da Costa, A. P., Leigh, A. E., Man, M. - S., & Kendrick, K. M. (2004). Face pictures reduce behavioural, autonomic, endocrine and neural indices of stress and fear in sheep. Proc. R. Soc. Lond. B., 271(1552), 2077–2084.
Abstract: Faces are highly emotive stimuli and we find smiling or familiar faces both attractive and comforting, even as young babies. Do other species with sophisticated face recognition skills, such as sheep, also respond to the emotional significance of familiar faces? We report that when sheep experience social isolation, the sight of familiar sheep face pictures compared with those of goats or inverted triangles significantly reduces behavioural (activity and protest vocalizations), autonomic (heart rate) and endocrine (cortisol and adrenaline) indices of stress. They also increase mRNA expression of activity–dependent genes (c–fos and zif/268) in brain regions specialized for processing faces (temporal and medial frontal cortices and basolateral amygdala) and for emotional control (orbitofrontal and cingulate cortex), and reduce their expression in regions associated with stress responses (hypothalamic paraventricular nucleus) and fear (central and lateral amygdala). Effects on face recognition, emotional control and fear centres are restricted to the right brain hemisphere. Results provide evidence that face pictures may be useful for relieving stress caused by unavoidable social isolation in sheep, and possibly other animal species, including humans. The finding that sheep, like humans, appear to have a right brain hemisphere involvement in the control of negative emotional experiences also suggests that functional lateralization of brain emotion systems may be a general feature in mammals.
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Peirce, J. W., Leigh, A. E., & Kendrick, K. M. (2000). Configurational coding, familiarity and the right hemisphere advantage for face recognition in sheep. Neuropsychologia, 38(4), 475–483.
Abstract: This study examined characteristics of visual recognition of familiar and unfamiliar faces in sheep using a 2-way discrimination task. Of particular interest were effects of lateralisation and the differential use of internal (configurational) vs external features of the stimuli. Animals were trained in a Y-maze to identify target faces from pairs, both of which were familiar (same flock as the subjects) or both of which were unfamiliar (different flock). Having been trained to identify the rewarded face a series of stimuli were presented to the sheep, designed to test for the use of each visual hemifield in the discriminations and the use of internal and external facial cues. The first experiment showed that there was a left visual hemifield (LVF) advantage in the identification of [`]hemifaces', and [`]mirrored hemifaces' and [`]chimeric' faces and that this effect was strongest with familiar faces. This represents the first evidence for visual field bias outside the primate literature. Results from the second experiment showed that, whilst both familiar and unfamiliar faces could be identified by the external features alone, only the familiar faces could be recognised by the internal features alone. Overall the results suggest separate recognition methods for socially familiar and unfamiliar faces, with the former being coded more by internal, configurational cues and showing a lateral bias to the left visual field.
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