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Ash, C., Chin, G., Pennisi, E., & Sugden, A. (2007). Living in Societies. Science, 317(5843), 1337–.
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Van Doorn G.S., Hengeveld G.M., & Weissing F.J. (2003). The Evolution of Social Dominance II: Multi-Player Models. Behavior, 140(10), 1333–1358.
Abstract: The social hierarchies observed in natural systems often show a high degree of transitivity. Transitive hierarchies do not only require rank differentiation within pairs of individuals but also a higher level ordering of relations within the group. Several authors have suggested that the formation of linear hierarchies at the group level is an emergent property of individual behavioural rules, referred to as winner and loser effects. Winner and loser effects occur if winners of previous conflicts are more likely to escalate the current conflict, whereas the losers of previous conflicts are less likely to do so. According to this idea, an individual's position in a hierarchy may not necessarily reflect its fighting ability, but may rather result from arbitrary historical asymmetries, in particular the history of victories and defeats. However, if this is the case, it is difficult to explain from an evolutionary perspective why a low ranking individual should accept its subordinate status. Here we present a game theoretical model to investigate whether winner and loser effects giving rise to transitive hierarchies can evolve and under which conditions they are evolutionarily stable. The main version of the model focuses on an extreme case in which there are no intrinsic differences in fighting ability between individuals. The only asymmetries that may arise between individuals are generated by the outcome of previous conflicts. We show that, at evolutionary equilibrium, these asymmetries can be utilized for conventional conflict resolution. Several evolutionarily stable strategies are based on winner and loser effects and these strategies give rise to transitive hierarchies.
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Outram, A. K., Stear, N. A., Bendrey, R., Olsen, S., Kasparov, A., Zaibert, V., et al. (2009). The Earliest Horse Harnessing and Milking. Science, 323(5919), 1332–1335.
Abstract: Horse domestication revolutionized transport, communications, and warfare in prehistory, yet the identification of early domestication processes has been problematic. Here, we present three independent lines of evidence demonstrating domestication in the Eneolithic Botai Culture of Kazakhstan, dating to about 3500 B.C.E. Metrical analysis of horse metacarpals shows that Botai horses resemble Bronze Age domestic horses rather than Paleolithic wild horses from the same region. Pathological characteristics indicate that some Botai horses were bridled, perhaps ridden. Organic residue analysis, using δ13C and δD values of fatty acids, reveals processing of mare's milk and carcass products in ceramics, indicating a developed domestic economy encompassing secondary products.
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Bugnyar, T., Stöwe, M., & Heinrich, B. (2004). Ravens, Corvus corax, follow gaze direction of humans around obstacles. Proc. Roy. Soc. Lond. B Biol. Sci., 271(1546), 1331–1336.
Abstract: The ability to follow gaze (i.e. head and eye direction) has recently been shown for social mammals, particularly primates. In most studies, individuals could use gaze direction as a behavioural cue without understanding that the view of others may be different from their own. Here, we show that hand–raised ravens not only visually co–orient with the look–ups of a human experimenter but also reposition themselves to follow the experimenter's gaze around a visual barrier. Birds were capable of visual co–orientation already as fledglings but consistently tracked gaze direction behind obstacles not before six months of age. These results raise the possibility that sub–adult and adult ravens can project a line of sight for the other person into the distance. To what extent ravens may attribute mental significance to the visual behaviour of others is discussed.
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Nguyen, N., Van Horn, R., Alberts, S., & Altmann, J. (2009). “Friendships” between new mothers and adult males: adaptive benefits and determinants in wild baboons (Papio cynocephalus). Behav. Ecol. Sociobiol., 63(9), 1331–1344.
Abstract: Close associations between adult males and lactating females and their dependent infants are not commonly described in non-monogamous mammals. However, such associations [sometimes called friendships (Smuts 1985)] are regularly observed in several primate species in which females mate with multiple males during the fertile period. The absence of mating exclusivity among friends suggests that males should invest little in infant care, raising questions about the adaptive significance of friendship bonds. Using data from genetic paternity analyses, patterns of behavior, and long-term demographic and reproductive records, we evaluated the extent to which friendships in four multi-male, multi-female yellow baboon (Papio cynocephalus) groups in Amboseli, Kenya represent joint parental care of offspring or male mating effort. We found evidence that mothers and infants benefited directly from friendships; friendships provided mother–infant dyads protection from harassment from other adult and immature females. In addition, nearly half of all male friends were the genetic fathers of offspring and had been observed mating with mothers during the days of most likely conception for those offspring. In contrast, nearly all friends who were not fathers were also not observed to consort with the mother during the days of most likely conception, suggesting that friendships between mothers and non-fathers did not result from paternity confusion. Finally, we found no evidence that prior friendship increased a male’s chances of mating with a female in future reproductive cycles. Our results suggest that, for many male–female pairs at Amboseli, friendships represented a form of biparental care of offspring. Males in the remaining friendship dyads may be trading protection of infants in exchange for some resources or services not yet identified. Our study is the first to find evidence that female primates gain social benefits from their early associations with adult males.
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Kenward, B., Rutz, C., Weir, A. A. S., & Kacelnik, A. (2006). Development of tool use in New Caledonian crows: inherited action patterns and social influences. Anim. Behav., 72(6), 1329–1343.
Abstract: New Caledonian crows, Corvus moneduloides, are the most advanced avian tool makers and tool users. We previously reported that captive-bred isolated New Caledonian crows spontaneously use twig tools and cut tools out of Pandanus spp. tree leaves, an activity possibly under cultural influence in the wild. However, what aspects of these behaviours are inherited and how they interact with individual and social experience remained unknown. To examine the interaction between inherited traits, individual learning and social transmission, we observed the ontogeny of twig tool use in hand-reared juveniles. Successful food retrieval was preceded by stereotyped object manipulation action patterns that resembled components of the mature behaviour, demonstrating that tool-oriented behaviours in this species are an evolved specialization. However, there was also an effect of social learning: juveniles that had received demonstrations of twig tool use by their human foster parent showed higher levels of handling and insertion of twigs than did their naive counterparts; a choice experiment showed that they preferred to handle objects that they had seen being manipulated by their human foster parent. Our observations are consistent with the hypothesis that individual learning, cultural transmission and creative problem solving all contribute to the acquisition of the tool-oriented behaviours in the wild, but inherited species-typical action patterns have a greater role than has been recognized.
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Jolly, A. (2007). BEHAVIOR: The Social Origin of Mind. Science, 317(5843), 1326–1327.
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Galef,, Bennett G. (1995). Why behaviour patterns that animals learn socially are locally adaptive. Anim. Behav., 49(5), 1325–1334.
Abstract: Recent models of the social transmission of behaviour by animals have repeatedly led their authors to the counterintuitive (and counterfactual) conclusion that traditional behaviour patterns in animals are often not locally adaptive. This deduction results from the assumption in such models that frequency of expression of socially learned behaviour patterns is not affected by rewards or punishments contingent upon their expression. An alternative approach to analysis of social learning processes, based on Staddon-Simmelhag's conditioning model, is proposed here. It is assumed that social interactions affect the probability of introduction of novel behaviour patterns into a naive individual's repertoire and that consequences of engaging in a socially learned behaviour determine whether that behaviour continues to be expressed. Review of several recently analysed instances of animal social learning suggests that distinguishing processes that introduce behaviour patterns into the repertoires of individuals from processes that select among behavioural alternatives aids in understanding observed differences in the longevity of various traditional behaviour patterns studied in both laboratory and field. Finally, implications of the present approach for understanding the role of social learning in evolutionary process are discussed.
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McGuigan, M. P., & Wilson, A. M. (2003). The effect of gait and digital flexor muscle activation on limb compliance in the forelimb of the horse Equus caballus. J Exp Biol, 206(Pt 8), 1325–1336.
Abstract: A horse's legs are compressed during the stance phase, storing and then returning elastic strain energy in spring-like muscle-tendon units. The arrangement of the muscle-tendon units around the lever-like joints means that as the leg shortens the muscle-tendon units are stretched. The forelimb anatomy means that the leg can be conceptually divided into two springs: the proximal spring, from the scapula to the elbow, and the distal spring, from the elbow to the foot. In this paper we report the results of a series of experiments testing the hypothesis that there is minimal scope for muscle contraction in either spring to adjust limb compliance. Firstly, we demonstrate that the distal, passive leg spring changes length by 127 mm (range 106-128 mm) at gallop and the proximal spring by 12 mm (9-15 mm). Secondly, we demonstrate that there is a linear relationship between limb force and metacarpo-phalangeal (MCP) joint angle that is minimally influenced by digital flexor muscle activation in vitro or as a function of gait in vivo. Finally, we determined the relationship between MCP joint angle and vertical ground-reaction force at trot and then predicted the forelimb peak vertical ground-reaction force during a 12 m s(-1) gallop on a treadmill. These were 12.79 N kg(-1) body mass (BM) (range 12.07-13.73 N kg(-1) BM) for the lead forelimb and 15.23 N kg(-1) BM (13.51-17.10 N kg(-1) BM) for the non-lead forelimb.
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Verheyen, K., Price, J., Lanyon, L., & Wood, J. (2006). Exercise distance and speed affect the risk of fracture in racehorses. Bone, 39(6), 1322–1330.
Abstract: In order to gain insight into those training regimens that can minimise the risk of fracture in athletic populations, we conducted a large epidemiological study in racehorses. Thoroughbred racehorses provide a suitable model for studying fracture development and exercise-related risk factors in physically active populations. They represent a homogeneous population, undertaking intensive exercise programmes that are sufficiently heterogeneous to determine those factors that influence injury risk. Daily exercise information was recorded for a cohort of 1178 thoroughbreds that were monitored for up to 2 years. A total of 148 exercise-induced fractures occurred in the study population. Results from a nested case-control study showed a strong interactive effect of exercise distances at different speeds on fracture risk. Horses that exceeded 44 km at canter (< or =14 m/s) and 6 km at gallop (>14 m/s) in a 30-day period were at particularly increased risk of fracture. These distances equate to ca. 7700 bone loading cycles at canter and 880 loading cycles at gallop. Fifty-six fractures occurred in the subset of study horses that were followed since entering training as yearlings, when skeletally immature (n = 335). Cohort analysis of this data set showed that, in previously untrained bones, accumulation of canter exercise increased the risk of fracture (P < or = 0.01), whereas accumulation of high-speed gallop exercise had a protective effect (P < 0.01). However, increasing distances at canter and gallop in short time periods (up to one month) were associated with an increasing fracture risk. All training exercise involves a balance between the risk of fracture inherent in exposure to loading and the beneficial effect that loading has by stimulating bone cells to produce a more robust architecture. Results from our study provide important epidemiological evidence of the effects of physical exercise on bone adaptation and injury risk and can be used to inform the design of safer exercise regimens in physically active populations.
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