Abstract: tEye and limb preferences were scored in the closest undomesticated relative of Equuscaballus using the same methods as used previously to study laterality in feral horses.Observations were made of 33 Przewalski horses (Equus ferus przewalskii) (male N = 20,female N = 13) living under natural social conditions on a large reserve in France. Signifi-cant left-eye/side biases were found in agonistic interactions within harem bands (M ± SEbias to left 58% ± 0.01 for threats, P < 0.001; 68% ± 0.05 for attacks; P < 0.001) and in stallionfights (threats, 52% ± 0.01 left, P < 0.001; attacks, 63% ± 0.02 left, P < 0.001): as many as 80%of the horses were significantly lateralized in attack responses within harem bands. Lat-erality of vigilance was measured as lifting up the head from grazing and turning it to theleft or right side: a directional bias to the left was found (M ± SE 53% ± 0.02 left, P < 0.001).Side bias in reactivity was calculated as the percent of head lifts above the level of thewithers on the left or right side and this was also left side biased (M ± SE 73% ± 0.03 left,P < 0.001). These results indicate right-hemisphere specialization for control of aggressionand responses to novelty. The left bias in attack scores within harem bands was strongerin males than females (P = 0.024) and in immature than adult horses (P = 0.032). Immaturehorses were also more strongly lateralized than adults in vigilance scores (P = 0.022), whichmay suggest that experience reduces these side biases. Our results show that Przewalskihorses exhibit left eye preferences, as do feral horses, and do so even more strongly thanferal horses. Considering feral and Przewalski horses together, we deduce that ancestralhorses had similar lateral biases. Also similar to feral horses, the Przewalski horses showedno significant forelimb preference at the group level or in the majority of horses at theindividual level, confirming the hypothesis that previously reported limb preferences indomestic breeds are entrained or generated by breed-specific selection.

Abstract: We investigated whether horses display greater reactivity to a novel stimulus presented in the left compared to the right monocular visual field, and whether a population bias exists for escape turning when the same stimulus was presented binocularly. Domestic horses (N=30) were tested on three occasions by a person opening an umbrella five metres away and then approaching. The distance each horse moved away before stopping was measured. Distance was greatest for approach on the left side, indicating right hemisphere control of flight behaviour, and thus followed the same pattern found previously in other species. When order of monocular presentation was considered, an asymmetry was detected. Horses tested initially on the left side exhibited greater reactivity for left approach, whereas horses tested on the right side first displayed no side difference in reactivity. Perhaps left hemisphere inhibition of flight response allowed horses to learn that the stimulus posed no threat and this information was transferred to the right hemisphere. No population bias existed for the direction of escape turning, but horses that turned to the right when approached from the front were found to exhibit longer flight distances than those that turned to the left.

Abstract: Stereokinetic illusions have never been investigated in non-human primates, nor in other mammalian species. These illusions consist in the perception of a 3D solid object when certain 2D stimuli are rotated slowly in the plane perpendicular to the line of sight. The ability to perceive the stereokinetic illusion was investigated in the common marmoset (Callithrix jacchus). Four adult marmosets were trained to discriminate between a solid cylinder and a solid cone for food reward. Once learning criterion was reached, the marmosets were tested in sets of eight probe trials in which the two solid objects used at training were replaced by two rotating 2D stimuli. Only one of these stimuli produced, at least to the human observer, the stereokinetic illusion corresponding to the solid object previously reinforced. At test, the general behaviour and the total time spent by the marmosets observing each stimulus were recorded. The subjects stayed longer near the stimulus producing the stereokinetic illusion corresponding to the solid object reinforced at training than they did near the illusion corresponding to the previously non-rewarded stimulus. Hence, the common marmosets behaved as if they could perceive stereokinetic illusions.

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: We investigated lateralisation in horses because it is likely to be important in training and athletic performance. Thoroughbred horses (n = 106) were observed every 60 s for 2 h, when they were at pasture, and the position of the forelimbs in relation to one another was recorded. There was a population bias skewed to standing with the left forelimb advanced over the right (i.e. directional lateralisation). Using the first 50 observations, the distribution of preferences was 43 significantly left, 10 significantly right with 53 being non-significant (i.e. ambidextextrous). The strength of motor bias increased with age, suggesting maturation or an influence of training. The horses were also presented with an olfactory stimulus (stallion faeces) to score the tendency to use one nostril rather than the other. A significant preference to use the right nostril first was shown in horses under 4 years of age (n = 61) but not in older horses. Of the 157 horses tested for nostril bias, 76 had been assessed for motor bias and so were used for analysis of the relationship between laterality in the two modalities. There was no significant relationship between direction of foreleg motor bias and first nostril used, total number of inhalations or laterality index of nostril use. The absence of a correlation between laterality of nostril use and motor bias indicates that lateralisation of the equine brain occurs on at least two levels of neural organisation--sensory and motor--a finding that is consistent with other examples of lateralisation in species that have been examined in more detail.