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Equity Plans - Stock-based Compensation Expense (Details) - USD ($)
$ in Thousands
3 Months Ended
Mar. 31, 2020
Mar. 31, 2019
Share-based Compensation Arrangement by Share-based Payment Award, Compensation Cost [Line Items]    
Total stock-based compensation expense $ 2,048 $ 1,979
Total unrecognized stock-based compensation costs related to stock options $ 16,200  
Period of unrecognized stock-based compensation costs to be recognized 3 years 1 month 6 days  
Research and Development    
Share-based Compensation Arrangement by Share-based Payment Award, Compensation Cost [Line Items]    
Total stock-based compensation expense $ 1,066 1,122
General and Administrative    
Share-based Compensation Arrangement by Share-based Payment Award, Compensation Cost [Line Items]    
Total stock-based compensation expense $ 982 $ 857