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Visser, E. K., van Reenen, C. G., Schilder, M. B. H., Barneveld, A., & Blokhuis, H. J. (2003). Learning performances in young horses using two different learning tests. Appl. Anim. Behav. Sci., 80, 311–326.
Abstract: To achieve optimal performance in equine sports as well as in leisure not only the physical abilities of the horse should be considered, but also the horse's personality. Besides temperamental aspects, like emotionality, or the horse's reactivity towards humans in handling situations, the learning ability of the horse is another relevant personality trait. To study whether differences in learning performance are consistent over time and whether individual learning performance differs between learning tests or is affected by emotionality, 39 young horses (Dutch Warmblood) were tested repeatedly in two learning tests. An aversive stimulus (AS) was used in one learning test (the avoidance learning test) and a reward was used in the other learning test (the reward learning test). During both learning tests behaviour as well as heart rate were measured. Each test was executed four times, twice when horses were 1 year of age, and twice when they were 2 years of age. Half of the horses received additional physical training from 6 months onwards. In both tests horses could be classified as either performers, i.e. completing the daily session, or as non-performers, i.e. returning to the home environment without having completed the daily session. There were some indications that emotionality might have caused non-performing behaviour, but these indications are not convincing enough to exclude other causes. Furthermore, there seem to be no simple relationships between measures of heart rate, behavioural responses putatively related to emotionality and learning performance. Horses revealed consistent individual learning performances within years in both tests, and in the avoidance learning test also between years. There was no significant correlation between learning performances in the avoidance learning test and the learning performances in the reward learning test. It is concluded that individual learning abilities are consistent over a short time interval for an avoidance learning test and a reward learning test and over a longer time for the avoidance learning test. Furthermore, results indicate that some horses perform better when they have to learn to avoid an aversive stimulus while others perform better when they are rewarded after a correct response. It is suggested that these differences may be relevant to design optimal individual training programmes and methods.
Keywords: Horse; Personality; Learning performance; Consistency; Emotionality
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Farmer, K., Krueger, K., & Byrne, R. (2010). Visual laterality in the domestic horse (Equus caballus) interacting with humans. Anim. Cogn., 13, 229–238.
Abstract: Most horses have a side on which they are easier to handle and a direction they favour when working on a circle, and recent studies have suggested a correlation between emotion and visual laterality when horses observe inanimate objects. As such lateralisation could provide important clues regarding the horse’s cognitive processes, we investigated whether horses also show laterality in association with people. We gave horses the choice of entering a chute to left or right, with and without the passive, non-interactive presence of a person unknown to them. The left eye was preferred for scanning under both conditions, but significantly more so when a person was present. Traditionally, riders handle horses only from the left, so we repeated the experiment with horses specifically trained on both sides. Again, there was a consistent preference for left eye scanning in the presence of a person, whether known to the horses or not. We also examined horses interacting with a person, using both traditionally and bilaterally trained horses. Both groups showed left eye preference for viewing the person, regardless of training and test procedure. For those horses tested under both passive and interactive conditions, the left eye was preferred significantly more during interaction. We suggest that most horses prefer to use their left eye for assessment and evaluation, and that there is an emotional aspect to the choice which may be positive or negative, depending on the circumstances. We believe these results have important practical implications and that emotional laterality should be taken into account in training methods.
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Siniscalchi, M., Sasso, R., Pepe, A. M., Dimatteo, S., Vallortigara, G., & Quaranta, A. (). Sniffing with the right nostril: lateralization of response to odour stimuli by dogs. Anim. Behav., In Press, Corrected Proof.
Abstract: Lateralization in dogs, Canis familiaris, has been reported for paw usage and response to visual and acoustic stimuli. Surprisingly, however, no investigation of possible lateralization for the most relevant sensory domain of dogs, namely olfaction, has been carried out. Here we investigated left and right nostril use in dogs freely sniffing different emotive stimuli in unrestrained conditions. When sniffing novel nonaversive stimuli (food, lemon, vaginal secretion and cotton swab odours), dogs showed initial preferential use of the right nostril and then a shift towards use of the left nostril with repeated stimulus presentation. When sniffing arousal stimuli such as adrenaline and veterinary sweat odorants, dogs showed a consistent right nostril bias all over the series of stimulus presentations. Results suggest initial involvement of the right hemisphere in processing of novel stimuli followed by the left hemisphere taking charge of control of routine behaviour. Sustained right nostril response to arousal stimuli appears to be consistent with the idea that the sympathetic hypothalamic-pituitary-adrenal axis is mainly under the control of the right hemisphere. The implications of these findings for animal welfare are discussed.
Keywords: animal welfare; Canis familiaris; dog; emotion; laterality olfaction; physiology
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Leliveld, L. M. C., Düpjan, S., Tuchscherer, A., & Puppe, B. (2016). Behavioural and physiological measures indicate subtle variations in the emotional valence of young pigs. Physiol. Behav., 157, 116–124.
Abstract: Abstract In the study of animal emotions, emotional valence has been found to be difficult to measure. Many studies of farm animals' emotions have therefore focussed on the identification of indicators of strong, mainly negative, emotions. However, subtle variations in emotional valence, such as those caused by rather moderate differences in husbandry conditions, may also affect animals' mood and welfare when such variations occur consistently. In this study, we investigated whether repeated moderate aversive or rewarding events could lead to measurable differences in emotional valence in young, weaned pigs. We conditioned 105 female pigs in a test arena to either a repeated startling procedure (sudden noises or appearances of objects) or a repeated rewarding procedure (applesauce, toy and straw) over 11 sessions. Control pigs were also regularly exposed to the same test arena but without conditioning. Before and after conditioning, we measured heart rate and its variability as well as the behavioural reactions of the subjects in the test arena, with a special focus on detailed acoustic analyses of their vocalisations. The behavioural and heart rate measures were analysed as changes compared to the baseline values before conditioning. A limited number of the putative indicators of emotional valence were affected by the conditioning. We found that the negatively conditioned pigs showed changes that were significantly different from those in control pigs, namely a decrease in locomotion and an increase in standing. The positively conditioned pigs, however, showed a stronger increase in heart rate and a smaller decrease in SDNN (a heart rate variability parameter indicating changes in autonomic regulation) compared to the controls. Compared to the negatively conditioned pigs, the positively conditioned pigs produced fewer vocalisations overall as well as fewer low-frequency grunts but more high-frequency grunts. The low-frequency grunts of the negatively conditioned pigs also showed lower frequency parameters (bandwidth, maximum frequency, 25% and 50% quartiles) compared to those of the positively conditioned pigs. In any of the statistically significant results, the conditioning accounted for 1.5–11.9% of variability in the outcome variable. Hence, we conclude that repeated moderate aversive and rewarding events have weak but measurable effects on some aspects of behaviour and physiology in young pigs, possibly indicating changes in emotional valence, which could ultimately affect their welfare. The combination of ethophysiological indicators, i.e., the concurrent examination of heart rate measures, behavioural responses and especially vocalisation patterns, as used in the current study, might be a useful way of examining subtle effects on emotional valence in further studies.
Keywords: Emotion; Heart rate; Vocalisation; Emotional valence; Animal welfare; Domestic pig
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McCoy, D. E., Schiestl, M., Neilands, P., Hassall, R., Gray, R. D., & Taylor, A. H. (2019). New Caledonian Crows Behave Optimistically after Using Tools. Current Biology, .
Abstract: Summary Are complex, species-specific behaviors in animals reinforced by material reward alone or do they also induce positive emotions? Many adaptive human behaviors are intrinsically motivated: they not only improve our material outcomes, but improve our affect as well [1, 2, 3, 4, 5, 6, 7, 8]. Work to date on animal optimism, as an indicator of positive affect, has generally focused on how animals react to change in their circumstances, such as when their environment is enriched [9, 10, 11, 12, 13, 14] or they are manipulated by humans [15, 16, 17, 18, 19, 20, 21, 22, 23], rather than whether complex actions improve emotional state. Here, we show that wild New Caledonian crows are optimistic after tool use, a complex, species-specific behavior. We further demonstrate that this finding cannot be explained by the crows needing to put more effort into gaining food. Our findings therefore raise the possibility that intrinsic motivation (enjoyment) may be a fundamental proximate cause in the evolution of tool use and other complex behaviors. Video Abstract
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Goursot, C., Düpjan, S., Puppe, B., & Leliveld, L. M. C. (2021). Affective styles and emotional lateralization: A promising framework for animal welfare research. Appl. Anim. Behav. Sci., 237, 105279.
Abstract: The growing recognition of animals as individuals has broader implications for farm animal welfare research. Even under highly standardized on-farm conditions, farm animals show heterogeneous but individually consistent behavioural patterns towards various stimuli, based on how they appraise these stimuli. As a result, animal welfare is likely to be highly individual as well, and studying the proximate mechanisms underlying distinct individual behaviour patterns and appraisal will improve animal welfare research. We propose to extend the framework of affective styles to bridge the gap between existing research fields on animal personality and affective states. Affective styles refer to consistent individual differences in emotional reactivity and regulation and can be predicted by baseline cerebral lateralization. Likewise, animals with consistent left or right motor biases--a proxy measure of individual patterns in cerebral lateralization--have been shown to differ in their personality, emotional reactivity, motivational tendencies or coping styles. In this paper, we present the current knowledge of the links between laterality and stable individual traits in behaviour and affect in light of hypotheses on emotional lateralization. Within our suggested framework, we make recommendations on how to investigate affective styles in non-human animals and give practical examples. This approach has the potential to promote a science of affective styles in nonhuman animals and significantly advance research on animal welfare.
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Preston, S. D., & de Waal, F. B. M. (2002). Empathy: Its ultimate and proximate bases. Behav Brain Sci, 25(1), 1–20; discussion 20–71.
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.
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Cattell, R. B., & Korth, B. (1973). The isolation of temperament dimensions in dogs. Behav Biol, 9(1), 15–30.
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Lansade, L., Bertrand, M., & Bouissou, M. - F. (2005). Effects of neonatal handling on subsequent manageability, reactivity and learning ability of foals. Appl. Anim. Behav. Sci., 92(1-2), 143–158.
Abstract: Behaviour is an important factor to be taken into account in the various uses of horses. Today horses are mainly used for sport and leisure activities. They should therefore be easy to manage, calm and not fearful. Early handling is known to improve manageability and learning ability and to reduce fearfulness in various species. It has become fashionable in the horse industry to use an early training procedure, referred to as “imprint training”, which is said to produce durable if not permanent effects. However, no studies concerning the long-term effects of such neonatal handling have been carried out in horses. The present study examines the short- and long-term effects of neonatal handling on manageability, general reactivity and learning ability of foals. Twenty-six Welsh foals were studied: 13 were handled daily for 14 days from birth and 13 were non-handled controls. The handling procedure consisted of fitting a halter, gently patting all parts of each foal's body, picking up feet and leading over 40 m. Two days, 3 months, 6 months and 1 year after the end of the handling period, foals underwent behavioural tests to measure their manageability and various aspects of their reactivity. The results showed that neonatal handling has only short-term effects on manageability: 2 days after the handling period, handled animals were significantly easier to handle than controls for the four parameters measured during this test (time to fit a halter, time to pick up feet, walk ratio that is time during which foal walks under constraint/total time measured during leading and number of defensive reactions). Two parameters (time to fit a halter and walk ratio) were still lower in handled foals than in non-handled foals 3 months later and only one 6 months later (walk ratio). One year later there was no difference between groups. In addition, there was no effect of handling on reactivity at any time of testing or in any of the tests (reaction to isolation from conspecifics, presence of a human, presence of a novel object and to a surprise effect). Finally, neonatal handling did not improve the spatial or discriminative learning abilities measured at 14 months of age. To conclude, the effects of neonatal handling are only temporary.
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Lansade, L., Bertrand, M., Boivin, X., & Bouissou, M. - F. (2004). Effects of handling at weaning on manageability and reactivity of foals. Appl. Anim. Behav. Sci., 87(1-2), 131–149.
Abstract: The horse's temperament, including its manageability and reactivity and/or fearfulness, is of importance as it can result in problems and can render horses unsuitable for inexperienced riders. Early experience, including handling during infancy, may influence the horse's adult behaviour and reduce its fear of humans and other potentially frigthening situations. In the various species studied, handling has generally been undertaken during the neonatal period. The aim of the present study was to test the effects of handling young horses around the time of weaning, a period which has been demonstrated to be effective in increasing ease of handling in cattle and goats. Sixteen Anglo-Arab foals were handled for 12 days either immediately following weaning (early handled: EH) or 21 days later (late handled: LH); eight additional non-handled foals served as controls (C). Handling consisted of haltering, gently petting all parts of the body, picking up feet and leading the foal over 120 m. During handling sessions, EH were easier to handle than LH: time taken to fit them with a halter, to pick up feet, and “walk-ratio” (time walking under constraint/total time walking) were significantly lower for EH. During subsequent tests conducted over 2 days, 4, and 7 months, as well as 10 months and to some extent 18 months after the end of handling period, EH and LH were easier to handle and less reactive than controls, although differences diminished with time. The period following weaning can therefore be qualified as an “optimal period” for handling. Some of the effects persist for at least 18 months.
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