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Equity Incentive Plans - Allocation of Share-based Compensation Expense (Details) - USD ($) $ in Thousands | 3 Months Ended | 9 Months Ended
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Sep. 30, 2020 | Sep. 30, 2019 | Sep. 30, 2020 | Sep. 30, 2019
Share-based Compensation Arrangement by Share-based Payment Award [Line Items] |  |  |  |
Stock-based compensation | $ 2,805 | $ 3,144 | $ 8,521 | $ 9,654
Research and development |  |  |  |
Share-based Compensation Arrangement by Share-based Payment Award [Line Items] |  |  |  |
Stock-based compensation | 926 | 1,580 | 3,030 | 4,469
General and administrative |  |  |  |
Share-based Compensation Arrangement by Share-based Payment Award [Line Items] |  |  |  |
Stock-based compensation | $ 1,879 | $ 1,564 | $ 5,491 | $ 5,185

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