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Stock-Based Compensation - Accounting for Stock-based Compensation (Details) - USD ($)
$ / shares in Units, $ in Thousands
3 Months Ended 9 Months Ended
Sep. 30, 2019
Sep. 30, 2018
Sep. 30, 2019
Sep. 30, 2018
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)     $ 1.65 $ 7.15
Stock-based compensation $ 963 $ 1,327 $ 2,868 $ 4,454
Research and development        
Share-based Compensation Arrangement by Share-based Payment Award        
Stock-based compensation 337 314 1,005 1,441
Research and development | Employee Stock Purchase Plans        
Share-based Compensation Arrangement by Share-based Payment Award        
Stock-based compensation 9 11 27 62
Research and development | Equity Incentive Plans        
Share-based Compensation Arrangement by Share-based Payment Award        
Stock-based compensation 328 303 978 1,379
General and administrative        
Share-based Compensation Arrangement by Share-based Payment Award        
Stock-based compensation 626 989 1,863 2,989
General and administrative | Employee Stock Purchase Plans        
Share-based Compensation Arrangement by Share-based Payment Award        
Stock-based compensation 4 8 18 40
General and administrative | Equity Incentive Plans        
Share-based Compensation Arrangement by Share-based Payment Award        
Stock-based compensation $ 622 981 $ 1,845 2,949
Restructuring costs        
Share-based Compensation Arrangement by Share-based Payment Award        
Stock-based compensation   24   24
Restructuring costs | Employee Stock Purchase Plans        
Share-based Compensation Arrangement by Share-based Payment Award        
Stock-based compensation   $ 24   $ 24