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Equity Incentive Plans - Stock-Based Compensation (Details) - USD ($)
$ in Thousands
3 Months Ended 9 Months Ended
Sep. 30, 2021
Sep. 30, 2020
Sep. 30, 2021
Sep. 30, 2020
Stock-based Compensation [Line Items]        
Stock-based compensation expense $ 2,262 $ 2,527 $ 8,568 $ 8,149
Research and development        
Stock-based Compensation [Line Items]        
Stock-based compensation expense 730 963 2,909 3,165
General and administrative        
Stock-based Compensation [Line Items]        
Stock-based compensation expense $ 1,532 $ 1,564 $ 5,659 $ 4,984