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Stockholders' Equity - Stock Option Plan Schedule of Stock-based Compensation Expense (Details) - USD ($)
$ in Thousands
3 Months Ended 9 Months Ended
Sep. 30, 2017
Sep. 30, 2016
Sep. 30, 2017
Sep. 30, 2016
Share-based Compensation Arrangement by Share-based Payment Award, Compensation Cost [Line Items]        
Total stock based compensation expense $ 3,146 $ 2,755 $ 9,820 $ 8,984
Research and Development        
Share-based Compensation Arrangement by Share-based Payment Award, Compensation Cost [Line Items]        
Total stock based compensation expense 1,534 1,124 4,341 4,325
General and Administrative        
Share-based Compensation Arrangement by Share-based Payment Award, Compensation Cost [Line Items]        
Total stock based compensation expense $ 1,612 $ 1,631 $ 5,479 $ 4,659