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Share-Based Compensation - (Tables)
6 Months Ended
Jun. 30, 2019
Share-based Payment Arrangement [Abstract]  
Schedule of Allocation of Share-based Compensation Costs by Award Type
Share-based compensation expense, net of estimated forfeitures, includes expenses related to stock options and RSUs granted to employees, non-employee directors and consultants, as well as options granted to employees to purchase shares under the ESPP. Stock-based compensation by award type was as follows:
 
Three Months Ended June 30,
 
Six Months Ended June 30,
 
2019
 
2018
 
2019
 
2018
Stock options
$
1,488

 
$
1,836

 
$
3,110

 
$
3,968

Restricted stock units
709

 
819

 
1,530

 
1,765

Employee stock purchase plan
10

 
7

 
37

 
11

Total
$
2,207

 
$
2,662

 
$
4,677

 
$
5,744

Schedule of Allocation of Share-based Compensation Costs by Financial Statement Line
The Company allocated stock-based compensation expense in the Company’s Consolidated Statements of Operations and Comprehensive Loss as follows:
 
Three Months Ended June 30,
 
Six Months Ended June 30,
 
2019
 
2018
 
2019
 
2018
Research and development
$
989

 
$
1,106

 
$
2,159

 
$
2,546

General and administrative
1,218

 
1,556

 
2,518

 
3,198

Total
$
2,207

 
$
2,662

 
$
4,677

 
$
5,744

Summary of the stock option activity including weighted average exercise prices, options outstanding and exercisable
A summary of the stock option activity, weighted average exercise prices, options outstanding, exercisable and expected to vest as of June 30, 2019 is as follows (in thousands except weighted average exercise price):
 
Number of Shares Underlying Options
 
Weighted
Average
Exercise
Price
Outstanding, December 31, 2018
5,903

 
$
13.72

Granted
137

 
$
1.35

Forfeited
(261
)
 
$
26.18

Expired
(89
)
 
$
12.72

Outstanding, June 30, 2019
5,690

 
$
12.89

Vested and exercisable, June 30, 2019
2,850

 
$
21.83

Vested and expected to vest, June 30, 2019
5,463

 
$
13.26

Summary of the Company's outstanding shares of RSU awards
The following table presents a summary of the Company's outstanding RSU awards granted as of June 30, 2019 (in thousands except weighted average grant-date fair value):
 
Restricted
Stock
Units
 
Weighted Average
Grant-Date
Fair Value
Outstanding, December 31, 2018
679

 
$
15.61

Awarded
10

 
$
1.38

Vested
(49
)
 
$
52.83

Forfeited
(5
)
 
$
38.98

Outstanding, June 30, 2019
635

 
$
12.40

Outstanding, Expected to vest
518

 
$
6.58