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Croney, C. C., & Millman, S. T. (2007). BOARD-INVITED REVIEW: The ethical and behavioral bases for farm animal welfare legislation. J. Anim Sci., 85(2), 556–565.
Abstract: Concerns about farm animal welfare vary among individuals and societies. As people increasingly consider the values underlying current farm animal production methods, farm animal welfare policy debates have escalated. Recent food animal protection policies enacted in the European Union have fueled highly contentious discussions about the need for similar legislative activity in the United States. Policymakers and scientists in the United States are apprehensive about the scientific assessment, validation, and monitoring of animal welfare, as well as the unforeseen consequences of moving too hastily toward legislating farm animal welfare. The potential impact of such legislation on producers, food prices, animals, and concerned citizens must also be considered. Balancing the interests of all stakeholders has therefore presented a considerable challenge that has stymied US policymaking. In this review, we examine the roles of ethics and science in policy decisions, discuss how scientific knowledge relative to animal behavior has been incorporated into animal welfare policy, and identify opportunities for additional refinement of animal welfare science that may facilitate ethical and policy decisions about animal care.
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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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Stomp, M., d'Ingeo, S., Henry, S., Cousillas, H., & Hausberger, M. (2021). Brain activity reflects (chronic) welfare state: Evidence from individual electroencephalography profiles in an animal model. Appl. Anim. Behav. Sci., 236, 105271.
Abstract: Assessing the animal welfare state is a challenge given the subjective individual cognitive and emotional processing involved. Electroencephalography (EEG) spectrum analysis has proved an ecologically valid recording situation to assess the link between brain processes and affective or cognitive states in humans: a higher slow wave/fast wave ratio has been associated with a positive internal state. In particular, a high production of theta power (3-8 Hz) has been related to positive emotions. On the other hand, it has been hypothesized that a left hemisphere (LH) dominance may be associated with a better welfare state. Here, we test the hypothesis that individual differences in the resting-state quantitative EEG power spectrum of adult horses (N = 18) and its lateralization pattern may reflect individual subjective perception of their conditions of life and welfare state. The results show clear individual differences in the proportions of the different waves and their inter-hemispheric distribution. Three different EEG power spectrum profiles were highlighted, from a bilateral predominance of theta waves in horses in a more positive welfare state to a bilateral predominance of beta waves in horses with clear expressions of compromised welfare. Interestingly, particular correlations were found between wave power activity and welfare parameters. We found a negative correlation between the number of stereotypic behaviours per hour and the median proportion of theta waves in the left hemisphere. and between the overall state (total chronic stress score) of welfare and gamma production in the right hemisphere (RH). These findings go along the hypothesis of a particular involvement of the left hemisphere for positive processing and of the right hemisphere for negative processing. However, the pattern of laterality did not appear as the most important feature here as both extreme clusters in terms of welfare showed bilateral predominance of one wave type. It is possible that hemispheric specialization makes more sense during acute emotion-inducing conditions rather than in this resting-state context (i.e. in absence of any high emotion-inducing stimulation), although the opposition gamma versus theta waves between both hemispheres in the horses with an intermediate welfare state is noticeable and intriguing. It seems that bilateral but also LH theta activity is a promising neurophysiological marker of good welfare in horses, while a bilateral or RH high production of gamma waves should alert about potential welfare alterations. Quantitative resting-state EEG power spectrum appears as a highly promising tool for exploring the brain processes involved in the subjective perception of chronic welfare, as a useful complementary tool for welfare assessment.
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Stomp, M., d'Ingeo, S., Henry, S., Cousillas, H., & Hausberger, M. (2021). Brain activity reflects (chronic) welfare state: Evidence from individual electroencephalography profiles in an animal model. Applied Animal Behaviour Science, 236, 105271.
Abstract: Assessing the animal welfare state is a challenge given the subjective individual cognitive and emotional processing involved. Electroencephalography (EEG) spectrum analysis has proved an ecologically valid recording situation to assess the link between brain processes and affective or cognitive states in humans: a higher slow wave/fast wave ratio has been associated with a positive internal state. In particular, a high production of theta power (3-8 Hz) has been related to positive emotions. On the other hand, it has been hypothesized that a left hemisphere (LH) dominance may be associated with a better welfare state. Here, we test the hypothesis that individual differences in the resting-state quantitative EEG power spectrum of adult horses (N = 18) and its lateralization pattern may reflect individual subjective perception of their conditions of life and welfare state. The results show clear individual differences in the proportions of the different waves and their inter-hemispheric distribution. Three different EEG power spectrum profiles were highlighted, from a bilateral predominance of theta waves in horses in a more positive welfare state to a bilateral predominance of beta waves in horses with clear expressions of compromised welfare. Interestingly, particular correlations were found between wave power activity and welfare parameters. We found a negative correlation between the number of stereotypic behaviours per hour and the median proportion of theta waves in the left hemisphere. and between the overall state (total chronic stress score) of welfare and gamma production in the right hemisphere (RH). These findings go along the hypothesis of a particular involvement of the left hemisphere for positive processing and of the right hemisphere for negative processing. However, the pattern of laterality did not appear as the most important feature here as both extreme clusters in terms of welfare showed bilateral predominance of one wave type. It is possible that hemispheric specialization makes more sense during acute emotion-inducing conditions rather than in this resting-state context (i.e. in absence of any high emotion-inducing stimulation), although the opposition gamma versus theta waves between both hemispheres in the horses with an intermediate welfare state is noticeable and intriguing. It seems that bilateral but also LH theta activity is a promising neurophysiological marker of good welfare in horses, while a bilateral or RH high production of gamma waves should alert about potential welfare alterations. Quantitative resting-state EEG power spectrum appears as a highly promising tool for exploring the brain processes involved in the subjective perception of chronic welfare, as a useful complementary tool for welfare assessment.
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Vallortigara, G., Chiandetti, C., & Sovrano, V. A. (2011). Brain asymmetry (animal). WIREs Cogn Sci, 2(2), 146–157.
Abstract: Once considered a uniquely human attribute, brain asymmetry has been proved to be ubiquitous among non-human animals. A synthetic review of evidence of animal lateralization in the motor, sensory, cognitive, and affective domains is provided, together with a discussion of its development and possible biological functions. It is argued that investigation of brain asymmetry in a comparative perspective may favor the link between classical neuropsychological studies and modern developmental and evolutionary biology approaches. WIREs Cogni Sci 2011 2 146–157 DOI: 10.1002/wcs.100 For further resources related to this article, please visit the WIREs website
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Neveu, P. J. (1993). Brain Lateralization and Immunomodulation. Int J Neurosci, 70(1-2), 135–143.
Abstract: The two sides of the brain may be differently involved in the modulation of immune responses as demonstrated by lesional and behavioral approaches in rodents. Lesions of right or left neocortex induced opposite effects on various immune parameters including mitogen-induced lymphoproliferation, interleukin-2 production, macrophage activation or natural killer cell activity. This animal model, useful to elucidate whereby the brain and the immune system can communicate, appears to be suitable for studying the immune perturbations observed during stroke in humans. Brain asymmetry in modulation of immune reactivity may also be demonstrated in intact animal using a behavioral paradigm. The direction of a lateralized motor behavior ie paw preference in a food reaching task, correlated with an asymmetrical brain organization, was shown to be associated with lymphocyte reactivity, natural killer cell activity and auto-antibody production. The association between paw preference and immune reactivity in mice varies according to the immune parameters tested and is a sex-dependent phenomenon in which genetic background may be involved. The experimental models for investigating asymmetrical brain modulation of the immune system should be useful for studying several physiological, pathological and genetic aspects of neuroimmunomodulation.
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Harvey, P. H., Clutton-Brock, T. H., & Mace, G. M. (1980). Brain size and ecology in small mammals and primates. PNAS, 77(7), 4387–4389.
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Weisbecker, V., & Goswami, A. (2010). Brain size, life history, and metabolism at the marsupial/placental dichotomy. Proc. Natl. Acad. Sci. U.S.A., 107(37), 16216–16221.
Abstract: The evolution of mammalian brain size is directly linked with the evolution of the brain's unique structure and performance. Both maternal life history investment traits and basal metabolic rate (BMR) correlate with relative brain size, but current hypotheses regarding the details of these relationships are based largely on placental mammals. Using encephalization quotients, partial correlation analyses, and bivariate regressions relating brain size to maternal investment times and BMR, we provide a direct quantitative comparison of brain size evolution in marsupials and placentals, whose reproduction and metabolism differ extensively. Our results show that the misconception that marsupials are systematically smaller-brained than placentals is driven by the inclusion of one large-brained placental clade, Primates. Marsupial and placental brain size partial correlations differ in that marsupials lack a partial correlation of BMR with brain size. This contradicts hypotheses stating that the maintenance of relatively larger brains requires higher BMRs. We suggest that a positive BMR–brain size correlation is a placental trait related to the intimate physiological contact between mother and offspring during gestation. Marsupials instead achieve brain sizes comparable to placentals through extended lactation. Comparison with avian brain evolution suggests that placental brain size should be constrained due to placentals’ relative precociality, as has been hypothesized for precocial bird hatchlings. We propose that placentals circumvent this constraint because of their focus on gestation, as opposed to the marsupial emphasis on lactation. Marsupials represent a less constrained condition, demonstrating that hypotheses regarding placental brain size evolution cannot be generalized to all mammals.
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Finarelli, J. A., & Flynn, J. J. (2009). Brain-size evolution and sociality in Carnivora. Proc. Natl. Acad. Sci. U.S.A., 106(23), 9345–9349.
Abstract: Increased encephalization, or larger brain volume relative to body mass, is a repeated theme in vertebrate evolution. Here we present an extensive sampling of relative brain sizes in fossil and extant taxa in the mammalian order Carnivora (cats, dogs, bears, weasels, and their relatives). By using Akaike Information Criterion model selection and endocranial volume and body mass data for 289 species (including 125 fossil taxa), we document clade-specific evolutionary transformations in encephalization allometries. These evolutionary transformations include multiple independent encephalization increases and decreases in addition to a remarkably static basal Carnivora allometry that characterizes much of the suborder Feliformia and some taxa in the suborder Caniformia across much of their evolutionary history, emphasizing that complex processes shaped the modern distribution of encephalization across Carnivora. This analysis also permits critical evaluation of the social brain hypothesis (SBH), which predicts a close association between sociality and increased encephalization. Previous analyses based on living species alone appeared to support the SBH with respect to Carnivora, but those results are entirely dependent on data from modern Canidae (dogs). Incorporation of fossil data further reveals that no association exists between sociality and encephalization across Carnivora and that support for sociality as a causal agent of encephalization increase disappears for this clade.
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Dugatkin, L. A. (1998). Breaking up fights between others: a model of intervention behaviour. Proc. R. Soc. Lond. B, 265(1394), 433–437.
Abstract: To examine when and why animals break up fights between others in their group, I modelled whether ‘winner’ and ‘loser’ effects might be one element driving the evolution of intervention behaviour. I considered one particular type of intervention: when the intervener simply breaks up fights between two others, but does not favour either party in so doing. When victories at time T + 1 are more likely given a victory at time T (i.e. winner effects), intervention is often favoured. Intervention is favoured in these circumstances because the intervening party in essence stops others from ‘getting on a roll’ and climbing up any hierarchy that exists. However, when loser effects alone are at work (defeats at time T + 1 are more likely given a defeat at time T), breaking up fights between others is never selected. If both winner and loser effects are operating simultaneously, then the likelihood of intervention behaviour evolving is a function of the relative strength of these two effects. The greater the winner effect relative to the loser effect, the more likely intervention behaviour is to evolve.
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