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Vallortigara, G., & Rogers, L. J. (2005). Survival with an asymmetrical brain: advantages and disadvantages of cerebral lateralization. Behav Brain Sci, 28(4), 575–89; discussion 589–633.
Abstract: Recent evidence in natural and semi-natural settings has revealed a variety of left-right perceptual asymmetries among vertebrates. These include preferential use of the left or right visual hemifield during activities such as searching for food, agonistic responses, or escape from predators in animals as different as fish, amphibians, reptiles, birds, and mammals. There are obvious disadvantages in showing such directional asymmetries because relevant stimuli may be located to the animal's left or right at random; there is no a priori association between the meaning of a stimulus (e.g., its being a predator or a food item) and its being located to the animal's left or right. Moreover, other organisms (e.g., predators) could exploit the predictability of behavior that arises from population-level lateral biases. It might be argued that lateralization of function enhances cognitive capacity and efficiency of the brain, thus counteracting the ecological disadvantages of lateral biases in behavior. However, such an increase in brain efficiency could be obtained by each individual being lateralized without any need to align the direction of the asymmetry in the majority of the individuals of the population. Here we argue that the alignment of the direction of behavioral asymmetries at the population level arises as an “evolutionarily stable strategy” under “social” pressures occurring when individually asymmetrical organisms must coordinate their behavior with the behavior of other asymmetrical organisms of the same or different species.
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Rogers, L. J. (2000). Evolution of hemispheric specialization: advantages and disadvantages. Brain Lang, 73(2), 236–253.
Abstract: Lateralization of the brain appeared early in evolution and many of its features appear to have been retained, possibly even in humans. We now have a considerable amount of information on the different forms of lateralization in a number of species, and the commonalities of these are discussed, but there has been relatively little investigation of the advantages of being lateralized. This article reports new findings on the differences between lateralized and nonlateralized chicks. The lateralized chicks were exposed to light for 24 h on day 19 of incubation, a treatment known to lead to lateralization of a number of visually guided responses, and the nonlateralized chicks were incubated in the dark. When they were feeding, the lateralized chicks were found to detect a stimulus resembling a raptor with shorter latency than nonlateralized chicks. This difference was not a nonspecific effect caused by the light-exposed chicks being more distressed by the stimulus. Instead, it appears to be a genuine advantage conferred by having a lateralized brain. It is suggested that having a lateralized brain allows dual attention to the tasks of feeding (right eye and left hemisphere) and vigilance for predators (left eye and right hemisphere). Nonlateralized chicks appear to perform these dual tasks less efficiently than lateralized ones. Reference is made to other species in discussing these results.
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Apfelbach, R., Blanchard, C. D., Blanchard, R. J., Hayes, R. A., & McGregor, I. S. (2005). The effects of predator odors in mammalian prey species: A review of field and laboratory studies. Neuroscience and Biobehavioral Reviews, 29(8), 1123–1144.
Abstract: Prey species show specific adaptations that allow recognition, avoidance and defense against predators. For many mammalian species this includes sensitivity towards predator-derived odors. The typical sources of such odors include predator skin and fur, urine, feces and anal gland secretions. Avoidance of predator odors has been observed in many mammalian prey species including rats, mice, voles, deer, rabbits, gophers, hedgehogs, possums and sheep. Field and laboratory studies show that predator odors have distinctive behavioral effects which include (1) inhibition of activity, (2) suppression of non-defensive behaviors such as foraging, feeding and grooming, and (3) shifts to habitats or secure locations where such odors are not present. The repellent effect of predator odors in the field may sometimes be of practical use in the protection of crops and natural resources, although not all attempts at this have been successful. The failure of some studies to obtain repellent effects with predator odors may relate to (1) mismatches between the predator odors and prey species employed, (2) strain and individual differences in sensitivity to predator odors, and (3) the use of predator odors that have low efficacy. In this regard, a small number of recent studies have suggested that skin and fur-derived predator odors may have a more profound lasting effect on prey species than those derived from urine or feces. Predator odors can have powerful effects on the endocrine system including a suppression of testosterone and increased levels of stress hormones such as corticosterone and ACTH. Inhibitory effects of predator odors on reproductive behavior have been demonstrated, and these are particularly prevalent in female rodent species. Pregnant female rodents exposed to predator odors may give birth to smaller litters while exposure to predator odors during early life can hinder normal development. Recent research is starting to uncover the neural circuitry activated by predator odors, leading to hypotheses about how such activation leads to observable effects on reproduction, foraging and feeding. © 2005 Elsevier Ltd. All rights reserved.
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McGreevy, P. D., & McLean, A. N. (2007). Roles of learning theory and ethology in equitation. Journal of Veterinary Behavior: Clinical Applications and Research, 2(4), 108–118.
Abstract: By definition, ethology is primarily the scientific study of animal behavior, especially as it occurs in a natural environment; applied ethology being the study of animal behavior in the human domain. The terms equine ethology and ethological training are becoming commonplace in the equestrian domain, yet they seem to be used with a conspicuous lack of clarity and with no mention of learning theory. Most of what we do to train horses runs counter to their innate preferences. This article summarizes the ethological challenges encountered by working horses and considers the merits and limitations of ethological solutions. It also questions the use of terms such as “alpha” and “leader” and examines aspects of learning theory, equine cognition, and ethology as applied to horse training and clinical behavior modification. We propose 7 training principles that optimally account for the horse's ethological and learning abilities and maintain maximal responsivity in the trained horse. These principles can be summarized as: (1) use learning theory appropriately; (2) train easy-to-discriminate signals; (3) train and subsequently elicit responses singularly; (4) train only one response per signal; (5) train all responses to be initiated and subsequently completed within a consistent structure; (6) train persistence of current operantly conditioned responses; and (7) avoid and disassociate flight responses. Adherence to these principles and incorporating them into all horse training methodologies should accelerate training success, reduce behavioral wastage of horses, and improve safety for both humans and horses.
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Takimoto, A., & Fujita, K. (2008). Are horses (Equus caballus) sensitive to human attentional states? In IESM 2008.
Abstract: The ability to reliably detect what others are attending to seems important for social species to interact with their partners. Domestic horses (Equus caballus) have lived with humans for over five thousand years, hence they might have developed sensitivity to human attention. In the present study, we investigated whether horses would discriminate the situation in which a human experimenter could see them from the situation in which she could not. Specifically, we tested whether horses understand the role of eyes in human attentional states, produce more visual gestures when the experimenter can see their begging behaviors and produce more auditory or tactile gestures when she can not. We used with a slight modification the paradigm that previously yielded support for chimpanzee understanding of human attention (Hostetter et al. 2007). Twelve horses were offered food by the experimenter who showed various attentional states in front of them. We scored frequency of begging behaviors by the horses. In experiment 1, we set three kinds of condition: hand over the eyes, hand over the mouth and away. In the last condition there was only a food in front of horses, which was a control condition. The results showed that horses produced more auditory or tactile begging behaviors when the experimenter“s eyes were not visible than when her eyes were visible, but there was no difference in visual begging behaviors. In experiment 2, we set two kinds of condition: eyes closed and eyes open. The horses also produced more auditory or tactile begging behaviors when the experimenter”s eyes were closed than when they were open. However, there was no difference in visual begging behaviors. These results show that horses discriminate the situation in which humans can see from that in which humans can not. Of special interest, horses increased only auditory or tactile behaviors, not all types of communicative behaviors, when the experimenter could not see their begging behaviors. This result suggests that horses are sensitive to human attentional states. Moreover, horses may do recognize the eyes as an important indicator of whether or not humans will respond to their behavior and they may be able to behave flexibly depending upon human attentional states
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Ransom, J. I., & Cade, B. S. (2008). Influences of immunocontraception on intraband social behavior in free-ranging feral horses, Equus caballus. In IESM 2008.
Abstract: Contraception is often considered for population control of wildlife that is otherwise protected from more traditional management strategies, though little is known about the behavioral consequences of contraception in wild populations of socially complex animals. Feral horses, Equus caballus, in the western United States are ideal candidates for contraceptive management due to broad scale federal protection, reported herd growth rates of 15-25% per year, and finite public land allocated for them. We investigated influences of the immunocontraceptive porcine zona pellucida (PZP) on social behavior within bands of feral horses in three discrete populations for 4 years. Each band consisted of a single stallion and a harem of adult females (>1 years old) and their foals, as well as associated yearling males and females. Four mutually exclusive behaviors (herding, reproductive, harem tending, and agonism) were analyzed to investigate the difference between behaviors initiated by band stallions toward control mares and contracepted mares within the bands. Additionally, spatial relationships between each stallion and each harem female were analyzed to assess possible passive interactions. A candidate set of 22 hierarchical mixed effects models, using the discrete populations as a random effect on various fixed effects, was analyzed by restricted maximum likelihood estimation. The most supported models were selected by corrected Akaike"s Information Criteria (AICc). Analyses were done on 3 female age cohorts based on distinct fecundity rates: 2 to 5 year olds, 6 to 14 year olds, and > 14 year olds. Stallions herded control mares in the 6 to 14 year cohort more than contracepted mares (n=128, P=0.037) with treatment being the most supported effect, but foal presence (dependent foal) also contributed significantly to the model. Contracepted mares received more reproductive behaviors than control mares in the 6 to 14 year cohort (n = 151, P=0.020). No differences were detected in herding or reproductive behaviors in
the least fecund groups, the 2 to 5 year olds and ≥ 15 year olds. The only independent variable in the most supported model for reproductive behavior was treatment, and the covariates of foal presence, band residence (resident or transient female), band size, and body condition did not contribute. There were no differences in any age cohort for harem tending or agonism. Spatially, stallions maintained closer relationships with 2 to 5 year old contracepted females than with the same age control females (n=136, P < 0.001) while the group was feeding (at its most dispersed structure), but there were no differences while the band was resting or in locomotion. There were no spatial differences detected in the other age cohorts. Given the polyestrous nature and high fecundity of feral horses, the observed difference in reproductive behaviors between treatment groups was not surprising; however, the difference detected in herding rates was an unexpected behavioral modification. This change in herding behavior suggests that further investigation is needed to determine if PZP immunocontraception has an underlying influence on mare social rank, band structure, and band stability, as well as the scope and long-term importance of these behavioral dynamics.
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McDonnell, S. M. (2008). Human-horse interactions: Where are the behaviorists in 2008? In IESM 2008.
Abstract: This presentation will include commentary on work presented at this meeting as representative of the rapidly growing body of equine behavior science evidence relevant to human-horse interaction and the welfare of domestic, feral, and wild horses.
A substantial literature has accumulated detailing the behavior patterns of wild and feral equid populations, and to some extent the behavior of horses in various domestic environments. Professor Klingel“s and Professor Houpt”s work, begun nearly 40 years ago, continues to inspire colleagues around the world to qualitatively and quantitatively describe behavior of equids in a variety of environments. Several examples of the importance of this type of work to human-horse interaction and horse welfare are evident in throughout the meeting. It forms the basis for assessing the disturbance of behavior in wildlife management projects such as fertility manipulations (Hopkins; Ransom & Cade) or species reintroduction (Kaczensky et al). Study of the apparent variability in observations among the populations will lead to a better understanding of environmental and other factors, which will have fruitful application to welfare of horses both in domestic and natural environments. Unfortunately, this work always brings to mind what I perceive as a nagging threat to horse welfare and quality of human-horse interaction is misinformation concerning natural horse behavior. Natural horse behavior seems to be of great general interest, but unfortunately inaccuracies and misinterpretations are pervasive in popular “horse culture” and continue to be a conspicuous influence on management and training of domestic horses. Unfortunately, this misinformation often makes its way and influences equine education, both lay and equine science/veterinary education. Comparative observational study of behavior of horses in all settings by trained behaviorists, along with research designed to address purported implications for management of domestic horses, along with initiatives to transfer knowledge to educators at all levels should be encouraged.
Included in this meeting is considerable work addressing questions of domestic management practices such as forced weaning, transportation, stabling, and arbitrary grouping and regrouping of horses. This adds to a growing body of applied physiology and behavior research that has established trained behaviorists as a critical resource on teams making decisions on humane management.
Also well represented in work presented at this meeting is the exploration of cognition, perception, and temperament in horses. It is personally pleasing to see examples of direct investigation of the ability of the horse to respond to subtle human posture and gesture, which for many of us has represented just annoying possible confounders of earlier cognition studies. Behaviorists trained in perception and learning will no doubt contribute enormously to this exciting area of investigation.
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Connor, R. C., Wells, R. S., Mann, J., & Read, A. J. (2000). The bottlenose dolphin: Social relationships in a fission-fusion society. In J. Mann, R. C. Connor, P. L. Tyack, & H. Whitehead (Eds.), Cetacean Societies: Field Studies of Dolphins and Whales. (pp. 91–126). Chicago: University of Chicago Press.
Abstract: Book Description
“Part review, part testament to extraordinary dedication, and part call to get involved, Cetacean Societies highlights the achievements of behavioral ecologists inspired by the challenges of cetaceans and committed to the exploration of a new world.”-from the preface by Richard Wrangham
Long-lived, slow to reproduce, and often hidden beneath the water's surface, whales and dolphins (cetaceans) have remained elusive subjects for scientific study even though they have fascinated humans for centuries. Until recently, much of what we knew about cetaceans came from commercial sources such as whalers and trainers for dolphin acts. Innovative research methods and persistent efforts, however, have begun to penetrate the depths to reveal tantalizing glimpses of the lives of these mammals in their natural habitats.
Cetacean Societies presents the first comprehensive synthesis and review of these new studies. Groups of chapters focus on the history of cetacean behavioral research and methodology; state-of-the-art reviews of information on four of the most-studied species: bottlenose dolphins, killer whales, sperm whales, and humpback whales; and summaries of major topics, including group living, male and female reproductive strategies, communication, and conservation drawn from comparative research on a wide range of species.
Written by some of the world's leading cetacean scientists, this landmark volume will benefit not just students of cetology but also researchers in other areas of behavioral and conservation ecology as well as anyone with a serious interest in the world of whales and dolphins.
Contributors are Robin Baird, Phillip Clapham, Jenny Christal, Richard Connor, Janet Mann, Andrew Read, Randall Reeves, Amy Samuels, Peter Tyack, Linda Weilgart, Hal Whitehead, Randall S. Wells, and Richard Wrangham.
Keywords: cetacean social behavior, male alliance formation, most cetacean species, platanistid river dolphins, cetacean sociality, strategies and social bonds, female cetaceans, many cetologists, most mysticetes, sperm whale calves, passive fishing nets, variant whistles, historical whaling records, cetacean systematics, stable matrilineal groups, peak calving season, suction cup tags, mutualistic groups, cetacean vocalizations, focal animal studies, larger odontocetes, predictive signaling, individual cetaceans, sperm whale clicks, resident killer whales
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Berger, J. (1983). Induced abortion and social factors in wild horses. Nature, 303(5912), 59–61.
Abstract: Much evidence now suggests that the postnatal killing of young in primates and carnivores, and induced abortions in some rodents, are evolved traits exerting strong selective pressures on adult male and female behaviour. Among ungulates it is perplexing that either no species have developed convergent tactics or that these behaviours are not reported, especially as ungulates have social systems similar to those of members of the above groups. Only in captive horses (Equus caballus) has infant killing been reported. It has been estimated that 40,000 wild horses live in remote areas of the Great Basin Desert of North America (US Department of Interior (Bureau of Land Management), unpublished report), where they occur in harems (females and young) defended by males. Here I present evidence that, rather than killing infants directly, invading males induce abortions in females unprotected by their resident stallions and these females are then inseminated by the new males.
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Henderson, A. J. Z. (2007). Don't fence me in: managing psychological well being for elite performance horses. J. Appl. Anim. Welf. Sci., 10(4), 309–329.
Abstract: This article posits that stereotypical behavior patterns and the overall psychological well being of today's performance horse could be substantially enhanced with care that acknowledges the relationship between domesticated horses and their forerunners. Feral horses typically roam in stable, social groups over large grazing territories, spending 16-20 hr per day foraging on mid- to poor-quality roughage. In contrast, today's elite show horses live in relatively small stalls, eat a limited-but rich-diet at specific feedings, and typically live in social isolation. Although the horse has been domesticated for more than 6000 years, there has been no selection for an equid who no longer requires an outlet for these natural behaviors. Using equine stereotypies as a welfare indicator, this researcher proposes that the psychological well being of today's performance horse is compromised. Furthermore, the article illustrates how minimal management changes can enhance horses' well being while still remaining compatible with the requirements of the sport-horse industry. The article discusses conclusions in terms of Fraser, Weary, Pajor, and Milligan's “integrative welfare model” (1997).
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