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Stock-Based Compensation (Tables)
6 Months Ended
Jun. 30, 2019
Disclosure of Compensation Related Costs, Share-based Payments [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,
2019201820192018
Research and development$2,453 $1,989 $4,267 $3,689 
General and administrative2,431 2,174 4,416 3,906 
Total stock-based compensation expense$4,884 $4,163 $8,683 $7,595 
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,
20192018
Expected dividend yield0%  0%  
Expected volatility73.7% - 75.3%67.8% - 72.2%
Risk-free interest rate1.9% - 2.6%2.4% - 3.1%
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, 2019:
SharesWeighted-
Average
Exercise Price
Weighted-Average
Remaining
Contractual Term
(Years)
Aggregate
Intrinsic
Value
(in thousands)
Outstanding, December 31, 20185,273,964 $22.23 6.8
Granted1,796,645 21.42 
Exercised(189,428)3.25 
Forfeited or expired(119,392)23.74 
Outstanding, June 30, 20196,761,789 22.52 7.3$8,883 
As of June 30, 2019:
Exercisable3,722,761 21.92 5.98,850 
Vested and expected to vest6,461,922 22.46 7.28,880