Abstract: Eyes play an important role in communication amongst humans and animals. However, relatively little is known about specific differences in eye morphology amongst primates and how these features might be associated with social structure and direction of gaze. We present a detailed study of gazing and eye morphology-exposed sclera and surrounding features in orangutans. We measured gazing in rehabilitating orangutans in two contexts: interspecific viewing of the experimenter (with video camera) and intraspecific gazing (between subjects). Our findings show that direct staring is avoided and social looking is limited to certain age/social categories: juveniles engage in more looking at other orangutans than do adults or infants. While orangutans use eye movements in social communication, they avoid the more prolonged mutual gaze that is characteristic of humans, and also apparent in chimpanzees and gorillas. Detailed frame-by-frame analysis of videotapes from field and zoo studies of orangutans revealed that they pay visual attention to both human observers and conspecifics by glancing sideways, with the head turned at an angle away from the subject being observed. Mutual gaze was extremely rare, and we have observed only two incidences of gaze following. Orangutans in captivity appear to use a more restricted pattern of gazes compared to free-living, rehabilitating ones, possibly suggesting the presence of a pathological condition (such as depression) in the captive subjects. Our findings have implications for further investigations of social communication and cognition in orangutans.

Abstract: Lateralization of the brain appeared early in evolution and many of its features appear to have been retained, possibly even in humans. We now have a considerable amount of information on the different forms of lateralization in a number of species, and the commonalities of these are discussed, but there has been relatively little investigation of the advantages of being lateralized. This article reports new findings on the differences between lateralized and nonlateralized chicks. The lateralized chicks were exposed to light for 24 h on day 19 of incubation, a treatment known to lead to lateralization of a number of visually guided responses, and the nonlateralized chicks were incubated in the dark. When they were feeding, the lateralized chicks were found to detect a stimulus resembling a raptor with shorter latency than nonlateralized chicks. This difference was not a nonspecific effect caused by the light-exposed chicks being more distressed by the stimulus. Instead, it appears to be a genuine advantage conferred by having a lateralized brain. It is suggested that having a lateralized brain allows dual attention to the tasks of feeding (right eye and left hemisphere) and vigilance for predators (left eye and right hemisphere). Nonlateralized chicks appear to perform these dual tasks less efficiently than lateralized ones. Reference is made to other species in discussing these results.

Abstract: The left and right sides of the brain are specialised to process information in different ways and to control different categories of behaviour. Research on a range of species has shown that the left hemisphere controls well-established patterns of behaviour performed in non-stressful situations, whereas the right hemisphere responds to unexpected stimuli and controls escape and other emergency responses. The known functions of each hemisphere are summarised in this paper. Then it is hypothesised that stressed animals rely on predominant use of the right hemisphere, and that a bias to use the right or left hemisphere, respectively, may explain the behavioural differences between animals with a negative cognitive bias and those with a positive cognitive bias. In some species of primates it has been shown that the preferred limb used to pick up food when the animal is in a relaxed state reflects the dominant hemisphere and may be an accessible measure indicating susceptibility to stress and tendency towards positive versus negative cognitive bias. Hence, limb preference might be a useful measure of such tendencies in domesticated species. Some difficulties in determining a relevant measure of limb preference in non-primate species are mentioned, followed by the suggestion that eye preferences for viewing certain stimuli may be a useful measure in species with laterally placed eyes. Finally, effects of experience on the development of hemispheric dominance are discussed, leading to a suggestion that the welfare of domestic animals may be enhanced by ensuring development of left hemisphere dominance (e.g. by exposing chick embryos to light) and by shifting right to left hemisphere dominance in animals with negative cognitive bias.

Abstract: We tested the responses of Bufo marinus to prey stimuli of varying visual complexity that were moved around the toads in either a clockwise or anticlockwise direction at 1.7 revolutions/min. Predatory responses directed at prey resembling an insect were frequent when the model insect moved clockwise across the visual midline into the right visual hemifield. In contrast, the toads tended to ignore such stimuli when they moved anticlockwise across the midline into the left hemifield. No such lateralization was found when a rectangular strip moved along its longest axis was presented in a similar way. The toads also directed more responses towards the latter stimulus than towards the insect prey. Hence, the results suggest that lateralized predatory responses occur for considered decisions on whether or not to respond to complex insect-like stimuli, but not for decisions on comparatively simple stimuli. We discuss similarities between the lateralized feeding responses of B. marinus and those of avian species, as support for the hypothesis that lateralized brain function in tetrapods may have arisen from a common lateralized ancestor.

Abstract: The influence of early experience on the development of lateralisation of hemispheric function was further investigated, using the chick as a model. A range of functions are lateralised in the chick and these correlate with asymmetry in the organisation of the visual projections. Chicks using the right eye and, therefore, primarily the left hemisphere are able to switch from pecking randomly at grain and pebbles to pecking mainly at grain, whereas those using the left eye and primarily the right hemisphere continue to peck at random. Exposure to light during the last days of incubation establishes this lateralisation in males, as a consequence of the embryo being oriented in the egg so that the left eye only is occluded. Males incubated in the dark peck at random when using either the right or left eye. Irrespective of light experience, females perform the same as darkincubated males: they are not influenced by light exposure. Monocular performance of the pebble-grain task is compared to binocular performance, and the sensitive period for the influence of light is delineated. The interactive effects of sex hormone levels on the differentiation of lateralisation are discussed and also the relevance of the results to other species, including humans.