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Item 7. Management’s Discussion and Analysis

2026-02-26 compared with 2025-03-04 · 216 added, 351 removed, 1293 unchanged (30% of the section changed)

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Our RNA medicines platform, PRISM ® , combines multiple modalities, chemistry innovation and deep insights into human genetics to deliver scientific breakthroughs that treat both rare and common disorders.

Our toolkit of RNA-targeting modalities includes RNA editing, splicing, silencing using RNA interference (“siRNA") and antisense silencing, providing us with unique capabilities for designing and sustainably delivering candidates that optimally address disease biology.

Our diversified pipeline includes clinical programs in obesity, alpha-1 antitrypsin deficiency (“AATD”), Duchenne muscular dystrophy (“DMD”), and Huntington’s disease (“HD”), as well as several preclinical programs utilizing our versatile RNA medicines platform.

Our toolkit of RNA-targeting modalities, including RNAi (SpiNA) and RNA editing (AIMers), provides us with unmatched capabilities for designing and sustainably delivering candidates that optimally address disease biology.

Our pipeline is focused on our obesity (WVE-007), alpha-1 antitrypsin deficiency (“AATD”) (WVE-006) and PNPLA3 I148M liver disease (WVE-008) programs, and also includes clinical programs for Duchenne muscular dystrophy (“DMD”) and Huntington’s disease (“HD”), as well as several preclinical programs utilizing our versatile RNA medicines platform.

We were founded on the recognition that there was a significant, untapped opportunity to use chemistry innovation to tune the pharmacological properties of oligonucleotides.

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We believe that we have built the most versatile toolkit of RNA-targeting modalities in the industry, with multiple means of repairing, restoring, or reducing proteins and designing best-fit solutions based on the unique biology of a given disease target.

We are actively advancing programs using four distinct modalities, including novel A-to-I RNA editing oligonucleotides (“AIMers”).

These modalities include:

• RNA editing, which uses AIMers that are designed to target single bases on an RNA transcript and recruit endogenous ADAR enzymes that naturally possess the ability to change an adenine (A) to an inosine (I), which cells read as guanine (G).

This approach enables both the correction of G-to-A point mutations and the modulation of RNA to either upregulate protein expression, modify protein-protein interactions, or alter RNA folding and processing.

AIMers are short in length, fully chemically modified, and use our novel chemistry, which make them distinct from other ADAR-mediated editing approaches.

• Antisense (silencing) , which uses our oligonucleotide designed to bind to a specific sequence in a target RNA strand that encodes a disease-associated protein or pathogenic RNA.

The resulting double-stranded molecule (“duplex”) is then recognized by a cellular enzyme called RNase H, which cleaves, or cuts, the target RNA in the duplex, thereby preventing the disease-associated protein from being made.

• RNA interference (RNAi ) (silencing) , which uses our double-stranded RNAs called siRNAs to engage the RNAi machinery known as the RNA-induced silencing complex (“RISC”) and to silence a target RNA that is either pathogenic itself or encodes a disease-associated protein, thereby preventing the accumulation of the pathogenic species (RNA or protein).

• Splicing / exon skipping , which is the processing of a nascent pre-mRNA transcript into mRNA by removing introns and joining exons together.

Exon skipping uses our oligonucleotide designed to bind to a particular sequence within a target pre-mRNA and direct the cellular machinery to alter the final composition of exons in mature mRNA by deleting, or splicing out, certain specific regions of that RNA.

We are actively advancing programs across modalities, including RNA interference (“RNAi”) (silencing), RNA editing, which uses novel A-to-I RNA editing oligonucleotides (“AIMers”), antisense silencing, and splicing.

We have also advanced novel bifunctional modalities designed to silence multiple targets or silence one target while simultaneously editing or upregulating another unique target.

We intentionally focus on targeting the transcriptome using oligonucleotides rather than other nucleic acid modalities such as gene therapy and DNA editing.

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• Leverage well-established industry manufacturing processes and regulatory, access, and reimbursement pathways.

We have a robust and diverse pipeline of potential first-or best-in-class programs addressing both rare and common diseases:

• GalNAc-conjugated oligonucleotides for hepatic and metabolic diseases including:

WVE-007 is a GalNAc-conjugated siRNA targeting inhibin βE (“INHBE”);

• Alpha-1 antitrypsin deficiency ("AATD"):

WVE-006 is a GalNAc-conjugated SERPINA1 AIMer;

• Liver disease:

GalNAc-conjugated AIMer targeting PNPLA3 I148M for correction;

• Heterozygous Familial Hypercholesterolemia (“HeFH”):

GalNAc-conjugated AIMer targeting low-density lipoprotein receptor (“LDLR”) for upregulation and GalNAc-conjugated AIMer targeting apolipoprotein B (“APOB”) for correction.

• Unconjugated oligonucleotides for muscle, CNS and other disease areas including:

• Duchenne muscular dystrophy ("DMD"):

WVE-N531 is an exon 53 splicing oligonucleotide;

• Huntington’s disease ("HD"):

WVE-003 is an allele-selective oligonucleotide designed to lower mutant huntingtin (“mHTT”) protein and preserve healthy, wild-type huntingtin (“wtHTT”) protein.

Our RNA editing capability affords us the dexterity to address both rare and common diseases, as well as those diseases impacting large patient populations.

AIMers are designed to target single bases on an RNA transcript and recruit proteins that exist in the body, called ADAR enzymes, which naturally possess the ability to change an adenine (A) to an inosine (I), which cells read as guanine (G).

AIMers enable simplified delivery and avoid the risk of permanent changes to the genome and irreversible off-target effects with DNA-targeting approaches.

AIMers are short in length, fully chemically modified, and use our novel chemistry, which make them distinct from other ADAR-mediated editing approaches.

GSK Collaboration

In December 2022, we announced a strategic collaboration with GlaxoSmithKline Intellectual Property (No.

3) (“GSK”) to advance transformative oligonucleotide therapeutics, including WVE-006.

The collaboration combines GSK’s novel genetic insights, as well as its global development and commercial capabilities, with our PRISM platform and oligonucleotide expertise.

The collaboration will enable us to continue building a pipeline of first-in-class oligonucleotide-based therapeutics and unlock new areas of disease biology, as well as realize the full value of WVE-006 as a potential best-in-class treatment for AATD that has the potential to simultaneously address both liver and lung manifestations of the disease.

Our GSK collaboration has three components:

(1) a discovery collaboration which enables us to advance up to three programs leveraging targets informed by GSK’s novel genetic insights;

(2) a discovery collaboration which enables GSK to advance up to eight programs leveraging PRISM and our oligonucleotide expertise and discovery capabilities;

(3) an exclusive global license for GSK to WVE-006, our AATD program, that uses our proprietary AIMer technology.

We will maintain development responsibilities for WVE-006 through completion of RestorAATion-2, at which point development and commercial responsibilities will transition to GSK.

Takeda Collaboration (expired in October 2024)

In February 2018, we entered into a global strategic collaboration with Takeda Pharmaceutical Company Limited (“Takeda”), pursuant to which we agreed to collaborate with Takeda on the research, development and commercialization of oligonucleotide therapeutics for disorders of the CNS.

On October 11, 2024, we were notified by Takeda that Takeda did not intend to exercise and therefore elected to terminate its option for the HD target under the collaboration.

As HD was the last active collaboration target under the collaboration, the collaboration expired with immediate effect.

As a result of the option termination, we are now free to advance WVE-003, our clinical-stage Huntington’s disease program, as well as any other programs targeting HTT, independently or with other partners.

We are currently prioritizing lead programs that use GalNAc delivery for hepatic and metabolic diseases, each of which have potential to translate powerful human genetic insights into potentially transformational RNA medicines:

• WVE-007 is a GalNAc-conjugated siRNA (SpiNA design) targeting inhibin βE (“INHBE”) for obesity;

• WVE-006 is a GalNAc-conjugated RNA editing oligonucleotide (AIMer) for AATD;

• WVE-008 is a GalNAc-conjugated RNA editing oligonucleotide (AIMer) for PNPLA3 I148M liver disease.

Our clinical-stage portfolio also includes WVE-N531, an exon 53 splicing oligonucleotide for DMD, and WVE-003, an allele-selective oligonucleotide designed to lower mutant huntingtin (“mHTT”) protein and preserve healthy, wild-type huntingtin (“wtHTT”) protein.

We are also advancing several emerging siRNA and RNA editing programs targeting both hepatic and extra-hepatic tissues.

Financial Operations Overview

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Research and Development Expenses

Research and development expenses consist primarily of costs incurred for our research activities, including our discovery efforts, and the development of our product candidates, which include:

Research and development expenses consist primarily of costs incurred for our research activities, including development of our RNA medicines platform, our discovery efforts, and the development of our product candidates, which include:

• compensation-related expenses, including employee salaries, bonuses, share-based compensation expense and other related benefits expenses for personnel in our research and development organization;

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Our primary research and development focus has been the development of our RNA medicines platform, PRISM.

We are using PRISM, which includes our novel chemistry modifications, to design, develop and commercialize a broad pipeline of first- or best-in class RNA medicines using our editing, splicing, RNAi, and antisense modalities.

We are using PRISM, which combines multiple modalities, chemistry innovation and deep insights in human genetics, to deliver scientific breakthroughs that treat both rare and common disorders, and advance our pipeline of RNA medicines.

Our research and development expenses consist primarily of expenses related to our CROs, CMOs, consultants, other external vendors and fees paid to global regulatory agencies to conduct our clinical trials, in addition to compensation-related expenses, internal manufacturing expenses, facility-related expenses and other general operating expenses.

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However, we do not allocate compensation-related expenses, internal manufacturing expenses, equipment repairs and maintenance expense, facility-related expenses or other operating expenses to specific programs.

These expenses, which are not allocated on a program-by-program basis, are included in the “Other research and development expenses (1) , including INHBE, RNA

editing, PRISM, others” category along with other external expenses related to our discovery and development programs, as well as platform development and identification of potential drug discovery candidates.

These expenses, which are not allocated on a program-by-program basis, are included in the “Other research and development expenses (1) , including PNPLA3, additional preclinical programs, PRISM” category along with other external expenses related to our discovery and development programs, as well as platform development and identification of potential drug discovery candidates.

Product candidates in later stages of clinical development generally have higher development costs than those in earlier stages of clinical development, primarily due to the increased size and duration of later-stage clinical trials.

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Other Income, Net

Other income, net is comprised of refundable tax credits from tax authorities, dividend and interest income earned on cash and cash equivalents balances, gains and losses on foreign currency transactions, and real estate taxes.

Other income, net is comprised primarily of interest income on cash and cash equivalents and refundable tax credits from tax authorities.

We recognize refundable tax credits when there is reasonable assurance that we will comply with the requirements of the refundable tax credit and that the refundable tax credit will be received.

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For the Year Ended December 31,

Increase (Decrease)

(in thousands)

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Loss before income taxes

Income tax benefit (provision)

Revenue for the years ended December 31, 2024 and 2023, was $108.3 million and $113.3 million, respectively, and was earned under the GSK Collaboration Agreement and the Takeda Collaboration Agreement.

The $5.0 million decrease in revenue year over year was driven by the revenue recognized under the GSK Collaboration Agreement, partially offset by the increase in revenue recognized under the Takeda Collaboration Agreement.

The $108.3 million in revenue recognized during the year ended December 31, 2024 was comprised of $37.0 million in revenue recognized under the GSK Collaboration Agreement and $71.3 million of revenue recognized under the Takeda Collaboration Agreement.

The $113.3 million in revenue recognized during the year ended December 31, 2023 was comprised of $66.3 million in revenue recognized under the GSK Collaboration Agreement and $47.0 million of revenue recognized under the Takeda Collaboration Agreement.

The increase in the Takeda Collaboration revenue earned year over year was primarily due to the termination of the collaboration in October 2024, which led to the recognition of the remainder of the deferred revenue related to the research and development services, as well as the license related to the HD program.

This was offset by the decrease in the GSK revenue earned year over year primarily related to the AATD program.

Income tax benefit

Revenue for the year ended December 31, 2025 was $42.7 million, and was earned under the GSK Collaboration Agreement.

Revenue for the year ended December 31, 2024 was $108.3 million, and was earned under the GSK Collaboration Agreement ($37.0 million) and the Takeda Collaboration Agreement ($71.3 million).

The $65.6 million decrease in revenue year over year was driven by the revenue recognized under the Takeda Collaboration Agreement in 2024, partially offset by the increase in revenue recognized under the GSK Collaboration Agreement.

The decrease in the Takeda Collaboration revenue earned year over year was primarily due to the termination of the collaboration agreement in October 2024, which led to the recognition of the remainder of the deferred revenue related to the research and development services, as well as the license related to the HD program.

Research and Development Expenses

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For the Year Ended December 31,

Increase (Decrease)

(in thousands)

Other research and development expenses (1) , including INHBE, RNA editing, PRISM, others

ALS and FTD programs (discontinued)

INHBE program

Other research and development expenses(1), including PNPLA3, additional preclinical programs, PRISM

Total research and development expenses

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The increase of $23.1 million was due to the following:

• an increase of $3.2 million in external expenses related to our AATD program, WVE-006 (RNA editing);

• an increase of $7.7 million in external expenses related to our DMD programs, including WVE-N531 (splicing);

• a decrease of $1.3 million in external expenses related to our HD programs, including WVE-003 (silencing);

• an increase of $28.4 million in other research and development expenses, including INHBE, RNA editing, PRISM, and other internal and external research and development expenses that are not allocated on a program-by-program basis or are related to other discovery and development programs, and the identification of potential drug discovery candidates, mainly due to increases in compensation-related expenses and facilities-related expenses, partially offset by decreases in other external research and development expenses;

• a decrease of $8.3 million in external expenses related to our discontinued ALS and FTD program, WVE-004.

• an increase of $6.4 million in external expenses related to our INHBE program, including WVE-007 (RNAi);

• a decrease of $5.9 million in external expenses related to our AATD program, WVE-006 (RNA editing);

• an increase of $3.9 million in external expenses related to our DMD program, including WVE-N531 (splicing);

• a decrease of $9.1 million in external expenses related to our HD program, including WVE-003 (silencing);

• an increase of $27.8 million in other research and development expenses, including PNPLA3, additional preclinical programs, PRISM, and internal and external research and development expenses that are not allocated on a program-by-program basis or are related to other discovery and development programs, and the identification of potential drug discovery candidates.

This is mainly due to increases in compensation-related expenses and facilities-related expenses, partially offset by decreases in other external research and development expenses.

General and Administrative Expenses

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Other income, net for the years ended December 31, 2025 and 2024 was $11.0 million and $13.4 million, respectively.

The increase of $3.6 million in other income, net was primarily driven by an increase in estimated refundable tax credits as well as an increase in dividend income during the year ended December 31, 2024.

The decrease of $2.4 million in other income, net was primarily driven by a decrease in estimated refundable tax credits during the year ended December 31, 2025.

Income Tax Benefit

During the years ended December 31, 2024 and 2023, we recorded no income tax benefit or provision and an income tax benefit of $0.7 million, respectively.

The income tax benefit for the year ended December 31, 2023 was due to a change in estimate in connection with U.S.

tax guidance relating to the capitalization of research and development expenditures.

During the years ended December 31, 2025 and 2024, we recorded no income tax benefit or provision.

Comparison of the Year Ended December 31, 2024 to the Year Ended December 31, 2023

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For the Year Ended December 31,

Increase (Decrease)

(in thousands)

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Loss before income taxes

Income tax benefit (provision)

Revenue for the year ended December 31, 2023 was $113.3 million and was earned under the GSK Collaboration Agreement and the Takeda Collaboration Agreement.

Revenue for year ended December 31, 2022 was $3.6 million and was earned primarily under the Takeda Collaboration Agreement, as the GSK Collaboration Agreement became effective in January 2023.

The $109.7 million increase in revenue year over year was driven by the revenue recognized under the new GSK Collaboration Agreement, after it became effective in January 2023, as well as the increase in revenue recognized under the Takeda Collaboration Agreement.

Income tax benefit

The $3.6 million in revenue recognized during the year ended December 31, 2022 was primarily related to the research and development services under the Takeda Collaboration Agreement related to the HD, C9, and SCA3 programs.

During the year ended December 31, 2023, the Company recognized revenue of $47.0 million under the Takeda Collaboration.

The increase in revenue earned year over year related to the Takeda Collaboration is primarily due to the termination of the C9 and SCA3 programs in 2023 which led to the recognition of the remainder of the deferred revenue related to the research and development services, as well as the options related to the C9 and SCA3 programs.

This was offset by the decrease in the GSK revenue earned year over year primarily related to the AATD program.

Research and Development Expenses

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For the Year Ended December 31,

Increase (Decrease)

(in thousands)

Other research and development expenses (1) , including INHBE, RNA editing, PRISM, others

ALS and FTD programs (discontinued)

INHBE program

Other research and development expenses(1), including PNPLA3, additional preclinical programs, PRISM

Total research and development expenses

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The increase of $29.7 million was due to the following:

• an increase of $9.1 million in external expenses related to our INHBE program, including WVE-007 (RNAi);

• an increase of $3.2 million in external expenses related to our AATD program, WVE-006 (RNA editing);

• an increase of $5.2 million in external expenses related to our DMD programs, including WVE-N531 (splicing);

• an increase of $5.1 million in external expenses related to our HD programs, including WVE-003 (silencing);

• an increase of $1.6 million in other research and development expenses, including INHBE, RNA editing, PRISM, and other internal and external research and development expenses that are not allocated on a program-by-program basis.

or are related to other discovery and development programs, and the identification of potential drug discovery candidates, mainly due to increases in compensation-related expenses and facilities-related expenses, partially offset by decreases in other external research and development expenses;

• a decrease of $2.5 million in external expenses related to our discontinued ALS and FTD program, WVE-004.

• an increase of $7.7 million in external expenses related to our DMD program, including WVE-N531 (splicing);

• a decrease of $1.3 million in external expenses related to our HD program, including WVE-003 (silencing);

• an increase of $11.0 million in other research and development expenses, including PNPLA3, additional preclinical programs, PRISM, and internal and external research and development expenses that are not allocated on a program-by-program basis or are related to other discovery and development programs, and the identification of potential drug discovery candidates.

This is mainly due to increases in compensation-related expenses and facilities-related expenses, partially offset by decreases in other external research and development expenses.

General and Administrative Expenses

General and administrative expenses were $59.0 million for the year ended December 31, 2024, compared to $51.3 million for the year ended December 31, 2023.

The increase of $0.8 million was primarily driven by increases in other general and administrative operating expenses, partially offset by a decrease in compensation-related expenses.

The increase of $7.7 million was primarily driven by increases in compensation related expenses and administrative expenses.

Other Income, Net

Other income, net for the years ended December 31, 2024 and 2023 was $13.4 million and $9.8 million, respectively.

The increase of $8.2 million in other income, net was primarily driven by an increase in dividend income as well as an increase in estimated refundable tax credits during the year ended December 31, 2023.

Income Tax Benefit (Provision)

During the years ended December 31, 2023 and 2022, we recorded an income tax benefit of $0.7 million and an income tax provision of $0.7 million, respectively.

The income tax benefit for the year ended December 31, 2023 was due to a change in estimate in connection with recent U.S.

The increase of $3.6 million in other income, net was primarily driven by an increase in estimated refundable tax credits as well as an increase in interest income during the year ended December 31, 2024.

Income Tax Benefit

During the years ended December 31, 2024 and 2023, we recorded no income tax benefit or provision and an income tax benefit of $0.7 million, respectively.

The income tax benefit for the year ended December 31, 2023 was due to a change in estimate in connection with U.S.

tax guidance relating to the capitalization of research and development expenditures.

The income tax provision for the year ended December 31, 2022 was primarily due to the requirement under the Tax Cuts and Jobs Act of 2017 for taxpayers to capitalize and amortize research and development expenditures over five or fifteen years pursuant to Section 174 of the Internal Revenue Code of 1986, as amended.

Liquidity and Capital Resources

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Through December 31, 2025, we have received an aggregate of approximately $2,076.7 million in net proceeds from these transactions, consisting of $1,450.5 million in net proceeds from public and other registered offerings of our ordinary shares and other securities, $536.9 million from our collaborations and $89.3 million in net proceeds from private placements of our debt and equity securities.

In January 2024, the representatives of the underwriters in connection with the previously disclosed underwritten public offering (the “December 2023 Offering”) exercised their option to purchase an additional 3,000,000 ordinary shares at a price of $5.00 per ordinary share as a part of the December 2023 Offering.

We received an additional $14.0 million in net proceeds from the December 2023 Offering in January 2024.

On September 27, 2024, we closed the September 2024 Offering in which we issued and sold 23,125,001 of our ordinary shares and the 2024 Pre-Funded Warrants to purchase up to 1,875,023 of our ordinary shares.

The gross proceeds to us from the September 2024 Offering were $200.0 million before deducting underwriting discounts and commissions and other offering expenses.

On October 1, 2024, the representatives of the underwriters exercised their option in full to purchase an additional 3,750,000 ordinary shares, for additional net proceeds to us of approximately $28.2 million.

On December 11, 2025, we closed an underwritten public offering (the "December 2025 Offering") in which we issued and sold 18,552,632 of our ordinary shares, including 2,763,157 ordinary shares issued and sold pursuant to the underwriter's exercise in full of their option to purchase additional shares, and pre-funded warrants to purchase up to 2,631,578 of our ordinary shares (the "2025 Pre-Funded Warrants").

The gross proceeds to us from the December 2025 Offering were approximately $402.5 million before deducting underwriting discounts and commissions and other offering expenses, and including gross proceeds from the exercise of the Underwriters' option to purchase the additional shares in full.

As of December 31, 2025, we had cash and cash equivalents of $602.1 million, restricted cash of $3.8 million and an accumulated deficit of $1,326.2 million.

We expect that our existing cash and cash equivalents will be sufficient to fund our operations for at least the next twelve months.

We expect that our existing cash and cash equivalents will be sufficient to fund our operations for at least the next twelve months from the issuance date of these financial statements.

We have based this expectation on assumptions that may prove to be incorrect, and we may use our available capital resources sooner than we currently expect.

In addition, we may elect to raise additional funds before we need them if the conditions for raising capital are

favorable due to market conditions or strategic considerations, even if we expect we have sufficient funds for our current or future operating plans.

In addition, we may elect to raise additional funds before we need them if the conditions for raising capital are favorable due to market conditions or strategic considerations, even if we expect we have sufficient funds for our current or future operating plans.

Our operating lease commitments as of December 31, 2025 total $19.5 million, of which $9.6 million is related to payments in 2026 and approximately $9.9 million is related to payments beyond 2026.

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2, dated as of March 3, 2022, and Amendment No.

3, dated November 12, 2024, (collectively, the “Sales Agreement”), for our “at-the-market” equity program.

For the three months ended December 31, 2024, we received $5.2 million in net proceeds from sales under our “at-the-market" equity program.

3, dated November

12, 2024, (collectively, the “Sales Agreement”), for our “at-the-market” equity program.

For the twelve months ended December 31, 2025, we received $94.6 million in net proceeds from sales under our “at-the-market" equity program.

Adequate additional financing may not be available to us on acceptable terms, or at all.

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Effect of foreign exchange rates on cash

Net increase (decrease) in cash, cash equivalents and restricted cash

Net increase in cash, cash equivalents and restricted cash

Operating Activities

During 2025, operating activities used $187.5 million of cash, primarily due to our net loss of $204.4 million and changes in our operating assets and liabilities of $16.9 million, partially offset by non-cash charges of $33.8 million The non-cash charges for 2025 related to share-based compensation expense of $25.0 million, amortization of right-of-use assets of $5.4 million, and depreciation expense of $3.4 million.

The largest change in operating assets and liabilities was a $19.8 million decrease in deferred revenue, mainly driven by revenue recognized under the GSK Collaboration Agreement.

During 2024, operating activities used $151.0 million of cash, primarily due to our net loss of $97.0 million, partially offset by non-cash charges of $21.8 million and changes in our operating assets and liabilities of $75.8 million.

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The largest change in operating assets and liabilities was a $54.3 million increase in deferred revenue, mainly driven by our GSK Collaboration Agreement, which became effective in January 2023, which was partially offset by the second largest change in operating assets and liabilities, the $21.1 million increase in accounts receivable primarily related to the achievement of a milestone under the GSK Collaboration Agreement.

During 2022, operating activities used $127.8 million of cash, primarily due to our net loss of $161.8 million, partially offset by non-cash charges of $27.3 million and changes in our operating assets and liabilities of $6.7 million.

The non-cash charges for 2022 related mainly to share-based compensation expense of $17.2 million and depreciation expense of $6.6 million.

The largest change in operating assets and liabilities was a $9.3 million increase in accounts payable.

Investing Activities

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During 2023, investing activities used $1.1 million of cash, primarily consisting of purchases of property and equipment.

Additionally, we purchased $75.0 million of short-term investments during 2022, all of which matured in 2022.

Financing Activities

During 2024, net cash provided by financing activities was $253.9 million, which was primarily due to the $215.8 million in net proceeds from the September 2024 Offering of ordinary shares and the 2024 Pre-Funded Warrants;

as well as the $14.0 million in net proceeds from the January 2024 exercise of the underwriters’ option to purchase an additional 3,000,000 shares under the December 2023 Offering.

During 2025, net cash provided by financing activities was $488.2 million, primarily due to the $377.8 million in net proceeds from the December 2025 Offering of ordinary shares and the 2025 Pre-Funded Warrants, $94.6 million in net proceeds from sales under our “at-the-market” equity program, as well as $14.9 million in proceeds from the exercise of stock options.

During 2024, net cash provided by financing activities was $253.9 million, primarily due to the $215.8 million in net proceeds from the September 2024 underwritten public offering of ordinary shares and the 2024 Pre-Funded Warrants (as defined below) (the "September 2024 Offering");

as well as the $14.0 million in net proceeds from the January 2024 exercise of the underwriters’ option

to purchase an additional 3,000,000 shares under the December 2023 Offering.

Additionally, we received $20.4 million in net proceeds from sales under our “at-the-market” equity program.

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Additionally, there were $3.1 million in net proceeds from our "at-the-market" equity program.

During 2022, net cash provided by financing activities was $67.2 million, primarily due to the $65.5 million in net proceeds from the underwritten offering we completed in June 2022, which was comprised of sales of ordinary shares and the 2022 Pre-Funded Warrants.

Additionally, there were $1.1 million in net proceeds from our "at-the-market" equity program.

Funding Requirements

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• seek and obtain regulatory approvals for our product candidates;

• respond to the impacts of local and global health epidemics, the conflict involving Russia and Ukraine, the conflict in the Middle East, global economic uncertainty, volatility in inflation, volatility in interest rates or market disruptions on our business;

• respond to the impacts of local and global health epidemics, geopolitical conflicts, global economic uncertainty, tariffs, rising inflation, rising interest rates or market disruptions on our business;

• establish and build capabilities to market, distribute and sell our product candidates.

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• the cost of manufacturing clinical supplies of our product candidates;

• whether and to what extent milestone events are achieved under our collaborations with Takeda and GSK or any potential future licensee or collaborator;

• whether and to what extent milestone events are achieved under our collaboration with GSK or any potential future licensee or collaborator;

• the costs, timing and outcome of regulatory review of our product candidates;

• our ability to obtain marketing approval for our product candidates;

• the impacts of local and global health epidemics, the conflict involving Russia and Ukraine, the conflict in the Middle East, global economic uncertainty, volatility in inflation, volatility in interest rates or market disruptions on our business;

• the impacts of local and global health epidemics, geopolitical conflicts, global economic uncertainty, tariffs, rising inflation, rising interest rates or market disruptions on our business;

• the costs and timing of future commercialization activities, including manufacturing, marketing, sales and distribution, for any of our product candidates for which we receive marketing approval;

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Adequate additional funds may not be available to us on acceptable terms when we need them, or at all.

We do not currently have any committed external source of funds, except for possible future payments from GSK under our collaborations with them.

We do not currently have any committed external source of funds, except for possible future payments from GSK under our collaboration with them.

To the extent that we raise additional capital through the sale of equity or convertible debt securities, the ownership interest of our existing shareholders will be diluted, and the terms may include liquidation or other preferences that adversely affect the rights of our shareholders.

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For research and development services that are combined with other promises, the Company utilizes judgment to assess the nature of the combined performance obligation to determine whether the combined performance obligation is satisfied over time or at a point in time and, if over time, the appropriate method of measuring progress for purposes of recognizing revenue.

Company evaluates the measure of progress each reporting period and, if necessary, adjusts the measure of performance and related revenue recognition.

The Company evaluates the measure of progress each reporting period and, if necessary, adjusts the measure of performance and related revenue recognition.

Customer options:

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Any such adjustments are recorded on a cumulative catch-up basis, which would affect revenues and earnings in the period of adjustment.

For arrangements that include sales-based royalties, including milestone payments based on a level of sales, and the license is deemed to be the predominant item to which the royalties relate, the Company recognizes revenue at the later of (i) when the related sales occur, or (ii) when the performance obligation to which some or all of the royalty has been allocated has been satisfied (or partially satisfied).

sales occur, or (ii) when the performance obligation to which some or all of the royalty has been allocated has been satisfied (or partially satisfied).

To date, the Company has not recognized any royalty revenue resulting from any of its licensing arrangements.

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Other Information

Singapore Goods and Services Tax (“GST”) Rate

The issue or transfer of ownership of our ordinary shares would be exempt from GST, although the sale of our ordinary shares by a GST-registered investor may be considered to be a taxable supply subject to GST at 0% if certain conditions are met.

Services consisting of arranging, brokering, underwriting or advising on the issue, allotment or transfer of ownership of our ordinary shares rendered by a GST-registered person to an investor belonging in Singapore for GST purposes in connection with the investor’s purchase, sale or holding of our ordinary shares will be subject to GST at the prevailing standard rate of 9%.

Similar services rendered by a GST-registered person contractually to an investor belonging outside Singapore and for the direct benefit of an investor belonging outside Singapore or a GST-registered person in Singapore should generally, subject to the satisfaction of certain conditions, be subject to GST at 0%.

Rule 10b5-1 Trading Plans

During the three months ended December 31, 2024, certain of our officers (as defined in Rule 16a-1(f) of the Exchange Act) entered into contracts, instructions or written plans (each, a “Rule 10b5-1 Trading Plan” and collectively, the “Rule 10b5-1 Trading Plans”) for the purchase or sale of our securities that are intended to satisfy the conditions specified in Rule 10b5-1(c) under the Exchange Act for an affirmative defense against liability for trading in securities on the basis of material nonpublic information.

During the three months ended December 31, 2025, certain of our officers (as defined in Rule 16a-1(f) of the Exchange Act) and directors entered into contracts, instructions or written plans (each, a “Rule 10b5-1 Trading Plan” and collectively, the “Rule 10b5-1 Trading Plans”) for the purchase or sale of our securities that are intended to satisfy the conditions specified in Rule 10b5-1(c) under the Exchange Act for an affirmative defense against liability for trading in securities on the basis of material nonpublic information.

We describe the material terms of these Rule 10b5-1 Trading Plans below.

On November 24, 2024 , Christian Henry , MBA, Chairman of our Board , adopted a Rule 10b5-1 Trading Plan providing for the sale of up to an aggregate of 105,670 of our ordinary shares pursuant to the terms of such Rule 10b5-1 Trading Plan.

10b5-1 Trading Plan is active until August 29, 2025 , or earlier, if and when all transactions under the Rule 10b5-1 Trading Plan are completed.

On November 25, 2024 , Kyle Moran , CFA, our Chief Financial Officer , adopted a Rule 10b5-1 Trading Plan providing for the sale of up to an aggregate of 196,000 of our ordinary shares pursuant to the terms of such Rule 10b5-1 Trading Plan.

Moran’s Rule 10b5-1 Trading Plan is active until June 30, 2025 , or earlier, if and when all transactions under the Rule 10b5-1 Trading Plan are completed.

On November 21, 2025 , Paul B.

Bolno , M.D., MBA, our President and Chief Executive Officer , adopted a Rule 10b5-1 Trading Plan providing for the sale of up to an aggregate of 1,480,900 of our ordinary shares pursuant to the terms of such Rule 10b5-1 Trading Plan.

Bolno's Rule 10b5-1 Trading Plan is active until June 16, 2026 , or earlier, if and when all transactions under the Rule 10b5-1 Trading Plan are completed.

On November 20, 2025 , Christopher Francis , Ph.D., our Senior Vice President , Corporate Development, Head of Emerging Areas, adopted a Rule 10b5-1 Trading Plan providing for the sale of up to an aggregate of 520,702 of our ordinary shares pursuant to the terms of such Rule 10b5-1 Trading Plan.

Francis' Rule 10b5-1 Trading Plan is active until November 3, 2025 , or earlier, if and when all transactions under the Rule 10b5-1 Trading Plan are completed.

On November 25, 2024 , Chandra Vargeese , Ph.D., our Chief Technology Officer , Head of Platform Discovery Sciences, adopted a Rule 10b5-1 Trading Plan providing for the sale of up to an aggregate of 86,972 of our ordinary shares pursuant to the terms of such Rule 10b5-1 Trading Plan.

Francis’ Rule 10b5-1 Trading Plan is active until August 21, 2026 , or earlier, if and when all transactions under the Rule 10b5-1 Trading Plan are completed.

On December 17, 2025 , Kyle Moran , CFA, our Chief Financial Officer , terminated a Rule 10b5-1 Trading Plan that was originally adopted on November 20, 2025 .

Moran’s former Rule 10b5-1 Trading Plan provided for the sale of up to an aggregate of 392,647 of our ordinary shares.

No ordinary shares were sold under Dr.

Moran’s Rule 10b5-1 Trading Plan prior to its termination.

On December 18, 2025 , Mr.

Moran adopted a Rule 10b5-1 Trading Plan providing for the sale of up to an aggregate of 196,647 of our ordinary shares pursuant to the terms of such Rule 10b5-1 Trading Plan.

Moran's Rule 10b5-1 Trading Plan is active until August 21, 2026 , or earlier, if and when all transactions under the Rule 10b5-1 Trading Plan are completed.

On December 16, 2025 , Chandra Vargeese , Ph.D., our Chief Technology Officer , Head of Platform Discovery Sciences, terminated a Rule 10b5-1 Trading Plan that was originally adopted on November 19, 2025 .

Vargeese’s former Rule 10b5-1 Trading Plan provided for the sale of up to an aggregate of 132,886 of our ordinary shares.

No ordinary shares were sold under Dr.

Vargeese’s Rule 10b5-1 Trading Plan prior to its termination.

On December 17, 2025 , Dr.

Vargeese adopted a Rule 10b5-1 Trading Plan providing for the sale of up to an aggregate of 82,886 of our ordinary shares pursuant to the terms of such Rule 10b5-1 Trading Plan.

Vargeese’s Rule 10b5-1 Trading Plan is active until August 20, 2026 , or earlier, if and when all transactions under the Rule 10b5-1 Trading Plan are completed.

Except as disclosed above, none of our directors or executive officers adopted , modified or terminated any contract, instruction or written plan for the purchase or sale of our securities that was intended to satisfy the affirmative defense conditions of Rule 10b5-1(c) or any “non-Rule 10b5-1 trading arrangement” as such term is defined in Item 408(a) of Regulation S-K, during the fiscal quarter ended December 31, 2025.

Disclosure Regarding Foreign Jurisdictions that Prevent Inspections

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Executi ve Compensation

The information required by this item is incorporated by reference to the information set forth in the section titled “Executive Officer and Director Compensation” in our Proxy Statement.

The information required by this item is incorporated by reference to the information set forth in the section titled “Executive Officer and Director Compensation,” Compensation Discussion and Analysis,” “Management and Corporate Governance—Compensation Committee Interlocks and Insider Participation,” and “Compensation Committee Report” in our Proxy Statement.

The section entitled “Pay Versus Performance” in our Proxy Statement is not incorporated by reference herein.

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(Exhibit 4.1)

Form of Pre-Funded Warrant (2025)

(Exhibit 4.1)

Share Purchase Agreement by and between the Registrant and C.P.

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(Exhibit 10.1)

Consulting Agreement by and between Ontorii, Inc.

(now Wave Life Sciences USA, Inc.) and Gregory Verdine, dated as of April 1, 2012

(Exhibit 10.16)

Equity and Other Compensation Plans

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Insider Trading Policy

(Exhibit 19.1)

List of Subsidiaries of the Registrant

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(*) The certification attached as Exhibit 32 that accompanies this Annual Report on Form 10-K is not deemed filed with the SEC and is not to be incorporated by reference into any filing of Wave Life Sciences Ltd.

under the Securities Act of 1933, as amended, or the

Securities Exchange Act of 1934, as amended, whether made before or after the date of this Form 10-K, irrespective of any general incorporation language contained in such filing.

under the Securities Act of 1933, as amended, or the Securities Exchange Act of 1934, as amended, whether made before or after the date of this Form 10-K, irrespective of any general incorporation language contained in such filing.

(+) Indicates management contract or compensatory plan or arrangement.

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Wave Life Sciences Ltd.

March 4, 2025

February 26, 2026

President and Chief Executive Officer

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( principal executive officer )

March 4, 2025

February 26, 2026

/s/ Kyle Moran

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principal accounting officer )

March 4, 2025

February 26, 2026

/s/ Christian Henry

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Chairman of the Board of Directors

March 4, 2025

February 26, 2026

/s/ Gregory L.

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Verdine, Ph.D.

March 4, 2025

February 26, 2026

/s/ Peter Kolchinsky, Ph.D.

Peter Kolchinsky, Ph.D.

March 4, 2025

February 26, 2026

/s/ Aik-Na Tan

March 4, 2025

February 26, 2026

/s/ Adrian Rawcliffe

Adrian Rawcliffe

March 4, 2025

February 26, 2026

/s/ Ken Takanashi

Ken Takanashi

March 4, 2025

February 26, 2026

Corrigan, M.D.

March 4, 2025

February 26, 2026

INDEX TO CONSOLIDATED FINANCIAL STATEMENTS

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We have audited the accompanying consolidated balance sheets of Wave Life Sciences Ltd.

and subsidiaries (the Company) as of December 31, 2024 and 2023, the related consolidated statements of operations and comprehensive loss, Series A preferred shares and shareholders’ equity (deficit), and cash flows for the each of the years in the three-year period ended December 31, 2024, and the related notes (collectively, the consolidated financial statements).

and subsidiaries (the Company) as of December 31, 2025 and 2024, the related consolidated statements of operations and comprehensive loss, Series A preferred shares and shareholders’ equity (deficit), and cash flows for each of the years in the three-year period ended December 31, 2025, and the related notes (collectively, the consolidated financial statements).

We also have audited the Company’s internal control over financial reporting as of December 31, 2025, based on criteria established in Internal Control – Integrated Framework (2013) issued by the Committee of Sponsoring Organizations of the Treadway Commission.

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Evaluation of revenue recognition for certain research and development services

As discussed in Note 5 to the consolidated financial statements, the Company is party to a collaboration agreement with GlaxoSmithKline (GSK) which has two performance obligations, including the promise to provide research and development (R&D) services.

As discussed in Note 5 to the consolidated financial statements, the Company is party to a collaboration agreement with GlaxoSmithKline (GSK) which has several performance obligations, including the promise to provide research and development (R&D) services.

The Company recognizes R&D services revenue over time using an input method.

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The following are the primary procedures we performed to address this critical audit matter.

For a selection of R&D performance obligations, we read the underlying contract with the customer, evaluated the determination of the method for measuring progress, and tested the Company’s estimate of total contract costs to be incurred by (1) comparing the Company’s initial estimates to actual costs incurred to assess the Company’s ability to estimate accurately, (2) inspecting underlying documentation and third-party evidence and comparing them to management’s assumptions and inputs, (3) inquiring of R&D personnel of the Company to evaluate factors related to the nature of the work to be performed and their impact on the total contract costs to be incurred, including progress to date and the estimate of remaining contract costs, and (4) assessing the Company’s history of estimating costs to be incurred in satisfying R&D performance obligations under similar contracts.

We evaluated the design and tested the operating effectiveness of certain internal controls related to revenue recognition for certain R&D services.

This included controls related to the Company’s process to develop and review the estimate of total costs expected to be incurred in satisfying certain R&D performance obligations, including the assessment of the nature of the work to be performed.

For a selection of R&D performance obligations, we read the underlying contract with the customer, evaluated the determination of the method for measuring progress, and tested the Company’s estimate of total contract costs to be incurred by (1) comparing the Company’s initial estimates to actual costs incurred to assess the Company’s ability to estimate accurately, (2) inspecting underlying documentation and third-party evidence and comparing them to management’s assumptions and inputs, and (3) inquiring of R&D personnel of the Company to evaluate factors related to the nature of the work to be performed and their impact on the total contract costs to be incurred, including progress to date and the estimate of remaining contract costs.

We have served as the Company’s auditor since 2015.

Boston, Massachusetts

March 4, 2025

February 26, 2026

WAVE LIFE SCIENCES LTD.

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Loss before income taxes

Income tax benefit (provision)

Income tax benefit

Net loss per share attributable to ordinary

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Other comprehensive loss:

Foreign currency translation

Foreign currency translation gain (loss)

Comprehensive loss

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Comprehensive

Income (Loss)

Shareholders’

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net of offering costs

Issuance of ordinary shares, pursuant

to the GSK Collaboration Agreement

Issuance of ordinary shares pursuant to

the at-the-market equity program, net

Issuance of pre-funded warrants,

net of offering costs

Share-based compensation

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net of offering costs

Issuance of ordinary shares, pursuant

to the GSK Collaboration Agreement

Issuance of ordinary shares pursuant to

the at-the-market equity program, net

Issuance of pre-funded warrants,

net of offering costs

Share-based compensation

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Share-based compensation expense

Loss on disposal of property and equipment

Changes in operating assets and liabilities:

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Purchases of property and equipment

Proceeds from the sale of property and equipment

Purchase of short-term investments

Proceeds from the maturity of short-term investments

Net cash used in investing activities

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Proceeds from the issuance of ordinary shares as a part of the

June 2022 Offering, net of offering costs

Proceeds from the issuance of ordinary shares as a part of the

December 2023 Offering, net of offering costs

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Proceeds from issuance pre-funded warrants as a part of the

June 2022 Offering, net of offering costs

Proceeds from issuance pre-funded warrants as a part of the,

September 2024 Offering, net of offering costs

Proceeds from the issuance of ordinary shares as a part of the

December 2025 Offering, net of offering costs

Proceeds from issuance pre-funded warrants as a part of the

December 2025 Offering, net of offering costs

Proceeds from issuance of ordinary shares pursuant to the

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Effect of foreign exchange rates on cash

Net increase (decrease) in cash, cash equivalents and restricted cash

Net increase in cash, cash equivalents and restricted cash

Cash, cash equivalents and restricted cash, beginning of period

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Offering costs in accounts payable at period end

Increase in operating lease right-of-use assets and

lease liabilities related to new lease

The accompanying notes are an integral part of the consolidated financial statements.

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Wave Life Sciences Ltd.

(together with its subsidiaries, “Wave” or the “Company”) is a clinical-stage biotechnology company focused on unlocking the broad potential of RNA medicines (also known as oligonucleotides), or those targeting RNA, to transform human health.

Wave’s RNA medicines platform, PRISM, combines multiple modalities, chemistry innovation and deep insights into human genetics to deliver scientific breakthroughs that treat both rare and common disorders.

The Company’s toolkit of RNA-targeting modalities includes RNA editing, splicing, silencing using siRNA and antisense silencing, providing us with unique capabilities for designing and sustainably delivering candidates that optimally address disease biology.

The Company’s diversified pipeline includes clinical programs in obesity, AATD, DMD, and HD, as well as several preclinical programs utilizing our versatile RNA medicines platform.

(together with its subsidiaries, “Wave” or the “Company”) is a clinical-stage biotechnology company focused on unlocking the broad potential of ribonucleic acid (“RNA”) medicines (also known as oligonucleotides), or those targeting RNA, to transform human health.

Our toolkit of RNA-targeting modalities, including RNAi silencing (SpiNA) and RNA editing (AIMers), provides us with unmatched capabilities for designing and sustainably delivering candidates that optimally address disease biology.

The Company was incorporated in Singapore on July 23, 2012 and has its principal U.S.

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As of December 31, 2025, the Company had cash and cash equivalents of $ 602.1 million.

The Company expects that its existing cash and cash equivalents will be sufficient to fund its operations for at least the next twelve months.

The Company expects that its existing cash and cash equivalents will be sufficient to fund its operations for at least the next twelve months from the issuance date of these financial statements.

The Company has based this expectation on the best information available, however the Company may use its available capital resources sooner than it currently expects.

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Restricted cash consists primarily of cash placed in separate restricted bank accounts as required under the terms of the Company’s lease agreements for its Cambridge, Massachusetts and Lexington, Massachusetts facilities (refer to Note 8).

As of December 31, 2024 and 2023, the Company had $ 3.8 million and $ 3.7 million of restricted cash, respectively, of which $ 2.8 million and $ 2.7 million related to the Lexington facility, respectively, and $ 1.0 million related to the Cambridge facility.

As of December 31, 2025 and 2024, the Company had $ 3.8 million of restricted cash, of which $ 2.8 million related to the Lexington facility, and $ 1.0 million related to the Cambridge facility.

Property and Equipment

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Share-Based Compensation

The Company measures and recognizes share-based compensation expense, for both employee and director option awards, based on the grant date fair value of the awards.

The Company measures and recognizes share-based compensation expense, for employees, consultants, and non-employee directors awards, based on the grant date fair value of the awards.

The Company calculates the fair value of awards based on the grant date fair value of the underlying ordinary shares.

The Company determines the fair value of share-based awards granted to non-employees as either the fair value of the consideration received or the fair value of the equity instruments issued, whichever is more reliably measurable.

Equity instruments issued to non-employees as consideration for goods or services received by the Company have been accounted for based on the fair value of the equity instruments issued.

Equity instruments issued to consultants and non-employee directors as consideration for goods or services received by the Company have been accounted for based on the fair value of the equity instruments issued.

The Company recognizes share-based compensation expense on a straight-line basis over the requisite service period of the awards, which is generally the vesting period.

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Recently Issued Accounting Pronouncements

In November 2023, the Financial Accounting Standards Board (the “FASB”) finalized Accounting Standards Update No.

2023-07, Segment Reporting (Topic 280):

Improvements to Reportable Segment Disclosures (“ASU 2023-07”).

ASU 2023-07 requires enhanced disclosures about reportable segments and the chief operating decision maker.

The Company adopted ASU 2023-07 for the Company's fiscal year 2024 annual reporting period and applied it retrospectively.

The adoption did not have a material impact on the Company's consolidated financial statements.

In December 2023, the FASB finalized Accounting Standards Update No.

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ASU 2023-09 requires a company's annual financial statements to include consistent categories and greater disaggregation of information in the rate reconciliation, and income taxes paid disaggregated by jurisdiction.

ASU 2023-09 is effective for the Company’s annual reporting periods beginning after December 15, 2025.

Adoption is either with a

prospective method or a fully retrospective method of transition.

Early adoption is permitted.

The Company is currently evaluating the effect that adoption of ASU 2023-09 will have on its consolidated financial statements.

The Company adopted ASU 2023-09 for the Company's fiscal year 2025 annual reporting period and applied it prospectively.

The adoption did not have a material impact on the Company's consolidated financial statements.

In November 2024, the FASB issued ASU 2024-03, Income Statement - Reporting Comprehensive Income - Expense Disaggregation Disclosures (Subtopic 220-40) (“ASU 2024-03”).

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Pursuant to the GSK Collaboration Agreement, Wave and GSK have agreed to collaborate on the research, development, and commercialization of oligonucleotide therapeutics, including an exclusive global license to WVE-006.

The discovery collaboration component has an initial four-year research term and combines Wave’s proprietary discovery and drug development platform, PRISM, with GSK’s unique genetic insights and its global

development and commercial capabilities.

On January 27, 2023, the GSK Collaboration Agreement became effective, and GSK paid Wave an upfront payment of $ 120.0 million.

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The GSK Equity Investment closed on January 26, 2023, following the completion of customary closing conditions.

The ordinary shares purchased by GGL in the GSK Equity Investment are subject to lock-up and standstill restrictions and carry certain registration rights, customary for transactions of this kind.

The ordinary shares purchased by GGL in the GSK Equity Investment carry certain registration rights, customary for transactions of this kind.

The Company did not incur any material costs in connection with the issuance of the ordinary shares under the SPA.

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Thereafter, GSK will be responsible for advancing WVE-006 through pivotal studies, registration, and global commercialization at GSK’s sole cost.

On February 2, 2026, the Company announced that it regained full rights to WVE-006, an investigational GalNAc-conjugated RNA editing oligonucleotide for alpha-1 antitrypsin deficiency, which was originally the subject of an exclusive global license to GSK under the GSK Collaboration Agreement.

Under the GSK Collaboration Agreement, each party grants to the other party certain licenses to the collaboration products to enable the other party to perform its obligations and exercise its rights under the GSK Collaboration Agreement, including license grants to enable each party to conduct research, development and commercialization activities pursuant to the terms of the GSK Collaboration Agreement.

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The Company will reevaluate the transaction price at the end of each reporting period, and as uncertain events are resolved or other changes in circumstances occur, the Company will adjust its estimate of the transaction price.

Under the GSK Collaboration Agreement, GSK can advance up to eight programs leveraging the Company's PRISM platform and multiple RNA-targeting modalities (RNA editing, splicing, siRNA, and antisense) with target validation work ongoing across multiple therapy areas.

GSK selected its first two programs to advance to development candidates following achievement of target validation in the three months ended June 30, 2024.

These programs utilize the Company's next generation GalNAc-siRNA format and are in hepatology.

Under the GSK Collaboration Agreement, GSK was required to provide an aggregate initiation payment of $ 12.0 million to the Company for these two oligonucleotide programs, which was received during the three months ended June 30, 2024.

The following table summarizes the allocation of the total transaction price to the identified performance obligation under the GSK Collaboration Agreement, and the amount of the transaction price unsatisfied as of December 31, 2024 (in thousands):

Transaction Price Allocated

Under the GSK Collaboration Agreement, GSK can advance up to eight programs (GSK Collaboration Programs) leveraging the Company's PRISM platform and multiple RNA-targeting modalities (RNAi, RNA editing, splicing, and antisense) with target validation work ongoing across multiple therapy areas.

The advancement to a development candidate following the achievement of target validation results in a GSK Collaboration Program performance obligation.

In September 2025, GSK selected a third program to advance to a development candidate following the achievement of target validation, resulting in three programs designated as GSK Collaboration Programs as of December 31, 2025.

Subsequent to December 31, 2025, GSK selected a fourth program to advance to development candidate following achievement of target validation.

Under the GSK Collaboration Agreement, GSK has paid an aggregate of $ 32.0 million in program initiation payments related to these four programs, of which $ 10.0 million was received after December 31, 2025.

The following table summarizes the allocation of the total transaction price to the identified performance obligations under the GSK Collaboration Agreement, and the amount of the transaction price unsatisfied as of December 31, 2025 (in thousands):

Transaction Price

Transaction Price Unsatisfied (1)

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Discovery Research Collaboration

GSK Collaboration Program

GSK Collaboration Programs

(1) The Unsatisfied transaction price will be recognized over the remaining applicable research or program term.

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Additional funding related to the Company’s research activities related to Discovery Research Collaboration will be recorded as accounts receivable when contractually enforceable and recorded as deferred revenue, or as revenue as the services are provided.

During the year ended December 31, 2023 , the Company achieved a developmental milestone which pertained to the initiation of dosing in healthy volunteers in the RestorAATion clinical trial program, triggering a $ 20.0 million milestone payment to the Company from GSK.

As of December 31, 2023 , the $ 20.0 million related to the achievement of the milestone was included in the current portion of accounts receivable and payment was received from GSK in the first quarter of 2024.

Under the GSK Collaboration Agreement, during the years ended December 31, 2024 and December 31, 2023, the Company recognized revenue of $ 37.0 million and $ 66.3 million, respectively, using the input method described above.

During the year ended December 31, 2023, the Company achieved a developmental milestone which pertained to the initiation of dosing in healthy volunteers in the RestorAATion clinical trial program, triggering a $ 20.0 million milestone payment to the Company from GSK, which was collected in the first quarter of 2024.

During the year ended December 31, 2025 , GSK acknowledged the Company’s achievement of another development milestone for the AATD program, triggering a $ 10.0 million milestone payment to the Company from GSK, which was collected during the fourth quarter of 2025.

Under the GSK Collaboration Agreement, during the years ended December 31, 2025, 2024, and 2023 , the Company recognized revenue of $ 42.7 million, $ 37.0 million, and $ 66.3 million, respectively, using the input method described above.

The aggregate amount of the transaction price allocated to the Company’s unsatisfied and partially unsatisfied performance obligations which are recorded in deferred revenue as of December 31, 2025 is approximately $ 52.2 million, of which approximately $ 44.4 million is included in current liabilities and $ 7.8 million is included in long-term liabilities.

The aggregate amount of the transaction price allocated to the Company’s unsatisfied and partially unsatisfied performance obligations which were recorded in deferred revenue as of December 31, 2024 was approximately $ 72.1 million, of which approximately $ 66.0 million was included in current liabilities and $ 6.1 million was included in long-term liabilities.

The Company expects to recognize the remaining deferred revenue related to the AATD performance obligation, which was fully satisfied upon the termination of the AATD license, in the first quarter of 2026.

Takeda Collaboration and Equity Agreements

In February 2018 , Wave USA and Wave UK entered into a global strategic collaboration (the “Takeda Collaboration”) with Takeda Pharmaceutical Company Limited (“Takeda”), pursuant to which Wave USA, Wave UK and Takeda agreed to collaborate on the research, development and commercialization of oligonucleotide therapeutics for disorders of the Central Nervous System (“CNS”).

The Takeda Collaboration provided the Company with at least $ 230.0 million in committed cash and Takeda with the option to co-develop and co-commercialize the Company’s CNS development programs in (1) Huntington’s disease (“HD”);

(2) amyotrophic lateral sclerosis (“ALS”) and frontotemporal dementia (“FTD”);

and (3) the Company’s discovery-stage program targeting ATXN3 for the treatment of spinocerebellar ataxia 3 (“SCA3”) (collectively, “Category 1 Programs”).

In addition, the Takeda Collaboration provided Takeda the right to exclusively license multiple preclinical programs for CNS disorders, including Alzheimer’s disease and Parkinson’s disease (collectively, “Category 2 Programs”).

In April 2018, the Takeda Collaboration became effective and Takeda paid the Company $ 110.0 million as an upfront payment.

Takeda also agreed to fund the Company’s research and preclinical activities in the amount of $ 60.0 million during the four-year research term and to reimburse the Company for any collaboration-budgeted research and preclinical expenses incurred by Wave that exceed that amount.

Simultaneously with Wave USA and Wave UK’s entry into the collaboration and license agreement with Takeda dated February 19, 2018, as amended (the “Takeda Collaboration Agreement”), the Company entered into a share purchase agreement with Takeda (the “Takeda Equity Agreement,” and together with the Takeda Collaboration Agreement, the “Takeda Agreements”) pursuant to which it agreed to sell to Takeda 1,096,892 of its ordinary shares at a purchase price of $ 54.70 per share.

In April 2018 , the Company closed the Takeda Equity Agreement and received aggregate cash proceeds of $ 60.0 million.

The Company did not incur any material costs in connection with the issuance of the shares.

With respect to Category 1 Programs, the Company was responsible for researching and developing products and companion diagnostics for Category 1 Programs through completion of the first proof of mechanism study for such products.

Takeda had an exclusive option for each target and all associated products and companion diagnostics for such target, which it could exercise at any time through completion of the proof of mechanism study.

If Takeda had exercised this option, the Company would have received an opt-in payment and would have led manufacturing and joint clinical co-development activities and Takeda would have led joint co-commercial activities in the United States and all commercial activities outside of the United States.

Global costs and potential profits would have been shared 50 :

50 and the Company would have been eligible to receive development and commercial milestone payments.

In addition to its 50% profit share, the Company was eligible to receive option exercise fees and development and commercial milestone payments for each of the Category 1 Programs.

With respect to Category 2 Programs, the Company granted Takeda the right to exclusively license multiple preclinical programs during a four-year research term (subject to limited extension for programs that were initiated prior to the expiration of the research term, in accordance with the Takeda Collaboration Agreement) (“Category 2 Research Term”).

During that term, the Takeda Collaboration provided that the parties may collaborate on preclinical programs for up to six targets at any one time .

The Company was responsible for researching and preclinically developing products and companion diagnostics directed to the agreed upon targets through completion of Investigational IND enabling studies in the first major market country.

Thereafter, Takeda would have an exclusive worldwide license to develop and commercialize products and companion diagnostics directed to such targets, subject to the Company’s retained rights to lead manufacturing activities for products directed to such targets.

Takeda agreed to fund the Company’s research and preclinical activities in the amount of $ 60.0 million during the research term and reimburse the Company for any collaboration-budgeted research and preclinical expenses incurred by the Company that exceeded that amount.

The Company was also eligible to receive tiered high single-digit to mid-teen royalties on Takeda’s global commercial sales of products from each Category 2 Program.

Under the Takeda Collaboration Agreement, each party granted to the other party specific intellectual property licenses to enable the other party to perform its obligations and exercise its rights under the Takeda Collaboration Agreement, including license grants to enable each party to conduct research, development and commercialization activities pursuant to the terms of the Takeda Collaboration Agreement.

The term of the Takeda Collaboration Agreement commenced on April 2, 2018 and, unless terminated earlier, would have continued until the date on which:

(i) with respect to each Category 1 Program target for which Takeda does not exercise its option, the expiration or termination of the development program with respect to such target;

(ii) with respect to each Category 1 Program target for which Takeda exercises its option, the date on which neither party is researching, developing or manufacturing any products or companion diagnostics directed to such target;

or (iii) with respect to each Category 2 Program target, the date on which royalties are no longer payable with respect to products directed to such target.

Takeda had the right to terminate the Takeda Collaboration Agreement for convenience on 180 days ’ notice, in its entirety or on a target-by-target basis.

Subject to certain exceptions, each party had the right to terminate the Takeda Collaboration Agreement on a target-by-target basis if the other party, or a third party related to such party, challenges the patentability, enforceability or validity of any patents within the licensed technology that cover any product or companion diagnostic that was subject to the Takeda Collaboration Agreement.

In the event of any material breach of the Takeda Collaboration Agreement by a party, subject to cure rights, the other party had the right to terminate the Takeda Collaboration Agreement in its entirety if the breach related to all targets or on a target-by-target basis if the breach related to a specific target.

In the event that Takeda and its affiliates ceased development, manufacturing and commercialization activities with respect to compounds or products subject to the Takeda Collaboration Agreement and directed to a particular target, the Company had the right to terminate the Takeda Collaboration Agreement with respect to such target.

Either party had the right to terminate the Takeda Collaboration Agreement for the other party’s insolvency.

In certain termination circumstances, the Company would have received a license from Takeda to continue researching, developing and manufacturing certain products, and companion diagnostics.

The Takeda Collaboration was managed by a joint steering committee in which both parties were represented equally.

The joint steering committee was tasked with overseeing the scientific progression of each Category 1 Program and, prior to the Amendment (discussed below), the Category 2 Programs.

The Company assessed this arrangement in accordance with ASC 606 and concluded that the contract counterparty, Takeda, is a customer for Category 1 Programs prior to Takeda exercising its option, and for Category 2 Programs during the Category 2 Research Term.

The Company identified the following material promises under the arrangement:

(1) the non-exclusive, royalty-free research and development license for each Category 1 Program;

(2) the research and development services for each Category 1 Program through completion of the first proof of mechanism study;

(3) the exclusive option to license, co-develop and co-commercialize each Category 1 Program;

(4) the right to exclusively license the Category 2 Programs;

and (5) the research and preclinical development services of the Category 2 Programs through completion of IND-enabling studies.

The research and development services for each Category 1 Program were determined to not be distinct from the research and development license and should therefore be combined into a single performance obligation for each Category 1 Program.

The research and preclinical development services for the Category 2 Programs were determined to not be distinct from the exclusive licenses for the Category 2 Programs and therefore were combined into a single performance obligation.

Additionally, the Company determined that the exclusive option for each Category 1 Program was priced at a discount and, as such, provide material rights to Takeda, representing three separate performance obligations.

Based on these assessments, the Company identified seven performance obligations in the Takeda Collaboration Agreement:

(1) research and development services through completion of the first proof of mechanism and non-exclusive research and development license for HD;

(2) research and development services through completion of the first proof of mechanism and non-exclusive research and development license for ALS and FTD;

(3) research and development services through completion of the first proof of mechanism and non-exclusive research and development license for SCA3;

(4) the material right provided for the exclusive option to license, co-develop and co-commercialize HD;

(5) the material right provided for the exclusive option to license, co-develop and co-commercialize ALS and FTD;

(6) the material right provided for the exclusive option to license, co-develop and co-commercialize SCA3;

and (7) the research and preclinical development services and right to exclusively license the Category 2 Programs.

At the outset of the arrangement, the transaction price included the $ 110.0 million upfront consideration received and the $ 60.0 million of committed research and preclinical funding for the Category 2 Programs.

The Company determined that the Takeda Collaboration Agreement did not contain a significant financing component.

The option exercise fees to license, co-develop and co-commercialize each Category 1 Program that could have been received were excluded from the transaction price until each customer option was exercised.

The potential milestone payments were excluded from the transaction price, as all milestone amounts were fully constrained at the inception of the Takeda Collaboration Agreement.

The Company would have reevaluated the transaction price at the end of each reporting period and, as uncertain events were resolved or other changes in circumstances occurred, if necessary, would have adjusted its estimate of the transaction price.

The Company allocated the transaction price to the performance obligations on a relative standalone selling price basis.

For the performance obligations associated with the research and development services through completion of the first proof of mechanism and non-exclusive research and development license for HD;

the research and development services through completion of the first proof of mechanism and non-exclusive research and development license for ALS and FTD;

the research and development services through completion of the first proof of mechanism and non-exclusive research and development license for SCA3;

and the research and preclinical development services and right to exclusively license the Category 2 Programs, the Company determined the

standalone selling price using estimates of the costs to perform the research and development services, including expected internal and external costs for services and supplies, adjusted to reflect a profit margin.

The total estimated cost of the research and development services reflected the nature of the services to be performed and the Company’s best estimate of the length of time required to perform the services.

For the performance obligations associated with the material right provided for the exclusive option to license, co-develop and co-commercialize HD;

the material right provided for the exclusive option to license, co-develop and co-commercialize ALS and FTD;

and the material right provided for the exclusive option to license, co-develop and co-commercialize SCA3, the Company estimated the standalone fair value of the option to license each Category 1 Program utilizing an adjusted market assessment approach, and determined that any standalone fair value in excess of the amounts to be paid by Takeda associated with each option represented a material right.

Revenue associated with the research and development services for each Category 1 Program performance obligation were recognized as the research and development services were provided using an input method, according to the costs incurred on each Category 1 Program and the total costs expected to be incurred to satisfy each Category 1 Program performance obligation.

Prior to the Amendment described below, revenue associated with the research and preclinical development services for the Category 2 Programs performance obligation was recognized as the research and preclinical development services that were provided using an input method, according to the costs incurred on Category 2 Programs and the total costs expected to be incurred to satisfy the performance obligation.

The amount allocated to the material right for each Category 1 Program option would have been recognized on the date that Takeda exercised each respective option, or immediately as each option expired unexercised.

The amounts received that were not yet recognized as revenue were recorded in deferred revenue on the Company’s consolidated balance sheet.

On October 15, 2021 , Wave USA, Wave UK and Takeda entered into the Second Amendment to the Takeda Collaboration Agreement (the “Amendment”), which discontinued the Category 2 component of the Takeda Collaboration.

The Category 1 Programs under the Collaboration Agreement remain in effect and are unchanged by the Amendment.

Pursuant to the Amendment, Takeda agreed to pay the Company an additional $ 22.5 million as full payment for reimbursable Category 2 Programs collaboration-budgeted research and preclinical expenses.

The Company received this payment from Takeda related to the Category 2 component and recognized the full amount as collaboration revenue in the year ended December 31, 2021.

During the year ended December 31, 2021, in addition to the revenue recognized related to the Amendment, the Company recognized another $ 18.5 million of collaboration revenue related to services pertaining to the Category 1 Programs and Category 2 Programs.

In May 2023, the Company announced its decision to discontinue clinical development of WVE-004 for C9orf72-associated ALS and FTD (“C9 for ALS/FTD”), one of the Category 1 Programs.

In July 2023, the joint steering committee that manages the Takeda Collaboration terminated C9 for ALS/FTD as a target under the collaboration (the “C9 Target”) and consequently Takeda and the Company’s rights and obligations under the Takeda Collaboration were terminated with respect to the C9 Target.

As a result of the termination of the C9 for ALS/FTD Category 1 Program, the Company recognized $ 28.0 million in revenue during the three months ended September 30, 2023, which represented the remainder of the deferred revenue for the C9 for ALS/FTD Category 1 Program as of June 30, 2023.

In the third quarter of 2023, the Company achieved a developmental milestone related to the HD Category 1 Program, which pertained to the positive results from a non-clinical study of WVE-003 in non-human primates (“NHPs”).

As a result of achieving the milestone, the Company recognized $ 7.0 million in revenue, which was not previously recorded in deferred revenue, as it was fully constrained at the inception of the Takeda Collaboration.

In December 2023, the joint steering committee that manages the Takeda Collaboration terminated the SCA3 Category 1 Program as a target under the collaboration and consequently Takeda and the Company’s rights and obligations under the Takeda Collaboration were terminated with respect to the SCA3 Category 1 Program.

As a result of the termination of the SCA3 Category 1 Program, the Company recognized $ 9.9 million in revenue during the three months ended December 31, 2023, which represented the remainder of the deferred revenue for the SCA3 Category 1 Program as of September 30, 2023.

In October 2024, the Company was notified by Takeda that Takeda did not intend to exercise and therefore elected to terminate its option (“Option Termination”) for the HD target under the Takeda Collaboration Agreement.

As HD was the last active collaboration target under the Takeda Collaboration Agreement, the Takeda Collaboration Agreement expired with immediate effect, and $ 70.2 million that was previously recorded as deferred revenue was recognized as revenue in the fourth quarter of 2024 related to this expiration.

During the years ended December 31, 2024, 2023, and 2022 , the Company recognized revenue of approximately $ 71.3 million, $ 47.0 million, and $ 3.3 million, respectively, under the Takeda Collaboration Agreement in the Company’s consolidated statements of operations and comprehensive loss.

Through December 31, 2024, the Company has recognized revenue of $ 199.5 million under the Takeda Collaboration Agreement as collaboration revenue in the Company’s consolidated statements of operations and comprehensive loss.

The aggregate amount of the transaction price allocated to the Company’s unsatisfied and partially unsatisfied performance obligations which are recorded in deferred revenue as of December 31, 2024 and 2023 , is $ 0.0 million and $ 71.3 million, respectively.

In February 2018, the Company entered into a global strategic collaboration with Takeda Pharmaceutical Company Limited (“Takeda”), pursuant to which the Company agreed to collaborate with Takeda on the research, development and commercialization of oligonucleotide therapeutics for disorders of the central nervous system (the “Takeda Collaboration Agreement”).

On October 11, 2024, the Company was notified by Takeda that Takeda did not intend to exercise and therefore elected to terminate its option for the HD target under the collaboration.

As HD was the last active collaboration target under the collaboration, the collaboration expired with immediate effect.

As a result of the option termination, the Company is now free to advance WVE-003, its clinical-stage HD program, as well as any other programs targeting Huntingtin (“HTT”), independently or with other partners.

During the year ended December 31, 2025 , the Company recognized no revenue and in the years ended December 31, 2024 and 2023 , the Company recognized $ 71.3 million and $ 47.0 million in revenue, respectively, under the Takeda Collaboration Agreement in the Company’s consolidated statements of operations and comprehensive loss.

SHARE CAPITAL

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During the years ended December 31, 2025, 2024, and 2023 , the Company sold 12,226,623 , 2,952,591 , and 751,688 ordinary shares, respectively, under its "at-the-market" equity program for aggregate net proceeds of $ 94.6 million, $ 20.4 million, and $ 3.1 million, respectively, after deducting commissions and offering expenses.

• On June 16, 2022, the Company closed an underwritten offering (the “June 2022 Offering”) in which the Company issued and sold 25,464,483 of the Company’s ordinary shares at a price of $ 2.15 per share and pre-funded warrants (the “2022 Pre-Funded Warrants”) to purchase up to 7,093,656 of the Company’s ordinary shares at an offering price of $ 2.1499 per 2022 Pre-Funded Warrant, which represents the per share offering price for the ordinary shares less the $ 0.0001 per share exercise price for each 2022 Pre-Funded Warrant.

These 2022 Pre-Funded Warrants were recorded as a component of shareholders’ equity within additional paid-in capital.

The gross proceeds to the Company from the June 2022 Offering were $ 70.0 million before deducting underwriting discounts and commissions and other offering expenses.

The net proceeds to the Company from the June 2022 Offering were approximately $ 65.5 million, after deducting underwriting commissions and offering expenses.

The 2022 Pre-Funded Warrants are exercisable at any time after their original issuance and on or prior to the five-year anniversary of the original issuance date.

A holder of 2022 Pre-Funded Warrants may not exercise the warrant if the holder, together with its affiliates, would beneficially own more than 19.99 % of the number of the Company’s ordinary shares outstanding or more than 19.99 % of the combined voting power of the Company’s securities outstanding immediately after giving effect to such exercise, unless and until shareholder approval is obtained.

• On December 11, 2023, the Company closed an underwritten public offering (the “December 2023 Offering” ) in which the Company issued and sold 20,000,000 of the Company's ordinary shares at a price of $ 5.00 per share.

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3, dated as of November 12, 2024, (as amended, the “Sales Agreement”), for its “at-the-market” equity program.

• On December 11, 2025, the Company closed an underwritten public offering (the “December 2025 Offering”) in which the Company issued and sold 18,552,632 of the Company’s ordinary shares at a price of $ 19.00 per share, including 2,763,157 ordinary shares issued and sold pursuant to the underwriter's exercise in full of their option to purchase additional shares, and pre-funded warrants (the “2025 Pre-Funded Warrants”) to purchase up to 2,631,578 of the Company’s ordinary shares at an offering price of $ 18.9999 per 2025 Pre-Funded Warrant, which represents the per share offering price for the ordinary shares less the $ 0.0001 per share exercise price for each 2025 Pre-Funded Warrant.

These 2025 Pre-Funded Warrants were recorded as a component of shareholders’ equity within additional paid-in capital.

The gross proceeds to the Company from the December 2025 Offering were approximately $ 402.5 million before deducting underwriting discounts and commissions and other offering expenses, and including gross proceeds from the exercise of the Underwriters’ option to purchase additional shares in full.

The 2025 Pre-Funded Warrants are exercisable at any time after their original issuance and on or prior to the five-year anniversary of the original issuance date.

A holder of the 2025 Pre-Funded Warrants may not exercise the warrant if the holder, together with its affiliates, would beneficially own more than 4.99 % (or at the election of such holder, 9.99 % or 19.99 %) of the number of the Company’s ordinary shares outstanding or more than 4.99 % (or at the election of such holder, 9.99 % or 19.99 %) of the combined voting power of the Company’s securities outstanding immediately after giving effect to such exercise, unless and until shareholder approval is obtained.

Features of the Series A Preferred Shares and Ordinary Shares

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The Wave Life Sciences Ltd.

2021 Equity Incentive Plan was approved by the Company’s shareholders and went into effect on August 10, 2021 and was amended effective as of August 9, 2022, August 1, 2023, and August 6, 2024 (as amended, the “2021 Plan”).

2021 Equity Incentive Plan was approved by the Company’s shareholders and went into effect on August 10, 2021 and was amended effective as of August 9, 2022, August 1, 2023, August 6, 2024, and August 5, 2025 (as amended, the “2021 Plan”).

The 2021 Plan serves as the successor to the Wave Life Sciences Ltd.

2014 Equity Incentive Plan, as amended (the “2014 Plan”), such that outstanding awards granted under the 2014 Plan continue to be governed by the terms of the 2014 Plan, but no awards may be made under the 2014 Plan after August 10, 2021.

The aggregate number of ordinary shares authorized for issuance of awards under the 2021 Plan was originally 5,450,000 ordinary shares, and was subsequently increased to 11,450,000, 17,950,000 , and 22,950,000 in August 2022, August 2023, and August 2024, respectively, plus the number of ordinary shares underlying any awards under the 2014 Plan that are forfeited, cancelled or otherwise terminated (other than by exercise or withheld by the Company to satisfy any tax withholding obligation) on or after August 10, 2021.

The 2021 Plan authorizes (and the 2014 Plan previously authorized) the board of directors or a committee of the board of directors to, among other things, grant non-qualified share options, restricted awards, which include restricted shares and restricted share units (“RSUs”), and performance awards to eligible employees and directors of the Company.

The Company accounts for grants to its board of directors as grants to employees.

The aggregate number of ordinary shares authorized for issuance of awards under the 2021 Plan was originally 5,450,000 ordinary shares, and was subsequently increased to 11,450,000, 17,950,000 , 22,950,000 , and 30,950,000 in August 2022, August 2023, August 2024, and August 2025 respectively, plus the number of ordinary shares underlying any awards under the 2014 Plan that are forfeited, cancelled or otherwise terminated (other than by exercise or withheld by the Company to satisfy any tax withholding obligation) on or after August 10, 2021.

The Company accounts for grants to its non-employee directors as grants to employees.

As of December 31, 2025 , 9,808,797 ordinary shares remained available for future grant under the 2021 Plan.

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The contractual life of options is generally five or ten years from the grant date.

The assumptions used in the Black-Scholes option pricing model to determine the fair value of share options granted to employees during the period were as follows:

There were no equity grants made to consultants during the years ended December 31, 2023, 2024, and one equity grant to a consultant in 2025.

The assumptions used in the Black-Scholes option pricing model to determine the fair value of share options granted to employees, consultants, and non-employee directors during the period were as follows:

For the Year Ended December 31,

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Expected dividend yield

In October 2022, the compensation committee of the Company’s board of directors (the "Compensation Committee") granted Dr.

Verdine, one of the Company’s founders and a member of the Company’s board of directors, a non-qualified share option for 163,467 ordinary shares ( “Verdine Scientific Advisory Grant”) as form of payment under Dr.

Verdine’s consulting agreement for scientific advisory services (as described in Note 13) for the service period of October 1, 2022 through December 31, 2024, the vesting of which is subject to Dr.

Verdine’s continued service under the consulting agreement.

The Verdine Scientific Advisory Grant was granted as a non-employee grant during the year ended December 31, 2022, and there were no equity grants made to non-employees during the years ended December 31, 2023 and 2024.

The assumptions used in the Black-Scholes option pricing model to determine the fair value of the Verdine Scientific Advisory Grant were as follows:

December 31, 2022

Risk-free interest rate

Expected term (in years)

Expected volatility

Expected dividend yield

RSU activity for the year ended December 31, 2025 is summarized as follows:

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RSUs that are forfeited are available to be granted again.

During the year ended December 31, 2024 , 379,100 time-based RSUs were granted to employees.

During the year ended December 31, 2025 , 1,320,515 time-based RSUs were granted to employees and non-employee directors.

Of the RSUs outstanding at December 31, 2025 , 1,489,849 are time-based RSUs and 287,567 are performance-based RSUs.

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As of December 31, 2025, the unrecognized compensation cost related to outstanding options was $ 47.1 million.

The unrecognized compensation cost related to outstanding options is expected to be recognized over a weighted-average period of approximately 2.67 years .

The unrecognized compensation cost related to outstanding options is expected to be recognized over a weighted-average period of approximatel y 2.37 years.

For the years ended December 31, 2025 and 2024 , the weighted-average grant date fair value per granted option was $ 8.12 and $ 3.38 , respectively.

The aggregate fair value of options that vested during the years ended December 31, 2025 and 2024 was $ 124.1 million and $ 11.4 million, respectively.

The unrecognized compensation costs related to outstanding time-based RSUs was $ 2.8 million as of December 31, 2024, and is expected to be recognized over a weighted-average period of approximately 2.25 years.

The unrecognized compensation costs related to outstanding time-based RSUs was $ 11.9 million as of December 31, 2025, and is expected to be recognized over a weighted-average period of approximatel y 2.80 years.

The total fair value of RSUs vested during the years ended December 31, 2025, and 2024 was $ 2.2 million and $ 0.7 million, respectively.

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Total share-based compensation expense

Of the total share-based compensation expense recorded for the years ended December 31, 2024, 2023, and 2022 , $ 0.3 million, $ 0.2 million, and less than $ 0.1 million, respectively, were related to non-employee option grants, specifically the Verdine Scientific Advisory Grant, and all of the related expense is included in research and development expenses on the consolidated statements of operations and comprehensive loss.

Lease Arrangements

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Throughout the term of the lease, the Company is responsible for paying certain costs and expenses, in addition to the rent, as specified in the lease, including a proportionate share of applicable taxes, operating expenses and utilities.

As required under the terms of the lease agreement, the Company has placed restricted cash of approximately $ 2.8 million and $ 2.7 million in a separate bank account as of December 31, 2024 and 2023, respectively.

As required under the terms of the lease agreement, the Company has placed restricted cash of approximately $ 2.8 million in a separate bank account as of December 31, 2025 and 2024.

In April 2015, the Company entered into a lease agreement for an office and laboratory facility in Cambridge, Massachusetts (the “Cambridge Lease”), which commenced in October 2015 with a term of 7.5 years with a five-year renewal option to extend the lease .

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Operating cash flows used for operating leases

Increase in operating right-of-use assets

Operating lease liabilities arising from

obtaining right-of-use assets

Weighted average remaining lease term

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NET LOSS PER ORDINARY SHARE

In connection with the September 2024 Offering, the Company sold 1,875,023 2024 Pre-Funded Warrants, which are included in the total vested and exercisable pre-funded warrants (the 2022 Pre-Funded Warrants and the 2024 Pre-Funded Warrants are referred to together as the “Pre-Funded Warrants”).

In connection with the December 2025 Offering, the Company sold 2,631,578 2025 Pre-Funded Warrants, which are included in the total vested and exercisable pre-funded warrants (the Company's pre-funded warrants outstanding are collectively referred to as the “Pre-Funded Warrants”).

As of December 31, 2025 and 2024 , there were 11,600,257 and 8,968,679 , respectively, vested and exercisable Pre-Funded Warrants outstanding to purchase ordinary shares for the exercise price of $ 0.0001 per share, provided that, unless and until the Company obtains shareholder approval for the issuance of the shares underlying the Pre-Funded Warrants, a holder will not be entitled to exercise any portion of any Pre-Funded Warrant, which, upon giving effect to such exercise, would cause (i) the aggregate number of our ordinary shares beneficially owned by the holder (together with its affiliates) to exceed, depending on the terms of the applicable Pre-Funded Warrants and in certain cases at the election of the holder, either 4.99 %, 9.99 % or 19.99 % of the number of our ordinary shares outstanding immediately after giving effect to the exercise, or (ii) the combined voting power of our securities beneficially owned by the holder (together with its affiliates) to exceed, depending on the terms of the applicable Pre-Funded Warrants and in certain cases at the election of the holder, either 4.99 %, 9.99 % or 19.99 % of the combined voting power of all of our securities then outstanding immediately after giving effect to the exercise, as such percentage ownership is determined in accordance with the terms of the applicable Pre-Funded Warrants.

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Loss before income taxes

During the years ended December 31, 2024, 2023, and 2022 , the Company recorded no income tax benefit or provision , an income tax benefit of $ 0.7 million, and an income tax provision of $ 0.7 million, respectively.

During the years ended December 31, 2025 and 2024, the Company recorded no income tax benefit or provision.

During the year ended December 31, 2023, the Company recorded an income tax benefit of $ 0.7 million.

The income tax benefit for the year ended December 31, 2023 was due to a change in estimate in connection with U.S.

tax guidance relating to the capitalization of research and development expenditures.

The income tax provision for the year ended December 31, 2022 was primarily due to the requirement

under the Tax Cuts and Jobs Act of 2017 for taxpayers to capitalize and amortize research and development expenditures over five or fifteen years pursuant to Section 174 of the Internal Revenue Code of 1986, as amended (the “Code”).

The components of the benefit (provision) for income taxes were as follows:

The components of the benefit for income taxes were as follows:

Year Ended December 31,

(in thousands)

Current benefit (provision) for income taxes:

Current benefit for income taxes:

Rest of world

Total current benefit (provision) for income taxes

Total current benefit for income taxes

Deferred benefit for income taxes:

Rest of world

Total deferred benefit (provision) for income taxes

Total benefit (provision) for income taxes

Total deferred benefit for income taxes

Total benefit for income taxes

A reconciliation of the Singapore statutory income tax rate to the Company’s effective income tax rate is as follows:

1 unchanged sentence

Singapore statutory income tax rate

Foreign tax effects

United States

Statutory tax rate difference between U.S.

federal and Singapore

Research and development tax credits

Orphan drug credit

Change in valuation allowances

Nontaxable or nondeductible items

Nondeductible executive compensation

Stock compensation

Statutory tax rate difference between the United Kingdom and Singapore

Research and development

Changes in valuation allowances

Other Foreign Jurisdictions

Changes in valuation allowances

Nontaxable or nondeductible items

Changes in unrecognized tax benefits

Effective income tax rate

(1) Includes United States state and local income taxes, net of federal income tax effect.

A reconciliation of the Singapore statutory income tax rate to the Company’s effective income tax rate is as follows:

Year Ended December 31,

Singapore statutory income tax rate

Federal and state tax credits

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As of December 31, 2025 , the Company had federal net operating loss carryforwards in the United States of $ 501.3 million, of which $ 500.5 million may be available to offset future U.S.

federal taxable income indefinitely, while $ 0.8 million of carryforwards may offset future U.S.

federal taxable income indefinitely, while $ 0.8 million of the carryforwards may offset future U.S.

federal taxable income through 2037 .

3 unchanged sentences

As of December 31, 2025 and 2024 , the Company had U.S.

federal research and development tax credit carryforwards of approximately $ 11.2 million and $ 6.1 million, respectively, available to offset future U.S.

federal research and development tax credit carryforwards of $ 15.1 million and $11 .2 million, respectively, available to offset future U.S.

federal income taxes and will begin to expire in 2042 .

As of December 31, 2025 and 2024 , the Company had U.S.

state research and development tax credit carryforwards of approximately $ 4.8 million and $ 2.6 million, respectively, available to offset future U.S.

state research and development tax credit carryforwards of $ 7.0 million and $4 .8 million, respectively, available to offset future U.S.

state income taxes and will begin to expire in 2037 .

As of December 31, 2024 , the Company had a U.S.

orphan drug credit carryforward of $ 0.8 million available to offset future U.S.

As of December 31, 2025 and 2024 , the Company had a U.S.

orphan drug credit carryforward of $ 3.7 million and $ 0.8 million, respectively, available to offset future U.S.

federal income taxes that will begin to expire in 2042 .

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Tax positions related to the current year

Tax position releases

Unrecognized tax benefit at the end of the year

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At December 31, 2025 , no ne of the net unrecognized tax benefits would affect the Company’s effective tax rate due to the Company's full valuation allowance.

The Company anticipates that $ 2.6 million of the total unrecognized tax benefits at December 31, 2024 will decrease within the next twelve months due to certain tax return filings.

The Company files income tax returns as prescribed by the tax laws of the jurisdictions in which it operates.

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Because of the complexity of Singapore and the rest-of-the-world tax rules applicable to the method of recovery of the investment in its subsidiaries, including distribution of earnings from its subsidiaries to Singapore, the determination of the unrecognized deferred tax liability is not practicable.

Utilization of the net operating loss carryforwards and research and development tax credit carryforwards in the United States may be subject to a substantial annual limitation under Section 382 of the Code, due to ownership changes that have occurred previously or that could occur in the future.

These ownership changes may limit the amount of carryforwards that can be utilized annually to offset future taxable income.

Utilization of the net operating loss carryforwards and research and development tax credit carryforwards in the United States may be subject to a substantial annual limitation under Section 382 and Section 383 of the Code, due to ownership changes that have occurred previously or that could occur in the future.

These ownership changes may limit the amount of carryforwards that can be utilized annually to offset future taxable income and tax.

In general, an ownership change, as defined by Section 382, results from transactions increasing the ownership of certain shareholders or public groups in the shares of a corporation by more than 50% over a three-year period.

The Company has completed Section 382 studies to assess whether there have been ownership changes since its formation through 2022.

The results of the studies indicated that the Company experienced ownership changes as defined by Section 382 of the Code, and, as such, the Company has adjusted its net operating losses and research and development credit carryforwards to reflect the limitations as a result of such ownership changes.

The Company previously completed Section 382 studies to assess whether there have been ownership changes since its formation through 2022.

The results of the studies indicated that the Company experienced ownership changes as defined by Section 382 of the Code, and, as such, the Company adjusted its net operating losses and research and development credit carryforwards to reflect the limitations as a result of such ownership changes.

The Company updated its Section 382 study to assess whether there have been ownership changes since December 31, 2022 through 2024 and the Company did not experience any additional ownership changes.

Should one or more ownership changes occur in the future, the Company’s ability to utilize its net operating losses and research and development credit carryforwards may be further limited.

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Under the 401(k) Plan, the Company may make discretionary contributions as approved by the board of directors.

The Company made contributions of $ 1.7 million and $ 1.4 million in the years ended December 31, 2024 and 2023 , respectively.

The Company made contributions of $ 2.0 million and $ 1.7 million i n the years ended December 31, 2025 and 2024 , respectively.

RELATED PARTIES

The Company had the following related party transactions for the periods presented in the accompanying consolidated financial statements:

• In 2012, the Company entered into a consulting agreement for scientific advisory services with Dr.

Verdine, one of the Company’s founders and a member of the Company’s board of directors.

The consulting agreement does not have a specific term and may be terminated by either party upon 14 days’ prior written notice.

Pursuant to the consulting agreement, the Company pays Dr.

Verdine approximately $ 13 thousand per month, plus reimbursement for certain expenses.

In October 2022, the Compensation Committee granted Dr.

Verdine a non-qualified share option for 163,467 ordinary shares in lieu of cash as payment under this consulting agreement for the service period of October 1, 2022 through December 31, 2024, the monthly vesting of which is subject to Dr.

Verdine’s continued service under the consulting agreement.

• In April 2023, the Company engaged Shin Nippon Biomedical Laboratories Ltd.

(“SNBL”), one of the Company’s shareholders, to provide approximately $ 2.8 million in certain NHPs contract research services to the Company.

During the years ended December 31, 2024 and 2023 , the Company made payments of $ 0.9 million and $ 1.4 million, respectively, to SNBL.

Through December 31, 2024 , the Company has paid $ 2.3 million to SNBL for the aforementioned NHP contract research services.

• Pursuant to the terms of various contract research service agreements with Shin Nippon Biomedical Laboratories Ltd., a contract research organization and one of the Company’s shareholders, and its affiliates (together, “SNBL”), the Company paid SNBL $ 1.1 million and $ 0.9 million, respectively, during each of the years ended December 31, 2025 and 2024.

Through December 31, 2025 , the Company has paid an aggregate of $ 3.4 million to SNBL for the aforementioned various service agreements.

During each of the years ended December 31, 2025 and 2024 , the Company incurred expenses of approximately $ 4.6 million and $ 1.0 million, respectively, related to the aforementioned various service agreements.

SEGMENT INFORMATION

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Research and development expenses:

Other research and development expenses (1) , including INHBE, RNA editing, PRISM, others

ALS and FTD programs (discontinued)

INHBE program

Other research and development expenses(1), including PNPLA3, additional preclinical programs, PRISM

Total research and development expenses

3 unchanged sentences

Compared sentence by sentence after normalising whitespace, quotation marks, case and digits, so re-formatting and restated figures do not read as changed language. Wording changes appear as one removal and one addition. The current filing and the prior one are authoritative.

What changed 10-K

Item 7. Management’s Discussion and Analysis