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Summary of Significant Accounting Policies - Adoption of 606 (Details) - USD ($)
$ / shares in Units, $ in Thousands
 3 Months Ended 12 Months Ended
Dec. 31, 2019
Sep. 30, 2019
Jun. 30, 2019
Mar. 31, 2019
Dec. 31, 2018
Sep. 30, 2018
Jun. 30, 2018
Mar. 31, 2018
Dec. 31, 2019
Dec. 31, 2018
Dec. 31, 2017
Jan. 01, 2018
Revenue, Initial Application Period Cumulative Effect Transition [Line Items]                        
Deferred revenue, current $ 10,700       $ 21,721       $ 10,700 $ 21,721    
Deferred revenue, net of current portion 9,153       19,001       9,153 19,001    
Accumulated deficit (642,082)       (490,271)       (642,082) (490,271)    
Revenues 25,192 $ 18,741 $ 10,593 $ 9,662 15,794 $ 20,798 $ 18,834 $ 4,695 64,188 60,121 $ 157,742  
Net loss $ (30,396) $ (44,631) $ (31,767) $ (45,017) (44,644) $ (34,029) $ (43,244) $ (49,536) $ (151,811) $ (171,453) $ (19,626)  
Basic and diluted net loss per common share (in usd per share)                 $ (3.16) $ (4.19) $ (0.54)  
Accounting Standards Update 2014-09                        
Revenue, Initial Application Period Cumulative Effect Transition [Line Items]                        
Deferred revenue, current         21,721         $ 21,721   $ 7,742
Deferred revenue, net of current portion         19,001         19,001   19,576
Accumulated deficit         (490,271)         (490,271)   (318,818)
Net loss                   $ (171,453)    
Basic and diluted net loss per common share (in usd per share)                   $ (4.19)    
Accounting Standards Update 2014-09 | Pre-Adoption                        
Revenue, Initial Application Period Cumulative Effect Transition [Line Items]                        
Deferred revenue, current         22,210         $ 22,210   7,202
Deferred revenue, net of current portion         12,573         12,573   13,637
Accumulated deficit         (484,332)         (484,332)   (312,340)
Net loss                   $ (171,993)    
Basic and diluted net loss per common share (in usd per share)                   $ (4.20)    
Accounting Standards Update 2014-09 | ASC 606 Adjustment                        
Revenue, Initial Application Period Cumulative Effect Transition [Line Items]                        
Deferred revenue, current         (489)         $ (489)   540
Deferred revenue, net of current portion         6,428         6,428   5,939
Accumulated deficit         $ (5,939)         (5,939)   $ (6,478)
Net loss                   $ (540)    
Basic and diluted net loss per common share (in usd per share)                   $ 0.01    
Revenue from collaborative agreements                        
Revenue, Initial Application Period Cumulative Effect Transition [Line Items]                        
Revenues                 $ 62,024 $ 58,644 $ 155,516  
Revenue from collaborative agreements | Accounting Standards Update 2014-09                        
Revenue, Initial Application Period Cumulative Effect Transition [Line Items]                        
Revenues                   58,644    
Revenue from collaborative agreements | Accounting Standards Update 2014-09 | Pre-Adoption                        
Revenue, Initial Application Period Cumulative Effect Transition [Line Items]                        
Revenues                   58,104    
Revenue from collaborative agreements | Accounting Standards Update 2014-09 | ASC 606 Adjustment                        
Revenue, Initial Application Period Cumulative Effect Transition [Line Items]                        
Revenues                   $ 540