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Stock-Based Compensation - Employee expense (Details) - USD ($)
$ in Thousands
 12 Months Ended
Dec. 31, 2021
Dec. 31, 2020
Dec. 31, 2019
Stock-based compensation      
Total employee, director and non-employee stock-based compensation expense $ 36,975 $ 31,619 $ 31,851
Options to purchase common stock      
Stock-based compensation      
Total employee, director and non-employee stock-based compensation expense 27,909 26,045 30,502
Employee stock purchase plan      
Stock-based compensation      
Total employee, director and non-employee stock-based compensation expense 992 804 687
Restricted stock units      
Stock-based compensation      
Total employee, director and non-employee stock-based compensation expense 8,074 4,770 662
General and administrative      
Stock-based compensation      
Total employee, director and non-employee stock-based compensation expense 12,813 10,769 8,854
Research and development      
Stock-based compensation      
Total employee, director and non-employee stock-based compensation expense $ 24,162 $ 20,850 $ 22,997