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Stock-Based Compensation - Accounting for Stock-based Compensation (Details) - USD ($)
$ / shares in Units, $ in Thousands
12 Months Ended
Dec. 31, 2019
Dec. 31, 2018
Dec. 31, 2017
Share-based Compensation Arrangement by Share-based Payment Award      
Percentage of share-based compensation expense 15.00%    
Weighted average grant date fair value of options granted during the period (per share) $ 1.64 $ 7.00 $ 8.08
Stock-based compensation $ 3,845 $ 5,674 $ 10,720
Research and development      
Share-based Compensation Arrangement by Share-based Payment Award      
Stock-based compensation 1,348 1,851 6,494
Research and development | Former Executive      
Share-based Compensation Arrangement by Share-based Payment Award      
Stock-based compensation     4,300
Research and development | Employee Stock Purchase Plans      
Share-based Compensation Arrangement by Share-based Payment Award      
Stock-based compensation 36 71 96
Research and development | Equity Incentive Plans      
Share-based Compensation Arrangement by Share-based Payment Award      
Stock-based compensation 1,312 1,780 6,398
General and administrative      
Share-based Compensation Arrangement by Share-based Payment Award      
Stock-based compensation 2,497 3,799 4,226
General and administrative | Employee Stock Purchase Plans      
Share-based Compensation Arrangement by Share-based Payment Award      
Stock-based compensation 20 48 63
General and administrative | Equity Incentive Plans      
Share-based Compensation Arrangement by Share-based Payment Award      
Stock-based compensation $ 2,477 3,751 $ 4,163
Restructuring costs      
Share-based Compensation Arrangement by Share-based Payment Award      
Stock-based compensation   24  
Restructuring costs | Equity Incentive Plans      
Share-based Compensation Arrangement by Share-based Payment Award      
Stock-based compensation   $ 24