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Stock-based Compensation - Expense (Details) - USD ($)
$ in Thousands
12 Months Ended
Dec. 31, 2021
Dec. 31, 2020
Dec. 31, 2019
Stock-based compensation expense      
Stock-based compensation expense $ 26,873 $ 20,730 $ 22,759
Research and development      
Stock-based compensation expense      
Stock-based compensation expense 10,698 8,833 8,277
General and administrative      
Stock-based compensation expense      
Stock-based compensation expense 16,175 11,897 14,482
Stock options      
Stock-based compensation expense      
Stock-based compensation expense 26,873 20,730 22,487
Unvested restricted common stock      
Stock-based compensation expense      
Stock-based compensation expense $ 0 $ 0 $ 272