Collaboration, Development and License Agreements - Cystic Fibrosis Foundation Development Agreement (Details) - USD ($)	1 Months Ended		3 Months Ended	12 Months Ended
	Dec. 31, 2016	Nov. 30, 2016	Jun. 30, 2020	Dec. 31, 2021	Dec. 31, 2020	Sep. 30, 2021	Mar. 31, 2019
Collaborative Arrangements and Non-collaborative Arrangement Transactions
Deferred revenue				$ 20,838,000	$ 18,748,000
Revenue				1,535,000	1,000,000
Phase 1 MAD
Collaborative Arrangements and Non-collaborative Arrangement Transactions
Amount of achieved consideration			$ 1,000,000.0
CFF
Collaborative Arrangements and Non-collaborative Arrangement Transactions
Amount of award	$ 2,900,000	$ 7,500,000
Upfront payment received from CFF	$ 200,000
Deferred revenue				476,000
Revenue					$ 1,000,000.0
CFF | Phase 1 SAD
Collaborative Arrangements and Non-collaborative Arrangement Transactions
Amount of probable transaction price							$ 1,700,000
Amount of probable variable consideration							$ 1,000,000.0
CFF | Phase 2a Clinical Trial
Collaborative Arrangements and Non-collaborative Arrangement Transactions
Gross transaction price						$ 1,000,000.0
Amount of variable consideration				3,800,000
Revenue				$ 524,000