Stahlbaum, C. C., & Houpt, K. A. (1989). The role of the Flehmen response in the behavioral repertoire of the stallion. Physiol. Behav., 45(6), 1207–1214.
Abstract: The role of the Flehmen response in equine behavior was investigated under field and laboratory conditions. In Experiment 1, a field study made of five stallions on pasture with between three and eighteen mares each during the season indicated the following: 1) The Flehmen response was most frequently preceded by nasal, rather than oral, investigation of substances; 2) The stallions' rate of Flehmen varied with the estrous cycles of the mares; 3) The rate of Flehmen response did not show a variation with time of day; and 4) The Flehmen response was most frequently followed by marking behaviors rather than courtship behaviors. The results suggest that the Flehmen response is not an immediate component of sexual behavior, e.g., courtship of the stallion but may be involved in the overall monitoring of the mare's estrous cycle. Therefore the Flehmen response may contribute to the chemosensory priming of the stallion for reproduction. In Experiment 2 stallions were presented with urine or feces of mares in various stages of the reproductive cycle as well as with their own or other males' urine or feces. The occurrence of sniffing and Flehmen was used to determine the discriminatory ability of the stallions. Stallions can differentiate the sex of a horse on the basis of its feces alone, but cannot differentiate on the basis of urine. This ability may explain the function of fecal marking behavior of stallions.
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Waters, A. J., Nicol, C. J., & French, N. P. (2002). Factors influencing the development of stereotypic and redirected behaviours in young horses: findings of a four year prospective epidemiological study. Equine Vet J, 34(6), 572–579.
Abstract: Stereotypies are invariant and repetitive behaviour patterns that seemingly have no function, which tend to develop in captive animals faced with insoluble problems and may be indicative of reduced welfare. A 4 year prospective study of the factors influencing the development of stereotypic and redirected behaviours (abnormal behaviour) in a population of 225 young Thoroughbred and part-Thoroughbred horses was conducted between 1995 and 1999. Abnormal behaviour affected 34.7% of the population. Multivariable analysis showed that foals of low- or middle-ranking mares were less likely to develop abnormal behaviour than foals of dominant mares (rate ratio (RR) 0.23, P<0.01; RR 0.48, P<0.01, respectively). Weaning by confinement in a stable or barn was associated with an increased rate of development of abnormal behaviour, compared with paddock-weaning (RR 2.19, P<0.05), and housing in barns, rather than at grass after weaning, was associated with a further increase (RR 2.54, P<0.01). Specific stereotypic and redirected behaviours were then considered as separate outcomes. Crib-biting was initiated by 10.5% of horses at median age 20 weeks, weaving by 4.6% of horses at median age 60 weeks, box-walking by 2.3% of horses at median age 64 weeks and wood-chewing by 30.3% of horses at median age 30 weeks. Wood-chewing developed at a lower rate in horses born to subordinate or mid-ranking mares than in horses born to dominant mares (RR 0.29, P<0.01; RR 0.41, P<0.01, respectively), and at a higher rate in horses kept in barns or stables rather than at grass after weaning (RR 4.49, P<0.001; RR 1A6, P<0.001, respectively). Feeding concentrates after weaning was associated with a 4-fold increase in the rate of development of crib-biting (RR 4.12, P = 0.02). The results of this study support the idea that simple changes in feeding, housing and weaning practices could substantially lower the incidence of abnormal behaviour in young horses.
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Spadavecchia, C., Arendt-Nielsen, L., Andersen, O. K., Spadavecchia, L., Doherr, M., & Schatzmann, U. (2003). Comparison of nociceptive withdrawal reflexes and recruitment curves between the forelimbs and hind limbs in conscious horses. Am J Vet Res, 64(6), 700–707.
Abstract: OBJECTIVE: To compare nociceptive withdrawal reflexes (NWRs) evoked from the distal aspect of the left forelimb and hind limb in conscious standing horses and to investigate NWR recruitment for graded electrical stimulation intensities. ANIMALS: 20 adult horses. PROCEDURE: Surface electromyographic (EMG) activity evoked by transcutaneous electrical stimulation of the digital palmar (or plantar) nerve was recorded from the common digital extensor and cranial tibial muscles. Stimuli consisted of 25-millisecond train-of-5 constant current pulses. Current intensity was gradually increased until NWR threshold intensity was reached. The EMG signal was analyzed for quantification of the NWR. Behavioral responses accompanying the reflex were scored (scale, 0 to 5). The NWR recruitment curves were determined at 0.9, 1.1, 1.2, and 1.3 times the NWR threshold intensity. RESULTS: The NWR threshold was significantly higher for the hind limb (median value, 6.6 mA; range, 3 to 10 mA) than the forelimb (median, 3 mA; range, 1.7 to 5.5 mA). The NWR of the hind limb had a significantly longer latency (median, 122.8 milliseconds; range, 106 to 172 milliseconds), compared with the forelimb (median, 98 milliseconds; range, 86 to 137 milliseconds), and it was associated with significantly stronger behavioral reactions. Gradual increase of NWR amplitude was evident at increasing stimulation intensities and supported by the behavioral observations. CONCLUSIONS AND CLINICAL RELEVANCE: We documented NWRs evoked from the forelimb and hind limb and their recruitment with stimuli of increasing intensity in horses. These results provide a basis for use of NWRs in studies on nociceptive modulation in horses.
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Grubb, T. L., Foreman, J. H., Benson, G. J., Thurmon, J. C., Tranquilli, W. J., Constable, P. D., et al. (1996). Hemodynamic effects of calcium gluconate administered to conscious horses. J Vet Intern Med, 10(6), 401–404.
Abstract: Calcium gluconate was administered to conscious horses at 3 different rates (0.1, 0.2, and 0.4 mg/kg/min for 15 minutes each). Serum calcium concentrations and parameters of cardiovascular function were evaluated. All 3 calcium administration rates caused marked increases in both ionized and total calcium concentrations, cardiac index, stroke index, and cardiac contractility (dP/dtmax). Mean arterial pressure and right atrial pressure were unchanged; heart rate decreased markedly during calcium administration. Ionized calcium concentration remained between 54% and 57% of total calcium concentration throughout the study. We conclude that calcium gluconate can safely be administered to conscious horses at 0.1 to 0.4 mg/kg/min and that administration will result in improved cardiac function.
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Tavernor, W. D., & Lees, P. (1968). A pharmacological investigation of the influence of suxamethonium on cardiac function in the horse. Experientia, 24(6), 582–583.
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Alexander, F., Davies, M. E., & Muir, A. R. (1970). Bacteriophage-like particles in the large intestine of the horse. Res Vet Sci, 11(6), 592–593.
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Weik, H., & Altmann, J. (1972). The effect of L(+)-lactate on rat and horse adipose tissue in vitro. Zentralbl Veterinarmed A, 19(6), 514–518.
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Hertsch, B., & Becker, C. (1986). [Occurrence of aseptic necrosis of the palmar and plantar ligament in the horse--a contribution to the differentiation of sesamoid bone diseases]. Dtsch Tierarztl Wochenschr, 93(6), 263–266.
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de Waal, F. B. (1999). The end of nature versus nurture. Sci Am, 281(6), 94–99.
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Gardner, A., West, S. A. (2004). Cooperation and Punishment, Especially in Humans. Americ. Natur., 164(6), 753–764.
Abstract: Explaining altruistic cooperation is one of the greatest
challenges faced by sociologists, economists, and evolutionary biologists.
The problem is determining why an individual would carry
out a costly behavior that benefits another. Possible solutions to this
problem include kinship, repeated interactions, and policing. Another
solution that has recently received much attention is the threat
of punishment. However, punishing behavior is often costly for the
punisher, and so it is not immediately clear how costly punishment
could evolve. We use a direct (neighbor-modulated) fitness approach
to analyze when punishment is favored. This methodology reveals
that, contrary to previous suggestions, relatedness between interacting
individuals is not crucial to explaining cooperation through punishment.
In fact, increasing relatedness directly disfavors punishing
behavior. Instead, the crucial factor is a positive correlation between
the punishment strategy of an individual and the cooperation it
receives. This could arise in several ways, such as when facultative
adjustment of behavior leads individuals to cooperate more when
interacting with individuals who are more likely to punish. More
generally, our results provide a clear example of how the fundamental
factor driving the evolution of social traits is a correlation between
social partners and how this can arise for reasons other than genealogical
kinship.
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