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Collaboration Agreements - Takeda (Details) | 3 Months Ended | 9 Months Ended | 45 Months Ended |  |
--- | --- | --- | --- | --- | ---
Sep. 30, 2021 USD ($) | Sep. 30, 2020 USD ($) | Sep. 30, 2021 USD ($) | Sep. 30, 2020 USD ($) | Sep. 30, 2021 USD ($) | Dec. 31, 2020 USD ($) | Jan. 31, 2018 program
Collaborative Arrangements and Non-collaborative Arrangement Transactions [Line Items] |  |  |  |  |  |  |  |  |
Collaboration revenue from customers | $ 5,285,000 | [1] | $ 9,388,000 | $ 36,143,000 | [1] | $ 18,751,000 |  |  |
Collaborative Arrangement | Takeda Pharmaceutical Company Limited |  |  |  |  |  |  |  |  |
Collaborative Arrangements and Non-collaborative Arrangement Transactions [Line Items] |  |  |  |  |  |  |  |  |
Transaction price, change | 0 |  |  | 0 |  |  |  |  |
Contract liability | 29,200,000 |  |  | 29,200,000 |  |  | $ 29,200,000 | $ 39,800,000 |
Number of performance obligations | program |  |  |  |  |  |  |  |  | 3
Preclinical milestone payment earned not yet received | $ 0 |  |  | $ 0 |  |  | 0 |  |
Collaborative Arrangement | Takeda Pharmaceutical Company Limited | Milestone Triggered |  |  |  |  |  |  |  |  |
Collaborative Arrangements and Non-collaborative Arrangement Transactions [Line Items] |  |  |  |  |  |  |  |  |
Collaboration revenue from customers |  |  |  |  |  |  | 31,000,000 |  |
Collaborative Arrangement | Takeda Pharmaceutical Company Limited | Product |  |  |  |  |  |  |  |  |
Collaborative Arrangements and Non-collaborative Arrangement Transactions [Line Items] |  |  |  |  |  |  |  |  |
Collaboration revenue from customers |  |  |  |  |  |  | $ 0 |  |
Collaborative Arrangement, PTV:PGRN | Takeda Pharmaceutical Company Limited |  |  |  |  |  |  |  |  |
Collaborative Arrangements and Non-collaborative Arrangement Transactions [Line Items] |  |  |  |  |  |  |  |  |
Preclinical milestone payment earned not yet received |  |  |  |  |  |  |  | $ 8,000,000 |
[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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