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Saslow, C. A. (2002). Understanding the perceptual world of horses. Appl. Anim. Behav. Sci., 78(2-4), 209–224.
Abstract: From the viewpoint of experimental psychology, there are two problems with our current knowledge of equine perception. The first is that the behavioral and neurophysiological research in this area has enormous gaps, reflecting that this animal is not a convenient laboratory subject. The second is that the horse, having been a close companion to humans for many millennia, entrenched anecdotal wisdom is often hard to separate from scientific fact. Therefore, any summary at present of equine perception has to be provisional. The horse appears to have developed a visual system particularly sensitive to dim light and movement, it may or may not have a weak form of color vision in part of the retina, it has little binocular overlap, and its best acuity is limited to a restricted horizontal band which is aimed primarily by head/neck movements. However, the total field of view is very large. Overall, as would be expected for a prey animal, horse vision appears to have evolved more for detection of predator approach from any angle than for accurate visual identification of stationary objects, especially those seen at a distance. It is likely that, as for most mammals except the primates, horses rely more heavily on their other senses for forming a view of their world. Equine high-frequency hearing extends far above that of humans, but horses may be less able to localize the point of origin of brief sounds. The horse's capacity for chemoreception and its reliance on chemical information for identification may more closely resemble that of the dog than of the human. Its tactile sensitivity is high, and the ability of its brain and body to regulate pain perception appears to be similar to that found in other mammals. There is room for a great deal of future research in both the area of equine perception and sensory-based cognition, but for the present time persons interacting with this animal should be made aware of the importance of the sounds they make, the movements of their bodies, the way they touch the animal, and the odors they emit or carry on their clothing.
Keywords: Horse; Perception; Vision; Olfaction; Touch; Hearing; Pain; Training; Psychophysics; Umwelt
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Judge, P. G., & Mullen, S. H. (2005). Quadratic postconflict affiliation among bystanders in a hamadryas baboon group. Anim. Behav., 69(6), 1345–1355.
Abstract: The tendency in primate groups for two opponents to affiliate shortly after a fight has been described as dyadic reconciliation. The response has been shown to restore disrupted relationships and curtail ongoing aggression. Rates of self-directed behaviour (e.g. scratching) are positively correlated with anxiety in primates and the rates decline after reconciliation, indicating that the response also functions to reduce postconflict tension. Third parties not involved in an aggressive interaction are also likely to affiliate with one of the combatants subsequent to a fight. Such `triadic' interactions may also promote conflict resolution when, for instance, the relatives of a victim affiliate with their relative's aggressor. Because aggression in a group influences a bystander's behaviour with combatants, we hypothesized that aggression between two animals would also influence a bystander's behaviour with other bystanders. Such `quadratic' postconflict interactions might also function to reduce postconflict tension or occur in patterns among kin subgroups to resolve conflict. We tested for quadratic interactions in an 18-member group of captive hamadryas baboons, Papio hamadryas hamadryas. Immediately following a fight, an uninvolved bystander was randomly selected for observation and its affiliative interactions with other bystanders and its displacement activities were recorded for 3 min. Rates of behaviour during these postconflict periods were compared to rates during 3-min baseline periods not preceded by aggression. Bystanders engaged in quadratic interactions by increasing affiliation with other bystanders following aggression. Bystanders directed affiliation to nonkin bystanders that were their preferred social partners. Displacement activities of bystanders were significantly higher during postconflict intervals compared to baseline intervals, and bystander displacement activity levels before affiliative contact with other bystanders were significantly higher than after contact. Apparently, bystanders become tense or anxious after witnessing aggression and affiliate with preferred partners to reduce the arousal.
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McLean, A. N. (2004). Short-term spatial memory in the domestic horse. Appl. Anim. Behav. Sci., 85(1-2), 93–105.
Abstract: This study investigates the ability of horses to recall a feeding event in a two-point choice apparatus. Twelve horses were individually tested whereby they were maintained immobile in a test arena and visually and aurally experienced the delivery of food into one of two feed goals. The horses were then released to make their choice in two experimental contexts: immediate release after experiencing the delivery of food, and release 10 s after food delivery. Each horse performed 40 immediate-release (IR) trials, followed by forty 10-s release trials over a 3-day period. In addition, the same horses were tested 3 months later in the spring with the same number and sequence of trials. Results were analysed by log-linear analysis of frequencies. Results showed that while horses were able to achieve the correct feed goal choice in the immediate-release trials, they were unsuccessful with the 10-s release trials. This suggests that there are limitations in recall abilities in horses, in that they may not possess a prospective type of memory. There are welfare and training implications in these findings concerning the effects of overestimating the mental abilities of horses during training and the effects of delays in reinforcements.
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Nicol, C. J. (2002). Equine learning: progress and suggestions for future research. Appl. Anim. Behav. Sci., 78(2-4), 193–208.
Abstract: Horses are well able to form classical and instrumental associations and so the focus of much recent research has been on the stimulus control of instrumental learning. Horses appear to discriminate using spatial cues more easily than other stimulus features, as indicated both by the speed of initial task acquisition and by the extent to which acquired discriminations can be reversed. Phenomena associated with discrimination learning in laboratory animals, including generalisation and peak shift, have been demonstrated in horses. However, the ability of horses to classify stimuli into categories is more controversial. Although there is some evidence that horses may be able to form categories based on similarities in the physical appearance of different stimuli, there is currently no evidence that they are able to develop abstract concepts. Their performance on social learning tasks has also been poor. Few correlations are observed between the learning ability of individual horses on different tasks, suggesting that it may not be possible to classify individual horses as `good' or `poor' learners. Better learning performance by horses that are naturally calm is probably due to reduced interference in the learning process. Correct handling procedures can lower reactivity levels in horses, and may facilitate learning in some circumstances. Future research on equine learning needs to take into account the complex nature of equine social interaction. Studies on the effects of stress on learning, and on social and spatial cognition, are also particularly needed.
Keywords: Learning; Horse; Equine; Discrimination; Training
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Miklósi, Á., Topál, J., & Csányi, V. (2004). Comparative social cognition: what can dogs teach us? Anim. Behav., 67(6), 995–1004.
Abstract: Research in comparative social cognition addresses how challenges of social living have formed the cognitive structures that control behaviours involved in communication, social learning and social understanding. In contrast to the traditional psychological approach, recent investigations take both evolutionary and functional questions into account, but the main emphasis is still on the mechanisms of behaviour. Although in traditional research `comparative' meant mainly comparisons between humans and other primates, ethological influences have led to a broadening of the spectrum of species under study. In this review, we evaluated how the study of dogs broadens our understanding of comparative social cognition. In the early days of ethology, dogs enjoyed considerable interest from ethologists, but during the last 20 years, dogs have rarely been studied by ethological methods. Through a complex evolutionary process, dogs became adapted for living in human society; therefore, the human environment and social setting now represents a natural ecological niche for this species. We have evidence that dogs have been selected for adaptations to human social life, and that these adaptations have led to marked changes in their communicative, social, cooperative and attachment behaviours towards humans. Until now, the study of dogs was hindered by the view that they represent an `artificial' species, but by accepting that dogs are adapted to their niche, as are other `natural' species, comparative investigations can be put into new light.
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Linklater, W. L., & Cameron, E. Z. (2000). Tests for cooperative behaviour between stallions. Anim. Behav., 60(6), 731–743.
Abstract: Breeding groups with multiple stallions occur sympatrically with single-stallion breeding groups in feral horse, Equus caballus, populations. Mutualism and reciprocal altruism between stallions have been proposed to explain the origin and functioning of multistallion bands. However, empirical support for these hypotheses is contradictory and incomplete. Furthermore, there are no explicit tests of the predictions that each hypothesis makes about stallion behaviour and social structure. We compared nine multistallion and 18 single-stallion bands in the Kaimanawa Ranges, New Zealand. Compared with agonistic behaviours, affiliative behaviours were relatively unimportant in the relationships between stallions within bands. The number of stallions in the band did not have a positive influence on mare group size, stability, home range quality or reproductive success in bands. Furthermore, there was a positive relationship between aggression ('intolerance') by the dominant towards subordinate stallions and the subordinates' effort in mare group defence ('helping') but a negative relationship between helping effort by subordinates and their proximity to, and mating with, the bands' mares. Therefore, the predictions of the mutualism and reciprocal altruism hypotheses were not supported. Indeed, for some of the predictions we found the opposite outcomes to be true. Multistallion bands had significantly poorer reproductive success, and dominant stallions were less tolerant of subordinates that helped most and reduced their access to mares. Nevertheless, in all other respects Kaimanawa stallions in multistallion bands behaved like those described elsewhere. Thus, we reject cooperative hypotheses for multimale breeding groups in horses and discuss the mate parasitism and consort hypotheses as better alternatives. Copyright 2000 The Association for the Study of Animal Behaviour.
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Cameron EZ, L. W. L. (2000). Individual mares bias investment in sons and daughters in relation to their condition. Anim. Behav., 60(3), 359–367.
Abstract: The Trivers-Willard hypothesis (TWH) predicts that a mother will treat a son or daughter differently depending on her ability to invest and the impact of her investment on offspring reproductive success. Although many studies have investigated the hypothesis, few have definitively supported or refuted it because of confounding factors or an inappropriate level of analysis. We studied maternal investment in sons and daughters in feral horses, Equus caballus, which meet the assumptions of the TWH with a minimum of confounding variables. Population level analyses revealed no differences in maternal behaviour towards sons and daughters. When we incorporated mare condition, we found that sons were more costly to mares in good condition, whereas daughters were more costly to mares in poor condition, although no differences in maternal behaviour were found. However, since the TWH makes predictions about individual mothers, we examined investment by mares who reared both a son and a daughter in different years of the study. Mares in good condition invested more in their sons in terms of maternal care patterns, costs to maternal body condition and costs to future reproduction. Conversely, mares in poor condition invested more in daughters. Therefore, with an appropriate level of analysis in a species in which confounding variables are minimal, the predictions of the Trivers-Willard hypothesis are supported. Copyright 2000 The Association for the Study of Animal Behaviour.
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Linklater, W. L., Cameron, E. Z., Minot, E. O., & Stafford, K. J. (1999). Stallion harassment and the mating system of horses. Anim. Behav., 58(2), 295–306.
Abstract: Feral horse, Equus caballus, breeding groups, called bands, usually include one but sometimes up to five stallions. We found that mares were loyal to single-stallion (SS) or multistallion (MS) bands or were social dispersers (maverick mares, Mv). The spacing and social behaviour of mares and stallions in single- and multistallion bands was measured. Indices of mare well-being were also measured including activity budgets (feeding: MS>SS=Mv; resting: MS<SS=Mv), band and mare travel (MS>SS), maternal effort in maintaining contact with foals (MS=Mv>SS), parasite levels in faeces (MS>Mv>SS), body condition (MS=Mv<SS), fecundity (Mv<MS<SS) and offspring mortality (Mv<MS<SS). We present evidence suggesting that the poorer well-being of maverick mares and multistallion band mares results from greater harassment by stallions. Stallion and mare behaviour and poor reproductive success in multistallion bands were not consistent with explanations for the existence of such bands based on cooperation or alternative mating strategies. We suggest an alternative explanation. Stable relationships between mares and a single stallion may enhance reproductive success by reducing aggression between individuals. Therefore, we propose that there is strong selection pressure for stable, long-term stallion-mare relationships, called consort relationships. We propose the consort hypothesis, that multistallion bands are an artefact of selection for stable relationships that occasionally result in more than one such relationship forming, because mares solicit more than one stallion and stallion dominance changes during band formation. Copyright 1999 The Association for the Study of Animal Behaviour.
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Cameron, E. Z., Stafford, K. J., Linklater, W. L., & Veltman, C. J. (1999). Suckling behaviour does not measure milk intake in horses, Equus caballus. Anim. Behav., 57(3), 673–678.
Abstract: Studies of parental investment in mammals have frequently used suckling behaviour to estimate energy transfer from mother to offspring, and consequently to measure maternal input. Such studies assume that the more an offspring sucks, the more milk it will receive. This assumption has been questioned, and a review of the literature found little support for it. To test if suckling behaviour provided an accurate index of milk or energy intake we used a radioactive isotope technique to label the milk of thoroughbred mares and to measure milk transfer to foals. We found no significant linear relationship between usual measures of suckling behaviour and milk or energy intake. No behaviours associated with suckling nor with characteristics of mares and foals improved the relationship; only the number of butts associated with each suck episode even approached significance. If we had used suckling behaviour to test theories on differential maternal investment our conclusions would have been in error. For example, female foals tended to suck for longer than males did but there was no difference in the amount of milk transferred. Consequently, we show that measures of suckling behaviour do not adequately predict milk intake in the domestic horse and we suggest that conclusions about differential maternal investment in mammals based on suckling behaviour are likely to be in error. Copyright 1999 The Association for the Study of Animal Behaviour.
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Cameron, E. Z. (1998). Is suckling behaviour a useful predictor of milk intake? A review. Anim. Behav., 56(3), 521–532.
Abstract: In studies on mammalian parental investment, time spent suckling is often used as a predictor of the milk transferred from mother to infant. It is assumed that the rate of milk transfer is positively correlated with the time spent suckling. However, this assumption has not been tested and empirical studies show conflicting results. Nevertheless, in species in which suckling can readily be observed, time spent suckling is still used to measure milk transfer, although an increasing number of workers recognize that the measure is potentially inaccurate. A meta-analysis on studies that have correlated measures of time spent suckling with milk intake estimates based on weight gain revealed a weak positive relationship and significant heterogeneity between studies. Isotope-labelling techniques for the measurement of milk transfer independent of behaviour have been in use since the 1970s, particularly in studies of species in which suckling is difficult to observe. Only one study has attempted to correlate behavioural measures with independent isotope measures, and it found no relationship between the two measures. I suggest that researchers have avoided such a test as it is unlikely that a strong relationship will be found between milk transfer and suckling behaviour, and I discuss the various factors that confound the relationship and contribute to high heterogeneity between studies. Consequently, the assumption that milk transfer can be measured by time spent suckling has inadequate empirical foundation, and needs to be tested using isotope-labelling methods. Copyright 1998 The Association for the Study of Animal Behaviour
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