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Emotional modulation of the postauricular reflex

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Emotion and <strong>the</strong> post-auricular <strong>reflex</strong> 431<br />

2000) and with motor deficits arising from Parkinson’s disease in<br />

humans (Heim et al., 2002).<br />

An alternative possibility is that increased post-auricular<br />

reactivity reflects enhanced orienting to <strong>the</strong> noise probe stimulus.<br />

Indeed, <strong>the</strong>re is evidence that unexpected noises can evoke<br />

observable movement <strong>of</strong> <strong>the</strong> post-auricular (pinna) muscle in<br />

primates in whom this muscle serves more than a vestigial<br />

function. Naturalistic observations <strong>of</strong> Hanuman langurs, a lower<br />

primate species, have revealed that in adult males, loud and<br />

surprising sounds yield brief (i.e., 0.2–0.3 s) movements <strong>of</strong> both<br />

<strong>the</strong> pinna and <strong>the</strong> head in <strong>the</strong> direction <strong>of</strong> <strong>the</strong> noise (Trivedi &<br />

Mohnot, 2002), suggesting that <strong>the</strong> pinna <strong>reflex</strong> serves an<br />

orienting function in this primate species. It may be that states <strong>of</strong><br />

attentional engagement, such as those associated with viewing <strong>of</strong><br />

pleasurable pictures, facilitate this phasic <strong>reflex</strong>ive orienting<br />

reaction. However, because pleasant and unpleasant pictures<br />

engage more attention than neutral (Lang et al., 1990, 1997), <strong>the</strong><br />

predicted attentional pattern would have been one <strong>of</strong> <strong>postauricular</strong><br />

<strong>reflex</strong> inhibition for both affective categories compared<br />

to neutral. Fur<strong>the</strong>rmore, juveniles and females <strong>of</strong> this species<br />

exhibit greater short-duration pinna movements primarily<br />

during play, feeding, and foraging for food (Trivedi & Mohnot,<br />

2002), suggesting heightened responsiveness <strong>of</strong> this <strong>reflex</strong>ive<br />

system specifically during pleasurable activities.<br />

Never<strong>the</strong>less, <strong>the</strong>re is evidence that unpleasant pictures also<br />

engage attention (Lang et al., 1997), so it is possible that some<br />

types <strong>of</strong> aversive pictures might likewise facilitate <strong>the</strong> <strong>postauricular</strong><br />

<strong>reflex</strong>. The current study focused on threat and physical<br />

attack pictures that tend to be perceived as fearful. Modulatory<br />

Ames, K. A., Merritt, N. P., Stout, K., & Hetrick, W. P. (2003).<br />

Differential effects <strong>of</strong> affective <strong>modulation</strong> on orbicularis and <strong>postauricular</strong><br />

indices <strong>of</strong> startle. Psychophysiology, 40, S22.<br />

Anthony, B. J., & Graham, F. K. (1985). Blink <strong>reflex</strong> modification by<br />

selective attention: Evidence for <strong>the</strong> <strong>modulation</strong> <strong>of</strong> automatic<br />

processing. Biological Psychology, 21, 43–59.<br />

Benning, S. D., Patrick, C. J., Hicks, B., Blonigen, D., & Lang, A. R.<br />

(2001). Affective <strong>modulation</strong> <strong>of</strong> blink startle and post-auricular<br />

<strong>reflex</strong>es to noise probes. Psychophysiology, 38, S24.<br />

Bonnet, M., Bradley, M. M., Lang, P. J., & Requin, J. (1995).<br />

Modulation <strong>of</strong> spinal <strong>reflex</strong>es: Arousal, pleasure, action. Psychophysiology,<br />

32, 367–372.<br />

Bradley, M. M., Codispoti, M., Cuthbert, B. N., & Lang, P. J. (2001).<br />

Emotion and motivation I: Defensive and appetitive reactions in<br />

picture processing. Emotion, 1, 276–298.<br />

Bradley, M. M., Cuthbert, B. N., & Lang, P. J. (1993). Emotion, novelty,<br />

and <strong>the</strong> startle <strong>reflex</strong>: Habituation in humans. Behavioral Neuroscience,<br />

107, 970–980.<br />

Bradley, M. M., Drobes, D., & Lang, P. J. (1996). A probe for all<br />

reasons: Reflex and RTmeasures in perception. Psychophysiology, 33,<br />

S25.<br />

Braff, D. L., Geyer, M. A., & Swerdlow, N. R. (2001). Human studies <strong>of</strong><br />

prepulse inhibition <strong>of</strong> startle: Normal subjects, patient groups, and<br />

pharmacological studies. Psychopharmacology, 156, 234–258.<br />

Cassella, J. V., & Davis, M. (1986). Habituation, prepulse inhibition, fear<br />

conditioning, and drug <strong>modulation</strong> <strong>of</strong> <strong>the</strong> acoustically elicited pinna<br />

<strong>reflex</strong> in rats. Behavioral Neuroscience, 100, 39–44.<br />

Cassella, J. V., Harty, P. T., & Davis, M. (1986). Fear conditioning, prepulse<br />

inhibition and drug <strong>modulation</strong> <strong>of</strong> a short latency startle<br />

response measured electromyographically from neck muscles in <strong>the</strong><br />

rat. Physiology and Behavior, 36, 1187–1191.<br />

Center for <strong>the</strong> Study <strong>of</strong> Emotion and Attention [CSEA-NIMH]. (1999).<br />

The international affective picture system: Digitized photographs.<br />

Gainesville, FL: The Center for Research in Psychophysiology,<br />

University <strong>of</strong> Florida.<br />

REFERENCES<br />

effects may differ for o<strong>the</strong>r aversive contents, such as sadness,<br />

disgust, or mutilation pictures. It will also be important in future<br />

research to examine effects for o<strong>the</strong>r pleasurable picture contents<br />

(e.g., babies, animals, food). In addition, <strong>the</strong>re are o<strong>the</strong>r<br />

limitations <strong>of</strong> <strong>the</strong> current study that need to be addressed before<br />

firm conclusions can be advanced regarding <strong>the</strong> post-auricular<br />

<strong>reflex</strong> as an index <strong>of</strong> emotion. The current sample included only<br />

men; thus, it will be important to replicate <strong>the</strong>se findings in<br />

women. Fur<strong>the</strong>rmore, it will be important to assess <strong>modulation</strong><br />

<strong>of</strong> <strong>the</strong> post-auricular <strong>reflex</strong> in emotional processing tasks o<strong>the</strong>r<br />

than picture viewing. A key question is whe<strong>the</strong>r potentiation <strong>of</strong><br />

<strong>the</strong> post-auricular <strong>reflex</strong> would be observed in appetitive<br />

conditioning or reward anticipation contexts, as is true for<br />

startle in corresponding aversive paradigms (e.g., Hamm,<br />

Greenwald, Bradley, & Lang, 1993; Patrick & Berthot, 1995).<br />

Affirmative findings would lend support to <strong>the</strong> idea that <strong>the</strong> <strong>postauricular</strong><br />

<strong>reflex</strong> is primed during appetitive states. Finally, in light<br />

<strong>of</strong> recent data indicating that hedonic valence and approach–<br />

withdrawal aspects <strong>of</strong> emotion are dissociable, it would be<br />

informative to assess whe<strong>the</strong>r <strong>the</strong> post-auricular <strong>reflex</strong> is<br />

enhanced or inhibited during a negatively valent approach state<br />

(e.g., anger; Harmon-Jones & Sigelman, 2001).<br />

Notwithstanding <strong>the</strong>se limitations, <strong>the</strong> findings <strong>of</strong> our study<br />

are intriguing. They suggest that <strong>the</strong> post-auricular <strong>reflex</strong>, which<br />

is evoked by <strong>the</strong> same acoustic probe as <strong>the</strong> startle <strong>reflex</strong>, may<br />

provide an index <strong>of</strong> appetitive system activation. If so, this<br />

<strong>reflex</strong>ive measure could provide a valuable tool for studying basic<br />

appetitive processes and investigating deficiencies in positive<br />

affect associated with psychopathology.<br />

Cook, E. W. III, Atkinson, L. S., & Lang, K. G. (1987). Stimulus control<br />

and data acquisition for IBM PCs and compatibles. Psychophysiology,<br />

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Cuthbert, B. N., Bradley, M. M., & Lang, P. J. (1996). Probing picture<br />

perception: Activation and emotion. Psychophysiology, 33, 103–111.<br />

Davis, M., Gendelman, D. S., Tischler, M. D., & Gendelman, P. M.<br />

(1982). A primary acoustic startle circuit: Lesion and stimulation<br />

studies. Journal <strong>of</strong> Neuroscience, 2, 791–805.<br />

Davis, M., Walker, D. L., & Lee, Y. (1999). Neurophysiology and<br />

neuropharmacology <strong>of</strong> startle and its affective <strong>modulation</strong>. In M. E.<br />

Dawson, M. E. Schell, & A. H. Bohmelt (Eds.), Startle modification:<br />

Implications for neuroscience, cognitive science, and clinical science (pp.<br />

95–117). New York: Cambridge University Press.<br />

Graham, F. K. (1975). The more or less startling effects <strong>of</strong> weak<br />

prestimulation. Psychophysiology, 2, 238–247.<br />

Gray, H. (1901/1995). Anatomy: Descriptive and surgical (15th edn). New<br />

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Hackley, S. A. (1993). An evaluation <strong>of</strong> <strong>the</strong> automaticity <strong>of</strong> sensory<br />

processing using event-related potentials and brain-stem <strong>reflex</strong>es.<br />

Psychophysiology, 30, 415–428.<br />

Hackley, S. A., Woldorff, M., & Hillyard, S. A. (1987). Combined use <strong>of</strong><br />

micro<strong>reflex</strong>es and event-related brain potentials as measures <strong>of</strong><br />

auditory selective attention. Psychophysiology, 24, 632–647.<br />

Hamm, A. O., Greenwald, M. K., Bradley, M. M., & Lang, P. J. (1993).<br />

<strong>Emotional</strong> learning, hedonic change, and <strong>the</strong> startle probe. Journal <strong>of</strong><br />

Abnormal Psychology, 102, 453–465.<br />

Harmon-Jones, E., & Sigelman, J. (2001). State anger and prefrontal<br />

brain activity: Evidence that insult-related relative left-prefrontal<br />

activation is associated with experienced anger and aggression.<br />

Journal <strong>of</strong> Personality and Social Psychology, 80, 797–803.<br />

Heim, C., Sova, L., Kurz, T., Kolasiewicz, W., Schwegler, H., & Sontag,<br />

K.-H. (2002). Partial loss <strong>of</strong> dopaminergic neurons in <strong>the</strong> substantia<br />

nigra, ventrotegmental area and <strong>the</strong> retrorubral area – Model <strong>of</strong> <strong>the</strong><br />

early beginning <strong>of</strong> Parkinson’s symptomatology? Journal <strong>of</strong> Neural<br />

Transmission, 109, 691–709.

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