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Income Taxes - Tax Effects (Details) - USD ($) $ in Thousands | 12 Months Ended
--- | ---
Dec. 31, 2016 | Dec. 31, 2015 | Dec. 31, 2014
Income Tax Disclosure [Abstract] |  |  |
Pretax loss | $ 31,400 | $ 33,700 | $ 29,400
Provision for income taxes | 0 | 0 | 0
Reconciliation of tax expense computed at the statutory federal rate and the Company's tax expense |  |  |
U.S. federal taxes at statutory rate | (10,662) | (11,459) | (9,987)
State tax (net of federal benefit) | 20 | (30) | 5
Permanent differences | 153 | 96 | 64
Incentive stock options | 1,095 | 789 | 672
Tax credits | (677) | (581) | (461)
Change in valuation allowance | 10,071 | 11,185 | 9,707
Total provision for income taxes | 0 | 0 | 0
Non-current deferred tax assets: |  |  |
Net operating loss carryforwards | 61,674 | 52,262 | 41,971
Research and development credits | 3,174 | 2,497 | 1,916
Stock-based compensation | 2,847 | 1,825 | 826
Genzyme co-promotion agreement | 0 | 330 | 995
Accruals, deferred rent and other | 4,511 | 4,698 | 3,381
Gross deferred tax assets | 72,206 | 61,612 | 49,089
Valuation allowance | (65,975) | (55,101) | (43,439)
Net deferred tax assets | 6,231 | 6,511 | 5,650
Deferred tax liabilities: |  |  |
Property and equipment | (1,180) | (1,215) | (60)
In-process research and development | (5,051) | (5,296) | (5,590)
Gross deferred tax liabilities | (6,231) | (6,511) | (5,650)
Net deferred tax liabilities | (6,231) | (6,511) | (5,650)
Increase in valuation allowance against deferred tax assets | 10,900 | $ 11,700 | $ 10,600
Operating loss carryforwards |  |  |
Federal and state net operating loss carryforwards related to stock-based compensation | 1,600 |  |
Federal |  |  |
Operating loss carryforwards |  |  |
Net operating loss carryforwards | 169,100 |  |
State |  |  |
Operating loss carryforwards |  |  |
Net operating loss carryforwards | $ 84,200 |  |

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