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Tobin, T., & Combie, J. D. (1982). Performance testing in horses: a review of the role of simple behavioral models in the design of performance experiments. J Vet Pharmacol Ther, 5(2), 105–118.
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Tebbich, S., Bshary, R., & Grutter, A. S. (2002). Cleaner fish Labroides dimidiatus recognise familiar clients. Anim. Cogn., 5(3), 139–145.
Abstract: Individual recognition has been attributed a crucial role in the evolution of complex social systems such as helping behaviour and cooperation. A classical example for interspecific cooperation is the mutualism between the cleaner fish Labroides dimidiatus and its client reef fish species. For stable cooperation to evolve, it is generally assumed that partners interact repeatedly and remember each other's past behaviour. Repeated interactions may be achieved by site fidelity or individual recognition. However, as some cleaner fish have more than 2,300 interactions per day with various individuals per species and various species of clients, basic assumptions of cooperation theory might be violated in this mutualism. We tested the cleaner L. dimidiatus and its herbivorous client, the surgeon fish Ctenochaetus striatus, for their ability to distinguish between a familiar and an unfamiliar partner in a choice experiment. Under natural conditions, cleaners and clients have to build up their relationship, which is probably costly for both. We therefore predicted that both clients and cleaners should prefer the familiar partner in our choice experiment. We found that cleaners spent significantly more time near the familiar than the unfamiliar clients in the first 2 minutes of the experiment. This indicates the ability for individual recognition in cleaners. In contrast, the client C. striatus showed no significant preference. This could be due to a sampling artefact, possibly due to a lack of sufficient motivation. Alternatively, clients may not need to recognise their cleaners but instead remember the defined territories of L. dimidiatus to achieve repeated interactions with the same individual.
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Suzuki, Y., & Toquenaga, Y. (2005). Effects of information and group structure on evolution of altruism: analysis of two-score model by covariance and contextual analyses. J. Theor. Biol., 232(2), 191–201.
Abstract: An altruistic individual has to gamble on cooperation to a stranger because it does not know whether the stranger is trustworthy before direct interaction. Nowak and Sigmund (Nature 393 (1998a) 573; J. Theor. Biol. 194 (1998b) 561) presented a new theoretical framework of indirect reciprocal altruism by image scoring game where all individuals are informed about a partner's behavior from its image score without direct interaction. Interestingly, in a simplified version of the image scoring game, the evolutionarily stability condition for altruism became a similar form of Hamilton's rule, i.e. inequality that the probability of getting correct information is more than the ratio of cost to benefit. Since the Hamilton's rule was derived by evolutionarily stable analysis, the evolutionary meaning of the probability of getting correct information has not been clearly examined in terms of kin and group selection. In this study, we applied covariance analysis to the two-score model for deriving the Hamilton's rule. We confirmed that the probability of getting correct information was proportional to the bias of altruistic interactions caused by using information about a partner's image score. The Hamilton's rule was dependent on the number of game bouts even though the information reduced the risk of cooperation to selfish one at the first encounter. In addition, we incorporated group structure to the two-score model to examine whether the probability of getting correct information affect selection for altruism by group selection. We calculated a Hamilton's rule of group selection by contextual analysis. Group selection is very effective when either the probability of getting correct information or that of future interaction, or both are low. The two Hamilton's rules derived by covariance and contextual analyses demonstrated the effects of information and group structure on the evolution of altruism. We inferred that information about a partner's behavior and group structure can produce flexible pathways for the evolution of altruism.
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Stock, K. F., & Distl, O. (2005). Prediction of breeding values for osseous fragments in fetlock and hock joints, deforming arthropathy in hock joints, and pathologic changes in the navicular bones of Hanoverian Warmblood horses. Livestock Production Science, 92(1), 77–94.
Abstract: The results of a standardised radiological examination of 5928 Hanoverian Warmblood horses selected for sale at auction were used to predict relative breeding values (RBV) in the 23,662 horses included in the last four generations of the probands' pedigrees. The distribution of the RBV was investigated in the whole four-generation pedigree, in the probands and in those stallions that contributed at least three offspring to the probands. The RBV of the probands' sires were further compared with the officially published performance-based relative breeding values, i.e., total indices dressage (TID) and jumping (TIJ). The sires' level of dressage indices was considerably higher (mean TID=110) than of the jumping indices (mean TIJ=98). Total indices radiographic findings (TIR) were calculated for the sires with varying weighting given to the individual RBV. In each case, this resulted in a mean TIR of 99. Finally, total indices were derived from TIR and TID and/or TIJ in order to develop different selection schemes for all-purpose breeding and for breeding focused on dressage and show jumping. All breeding values under consideration increased by between 1% and 19% when radiographic findings were weighted with between 30% and 60% complementary to the corresponding performance parameters, and when only sires were selected with above average total indices. At the same time, the prevalences of the radiographic findings investigated here were lowered by up to 10% each. When only one radiographic finding was considered at a time, the maximum attainable response to selection was a 16-23% increase in the relative breeding values and a relative decrease in prevalences of radiographic findings of between 31% and 52%. The results of this study indicate that it is possible in horse breeding to consider simultaneously health and performance traits. Medical data should be included in the prediction of breeding values in order to improve the radiological status of today's riding 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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Smith, L. A., Wells, K. L., Marion, G., Swain, D. L., & Hutchings, M. R. (). Effects of group composition on the grazing behaviour of herbivores. Anim. Behav., In Press, Corrected Proof.
Abstract: Animal behaviour is often a function of the animal's physiological state. Groups of animals will often contain individuals with a range of physiological states and the grazing behaviour of herbivores is affected by their physiological state. This study compared the grazing decisions of animals in groups of single and mixed physiological states. Using a grazing model that simulated individual herbivore behaviour in relation to environmental distributions of forage resource (grass) and parasites (faeces), we tested the hypothesis that an animal's level of parasite exposure via the faecal-oral route is affected by the composition of physiological states in the group. Four physiological states were considered: parasite-naïve, parasitized, lactating and parasite-immune animals. Baseline parasite exposure levels for each state were generated by simulating single-state groups and were compared to simulations of each of the six two-state combinations. In single-state groups parasitized animals had the least and lactating animals had the greatest levels of parasite exposure. When co-grazing with lactating animals, parasitized, immune and naïve animals increased their parasite exposure, relative to single-state groups. When co-grazing with parasitized animals, lactating, immune and naïve animals reduced their parasite exposure, relative to single-state groups. There was no difference in parasite exposure of the immune or naïve animals co-grazing together when compared to the single-state groups. These results highlight the need to recognize the impact of the individual when studying group-living animals.
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Smith, J. E., Kolowski, J. M., Graham, K. E., Dawes, S. E., & Holekamp, K. E. (2008). Social and ecological determinants of fission-fusion dynamics in the spotted hyaena. Anim. Behav., 76(3), 619–636.
Abstract: Theory predicts that individuals living in fission-fusion societies, in which group members frequently change subgroups, should modify grouping patterns in response to varying social and environmental conditions. Spotted hyaenas, Crocuta crocuta, are long-lived carnivores that reside in permanent social groups called clans. Clans are complex, fission-fusion societies in which individual members travel, rest and forage in subgroups that frequently change composition. We studied two clans in Kenya to provide the first detailed description of fission-fusion dynamics in this species. Because social and ecological circumstances can influence the cohesiveness of animal societies, we evaluated the extent to which specific circumstances promote the formation of subgroups of various sizes. We found that cooperative defence of shared resources during interclan competition and protection from lions were cohesive forces that promoted formation of large subgroups. We also tested hypotheses suggesting factors limiting subgroup size. Mothers with small cubs avoided conspecifics, thereby reducing infanticide risk. Victims of aggression either reconciled fights or separated from former opponents to reduce the immediate costs of escalated aggression in the absence of food. As predicted by the ecological constraints hypothesis, hyaenas adjusted their grouping patterns over both short and long time scales in response to feeding competition. Crocuta were most gregarious during periods of abundant prey, joined clanmates at ephemeral kills in numbers that correlated with the energetic value of the prey and gained the most energy when foraging alone because cooperative hunting attracted numerous competitors. Overall, our findings indicate that resource limitation constrains grouping in this species.
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Sloet van Oldruitenborgh-Oosterbaan, M. M., Blok, M. B., Begeman, L., Kamphuis, M. C. D., Lameris, M. C., Spierenburg, A. J., et al. (2006). Workload and stress in horses: comparison in horses ridden deep and round ('rollkur') with a draw rein and horses ridden in a natural frame with only light rein contact. Tijdschr Diergeneeskd, 131(5), 152–157.
Abstract: 'Rollkur' or 'overbending' is the low and deep riding of a dressage horse during training or warming up. Lately, this technique has been criticized, and not necessarily objectively, on welfare grounds. To be able to evaluate these criticisms, more needs to be known about the workload and stress of horses being ridden 'rollkur'. The aim of the present study was to compare the workload of eight riding-school horses when being ridden deep and round with a draw rein ('rollkur') and when being ridden in a natural frame with only light rein contact ('free'). Workload (as measured by heart rate and blood lactate concentration) was slightly higher when horses were ridden 'rollkur' than when they were ridden 'free'. There were no differences in packed cell volume, or glucose and cortisol concentrations. No signs of uneasiness or stress could be determined when the horses were ridden 'rollkur'. Subjectively, all horses improved their way of moving during 'rollkur' and were more responsive to their rider.
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Skov-Rackette, S. I., & Shettleworth, S. J. (2005). What do rats learn about the geometry of object arrays? Tests with exploratory behavior. J Exp Psychol Anim Behav Process, 31(2), 142–154.
Abstract: Six experiments using habituation of exploratory behavior tested whether disoriented rats foraging in a large arena encode the shapes of arrays of objects. Rats did not respond to changes in position of a single object, but they responded to a change in object color and to a change in position of 1 object in a square array, as in previous research (e.g., C. Thinus-Blanc et al., 1987). Rats also responded to an expansion of a square array, suggesting that they encoded sets of interobject distances rather than overall shape. In Experiments 4-6, rats did not respond to changes in sense of a triangular array that maintained interobject distances and angles. Shapes of object arrays are encoded differently from shapes of enclosures.
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Shettleworth, S. J., & Sutton, J. E. (2005). Multiple systems for spatial learning: dead reckoning and beacon homing in rats. J Exp Psychol Anim Behav Process, 31(2), 125–141.
Abstract: Rats homed with food in a large lighted arena. Without visual cues, they used dead reckoning. When a beacon indicated the home, rats could also use the beacon. Homing did not differ in 2 groups of rats, 1 provided with the beacon and 1 without it; tests without the beacon gave no evidence that beacon learning overshadowed dead reckoning (Experiment 1). When the beacon was at the home for 1 group and in random locations for another, there was again no evidence of cue competition (Experiment 2). Dead reckoning experience did not block acquisition of beacon homing (Experiment 3). Beacon learning and dead reckoning do not compete for predictive value but acquire information in parallel and are used hierarchically.
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