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Stock-Based Compensation (Tables)
6 Months Ended
Jun. 30, 2020
Share-based Payment Arrangement [Abstract]  
Stock-Based Compensation Expense
The following stock-based compensation expense was recognized for the periods indicated (in thousands):
Three Months Ended June 30,Six Months Ended June 30,
2020201920202019
Research and development$2,589  $2,453  $5,023  $4,267  
General and administrative2,506  2,431  4,564  4,416  
Total stock-based compensation expense$5,095  $4,884  $9,587  $8,683  
Valuation Assumptions Using the Black-Scholes Option-Pricing Model
The fair value of each option award is estimated on the date of grant using the Black-Scholes option-pricing model using the assumptions in the following table for options issued during the period indicated:
Six Months Ended June 30,
20202019
Expected dividend yield0%0%
Expected volatility
67.3% - 108.8%
73.7% - 75.3%
Risk-free interest rate
0.5% - 1.8%
1.9% - 2.6%
Expected term6.25 years6.25 years
Stock Option and Restricted Stock Unit Activity
The following table summarizes stock option activity during the six months ended June 30, 2020:
SharesWeighted-
Average
Exercise Price
Weighted-Average
Remaining
Contractual Term
(Years)
Aggregate
Intrinsic
Value
(in thousands)
Outstanding, December 31, 20196,706,994  $22.33  6.9
Granted1,547,627  12.67  
Exercised(329,592) 7.31  
Forfeited or expired(650,489) 20.90  
Outstanding, June 30, 20207,274,540  21.07  7.2$54,434  
As of June 30, 2020:
Exercisable4,134,978  23.32  5.923,276  
Vested and expected to vest6,954,436  21.17  7.151,498  
Schedule of RSU Activity
The following table summarizes RSU activity during the six months ended June 30, 2020:

Shares
Weighted-Average
Grant Date Fair Value
Outstanding, December 31, 2019452,500  $15.32  
Granted15,000  23.68  
Exercised—  —  
Forfeited(29,100) 15.32  
Outstanding, June 30, 2020438,400  15.61