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Equity Incentive Plans - Stock-Based Compensation (Details) - USD ($)
$ in Thousands
3 Months Ended 6 Months Ended
Jun. 30, 2021
Jun. 30, 2020
Jun. 30, 2021
Jun. 30, 2020
Stock-based Compensation [Line Items]        
Stock-based compensation expense $ 3,219 $ 2,674 $ 6,306 $ 5,622
Research and development        
Stock-based Compensation [Line Items]        
Stock-based compensation expense 1,087 1,144 2,179 2,202
General and administrative        
Stock-based Compensation [Line Items]        
Stock-based compensation expense $ 2,132 $ 1,530 $ 4,127 $ 3,420