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Crowell-Davis, S. L. (1985). Nursing behaviour and maternal aggression among Welsh ponies (Equus caballus). Appl Anim Behav Sci, 14(1), 11–25.
Abstract: Nursing behaviour and related aggression of mare-foal pairs was studied from birth (n = 21) to 24 weeks of age (n = 15) of the foal. Foals exhibited a decreasing length and frequency of nursing as they grew older. Mares rarely aggressed against their foals during nursing in the foal's first 4 weeks of life, but did so increasingly through Weeks 13-16, after which the rate of aggression during nursing decreased. Mares terminated nursing primarily by moving away, and were most likely to do so during the foal's first 4 weeks of life. They became gradually less likely to do so as the foal grew older. It was concluded that mares sometimes flex their hind limb on the side opposite the foal during nursing in order to conserve energy in a situation in which they would be remaining still anyway. There was no difference between colts and fillies in the frequency or duration of nursing or in the frequency with which their mothers aggressed against them or terminated nursing.
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Kronfeld, D. S., Custalow, S. E., Ferrante, P. L., Taylor, L. E., Wilson, J. A., & Tiegs, W. (1998). Acid-base responses of fat-adapted horses: relevance to hard work in the heat. Appl. Anim. Behav. Sci., 59(1-3), 61–72.
Abstract: Feeding and training may affect acid-base responses to strenuous exercise. Acidosis usually correlates with higher blood lactate concentrations during intense exercise, but alkalosis has been found in several studies of horses, and higher lactate responses during sprints have been found in fat adapted horses. To elucidate these unexpected findings, we applied a comprehensive physicochemical approach to evaluate acid-base responses during exercise in fat adapted horses. In incremental tests and repeated sprints, changes in blood [H+] were dependent upon corresponding changes in pCO2 but not strong ion difference (SID, the algebraic sum of ions of sodium, potassium, chloride and lactate). The influence of changes in [Lac-] were largely offset by changes in [Na+], [K+] and [Cl-], so that SID was unchanged and did not contribute to the exercise induced acidemia, so it may be inaccurate to term this a lacticacidosis. During repeated sprints, central venous [H+] increased (acidosis) but arterial [H+] decreased (alkalosis). These changes were consistent with concurrent changes in venous and arterial pCO2 but not SID. Fat adaptation decreased mixed venous pCO2 during repeated sprints, which is consistent with the lower respiratory quotient associated with fat oxidation. Less pulmonary work to eliminate CO2 could benefit horses under hot and humid conditions, especially those with mildly reduced pulmonary function. The blood lactate response was decreased during aerobic tests but increased during anaerobic tests on fat adapted horses. Fat adaptation appears to facilitate the metabolic regulation of glycolysis, by sparing glucose and glycogen at work of low intensity, but by promoting glycolysis when power is needed for high intensity exercise. The blood lactate response to repeated sprints was increased more by the combination of fat adaptation and oral supplementation of sodium bicarbonate than by the sum of the responses to fat alone or bicarbonate alone. This synergism suggests that need for further studies of the interaction of fat adaptation with dietary cation-anion balance, especially under hot conditions. These results integrate harmoniously with previous findings of lower feed intake and fecal output, lower loads of heat and CO2, lower water losses in the feces and by evaporation, and less spontaneous activity and reactivity in fat adapted horses. Thus fat adaptation confers several advantages on horses and presumably other equids used for hard work, especially in the heat.
Keywords: Acid-base; Strong ion difference; pCO2; Exercise; Fat adaptation; Corn oil; Ambient heat; Horse
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McLean, A. N. (2001). Cognitive abilities -- the result of selective pressures on food acquisition? Appl. Anim. Behav. Sci., 71(3), 241–258.
Abstract: Locating and capturing food are suggested as significant selection pressures for the evolution of various cognitive abilities in mammals and birds. The hypothesis is proposed that aspects of food procuring behaviour should be strongly indicative of particular cognitive abilities. Experimental data concerning higher mental abilities in mammals and birds are reviewed. These data deal with self-recognition studies, rule-learning experiments, number concept, deceptive abilities, tool-use and observational learning. A Darwinian approach reveals: (1) the adaptiveness of particular abilities for particular niches, (2) that in complex foraging environments, increases in foraging efficiencies in animals should result from the evolution of particular cognitive abilities, (3) that phenomena such as convergent mental evolution should be expected to have taken place across taxonomic groups for species exploiting similar niches, (4) that divergence in mental ability should also have taken place where related species have exploited dissimilar niches. Experimental data of higher mental abilities in animals concur with a Darwinian explanation for the distribution of these cognitive abilities and no anomalies have been found. There are, as a consequence, significant implications for the welfare of animals subject to training when training methodology gives little or no consideration to the various mental abilities of species.
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Araba, B. D., & Crowell-Davis, S. L. (1994). Dominance relationships and aggression of foals (Equus caballus). Appl. Anim. Behav. Sci., 41(1-2), 1–25.
Abstract: Studied a herd of 15 Belgian brood-mares and 10 foals. Specific aspects of social structure studied were dominance-subordinance relationships, preferred associates, social spacing, aggression rates, the frequency of aggressions administered down the dominance hierarchy, and interactive play bouts. The rank order of the foals, both before and after weaning, was positively correlated with the rank order of their dams. There was also a significant relationship between a foal's rank and its total aggression or aggression rate per subordinate post-weaning. Higher ranking foals had higher rates of aggression. Over 80% of threats were directed down the dominance hierachy. The play-rank order of the foals, scored by the number of times foal left a play bout, was not significantly correlated with the rank order as scored by agonistic interactions. -from Authors
Keywords: aggression; dominance; horse; Equus caballus
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Ahrendt, L. P., Labouriau, R., Malmkvist, J., Nicol, C. J., & Christensen, J. W. (2015). Development of a standard test to assess negative reinforcement learning in horses. Appl. Anim. Behav. Sci., 169, 38–42.
Abstract: Most horses are trained by negative reinforcement. Currently, however, no standardised test for evaluating horses' negative reinforcement learning ability is available. The aim of this study was to develop an objective test to investigate negative reinforcement learning in horses. Twenty-four Icelandic horses (3 years old) were included in this study. The horses were tested in a pressure-release task on three separate days with 10, 7 and 5 trials on each side, respectively. Each trial consisted of pressure being applied on the hindquarter with an algometer. The force of the pressure was increased until the horse moved laterally away from the point of pressure. There was a significant decrease in required force over trials on the first test day (P<0.001), but not the second and third day. The intercepts on days 2 and 3 differed significantly from day 1 (P<0.001), but not each other. Significantly stronger force was required on the right side compared to the left (P<0.001), but there was no difference between first and second side tested (P=0.56). Individual performance was evaluated by median-force and the change in force over trials on the first test day. These two measures may explain different characteristics of negative reinforcement learning. In conclusion, this study presents a novel, standardised test for evaluating negative reinforcement learning ability in horses.
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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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Watanabe, S. (2007). How animal psychology contributes to animal welfare. Appl. Anim. Behav. Sci., 106(4), 193–202.
Abstract: This article explores the contribution of animal psychology to animal welfare. Since animal welfare includes subjective welfare, it is crucial to know the subjective world of animals. Analysis of the concept of anthropomorphism is particularly important because it is a basic idea of animal ethics. The history of animal psychology, focusing on anthropomorphism and behaviourism, is briefly described, and then measurement of the subjective experience of animals in two ways, namely animal cognition and pleasure or reinforcing effects, is reported. Finally, it is suggested that animal welfare is not a permanently fixed idea, but a socially constructed one that can be changed. To gain widespread agreement about a socially constructed idea, it is important to know in which circumstances ordinary people employ metaphorical extension to an understanding of animal behaviour. In other words, a survey of “folk animal psychology” is important in order to establish a consensus about animal welfare.
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Burn, C. C., Dennison, T. L., & Whay, H. R. (2010). Relationships between behaviour and health in working horses, donkeys, and mules in developing countries. Appl. Anim. Behav. Sci., 126(3-4), 109–118.
Abstract: Recent studies raise serious welfare concerns regarding the estimated 93.6 million horses, donkeys and mules in developing countries. Most equids are used for work in poor communities, and are commonly afflicted with wounds, poor body condition, respiratory diseases, parasites, dental problems, and lameness. Non-physical welfare problems, such as fear of humans, are also of concern. Interventions to improve working equine welfare aim to prioritise the conditions that cause the most severe impositions on the animals' subjectively experienced welfare, but data identifying which conditions these may be, are lacking. Here we describe a stage in the validation of behavioural welfare indicators that form part of a working equine welfare assessment protocol. Over 4 years, behavioural and physical data were collected from 5481 donkeys, 4504 horses, and 858 mules across nine developing countries. Behaviours included the animals' general alertness, and their responses to four human-interaction tests, using the unfamiliar observer as the human stimulus. Avoidance behaviours correlated significantly with each other across the human-interaction tests, with 21% of animals avoiding the observer, but they showed no associations with likely anthropogenic injuries. Over 13% of equids appeared [`]apathetic': lethargic rather than alert. Measures of unresponsiveness correlated with each other across the five tests, and were associated with poor body condition, abnormal mucous membrane colour, faecal soiling, eye abnormalities, more severe wounds, and older age, depending on the equine species. This suggests that working equids in poor physical health show an unresponsive behavioural profile, consistent with sickness behaviour, exhaustion, chronic pain, or depression-like states.
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Newberry, R. C., & Swanson, J. C. (2008). Implications of breaking mother-young social bonds. Appl. Anim. Behav. Sci., 110(1-2), 3–23.
Abstract: Whereas mammalian mothers and young may retain long-term social affiliations in nature, the management of animals in captivity typically dictates that offspring are abruptly and permanently separated from their mothers at a relatively early age, often prior to the time of natural weaning. For animal breeders, this strategy can enhance the yield of offspring from a breeding population. Morbidity and mortality can also precipitate severance of mother-young bonds. Although it is recognized that early weaning provides nutritional challenges for the young, relatively little attention has been paid to the psychological consequences and long-term impacts of breaking the mother-young bond in non-human mammals. Furthermore, whereas great strides are being made in our understanding of the neurobiological and genetic underpinnings of social bonding, the mechanisms underlying the process of detachment following establishment of a mother-young bond remain relatively unexplored, although parallels can be drawn with processes involved in withdrawal from addictive substances. In this review, we outline mechanisms involved in social bonding. We consider the diversity in extent and duration of mother-young attachment across mammalian lineages and implications for predicting the outcome of severing ties between mothers and young at different times post-partum. We identify characteristics signalling emotional distress resulting from separation of mothers and young and discuss strategies for mitigating separation-induced distress. These include postponement of separation, ensuring high-quality maternal care of young prior to separation, providing bonded individuals with opportunities to separate voluntarily for brief periods prior to permanent separation, use of anti-suck devices prior to separation, allowing a period of partial (fence line) contact prior to full separation, providing substitutes for stimuli previously exchanged between mother and young, providing social buffers, gradual introduction to new housing arrangements, and pharmacological intervention. Areas for future research are proposed, including the use of functional neuroimaging technologies and functional genomics approaches, in combination with behavioural assessments of reinstatement motivation, individual recognition memory and long-term consequences of early separation, to shed further light on the nature of mother-young bonding and detachment in animals.
Keywords: Animal welfare; Emotion; Separation; Social attachment; Weaning
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Strand, S. C., Tiefenbacher, S., Haskell, M., Hosmer, T., McDonnell, S. M., & Freeman, D. A. (2002). Behavior and physiologic responses of mares to short-term isolation. Appl. Anim. Behav. Sci., 78(2-4), 145–157.
Abstract: The aim of this study was to evaluate the behavior and physiologic responses of mares to removal from an established pasture herd and to isolation in a pasture setting for 6 h (Group I, n=5). Responses of mares in Group I were compared to mares that were transported and returned to the herd (Group T, n=5) and to mares moved to the isolation pasture with a companion (Group C, n=5). Behavior was recorded continuously for 6 h on the day before the isolation procedures (baseline, Day 0) and again on the day of the procedure (test, Day 1). Plasma cortisol, white blood cell count (WBC), neutrophil:lymphocyte ratio (N:L), and hematocrit (HCT) were measured once on Day 0 (a.m.) and twice on Day 1 (a.m. and p.m.). Heart rate (HR) was monitored continuously during Day 0 and Day 1. Intradermal response to phytohemagglutinin (PHA) injection was measured 18 h following injection, which was administered at the end of Day 1. Average time spent standing alert increased (P<0.05) in Groups I and C and average time spent grazing decreased (P<0.05) in Group C from Day 0 to Day 1. Also, there was a significant difference between groups (based on a calculated χ2-square value) in the proportion of mares that autogroomed, defecated, urinated, rolled, and whinnied on Day 1. Activity shift rate (ASR) and temperament scores increased significantly in Groups I and C from Day 0 to Day 1 (P<0.05). Plasma cortisol increased significantly in all groups from Day 0 to Day 1, a.m. (P<0.05) and decreased significantly from Day 1, a.m. to Day 1, p.m. (P<0.05). HCT significantly increased in all three groups from Day 0 to Day 1, a.m. (P<0.05). WBC significantly increased in Group T from Day 0 to Day 1, a.m. (P<0.05). N:L ratio significantly increased in Groups I and C from Day 0 and Day 1, a.m. to Day 1, p.m. (P<0.05). A variety of measures did indicate a response to removal from the pasture group, however, the overall, short-term response was minimal. Since the responses of Groups I and C were similar, the effects of isolation versus a novel environment or separation from the established herd could not be differentiated.
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