Abstract: There is disagreement in the literature about the exact nature of the phenomenon of empathy. There are emotional, cognitive, and conditioning views, applying in varying degrees across species. An adequate description of the ultimate and proximate mechanism can integrate these views. Proximately, the perception of an object's state activates the subject's corresponding representations, which in turn activate somatic and autonomic responses. This mechanism supports basic behaviors (e.g., alarm, social facilitation, vicariousness of emotions, mother-infant responsiveness, and the modeling of competitors and predators) that are crucial for the reproductive success of animals living in groups. The Perception-Action Model (PAM), together with an understanding of how representations change with experience, can explain the major empirical effects in the literature (similarity, familiarity, past experience, explicit teaching, and salience). It can also predict a variety of empathy disorders. The interaction between the PAM and prefrontal functioning can also explain different levels of empathy across species and age groups. This view can advance our evolutionary understanding of empathy beyond inclusive fitness and reciprocal altruism and can explain different levels of empathy across individuals, species, stages of development, and situations.

Abstract: I investigated the behavioural background of the way horses learn to follow humans in the “round pen technique” suggested by “horse whisperers” as a gentle method for initial horse training. Though the practicability of this technique has been adequately demonstrated in the past, the horses' behaviour during such training has not yet been documented in detail. In a riding arena, horses, that did not follow the trainer immediately, were chased away so that they galloped around the trainer. Galloping horses showed specific behaviour such as turning the ear to the trainer, chewing, licking, and stretching head and throat downwards. In subsequent trials horses needed to be chased for less time and finally followed immediately, even when conditions were changed or the trainer was replaced by another person. This suggests that horses learn to follow in this particular situation and also show some generalisation. However, following did not occur on a pasture even after several successful trials in the riding arena.

Abstract: The influence of age, dominance rank, kinship and aggressiveness over affiliative relationships and sexual behaviours were analysed in a herd of Sorraia horses, Equus caballus, kept under extensive management. Subjects were 10 adult mares 5-18 years old that had known each other since birth, and a stallion introduced into the group for breeding for the first time. Kinship coefficient and dominance rank were the most important factors affecting affiliative relationships. Bonds were reciprocal and stronger among mares with higher kinship. Mares spent more time in proximity to close-ranking and lower-ranking females. Mares with stronger affiliative relationships or higher relatedness were not less aggressive towards each other. Affiliative relationships between the stallion and the mares were not reciprocal: lower-ranking mares formed stronger bonds with the stallion but he preferred the less genetically related mares for proximity. However, the stallion was involved in sexual behaviours more frequently with the mares that were more genetically related to him. These results suggest that kinship beyond close relatives may affect affiliative relationships both among familiar and among unfamiliar horses. However, the influence of kinship does not imply that horses possess a kin recognition system and alternative explanations are discussed.

Abstract: We studied experimentally whether horse dams influenced foals' relationships with humans. We investigated the influence of the establishment of positive human-mare relationships on foals' behaviour toward humans. Forty-one foals and their dams were involved in this experiment. Half of the mares were softly brushed and fed by hand during a short period (total of 1.25 h) during the first 5 days of their foals' lives (experimental group, n = 21). The other mares were not handled experimentally and their foals received no contact with the experimenter (control group, n = 20). The reactions of both experimental and control foals were recorded under various conditions, first, for 5 min in the presence of a motionless experimenter, when foals were 15 and 30-35 days old, then in an approach test when they were 15 days old and in a saddle-pad tolerance test when they were 30-35 days old. Finally, approach-stroking tests were performed successively by the familiar experimenter when foals were 11-13 months old and by an unfamiliar person when they were 13-15 months old. Several observations strongly suggest that mares can influence their foals' behaviour toward humans: (1) during the handling procedure, experimental foals of protective mares were further from the handler than foals of calm mares (p < 0.001); (2) experimental foals remained, at all ages, closer to the experimenter (p < 0.05) and initiated more physical contacts (sniffing, licking, etc.) with the experimenter (p < 0.05) than control foals; (3) avoidance and flight responses of experimental foals were considerably reduced during approaches by the experimenter (p < 0.01) and they accepted saddle-pads on their backs more easily (p < 0.01) and more quickly (p < 0.01) than control foals. Lastly, the consequences of handling mares had effects that lasted at least until foals were one year old (p < 0.05) and became generalized from experimenter to unfamiliar humans, who could approach and stroke experimental foals rapidly during a test (p < 0.05). This is the first report of an attempt to use observation of mother by foals to facilitate human-foal relationships. The procedure is simple, takes little time and can easily be applied to any dam-foal pair, as it is not intrusive and presents no risks of disrupting mare-foal bonds.

Abstract: The Sir Philip Sidney game has been used by numerous authors to show how signal cost can facilitate honest signaling among relatives. Here, we demonstrate that, in this game, honest cost-free signals are possible as well, under very general conditions. Moreover, these cost-free signals are better for all participants than the previously explored alternatives. Recent empirical evidence suggests that begging is energetically inexpensive for nestling birds; this finding led some researchers to question the applicability of the costly signaling framework to nestling begging. Our results show that cost-free or inexpensive signals, as observed empirically, fall within the framework of signaling theory.