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Basis of Presentation and Significant Accounting Policies (Details) - USD ($)
$ / shares in Units, $ in Thousands
 3 Months Ended
Mar. 31, 2018
Mar. 31, 2017
Jan. 01, 2018
Dec. 31, 2017
Revenue, Initial Application Period Cumulative Effect Transition [Line Items]        
Deferred revenue, current $ 8,121   $ 7,742 $ 7,202
Deferred revenue, net of current portion 17,762   19,576 13,637
Accumulated deficit 368,355   318,819 312,340
Revenue from collaborative agreements 4,501 $ 1,278    
Net loss $ (49,536) $ (37,655)    
Basic and diluted net loss per common share (in usd per share) $ (1.34) $ (1.08)    
Change in Contract with Customer, Liability [Roll Forward] [Roll Forward]        
Beginning balance $ 27,318      
Additions 500      
Deductions (1,935)      
Ending balance 25,883      
Calculated under Revenue Guidance in Effect before Topic 606        
Revenue, Initial Application Period Cumulative Effect Transition [Line Items]        
Deferred revenue, current 7,417     7,202
Deferred revenue, net of current portion 12,122     13,637
Accumulated deficit 362,011     $ 312,340
Revenue from collaborative agreements 4,366      
Net loss $ (49,671)      
Basic and diluted net loss per common share (in usd per share) $ (1.35)      
Accounting Standards Update 2014-09 | Difference between Revenue Guidance in Effect before and after Topic 606        
Revenue, Initial Application Period Cumulative Effect Transition [Line Items]        
Deferred revenue, current $ 704   540  
Deferred revenue, net of current portion 5,640   5,939  
Accumulated deficit 6,344   $ 6,479  
Revenue from collaborative agreements 135      
Net loss $ (135)      
Basic and diluted net loss per common share (in usd per share) $ 0.01