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Hagen, S. J., & Eaton, W. A. (2000). Two-state expansion and collapse of a polypeptide. J Mol Biol, 301(4), 1019–1027.
Abstract: The initial phase of folding for many proteins is presumed to be the collapse of the polypeptide chain from expanded to compact, but still denatured, conformations. Theory and simulations suggest that this collapse may be a two-state transition, characterized by barrier-crossing kinetics, while the collapse of homopolymers is continuous and multi-phasic. We have used a laser temperature-jump with fluorescence spectroscopy to measure the complete time-course of the collapse of denatured cytochrome c with nanosecond time resolution. We find the process to be exponential in time and thermally activated, with an apparent activation energy approximately 9 k(B)T (after correction for solvent viscosity). These results indicate that polypeptide collapse is kinetically a two-state transition. Because of the observed free energy barrier, the time scale of polypeptide collapse is dramatically slower than is predicted by Langevin models for homopolymer collapse.
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Quaresmini, C., Forrester, G. S., Spiezio, C., & Vallortigara, G. (2014). Social environment elicits lateralized behaviors in gorillas (Gorilla gorilla gorilla) and chimpanzees (Pan troglodytes). Journal of Comparative Psychology, 128(3), 276–284.
Abstract: The influence of the social environment on lateralized behaviors has now been investigated across a wide variety of animal species. New evidence suggests that the social environment can modulate behavior. Currently, there is a paucity of data relating to how primates navigate their environmental space, and investigations that consider the naturalistic context of the individual are few and fragmented. Moreover, there are competing theories about whether only the right or rather both cerebral hemispheres are involved in the processing of social stimuli, especially in emotion processing. Here we provide the first report of lateralized social behaviors elicited by great apes. We employed a continuous focal animal sampling method to record the spontaneous interactions of a captive zoo-living colony of chimpanzees (Pan troglodytes) and a biological family group of peer-reared western lowland gorillas (Gorilla gorilla gorilla). We specifically focused on which side of the body (i.e., front, rear, left, right) the focal individual preferred to keep conspecifics. Utilizing a newly developed quantitative corpus-coding scheme, analysis revealed both chimpanzees and gorillas demonstrated a significant group-level preference for focal individuals to keep conspecifics positioned to the front of them compared with behind them. More interestingly, both groups also manifested a population-level bias to keep conspecifics on their left side compared with their right side. Our findings suggest a social processing dominance of the right hemisphere for context-specific social environments. Results are discussed in light of the evolutionary adaptive value of social stimulus as a triggering factor for the manifestation of group-level lateralized behaviors. (PsycINFO Database Record (c) 2016 APA, all rights reserved)
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Pawluski, J., Jego, P., Henry, S., Bruchet, A., Palme, R., Coste, C., et al. (2017). Low plasma cortisol and fecal cortisol metabolite measures as indicators of compromised welfare in domestic horses (Equus caballus). Plos One, 12(9), e0182257.
Abstract: The hypothalamic-pituitary-adrenal (HPA) axis response to chronic stress is far from straight forward, particularly with regards to animal welfare. There are reports of no effect as well as both decreases and increases in cortisol after chronic stressors. Therefore, the first aim of the present study was to determine how measures of compromised welfare, such as chronic pain and haematological anomalies, related to cortisol levels in domestic horses (Equus caballus). Domestic horses are an informative model to investigate the impact of chronic stress (due to environment, pain, work, housing conditions...) on the HPA axis. The second aim was to determine whether levels of fecal cortisol metabolites (FCM) may be used as an indicator of welfare measures. The present study used fifty-nine horses (44 geldings and 15 mares), from three riding centres in Brittany, France. The primary findings show that horses whose welfare was clearly compromised (as indicated by an unusual ears backward position, presence of vertebral problems or haematological anomalies, e.g. anaemia) also had lower levels of both FCM and plasma cortisol. This work extends our previous findings showing that withdrawn postures, indicators of depressive-like behavior in horses, are associated with lower plasma cortisol levels. We also found that evening plasma cortisol levels positively correlated with FCM levels in horses. Future research aims to determine the extent to which factors of influence on welfare, such as living conditions (e.g. single stalls versus group housing in pasture or paddocks), early life factors, and human interaction, act as mediators of cortisol levels in horses.
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Cozzi, B., Povinelli, M., Ballarin, C., & Granato, A. (2014). The Brain of the Horse: Weight and Cephalization Quotients. Brain Behav Evol, 83(1), 9–16.
Abstract: The horse is a common domestic animal whose anatomy has been studied since the XVI century. However, a modern neuroanatomy of this species does not exist and most of the data utilized in textbooks and reviews derive from single specimens or relatively old literature. Here, we report information on the brain of Equus caballus obtained by sampling 131 horses, including brain weight (as a whole and subdivided into its constituents), encephalization quotient (EQ), and cerebellar quotient (CQ), and comparisons with what is known about other relevant species. The mean weight of the fresh brains in our experimental series was 598.63 g (SEM ± 7.65), with a mean body weight of 514.12 kg (SEM ± 15.42). The EQ was 0.78 and the CQ was 0.841. The data we obtained indicate that the horse possesses a large, convoluted brain, with a weight similar to that of other hoofed species of like mass. However, the shape of the brain, the noteworthy folding of the neocortex, and the peculiar longitudinal distribution of the gyri suggest an evolutionary specificity at least partially separate from that of the Cetartiodactyla (even-toed mammals and cetaceans) with whom Perissodactyla (odd-toed mammals) are often grouped.
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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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Bergmüller, R., & Taborsky, M. (2010). Animal personality due to social niche specialisation. Trends in Ecology & Evolution, 25(9), 504–511.
Abstract: The existence of 'animal personality', i.e. consistent individual differences in behaviour across time and contexts, is an evolutionary puzzle that has recently generated considerable research interest. Although social factors are generally considered to be important, it is as yet unclear how they might select for personality. Drawing from ecological niche theory, we explore how social conflict and alternative social options can be key factors in the evolution and development of consistent individual differences in behaviour. We discuss how animal personality research might benefit from insights into the study of alternative tactics and illustrate how selection can favour behavioural diversification and consistency due to fitness benefits resulting from conflict reduction among social partners.
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Frère, C. H., Krützen, M., Mann, J., Connor, R. C., Bejder, L., & Sherwin, W. B. (2010). Social and genetic interactions drive fitness variation in a free-living dolphin population. Proc. Natl. Acad. Sci. U.S.A., 107(46), 19949–19954.
Abstract: The evolutionary forces that drive fitness variation in species are of considerable interest. Despite this, the relative importance and interactions of genetic and social factors involved in the evolution of fitness traits in wild mammalian populations are largely unknown. To date, a few studies have demonstrated that fitness might be influenced by either social factors or genes in natural populations, but none have explored how the combined effect of social and genetic parameters might interact to influence fitness. Drawing from a long-term study of wild bottlenose dolphins in the eastern gulf of Shark Bay, Western Australia, we present a unique approach to understanding these interactions. Our study shows that female calving success depends on both genetic inheritance and social bonds. Moreover, we demonstrate that interactions between social and genetic factors also influence female fitness. Therefore, our study represents a major methodological advance, and provides critical insights into the interplay of genetic and social parameters of fitness.
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Dalla Costa, E., Dai, F., Lebelt, D., Scholz, P., Barbieri, S., Canali, E., et al. (2016). Welfare assessment of horses: the AWIN approach. Anim. Welf., 25(4), 481–488.
Abstract: The EU-funded Animal Welfare Indicators (AWIN) research project (2011-2015) aimed to improve animal welfare through the development of practical on-farm animal welfare assessment protocols. The present study describes the application of the AWIN approach to the development of a welfare assessment protocol for horses (Equus caballus). Its development required the following steps: (i) selection of potential welfare indicators; (ii) bridging gaps in knowledge; (iii) consulting stakeholders; and (iv) testing a prototype protocol on-farm. Compared to existing welfare assessment protocols for other species, the AWIN welfare assessment protocol for horses introduces a number of innovative aspects, such as implementation of a two-level strategy focused on improving on-farm feasibility and the use of electronic tools to achieve standardised data collection and so promote rapid outcomes. Further refinement to the AWIN welfare assessment protocol for horses is needed in order to firstly gather data from a larger reference population and, secondly, enhance the welfare assessment protocol with reference to different horse housing and husbandry conditions.
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Gaunitz, C., Fages, A., Hanghøj, K., Albrechtsen, A., Khan, N., Schubert, M., et al. (2018). Ancient genomes revisit the ancestry of domestic and Przewalski's horses. Science, 360(6384), 111–114.
Abstract: The Eneolithic Botai culture of the Central Asian steppes provides the earliest archaeological evidence for horse husbandry, ~5,500 ya, but the exact nature of early horse domestication remains controversial. We generated 42 ancient horse genomes, including 20 from Botai. Compared to 46 published ancient and modern horse genomes, our data indicate that Przewalski's horses are the feral descendants of horses herded at Botai and not truly wild horses. All domestic horses dated from ~4,000 ya to present only show ~2.7% of Botai-related ancestry. This indicates that a massive genomic turnover underpins the expansion of the horse stock that gave rise to modern domesticates, which coincides with large-scale human population expansions during the Early Bronze Age.
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Christensen, J. W., Beekmans, M., van Dalum, M., & VanDierendonck, M. (2014). Effects of hyperflexion on acute stress responses in ridden dressage horses. Physiol. Behav., 128, 39–45.
Abstract: The effects of hyperflexion on the welfare of dressage horses have been debated. This study aimed to investigate acute stress responses of dressage horses ridden in three different Head-and-Neck-positions (HNPs). Fifteen dressage horses were ridden by their usual rider in a standardised 10-min dressage programme in either the competition frame (CF), hyperflexion (“Low-Deep-and-Round”; LDR) or a looser frame (LF) in a balanced order on three separate test days. Heart rate (HR), heart rate variability parameters (HRV), behaviour and rein tension were recorded during the test. Salivary cortisol concentrations were measured 60min before and 0, 5, 15 and 30min after the test. Rein tension was significantly lower in LF and did not differ between CF and LDR; however approx. 15% of recordings in CF and LDR were above the sensor detection limit of 5kg. The horses had significantly higher cortisol concentrations directly after LDR compared to LF. In addition, the horses showed more distinctive head movements, including head waving, during LDR. There were no significant treatment effects on HR and HRV. In conclusion, the results indicate that LDR may be more stressful to these horses during riding.
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