Abstract: The ability of horses to learn and remember new tasks is fundamentally important for their use by humans. Fearfulness may, however, interfere with learning, because stimuli in the environment can overshadow signals from the rider or handler. In addition, prolonged high levels of stress hormones can affect neurons within the hippocampus; a brain region central to learning and memory. In a series of experiments, we aimed to investigate the link between performance in two learning tests, the baseline level of stress hormones, measured as faecal cortisol metabolites (FCM), fearfulness, and social rank. Twenty-five geldings (2 or 3 years old) pastured in one group were included in the study. The learning tests were performed by professional trainers and included a number of predefined stages during which the horses were gradually trained to perform exercises, using either negative (NR) or positive reinforcement (PR). Each of the learning tests lasted 3 days; 7min/horse/day. The NR test was repeated in a novel environment. Performance, measured as final stage in the training programme, and heart rate (HR) were recorded. Faeces were collected on four separate days where the horses had been undisturbed at pasture for 48h. Social rank was determined through observations of social interactions during feeding. The fear test was a novel object test during which behaviour and HR were recorded. Performance in the NR and PR learning tests did not correlate. In the NR test, there was a significant, negative correlation between performance and HR in the novel environment (rS=-0.66, P<0.001, i.e. nervous horses had reduced performance), whereas there was no such correlation in the home environment (both NR and PR). Behavioural reactions in the fear test correlated significantly with performance in the NR test in the novel environment (e.g. object alertness and final stage: rS=-0.43, P=0.04), suggesting that performance under unfamiliar, stressful conditions may be predicted by behavioural responses in a fear test. There was a negative correlation between social rank and baseline stress hormones (rS=-0.43, P=0.04), i.e. high rank corresponded to low FCM concentrations, whereas neither rank nor FCM correlated with fearfulness or learning performance. We conclude that performance under stressful conditions is affected by activation of the sympathetic nervous system during training and related to behavioural responses in a standardised fear test. Learning performance in the home environment, however, appears unrelated to fearfulness, social rank and baseline FCM levels.

Abstract: The ability to innovate and the social transmission of innovations have played a central role in human evolution. However, innovation is also crucial for other animals, by allowing them to cope with novel socioecological challenges. Although innovation plays such a central role in animals' lives, we still do not know the conditions required for innovative behaviour to emerge. Here, we focused on interindividual differences in innovation by (1) extensively reviewing existing literature on innovative behaviour in animals and (2) quantitatively testing the different evolutionary hypotheses that have been proposed to explain interindividual variation in innovation propensity during foraging tasks. We ran a series of phylogenetically controlled mixed-effects meta-regression models to determine which hypotheses (if any) are supported by currently available empirical studies. Our analyses show that innovation is more common in individuals that are older and belong to the larger sex, but also in more neophilic and/or explorative individuals. Moreover, these effects change depending on the study setting (i.e. wild versus captive). Our results provide no clear support to the excess of energy or the bad competitor hypotheses and suggest that study setting and interindividual differences in traits related to personality are also important predictors of innovation.