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Pay vs Performance Disclosure - USD ($) | 12 Months Ended
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Dec. 31, 2024 | Dec. 31, 2023 | Dec. 31, 2022
Pay vs Performance Disclosure |  |  |
Pay vs Performance Disclosure, Table | ​
​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​
​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | Value of Initial | ​ | ​ | ​
​ | ​ | ​ | ​ | ​ | ​ | Average | ​ | Average | ​ | Fixed $100 | ​ | ​ | ​
​ | ​ | ​ | ​ | ​ | ​ | Summary | ​ | Compensation | ​ | Investment Based | ​ | ​ | ​
​ | ​ | Summary | ​ | Compensation | ​ | Compensation | ​ | Actually | ​ | on Total | ​ | ​ | ​
​ | ​ | Compensation | ​ | Actually | ​ | Table for | ​ | Paid to the | ​ | Shareholder | ​ | Net Income (Loss)
​ | ​ | Total for PEO | ​ | Paid to PEO | ​ | Non-PEO NEOs | ​ | Non-PEO NEOs | ​ | Return | ​ | (in thousands)
Year |  | ($)(1) |  | ($)(2) |  | ($)(1) |  | ($)(2) |  | ($)(3) |  | ($)(4)
2024 | ​ | 6,875,603 | ​ | 6,053,171 | ​ | 4,359,533 | ​ | 3,827,994 | ​ | ​ | 25.65 | ​ | ​ | (147,791)
2023 |  | 6,336,832 |  | 5,837,614 |  | 3,740,448 |  | 3,465,959 | ​ | ​ | 29.57 | ​ | ​ | (84,027)
2022 |  | 7,715,156 |  | (1,124,733) |  | 5,291,480 |  | (1,332,788) | ​ | ​ | 27.46 | ​ | ​ | (129,615)
(1) | During 2024, 2023 and 2022, our PEO was Alexandria Forbes, Ph.D. and our non-PEO NEOs consisted of Richard Giroux and Stuart Naylor, Ph.D. The amounts shown in these columns represent the total compensation for the PEO and an average of the total compensation for the non-PEO NEOs as reported in the Summary Compensation Table for 2024, 2023 and 2022.
(2) | The dollar amounts reported are adjusted to calculate “compensation actually paid” to the PEO and the average “compensation actually paid” to the two non-PEO NEOs as computed in accordance with Item 402(v) of Regulation S-K. The dollar amounts do not reflect the actual amount of compensation earned by or paid to the NEOs during the applicable year. In accordance with the requirements of Item 402(v) of Regulation S-K, the following adjustments were made to the PEO and non-PEO NEOs’ total compensation reported in the Summary Compensation Table for fiscal years 2024, 2023 and 2022 to determine the “compensation actually paid”:
​
​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​
​ | ​ | 2024 | ​ | 2023 | ​ | 2022
​ | ​ | ​ | ​ | Average Non- | ​ | ​ | ​ | Average Non- | ​ | ​ | ​ | Average Non-
​ | ​ | ​ | ​ | PEO | ​ | ​ | ​ | PEO | ​ | ​ | ​ | PEO
Adjustments |  | PEO ($) |  | NEOs ($) |  | PEO ($) |  | NEOs ($) |  | PEO ($) |  | NEOs ($)
Deduction for amounts reported under the “Stock Awards” and “Option Awards” columns in the Summary Compensation Table |  | (3,819,200) | ​ | (2,371,600) | ​ | (3,448,250) |  | (2,139,975) |  | (5,382,500) |  | (3,875,400)
Increase for fair value of awards granted during year that remained unvested at year-end |  | 3,775,800 | ​ | 2,344,650 | ​ | 2,751,975 |  | 1,714,043 |  | 1,630,000 |  | 1,173,600
Change in fair value from prior year-end to year-end of awards granted in a prior year that were outstanding and unvested at year-end |  | (491,309) | ​ | (324,412) | ​ | 177,685 |  | 132,775 |  | (4,220,708) |  | (3,287,749)
Change in fair value from prior year-end to vesting date of awards granted in a prior year that vested during year |  | (287,722) | ​ | (180,177) | ​ | 19,372 |  | 18,669 |  | (866,681) |  | (634,719)
Total adjustments |  | (822,432) | ​ | (531,539) | ​ | (499,218) |  | (274,489) |  | (8,839,889) |  | (6,624,268)
​
(3) | Cumulative Total Shareholder Return (“TSR”) is the calculated difference between our share price at the end and the beginning of the applicable measurement period, and our share price at the beginning of the measurement period. To perform this calculation, we assume a $100 investment on December 31, 2021.
(4) | The dollar amount reported represents the amount of net income (loss) reflected in our audited financial statements for the applicable year.
Named Executive Officers, Footnote |
(1) | During 2024, 2023 and 2022, our PEO was Alexandria Forbes, Ph.D. and our non-PEO NEOs consisted of Richard Giroux and Stuart Naylor, Ph.D. The amounts shown in these columns represent the total compensation for the PEO and an average of the total compensation for the non-PEO NEOs as reported in the Summary Compensation Table for 2024, 2023 and 2022.
PEO Total Compensation Amount | $ 6,875,603 | $ 6,336,832 | $ 7,715,156
PEO Actually Paid Compensation Amount | $ 6,053,171 | 5,837,614 | (1,124,733)
Adjustment To PEO Compensation, Footnote |
(2) | The dollar amounts reported are adjusted to calculate “compensation actually paid” to the PEO and the average “compensation actually paid” to the two non-PEO NEOs as computed in accordance with Item 402(v) of Regulation S-K. The dollar amounts do not reflect the actual amount of compensation earned by or paid to the NEOs during the applicable year. In accordance with the requirements of Item 402(v) of Regulation S-K, the following adjustments were made to the PEO and non-PEO NEOs’ total compensation reported in the Summary Compensation Table for fiscal years 2024, 2023 and 2022 to determine the “compensation actually paid”:
​
​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​
​ | ​ | 2024 | ​ | 2023 | ​ | 2022
​ | ​ | ​ | ​ | Average Non- | ​ | ​ | ​ | Average Non- | ​ | ​ | ​ | Average Non-
​ | ​ | ​ | ​ | PEO | ​ | ​ | ​ | PEO | ​ | ​ | ​ | PEO
Adjustments |  | PEO ($) |  | NEOs ($) |  | PEO ($) |  | NEOs ($) |  | PEO ($) |  | NEOs ($)
Deduction for amounts reported under the “Stock Awards” and “Option Awards” columns in the Summary Compensation Table |  | (3,819,200) | ​ | (2,371,600) | ​ | (3,448,250) |  | (2,139,975) |  | (5,382,500) |  | (3,875,400)
Increase for fair value of awards granted during year that remained unvested at year-end |  | 3,775,800 | ​ | 2,344,650 | ​ | 2,751,975 |  | 1,714,043 |  | 1,630,000 |  | 1,173,600
Change in fair value from prior year-end to year-end of awards granted in a prior year that were outstanding and unvested at year-end |  | (491,309) | ​ | (324,412) | ​ | 177,685 |  | 132,775 |  | (4,220,708) |  | (3,287,749)
Change in fair value from prior year-end to vesting date of awards granted in a prior year that vested during year |  | (287,722) | ​ | (180,177) | ​ | 19,372 |  | 18,669 |  | (866,681) |  | (634,719)
Total adjustments |  | (822,432) | ​ | (531,539) | ​ | (499,218) |  | (274,489) |  | (8,839,889) |  | (6,624,268)
Non-PEO NEO Average Total Compensation Amount | $ 4,359,533 | 3,740,448 | 5,291,480
Non-PEO NEO Average Compensation Actually Paid Amount | $ 3,827,994 | 3,465,959 | (1,332,788)
Adjustment to Non-PEO NEO Compensation Footnote |
(2) | The dollar amounts reported are adjusted to calculate “compensation actually paid” to the PEO and the average “compensation actually paid” to the two non-PEO NEOs as computed in accordance with Item 402(v) of Regulation S-K. The dollar amounts do not reflect the actual amount of compensation earned by or paid to the NEOs during the applicable year. In accordance with the requirements of Item 402(v) of Regulation S-K, the following adjustments were made to the PEO and non-PEO NEOs’ total compensation reported in the Summary Compensation Table for fiscal years 2024, 2023 and 2022 to determine the “compensation actually paid”:
​
​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​
​ | ​ | 2024 | ​ | 2023 | ​ | 2022
​ | ​ | ​ | ​ | Average Non- | ​ | ​ | ​ | Average Non- | ​ | ​ | ​ | Average Non-
​ | ​ | ​ | ​ | PEO | ​ | ​ | ​ | PEO | ​ | ​ | ​ | PEO
Adjustments |  | PEO ($) |  | NEOs ($) |  | PEO ($) |  | NEOs ($) |  | PEO ($) |  | NEOs ($)
Deduction for amounts reported under the “Stock Awards” and “Option Awards” columns in the Summary Compensation Table |  | (3,819,200) | ​ | (2,371,600) | ​ | (3,448,250) |  | (2,139,975) |  | (5,382,500) |  | (3,875,400)
Increase for fair value of awards granted during year that remained unvested at year-end |  | 3,775,800 | ​ | 2,344,650 | ​ | 2,751,975 |  | 1,714,043 |  | 1,630,000 |  | 1,173,600
Change in fair value from prior year-end to year-end of awards granted in a prior year that were outstanding and unvested at year-end |  | (491,309) | ​ | (324,412) | ​ | 177,685 |  | 132,775 |  | (4,220,708) |  | (3,287,749)
Change in fair value from prior year-end to vesting date of awards granted in a prior year that vested during year |  | (287,722) | ​ | (180,177) | ​ | 19,372 |  | 18,669 |  | (866,681) |  | (634,719)
Total adjustments |  | (822,432) | ​ | (531,539) | ​ | (499,218) |  | (274,489) |  | (8,839,889) |  | (6,624,268)
Compensation Actually Paid vs. Total Shareholder Return |  |  |
Compensation Actually Paid vs. Net Income |  |  |
Total Shareholder Return Amount | $ 25.65 | 29.57 | 27.46
Net Income (Loss) | $ (147,791,000) | (84,027,000) | (129,615,000)
PEO Name | Alexandria Forbes |  |
Equity Awards Adjustments, Footnote | In accordance with the requirements of Item 402(v) of Regulation S-K, the following adjustments were made to the PEO and non-PEO NEOs’ total compensation reported in the Summary Compensation Table for fiscal years 2024, 2023 and 2022 to determine the “compensation actually paid”: ​
​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​ | ​
​ | ​ | 2024 | ​ | 2023 | ​ | 2022
​ | ​ | ​ | ​ | Average Non- | ​ | ​ | ​ | Average Non- | ​ | ​ | ​ | Average Non-
​ | ​ | ​ | ​ | PEO | ​ | ​ | ​ | PEO | ​ | ​ | ​ | PEO
Adjustments |  | PEO ($) |  | NEOs ($) |  | PEO ($) |  | NEOs ($) |  | PEO ($) |  | NEOs ($)
Deduction for amounts reported under the “Stock Awards” and “Option Awards” columns in the Summary Compensation Table |  | (3,819,200) | ​ | (2,371,600) | ​ | (3,448,250) |  | (2,139,975) |  | (5,382,500) |  | (3,875,400)
Increase for fair value of awards granted during year that remained unvested at year-end |  | 3,775,800 | ​ | 2,344,650 | ​ | 2,751,975 |  | 1,714,043 |  | 1,630,000 |  | 1,173,600
Change in fair value from prior year-end to year-end of awards granted in a prior year that were outstanding and unvested at year-end |  | (491,309) | ​ | (324,412) | ​ | 177,685 |  | 132,775 |  | (4,220,708) |  | (3,287,749)
Change in fair value from prior year-end to vesting date of awards granted in a prior year that vested during year |  | (287,722) | ​ | (180,177) | ​ | 19,372 |  | 18,669 |  | (866,681) |  | (634,719)
Total adjustments |  | (822,432) | ​ | (531,539) | ​ | (499,218) |  | (274,489) |  | (8,839,889) |  | (6,624,268)
PEO | Equity Awards Adjustments |  |  |
Pay vs Performance Disclosure |  |  |
Adjustment to Compensation, Amount | $ (822,432) | (499,218) | (8,839,889)
PEO | Aggregate Grant Date Fair Value of Equity Award Amounts Reported in Summary Compensation Table |  |  |
Pay vs Performance Disclosure |  |  |
Adjustment to Compensation, Amount | (3,819,200) | (3,448,250) | (5,382,500)
PEO | Year-end Fair Value of Equity Awards Granted in Covered Year that are Outstanding and Unvested |  |  |
Pay vs Performance Disclosure |  |  |
Adjustment to Compensation, Amount | 3,775,800 | 2,751,975 | 1,630,000
PEO | Year-over-Year Change in Fair Value of Equity Awards Granted in Prior Years That are Outstanding and Unvested |  |  |
Pay vs Performance Disclosure |  |  |
Adjustment to Compensation, Amount | (491,309) | 177,685 | (4,220,708)
PEO | Change in Fair Value as of Vesting Date of Prior Year Equity Awards Vested in Covered Year |  |  |
Pay vs Performance Disclosure |  |  |
Adjustment to Compensation, Amount | (287,722) | 19,372 | (866,681)
Non-PEO NEO | Equity Awards Adjustments |  |  |
Pay vs Performance Disclosure |  |  |
Adjustment to Compensation, Amount | (531,539) | (274,489) | (6,624,268)
Non-PEO NEO | Aggregate Grant Date Fair Value of Equity Award Amounts Reported in Summary Compensation Table |  |  |
Pay vs Performance Disclosure |  |  |
Adjustment to Compensation, Amount | (2,371,600) | (2,139,975) | (3,875,400)
Non-PEO NEO | Year-end Fair Value of Equity Awards Granted in Covered Year that are Outstanding and Unvested |  |  |
Pay vs Performance Disclosure |  |  |
Adjustment to Compensation, Amount | 2,344,650 | 1,714,043 | 1,173,600
Non-PEO NEO | Year-over-Year Change in Fair Value of Equity Awards Granted in Prior Years That are Outstanding and Unvested |  |  |
Pay vs Performance Disclosure |  |  |
Adjustment to Compensation, Amount | (324,412) | 132,775 | (3,287,749)
Non-PEO NEO | Change in Fair Value as of Vesting Date of Prior Year Equity Awards Vested in Covered Year |  |  |
Pay vs Performance Disclosure |  |  |
Adjustment to Compensation, Amount | $ (180,177) | $ 18,669 | $ (634,719)

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