Equity Incentive Plans - Summary of Stock-based Compensation Expense Recognized (Details) - USD ($) $ in Thousands	12 Months Ended
	Dec. 31, 2021	Dec. 31, 2020	Dec. 31, 2019
Share Based Compensation Arrangement By Share Based Payment Award [Line Items]
Stock-based compensation	$ 10,264	$ 8,060	$ 6,466
2010 Plan, 2014 Plan and 2018 Inducement Plan
Share Based Compensation Arrangement By Share Based Payment Award [Line Items]
Stock-based compensation	10,105	7,609	6,109
2010 Plan, 2014 Plan and 2018 Inducement Plan | Research and Development
Share Based Compensation Arrangement By Share Based Payment Award [Line Items]
Stock-based compensation	5,462	3,742	3,223
2010 Plan, 2014 Plan and 2018 Inducement Plan | General and Administrative
Share Based Compensation Arrangement By Share Based Payment Award [Line Items]
Stock-based compensation	$ 4,643	$ 3,867	$ 2,886