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Stock-Based Compensation - Accounting for Stock-based Compensation (Details) - USD ($)
$ / shares in Units, $ in Thousands
3 Months Ended
Mar. 31, 2018
Mar. 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) $ 1.24 $ 0.99
Stock-based compensation $ 1,589 $ 1,784
Research and development    
Share-based Compensation Arrangement by Share-based Payment Award    
Stock-based compensation 578 736
Research and development | Employee Stock Purchase Plans    
Share-based Compensation Arrangement by Share-based Payment Award    
Stock-based compensation 22 22
Research and development | Equity Incentive Plans    
Share-based Compensation Arrangement by Share-based Payment Award    
Stock-based compensation 556 714
General and administrative    
Share-based Compensation Arrangement by Share-based Payment Award    
Stock-based compensation 1,011 1,048
General and administrative | Employee Stock Purchase Plans    
Share-based Compensation Arrangement by Share-based Payment Award    
Stock-based compensation 14 19
General and administrative | Equity Incentive Plans    
Share-based Compensation Arrangement by Share-based Payment Award    
Stock-based compensation $ 997 $ 1,029