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Laut, J. E., Houpt, K. A., Hintz, H. F., & Houpt, T. R. (1985). The effects of caloric dilution on meal patterns and food intake of ponies. Physiol. Behav., 35(4), 549–554.
Abstract: In order to determine if horses will increase their intake in response to caloric dilution, four pony geldings were fed ad lib a mixed grain diet either undiluted (3.4 Mcal/kg of digestible energy) or diluted (wt/wt) with 25% sawdust (2.6 Mcal/kg) or with 50% sawdust (1.7 Mcal/kg). The mean daily caloric intake was 17,457 kcal (3.4 Mcal diet), 17,546 kcal (2.6 Mcal diet) and 12,844 kcal (1.7 Mcal). The mean time spent eating was 246 (3.4 Mcal), 351 (2.6 Mcal), and 408 (1.7 Mcal) minutes/day. Meal size increased and meal frequency decreased with increasing dilution. The median long survivorships of intermeal intervals were 6.4 min (3.4 Mcal), 3.95 min (2.6 Mcal) and 4.91 min (1.7 Mcal). Ponies responded to caloric dilution by increasing the volume of intake to maintain caloric intake when the diet had 25% diluent. When the diet was diluted by 50%, intake was increased, but not at a rate adequate to maintain caloric intake. However, the ponies were able to maintain body weight.
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Visser, E. K., van Reenen, C. G., van der Werf, J. T. N., Schilder, M. B. H., Knaap, J. H., Barneveld, A., et al. (2002). Heart rate and heart rate variability during a novel object test and a handling test in young horses. Physiol. Behav., 76(2), 289–296.
Abstract: Forty-one Dutch Warmblood immature horses were used in a study to quantify temperamental traits on the basis of heart rate (HR) and heart rate variability (HRV) measures. Half of the horses received additional training from the age of 5 months onwards; the other half did not. Horses were tested at 9, 10, 21 and 22 months of age in a novel object and a handling test. During the tests, mean HR and two heart variability indices, e.g. standard deviation of beat-to-beat intervals (SDRR) and root mean square of successive beat-to-beat differences (rMSSD), were calculated and expressed as response values to baseline measures. In both tests, horses showed at all ages a significant increase in mean HR and decrease in HRV measures, which suggests a marked shift of the balance of the autonomic nervous system towards a sympathetic dominance. In the novel object test, this shift was more pronounced in horses that had not been trained. Furthermore, statistical analysis showed that the increase in mean HR could not be entirely explained by the physical activity. The additional increase in HR, the nonmotor HR, was more pronounced in the untrained horses compared to the trained. Hence, it is suggested that this nonmotor HR might be due to the level of emotionality. HR variables showed consistency between years, as well as within the second year. These tests bring about a HR response in horses, part of which may indicate a higher level of emotionality; and horses show individual consistency of these HR variables over ages. Therefore, it is concluded that mean HR and HRV measures used with these tests quantify certain aspects of a horse's temperament.
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Griffiths, D. P., & Clayton, N. S. (2001). Testing episodic memory in animals: A new approach. Physiol. Behav., 73(5), 755–762.
Abstract: Episodic memory involves the encoding and storage of memories concerned with unique personal experiences and their subsequent recall, and it has long been the subject of intensive investigation in humans. According to Tulving's classical definition, episodic memory “receives and stores information about temporally dated episodes or events and temporal-spatial relations among these events.” Thus, episodic memory provides information about the `what' and `when' of events (`temporally dated experiences') and about `where' they happened (`temporal-spatial relations'). The storage and subsequent recall of this episodic information was thought to be beyond the memory capabilities of nonhuman animals. Although there are many laboratory procedures for investigating memory for discrete past episodes, until recently there were no previous studies that fully satisfied the criteria of Tulving's definition: they can all be explained in much simpler terms than episodic memory. However, current studies of memory for cache sites in food-storing jays provide an ethologically valid model for testing episodic-like memory in animals, thereby bridging the gap between human and animal studies memory. There is now a pressing need to adapt these experimental tests of episodic memory for other animals. Given the potential power of transgenic and knock-out procedures for investigating the genetic and molecular bases of learning and memory in laboratory rodents, not to mention the wealth of knowledge about the neuroanatomy and neurophysiology of the rodent hippocampus (a brain area heavily implicated in episodic memory), an obvious next step is to develop a rodent model of episodic-like memory based on the food-storing bird paradigm. The development of a rodent model system could make an important contribution to our understanding of the neural, molecular, and behavioral mechanisms of mammalian episodic memory.
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Penn, D., & Potts, W. K. (1998). Untrained mice discriminate MHC-determined odors. Physiol. Behav., 64(3), 235–243.
Abstract: PENN, D. AND W. K. POTTS. Untrained mice distinguish MHC-determined odors. PHYSIOL BEHAV 64(3) 235-243, 1998.--Immune recognition occurs when foreign antigens are presented to T-lymphocytes by molecules encoded by the highly polymorphic genes of the major histocompatibility complex (MHC). House mice (Mus musculus) prefer to mate with individuals that have dissimilar MHC genes. Numerous studies indicate that mice recognize MHC identity through chemosensory cues; however, it is unclear whether odor is determined by classical, antigen-presenting MHC loci or closely linked genes. Previous studies have relied on training laboratory mice and rats to distinguish MHC-associated odors, but there are several reasons why training experiments may be inappropriate assays for testing if MHC genes affect odor. The aim of this study was to determine whether classical MHC genes affect individual odors and whether wild-derived mice can detect MHC-associated odors without training. In the first experiment, we found that wild-derived mice can be trained in a Y-maze to detect the odors of mice that differ genetically only in the MHC region. In the second and third experiments, we used a naturalistic habituation assay and found that wild-derived mice can, without training, distinguish the odors of mice that differ genetically only at one classical MHC locus (dm2 mutants).
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Mormède, P., Andanson, S., Aupérin, B., Beerda, B., Guémené, D., Malmkvist, J., et al. (2007). Exploration of the hypothalamic-pituitary-adrenal function as a tool to evaluate animal welfare. Physiol. Behav., 92(3), 317–339.
Abstract: Measuring HPA axis activity is the standard approach to the study of stress and welfare in farm animals. Although the reference technique is the use of blood plasma to measure glucocorticoid hormones (cortisol or corticosterone), several alternative methods such as the measurement of corticosteroids in saliva, urine or faeces have been developed to overcome the stress induced by blood sampling itself. In chronic stress situations, as is frequently the case in studies about farm animal welfare, hormonal secretions are usually unchanged but dynamic testing allows the demonstration of functional changes at several levels of the system, including the sensitization of the adrenal cortex to ACTH and the resistance of the axis to feedback inhibition by corticosteroids (dexamethasone suppression test). Beyond these procedural aspects, the main pitfall in the use of HPA axis activity is in the interpretation of experimental data. The large variability of the system has to be taken into consideration, since corticosteroid hormone secretion is usually pulsatile, follows diurnal and seasonal rhythms, is influenced by feed intake and environmental factors such as temperature and humidity, age and physiological state, just to cite the main sources of variation. The corresponding changes reflect the important role of glucocorticoid hormones in a number of basic physiological processes such as energy metabolism and central nervous system functioning. Furthermore, large differences have been found across species, breeds and individuals, which reflect the contribution of genetic factors and environmental influences, especially during development, in HPA axis functioning. Usually, these results will be integrated with data from behavioral observation, production and pathology records in a comprehensive approach of farm animal welfare.
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Forkman, B., Boissy, A., Meunier-Salaün, M. - C., Canali, E., & Jones, R. B. (2007). A critical review of fear tests used on cattle, pigs, sheep, poultry and horses. Physiol. Behav., 92(3), 340–374.
Abstract: FORKMAN, B., A., BOISSY, M.-C., SALAUN, E., CANALI, AND R.B., JONES. A critical review of fear tests used on cattle, pigs, sheep, poultry and horses. PHYSIOL. BEHAV. 000-000, 2007. Fear is arguably the most commonly investigated emotion in domestic animals. In the current review we attempt to establish the level of repeatability and validity found for fear tests used on cattle, pigs, sheep and goats, poultry and horses. We focus the review on the three most common types of fear tests: the arena test (open field), the novel object test, and the restraint test. For some tests, e.g. tonic immobility in poultry, there is a good and broad literature on factors that affect the outcome of the test, the validity of the test and its age dependency. However, there are comparatively few of these well defined and validated tests and what is especially missing for most tests is information on the robustness, i.e., what aspects can be changed without affecting the validity of the tests. The relative absence of standardized tests hampers the development of applied ethology as a science.
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König von Borstel, U., Euent, S., Graf, P., König, S., & Gauly, M. (2011). Equine behaviour and heart rate in temperament tests with or without rider or handler. Physiol. Behav., 104(3), 454–463.
Abstract: The aim of the present study was to compare horses' heart rate (HR), heart rate variability (RMSSD, pNN50) and behaviour in the same temperament test when being ridden, led, and released free. Behavioural measurements included scores and linear measurements for reactivity (R), activity (A), time to calm down (T) and emotionality (E), recorded during the approach (1) and/or during confrontation with the stimulus (2). Sixty-five horses were each confronted 3 times (1 ridden, 1 led, 1 free running in balanced order) with 3 novel and/or sudden stimuli. Mixed model analysis indicated that leading resulted in the lowest (P < 0.05 throughout) reactions as measured by A1, A2, E1, E2, R2, and pNN50 while riding produced the strongest (A1, T2, HR, RMSSD, pNN50) or medium (E1, E2, R2) reactions. Free running resulted either in the strongest (A2, E1, E2, R2) or in the lowest (A1, T2, HR, RMSSD, pNN50) reactions. The repeatability across tests for HR (0.57), but not for RMSSD (0.23) or pNN50 (0.25) was higher than for any behavioural measurement: the latter ranged from values below 0.10 (A1, A2, T2) to values between 0.30 and 0.45 (E1, E2, R2). Overall, the results show that a rider or handler influences, but not completely masks, the horses' intrinsic behaviour in a temperament test, and this influence appeared to be stronger on behavioural variables and heart rate variability than on the horses' heart rates. Taking both practical considerations and repeatabilities into account, reactivity appears to be the most valuable parameter. Emotionality and heart rate can also yield valid results reflecting additional dimensions of temperament although their practical relevance may be less obvious. If a combination of observed variables is chosen with care, a valid assessment of a horse's temperament may be possible in all types of tests. However, in practice, tests that resemble the practical circumstances most closely, i.e. testing riding horses under a rider, should be chosen.
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Sankey, C., Henry, S., Clouard, C., Richard-Yris, M. - A., & Hausberger, M. (2011). Asymmetry of behavioral responses to a human approach in young naive vs. trained horses. Physiology & Behavior, 104(3), 464–468.
Abstract: The aim of this study was to investigate the impact of training experience on young horses (Equus caballus)’ lateralized responses to an approaching human. The results show that the one year old untrained horses display asymmetrical responses to an approaching human, with more negative reactions (escapes, threats) when approached from the left side, while approaches towards the right shoulder elicited more positive behaviors. On the contrary, two years old trained horses reacted equally positively to approaches and contact on both sides. Our findings support those of previous studies investigating a link between emotionality and laterality and confirm the role of the left hemisphere in the processing of novel or negative stimuli. Moreover, the data underline the impact work and training can have on this laterality in horses.
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Henry, S., Zanella, A. J., Sankey, C., Richard-Yris, M. - A., Marko, A., & Hausberger, M. (2012). Adults may be used to alleviate weaning stress in domestic foals (Equus caballus). Physiology & Behavior, 106(4), 428–438.
Abstract: The present study aims to investigate whether the presence of unrelated adult horses at weaning would reduce the social stress of weaning and the emergence of undesirable behaviours. We tested this hypothesis in 32 domestic foals by comparing short and medium term behavioural and physiological responses to weaning in foals maintained in homogeneous groups of peers (PW) to those of foals grouped with both peers and unrelated adults (AW). In total, three trials were conducted, which each trial consisting of one AW group and one PW group. In all foals, weaning was followed by increased vocalization, increased locomotion and increased salivary cortisol concentration. However, signs of stress were less pronounced and shorter in duration in weanlings housed with unrelated adults (e.g. whinnies: p < 0.05; salivary cortisol: p < 0.05). Only foals without adults exhibited increased aggressiveness towards peers (p < 0.05) and abnormal behaviours (p < 0.05) such as excessive wood-chewing and redirected sucking towards peers. In conclusion, introducing adults to minimize weaning stress in foals and later on aggressiveness and abnormal behaviours appears as the most promising approach to date.
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Wolter, R., Pantel, N., Stefanski, V., Möstl, E., & Krueger, K. (2014). The role of an alpha animal in changing environmental conditions. Physiol. Behav., 133, 236–243.
Abstract: Abstract The maintenance and development of conservation areas by grazing of large herbivores, such as Przewalski's horses, is common practice. Several nature conservation areas house male bachelor groups of this species. When males are needed for breeding they are removed from the groups, often without considering group compositions and individual social positions. However, alpha animals are needed for ensuring group stability and decision making in potentially dangerous situations in several species. To investigate the role of the alpha male in a bachelor group, we observed the behaviour of five Przewalski's horse males during the enlargement of their enclosure. We analyzed the group's social structure and movement orders, as well as the animals' connectedness, activity budgets, and whether they moved with preferred group members and how factors such as social rank influenced the horses' behaviour. We also investigated the excretion of glucocorticoid metabolites (GCM) via faeces of the horses while exploring a new area as a parameter of glucocorticoid production. Our results show that the alpha male is important for a bachelor group in changing environmental conditions. The alpha male had the highest level of connectedness within the group. When exploring the new environment, its position in the group changed from previously being the last to being the first. Furthermore the whole group behaviour changed when exploring the new area. The stallions showed reduced resting behavior, increased feeding and did not stay close to each other. We found that the excretion of glucocorticoid metabolites of most horses rose only marginally during the first days on the new area while only the alpha male showed a significant increased amount of glucocorticoid production during the first day of the enclosure enlargement.
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