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Whiten, A., Horner, V., Litchfield, C. A., & Marshall-Pescini, S. (2004). How do apes ape? Learn. Behav., 32(1), 36–52.
Abstract: In the wake of telling critiques of the foundations on which earlier conclusions were based, the last 15 years have witnessed a renaissance in the study of social learning in apes. As a result, we are able to review 31 experimental studies from this period in which social learning in chimpanzees, gorillas, and orangutans has been investigated. The principal question framed at the beginning of this era, Do apes ape? has been answered in the affirmative, at least in certain conditions. The more interesting question now is, thus, How do apes ape? Answering this question has engendered richer taxonomies of the range of social-learning processes at work and new methodologies to uncover them. Together, these studies suggest that apes ape by employing a portfolio of alternative social-learning processes in flexibly adaptive ways, in conjunction with nonsocial learning. We conclude by sketching the kind of decision tree that appears to underlie the deployment of these alternatives.
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Whiten, A., Custance, D. M., Gomez, J. C., Teixidor, P., & Bard, K. A. (1996). Imitative learning of artificial fruit processing in children (Homo sapiens) and chimpanzees (Pan troglodytes). J Comp Psychol, 110(1), 3–14.
Abstract: Observational learning in chimpanzees and young children was investigated using an artificial fruit designed as an analog of natural foraging problems faced by primates. Each of 3 principal components could be removed in 2 alternative ways, demonstration of only one of which was watched by each subject. This permitted subsequent imitation by subjects to be distinguished from stimulus enhancement. Children aged 2-4 years evidenced imitation for 2 components, but also achieved demonstrated outcomes through their own techniques. Chimpanzees relied even more on their own techniques, but they did imitate elements of 1 component of the task. To our knowledge, this is the first experimental evidence of chimpanzee imitation in a functional task designed to simulate foraging behavior hypothesized to be transmitted culturally in the wild.
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Whiten, A. (2000). Social complexity and social intelligence. In Novartis Foundation Symposium (Vol. 233, pp. 185–96; discussion pp. 196–201).
Abstract: When we talk of the 'nature of intelligence', or any other attribute, we may be referring to its essential structure, or to its place in nature, particularly the function it has evolved to serve. Here I examine both, from the perspective of the evolution of intelligence in primates. Over the last 20 years, the Social (or 'Machiavellian') Intelligence Hypothesis has gained empirical support. Its core claim is that the intelligence of primates is primarily an adaptation to the special complexities of primate social life. In addition to this hypothesis about the function of intellect, a secondary claim is that the very structure of intelligence has been moulded to be 'social' in character, an idea that presents a challenge to orthodox views of intelligence as a general-purpose capacity. I shall outline the principal components of social intelligence and the environment of social complexity it engages with. This raises the question of whether domain specificity is an appropriate characterization of social intelligence and its subcomponents, like theory of mind. As a counter-argument to such specificity I consider the hypothesis that great apes exhibit a cluster of advanced cognitive abilities that rest on a shared capacity for second-order mental representation.
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Whiten, A. (1998). Imitation of the sequential structure of actions by chimpanzees. J Comp Psychol, 11.
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Whiten A., & Byrne, R. W. (Eds.). (1997). Machiavellian Intelligence II – Extensions and Evaluations. Cambridge: Cambridge University Press.
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White, A. M., Swaisgood, R. R., & Czekala, N. (2007). Ranging patterns in white rhinoceros, Ceratotherium simum simum: implications for mating strategies. Appl. Anim. Behav. Sci., 74(2), 349–356.
Abstract: How animals use space has important consequences for feeding ecology, social organization, mating strategies and conservation management. In white rhinoceros, female home ranges are much larger than male territories, suggesting that movement patterns are influenced by factors other than resource distribution. In this study we placed radiotransmitters on 15 female white rhinoceros, recording 1758 locations and collecting behavioural data during 1671 observation sessions, making this the largest data set of its kind in this species. We investigated how habitat variables and male territories influenced female movement and reproductive behaviour. Female home ranges were approximately 20 km2 and core areas were 5 km2, with male territories roughly the same size as female core areas. Female range size did not vary with season, but the pattern of space use did vary. Females used grassland habitat preferentially, utilizing these areas significantly more than expected based on availability. Findings relevant to the mating strategy include: (1) the amount of grassland in a male's territory predicted female use of the territory; (2) the time that a female spent in a male's territory was a significant predictor of reproductive activity with the male, indicating that females probably mate with the most familiar male; and (3) the temporal pattern of female space use suggests that females did not increase mate sampling behaviour nor did they become more choosy about which males they visited when reproductively active. These findings suggest that males may maximize reproductive success by defending areas containing more grassland habitat.
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Whalen, A., Cownden, D., & Laland, K. (2015). The learning of action sequences through social transmission. Anim. Cogn., 18(5), 1093–1103.
Abstract: Previous empirical work on animal social learning has found that many species lack the ability to learn entire action sequences solely through reliance on social information. Conversely, acquiring action sequences through asocial learning can be difficult due to the large number of potential sequences arising from even a small number of base actions. In spite of this, several studies report that some primates use action sequences in the wild. We investigate how social information can be integrated with asocial learning to facilitate the learning of action sequences. We formalize this problem by examining how learners using temporal difference learning, a widely applicable model of reinforcement learning, can combine social cues with their own experiences to acquire action sequences. The learning problem is modeled as a Markov decision process. The learning of nettle processing by mountain gorillas serves as a focal example. Through simulations, we find that the social facilitation of component actions can combine with individual learning to facilitate the acquisition of action sequences. Our analysis illustrates that how even simple forms of social learning, combined with asocial learning, generate substantially faster learning of action sequences compared to asocial processes alone, and that the benefits of social information increase with the length of the action sequence and the number of base actions.
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Werhahn, H., Hessel, E. F., & Van den Weghe, H. F. A. (2012). Competition Horses Housed in Single Stalls (II): Effects of Free Exercise on the Behavior in the Stable, the Behavior during Training, and the Degree of Stress. Journal of Equine Veterinary Science, 32(1), 22–31.
Abstract: Although housing horses in single stalls limits their natural behavior to a great extent, this housing system is widespread in Germany, especially for competition horses. To improve the welfare of this system, free exercise on pastures or paddocks is deemed suitable, but it is also feared because of injuries and decreased willingness or motivation to perform. In the present study, three treatments were investigated with regard to their effect on the behavior of six competition horses in the stable, behavior during training, and on their degree of stress: daily training without free exercise (no turnout [NT]), solitary turnout for 2 hours after training, and 2-hour turnout in groups of two after training (group turnout). The horses' behavior in the stable was continuously analyzed through video recordings (2 pm to 6 am) on 3 days at the end of each treatment. The degree of stress was evaluated daily by heart rate variability at rest. The behavior during training was evaluated by a questionnaire answered by the riders, and the distance covered during training was measured by global positioning system. When NT was allowed, the horses showed less lying in the stable compared with the treatments with turnout. Heart rate variability measurements resulted in great individual differences, but generally, there was a higher degree of stress shown with the treatment NT according to the following parameters: standard deviation of inter-beat-intervals (SDNN), square root of the mean of the sum of the squares of differences between successive inter-beat-intervals (RMSSD), and ratio between low frequency and high frequency (LF/HF). The willingness to perform was evaluated as being slightly better in the treatments with turnout than in the treatment without turnout.
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Weishaupt, M. A., Wiestner, T., von Peinen, K., Waldern, N., Roepstorff, L., van Weeren, R., et al. (2006). Effect of head and neck position on vertical ground reaction forces and interlimb coordination in the dressage horse ridden at walk and trot on a treadmill. Equine Vet J Suppl, (36), 387–392.
Abstract: REASONS FOR PERFORMING STUDY: Little is known in quantitative terms about the influence of different head-neck positions (HNPs) on the loading pattern of the locomotor apparatus. Therefore it is difficult to predict whether a specific riding technique is beneficial for the horse or if it may increase the risk for injury. OBJECTIVE: To improve the understanding of forelimb-hindlimb balance and its underlying temporal changes in relation to different head and neck positions. METHODS: Vertical ground reaction force and time parameters of each limb were measured in 7 high level dressage horses while being ridden at walk and trot on an instrumented treadmill in 6 predetermined HNPs: HNP1 – free, unrestrained with loose reins; HNP2 – neck raised, bridge of the nose in front of the vertical; HNP3 – neck raised, bridge of the nose behind the vertical; HNP4 – neck lowered and flexed, bridge of the nose considerably behind the vertical; HNP5 – neck extremely elevated and bridge of the nose considerably in front of the vertical; HNP6 – neck and head extended forward and downward. Positions were judged by a qualified dressage judge. HNPs were assessed by comparing the data to a velocity-matched reference HNP (HNP2). Differences were tested using paired t test or Wilcoxon signed rank test (P<0.05). RESULTS: At the walk, stride duration and overreach distance increased in HNP1, but decreased in HNP3 and HNP5. Stride impulse was shifted to the forehand in HNP1 and HNP6, but shifted to the hindquarters in HNP5. At the trot, stride duration increased in HNP4 and HNP5. Overreach distance was shorter in HNP4. Stride impulse shifted to the hindquarters in HNP5. In HNP1 peak forces decreased in the forelimbs; in HNP5 peak forces increased in fore- and hindlimbs. CONCLUSIONS: HNP5 had the biggest impact on limb timing and load distribution and behaved inversely to HNP1 and HNP6. Shortening of forelimb stance duration in HNP5 increased peak forces although the percentage of stride impulse carried by the forelimbs decreased. POTENTIAL RELEVANCE: An extremely high HNP affects functionality much more than an extremely low neck.
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Weckerly, F. W. (1999). Social bonding and aggression in female Roosevelt elk. Can J Zool, 77(9), 1379–1384.
Abstract: Abstract: The relationship between degree of social bonding (extent of association among individuals) and level of aggression in ruminants is unclear. I examined social bonding and aggression in three groups of female Roosevelt elk (Cervus elaphus roosevelti) over 2 years. I hypothesized that when animals are socially bonded, bouts of aggression will be won by the individual initiating the aggression, occur quickly, and involve little physical contact, and the level of aggression does not correlate with group size. The degree of social bonding was high among individuals in all groups. Dyads of known individuals were together >80% of the time. A permutation analysis indicated that groups with the observed sizes had <0.001 chance of random association, except on one occasion when the probability was 0.72 for one group. Using focal-animal sampling, aggressive interactions were won 72% of the time by the initiator, occurred quickly (<5 s), and involved little physical contact, and the level of aggression was not correlated with group size. The level of aggression was, however, significantly lower in one of the groups. This group may have had access to more abundant food resources than the other groups. Socially bonded elk conducted aggressive interactions in a fashion that did not disrupt social stability. Résumé : La relation entre le degré de liaison sociale (importance des associations entre individus) et l`agressivité n`est pas claire chez les ruminants. J`ai étudié les liaisons sociales et l`agressivité chez trois groupes de femelles du Cerf de Roosevelt (Cervus elaphus roosevelti) pendant 2 ans. J`ai posé en hypothèse que, chez les animaux liés socialement, la victoire devrait être emportée par l`individu qui entreprend l`agression, l`agression devrait être de courte durée, se faire avec peu de contacts physiques et la fréquence des agressions ne devrait pas être liée à la taille du groupe. Des paires d`individus passaient plus de 80% de leur temps ensemble. Une analyse des permutations a démontré que, chez les groupes des tailles observées, la probabilité d`une association aléatoire était de moins de 0,001, sauf en un cas où cette probabilité a été évaluée à 0,72 chez un groupe. Par échantillonnage directionnel, j`ai observé que les interactions agressives étaient gagnées par l`individu attaquant 72% du temps, étaient de courte durée (<5 s), se faisaient avec peu de contacts physiques et leur fréquence n`était pas reliée à la taille du groupe. Il y avait cependant moins d`agressivité chez l`un des groupes. Il se peut que ce groupe ait eu accès à plus de ressources alimentaires que les autres. Chez les cerfs liés par des liens sociaux, l`agressivité ne se manifeste pas de façon à déséquilibrer la stabilité sociale.
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