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Nature Precedings : hdl:10101/npre.2009.3683.1 : Posted 24 Aug 2009
Nature Precedings : hdl:10101/npre.2009.3683.1 : Posted 24 Aug 2009
Nature Precedings : hdl:10101/npre.2009.3683.1 : Posted 24 Aug 2009
Nature Precedings : hdl:10101/npre.2009.3683.1 : Posted 24 Aug 2009
Table 1.
Angry sentences
Reeling from the fight with that stubborn bigot, you slam the car door.
The pushy telemarketer won't let you return to your dinner.
Nature Precedings : hdl:10101/npre.2009.3683.1 : Posted 24 Aug 2009
The workload from your pompous professor is unreasonable.
Happy sentences
The water park is refreshing on the hot summer day.
Finally, you reach the summit of the tall mountain.
You spring up the stairs to your lover’s apartment.
Sad sentences
You hold back your tears as you enter the funeral home.
You open your email in-box on your birthday to find no new emails.
Your closest friend has just been hospitalized for a mental illness.
Table 2.
Nature Precedings : hdl:10101/npre.2009.3683.1 : Posted 24 Aug 2009
Session
Angry sentences Sad sentences Happy sentences
Session 1 .90 (.306) .92 (.271) .91 (.286)
Session 2 .86 (.344) .85 (.354) .92 (.266)
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