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Equity Plans - Stock-based Compensation Expense (Details) - USD ($)
$ in Thousands
3 Months Ended 6 Months Ended
Jun. 30, 2020
Jun. 30, 2019
Jun. 30, 2020
Jun. 30, 2019
Share-based Compensation Arrangement by Share-based Payment Award, Compensation Cost [Line Items]        
Total stock-based compensation expense $ 1,996 $ 2,013 $ 4,044 $ 3,992
Total unrecognized stock-based compensation costs related to stock options 14,700   $ 14,700  
Period of unrecognized stock-based compensation costs to be recognized     2 years 10 months 24 days  
Research and Development        
Share-based Compensation Arrangement by Share-based Payment Award, Compensation Cost [Line Items]        
Total stock-based compensation expense 1,026 977 $ 2,092 2,100
General and Administrative        
Share-based Compensation Arrangement by Share-based Payment Award, Compensation Cost [Line Items]        
Total stock-based compensation expense $ 970 $ 1,036 $ 1,952 $ 1,892