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Feist, J. D., & McCullough, D. R. (1976). Behavior patterns and communication in feral horses. Z. Tierpsychol., 41(4), 337–371.
Abstract: The social behavior of feral horses was studied in the western United States. Stable harem groups with a dominant stallion and bachelor hermaphrodite hermaphrodite groups occupied overlapping home ranges. Groups spacing, but not territoriality, was expressed. Harem group, stability resulted from strong dominance by dominant stallions, and fidelity of group members. Eliminations of group members were usually marked by urine of the dominant stallion. Hermaphrodite-hermaphrodite aggression involved spacing between harems and dominance in bachelor groups. Marking with feces was important in hermaphrodite-hermaphrodite interactions. Foaling occurred in May and early June, following the post-partum estrous. All breeding was done by harem stallions. Young were commonly nursed through yearling age. These horses showed social organizations similar to other feral horses and plains zebras.
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Edwards, D. H., & Spitzer, N. (2006). 6. Social dominance and serotonin receptor genes in crayfish. Curr Top Dev Biol, 74, 177–199.
Abstract: Gene expression affects social behavior only through changes in the excitabilities of neural circuits that govern the release of the relevant motor programs. In turn, social behavior affects gene expression only through patterns of sensory stimulation that produce significant activation of relevant portions of the nervous system. In crayfish, social interactions between pairs of animals lead to changes in behavior that mark the formation of a dominance hierarchy. Those changes in behavior result from changes in the excitability of specific neural circuits. In the new subordinate, circuits for offensive behavior become less excitable and those for defensive behavior become more excitable. Serotonin, which is implicated in mechanisms for social dominance in many animals, modulates circuits for escape and avoidance responses in crayfish. The modulatory effects of serotonin on the escape circuits have been found to change with social dominance, becoming excitatory in dominant crayfish and inhibitory in subordinates. These changes in serotonin's effects on escape affect the synaptic response to sensory input of a single cell, the lateral giant (LG) command neuron for escape. Moreover, these changes occur over a 2-week period and for the subordinate are reversible at any time following a reversal of the animal's status. The results have suggested that a persistent change in social status leads to a gradual change in the expression of serotonin receptors to a pattern that is more appropriate for the new status. To test that hypothesis, the expression patterns of crayfish serotonin receptors must be compared in dominant and subordinate animals. Two of potentially five serotonin receptors in crayfish have been cloned, sequenced, and pharmacologically characterized. Measurements of receptor expression in the whole CNS of dominant and subordinate crayfish have produced inconclusive results, probably because each receptor is widespread in the nervous system and is likely to experience opposite expression changes in different areas of the CNS. Both receptors have recently been found in identified neurons that mediate escape responses, and so the next step will be to measure their expression in these identified cells in dominant and subordinate animals.
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Levy, J. (1977). The mammalian brain and the adaptive advantage of cerebral asymmetry. Ann N Y Acad Sci, 299, 264–272.
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Friederici, A. D., & Alter, K. (2004). Lateralization of auditory language functions: a dynamic dual pathway model. Brain Lang, 89(2), 267–276.
Abstract: Spoken language comprehension requires the coordination of different subprocesses in time. After the initial acoustic analysis the system has to extract segmental information such as phonemes, syntactic elements and lexical-semantic elements as well as suprasegmental information such as accentuation and intonational phrases, i.e., prosody. According to the dynamic dual pathway model of auditory language comprehension syntactic and semantic information are primarily processed in a left hemispheric temporo-frontal pathway including separate circuits for syntactic and semantic information whereas sentence level prosody is processed in a right hemispheric temporo-frontal pathway. The relative lateralization of these functions occurs as a result of stimulus properties and processing demands. The observed interaction between syntactic and prosodic information during auditory sentence comprehension is attributed to dynamic interactions between the two hemispheres.
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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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Seyfarth, R. M., & Cheney, D. L. (2001). Cognitive strategies and the representation of social relations by monkeys. Nebr Symp Motiv, 47, 145–177.
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Hemelrijk, C. K., & Wantia, J. (2005). Individual variation by self-organisation. Neurosci Biobehav Rev, 29(1), 125–136.
Abstract: In this paper, we show that differences in dominance and spatial centrality of individuals in a group may arise through self-organisation. Our instrument is a model, called DomWorld, that represents two traits that are often found in animals, namely grouping and competing. In this model individual differences grow under the following conditions: (1) when the intensity of aggression increases and grouping becomes denser, (2) when the degree of sexual dimorphism in fighting power increases. In this case the differences among females compared to males grow too, (3) when, upon encountering another individual, the tendency to attack is 'obligate' and not conditional, namely 'sensitive to risks'. Results resemble phenomena described for societies of primates, mice, birds and pigs.
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Overli, O., Sorensen, C., Pulman, K. G. T., Pottinger, T. G., Korzan, W., Summers, C. H., et al. (2007). Evolutionary background for stress-coping styles: relationships between physiological, behavioral, and cognitive traits in non-mammalian vertebrates. Neurosci Biobehav Rev, 31(3), 396–412.
Abstract: Reactions to stress vary between individuals, and physiological and behavioral responses tend to be associated in distinct suites of correlated traits, often termed stress-coping styles. In mammals, individuals exhibiting divergent stress-coping styles also appear to exhibit intrinsic differences in cognitive processing. A connection between physiology, behavior, and cognition was also recently demonstrated in strains of rainbow trout (Oncorhynchus mykiss) selected for consistently high or low cortisol responses to stress. The low-responsive (LR) strain display longer retention of a conditioned response, and tend to show proactive behaviors such as enhanced aggression, social dominance, and rapid resumption of feed intake after stress. Differences in brain monoamine neurochemistry have also been reported in these lines. In comparative studies, experiments with the lizard Anolis carolinensis reveal connections between monoaminergic activity in limbic structures, proactive behavior in novel environments, and the establishment of social status via agonistic behavior. Together these observations suggest that within-species diversity of physiological, behavioral and cognitive correlates of stress responsiveness is maintained by natural selection throughout the vertebrate sub-phylum.
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Mazurek, M., McGee, M., Minchin, W., Crowe, M. A., & Earley, B. (2011). Is the avoidance distance test for the assessment of animals' responsiveness to humans influenced by either the dominant or flightiest animal in the group? Appl. Anim. Behav. Sci., 132(3-4), 107–113.
Abstract: A previously described (Windschnurer et al., 2009) avoidance distance test was used to assess animals’ fear of humans in order to quantify the human–animal relationship (HAR). This study investigated the influence of the dominant and flightiest animals within a group on the responsiveness of animals during the avoidance distance test. Eighty-eight pregnant heifers comprised of four different genotypes were used (22 animals per genotype): Limousin × Holstein-Friesian, Limousin × Simmental, Charolais × Limousin, and Charolais × Simmental. Sixty of the 88 heifers were group housed (n = 5) into 12 pens with 3 pens per breed, while 28 heifers were singly housed (seven heifers per breed). A reactivity test was performed on days 10, 18, 25 and 30 post-housing on the singly housed heifers, and then on the group housed heifers, on the same days, to calculate a reactivity score. On days 33 and 37 flight and dominance tests, respectively, were performed to identify the flightiest and the dominant animal within each group. On day 41, an avoidance test, measuring both the avoidance distance towards a familiar and an unfamiliar human, was performed on all heifers. No difference (P > 0.05) in reactivity scores was found between the genotypes, between pens for the group housed heifers or between singly housed and group housed heifers (P = 0.28). The avoidance distance (AD) of singly (S) housed heifers towards a familiar (F) (ADSF) human was shorter (P < 0.001) than the avoidance distance of group (G) housed heifers towards an unfamiliar human (ADSU). The ADSF and ADGF were correlated with the ADSU and ADGU (R = 0.87 for singly housed heifers; R = 0.61 for group housed heifers, P < 0.001). For the singly housed heifers, no correlation was observed between reactivity score and ADSF (R = 0.36, P = 0.18), whereas the reactivity score and ADSU were correlated (R = 0.68, P = 0.004). For the group housed heifers no significant correlation was detected between the reactivity score and ADGF (R = 0.18, P = 0.22) or ADGU (R = −0.11, P = 0.39). No influence of the most dominant animal and the flightiest animals was found on the behaviour of the group in term of avoidance distance and reactivity (P > 0.05). It is concluded that the assessment of the fear of the animals towards humans using the avoidance test at the feed bunk may be useful for singly and group housed heifers and that the leaders of a group such as the flightiest animal or the dominant animal did not influence the avoidance distance test.
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Goursot, C., Düpjan, S., Puppe, B., & Leliveld, L. M. C. (2021). Affective styles and emotional lateralization: A promising framework for animal welfare research. Appl. Anim. Behav. Sci., 237, 105279.
Abstract: The growing recognition of animals as individuals has broader implications for farm animal welfare research. Even under highly standardized on-farm conditions, farm animals show heterogeneous but individually consistent behavioural patterns towards various stimuli, based on how they appraise these stimuli. As a result, animal welfare is likely to be highly individual as well, and studying the proximate mechanisms underlying distinct individual behaviour patterns and appraisal will improve animal welfare research. We propose to extend the framework of affective styles to bridge the gap between existing research fields on animal personality and affective states. Affective styles refer to consistent individual differences in emotional reactivity and regulation and can be predicted by baseline cerebral lateralization. Likewise, animals with consistent left or right motor biases--a proxy measure of individual patterns in cerebral lateralization--have been shown to differ in their personality, emotional reactivity, motivational tendencies or coping styles. In this paper, we present the current knowledge of the links between laterality and stable individual traits in behaviour and affect in light of hypotheses on emotional lateralization. Within our suggested framework, we make recommendations on how to investigate affective styles in non-human animals and give practical examples. This approach has the potential to promote a science of affective styles in nonhuman animals and significantly advance research on animal welfare.
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