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Shettleworth, S. J., & Juergensen, M. R. (1980). Reinforcement and the organization of behavior in golden hamsters: brain stimulation reinforcement for seven action patterns. J Exp Psychol Anim Behav Process, 6(4), 352–375.
Abstract: Golden hamsters were reinforced with intracranial electrical stimulation of the lateral hypothalamus (ICS) for spending time engaging in one of seven topographically defined action patterns (APs). The stimulation used as reinforcer elicited hoarding and/or feeding and supported high rates of bar pressing. In Experiment 1, hamsters were reinforced successively for digging, open rearing, and face washing. Digging increased most in time spent, and face washing increased least. Experiments 2-5 examined these effects further and also showed that “scrabbling,” like digging, was performed a large proportion of the time, almost without interruption, for contingent ICS but that scratching the body with a hindleg and scent-marking showed relatively little effect of contingent ICS, the latter even in an environment that facilitated marking. In Experiment 6, naive hamsters received ICS not contingent on behavior every 30 sec (fixed-time 30-sec schedule). Terminal behaviors that developed on this schedule were APs that were easy to reinforce in the other experiments, but a facultative behavior, face washing, was one not so readily reinforced. Experiment 7 confirmed a novel prediction from Experiment 6--that wall rearing, a terminal AP, would be performed at a high level for contingent ICS. All together, the results point to both motivational factors and associative factors being involved in the considerable differences in performance among different reinforced activities.
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Shoshani, J., Kupsky, W. J., & Marchant, G. H. (2006). Elephant brain. Part I: gross morphology, functions, comparative anatomy, and evolution. Brain Res Bull, 70(2), 124–157.
Abstract: We report morphological data on brains of four African, Loxodonta africana, and three Asian elephants, Elephas maximus, and compare findings to literature. Brains exhibit a gyral pattern more complex and with more numerous gyri than in primates, humans included, and in carnivores, but less complex than in cetaceans. Cerebral frontal, parietal, temporal, limbic, and insular lobes are well developed, whereas the occipital lobe is relatively small. The insula is not as opercularized as in man. The temporal lobe is disproportionately large and expands laterally. Humans and elephants have three parallel temporal gyri: superior, middle, and inferior. Hippocampal sizes in elephants and humans are comparable, but proportionally smaller in elephant. A possible carotid rete was observed at the base of the brain. Brain size appears to be related to body size, ecology, sociality, and longevity. Elephant adult brain averages 4783 g, the largest among living and extinct terrestrial mammals; elephant neonate brain averages 50% of its adult brain weight (25% in humans). Cerebellar weight averages 18.6% of brain (1.8 times larger than in humans). During evolution, encephalization quotient has increased by 10-fold (0.2 for extinct Moeritherium, approximately 2.0 for extant elephants). We present 20 figures of the elephant brain, 16 of which contain new material. Similarities between human and elephant brains could be due to convergent evolution; both display mosaic characters and are highly derived mammals. Humans and elephants use and make tools and show a range of complex learning skills and behaviors. In elephants, the large amount of cerebral cortex, especially in the temporal lobe, and the well-developed olfactory system, structures associated with complex learning and behavioral functions in humans, may provide the substrate for such complex skills and behavior.
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Shultz, S., & Dunbar, R. I. M. (2006). Both social and ecological factors predict ungulate brain size. Proc Biol Sci, 273(1583), 207–215.
Abstract: Among mammals, the members of some Orders have relatively large brains. Alternative explanations for this have emphasized either social or ecological selection pressures favouring greater information-processing capacities, including large group size, greater foraging efficiency, higher innovation rates, better invasion success and complex problem solving. However, the focal taxa for these analyses (primates, carnivores and birds) often show both varied ecological competence and social complexity. Here, we focus on the specific relationship between social complexity and brain size in ungulates, a group with relatively simple patterns of resource use, but extremely varied social behaviours. The statistical approach we used, phylogenetic generalized least squares, showed that relative brain size was independently associated with sociality and social complexity as well as with habitat use, while relative neocortex size is associated with social but not ecological factors. A simple index of sociality was a better predictor of both total brain and neocortex size than group size, which may indicate that the cognitive demands of sociality depend on the nature of social relationships as well as the total number of individuals in a group.
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Simpson, B. S. (2002). Neonatal foal handling. Appl. Anim. Behav. Sci., 78(2-4), 303–317.
Abstract: Recent interest has focused on the advantage of intensively handling young foals as a means of producing more tractable horses, accustomed to humans and receptive to training. To date, the effect of this intensive handling, dubbed “imprint training” in the popular literature, has not been tested. The present study compares seven foals handled intensively as neonates with eight untreated controls. The handling protocol started from 2-8 h after birth and continued daily for a total of 5 days. The protocol consisted of a series of stimuli and experiences that were each repeated until the foal no longer resisted or reacted negatively. Subsequently, foals were tested before weaning, at 4 months of age. Results indicated that handled foals (HF) ranked higher than control foals (CF) in subjective ratings of calmness (*P<0.0117) and friendliness (*P<0.0001) and in several specific handling tasks (venipuncture *P<0.0220; placing in stock *P<0.0128). Although, in approach tests all foals but one allowed approach of a person to 4 m, significantly more HF approached the person than CF (P<0.0080). In stimulus tests, foals were presented specific stimuli to which they had been tested as neonates. Two of eight CF were too unruly and dangerous to test. Of foals that could be tested, CF required significantly more time to hook-up a heart rate monitor (**P<0.0055). Split-plot analysis indicated that HF had lower heart rates to initial left-sided stimuli, presented first, than CF (*P<0.0421). In response to right-sided stimuli, heart rate scores of CF were not significantly different from HF (P<0.2259), suggesting reduced reactivity over time due to a learning effect. Behavioral responses to specific stimuli did not differ between CF and HF, suggesting that neonatal handling has a general rather than specific effect on subsequent behavior. Cortisol concentrations were measured before and after testing and the difference calculated. All foals had higher post-testing levels than pre-testing levels. There was a significant difference between HF and CF, indicating greater reactivity among the CF (*P<0.050). In general, the results indicated that foals handled as neonates were more tractable and less reactive. Specific neonatal handling tasks, such as sticking a finger up the foal's nose or patting the bottom of the foot, seemed to have no beneficial effect on related tasks such as passing a nasogastric tube or tapping with a farrier's hammer at 4 months of age. Mechanisms for the observed effect of neonatal handling require further investigation.
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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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Sondergaard, E., & Ladewig, J. (2004). Group housing exerts a positive effect on the behaviour of young horses during training. Appl. Anim. Behav. Sci., 87(1-2), 105–118.
Abstract: In an experiment on the effects of social environment and training on the human-animal relationship, 20 horses were handled according to a defined schedule. Eight horses were housed singly and 12 horses were housed in four groups of 3 horses. Horses were handled three times per week in 10 min sessions from an age of 6 months until 2 years of age during two winter periods. A total of 50 and 70 sessions were given in the first and second period, respectively. Five randomly allocated people performed the training. The training scheme involved leading, tying up, touching, lifting feet, etc. in 43 stages. The horse had to fulfil the performance criteria of each stage in order to get to the next stage. In the first winter period, horses were led to the stable when they had “passed” a stage or after 10 min of training. In the second winter period, horses would start off at stage 1 again, and when they “passed” a stage they went on to the next stage within the same training session. Because of the change in training procedure results were analysed separately for the two winter periods. There was a significant difference between trainers in the number of times they allowed a horse to “pass” a stage within each winter period (χ32, P<0.05; χ32, P<0.001 for the first and the second winter period, respectively). Group housed horses “passed” more stages than single housed horses (17 versus 14; 27 versus 18 in the first and second winter period, respectively; P<0.05 for the interaction). Singly housed horses bit the trainer more frequently than did group housed horses (P<0.01). The responses of group housed horses to training clearly demonstrate the benefits of raising young horses in groups.
Keywords: Horses; Housing; Human-animal relationship; Social environment; Training; Learning
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Spier, S. J., Berger Pusterla, J., Villarroel, A., & Pusterla, N. (2004). Outcome of tactile conditioning of neonates, or “imprint training” on selected handling measures in foals. The Veterinary Journal, 168(3), 252–258.
Abstract: Behavioural reactions to selected handling procedures were compared between conditioned, or imprint-trained, and untrained foals raised on the same farm. Nineteen randomly chosen healthy foals were imprint trained at birth and 24 h later (Group A). Twenty-one similar foals that were not imprint-trained served as age-matched controls (Group B). Training began within 10 min of birth and consisted of touch desensitization by gentle rubbing. Each tactile stimulus was repeated 30-50 times over 45-60 min, until the foal no longer resisted the procedure and appeared relaxed. The procedure was then repeated at 24 h of age. At that time a physical examination and blood analysis were performed to assess the foals' health status. Group B animals were not handled except for a brief physical examination and blood analysis at 24 h of age. Thereafter all foals were kept on pastures with their dams with no further handling until they were three months of age. Any foals handled for other reasons before that time were excluded from the study. At three months, each of the 28 foals that completed the study experienced the following handling procedures: acceptance of restraint, haltering, complete physical examination, acceptance of a plastic rebreathing bag, touching the whole body, intramuscular vaccination in the neck, intranasal vaccination, and deworming with oral paste. Response to each procedure was scored (1=not resistant, 2=low resistance, 3=strong resistance, 4=not possible without major physical restraint). Conditioned foals (Group A) were significantly less resistant to touching the front and hind legs and picking up the hind feet (P<0.05). The administration of vaccines and paste dewormer and the collection of blood were tolerated by the majority of the foals of both groups with no or low resistance. It appeared that neonatal imprint training resulted in a learned behaviour that resulted in decreased self-defence responses towards handling the limbs at three months of age.
Keywords: Early handling; Imprint train; Neonatal foals; Horse
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Staunton, H. (2005). Mammalian sleep. Naturwissenschaften, 92(5), 203–220.
Abstract: This review examines the biological background to the development of ideas on rapid eye movement sleep (REM sleep), so-called paradoxical sleep (PS), and its relation to dreaming. Aspects of the phenomenon which are discussed include physiological changes and their anatomical location, the effects of total and selective sleep deprivation in the human and animal, and REM sleep behavior disorder, the latter with its clinical manifestations in the human. Although dreaming also occurs in other sleep phases (non-REM or NREM sleep), in the human, there is a contingent relation between REM sleep and dreaming. Thus, REM is taken as a marker for dreaming and as REM is distributed ubiquitously throughout the mammalian class, it is suggested that other mammals also dream. It is suggested that the overall function of REM sleep/dreaming is more important than the content of the individual dream; its function is to place the dreamer protagonist/observer on the topographical world. This has importance for the developing infant who needs to develop a sense of self and separateness from the world which it requires to navigate and from which it is separated for long periods in sleep. Dreaming may also serve to maintain a sense of 'I'ness or “self” in the adult, in whom a fragility of this faculty is revealed in neurological disorders.
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Valderrabano-Ibarra, C., Brumon, I., & Drummond, H. (2007). Development of a linear dominance hierarchy in nestling birds. Anim. Behav., 74(6), 1705–1714.
Abstract: Theoreticians propose that trained winning and losing are important processes in creating linear animal dominance hierarchies, and experiments have shown that both processes can occur in animals, but their actual roles in creating natural hierarchies are unknown. We described agonism in 18 broods of three blue-footed boobies, Sula nebouxii, a species for which trained winning and losing have been demonstrated, to infer how these processes generate and maintain a natural hierarchy. Ranks in the linear hierarchy that emerged in every brood were initially assigned by asymmetries in age, size and maturity, which led to differences between broodmates in levels of expressed and received aggression and, consequently, to differences in the training of their aggressiveness and submissiveness. Later, ranks appeared to be maintained by the chicks' acquired aggressive and submissive tendencies combined with ongoing effects of persisting differences in size and maturity. Our results suggest that trained winning and trained losing are important in the construction of booby hierarchies but that these two axes of learning are largely independent. Increase in submissiveness occurs over a period of about 10-20 days, and the level of submissiveness reached varies with the amount of aggression received. After training, submissiveness is apparently maintained by a lower level of aggression and increasing use of threats. Threats become increasingly effective as chicks age, but are never as effective as attacks.
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Versace, E., Morgante, M., Pulina, G., & Vallortigara, G. (2007). Behavioural lateralization in sheep (Ovis aries). Behav. Brain. Res., 184(1), 72–80.
Abstract: This study investigates behavioural lateralization in sheep and lambs of different ages. A flock was tested in a task in which the animals were facing an obstacle and should avoid it on either the right or left side to rejoin flock-mates (adult sheep) or their mothers (lambs). A bias for avoiding the obstacle on the right side was observed, with lambs apparently being more lateralized than sheep. This right bias was tentatively associated with the left-hemifield laterality in familiar faces recognition which has been documented in this species. Differences between adult sheep and lambs were likely to be due to differences in social reinstatement motivation elicited by different stimuli (flock-mates or mothers) at different ages. Preferential use of the forelegs to step on a wood-board and direction of jaw movement during rumination was also tested in adult animals. No population bias nor individual-level lateralization was observed for use of the forelegs. At the same time, however, there was a large number of animals showing individual-level lateralization for the direction of jaw movement during rumination even though there was no population bias. These findings highlight that within the same species individual- and population-level lateralization can be observed in different tasks. Moreover, the results fit the general hypothesis that population-level asymmetries are more likely to occur in tasks that require social coordination among behaviourally asymmetric individuals.
Keywords: Lateralization; Laterality; Brain asymmetry; Hemisphere; Sheep; Lamb; Strength of lateralization
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