties shall consult with one another on such proposed press release
or public statement and work in good faith to prepare a mutually acceptable press release or public statement. The Parties may repeat
any information relating to this Agreement that has already been publicly disclosed in accordance with this Section 12.2, provided
such information continues as of such time to be accurate.

 

12.3         Publication.
Lilly shall be entitled to issue scientific publications and make presentations with respect to the Research Program, the Project Targets,
the Lilly Targets, the Products, and their testing in accordance with Lilly’s internal guidelines without approval by ProQR, and
Lilly shall be in control of any publications or scientific presentations regarding the Products or their testing subject to this Section 12.3.
ProQR shall not issue any scientific publications regarding the Project Targets, the Lilly Targets, the Products or their testing without
Lilly’s prior written consent.

 

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Article 13

 

TERM & TERMINATION

 

13.1         Term.
This Agreement commences on the Effective Date and, unless terminated earlier as provided in this Article 13, shall continue on
a Product-by-Product basis until the expiration of the last Royalty Term in the Territory for such Product (the “Term”).

 

13.2         Termination
for Material Breach.

 

13.2.1           Termination.
Either Party may terminate this Agreement upon written notice to the other Party if such other Party materially breaches its obligations
under this Agreement and, after receiving written notice from the non-breaching Party identifying such material breach in reasonable
detail, fails to cure such material breach within [***] from the date of such notice; provided that if a non-payment related breach is
not reasonably capable of cure within such [***] period, the breaching Party may submit, prior to the end of such [***] period, a reasonable
plan to cure the breach within an additional [***], in which case the other Party may not terminate this Agreement for so long as the
breaching Party is using Commercially Reasonable Efforts to implement such cure plan within such additional [***].

 

13.2.2           Dispute.
If the alleged breaching Party disputes in good faith the existence or materiality of a breach specified in a notice provided by the
other Party in accordance with Section 13.2.1, and such alleged breaching Party provides the other Party notice of such dispute
within such [***]-day period, then the non-breaching Party may not terminate this Agreement under Section 13.2.1 unless and until
it has been finally determined pursuant to Article 14 that the alleged breaching Party has materially breached this Agreement and
such Party fails to cure such breach within [***] days following such court’s decision. During the pendency of such dispute, all
of the terms and conditions of this Agreement shall remain in effect and the Parties shall continue to perform all of their respective
obligations hereunder.

 

13.2.3           Lilly
Option to Continue Agreement. Notwithstanding anything to the contrary under this Agreement, if ProQR materially breaches this Agreement,
as finally determined under Article 14, during the Research Term such that Lilly would otherwise have the right to terminate this
Agreement under Section 13.2, Lilly shall have the option to cause the Research Transfer Scenario to occur by written notice to
ProQR. Lilly shall also be entitled, in lieu of terminating this Agreement, to set-off any damages finally determined under Article 14
to have resulted from ProQR’s material breach from any Milestone Payments and Royalties payable by Lilly to ProQR. For clarity,
in such scenario, the Agreement shall continue in accordance with its terms, save as expressly set forth in this Section 13.2.3.

 

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13.3         Termination
by Lilly.

 

13.3.1           Partial
Termination. Lilly may, at any time in its sole discretion and without cause, terminate this Agreement on a Project Target-by-Project
Target, Project-by-Project, or Product-by-Product basis upon [***] days’ prior written notice to ProQR.

 

13.3.2           Entire
Agreement. Lilly may, in its sole discretion, terminate this Agreement in its entirety at any time and without cause upon [***] days’
prior written notice to ProQR.

 

13.4         Termination
for Patent Challenges. Except to the extent the following is unenforceable under the Applicable Law of a jurisdiction, then:

 

13.4.1           if
Lilly, its Affiliates, or Sublicensees, directly or indirectly: (a) initiate or request an interference or opposition proceeding
with respect to any ProQR Patents; (b) make, file, or maintain any claim, demand, lawsuit, or cause of action to challenge the validity
or enforceability of any ProQR Patents; or (c) oppose any extension of, or the grant of a supplementary protection certificate with
respect to, any ProQR-Controlled Patent, in each case other than in response to a threat of an infringement claim or as necessary to
secure allowance of a ProQR-owned patent claim, then ProQR may terminate this Agreement solely with respect to the challenged ProQR Patent(s) with
respect to any Projects or Products to which such patent challenge relates upon thirty (30) days’ prior written notice to Lilly;
and

 

13.4.2           if
ProQR, its Affiliates, or sublicensees, directly or indirectly, (a) initiate or request an interference or opposition proceeding
with respect to any Lilly Patents, (b) make, file, or maintain any claim, demand, lawsuit, or cause of action to challenge the validity
or enforceability of any Lilly Patents, or (c) oppose any extension of, or the grant of a supplementary protection certificate with
respect to, any Lilly Patents, in each case other than in response to a threat of an infringement claim or as necessary to secure allowance
of a Lilly-owned patent claim, then Lilly may terminate this Agreement with respect to the challenged Lilly Patent(s) with respect
to any Projects or Products to which such patent challenge relates upon thirty (30) days’ prior written notice to ProQR.

 

13.5         Effects
of Termination. Upon any termination of this Agreement, the provisions of this Section 13.5 will apply, provided that if this
Agreement is terminated only with respect to specified Products (“Terminated Products”) or Projects (“Terminated
Projects”) and not in its entirety, then the following will apply to such Terminated Products or Terminated Projects only,
and if this Agreement is terminated in its entirety, then all Products will be deemed Terminated Products and all Projects will be deemed
Terminated Projects. If this Agreement is terminated with respect to a Project Target, all Products Directed To such Project Target will
be deemed Terminated Products and the Project relating to such Project Target will be deemed a Terminated Project. If this Agreement
is terminated solely with respect to specified Projects or a Project Target, the applicable Project Target (the subject of such Project)
shall be deemed to no longer be a Project Target and the Terminated Products shall be deemed to no longer be Compounds or Products, in
each case, except for the purposes of the provisions of this Agreement relating to the effects of such termination.

 

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13.5.1           Termination
of Licenses. All licenses for Terminated Products granted by ProQR under Article 6 shall terminate automatically as of the termination
effective date; provided that, if Lilly (or its Affiliates or Sublicensees) has inventory of usable Product(s) as of the effective
date of termination, then Lilly (and its Affiliates and Sublicensees) may continue to sell off such inventory of Products in the Field
in the Territory (and fulfill customer orders therefor, including to manufacture Products for customer orders placed prior to the effective
date of termination) until the earlier to occur of [***] days after the effective date of termination and the date on which Lilly (or
its Affiliates or Sublicensees) no longer has such inventory of Product(s) and shall pay ProQR any applicable payments due based
on such sales. Any permitted sublicense granted by Lilly or its Affiliate to a Third Party under the licenses granted to Lilly under
this Agreement shall survive the termination of this Agreement, provided that, in the case where termination of this Agreement for Lilly’s
uncured material breach pursuant to Section 13.2, such Sublicensee did not cause such uncured material breach.

 

13.5.2           Destruction
of Confidential Proprietary Information. Subject to the potential transfer of any data and information covered below in Section 13.5.3,
each Receiving Party shall destroy (at the Disclosing Party’s written request) all such Confidential Proprietary Information of
the Receiving Party in its possession as of the effective date of expiration or termination (with the exception of one copy of such Confidential
Proprietary Information, which may be retained by the legal department of the Receiving Party to confirm compliance with the non-use
and non-disclosure provisions of this Agreement), and any Confidential Proprietary Information of the Disclosing Party contained in its
laboratory notebooks or databases, provided that each Receiving Party may retain and continue to use such Confidential Proprietary Information
of the Disclosing Party to the extent necessary to exercise any surviving rights, licenses or obligations under this Agreement. Notwithstanding
the foregoing, a Receiving Party shall not be required to destroy any computer files created during automatic system back up that are
subsequently stored securely by it and not readily accessible to its employees, consultants, or others who received the Disclosing Party’s
Confidential Proprietary Information under this Agreement.

 

13.5.3           Terminated
Product Reversion. Except in connection with any termination by Lilly pursuant to Section 13.2 or Section 13.4.2, in the
event of any termination of this Agreement in its entirety or with respect to a Project, if requested by ProQR, the Parties shall negotiate
in good faith to enter into a separate agreement detailing the potential transition to ProQR of Lilly’s rights and obligations
(or portions thereof) with respect to any Terminated Product or Terminated Project, in each case that is Covered by a Patent contained
in the Lilly or Joint Program IP, which agreement may provide for the payment of royalties or other compensation by ProQR to Lilly for
the Commercialization by ProQR of any such Terminated Product; and provided that: (a) Lilly shall have no obligation to negotiate
or grant a license to any Excluded Technology; (b) Lilly shall have no obligation to provide ProQR any Lilly Background IP or Excluded
Technology used in such Terminated Products (or any rights to any such Excluded Technology); and (c) with respect to any Lilly Technology
that is licensed to Lilly from a Third Party, Lilly shall have no obligation to negotiate with such Third Party for, or grant, any sublicense
rights to ProQR, but shall advise ProQR of the identity of such Third Party licensor and the nature of the relevant Lilly Technology,
and ProQR shall be solely responsible, at its sole cost and expense, for obtaining and negotiating for any rights to such Third Party’s
technology or Intellectual Property Rights. Notwithstanding the foregoing, if the Parties are unable to agree on the terms of such a
transition agreement within ninety (90) days of commencement of discussions with respect thereto despite their good faith efforts, Lilly
shall have no further obligation to enter into such an agreement or negotiate with ProQR with respect thereto.

 

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13.6           Survival.
Expiration or termination of this Agreement shall not relieve the Parties of any obligation or right accruing prior to such expiration
or termination. Except as set forth below or elsewhere in this Agreement, the obligations and rights of the Parties under the following
provisions of this Agreement shall survive expiration or termination of this Agreement: Articles 1 (to the extent defined terms are used
in other surviving provisions), 11, and 14, and Sections 6.3, 6.5, 6.6, 8.3.1 (solely with respect to the conversion of the license granted
in Section 6.1 in the event of the expiration of this Agreement), 8.3.7 and 8.4 (both only if and as needed for Lilly to provide
a final Royalty payment and report), 8.5 (for a period of [***] to permit ProQR to conduct a final audit if applicable), 9.1-9.4 (inclusive),
9.7.4, 9.8.5, 9.8.6 and 9.8.7 (both solely with respect to Joint Patents), 9.10 (solely with respect to Joint Patents), 12.1.7, 13.5,
13.6, 15.2, 15.4, 15.6, 15.11 and 15.17.

 

13.7           Bankruptcy
Code. If this Agreement is rejected or disclaimed by a Party as a debtor under Section 365 of the United States Bankruptcy Code,
the Dutch Bankruptcy Act (Faillissementswet) and the Insolvency Regulation (and implementations thereof) or similar provisions
in the bankruptcy, insolvency, reorganization or debtor relief laws of another jurisdiction (collectively, the “Code”),
then, notwithstanding anything else in this Agreement to the contrary, all licenses and rights to licenses granted under or pursuant
to this Agreement by the Party in bankruptcy to the other Party are, and shall otherwise be deemed to be, for purposes of Section 365(n) of
the Code (or similar provision in the bankruptcy laws of the jurisdiction), licenses of rights to “intellectual property”
as defined under Section 101(35A) of the Code (or similar provision in the bankruptcy laws of another applicable jurisdiction).
The Parties agree that a Party that is a licensee of rights under this Agreement shall retain and may fully exercise all of its rights
and elections under the Code, and that upon commencement of a bankruptcy proceeding by or against a Party under the Code, the other Party
shall be entitled to a complete duplicate of, or complete access to (as such other Party deems appropriate), any such intellectual property
and all embodiments of such intellectual property, if not already in such other Party’s possession, shall be promptly delivered
to such other Party: (a) upon any such commencement of a bankruptcy proceeding upon written request therefor by such other Party,
unless the bankrupt Party elects to continue to perform all of its obligations under this Agreement; or (b) if not delivered under
the foregoing subclause (a), upon the rejection of this Agreement by or on behalf of the bankrupt Party upon written request therefor
by the other Party. The foregoing provisions of this Section 13.7 are without prejudice to any rights a Party may have arising under
the Code.

 

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Article 14

 

GOVERNING LAW; DISPUTE RESOLUTION

 

14.1           Governing
Law. This Agreement is governed by and will be construed in accordance with the laws of the State of New York, without reference
to its conflict of laws principles. The United Nations Convention of International Contracts on the Sale of Goods (the Vienna Convention)
does not apply to this Agreement.

 

14.2           Disputes.
The Parties recognize that controversies or claims arising out of, relating to, or in connection with this Agreement may arise from time
to time. It is the objective of the Parties to establish procedures to facilitate the resolution of disputes in an expedient manner by
mutual cooperation and without resort to litigation. To accomplish this objective, the Parties shall follow the procedures set forth
in this Article 14 to resolve any dispute. If any dispute, claim or controversy of any nature arising out of or relating to this
Agreement, including any action or claim based on tort, contract or statute, or concerning the interpretation, effect, termination, validity,
performance or breach of this Agreement (each, a “Dispute”), arises between the Parties, either Party may request
in writing that the Dispute be submitted to the Executive Officers of each Party for resolution within [***] of such written notice.
If the Executive Officers have not succeeded in negotiating a resolution of the Dispute within [***] after the notice of Dispute, and
a Party wishes to pursue the matter, the Parties may seek to resolve the Dispute in any federal court having jurisdiction thereof located
in New York, New York as further described in Section 14.3.

 

14.3           Litigation;
Equitable Relief. The Federal courts located in New York, New York shall have exclusive jurisdiction over, and shall be the exclusive
venue for resolution of, any Dispute not resolved through the informal Dispute-resolution procedures described in Section 14.2.
If, within [***] following a notice by either Party to the other that it does not believe the Dispute can be resolved through the Executive
Officers, neither Party has commenced proceedings seeking to resolve such Dispute in any federal court having jurisdiction, then such
Dispute and all related rights, demands, claims, actions, causes of action, suits, proceedings and Losses of every kind and nature shall
be deemed to have been irrevocably waived and released, to the fullest extent permitted under Applicable Laws. Notwithstanding anything
to the contrary in this Agreement, either Party may, at any time and without waiving any remedy under this Agreement, seek from any court
having jurisdiction any temporary injunctive or provisional relief necessary to protect the rights or property of that Party. Any final
judgment resolving a Dispute may be enforced by either Party in any court having appropriate jurisdiction.

 

14.4           Waiver
Of Jury Trial.

 

EXCEPT AS LIMITED BY APPLICABLE LAW, EACH PARTY
HERETO HEREBY IRREVOCABLY WAIVES ALL RIGHT TO TRIAL BY JURY IN ANY ACTION, SUIT, PROCEEDING, OR COUNTERCLAIM (WHETHER BASED IN CONTRACT,
TORT, OR OTHERWISE) ARISING OUT OF OR RELATING TO THIS AGREEMENT OR THE ACTIONS OF ANY PARTY HERETO IN THE NEGOTIATION, ADMINISTRATION,
PERFORMANCE, AND ENFORCEMENT HEREOF. THE PARTIES AGREE THAT ANY OF THEM MAY FILE A COPY OF THIS ARTICLE WITH ANY COURT AS WRITTEN
EVIDENCE OF THE KNOWING, VOLUNTARY, AND BARGAINED-FOR AGREEMENT BETWEEN THE PARTIES IRREVOCABLY TO WAIVE ITS RIGHT TO TRIAL BY JURY IN
ANY ACTION, SUIT, PROCEEDING WHATSOEVER BETWEEN THEM RELATING TO THIS AGREEMENT WHICH SHALL INSTEAD BE TRIED IN A COURT OF COMPETENT
JURISDICTION BY A JUDGE SITTING WITHOUT A JURY.

 

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Article 15

 

MISCELLANEOUS

 

15.1           Entire
Agreement; Amendment. This Agreement, including the Exhibits hereto, sets forth the complete, final and exclusive agreement and all
the covenants, promises, agreements, warranties, representations, conditions and understandings between the Parties hereto with respect
to the subject matter hereof and supersedes, as of the Effective Date, all prior and contemporaneous agreements and understandings between
the Parties with respect to the subject matter hereof, including the Confidentiality Agreement. The foregoing may not be interpreted
as a waiver of any remedies available to either Party as a result of any breach, prior to the Effective Date, by the other Party of its
obligations under the Confidentiality Agreement. No subsequent alteration, amendment, change or addition to this Agreement shall be binding
upon the Parties unless reduced to writing and signed by an authorized officer of each Party.

 

15.2           Limitation
of Liability. NEITHER PARTY MAY RECOVER FROM THE OTHER PARTY ANY SPECIAL, INCIDENTAL, CONSEQUENTIAL OR PUNITIVE DAMAGES
IN CONNECTION WITH THIS AGREEMENT, REGARDLESS OF ANY NOTICE OF THE POSSIBILITY OF SUCH DAMAGES; PROVIDED, HOWEVER, THAT THIS SECTION 15.2
SHALL NOT BE CONSTRUED TO LIMIT EITHER PARTY’S INDEMNIFICATION OBLIGATIONS UNDER Article 11,
EITHER PARTY’S LIABILITY FOR BREACH OF ITS EXCLUSIVITY OBLIGATIONS UNDER Article 7 OR CONFIDENTIALITY OBLIGATIONS UNDER Article 12
OR LIABILITY OF A PARTY FOR ITS INFRINGEMENT OR MISAPPROPRIATION OF ANY INTELLECTUAL PROPERTY RIGHTS OR FOR A PARTY’S GROSS
NEGLIGENCE, WILLFUL MISCONDUCT OR FRAUD.

 

15.3           Independent
Contractors. The relationship between Lilly and ProQR created by this Agreement is solely that of independent contractors. This Agreement
does not create any agency, distributorship, employee-employer, partnership, joint venture or similar business relationship between the
Parties. Neither Party is a legal representative of the other Party, nor can either Party assume or create any obligation, representation,
warranty, or guarantee, express or implied, on behalf of the other Party.

 

15.4           Notice.
Any notice required or permitted to be given by this Agreement must be in writing, in English. Any and all notices or other communications
or deliveries required or permitted to be provided hereunder must be in writing and will be deemed given and effective if: (a) delivered
by hand or by overnight courier with tracking capabilities; (b) mailed postage prepaid by first class, registered, or certified
mail; or (c) delivered by electronic mail, in each case, addressed as set forth below unless changed by notice so given:

 

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	If to ProQR:	ProQR Therapeutics N.V.

    Zernikedreef 9

    2333 CK Leiden, the Netherlands

    Attn: Business Development

    [***]

     

    With a copy to (which will not constitute notice):

     

    ProQR Therapeutics VIII B.V.

    Zernikedreef 9

    2333 CK Leiden, the Netherlands

    Attn: Legal Department

    [***]

	 	 
	If to Lilly:	Eli Lilly and Company 

    Lilly Corporate Center

    Indianapolis, Indiana 46285

    Attn:  Vice President, Corporate Business Development 

    [***]

 

Each Party shall also provide
a copy of any notice (via e-mail if available) to the other Party’s Alliance Manager.

 

15.5           Severability.
If, for any reason, any part of this Agreement is adjudicated invalid, unenforceable, or illegal by a court of competent jurisdiction:
(a) such adjudication shall not, to the extent feasible, affect or impair, in whole or in part, the validity, enforceability, or
legality of any remaining portions of this Agreement; (b) this Agreement shall be construed and enforced as if such invalid, unenforceable
or illegal provision had never comprised a part hereof; and (c) all remaining portions will remain in full force and effect and
shall not be affected by the invalid, unenforceable or illegal provision or by its severance herefrom.

 

15.6           Non-Use
of Names. ProQR shall not use the name, trademark, logo, or physical likeness of Lilly or its respective officers, directors or employees,
or any adaptation of any of them, in any advertising, promotional or sales literature, without Lilly’s prior written consent. ProQR
shall require its Affiliates to comply with the foregoing. Lilly shall not use the name, trademark, logo, or physical likeness of ProQR
or its officers, directors or employees, or any adaptation of any of them, in any advertising, promotional or sales literature, without
ProQR’s prior written consent. Lilly shall require its Affiliates and Sublicensees to comply with the foregoing.

 

15.7           Assignment.
Neither Party may assign or transfer this Agreement or any rights or obligations hereunder without the prior written consent of the other,
except that a Party may make such an assignment or transfer without the other Party’s consent to: (a) its Affiliate, provided
that such Party shall remain primarily liable for any acts or omissions of such Affiliate; or (b) to an Acquirer in connection with
a Change of Control, subject to Section 15.8. Any permitted assignee shall, in writing to the non-assigning Party, expressly assume
performance of such assigning Party’s rights and obligations. Any permitted assignment is binding on the successors of the assigning
Party. Any assignment or attempted assignment by either Party in violation of the terms of this Section 15.7 is null, void and of
no legal effect.

 

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15.8         ProQR
Change of Control.

 

15.8.1           Notification
of Change of Control. ProQR shall provide Lilly with written notice of any Change of Control of ProQR promptly [***].

 

15.8.2           Effects
of Change of Control. Except in the scenario set forth in Section 15.8.4(b) below (in which case, the terms of Section 15.8.4(b) shall
apply to such scenario), following a Change of Control of ProQR at any time before expiration of the Research Term, Lilly may elect whether
the Research Transfer Scenario (in subsection (a) below) or the Research Continuance Scenario (in subsection (b) below) shall
apply to such Change of Control, as provided in Section 15.8.3 depending on whether or not such Acquirer is engaged in a Competing
Program as of the closing of the Change of Control transaction.

 

(a)           Research
Transfer Scenario. The “Research Transfer Scenario” means [***]:

 

(i)           [***];

 

(ii)          [***];
and

 

(iii)         [***]

 

(b)           Research
Continuance Scenario. The “Research Continuance Scenario” means [***].

 

15.8.3           Acquirer
Engaged in Competing Program. [***]

 

15.8.4           Acquirer
Not Engaged in Competing Program. [***]

 

15.8.5           Acquirer
Patent Rights. Following any Change of Control of ProQR to an Acquirer that owns or controls any Patent rights Covering Products,
if Lilly thereafter makes any payments to the Acquirer or owes any amounts to the Acquirer on account of such Patent rights and Lilly’s
exploitation of rights hereunder with respect to Products, Lilly may deduct [***] percent ([***]%) of such amounts. from any payments
otherwise due to ProQR under Article 8 For avoidance of doubt, this right shall not apply to amounts due under any agreement between
Lilly and the Acquirer that was in effect prior to the Change of Control (which may nonetheless be subject to Section [***]).

 

15.8.6           Firewalled
Programs. Promptly following the first to occur of any of the following events in relation to an Acquirer of ProQR: (a) [***],
(b) [***], or (c) [***].

 

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15.8.7           Firewall
Audits. Lilly shall have the right, through a designated Third Party auditor reasonably acceptable to ProQR, to audit ProQR’s
(and, as applicable, its Affiliates’) obligations under this Agreement regarding implementation and enforcement of Firewalls under
this Section 15.8 for purposes of confirming compliance with the Firewalls, identifying any vulnerabilities or breaches and requiring
ProQR (or its Affiliates) to promptly remediate any non-compliance identified by such audit. In connection with such audit, duly authorized
representatives of Lilly’s designated auditor may make an on-site visit to ProQR (or its Affiliate) for the purpose of conducting
such audit. Lilly may conduct such audits from time to time as reasonably necessary to confirm ProQR’s compliance with such Firewall
requirements no more than [***] or more frequently if Lilly reasonably believes at any time that ProQR is not in compliance with such
Firewall requirements; provided that if the auditor identifies a breach of the Firewall, Lilly will be entitled to [***] additional audits
within the same [***] to verify that appropriate action has been taken to remedy the breach of the Firewall. Any audits described under
this Section 15.8.7 shall be conducted during ProQR’s regular business hours, for a duration only as reasonably necessary
to confirm ProQR’s compliance with the applicable Firewall requirements, and shall not unreasonably interfere with or impede ProQR’s
business operations. Lilly shall provide ProQR with written notice of such audit at least [***] prior to such requested audit (or such
shorter period as may be designated by Lilly if Lilly reasonably believes at any time that ProQR is not in compliance with such Firewall
requirements). All such audits shall be conducted at Lilly’s cost and expense. If the auditor identifies any breach of the Firewall,
Lilly and/or the auditor will notify ProQR, and ProQR will promptly (and will use reasonable efforts to ensure its Affiliates promptly)
take all action necessary to remedy such breach, and will provide Lilly with reasonable assurance that such action has been taken, at
ProQR’s sole expense.

 

15.9           Waivers.
The failure of a Party to insist upon strict performance of any provision of this Agreement or to exercise any right arising out of this
Agreement shall neither impair that provision or right nor constitute a waiver of that provision or right, in whole or in part, in that
instance or in any other instance. Any waiver by a Party of a particular provision or right shall be in writing, shall be as to a particular
matter and, if applicable, for a particular period of time and shall be signed by such Party.

 

15.10           Force
Majeure. Neither Party shall be responsible to the other for any failure or delay in performing any of its obligations under this
Agreement or for other nonperformance hereunder (excluding, in each case, the obligation to make payments when due) if such delay or
nonperformance is caused by strike, fire, flood, earthquake, accident, war, act of terrorism, epidemics, pandemics, quarantines,
act of God or of the government of any country or of any local government, or by any other cause unavoidable or beyond the control of
any Party hereto. In such event, such affected Party shall use Commercially Reasonable Efforts to resume performance of its obligations
and will keep the other Party informed of actions related thereto.

 

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15.11           Interpretation.
The captions and headings to this Agreement are for convenience only, and are to be of no force or effect in construing or interpreting
any of the provisions of this Agreement. Unless specified to the contrary, references to Articles, Sections, Appendices or Exhibits mean
the particular Articles, Sections, Appendices or Exhibits to this Agreement and references to this Agreement include all Exhibits
hereto. In the event of any conflict between the main body of this Agreement and any Exhibit hereto, the main body of this Agreement
shall prevail. Unless context otherwise clearly requires, whenever used in this Agreement: (a) the words “include” or
 “including” shall be construed as incorporating, also, “but not limited to” or “without limitation”;
(b) the word “day” or “year” means a calendar day or year unless otherwise specified; (c) the word
 “notice” means notice in writing (whether or not specifically stated) and shall include notices, consents, approvals and
other written communications contemplated under this Agreement; (d) the words “hereof,” “herein,” “hereby”
and derivative or similar words refer to this Agreement as a whole and not merely to the particular provision in which such words appear;
(e) the words “shall” and “will” have interchangeable meanings for purposes of this Agreement; (f) provisions
that require that a Party, the Parties or a committee hereunder “agree,” “consent” or “approve” or
the like shall require that such agreement, consent or approval be specific and in writing, whether by written agreement, letter, approved
minutes, email or otherwise; (g) words of any gender include the other gender; (h) words using the singular or plural number
also include the plural or singular number, respectively; (i) references to any specific law, rule or regulation, or article,
section or other division thereof, shall be deemed to include the then-current amendments thereto or any replacement law, rule or
regulation thereof; (j) the phrase “non-refundable” shall not prohibit, limit or restrict either Party’s right
to obtain damages in connection with a breach of this Agreement; and (k) neither Party shall be deemed to be acting on behalf of
the other Party.

 

15.12           Counterparts;
Electronic Signatures. This Agreement may be executed in any number of counterparts, each of which is deemed an original, but all
of which together constitute one instrument. This Agreement may be executed and delivered electronically and upon such delivery such
electronic signature will be deemed to have the same effect as if the original signature had been delivered to the other Party.

 

15.13           Expenses.
Each Party shall pay its own costs, charges and expenses incurred in connection with the negotiation, preparation and execution of this
Agreement.

 

15.14           Further
Assurances. Lilly and ProQR hereby covenant and agree without the necessity of any further consideration, to execute, acknowledge
and deliver any and all documents and take any action as may be reasonably necessary to carry out the intent and purposes of this Agreement.

 

15.15           No
Third Party Beneficiary Rights. This Agreement is not intended to and shall not be construed to give any Third Party any interest
or rights (including any Third Party beneficiary rights) with respect to or in connection with any agreement or provision contained herein
or contemplated hereby, except for Lilly Indemnitees and ProQR Indemnitees as expressly provided in Section 11.1 or as otherwise
expressly provided for in this Agreement.

 

15.16           Construction.
The Parties hereto acknowledge and agree that: (a) each Party and its counsel reviewed and negotiated the terms and provisions of
this Agreement and have contributed to its revision; (b) the rule of construction to the effect that any ambiguities are resolved
against the drafting Party shall not be employed in the interpretation of this Agreement; and (c) the terms and provisions of this
Agreement shall be construed fairly as to all Parties hereto and not in a favor of or against any Party, regardless of which Party was
generally responsible for the preparation of this Agreement.

 

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15.17           Cumulative
Remedies. No remedy referred to in this Agreement is intended to be exclusive unless explicitly stated to be so, but each shall be
cumulative and in addition to any other remedy referred to in this Agreement or otherwise available under law.

 

15.18           Extension
to Affiliates. Except as expressly set forth otherwise in this Agreement, each Party shall have the right to extend the rights and
immunities granted in this Agreement to one or more of its Affiliates. All applicable terms and provisions of this Agreement, except
this right to extend, shall apply to any such Affiliate to which this Agreement has been extended to the same extent as such terms and
provisions apply to the Party extending such rights and immunities. For clarity, Lilly extending the rights and immunities granted hereunder
shall remain primarily liable for any acts or omissions of its Affiliates.

 

15.19           ProQR
Corporate Group. ProQR shall ensure that the multiple entities within the definition of “ProQR” remain Affiliates of
each other throughout the Term and each such entity shall be jointly and severally liable for the performance or failure to perform of
any such other entity hereunder. Lilly may take action to enforce this Agreement against any or all such entities and no such entity
shall raise as a defense that the performance or failure to perform was an obligation of a different entity within the definition of
ProQR.

 

[signature page follows]

 

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In
Witness Whereof, the Parties have caused this Agreement to be executed as of the Effective Date by their duly authorized representatives.

 

	 	 
	Eli Lilly and Company	 
	 	 
	 	 
	By:	/s/David
    A. Ricks	 
	Name:	     David A. Ricks	 
	Title:	     Chairman and
    Chief Executive Officer	 
	 	 
	 	 
	ProQR Therapeutics
    N.V.	 
	 	 
	 	 
	By:	/s/Daniel de Boer	 
	Name:	     Daniel de Boer	 
	Title:	     Chief Executive
    Officer	 
	 	 
	 	 
	ProQR Therapeutics
    VIII B.V.	 
	 	 
	 	 
	By:	/s/Daniel de Boer	 
	Name:	     Daniel de Boer	 
	Title:	     Chief Executive
    Officer	 

 

[Signature
Page to Research and Collaboration Agreement] 

 

    

     

    

 

Exhibit 1.115 - ProQR Platform Patents

 

[***]

 

    

     

    

 

Exhibit 3.3 − Reserved Targets

 

Gene (Disease/Protein) – Uniprot ID

 

[***]

 

    

     

    

 

Exhibit 4.1 – Research Program
Outline

 

[***]

 

    

     

    

 

Exhibit 4.4.2 - Initial Workplan

 

[***]

 

    

     

    

 

Exhibit 4.7 Part A – Good
Research Practices

 

Good Research Practice Expectations for External
Partners

 

[***]

 

    

     

    

 

Exhibit 4.7 Part B – Animal
Care

 

Lilly Principles for Animal Care and Use for
Third Party Organizations

 

[***]

 

     

     

    

 

Exhibit 4.9 - Materials Transfer Record

 

[***]

 

     

     

    

 

Exhibit 10.2 – Disclosures Regarding
ProQR Representations and Warranties

 

[***]

 

     

     

    

 

Exhibit 10.2.4 - Existing Patents

 

[***]

 

     

     

    

 

Exhibit 12.2 – Publicity

 

[***]Exhibit 10.2

 

EXECUTION
VERSION

 

 

SHARE PURCHASE AGREEMENT

 

This Share
Purchase Agreement (this “Agreement”) is entered into as of September 3,
2021 (the “Execution Date”), by and between ProQR Therapeutics N.V.,
a public company with limited liability (naamloze vennootschap) incorporated under the laws of The Netherlands (“ProQR”),
and Eli Lilly and Company, a corporation organized and existing under the laws of
Indiana, with its principal business office located at Lilly Corporate Center, Indianapolis, Indiana 46285, U.S.A. (“Lilly”).
ProQR and Lilly are each hereafter referred to individually as a “Party” and together as the “Parties.”
The capitalized terms used herein and not otherwise defined have the meanings given to them in Appendix 1 attached hereto or the
Collaboration Agreement.

 

RECITALS

 

Whereas,
the Parties are entering into that certain Research and Collaboration Agreement of even date herewith (the “Collaboration Agreement”);
and

 

Whereas,
pursuant to the terms and subject to the conditions set forth in this Agreement, ProQR desires to issue and sell to Lilly, and Lilly desires
to purchase from ProQR, ProQR’s ordinary shares, nominal value € 0.04 per share (“Ordinary Shares”).

 

Now,
Therefore, in consideration of the following mutual promises and obligations, and for good and valuable consideration, the
adequacy and sufficiency of which are hereby acknowledged, the Parties agree as follows:

 

Article 1

 

SALE AND PURCHASE OF SHARES

 

1.1            Purchase
of Shares. Subject to the terms and conditions of this Agreement, at the Closing, ProQR will
issue and sell to Lilly, and Lilly will purchase from ProQR, 3,989,976 Ordinary Shares (the “Shares”)
for an aggregate purchase price of $30,000,003 (the “Purchase Price”).

 

1.2            Payment.
At the Closing, Lilly will pay the Purchase Price by wire transfer or electronic funds transfer of immediately available funds to an account
designated by ProQR, which account ProQR shall designate to Lilly no less than four (4) Business Days prior to the Closing Date (or
on such later date as may be permitted by Lilly). ProQR consents to payment of the Purchase Price in United States dollars.

 

1.3            Closing.
(a) The closing of the purchase and sale of the Shares hereunder (the “Closing”)
shall be held on the second (2nd) Business Day after
the satisfaction or waiver of the conditions to Closing set forth in Article 5 (other than those conditions that by their
nature are to be satisfied at the Closing, but subject to the satisfaction or waiver of such conditions at the Closing), at 10:00 a.m. Eastern
Time, remotely via the exchange of documents and signatures, or at such other time, date and location as the Parties may agree orally
or in writing. The date the Closing occurs is hereinafter referred to as the “Closing Date.”

 

(b)            ProQR
shall instruct its transfer agent at the Closing to register the Shares in book-entry in the name of Lilly, and ProQR shall cause its
transfer agent to deliver written confirmation of the book-entry delivery of the Shares to Lilly. ProQR will also deliver to Lilly at
the Closing a certificate in form and substance reasonably satisfactory to Lilly and duly executed on behalf of ProQR by an authorized
officer of ProQR, certifying that the conditions to the Closing set forth in Section 5.3 of this Agreement have been satisfied;

 

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(c)            At
Closing, Lilly shall deliver to ProQR a certificate in form and substance reasonably satisfactory to ProQR and duly executed on behalf
of Lilly by an authorized officer of Lilly, certifying that the conditions to Closing set forth in Section 5.2 of this Agreement
have been satisfied; and

 

(d)            At
the Closing, subject to receipt by ProQR of the Purchase Price from Lilly as contemplated by Section 1.2, ProQR shall deliver
or cause to be delivered to Lilly a true and correct copy of the irrevocable instructions to the Transfer Agent instructing the Transfer
Agent to deliver the Shares to Lilly on an expedited basis and record the Shares in the register of the Transfer Agent in book entry form.

 

Article 2

 

REPRESENTATIONS AND WARRANTIES OF PROQR

 

Except as otherwise specifically
contemplated by this Agreement, ProQR hereby represents and warrants as of the Execution Date and the Closing Date to Lilly that:

 

2.1            Private
Placement. Subject to the accuracy of the representations and warranties made by Lilly in
Article 3, the Shares will be issued and sold to Lilly in compliance with applicable exemptions from the registration and
prospectus delivery requirements of the Securities Act and the registration and qualification requirements of all applicable securities
laws of the states of the United States.

 

2.2            Organization
and Qualification. ProQR is an entity duly incorporated and validly existing as a public
company with limited liability (naamloze vennootschap) under the laws of the Netherlands, with the requisite corporate power and
authority to own or lease and use its properties and assets, to execute and deliver the Transaction Documents, to carry out the provisions
of the Transaction Documents and to issue and sell the Shares. True and correct copies of ProQR’s Amended Articles of Association
(the “Organizational Documents”), as amended and in effect on the Effective Date, are filed or incorporated by reference
as exhibits to the SEC Documents (defined below).

 

2.3            Authorization.
ProQR has the requisite corporate power and authority and has taken all requisite corporate action necessary for, and no further action
on the part of ProQR, its officers, directors or shareholders is necessary for, (i) the authorization, execution and delivery of
the Transaction Documents, (ii) the authorization of the performance of all obligations of ProQR hereunder or thereunder, and (iii) the
sale, issuance and delivery of the Shares. Each of the Transaction Documents, upon execution and delivery by ProQR, assuming due authorization,
execution and delivery by Lilly, constitute valid and binding obligations of ProQR, enforceable in accordance with their terms, except
(a) as limited by applicable bankruptcy, insolvency, reorganization, moratorium or other laws of general application affecting enforcement
of creditors’ rights and (b) general principles of equity that restrict the availability of equitable remedies.

 

    2 

     

    

 

2.4            Issuance
of Shares. When issued, the Shares will be duly and validly issued and, when paid for in
accordance with this Agreement, will be fully paid and non-assessable, and, when delivered to Lilly at the Closing, shall be free and
clear of all encumbrances and restrictions including, but not limited to, liens, encumbrances or restrictions on transfer, including preemptive
rights, rights of first refusal, purchase options, call options, subscription rights or other similar rights of shareholders of ProQR,
except for restrictions on transfer set forth in the Transaction Documents or imposed by applicable securities laws. Assuming the accuracy
of the representations and warranties of Lilly in this Agreement, the Shares will be issued in compliance with all applicable federal
and state securities laws of Indiana, the state in which Lilly’s principal place of business is located. No stop order or suspension
of trading of the Ordinary Shares has been imposed by Nasdaq or the SEC and remains in effect.

 

2.5            SEC
Documents, Financial Statements. (a) ProQR has (i) timely filed or furnished, as
applicable, all reports, schedules, forms, statements and other documents required to be filed or furnished by it with the United States
Securities and Exchange Commission (the “SEC”) since January 1, 2019, pursuant to the reporting requirements of
the Exchange Act (all of the foregoing and all exhibits included therein and financial statements and schedules thereto and documents
incorporated by reference therein, collectively, the “SEC Documents”) and (ii) delivered or made available (by
filing on the SEC’s electronic data gathering and retrieval system (EDGAR)) to Lilly complete copies of the SEC Documents, including,
but not limited to, its Annual Report on Form 20-F for the year ended December 31, 2020 (the “Form 20-F”).
As of its date, or if amended, as of the date of the last such amendment, each SEC Document complied in all material respects with the
requirements of the Exchange Act applicable to such SEC Documents, and, as of its date, or if amended, as of the date of the last such
amendment, such SEC Document did not contain any untrue statement of a material fact or omit to state a material fact required to be stated
therein or necessary in order to make the statements therein, in light of the circumstances under which they were made, not misleading.

 

(b)            (i) As
of the respective dates and for the respective periods indicated, the audited consolidated financial statements (including the notes thereto)
of ProQR included in the Form 20-F comply as to form in all material respects with the published rules and regulations of the
SEC with respect thereto, have been prepared in accordance with IFRS applied on a consistent basis during the periods involved (except
as may be indicated in the notes thereto) and fairly present in all material respects, in accordance with IFRS, the consolidated financial
position of ProQR as of the dates thereof and the consolidated results of its operations and cash flows for the periods then ended.

 

(ii)            As
of the respective dates and for the respective periods indicated, the unaudited consolidated financial statements (including the notes
thereto) of ProQR included in the Form 6-K filed on May 6, 2021 with the SEC comply as to form in all material respects with
the published rules and regulations of the SEC with respect thereto, have been prepared in accordance with IFRS applied on a consistent
basis during the periods involved (except as may be indicated in the notes thereto) and fairly present in all material respects, in accordance
with IFRS, the consolidated financial position of ProQR as of the dates thereof and the consolidated results of its operations and cash
flows for the periods then ended.

 

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(c)            The
Ordinary Shares are listed on Nasdaq and registered pursuant to Section 12(b) of the Exchange Act, and ProQR has taken no action
designed to or reasonably likely to have the effect of terminating the registration of the Ordinary Shares under the Exchange Act or delisting
the Ordinary Shares from Nasdaq. As of the Execution Date, ProQR has not received any written notification that, and has no Knowledge
that, the SEC or Nasdaq is contemplating terminating such registration or listing. ProQR is in compliance in all material respects with
the requirements of Nasdaq for continued listing of the Ordinary Shares thereon.

 

2.6            Internal
Controls; Disclosure Controls and Procedures. ProQR has established and maintains internal
control over financial reporting as defined in Rule 13a-15(f) under the Exchange Act. ProQR has implemented the “disclosure
controls and procedures” (as defined in Rules 13a-15(e) and 15d-15(e) under the Exchange Act) that (a) are required
in order for the principal executive officer and principal financial officer of ProQR to engage in the review and evaluation process mandated
by the Exchange Act, (b) have been evaluated by management of ProQR for effectiveness as of December 31, 2020 and (c) are,
to the Knowledge of ProQR, effective at a reasonable assurance level. To the Knowledge of ProQR, as of the Execution Date, ProQR is in
compliance with such disclosure controls and procedures in all material respects. Each of the principal executive officer and the principal
financial officer of ProQR has made all certifications required by Sections 302 and 906 of the Sarbanes-Oxley Act of 2002 with respect
to all reports, schedules, forms, statements and other documents required to be filed by ProQR with the SEC. From January 1, 2019
through the Execution Date, to the Knowledge of ProQR, there have been no (a) significant deficiencies or material weaknesses in
ProQR’s internal control over financial reporting (whether or not remediated), (b) change in ProQR’s internal control
over financial reporting that has materially affected, or is reasonably likely to materially affect, ProQR’s internal control over
financial reporting and (c) fraud that involves management or other employees who have a significant role in ProQR’s internal
control over financial reporting.

 

2.7            Capitalization
and Voting Rights. (a) As of the Effective Date, the capitalization of ProQR is as set
forth in Schedule 2.7. All of the issued and outstanding Ordinary Shares (A) have been duly authorized and validly issued, (B) are
fully paid and nonassessable and (C) were issued in material compliance with applicable federal and state securities laws and not
in violation of any preemptive rights.

 

(b)            As
of the Effective Date, all of the issued and outstanding Ordinary Shares are entitled to one (1) vote per share.

 

(c)            Except
as disclosed in the SEC Documents, there are no (i) outstanding equity securities, options, warrants, rights (including conversion
or preemptive rights, rights of first refusal, rights of first purchase, purchase options, call options or subscription rights) or other
agreements pursuant to which ProQR is or may become obligated to issue or sell, any of its share capital or any other securities of ProQR
other than equity securities that may have been granted pursuant to its share incentive plans, which plans are described in the SEC Documents,
(ii) restrictions on the transfer of share capital of ProQR other than pursuant to federal or state securities laws or as set forth
in this Agreement or (iii) obligation (contingent or otherwise) on the part of ProQR to repurchase, redeem or otherwise acquire any
of its equity securities or any interests therein or to pay any dividend or make any distribution in respect thereof.

 

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(d)            Except
as disclosed in the SEC Documents, ProQR is not a party to or subject to any agreement or understanding relating to the voting of the
share capital of ProQR or the giving of written consents by a shareholder or director of ProQR or relating to the registration of the
share capital of ProQR under the 1933 Act.

 

(e)            Except
as disclosed in the SEC Documents, ProQR does not have outstanding any shareholder rights plans or “poison pill” or any similar
arrangement in effect giving any Person the right to purchase any equity interest in ProQR upon the occurrence of certain events.

 

2.8            No
Conflicts; Government Consents and Permits. (a) The execution, delivery and performance
of the Transaction Documents by ProQR and the consummation by ProQR of the transactions contemplated thereby (including the issuance of
the Shares) will not (i) conflict with or result in a violation of any provision of ProQR’s Organizational Documents, (ii) result
in any encumbrance upon any of the Shares, other than restrictions on resale pursuant to securities laws or as set forth in this Agreement,
(iii) materially violate or conflict with, or result in a material breach, default, modification, acceleration of payment or termination
under of any provision of, or constitute a default under, any Material Contract, or (iv) result in a material violation of any law,
rule, regulation, order, judgment or decree (including United States federal and state securities laws and regulations and regulations
of any self-regulatory organizations) applicable to ProQR as of the Execution Date.

 

(b)            ProQR
is not required to obtain any consent, authorization or order of, or make any filing or registration with, any Governmental Authority
in order for it to execute, deliver and perform its obligations under this Agreement in accordance with the terms hereof, or to issue
and sell the Shares in accordance with the terms hereof, other than such as have been made or obtained, and except for (i) any post-Closing
filings required to be made under federal or state “blue sky” or securities laws or (ii) any required filings or notifications
regarding the issuance or listing of additional shares with Nasdaq.

 

2.9            Litigation.
Other than as set forth in the SEC Documents, there is no material action, suit, proceeding or investigation pending (of which ProQR has
received written notice or otherwise has Knowledge) or, to ProQR’s Knowledge, threatened, against ProQR or which ProQR intends to
initiate.

 

2.10            Licenses
and Other Rights; Compliance with Laws. Each of ProQR and its subsidiaries possesses such
valid and current certificates, authorizations or permits required by state, federal, provincial or foreign regulatory agencies or bodies
to conduct its business as currently conducted and as described in the SEC Documents, except where the failure to so possess would not
reasonably be expected to be materially adverse to ProQR and its subsidiaries, taken as a whole (“Permits”). Each of
ProQR and its subsidiaries is not in violation of, or in default under, any of the Permits, except for such violations or defaults that
would not reasonably be expected to be materially adverse to ProQR and its subsidiaries, taken as a whole. Neither ProQR nor any of its
subsidiaries has received any notice of proceedings relating to the revocation or modification of, or non-compliance with, any such Permits,
which if the subject of an unfavorable decision, ruling, or finding would reasonably be expected to be materially adverse to ProQR and
its subsidiaries, taken as a whole.

 

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2.11            Intellectual
Property. The representations and warranties of ProQR contained in Section 10.2
of the Collaboration Agreement are, subject to the exceptions and qualifications contained therein and disclosures related thereto, true,
correct and complete.

 

2.12            Taxes
and Tax Returns. Each of ProQR and each of its subsidiaries has timely filed (taking into
account all applicable extensions) all material Tax Returns required to be filed by it; all such Tax Returns were correct and complete
in all material respects; and each of ProQR and each of its subsidiaries has paid (or has had paid on its behalf) to the appropriate Governmental
Authority all material Taxes that have been required to be paid by it; except, in each case, with respect to matters contested (or that
could be contested) in good faith or for which adequate reserves have been established in accordance with the requirements of IFRS, if
any. There are no disputes pending or, to the Knowledge of ProQR, claims asserted in writing in respect of Taxes of ProQR or any of its
subsidiaries for which reserves that are required to be established under IFRS have not been established.

 

2.13            Absence
of Certain Changes. Since December 31, 2020 through the Execution Date, except as set
forth in the SEC Documents or as contemplated by this Agreement or the Collaboration Agreement, there has not been:

 

(a)            Any
change, development, occurrence or event that has had or would reasonably be expected to have, individually or in the aggregate, a Material
Adverse Effect on ProQR;

 

(b)            any
declaration, setting aside or payment of any dividends, or authorization or making of any distribution upon or with respect to any class
or series of ProQR’s share capital, (ii) sale, exchange or other disposition of any material assets or rights outside the ordinary
course of business of ProQR or its subsidiaries, or (iii) repurchase, redemption or other acquisition of any outstanding share capital
of ProQR;

 

(c)            any
admission by ProQR in writing of its inability to pay its debts generally as they become due, filing or consent to the filing against
it of a petition in bankruptcy or a petition to take advantage of any insolvency act, made an assignment for the benefit of creditors,
consent to the appointment of a receiver for itself or for the whole or any substantial part of its property, or any petition in bankruptcy
filed against it, been adjudicated a bankrupt or filed a petition or answer seeking reorganization or arrangement under the federal bankruptcy
laws or any other laws of the United States or any other jurisdiction;

 

(d)            any
material Tax election made or changed, any audit settled or any amended Tax Returns filed of ProQR;

 

(e)            any
material damage, destruction or loss (whether or not covered by insurance) involving any material asset or right of ProQR and its subsidiaries;

 

    6 

     

    

 

(f)            any
sale, assignment or transfer, or any agreement to sell, assign or transfer, any material asset, liability, property, obligation or right
of ProQR or any of its subsidiaries to any Person, in each case, other than in the ordinary course of business;

 

(g)            any
material obligation or liability incurred, or any material loans or advances made, by ProQR or any of its subsidiaries to any of its or
their other Affiliates, other than in the ordinary course of business;

 

(h)            any
purchase or acquisition, or agreement, plan or arrangement to purchase or acquire, any material property, rights or assets other than
in the ordinary course of business by ProQR or any of its subsidiaries;

 

(i)            any
material waiver of any material rights or claims of ProQR or any of its subsidiaries;

 

(j)            any
material lien upon, or adversely affecting, any material property or other material assets of ProQR or any of its subsidiaries; or

 

(k)            any
Contract entered into by ProQR or any of its subsidiaries to do any of the foregoing.

 

2.14            No
Undisclosed Material Liabilities. ProQR and its subsidiaries do not have any liabilities
or obligations of any nature (whether accrued, absolute, contingent or otherwise), except for liabilities or obligations (a) reflected
or reserved against on the most recent consolidated balance sheet of ProQR included in the SEC Documents or the notes thereto, (b) incurred
since the latest date of such balance sheet in the ordinary course of business or (c) that are not material to ProQR.

 

2.15            Material
Contracts. Each Material Contract is included as an exhibit in the SEC Documents. Each Material
Contract is the legal, valid and binding obligation of ProQR, enforceable against ProQR in accordance with its terms, and, to the Knowledge
of ProQR, is the legal, valid and binding obligation of the other party thereto, enforceable against each other party thereto in accordance
with its terms, except in each case to the extent that (a) enforceability may be limited by bankruptcy, insolvency, reorganization,
moratorium or similar laws affecting creditors’ rights generally and by general equitable principles and (b) the indemnification
provisions of certain agreements may be limited by federal or state securities laws or public policy considerations in respect thereof.
ProQR is not in material breach, violation or default under any such Material Contract or, to ProQR’s Knowledge, is any other Person
counterparty to such Material Contract. ProQR has not been notified in writing that any Third Party to any Material Contract has indicated
that such Third Party intends to cancel, terminate or not renew any Material Contract.

 

2.16            Brokers’
or Finders’ Fees. No broker, finder, investment banker, or other Person is entitled
to any brokerage, finder’s or other similar fee or commission from ProQR in connection with the transactions contemplated by this
Agreement or the Collaboration Agreement.

 

    7 

     

    

 

2.17            Not
an Investment Company. ProQR is not, and solely after receipt of the Purchase Price, will
not be, required to register as an “investment company” as defined in the Investment Company Act of 1940, as amended.

 

2.18            No
Integration. Assuming the accuracy of the representations and warranties of Lilly, the offer,
sale and issuance of the Shares will be exempt from the registration requirements of the 1933 Act, and will have been registered or qualified
(or are exempt from registration and qualification) under the registration, permit or qualification requirements of all applicable state
securities laws of Indiana, the state in which Lilly’s principal place of business is located. Neither ProQR nor any of its Affiliates
or any Person or agent on its or their behalf has engaged in any form of general solicitation or general advertising with respect to the
Shares nor have any of such Person or agent sold, offered for sale, solicited offers to sell or will solicit any offers to sell or has
offered to sell or will offer to sell all or any part of the Shares to any Person or Persons so as to (a) bring the sale of such
Shares by ProQR within the registration requirements of the 1933 Act or the securities laws of the Netherlands or (b) cause this
offering of Shares to be integrated with any prior offering of securities of ProQR for purposes of the 1933 Act or any applicable shareholder
approval provision of Nasdaq, nor will ProQR take any actions or steps that would cause the offering or issuance of the Shares to be integrated
with other offerings.

 

2.19            Foreign
Corrupt Practices Act. Neither ProQR nor any of its subsidiaries nor, to ProQR’s Knowledge,
any director, officer, agent, employee or other Person acting on behalf of ProQR or any of its subsidiaries has, in the course of its
actions for, or on behalf of, ProQR or any of its subsidiaries (a) used any corporate funds for any unlawful contribution, gift,
entertainment or other unlawful expenses relating to political activity; (b) made any direct or indirect unlawful payment to any
domestic government official, “foreign official” (as defined in the U.S. Foreign Corrupt Practices Act of 1977, as amended,
and the rules and regulations thereunder (collectively, the “FCPA”)) or employee from corporate funds; (c) violated
or is in violation of any provision of the FCPA or, to ProQR’s Knowledge, any applicable non-U.S. anti-bribery statute or regulation;
or (d) made any unlawful bribe, rebate, payoff, influence payment, kickback or other unlawful payment to any domestic government
official, such foreign official or employee. ProQR and its subsidiaries have conducted their respective businesses in compliance with
the FCPA and have instituted and maintain policies and procedures designed to ensure continued compliance therewith.

 

2.20            Money
Laundering Laws. The operations of ProQR and its subsidiaries are, and have been conducted
at all times, in compliance with applicable financial recordkeeping and reporting requirements of the Currency and Foreign Transactions
Reporting Act of 1970, as amended, and to ProQR’s Knowledge, the money laundering statutes of all applicable jurisdictions, the
rules and regulations thereunder and any related or similar applicable rules, regulations or guidelines, issued, administered or
enforced by any Governmental Authority.

 

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2.21            OFAC.
Neither ProQR nor any of its subsidiaries nor, to ProQR’s Knowledge, any director, officer, agent, employee or Person acting on
behalf of ProQR or any of its subsidiaries is currently subject to any U.S. sanctions administered by the Office of Foreign Assets Control
of the U.S. Treasury Department (“OFAC”); and ProQR will not directly or indirectly use the proceeds from the sale
of the Shares, or lend, contribute or otherwise make available such proceeds to any subsidiary or any joint venture partner or other Person,
for the purpose of financing the activities of or business with any Person, or in any country or territory, that currently is subject
to any U.S. sanctions administered by OFAC or in any other manner that will result in a violation by ProQR or any of its subsidiaries
of U.S. sanctions administered by OFAC.

 

2.22            Preclinical
and Clinical Data and Regulatory Compliance. Except as set forth in the SEC Documents (excluding
any forward-looking disclosures set forth in any “risk factors” section or “forward-looking statements” section
thereof), as of the Execution Date, the preclinical tests and clinical trials (collectively, “Studies”) that are described
in, or the results of which are referred to in, the SEC Documents were and, if still pending, are being conducted in all material respects
in accordance with the protocols, procedures and controls designed and approved for such Studies, except in each case as would not, individually
or in the aggregate, reasonably be expected to be materially adverse to ProQR. Except as set forth in the SEC Documents, as of the Execution
Date, neither ProQR nor any of its subsidiaries has received any written notice of, or correspondence from, any Regulatory Authority (as
defined below) or institutional review board requiring the termination, suspension or material modification of any Studies that are described
or referred to in the SEC Documents and ProQR and each such subsidiary have operated and currently are in compliance in all material respects
with applicable laws, rules, regulations and policies of the federal, state, local or foreign agencies or bodies engaged in the regulation
of pharmaceuticals and biological products such as those being developed by ProQR (collectively, “Regulatory Authorities”),
including current Good Laboratory Practices and current Good Clinical Practices, except in each case as would not, individually or in
the aggregate, reasonably be expected to be materially adverse to ProQR and such subsidiaries, taken as a whole.

 

2.23            Regulatory
Permits. Except as set forth in the SEC Documents, (a) ProQR and each of its subsidiaries
have such material permits, licenses, certificates, approvals, clearances, authorizations or amendments thereto (the “Regulatory
Permits”) issued by the appropriate federal, state, local or foreign regulatory agencies or bodies necessary to conduct the
business of ProQR as currently conducted and as described in the SEC Documents, including, without limitation, any Investigational New
Drug Application as required by the United States Food and Drug Administration (“FDA”) or authorizations issued by
Regulatory Authorities; (b) ProQR and each such subsidiary are in compliance in all material respects with the requirements of the
Regulatory Permits, and all of the Regulatory Permits are valid and in full force and effect, in each case in all material respects; (c) ProQR
has not received any notice of proceedings relating to the revocation, termination, modification or impairment of any of the Regulatory
Permits; (d) neither ProQR nor any such subsidiary has failed to file with the FDA or any other Regulatory Authority any material
required application, submission, report, document, notice, supplement or amendment, and all such filings were in material compliance
with applicable laws when filed and have been supplemented as necessary to remain in material compliance with applicable laws; and (e) no
material deficiencies have been asserted by the FDA or any other Regulatory Authority with respect to any such filings; except, in each
case ((a)-(e)), as would not, individually or in the aggregate, reasonably be expected to be material to ProQR.

 

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2.24            Related-Party
Transactions. The SEC Documents disclose all transactions between ProQR and any related parties
as required to be disclosed in the Form 20-F.

 

2.25            Subsidiaries.
All of the issued and outstanding share capital of each Person, of which ProQR owns a majority of its outstanding voting equity securities
are, where applicable, validly issued, fully paid, non-assessable and free of preemptive and similar rights to subscribe for or purchase
securities. Other than the Persons listed on Exhibit 8.1 to the Form 20-F, ProQR does not currently own or control, directly
or indirectly, any interest in any other corporation, partnership, trust, joint venture, limited liability company, association, or other
business entity. Except as disclosed in the SEC Reports, ProQR is not a participant in any material joint venture, partnership or similar
arrangement.

 

Article 3

 

REPRESENTATIONS AND WARRANTIES OF LILLY

 

Except as otherwise specifically
contemplated by this Agreement, Lilly hereby represents and warrants as of the Execution Date and Closing Date to ProQR that:

 

3.1            Authorization;
Enforcement. Lilly is a corporation duly organized, validly existing and in good standing
under the laws of Indiana. Lilly has all requisite corporate power and authority to enter into this Agreement and to consummate the transactions
contemplated hereby. Lilly has taken all necessary corporate action to authorize the execution, delivery and performance of this Agreement.
Upon the execution and delivery of this Agreement, this Agreement will constitute a valid and binding obligation of Lilly, enforceable
against Lilly in accordance with its terms, except (a) as limited by applicable bankruptcy, insolvency, reorganization, moratorium
or other laws of general application affecting enforcement of creditors’ rights and (b) general principles of equity that restrict
the availability of equitable remedies.

 

3.2            No
Conflicts; Government Consents. (a) The execution, delivery and performance of this
Agreement by Lilly and the consummation by Lilly of the transactions contemplated hereby (including the purchase of the Shares) will not
(i) conflict with or result in a violation of any provision of Lilly’s amended articles of incorporation or amended bylaws,
(ii) materially violate or conflict with, or result in a material breach of any provision of, or constitute a default under, any
agreement, indenture or instrument to which Lilly is a party, or (iii) result in a material violation of any law, rule, regulation,
order, judgment or decree (including United States federal and state securities laws and regulations and regulations of any self-regulatory
organizations) applicable to Lilly.

 

(b)            Lilly
is not required to obtain any consent, authorization or order of, or make any filing or registration with, any Governmental Authority
in order for it to execute, deliver or perform any of its obligations under this Agreement in accordance with the terms hereof, or to
purchase the Shares in accordance with the terms hereof.

 

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3.3            Investment
Purpose; Investment Experience. Lilly is purchasing the Shares for its own account and not
with a present view toward the public distribution thereof and has no arrangement or understanding with any other Persons regarding the
distribution of the Shares. Lilly will not, directly or indirectly, offer, sell, pledge, transfer or otherwise dispose of (or solicit
any offers to buy, purchase or otherwise acquire or take a pledge of) any of the Shares except in accordance with the Securities Act
and to the extent permitted by Sections 4.2 and 4.3 of this Agreement. Lilly is an “accredited investor” (as
defined in Regulation D under the Securities Act). Lilly has conducted its own due diligence on ProQR to its satisfaction and has such
knowledge and experience in financial or business matters that it is capable of evaluating the merits and risks of the investment in
the Shares to be purchased hereunder.

 

3.4            Reliance
on Exemptions. Lilly has not taken any of the actions set forth in, and is not subject to,
the disqualification provisions of Rule 506(d)(1) under the Securities Act and did not learn of the investment in the Shares
as a result of any general solicitation or advertising. As of the Execution Date and immediately prior to the Closing, neither Lilly nor
any of its controlled Affiliates beneficially owns, or will beneficially own any securities of ProQR. Lilly understands that ProQR intends
for the Shares to be offered and sold to it in reliance upon specific exemptions from the registration requirements of United States federal
and state securities laws and that ProQR is relying upon the truth and accuracy of, and Lilly’s compliance with, the representations,
warranties, agreements, acknowledgments and understandings of Lilly set forth herein (including in this Section 3.4) in order
to determine the availability of such exemptions and the eligibility of Lilly to acquire the Shares. For the avoidance of doubt, the representation
and warranty set forth in the second sentence of this Section 3.4 shall be deemed not to apply to (i) investment funds
or (ii) pension or other employee benefit plan administrator for any pension or other employee benefit plan for Lilly’s or
its Affiliates’ employees that, in the case of (i) and (ii) are not directed by Lilly, are conducted without the intent
or objective of effecting a change of control of ProQR or otherwise influencing the management or policy of ProQR.

 

3.5            Governmental
Review. Lilly understands that no United States federal or state agency or any other Governmental
Authority has passed upon or made any recommendation or endorsement of the Shares or an investment therein.

 

Article 4

 

COVENANTS AND AGREEMENTS

 

4.1            Market
Listing. From the Execution Date through the Closing, ProQR shall use best efforts to (i) maintain
the listing and trading of the Ordinary Shares on Nasdaq and (ii) effect the listing of the Shares on Nasdaq.

 

4.2            Transfer
or Resale. Lilly understands that:

 

(a)            the
Shares have not been and are not being registered under the Securities Act or any applicable state securities laws and, consequently,
Lilly may have to bear the risk of owning the Shares for an indefinite period of time because the Shares may not be transferred unless
(i) the resale of the Shares is registered pursuant to an effective registration statement under the Securities Act; (ii) Lilly
has delivered to ProQR an opinion of counsel (in form, substance and scope customary for opinions of counsel in comparable transactions)
to the effect that the Shares to be sold or transferred may be sold or transferred pursuant to an exemption from such registration; or
(iii) the Shares are sold or transferred pursuant to Rule 144 (provided, that Lilly provides ProQR with reasonable assurances
(including in the form of seller and broker representation letters) that the Shares may be sold pursuant to such rule).

 

    11 

     

    

 

(b)            any
sale of the Shares made in reliance on Rule 144 may be made only in accordance with the terms of Rule 144 and, if Rule 144
is not applicable, any resale of the Shares under circumstances in which the seller (or the Person through whom the sale is made) may
be deemed to be an underwriter (as that term is defined in the Securities Act) may require compliance with some other exemption under
the Securities Act.

 

(c)            ProQR
acknowledges and agrees that the combination of Lilly’s purchase of the Shares pursuant to this Agreement and its rights pursuant
to the Collaboration Agreement, taken alone and assuming no further acquisitions of Ordinary Shares by Lilly or any Affiliates of Lilly
or other changes to the relationship of the Parties, do not result in Lilly’s being an affiliate of ProQR for purposes of Rule 144
and that ProQR shall direct its transfer agent to remove the restrictive legends referred to in Section 4.3 hereof (and any
stop-transfer orders placed against the transfer of the Shares) no later than the first anniversary of the Closing Date. Subject to the
accuracy of the representations and warranties made by Lilly in Article 3, other than the restrictive legends referred to
in Section 4.3 hereof, ProQR may not place any other legends on the Shares.

 

(d)            For
as long as Lilly or any of its Affiliates beneficially owns any Shares, to the extent it shall be required to do so under the Exchange
Act, ProQR shall use reasonably best efforts to timely file the reports required to be filed by it under the Exchange Act or the Securities
Act (including reports under Sections 13 and 15(d) of the Exchange Act referred to in subparagraph (c)(1) of Rule 144),
and shall use reasonable efforts to take such further necessary action as Lilly may reasonably request in connection with the removal
of any restrictive legend on the Shares being sold, all to the extent required from time to time to enable Lilly to sell the Shares without
registration under the Securities Act within the limitations of the exemption provided by Rule 144.

 

4.3            Legends.
Lilly understands that the Shares will bear restrictive legends in substantially the following form (and a stop-transfer order may be
placed against transfer of the Shares):

 

THESE SECURITIES HAVE NOT
BEEN REGISTERED UNDER THE SECURITIES ACT OF 1933, AS AMENDED (THE “SECURITIES ACT”), AND MAY NOT BE SOLD, OFFERED
FOR SALE, PLEDGED OR HYPOTHECATED IN THE ABSENCE OF A REGISTRATION STATEMENT IN EFFECT WITH RESPECT TO THE SECURITIES UNDER THE SECURITIES
ACT OR AN OPINION OF COUNSEL (WHICH COUNSEL SHALL BE REASONABLY SATISFACTORY TO PROQR THERAPEUTICS N.V.) THAT SUCH REGISTRATION IS NOT
REQUIRED OR UNLESS SOLD PURSUANT TO RULE 144 OF THE SECURITIES ACT.

 

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If such Shares are transferred
pursuant to Sections 4.2 of this Agreement, Lilly may request that ProQR remove, and if so requested, ProQR shall agree to authorize
and instruct (including by causing any required legal opinion to be provided) the removal of any legend from the Shares, if permitted
by applicable securities law, within two (2) Business Days of any such request; provided, however, each Party will
be responsible for any fees it incurs in connection with such request and removal.

 

4.4            Registration
Rights. ProQR hereby provides Lilly with the registration rights set forth on Appendix
2 attached hereto, which is hereby incorporated in and made a part of this Agreement as if set forth in full herein.

 

4.5            Information
Rights. (a) Until Lilly no longer holds Shares representing beneficial ownership of
at least five percent (5%) of the outstanding Ordinary Shares, Lilly shall be entitled to consult with ProQR’s Chief Executive Officer,
who shall make himself or herself available quarterly on a reasonable basis and who shall respond to the reasonable information and other
requests made by Lilly on a quarterly basis with respect to ProQR’s business and financial matters.

 

(b)            Without
limiting any other obligations of confidentiality that Lilly has to ProQR under the Collaboration Agreement or otherwise, Lilly agrees
that it will keep confidential and will not disclose, divulge or use for any purpose (other than to monitor its investment in ProQR pursuant
to this Agreement) any confidential information obtained from ProQR pursuant to the terms of this Agreement, including Section 4.5(a) of
this Agreement, unless such confidential information is known or becomes generally known to the public in general (other than as a result
of a breach of this Section 4.5(b) of this Agreement). In addition, Lilly understands and acknowledges that the securities
laws of the United States and of the Netherlands restrict any person who has material, non-public information about a company from purchasing
or selling any securities of such company while in possession of such information, to the extent material, non-public information about
ProQR is conveyed to Lilly.

 

4.6            Right
to Conduct Activities. ProQR hereby agrees and acknowledges that Lilly is a public company
with numerous business lines (the “Existing Lilly Business”) and an active investment and acquisition program. ProQR
hereby agrees that none of Lilly or any of its Affiliates (together, the “Lilly Group”) shall be liable to ProQR or
any of its Affiliates for any claim arising out of, or based upon, (a) the investment by the Lilly Group in any entity competitive
with ProQR, (b) actions taken by any partner, officer or other representative of the Lilly Group to assist any such competitive company,
whether or not such action was taken as a member of the board of directors of such competitive company or otherwise, and whether or not
such action has a detrimental effect on ProQR, or (c) with respect to the Lilly Group, the Lilly Group engaging in Existing Lilly
Business; provided, however, that the foregoing shall not limit any of Lilly’s or any of its Affiliates’ obligations
under this Agreement or the Collaboration Agreement or otherwise relieve Lilly or any Affiliate of Lilly from liability associated with
the breach by Lilly of any representation, warranty, covenant, agreement or obligation set forth in this Agreement or the Collaboration
Agreement, including (for the avoidance of doubt) Lilly’s obligations of confidentiality and non-use under this Agreement and the
Collaboration Agreement.

 

    13 

     

    

 

4.7            Securities
Law Disclosure; Publicity. No public release, public disclosure or announcement concerning
the transactions contemplated hereby shall be made by either Party without the consent of the other Party (which consent shall not be
unreasonably withheld, conditioned or delayed), except as provided for in the Collaboration Agreement.

 

4.8            Use
of Proceeds. The proceeds to be received by ProQR at the Closing shall be used for general
corporate purposes at the direction of the Board.

 

4.9            Participation
in Future Financings. For so long as Lilly holds one hundred percent (100%) of the Shares
purchased by it pursuant to this Agreement and such Shares equal or exceed at least five percent (5%) of ProQR’s outstanding Ordinary
Shares, ProQR will use its commercially reasonable efforts to allow Lilly to participate (pro rata with its percentage ownership of the
outstanding Ordinary Shares) in public offerings or private placements of its Ordinary Shares to financial, non-strategic institutional
investors primarily for capital raising purposes, subject to any limitations (a) imposed by ProQR’s underwriters or investment
bankers or (b) arising under securities or other applicable laws, including, for the avoidance of doubt, the laws of the Netherlands.
ProQR may undertake such commercially reasonable efforts by notifying Lilly of the proposed financing transaction or instructing its underwriters,
investment bankers or other financial advisors (as applicable) to do so. If such participation is in the form of a public offering, Lilly
understands and acknowledges that ProQR and/or its underwriters or investment bankers may utilize customary “wall-cross” procedures
to notify Lilly of such opportunity to participate in such offering, or alternatively notify Lilly after initiation of such offering has
been publicly disclosed. If such offering is in the form of a private placement, ProQR may notify Lilly prior to the public disclosure
of such private placement utilizing customary “wall-cross” procedures of such opportunity to participate in such private placement.
Notwithstanding the foregoing, in the event, despite ProQR’s commercially reasonable efforts, such as in the event Lilly declines
to receive such information on a “wall-cross” basis, and Lilly is not provided the opportunity to participate in private placements
referenced in this Section 4.9, ProQR will arrange, as promptly as possible thereafter, to permit Lilly to participate in
a separate and subsequent private placement on substantially the same terms. Notwithstanding the foregoing, the opportunity for Lilly
to participate in the financings described in this Section 4.9 shall not apply to (i) “at-the-market” offerings
as defined in Rule 415(a)(4) promulgated under the 1933 Act; (ii) commercial debt in the form of customary credit facilities,
convertible debt, venture debt or similar transactions, provided that such transactions are primarily structured and issued as debt instruments
(regardless of whether such transactions include equity or equity-linked features such as warrants or units to purchase Ordinary Shares
or other instruments exercisable for or convertible into Ordinary Shares); and provided, further, the exception in this clause (ii) shall
not apply to public offerings of debt conducted by ProQR, for which offerings ProQR may utilize the procedures described in the second
and third sentences of this Section 4.9 above to invite Lilly’s participation on the same terms and conditions as the other
purchasers in such public debt offerings; or (iii) strategic partnerships, joint ventures, licenses, collaborations or similar transactions.

 

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4.10            Lockup.
During the period commencing on the Closing Date and until the earlier of (a) the date that is six (6) months after the Closing
Date and (b) the date that the Collaboration Agreement is terminated as provided in Article 13 thereof (collectively, the “Lockup
Period”), without the prior approval of ProQR, Lilly shall not Dispose of (x) any of the Shares, together with any Ordinary
Shares issued in respect thereof as a result of any stock split, stock dividend, share exchange, merger, consolidation or similar recapitalization,
and (y) any Ordinary Shares issued as (or issuable upon the exercise of any warrant, right or other security that is issued as) a
dividend or other distribution with respect to, or in exchange or in replacement of, the Ordinary Shares described in clause (x) of
this sentence (collectively, “Lockup Shares”); provided, however, that the foregoing shall not prohibit (a) Lilly
from transferring any Lockup Shares to (i) a Permitted Transferee (provided, that the Permitted Transferee agrees to be bound in
writing by the restrictions set forth herein), or (ii) to ProQR; (b) the Disposition of Lockup Shares with the prior written
consent of ProQR; and (c) the Disposition of Lockup Shares pursuant to a Third Party Tender/Exchange Offer, and any Disposition effected
pursuant to any business combination, merger, consolidation or similar transaction consummated by ProQR; provided, further, that, in the
event ProQR enters into any definitive agreement with a Third Party during the Lock-Up Period contemplating (x) a Third Party Tender/Exchange
Offer or (y) a business combination, merger, consolidation or similar transaction to which ProQR is a constituent corporation, then
the restrictions on the Lock-Up Shares automatically shall be terminated and of no further force or effect.

 

4.11            Notice
on Sale. Prior to the proposed sale of any Shares by Lilly in open market transactions that
exceed 20% of the daily average trading volume of the Ordinary Shares over the 30 trading days immediately preceding such sale, Lilly
agrees to use commercially reasonable efforts to notify ProQR of such proposed sale.

 

Article 5

 

CONDITIONS TO CLOSING

 

5.1            Mutual
Conditions to Closing. The obligations of ProQR and Lilly to consummate the Closing are subject
to the satisfaction or waiver of the following conditions at or prior to the Closing:

 

(a)            Absence
of Litigation. No proceeding challenging this Agreement or the transactions contemplated hereby, or seeking to prohibit, alter, prevent
or materially delay the Closing, will have been instituted or be pending before any Governmental Authority.

 

(b)            No
Governmental Prohibition. The sale of the Shares by ProQR and the purchase of the Shares by Lilly will not be prohibited by any applicable
law or Governmental Authority.

 

(c)            Collaboration
Agreement. The Collaboration Agreement shall be in full force and effect.

 

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5.2           Conditions
to Obligations of ProQR to Close. ProQR’s obligation to complete the purchase and sale
of the Shares and deliver the Shares to Lilly is subject to the satisfaction or waiver of the following conditions at or prior to the
Closing:

 

(a)            Receipt
of Funds. ProQR will have received immediately available funds in the full amount of the Purchase Price for the Shares.

 

(b)            Representations
and Warranties. The representations and warranties made by Lilly in Article 3 will be true and correct as of the Closing
Date, except to the extent such representations and warranties are specifically made as of a particular date, in which case such representations
and warranties will be true and correct as of such other date, except in each case where the failure of such representations and warranties
to be so true and correct (without giving effect to any limitation as to “materiality” set forth therein) would not reasonably
be expected to have a material adverse effect on Lilly’s ability to perform its obligations hereunder or consummate the transactions
contemplated hereby.

 

(c)            Covenants.
All covenants and agreements contained in this Agreement to be performed or complied with by Lilly on or prior to the Closing Date shall
have been performed or complied with in all material respects.

 

(d)            Closing
Deliverables. All Closing deliverables required to be delivered by Lilly to ProQR under Section 1.3(c) of this Agreement
shall have been so delivered.

 

5.3           Conditions
to Lilly’s Obligations to Close. Lilly’s obligation to complete the purchase
and sale of the Shares is subject to the satisfaction or waiver of the following conditions at or prior to the Closing:

 

(a)            Representations
and Warranties. The representations and warranties made by ProQR in Article 2 of this Agreement will be true and correct
as of the Closing Date, except to the extent such representations and warranties are specifically made as of a particular date, in which
case such representations and warranties will be true and correct as of such other date, except in each case where the failure of such
representations and warranties to be so true and correct (without giving effect to any limitation as to “materiality” set
forth therein) would not reasonably be expected to have a material adverse effect on ProQR’s ability to perform its obligations
hereunder or consummate the transactions contemplated hereby.

 

(b)            Covenants.
All covenants and agreements contained in this Agreement to be performed or complied with by ProQR on or prior to the Closing Date shall
have been performed or complied with in all material respects.

 

(c)            Closing
Deliverables. All Closing deliverables as required to be delivered by ProQR to Lilly under Section 1.3(b) and Section 1.3(d) of
this Agreement shall have been so delivered.

 

(d)            Nasdaq
Matters.

 

(i)            Prior
to the Closing, ProQR shall have taken all actions that are reasonably necessary, including providing appropriate notice to Nasdaq of
the transactions contemplated by this Agreement, for the Shares to be listed on Nasdaq and shall have complied with all listing, reporting,
filing and other obligations under the rules of Nasdaq and of the SEC with respect to the matters contemplated by this Agreement.

 

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(ii)           The
Ordinary Shares shall not have been suspended, as of the Closing Date, by the SEC or Nasdaq from trading on Nasdaq nor shall any such
suspension by the SEC or Nasdaq have been threatened, as of the Closing Date, in writing by the SEC or Nasdaq.

 

(e)            No
Material Adverse Effect. Since the Execution Date, there shall not have been any change, development, occurrence or event that has
had or would reasonably be expected to have a Material Adverse Effect on ProQR.

 

Article 6

 

TERMINATION

 

6.1           Ability
to Terminate. This Agreement may be terminated prior to the Closing by:

 

(a)            mutual
written consent of ProQR and Lilly;

 

(b)            either
ProQR or Lilly, upon written notice to the other, if any of the conditions to the Closing set forth in Section 5.1 of this
Agreement shall have become incapable of fulfillment by the Termination Date and such conditions shall not have been waived by the other
Party within ten (10) Business Days after receipt of notice of an intention to terminate pursuant to this Section 6.1(b) of
this Agreement; provided, however, that the right to terminate this Agreement under this Section 6.1(b) shall
not be available to any Party whose failure to fulfill any obligation under this Agreement or the Collaboration Agreement has been the
cause of, or resulted in, the failure to consummate the transactions contemplated hereby prior to the Termination Date;

 

(c)            ProQR,
upon written notice to Lilly, so long as ProQR is not then in breach of its representations, warranties, covenants or agreements under
this Agreement such that any of the conditions set forth in Section 5.3(a) or (b), as applicable, could not be
satisfied by the Termination Date, (i) upon a breach of any covenant or agreement on the part of Lilly set forth in this Agreement
or (ii) if any representation or warranty of Lilly shall have been or become untrue, in each case such that any of the conditions
set forth in Section 5.2(b) or (c), as applicable, could not be satisfied by the Termination Date; and

 

(d)            Lilly,
upon written notice to ProQR, so long as Lilly is not then in breach of its representations, warranties, covenants or agreements under
this Agreement such that any of the conditions set forth in Section 5.2(b) or (c), as applicable, could not be
satisfied by the Termination Date, as applicable, could not be satisfied by the Termination Date, (i) upon a breach of any covenant
or agreement on the part of ProQR set forth in this Agreement or (ii) if any representation or warranty of ProQR shall have been
or become untrue, in each case such that any of the conditions set forth in Section 5.3(a) or (b), as applicable,
could not be satisfied by the Termination Date.

 

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6.2           Effect
of Termination. In the event of the termination of this Agreement pursuant to Section 6.1,
(a) this Agreement (except for this Section 6.2 and Article 7, and any definitions set forth in this Agreement
and used in this Section 6.2 or Article 7) shall forthwith become void and have no effect, without any liability
on the part of either Party, and (b) all filings, applications and other submissions made pursuant to this Agreement, to the extent
practicable, shall be withdrawn from the agency or other Person to which they were made or appropriately amended to reflect the termination
of the transactions contemplated hereby; provided, however, that nothing contained in this Section 6.2 shall
relieve either Party from liability for fraud or any intentional or willful breach of this Agreement or the Collaboration Agreement.

 

Article 7

 

MISCELLANEOUS

 

7.1           Entire
Agreement; Amendment. This Agreement, together with the Collaboration Agreement, sets forth
the complete, final and exclusive agreement and all the covenants, promises, agreements, warranties, representations, conditions and
understandings between the Parties with respect to the subject matter hereof and supersedes, as of the Execution Date, all prior and
contemporaneous agreements and understandings between the Parties with respect to the subject matter hereof. No subsequent alteration,
amendment, change or addition to this Agreement shall be binding upon the Parties unless reduced to writing and signed by an authorized
officer of each Party.

 

7.2           Survival.
The representations, warranties, covenants and agreements contained in this Agreement shall survive the Closing of the transactions contemplated
by this Agreement.

 

7.3           Notice.
Any notice required or permitted to be given by this Agreement must be in writing, in English. Any and all notices or other communications
or deliveries required or permitted to be provided hereunder must be in writing and will be deemed given and effective if (a) delivered
by hand or by overnight courier with tracking capabilities; (b) mailed postage prepaid by first class, registered or certified mail;
or (c) delivered by electronic mail, in each case, addressed as set forth below unless changed by notice so given:

 

If to ProQR:

 

ProQR Therapeutics N.V.

Zernikedreef 9

2333 CK Leiden, The Netherlands

Attn: Business Development

E-mail: AMorris@proqr.com

 

with a copy (which shall not constitute notice) to:

 

ProQR Therapeutics N.V.

Zernikedreef 9

2333 CK Leiden, the Netherlands

Attn: Legal Department

Email: legal@proqr.com

 

    18 

     

    

 

If to Lilly:

 

Eli Lilly and Company

Lilly Corporate Center

Indianapolis, Indiana 46285

Attn: Senior Vice President, Corporate Business Development

Fax: (317) 651-3051

E-mail: custer_kenneth_1@lilly.com

 

with a copy (which shall not constitute notice) to:

 

Eli Lilly and Company

Lilly Corporate Center

Indianapolis, IN 46285

Attn: General Counsel

Fax: (317) 433-3000

E-mail: hakim_anat@lilly.com

 

7.4           Severability.
If, for any reason, any part of this Agreement is adjudicated invalid, unenforceable or illegal by a court of competent jurisdiction,
(a) such adjudication shall not, to the extent feasible, affect or impair, in whole or in part, the validity, enforceability or
legality of any remaining portions of this Agreement, (b) this Agreement shall be construed and enforced as if such invalid, unenforceable
or illegal provision had never comprised a part hereof, (c) all remaining portions will remain in full force and effect and shall
not be affected by the invalid, unenforceable or illegal provision or by its severance herefrom, and (d) in lieu of such invalid,
unenforceable or illegal provision, the Parties shall use reasonable efforts to seek and agree on an alternative valid and enforceable
provision that preserves the original purpose and intent of this Agreement.

 

7.5           Successors
and Assigns. This Agreement is binding upon and inures to the benefit of the Parties and
their respective successors and permitted assigns. Except for an assignment by Lilly of this Agreement or any rights hereunder to a Permitted
Transferee (which assignment will not relieve Lilly of any obligation hereunder), neither Party may assign this Agreement or any rights
or obligations hereunder without the prior written consent of the other Party.

 

7.6           Waivers.
The failure of a Party to insist upon strict performance of any provision of this Agreement or to exercise any right arising out of this
Agreement shall neither impair that provision or right nor constitute a waiver of that provision or right, in whole or in part, in that
instance or in any other instance. Any waiver by a Party of a particular provision or right shall be in writing, shall be as to a particular
matter and, if applicable, for a particular period of time and shall be signed by such Party.

 

7.7           Interpretation.
The captions and headings to this Agreement are for convenience only and are to be of no force or effect in construing or interpreting
any of the provisions of this Agreement. Unless specified to the contrary, references to Articles, Sections or Appendices mean the particular
Articles, Sections or Appendices to this Agreement. Unless the context otherwise clearly requires, whenever used in this Agreement: (a) the
words “include” or “including” shall be construed as incorporating, also, “but not limited to” or
 “without limitation”; (b) the word “day” or “year” means a calendar day or year unless otherwise
specified; (c) the word “notice” means notice in writing (whether or not specifically stated) and shall include notices,
consents, approvals and other written communications contemplated under this Agreement; (d) the words “hereof,” “herein,”
 “hereby” and derivative or similar words refer to this Agreement as a whole and not merely to the particular provision in
which such words appear; (e) the words “shall” and “will” have interchangeable meanings for purposes of
this Agreement; (f) provisions that require that a Party or the Parties “agree,” “consent” or “approve”
or the like shall require that such agreement, consent or approval be specific and in writing, whether by written agreement, letter,
approved minutes or otherwise; (g) words of any gender include the other gender; (h) words using the singular or plural number
also include the plural or singular number, respectively; (i) references to any specific law, rule or regulation, or article,
section or other division thereof, shall be deemed to include the then-current amendments thereto or any replacement law, rule or
regulation thereof; and (j) neither Party shall be deemed to be acting on behalf of the other Party.

 

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7.8           Counterparts;
Electronic Signatures. This Agreement may be executed in any number of counterparts, each
of which is deemed an original, but all of which together constitute one instrument. This Agreement may be executed and delivered electronically
and upon such delivery such electronic signature will be deemed to have the same effect as if the original signature had been delivered
to the other Party.

 

7.9           Expenses.
Each Party shall pay its own costs, charges and expenses incurred in connection with the negotiation, preparation, execution and performance
of this Agreement.

 

7.10         Further
Assurances. The Parties hereby covenant and agree, without the necessity of any further
consideration, to execute, acknowledge and deliver any and all documents and take any action as may be reasonably necessary to carry
out the intent and purposes of this Agreement.

 

7.11         No
Third Party Beneficiary Rights. This Agreement is not intended to and shall not be construed
to give any Third Party any interest or rights (including any Third Party beneficiary rights) with respect to or in connection with any
agreement or provision contained herein or contemplated hereby.

 

7.12         Construction.
The Parties acknowledge and agree that (a) each Party and its counsel reviewed and negotiated the terms and provisions of this Agreement
and have contributed to its revision; (b) the rule of construction to the effect that any ambiguities are resolved against
the drafting Party shall not be employed in the interpretation of this Agreement; and (c) the terms and provisions of this Agreement
shall be construed fairly as to each Party and not in a favor of or against either Party, regardless of which Party was generally responsible
for the preparation of this Agreement.

 

7.13         Governing
Law; Jurisdiction; Waiver of Jury Trial. This Agreement shall be governed by and construed
in accordance with the laws of the State of New York, without regard to the conflict of laws principles thereof that would require the
application of the laws of any other jurisdiction. Any action brought under or arising out of this Agreement shall be brought in the
federal courts of the United States of America located in the City and County of New York, Borough of Manhattan or (ii) the courts
of the State of New York located in the City and County of New York, Borough of Manhattan (collectively, the “Specified Courts”).
Each Party hereby irrevocably submits to the exclusive jurisdiction of such Specified Courts in respect of any claim relating to the
validity, interpretation and enforcement of this Agreement and hereby waives, and agrees not to assert, as a defense in any action, suit
or proceeding in which any such claim is made that it is not subject to the jurisdiction of such court or that such action, suit or proceeding
may not be brought or is not maintainable in such court, or that the venue thereof may not be appropriate or that this agreement may
not be enforced in or by such court. Each Party hereby consents to and grants the Specified Courts jurisdiction over such Party and over
the subject matter of any such claim and agree that mailing of process or other papers in connection with any such action, suit or proceeding
in the manner provided in Section 7.3 of this Agreement or in such other manner as may be permitted by law, shall be valid
and sufficient. Each of ProQR and Lilly hereby irrevocably waives, to the fullest extent permitted by applicable law, any and all right
to trial by jury in any legal proceeding arising out of or relating to this Agreement or the transactions contemplated hereby.

 

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7.14         Equitable
Relief. Each Party acknowledges and agrees that if it fails to perform any of its covenants
or agreements or discharge any of its obligations under this Agreement, irreparable damage could occur and any remedy at law may prove
to be inadequate relief for the other Party. Accordingly, notwithstanding anything herein to the contrary, each Party shall be entitled
(without any requirement to post bond) to seek injunctive relief and specific performance (including any relief or recovery under this
Agreement) in any court of competent jurisdiction anywhere in the world (including the court designated in Section 7.13 of
this Agreement).

 

7.15         Cumulative
Remedies. No remedy referred to in this Agreement is intended to be exclusive unless explicitly
stated to be so, but each shall be cumulative and in addition to any other remedy referred to in this Agreement or otherwise available
under law.

 

[Remainder of page intentionally left
blank.]

 

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In
Witness Whereof, the Parties have caused this Agreement to be executed as of the Execution Date by their duly authorized representatives.

 

	 	PROQR THERAPEUTICS N.V.
	 	 
	 	By:	/s/ Daniel de Boer
	 	 	Name: Daniel de Boer
	 	 	Title: Chief Executive Officer
	 	 	 
	 	 	 
	 	ELI LILLY AND COMPANY
	 	 
	 	By:	/s/ David A. Ricks
	 	 	Name: David A. Ricks
	 	 	Title: Chairman and Chief Executive Officer

 

[Signature page to Share Purchase Agreement]

 

    

     

    

 

APPENDIX 1

 

DEFINED TERMS

 

“Affiliate”
means, with respect to any Person, any entity that, at the relevant time (whether as of the Execution Date or thereafter), directly or
indirectly through one or more intermediaries, controls, is controlled by, or is under common control with such Person, for so long as
such control exists. For purposes of this definition, “control” means (i) to possess, directly or indirectly, the power
to direct or cause the direction of the management or policies of an entity, whether through ownership of voting securities or by contract
relating to voting rights or corporate governance, or (ii) direct or indirect ownership of 50% (or such lesser percentage that is
the maximum allowed to be owned by a foreign entity in a particular jurisdiction) or more of the voting share capital or other equity
interest in such entity.

 

“Board”
means the supervisory board of directors of ProQR.

 

“Business Day”
means any day, other than any Saturday, Sunday or any day that banks are authorized or required to be closed in Amsterdam, The Netherlands
or Indianapolis, Indiana.

 

“Contract”
means, with respect to any Person, any legally binding written or oral contracts, agreements, indentures, bonds, loans, leases, subleases,
licenses, sublicenses, instruments, notes and arrangements to which such Person is a party or by which any of its properties or assets
are subject.

 

“Disposition”
or “Dispose of” means (a) pledge, sale, contract to sell, sale of any option or Contract to purchase, purchase
of any option or Contract to sell, grant of any option, right or warrant for the sale of, or other disposition of or transfer of any
Ordinary Shares, or any options, warrants or other securities or rights convertible into or exercisable or exchangeable for, Ordinary
Shares, including, without limitation, any “short sale” or similar arrangement, or (b) swap, hedge, derivative instrument
or any other agreement or any transaction that transfers, in whole or in part, directly or indirectly, the economic consequence of ownership
of the Ordinary Shares, whether any such swap or transaction is to be settled by delivery of securities, in cash or otherwise.

 

“Exchange Act”
means the Securities Exchange Act of 1934, as amended, and the rules and regulations of the SEC thereunder.

 

“Governmental Authority”
means any national, international, federal, state, provincial or local government, or political subdivision thereof, or any multinational
organization or any authority, agency or commission entitled to exercise any administrative, executive, judicial, legislative, police,
regulatory or taxing authority or power, and any court or tribunal (or any department, bureau or division thereof, or any governmental
arbitrator or arbitral body).

 

“IFRS”
means International Financial Reporting Standards.

 

    Appendix 1-1

     

    

 

“Material Adverse
Effect” means any change, event or occurrence that, individually or in the aggregate with any other changes, events or circumstances,
has had or would reasonably be expected to have (i) a material adverse effect on the business, financial condition, assets or results
of operations of ProQR or its subsidiaries, taken as a whole, or (ii) a material adverse effect on ProQR’s ability to perform
its obligations hereunder or consummate the transactions contemplated hereby; provided, however, that in no event shall any of
the following occurring after the date hereof, alone or in combination, be deemed to constitute, or be taken into account in determining
whether a Material Adverse Effect has occurred: (a) changes in ProQR’s industry generally or in conditions in the United States
or global economy or capital or financial markets generally, including changes in interest or exchange rates, (b) any change, event
or occurrence caused by the announcement or pendency of the transactions contemplated hereby or by the Collaboration Agreement (c) the
performance of this Agreement, the Collaboration Agreement and the transactions contemplated hereby and thereby, or any action taken
or omitted to be taken by ProQR at the request or with the prior consent of Lilly, (d) changes in general legal, regulatory, political,
economic or business conditions occurring after the date hereof that, in each case, generally affect the biotechnology or biopharmaceutical
industries, (e) acts of war, sabotage or terrorism occurring after the date hereof, or any escalation or worsening of any such acts
of war, sabotage or terrorism, or (f) earthquakes, hurricanes, floods or other natural disasters occurring after the date hereof;
provided, however, that with respect to clauses (a), (d), (e) and (f), such effects, alone or in combination, may be deemed to constitute,
or be taken into account in determining whether a Material Adverse Effect has occurred, but only to the extent such effects disproportionately
affect ProQR compared to other companies in the biotechnology or biopharmaceutical industries.

 

“Material Contract”
means all Contracts in effect as of the Execution Date that are required to be filed as exhibits to ProQR’s annual report on Form 20-F.

 

“Nasdaq”
means The Nasdaq Stock Market LLC.

 

“Permitted Transferee”
means an Affiliate of Lilly; provided, however, that no such Person shall be deemed a Permitted Transferee for any purpose
under this Agreement unless: (a) the Permitted Transferee shall have agreed in writing to be subject to and bound by the terms of
this Agreement as though it were “Lilly” hereunder, and (b) Lilly acknowledges that it continues to be bound by the
terms of this Agreement.

 

“Person”
means any corporation, limited or general partnership, limited liability company, joint venture, trust, unincorporated association, governmental
body, authority, bureau or agency, any other entity or body, or an individual.

 

“Proportionate Interest”
at any given time shall mean the ratio of (a) the number of Ordinary Shares then owned by Lilly to (b) the total number of
outstanding Ordinary Shares then owned by all shareholders of ProQR.

 

“Qualified Securities”
means any Ordinary Shares or any other securities, warrants, options or rights of any nature that are convertible into or exchangeable
or exercisable for, or otherwise give the holder thereof any rights in respect of (whether or not the right to convert, exchange or exercise
is subject to the passage of time, contingencies or contractual restrictions or any combination thereof) any Ordinary Share, including
those issued in connection with or attached to any debt financing transaction.

 

    Appendix 1-2

     

    

 

“Registrable Securities”
means the Shares; provided, that any Shares will cease to be Registrable Securities when such Shares have been sold or otherwise Disposed
of pursuant to an effective Registration Statement or otherwise.

 

“Register,”
 “Registered” and “Registration” means a registration effected by preparing and filing (a) a
Registration Statement in compliance with the Securities Act (and any post effective amendments filed or required to be filed) and the
declaration or ordering of effectiveness of such Registration Statement, or (b) a Prospectus and/or Prospectus supplement in respect
of an appropriate effective Registration Statement.

 

“Registration Statement”
means a registration statement of ProQR that covers the resale of any Registrable Securities pursuant to the provisions of Appendix 2
filed with, or to be filed with, the SEC under the rules and regulations promulgated under the Securities Act, including the related
Prospectus, amendments and supplements to each such registration statement or Prospectus, including pre- and post-effective amendments,
all exhibits thereto, financial information and all other material incorporated by reference or deemed to be incorporated by reference
in such registration statement.

 

“SEC”
means the United States Securities and Exchange Commission or any successor entity.

 

“Securities Act”
means the Securities Act of 1933, as amended, and the rules and regulations of the SEC thereunder.

 

“Shelf Registration
Statement” means a “shelf” registration statement of ProQR that covers all Registrable Securities on Form F-3
and under Rule 415 under the Securities Act or, if ProQR is not then eligible to file on Form F-3, on another eligible form
under the Securities Act, or any successor rule that may be adopted by the SEC, including without limitation any such registration
statement filed pursuant to Appendix 2 and all amendments and supplements to such “shelf” registration statement, including,
post-effective amendments, in each case, including the Prospectus contained therein, all exhibits thereto and any document incorporated
by reference therein.

 

“Tax”
or “Taxes” shall mean all federal, state, local, and foreign income, excise, gross receipts, gross income, ad valorem,
profits, gains, property, capital, sales, transfer, use, payroll, employment, severance, withholding, duties, intangibles, franchise,
backup withholding, value- added, and other taxes imposed by a Governmental Authority, together with all interest, penalties and additions
to tax imposed with respect thereto.

 

“Tax Return”
shall mean a report, return or other document (including any amendments thereto) required to be supplied to a Governmental Authority
with respect to Taxes.

 

“Third Party”
means any Person other than Lilly or ProQR (or its respective Affiliates).

 

“Third Party Tender/Exchange
Offer” means any tender or exchange offer made to all of the holders of Ordinary Shares by a Third Party (other than a Third
Party acting on behalf of or as part of a group or in concert with Lilly).

 

    Appendix 1-3

     

    

 

“Transaction Documents”
means this Agreement.

 

“Underwriter”
means, with respect any Underwritten Offering, a securities dealer who purchases any Registrable Securities as a principal in connection
with a distribution of such Registrable Securities.

 

“Underwritten Offering”
means a public offering of securities Registered under the Securities Act in which an Underwriter participates in the distribution of
such securities, including on a firm commitment basis for reoffer and resale to the public, including any such offering that is a “bought
deal” or a block trade.

 

    Appendix 1-4

     

    

 

APPENDIX 2

 

Registration
Rights

 

1.             Resale
Registration.

 

1.1            On
or prior to the first (1st) Business Day following the expiration of the Lockup Period, ProQR will file a Shelf Registration Statement
registering for resale the Registrable Securities under the Securities Act. The Company shall use its commercially reasonable efforts
to cause such Shelf Registration Statement to become effective as promptly as practicable after filing. Until the earlier of such time
as (i) all Registrable Securities cease to be Registrable Securities or (ii) ProQR is no longer eligible to maintain a Shelf
Registration Statement, ProQR will keep current and effective such Shelf Registration Statement and file such supplements or amendments
to such Shelf Registration Statement (or file a new Shelf Registration Statement when such preceding Shelf Registration Statement expires
pursuant to the rules of the SEC) as may be necessary or appropriate in order to keep such Shelf Registration Statement continuously
effective and useable for the resale of Registrable Securities under the Securities Act in order to fulfill a Shelf Underwritten Offering
Request (as defined below). The Shelf Registration Statement shall include the Plan of Distribution attached hereto as Annex A.

 

1.2            Lilly
may use the Shelf Registration Statement to dispose of Registrable Securities pursuant to an Underwritten Offering from which it reasonably
expects to receive gross proceeds of at least $15.0 million in the aggregate from such Underwritten Offering. Upon written notice from
Lilly to ProQR of Lilly’s intention to sell Registrable Securities in such manner, ProQR shall, at Lilly’s request (a “Shelf
Underwritten Offering Request”), enter into an underwriting agreement in a form as is customary in Underwritten Offerings of
securities by ProQR with the underwriter or underwriters selected by Lilly and reasonably acceptable to ProQR and shall take all such
other reasonable actions as are requested by the managing underwriter of such Underwritten Offering and/or Lilly in order to expedite
or facilitate the disposition of such Registrable Securities (“Shelf Underwritten Offering”); provided, that in no
event shall ProQR have any obligation to facilitate or participate in more than two (2) Shelf Underwritten Offerings.

 

1.3            If
the filing, initial effectiveness or use of the Shelf Registration Statement at any time would require ProQR to make a public disclosure
of material non-public information that ProQR has a bona fide business purpose, in good faith, for not disclosing publicly at such time,
ProQR may, upon giving prompt written notice of such action to Lilly, delay the filing or initial effectiveness of, or suspend use of,
the Shelf Registration Statement (a “Suspension”). ProQR shall use commercially reasonable efforts to make
routine public disclosures about its business in the ordinary course consistent with past practice and subject to and in compliance with
applicable law. In the case of a Suspension, Lilly agrees to suspend use of the applicable Prospectus in connection with any sale or
purchase, or offer to sell or purchase, Shares, upon receipt of the notice referred to above. The Company shall immediately notify Lilly
in writing upon the termination of any Suspension, amend or supplement the Prospectus, if necessary, so it does not contain any untrue
statement or omission and furnish to Lilly such numbers of copies of the Prospectus as so amended or supplemented as Lilly may reasonably
request. The Company shall, if necessary, supplement or amend the Shelf Registration Statement, if required by law or as may reasonably
be requested by Lilly.

 

    

     

    

 

 

2.             Information.
The Company may require Lilly to furnish to ProQR such information regarding the distribution of the Shares and such other information
relating to Lilly and its ownership of Shares as ProQR may from time to time reasonably request in writing to the extent that such information
is required to be included in the Shelf Registration Statement.

 

3.             Expenses.
All expenses incurred by the parties with respect to each Shelf Underwritten Offering (including, for the avoidance of doubt, expenses
relating to the preparation, filing and effectiveness of the Shelf Registration Statement) shall be borne 50% by ProQR on the one hand
and 50% by Lilly on the other hand, and each party shall account for its own expenses and promptly submit to the other one or more requests
for reimbursement in accordance with such cost-sharing arrangement; provided that in the event that ProQR incurs reasonable and documented
expenses in excess of $75,000, Lilly shall promptly, upon written notice by ProQR, reimburse ProQR for any such excess amounts. The expenses
described in the foregoing sentence shall include (a) all registration and filing fees, and any other fees and expenses associated
with filings required to be made with the SEC or Financial Industry Regulatory Authority, (b) all fees and expenses in connection
with compliance with any securities or “Blue Sky” laws (including reasonable fees and disbursements of counsel for the Underwriters
in connection with blue sky qualifications of the Shares), (c) all printing, duplicating, word processing, messenger, telephone,
facsimile and delivery expenses (including expenses of printing certificates for the Shares in a form eligible for deposit with The Depository
Trust Company and of printing Prospectuses), (d) all fees and disbursements of counsels (including non-U.S. counsels) for ProQR and
of all independent certified public accountants or independent auditors of ProQR and any of its Subsidiaries (including the expenses of
any special audit and comfort letters required by or incident to such performance), (e) Securities Act liability insurance or similar
insurance if ProQR so desires, (f) all fees and expenses incurred in connection with the listing of the Shares on any securities
exchange or quotation of the Shares on any inter-dealer quotation system, and (g) all fees and expenses of any special experts or
other Persons retained by ProQR in connection with any registration. In addition, notwithstanding the foregoing, ProQR shall not be required
to pay any underwriting discounts and commissions and transfer Taxes, if any, attributable to the sale of the Shares.

 

4.             Notice.
The Company shall notify Lilly immediately upon (a) any request by the SEC or any other Federal or state Governmental Authority for
amendments or supplements to a Shelf Registration Statement or for additional information that pertains to Lilly as a selling stockholder;
(b) the issuance by the SEC of any stop order suspending the effectiveness of the Shelf Registration Statement or any order by the
SEC or any other regulatory authority preventing or suspending the use of any Prospectus or the initiation or threatening of any proceedings
for such purposes, (c) receipt by ProQR of any notification with respect to the suspension of the qualification of the Shares for
offering or sale in any jurisdiction or the initiation or threatening of any proceeding for such purpose, or (d) ProQR becoming aware
that the Shelf Registration Statement or the related Prospectus contains any untrue statement of a material fact or omits to state a material
fact necessary to make the statements therein (in the case of such Prospectus, in light of the circumstances under which they were made)
not misleading.

 

     

     

    

 

5.             Indemnification.

 

5.1           To
the extent permitted by Law, ProQR will indemnify and hold harmless Lilly, its officers, directors, agents, partners, members, stockholders
and employees, as applicable, and each Person who controls Lilly (within the meaning of the Securities Act or the Exchange Act), and the
officers, directors, agents, partners, members, stockholders and employees of each such controlling Person, from and against any and all
losses, claims, liabilities, damages, deficiencies, assessments, fines, judgments, fees, costs (including, without limitation, reasonable
costs of preparation and reasonable attorneys’ fees) and expenses (collectively “Losses”) (joint or several),
as incurred, to which they may become subject under the Securities Act, the Exchange Act or other federal or state law, insofar as such
Losses (or actions in respect thereof) arise out of, relate to, or are based upon any of the following statements, omissions or violations
(collectively a “Violation”) by ProQR: (a) any untrue statement or alleged untrue statement of a material
fact contained in the Shelf Registration Statement or incorporated by reference therein, including any Prospectus contained therein or
any amendments or supplements thereto, (b) the omission or alleged omission to state therein a material fact required to be stated
therein, or necessary to make the statements therein not misleading, or (c) any violation or alleged violation by ProQR of the Securities
Act, the Exchange Act, any state securities Law, or any rule or regulation promulgated under the Securities Act, the Exchange Act
or any state securities Law in connection with the Shelf Registration Statement; and ProQR will reimburse each such indemnified party
for any legal or other expenses reasonably incurred by them in connection with investigating or defending any such Loss or action; provided,
however, that the indemnity agreement contained in this Section 5.1 will not apply to amounts paid in settlement of any
such Loss or action if such settlement is effected without ProQR’s consent, nor will ProQR be liable in any such case for any such
Loss to the extent that it arises out of or is based upon a Violation which occurs in reliance upon and in conformity with written information
furnished by Lilly and stated to be expressly for use in connection with the Shelf Registration Statement or an applicable Prospectus.

 

5.2           To
the extent permitted by Law, Lilly will indemnify and hold harmless ProQR and each of its directors and its officers against any Losses
(joint or several) to which ProQR or any such director, officer, controlling Person, Underwriter or other Third Party who may become subject
under the Securities Act, the Exchange Act or other federal or state law, insofar as such Losses (or actions in respect thereto) arise
out of or are based upon any of the following statements: (a) any untrue statement or alleged untrue statement of a material fact
contained in any Registration Statement or any other document incorporated reference therein, including any preliminary Prospectus or
final Prospectus contained therein or any amendments or supplements thereto, or (b) the omission or alleged omission to state therein
a material fact required to be stated therein, or necessary to make the statements therein not misleading (collectively, a “Lilly
Violation”), in each case to the extent (and only to the extent) that such Lilly Violation occurs in reliance upon and in
conformity with written information furnished by Lilly under an instrument duly executed by Lilly, or written information furnished by
Lilly and stated to be expressly for use in connection with the Shelf Registration Statement or an applicable Prospectus; and Lilly will
reimburse any legal or other expenses reasonably incurred by ProQR or any such director, officer, controlling Person, Underwriter or other
Third Party in connection with investigating or defending any such Loss or action if it is judicially determined that there was such a
Lilly Violation; provided, however, that the indemnity agreement contained in this Section 5.2 will not apply
to amounts paid in settlement of any such Loss or action if such settlement is effected without Lilly’s consent; provided,
further, that the obligations of Lilly hereunder shall be limited to an amount equal to the net proceeds it receives in such Registration.

 

     

     

    

 

5.3           Promptly
after receipt by an indemnified party under this Section 5 of notice of the commencement of any action (including any governmental
action), such indemnified party will, if a claim in respect thereof is to be made against any indemnifying party under this Section 5,
deliver to the indemnifying party a written notice of the commencement thereof and the indemnifying party will have the right to participate
in, and, to the extent the indemnifying party so desires, jointly with any other indemnifying party similarly noticed, to assume the defense
thereof with counsel mutually satisfactory to the parties; provided, however, that an indemnified party will have the right
to retain its own counsel, with the fees and expenses thereof to be paid by the indemnifying party, 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
such indemnified party and any other party represented by such counsel in such proceeding. The failure to deliver written notice to the
indemnifying party within a reasonable time of the commencement of any such action will relieve such indemnifying party of any liability
to the indemnified party under this Section 5 to the extent, and only to the extent, prejudicial to its ability to defend
such action, but the omission so to deliver written notice to the indemnifying party will not relieve it of any liability that it may
have to any indemnified party otherwise than under this Section 5.

 

5.4           If
the indemnification provided for in this Section 5 is held by a court of competent jurisdiction to be unavailable to an indemnified
party with respect to any Losses referred to herein, the indemnifying party, in lieu of indemnifying such indemnified party thereunder,
will to the extent permitted by applicable Law contribute to the amount paid or payable by such indemnified party as a result of such
Loss in such proportion as is appropriate to reflect the relative fault of the indemnifying party, on the one hand, and of the indemnified
party, on the other, in connection with the Violation(s) or Lilly Violation(s), as applicable, that resulted in such Loss, as well
as any other relevant equitable considerations. The relative fault of the indemnifying party and of the indemnified party will be determined
by a court of law by reference to, among other things, whether the untrue or alleged untrue statement of a material fact or the omission
to state a material fact relates to information supplied by the indemnifying party or by the indemnified party and the parties’
relative intent, knowledge, access to information and opportunity to correct or prevent such statement or omission; provided, however,
that the obligations of Lilly hereunder shall be limited to an amount equal to the net proceeds it receives in such Registration; and
provided, further, 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.

 

5.5           The
obligations of ProQR and Lilly under this Section 5 will survive termination of this Agreement and the expiration or withdrawal
of the Shelf Registration Statement. No indemnifying party, in the defense of any such claim or litigation, will, 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 to such claim or
litigation.

 

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ANNEX A

 

PLAN OF DISTRIBUTION

 

The selling securityholders,
including their pledgees, donees, transferees, distributees, beneficiaries or other successors in interest, may from time to time offer
some or all of the ordinary shares (collectively, “Securities”) covered by this prospectus. To the extent required,
this prospectus may be amended and supplemented from time to time to describe a specific plan of distribution.

 

The selling securityholders
may sell the Securities covered by this prospectus from time to time, and may also decide not to sell all or any of the Securities that
they are allowed to sell under this prospectus. We will not receive any proceeds from the sale of Securities. The selling securityholders
will act independently of us in making decisions regarding the timing, manner and size of each sale. These dispositions may be at fixed
prices, at market prices prevailing at the time of sale, at prices related to such prevailing market prices, at varying prices determined
at the time of sale, or at privately negotiated prices. Sales may be made by the selling securityholders in one or more types of transactions,
which may include:

 

		·	purchases by underwriters, dealers and agents who may receive compensation in the form of underwriting discounts, concessions or commissions
from the selling securityholders and/or the purchasers of the Securities for whom they may act as agent;

 

		·	one or more block transactions, including transactions in which the broker or dealer so engaged will attempt to sell the Securities
as agent but may position and resell a portion of the block as principal to facilitate the transaction, or in crosses, in which the same
broker acts as an agent on both sides of the trade;

 

		·	ordinary brokerage transactions or transactions in which a broker solicits purchases;

 

		·	purchases by a broker-dealer or market maker, as principal, and resale by the broker-dealer for its account;

 

		·	the pledge of Securities for any loan or obligation, including pledges to brokers or dealers who may from time to time effect distributions
of Securities, and, in the case of any collateral call or default on such loan or obligation, pledges or sales of Securities by such pledgees
or secured parties;

 

		·	short sales or transactions to cover short sales relating to the Securities;

 

		·	one or more exchanges or over the counter market transactions;

 

		·	through distribution by a selling securityholder or its successor in interest to its members, general or limited partners or shareholders
(or their respective members, general or limited partners or shareholders);

 

     

     

    

 

		·	privately negotiated transactions;

 

		·	the writing of options, whether the options are listed on an options exchange or otherwise;

 

		·	distributions to creditors and equity holders of the selling securityholders; and

 

		·	any combination of the foregoing, or any other available means allowable under applicable law.

 

A selling securityholder may
also resell all or a portion of its Securities in open market transactions in reliance upon Rule 144 under the Securities Act of
1933, as amended (the “Securities Act”) provided it meets the criteria and conforms to the requirements of Rule 144
under the Securities Act and all applicable laws and regulations.

 

The selling securityholders
may enter into sale, forward sale and derivative transactions with third parties, or may sell securities not covered by this prospectus
to third parties in privately negotiated transactions. In connection with those sale, forward sale or derivative transactions, the third
parties may sell Securities covered by this prospectus, including in short sale transactions and by issuing securities that are not covered
by this prospectus but are exchangeable for or represent beneficial interests in the ordinary shares. The third parties also may use ordinary
shares received under those sale, forward sale or derivative arrangements or ordinary shares pledged by the selling securityholder or
borrowed from the selling securityholders or others to settle such third-party sales or to close out any related open borrowings of ordinary
shares. The third parties may deliver this prospectus in connection with any such transactions. Any third party in such sale transactions
will be an underwriter and will be identified in a supplement or a post- effective amendment to the registration statement of which this
prospectus is a part, as may be required.

 

In addition, the selling securityholders
may engage in hedging transactions with broker- dealers in connection with distributions of Securities or otherwise. In those transactions,
broker- dealers may engage in short sales of securities in the course of hedging the positions they assume with selling securityholders.
The selling securityholders may also sell securities short and redeliver securities to close out such short positions. The selling securityholders
may also enter into option or other transactions with broker-dealers which require the delivery of securities to the broker-dealer. The
broker-dealer may then resell or otherwise transfer such securities pursuant to this prospectus. The selling securityholders also may
loan or pledge Securities, and the borrower or pledgee may sell or otherwise transfer the Securities so loaned or pledged pursuant to
this prospectus. Such borrower or pledgee also may transfer those Securities to investors in our securities or the selling securityholders’
securities in connection with the offering of other securities not covered by this prospectus.

 

To the extent necessary, the
specific terms of the offering of Securities, including the specific Securities to be sold, the names of the selling securityholders,
the respective purchase prices and public offering prices, the names of any underwriter, broker-dealer or agent, if any, and any applicable
compensation in the form of discounts, concessions or commissions paid to underwriters or agents or paid or allowed to dealers will be
set forth in a supplement to this prospectus or a post-effective amendment to this registration statement of which this prospectus forms
a part. The selling securityholders may, or may authorize underwriters, dealers and agents to, solicit offers from specified institutions
to purchase Securities from the selling securityholders. These sales may be made under “delayed delivery contracts” or other
purchase contracts that provide for payment and delivery on a specified future date. If necessary, any such contracts will be described
and be subject to the conditions set forth in a supplement to this prospectus or a post- effective amendment to this registration statement
of which this prospectus forms a part.

 

     

     

    

 

Broker-dealers or agents may
receive compensation in the form of commissions, discounts or concessions from the selling securityholders. Broker-dealers or agents may
also receive compensation from the purchasers of Securities for whom they act as agents or to whom they sell as principals, or both. Compensation
to a particular broker-dealer might be in excess of customary commissions and will be in amounts to be negotiated in connection with transactions
involving securities. In effecting sales, broker-dealers engaged by the selling securityholders may arrange for other broker-dealers to
participate in the resales.

 

In connection with sales of
Securities covered hereby, the selling securityholders and any underwriter, broker-dealer or agent and any other participating broker-dealer
that executes sales for the selling securityholders may be deemed to be an “underwriter” within the meaning of the Securities
Act. Accordingly, any profits realized by the selling securityholders and any compensation earned by such underwriter, broker-dealer or
agent may be deemed to be underwriting discounts and commissions. Selling securityholders who are “underwriters” under the
Securities Act must deliver this prospectus in the manner required by the Securities Act. This prospectus delivery requirement may be
satisfied through the facilities of the national exchange on which the Securities are then traded in accordance with Rule 153 under
the Securities Act or satisfied in accordance with Rule 174 under the Securities Act.

 

We and the selling securityholders
have agreed to indemnify each other against certain liabilities, including liabilities under the Securities Act. In addition, we or the
selling securityholders may agree to indemnify any underwriters, broker-dealers and agents against or contribute to any payments the underwriters,
broker-dealers or agents may be required to make with respect to, civil liabilities, including liabilities under the Securities Act. Underwriters,
broker- dealers and agents and their affiliates are permitted to be customers of, engage in transactions with, or perform services for
us and our affiliates or the selling securityholders or their affiliates in the ordinary course of business.

 

In order to comply with applicable
securities laws of some states or countries, the Securities may only be sold in those jurisdictions through registered or licensed brokers
or dealers and in compliance with applicable laws and regulations. In addition, in certain states or countries the Securities may not
be sold unless they have been registered or qualified for sale in the applicable state or country or an exemption from the registration
or qualification requirements is available. In addition, any Securities of a selling securityholder covered by this prospectus that qualify
for sale pursuant to Rule 144 under the Securities Act may be sold in open market transactions under Rule 144 rather than pursuant
to this prospectus.

 

In connection with an offering
of Securities under this prospectus, the underwriters may purchase and sell securities in the open market. These transactions may include
short sales, stabilizing transactions and purchases to cover positions created by short sales. Short sales involve the sale by the underwriters
of a greater number of securities than they are required to purchase in an offering. Stabilizing transactions consist of certain bids
or purchases made for the purpose of preventing or retarding a decline in the market price of the securities while an offering is in progress.

 

     

     

    

 

The underwriters also may
impose a penalty bid. This occurs when a particular underwriter repays to the underwriters a portion of the underwriting discount received
by it because the underwriters have repurchased securities sold by or for the account of that underwriter in stabilizing or short-covering
transactions.

 

These activities by the underwriters
may stabilize, maintain or otherwise affect the market price of the Securities offered under this prospectus. As a result, the price of
the Securities may be higher than the price that otherwise might exist in the open market. If these activities are commenced, they may
be discontinued by the underwriters at any time. These transactions may be effected on the Nasdaq Stock Market or another securities exchange
or automated quotation system, or in the over-the-counter market or otherwise.

 

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