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Collaboration Agreements - Biogen (Details)	3 Months Ended			9 Months Ended
	Sep. 30, 2021 USD ($) milestone		Sep. 30, 2020 USD ($)	Sep. 30, 2021 USD ($) milestone		Sep. 30, 2020 USD ($)	Dec. 31, 2020 USD ($)
Collaborative Arrangements and Non-collaborative Arrangement Transactions [Line Items]
Offset to research and development expense, related party cost reimbursement	$ 1,200,000			$ 5,300,000
Cost sharing reimbursements due from related party	1,194,000			1,194,000			$ 5,674,000
Collaboration revenue from customers	5,285,000	[1]	$ 9,388,000	36,143,000	[1]	$ 18,751,000
Biogen Collaboration Agreement | Related Party
Collaborative Arrangements and Non-collaborative Arrangement Transactions [Line Items]
Contract liability	294,900,000			294,900,000
Biogen Collaborative Arrangement | Biogen Collaboration Agreement
Collaborative Arrangements and Non-collaborative Arrangement Transactions [Line Items]
Transaction price, change	0			0
Offset to research and development expense, related party cost reimbursement	1,200,000			5,300,000
Cost sharing reimbursements due from related party	$ 1,200,000			$ 1,200,000			$ 5,700,000
Number of milestones achieved | milestone	0			0
Biogen Collaborative Arrangement | Biogen Collaboration Agreement | Option Research Services | Related Party
Collaborative Arrangements and Non-collaborative Arrangement Transactions [Line Items]
Contract liability	$ 6,000,000.0			$ 6,000,000.0
Biogen Collaborative Arrangement | Biogen Collaboration Agreement | Product
Collaborative Arrangements and Non-collaborative Arrangement Transactions [Line Items]
Collaboration revenue from customers				0
ROFN and Option Agreement | Biogen Collaboration Agreement | Related Party
Collaborative Arrangements and Non-collaborative Arrangement Transactions [Line Items]
Contract liability	$ 288,900,000			$ 288,900,000
[1] Includes related party collaboration revenue from a customer of $0.9 million and $2.5 million for the three and nine months ended September 30, 2021, respectively.

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