Abstract: There is evidence that more than 47% of the sports horse population in normal work may be lame, but the lameness is not recognized by owners or trainers. An alternative means of detecting pain may be recognition of behavioral changes in ridden horses. It has been demonstrated that there are differences in facial expressions in nonlame and lame horses. The purpose of this study was to develop a whole horse ethogram for ridden horses and to determine whether it could be applied repeatedly by 1 observer (repeatability study, 9 horses) and if, by application of a related pain behavior score, lame horses (n = 24) and nonlame horses (n = 13) could be differentiated. It was hypothesized that there would be some overlap in pain behavior scores among nonlame and lame horses; and that overall, nonlame horses would have a lower pain behavior score than lame horses. The ethogram was developed with 117 behavioral markers, and the horses were graded twice in random order by a trained specialist using video footage. Overall, there was a good correlation between the 2 assessments (P < 0.001; R2 = 0.91). Behavioral markers that were not consistent across the 2 assessments were omitted, reducing the ethogram to 70 markers. The modified ethogram was applied to video recordings of the nonlame horses and lame horses (ethogram evaluation). There was a strong correlation between 20 behavioral markers and the presence of lameness. The ethogram was subsequently simplified to 24 behavioral markers, by the amalgamation of similar behaviors which scored similarly and by omission of markers which showed unreliable results in relation to lameness. Following this, the maximum individual occurrence score for lame horses was 14 (out of 24 possible markers), with a median and mean score of 9 (±2 standard deviation) compared with a maximum score of 6 for nonlame horses, with a median and mean score of 2 (±1.4). For lame horses, the following behaviors occurred significantly more (P < 0.05, chi-square): ears back, mouth opening, tongue out, change in eye posture and expression, going above the bit, head tossing, tilting the head, unwillingness to go, crookedness, hurrying, changing gait spontaneously, poor quality canter, resisting, and stumbling and toe dragging. Recognition of these features as potential indicators of musculoskeletal pain may enable earlier recognition of lameness and avoidance of punishment-based training. Further research is necessary to verify this new ethogram for assessment of pain in ridden horses.

Abstract: Summary For humans, facial expressions are important social signals, and how we perceive specific individuals may be influenced by subtle emotional cues that they have given us in past encounters. A wide range of animal species are also capable of discriminating the emotions of others through facial expressions [1, 2, 3, 4, 5], and it is clear that remembering emotional experiences with specific individuals could have clear benefits for social bonding and aggression avoidance when these individuals are encountered again. Although there is evidence that non-human animals are capable of remembering the identity of individuals who have directly harmed them [6, 7], it is not known whether animals can form lasting memories of specific individuals simply by observing subtle emotional expressions that they exhibit on their faces. Here we conducted controlled experiments in which domestic horses were presented with a photograph of an angry or happy human face and several hours later saw the person who had given the expression in a neutral state. Short-term exposure to the facial expression was enough to generate clear differences in subsequent responses to that individual (but not to a different mismatched person), consistent with the past angry expression having been perceived negatively and the happy expression positively. Both humans were blind to the photograph that the horses had seen. Our results provide clear evidence that some non-human animals can effectively eavesdrop on the emotional state cues that humans reveal on a moment-to-moment basis, using their memory of these to guide future interactions with particular individuals.

Abstract: Intelligence in large-brained vertebrates might have evolved through independent, yet similar processes based on comparable socioecological pressures and slow life histories. This convergent evolutionary route, however, cannot explain why cephalopods developed large brains and flexible behavioural repertoires: cephalopods have fast life histories and live in simple social environments. Here, we suggest that the loss of the external shell in cephalopods (i) caused a dramatic increase in predatory pressure, which in turn prevented the emergence of slow life histories, and (ii) allowed the exploitation of novel challenging niches, thus favouring the emergence of intelligence. By highlighting convergent and divergent aspects between cephalopods and large-brained vertebrates we illustrate how the evolution of intelligence might not be constrained to a single evolutionary route.

Abstract: Learning through the observation of others allows the transfer of information without the costs incurred during individual trial and error learning. Horses (Equus caballus) are a highly social species, which might be expected to be capable of learning from others, but experimental findings are inconsistent, and potentially confounded by social facilitation effects not related directly to the learning of the task. We refined the methods used in previous equine social learning studies, to examine and distinguish specific social influences on learning of a task: we used predefined group leaders rather than agonistically dominant individuals to demonstrate a detour task to familiar conspecific observers; in addition we had two control groups: a non-observer (true control) and a group with the demonstrator simply present at the goal (social facilitation control). 44 socially kept horses were allocated to one of the three test conditions and took part in five trials each. Success rate, latency and detour direction were recorded. There was no significant difference between the three groups in the likelihood of them succeeding in the task nor latency to succeed; however there was a significant difference in the route chosen by the groups, with the true control choosing the side with the entrance gate significantly more than either the observer group or social facilitation group. Both of the latter two groups chose to go in the same direction relative to themselves, regardless of which side the gate was. Seven out of nine horses in the observer group chose the same direction as their demonstrator every time. Our results show a significant role of social facilitation on detour behaviour and highlight the importance of including adequate controls for simpler cognitive influences on behaviour before claims can be made about the specific learning of motor actions or goal directed behaviour. Social cues may be important to horses if the task is sufficiently challenging and motivationally important, so future work should consider more demanding, but ecologically relevant situations, in order to maximise the potential revelation of social learning effects which do not depend on simple local or stimulus enhancement effects.

Abstract: Equine welfare can be enhanced by applying species specific training. This may incorporate social learning, as horses are highly social and social stimuli are of primary importance. Social learning is comparable to individual learning in its learning mechanisms, differing primarily in the way it is stimulated. Our initial study showed that horses of different breeds (N = 38) follow humans after observing other horses doing so, but only if the observed horse was familiar to and higher ranking than the observer (Fisher's exact test: N = 12, P = 0.003). A second study showed that horses and ponies (N = 25) learned to pull a rope to open a feeding apparatus after observing demonstrations by conspecifics, again, only if the demonstrating horse was older and higher ranking than the observer (Fisher's combination test, N = 3, v2 = 27.71, p = 0.006). Our third approach showed that horses and ponies (N = 24) learned to press a switch to open a feeding apparatus after observing a familiar person (GzLM: N = 24, z = 2.33, P = 0.02). Most recently, we confronted horses and ponies (N = 50) with persons demonstrating different techniques for opening a feeding apparatus. In this study we investigated whether the horses would copy the demonstrators' techniques or apply their own. Here only some horses copied the technique, and most of the successful learners used their mouths irrespective of the demonstrators' postures (Chi Square Test: N = 40, df = 2, &#967;2 = 31.4, p < 0.001). In all the approaches social stimuli elicited learning processes in the test horses, while only a few individuals in the control groups mastered the tasks by individual learning. The following behaviour observed in the initial study may have been facilitated by a social stimuli (social facilitation), and the opening of the feed boxes in the subsequent studies appear to be mostly the result of enhancement (social enhancement). Some horses may have used the social stimuli at first and continued their learning process by individual trial and error. However, the horses were also selective in whom and some in how to copy. This may have been conditioned (socially conditioned) or the result of simple forms of reasoning on the reliability of the particular information provided by demonstrators of certain social ranks or social positions, as high ranking and familiar horses and familiar persons were copied and some imitated exactly.
Lay person message: Traditional riding instructions suggest that horses learn by observing other horses. For example, older, more experienced driving horses are used for initial training of young driving horses. We have shown that horses indeed use learning stimuli provided by other horse, as well as by humans. Horses readily accept stimuli observed in high ranking and familiar horses, and familiar persons. Such stimuli elicit learning processes which are comparable to individual learning. We suggest applying social learning whenever possible, as it is much faster and less stressful than individual learning, where learners experience negative outcomes in trial and error learning.