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Rands, S. A., Cowlishaw, G., Pettifor, R. A., Rowcliffe, J. M., & Johnstone, R. A. (2003). Spontaneous emergence of leaders and followers in foraging pairs. Nature, 423(6938), 432–434.
Abstract: Animals that forage socially often stand to gain from coordination of their behaviour. Yet it is not known how group members reach a consensus on the timing of foraging bouts. Here we demonstrate a simple process by which this may occur. We develop a state-dependent, dynamic game model of foraging by a pair of animals, in which each individual chooses between resting or foraging during a series of consecutive periods, so as to maximize its own individual chances of survival. We find that, if there is an advantage to foraging together, the equilibrium behaviour of both individuals becomes highly synchronized. As a result of this synchronization, differences in the energetic reserves of the two players spontaneously develop, leading them to adopt different behavioural roles. The individual with lower reserves emerges as the 'pace-maker' who determines when the pair should forage, providing a straightforward resolution to the problem of group coordination. Moreover, the strategy that gives rise to this behaviour can be implemented by a simple 'rule of thumb' that requires no detailed knowledge of the state of other individuals.
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Brinkmann, L., Gerken, M., & Riek, A. (2015). Energetic adaptations of Shetland pony mares. In Proceedings of the 3. International Equine Science Meeting.
Abstract: Recent results suggest that wild Northern herbivores exhibit signs of a hypometabolism during times of low ambient temperature and food shortage in order to reduce their energetic needs. However, there are speculations that domestic animals lost the ability to reduce energy expenditure. To examine energetic and behavioural responses 10 Shetland pony mares were exposed to different environmental conditions (summer and winter). During winter ponies were allocated into two groups receiving two different food quantities (60% and 100% of maintenance energy requirement). We measured the field metabolic rate, water turn over, body temperature, locomotor activity, lying time, resting heart rate, body mass and body condition score.
In summer, the field metabolic rate of all ponies (FMR; 63.4±15.0 MJ/day) was considerably higher compared with food restricted and control animals in winter (24.6±7.8 and 15.0±1.1 MJ/day, respectively). Furthermore, during summer, locomotor activity, resting heart rate and total water turnover were significantly elevated (P<0.001) compared with winter. Animals receiving a reduced amount of food (N=5) reduced their FMR by 26% compared with control animals (N=5) to compensate for the decreased energy supply. Furthermore, resting heart rate, body mass and body condition score were lower(29.2±2.7 beats/min, 140±22 kg and 3.0±1.0 points, respectively) than in control animals (36.8±41 beats/min, 165±31 kg, 4.4±0.7 points; P<0.05). While no difference could be found in the observed behaviour, nocturnal hypothermia was elevated in restrictively fed animals. Our results indicate that ponies adapt to different climatic conditions by changing their metabolic rate, behaviour and some physiological parameters. When exposed to energy shortage, ponies, like wild herbivores, exhibited hypometabolism and nocturnal hypothermia. Keywords: Body temperature, Energy expenditure, Food restriction, Hypometabolism, Locomotor activity, Shetland pony |
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Shaw, E. B., Houpt, K. A., & Holmes, D. F. (1988). Body temperature and behaviour of mares during the last two weeks of pregnancy. Equine Vet J, 20(3), 199–202.
Abstract: Average daily core body temperature and behavioural patterns of pregnant mares were studied, in search of definitive signs of parturition within 24 h of the event. Nineteen pony mares were sampled twice daily for core body temperature. A significant temperature drop, averaging 0.1 degrees C (0.2 degrees F) was observed during the day prior to parturition. Between 18.00 h and 06.00 h, during the two weeks before parturition, Thoroughbred and Standardbred mares (n = 52) spent an average 66.8 per cent of their time standing, 27.0 per cent eating, 4.9 per cent lying in sternal recumbency, 1.0 per cent lying in lateral recumbency, and 0.3 per cent walking. On the night before parturition, mares spent significantly less time lying in sternal recumbency than on previous nights and on the night of parturition all behaviour patterns except eating were significantly different from the nights of the two weeks before parturition. There was an increase in walking (5.3 per cent), lying in sternal recumbency (8 per cent) and lying in lateral recumbency (5.3 per cent) whereas standing (53.3 per cent) was decreased. In 58 observed pregnancies, 54 mares (97 per cent) foaled in a recumbent position and 50 mares (86 per cent) foaled between 18.00 h and 06.00 h.
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Shettleworth, S. J. (1972). Stimulus relevance in the control of drinking and conditioned fear responses in domestic chicks (Gallus gallus). J Comp Physiol Psychol, 80(2), 175–198.
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Viscido, S. V., Miller, M., & Wethey, D. S. (2002). The dilemma of the selfish herd: the search for a realistic movement rule. J. Theor. Biol., 217(2), 183–194.
Abstract: The selfish herd hypothesis predicts that aggregations form because individuals move toward one another to minimize their own predation risk. The “dilemma of the selfish herd” is that movement rules that are easy for individuals to follow, fail to produce true aggregations, while rules that produce aggregations require individual behavior so complex that one may doubt most animals can follow them. If natural selection at the individual level is responsible for herding behavior, a solution to the dilemma must exist. Using computer simulations, we examined four different movement rules. Relative predation risk was different for all four movement rules (p<0.05). We defined three criteria for measuring the quality of a movement rule. A good movement rule should (a) be statistically likely to benefit an individual that follows it, (b) be something we can imagine most animals are capable of following, and (c) result in a centrally compact flock. The local crowded horizon rule, which allowed individuals to take the positions of many flock-mates into account, but decreased the influence of flock-mates with distance, best satisfied these criteria. The local crowded horizon rule was very sensitive to the animal's perceptive ability. Therefore, the animal's ability to detect its neighbors is an important factor in the dynamics of group formation.
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Boyd, L. (1986). Behavior problems of equids in zoos. Vet Clin North Am Equine Pract, 2(3), 653–664.
Abstract: Behavior problems in zoo equids commonly result from a failure to provide for needs basic to equine nature. Equids are gregarious, and failure to provide companions may result in pacing. Wild equids spend 60 to 70 per cent of their time grazing, and failure to provide ad libitum roughage contributes to the problems of pacing, cribbing, wood chewing, and coprophagia. Mimicking the normal processes of juvenile dispersal, bachelor-herd formation, and mate acquisition reduces the likelihood of agonistic and reproductive behavior problems. Infanticide can be avoided by introducing new stallions to herds containing only nonpregnant mares and older foals.
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Dyer, F. C. (2002). Animal behaviour: when it pays to waggle (Vol. 419).
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McCutcheon, L. J., & Geor, R. J. (2000). Influence of training on sweating responses during submaximal exercise in horses. J Appl Physiol, 89(6), 2463–2471.
Abstract: Sweating responses were examined in five horses during a standardized exercise test (SET) in hot conditions (32-34 degrees C, 45-55% relative humidity) during 8 wk of exercise training (5 days/wk) in moderate conditions (19-21 degrees C, 45-55% relative humidity). SETs consisting of 7 km at 50% maximal O(2) consumption, determined 1 wk before training day (TD) 0, were completed on a treadmill set at a 6 degrees incline on TD0, 14, 28, 42, and 56. Mean maximal O(2) consumption, measured 2 days before each SET, increased 19% [TD0 to 42: 135 +/- 5 (SE) to 161 +/- 4 ml. kg(-1). min(-1)]. Peak sweating rate (SR) during exercise increased on TD14, 28, 42, and 56 compared with TD0, whereas SRs and sweat losses in recovery decreased by TD28. By TD56, end-exercise rectal and pulmonary artery temperature decreased by 0.9 +/- 0.1 and 1.2 +/- 0.1 degrees C, respectively, and mean change in body mass during the SET decreased by 23% (TD0: 10.1 +/- 0.9; TD56: 7.7 +/- 0.3 kg). Sweat Na(+) concentration during exercise decreased, whereas sweat K(+) concentration increased, and values for Cl(-) concentration in sweat were unchanged. Moderate-intensity training in cool conditions resulted in a 1.6-fold increase in sweating sensitivity evident by 4 wk and a 0.7 +/- 0.1 degrees C decrease in sweating threshold after 8 wk during exercise in hot, dry conditions. Altered sweating responses contributed to improved heat dissipation during exercise and a lower end-exercise core temperature. Despite higher SRs for a given core temperature during exercise, decreases in recovery SRs result in an overall reduction in sweat fluid losses but no change in total sweat ion losses after training.
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Ratzlaff, M. H., Wilson, P. D., Hyde, M. L., Balch, O. K., & Grant, B. D. (1993). Relationship between locomotor forces, hoof position and joint motion during the support phase of the stride of galloping horses. Acta Anat (Basel), 146(2-3), 200–204.
Abstract: Three methods were used simultaneously to determine the relationships between the vertical forces exerted on the hooves and the positions of the limbs and hooves at the times of peak vertical forces from 2 horses galloping on a track straightaway. Vertical forces were recorded from an instrumented shoe, fetlock joint motion was measured with an electrogoniometer and the angles of the carpus, fetlock and hoof were determined from slow-motion films. At hoof contact, the mean angles of the carpus and fetlock were 181-182 degrees and 199-206 degrees, respectively. Peak vertical forces on the heel occurred at or near maximum extension of the carpal and fetlock joints. Peak forces on the toe occurred during flexion of the fetlock joint and at mean hoof angles of 28-31 degrees from the horizontal. The mean angles of the hoof from the horizontal at the time of heel contact were 6-7 degrees. Hoof lift occurred at mean carpal angles of 173-174 degrees and mean fetlock angles of 199-200 degrees.
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Gill, J. (1991). A new method for continuous recording of motor activity in horses. Comp Biochem Physiol A, 99(3), 333–341.
Abstract: 1. The use of an electronic recorder for the horse motor activity was described. 2. Examples of different types of motor activities are given in Figs 1-8. 3. The ultradian pattern of activity in all records was stressed. 4. The possibility of receiving of more physiological informations by this type of apparatus is discussed.
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