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McLean, A. N., & Christensen, J. W. (2017). The application of learning theory in horse training. Appl. Anim. Behav. Sci., 190, 18–27.
Abstract: The millennia-old practices of horse training markedly predate and thus were isolated from the mid-twentieth century revelation of animal learning processes. From this standpoint, the progress made in the application and understanding of learning theory in horse training is reviewed including a discussion of how learning processes are employed or otherwise under-utilised in training. This review describes the process of habituation and the most commonly applied desensitisation techniques (systematic desensitisation, counter-conditioning, overshadowing, response prevention) and propose two additional techniques (approach conditioning and stimulus blending). The salience of different types of cues, the interaction of operant and classical conditioning and the impact of stress are also discussed. This paper also exposes the inflexibility and occasional inadequacy of the terminology of learning theory when translated from the research laboratory situation to the practical setting in horse training. While learning theory provides a rich toolbox for riders and trainers, the training process is subject to the simultaneous use of multiple learning processes. In addition, learning/behavioural outcomes and trained responses are not just the result of simple stimulus-response based interactions but are further shaped by arousal, affective and attachment states. More research is needed in these areas. For the field of equitation science to progress and to improve clarity and use of learning processes, changes in nomenclature are required. In particular, the use of the terms 'positive' and 'negative' as descriptive labels in both reinforcement and punishment modalities are unacceptably misleading for everyday use. These labels inhibit the understanding and recognition of the learning processes that these terms supposedly represent, yet the learning processes they describe are vital for horse riders, handlers and trainers to understand. We therefore propose that these labels should be re-labelled more appropriately as 'addition' or 'subtraction' reinforcement/punishment. This would enlighten trainers on the correct application of learning theory, and safety and welfare benefits for people and horses would follow. Finally it is also proposed that the term 'conflict theory' be taken up in equitation science to facilitate diagnosis of training-related behaviour disorders and thus enable the emergence of improved training practices. The optimal use of learning theory should be established as a fundamental principle in equestrian education.
Keywords: Behaviour; Conflict theory; Desensitisation; Habituation; Reinforcement; Stress
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Pick, D. K., B., & Steciuch, C. (2015). The Familiarity Heuristic in the Horse (Equus caballus). In , & K. Krueger (Ed.), Proceedings of the 3. International Equine Science Meeting. Wald: Xenophon Publishing.
Abstract: This study replicated an unreported finding observed in a color perception experiment (Pick, Lovell, Brown, & Dail, 1994) where, after using the method of successive approximations to train a blue-gray discrimination, red-gray trials were initiated without further training. Although a gray choice had never been reinforced, the subject chose gray on the first 20 trials (p < .000001). In the study reported here, a horse was trained to approach a red feed bucket and not a green feed bucket. After the subject mastered the discrimination, a blue bucket was substituted for the previously reinforced red bucket. With double-blind controls in place, the subject chose the unreinforced green bucket on 15 out of the first 20 blue-green trials yielding a binomial p = 0.0148 that this outcome could be due to chance alone. These results are contrary to all behavioristic psychological learning theories, but consistent with prospect theory (Kahneman & Tversky, 1979). Prospect theory predicts that given a choice between two previously unreinforced stimuli, one familiar and the other novel, humans will choose the familiar. It is argued that the bias toward the familiar is the basis to a heuristic that has a genetic origin and should exist in other animals on the phylogenetic scale. The results of this study indicate that the heuristic is available at least as far down the scale as the horse. Conceptual replications using shape stimuli and sound stimuli are in progress.
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Hasenjager, M. J., & Dugatkin, L. A. Social Network Analysis in Behavioral Ecology. Advances in the Study of Behavior. Academic Press.
Abstract: Abstract In recent years, behavioral ecologists have embraced social network analysis (SNA) in order to explore the structure of animal societies and the functional consequences of that structure. We provide a conceptual introduction to the field that focuses on historical developments, as well as on novel insights generated by recent work. First, we discuss major advances in the analysis of nonhuman societies, culminating in the use of SNA by behavioral ecologists. Next, we discuss how network-based approaches have enhanced our understanding of social structure and behavior over the past decade, focusing on: (1) information transmission, (2) collective behaviors, (3) animal personality, and (4) cooperation. These behaviors and phenomena possess several features—e.g., indirect effects, emergent properties—that network analysis is well equipped to handle. Finally, we highlight recent developments in SNA that are allowing behavioral ecologists to address increasingly sophisticated questions regarding the structure and function of animal sociality.
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Newton-Fisher, N. E., & Lee, P. C. (2011). Grooming reciprocity in wild male chimpanzees. Anim. Behav., 81(2), 439–446.
Abstract: Understanding cooperation between unrelated individuals remains a central problem in animal behaviour; evolutionary mechanisms are debated, and the importance of reciprocity has been questioned. Biological market theory makes specific predictions about the occurrence of reciprocity in social groups; applied to the social grooming of mammals, it predicts reciprocity in the absence of other benefits for which grooming can be exchanged. Considerable effort has been made to test this grooming trade model in nonhuman primates; such studies show mixed results, but may be confounded by kin effects. We examined patterns of reciprocity within and across bouts, and tested predictions of the grooming trade model, among wild male chimpanzees, Pan troglodytes: a system with negligible kin effects. In accord with the model's expectations, we found that some grooming was directed by lower- to higher-ranked individuals, and that, on average, higher-ranked individuals groomed more reciprocally. We found no support, however, for a prediction that more reciprocity should occur between individuals close in rank. For most dyads, reciprocity of effort occurred through unbalanced participation in grooming bouts, but reciprocity varied considerably between dyads and only a small proportion showed strongly reciprocal grooming. Despite this, each male had at least one reciprocal grooming relationship. In bouts where both individuals groomed, effort was matched through mutual grooming, not alternating roles. Our results provide mixed support for the current grooming trade, biological market model, and suggest that it needs to incorporate risks of currency inflation and cheating for species where reciprocity can be achieved through repeated dyadic interactions.
Keywords: altruism; biological market theory; Budongo; chimpanzee; grooming; Pan troglodytes
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Mesterton-Gibbons, M., Gavrilets, S., Gravner, J., & Akçay, E. (2011). Models of coalition or alliance formation. J. Theor. Biol., 274(1), 187–204.
Abstract: More than half a century has now elapsed since coalition or alliance formation theory (CAFT) was first developed. During that time, researchers have amassed a vast amount of detailed and high-quality data on coalitions or alliances among primates and other animals. But models have not kept pace, and more relevant theory is needed. In particular, even though CAFT is primarily an exercise in polyadic game theory, game theorists have devoted relatively little attention to questions that motivate field research, and much remains largely unexplored. The state of the art is both a challenge and an opportunity. In this review we describe a variety of game-theoretic and related modelling approaches that have much untapped potential to address the questions that field biologists ask.
Keywords: Game theory; Cooperation
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Sterck, E., Watts, D., & van Schaik, C. (1997). The evolution of female social relationships in nonhuman primates. Behav. Ecol. Sociobiol., 41(5), 291–309.
Abstract: Considerable interspeci®c variation in female social relationships occurs in gregarious primates, particularly with regard to agonism and cooperation between females and to the quality of female relationships with males. This variation exists alongside variation in female philopatry and dispersal. Socioecological theories have tried to explain variation in female-female social relationships from an evolutionary perspective focused on ecological factors, notably predation and food distribution. According to the current ``ecological model'', predation risk forces females of most diurnal primate species to live in groups; the strength of the contest component of competition for resources within and between groups then largely determines social relationships between females. Social elationships among gregarious females are here characterized as DispersalEgalitarian, Resident-Nepotistic, Resident-Nepotistic-Tolerant, or Resident-Egalitarian. This ecological model has successfully explained i€erences in the occurrence of formal submission signals, decided dominance relation ships, coalitions and female philopatry. Group size and female rank generally a€ect female reproduction success as the model predicts, and studies of closely related species in di€erent ecological circumstances underscore the importance of the model. Some cases, however, can only be explained when we extend the model to incorporate the e€ects of infanticide risk and habitat saturation. We review evidence in support of the ecological model and test the power of alternative models that invoke between-group competition, forced female philopatry, demographic female recruitment, male interventions into female aggression, and male harassment.
Not one of these models can replace the ecological model, which already encompasses the between-group competition. Currently the best model, which explains several phenomena that the ecological model does not, is a ``socioecological model'' based on the combined importance of ecological factors, habitat saturation and infanticide avoidance. We note some points of similarity and divergence with other mammalian taxa; these remain to be explored in detail. Keywords: ecology; matrilocal; primate; social; theory
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Newman, M. E. J. (2003). The Structure and Function of Complex Networks. SIAM Rev., 45(2), 167–256.
Abstract: Inspired by empirical studies of networked systems such as the Internet, social networks, and biological networks, researchers have in recent years developed a variety of techniques and models to help us understand or predict the behavior of these systems. Here we review developments in this field, including such concepts as the small-world effect, degree distributions, clustering, network correlations, random graph models, models of network growth and preferential attachment, and dynamical processes taking place on networks.
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Conradt, L., Krause, J., Couzin, I. D., & Roper, T. J. (2009). “Leading According to Need” in Self-Organizing Groups. Am Nat, 173(3), 304–312.
Abstract: Self‐organizing‐system approaches have shed significant light on the mechanisms underlying synchronized movements by large groups of animals, such as shoals of fish, flocks of birds, or herds of ungulates. However, these approaches rarely consider conflicts of interest between group members, although there is reason to suppose that such conflicts are commonplace. Here, we demonstrate that, where conflicts exist, individual members of self‐organizing groups can, in principle, increase their influence on group movement destination by strategically changing simple behavioral parameters (namely, movement speed, assertiveness, and social attraction range). However, they do so at the expense of an increased risk of group fragmentation and a decrease in movement efficiency. We argue that the resulting trade‐offs faced by each group member render it likely that group movements are led by those members for which reaching a particular destination is most crucial or group cohesion is least important. We term this phenomenon leading according to “need” or “social indifference,” respectively. Both kinds of leading can occur in the absence of knowledge of or communication about the needs of other group members and without the assumption of altruistic cooperation. We discuss our findings in the light of observations on fish and other vertebrates.
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Conradt, L., & Roper, T. J. (2010). Deciding group movements: Where and when to go. Behav. Process., 84(3), 675–677.
Abstract: A group of animals can only move cohesively, if group members “somehow” reach a consensus about the timing (e.g., start) and the spatial direction/destination of the collective movement. Timing and spatial decisions usually differ with respect to the continuity of their cost/benefit distribution in such a way that, in principle, compromises are much more feasible in timing decision (e.g. median preferred time) than they are in spatial decisions. The consequence is that consensus costs connected to collective timing decisions are usually less skewed amongst group members than are consensus costs connected to spatial decisions. This, in turn, influences the evolution of decision sharing: sharing in timing decisions is most likely to evolve when conflicts are high relative to group cohesion benefits, while sharing in spatial decisions is most likely to evolve in the opposite situation. We discuss the implications of these differences for the study of collective movement decisions.
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Goodwin, D., McGreevy, P., Waran, N., & McLean, A. (2009). How equitation science can elucidate and refine horsemanship techniques. Special Issue: Equitation Science, 181(1), 5–11.
Abstract: The long-held belief that human dominance and equine submission are key to successful training and that the horse must be taught to [`]respect' the trainer infers that force is often used during training. Many horses respond by trialling unwelcome evasions, resistances and flight responses, which readily become established. When unable to cope with problem behaviours, some handlers in the past might have been encouraged to use harsh methods or devices while others may have called in a so-called [`]good horseman' or [`]horse whisperer' to remediate the horse. Frequently, the approaches such practitioners offer could not be applied by the horse's owner or trainer because of their lack of understanding or inability to apply the techniques. Often it seemed that these [`]horse-people' had magical ways with horses (e.g., they only had to whisper to them) that achieved impressive results although they had little motivation to divulge their techniques. As we begin to appreciate how to communicate with horses sensitively and consistently, misunderstandings and misinterpretations by horse and trainer should become less common. Recent studies have begun to reveal what comprises the simplest, most humane and most effective mechanisms in horse training and these advances are being matched by greater sharing of knowledge among practitioners. Indeed, various practitioners of what is referred to here as [`]natural horsemanship' now use techniques similar to the [`]whisperers' of old, but they are more open about their methods. Reputable horse trainers using natural horsemanship approaches are talented observers of horse behaviour and respond consistently and swiftly to the horse's subtle cues during training. For example, in the roundpen these trainers apply an aversive stimulus to prompt a flight response and then, when the horse slows down, moves toward them, or offers space-reducing affiliative signals, the trainer immediately modifies his/her agonistic signals, thus negatively reinforcing the desired response. Learning theory and equine ethology, the fundamentals of the emerging discipline of equitation science, can be used to explain almost all the behaviour modification that goes on in these contexts and in conventional horsemanship. By measuring and evaluating what works and what does not, equitation science has the potential to have a unifying effect on traditional practices and developing branches of equitation.
Keywords: Horse; Training; Equitation; Learning theory; Ethology; Cognition
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