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Leliveld, L. M. C., Düpjan, S., Tuchscherer, A., & Puppe, B. (2016). Behavioural and physiological measures indicate subtle variations in the emotional valence of young pigs. Physiol. Behav., 157, 116–124.
Abstract: Abstract In the study of animal emotions, emotional valence has been found to be difficult to measure. Many studies of farm animals' emotions have therefore focussed on the identification of indicators of strong, mainly negative, emotions. However, subtle variations in emotional valence, such as those caused by rather moderate differences in husbandry conditions, may also affect animals' mood and welfare when such variations occur consistently. In this study, we investigated whether repeated moderate aversive or rewarding events could lead to measurable differences in emotional valence in young, weaned pigs. We conditioned 105 female pigs in a test arena to either a repeated startling procedure (sudden noises or appearances of objects) or a repeated rewarding procedure (applesauce, toy and straw) over 11 sessions. Control pigs were also regularly exposed to the same test arena but without conditioning. Before and after conditioning, we measured heart rate and its variability as well as the behavioural reactions of the subjects in the test arena, with a special focus on detailed acoustic analyses of their vocalisations. The behavioural and heart rate measures were analysed as changes compared to the baseline values before conditioning. A limited number of the putative indicators of emotional valence were affected by the conditioning. We found that the negatively conditioned pigs showed changes that were significantly different from those in control pigs, namely a decrease in locomotion and an increase in standing. The positively conditioned pigs, however, showed a stronger increase in heart rate and a smaller decrease in SDNN (a heart rate variability parameter indicating changes in autonomic regulation) compared to the controls. Compared to the negatively conditioned pigs, the positively conditioned pigs produced fewer vocalisations overall as well as fewer low-frequency grunts but more high-frequency grunts. The low-frequency grunts of the negatively conditioned pigs also showed lower frequency parameters (bandwidth, maximum frequency, 25% and 50% quartiles) compared to those of the positively conditioned pigs. In any of the statistically significant results, the conditioning accounted for 1.5–11.9% of variability in the outcome variable. Hence, we conclude that repeated moderate aversive and rewarding events have weak but measurable effects on some aspects of behaviour and physiology in young pigs, possibly indicating changes in emotional valence, which could ultimately affect their welfare. The combination of ethophysiological indicators, i.e., the concurrent examination of heart rate measures, behavioural responses and especially vocalisation patterns, as used in the current study, might be a useful way of examining subtle effects on emotional valence in further studies.
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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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Palme, R. (2019). Non-invasive measurement of glucocorticoids: Advances and problems. Physiol. Behav., 199, 229–243.
Abstract: Glucocorticoids (GCs; i.e. cortisol/corticosterone) are a central component of the stress response and thus their measurement is frequently used to evaluate the impact of stressful situations. Their metabolites from faeces of various animal species are more and more taken as a non-invasive aid to assess GC release and thus adrenocortical activity. The current literature review includes an extensive collection (1327 papers) and evaluation (see also Supplementary Tables) of the literature on faecal cortisol/corticosterone metabolite (FCM) analysis published to date. It aims at giving reference for researchers interested in implementing FCM analysis into their study or seeking to improve such methods by providing background knowledge on GC metabolism and excretion, conveying insights into methodological issues and stating caveats of FCM analysis and by highlighting prerequisites for and some examples of a successful application of such methods. Collecting faecal samples and analysing FCMs may appear simple and straightforward, but researchers have to select and apply methods correctly. They also need to be aware of the many pitfalls and potentially confounding factors and, last but not least, have to carefully interpret results. Applied properly, measurement of FCMs is a powerful non-invasive tool in a variety of research areas, such as (stress) biology, ethology, ecology, animal conservation and welfare, but also biomedicine.
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Yarnell, K., Hall, C., & Billett, E. (2013). An assessment of the aversive nature of an animal management procedure (clipping) using behavioral and physiological measures. Physiol. Behav., 118, 32–39.
Abstract: Animal management often involves procedures that, while unlikely to cause physical pain, still cause aversive responses. The domestic horse (Equus caballus) regularly has excessive hair clipped off to facilitate its use as a riding/driving animal and this procedure causes adverse behavioral responses in some animals. The aim of this study was to compare behavioral and physiological measures to assess the aversive effect of this procedure. Ten horses were selected on the basis of being either compliant (C: n=5) or non-compliant (NC: n=5) during this procedure. The horses were subjected to a sham clipping procedure (SC: where the blades had been removed from the clippers) for a period of ten minutes. Measures were taken pre, during and post SC (-10min to +30min) and mean values calculated for ALL horses and for C and NC separately. Behavioral activity was scored (scale 1-5) by twenty students from video footage in (phase/group-blind scoring). Heart rate (HR), salivary cortisol and eye temperature were monitored throughout the procedure. The NC horses were found to be significantly more behaviorally active/less relaxed throughout the trial than C horses (p<0.05) with the greatest difference occurring during the SC procedure (p<0.01). NC horses were more active/less relaxed during, compared with pre or post SC (p<0.05), but showed no behavioral difference pre and post SC. HR of the NC horses was higher than that of the C horses throughout the trial but only significantly so after 10min of SC (p<0.01). ALL horses showed a significant increase in HR between +5 and +10min into the procedure (p<0.05). There was a significant increase in salivary cortisol concentration in ALL horses post procedure (p<0.01) with levels peaking at 20minute post SC. No significant differences in salivary cortisol concentration between C and NC were found at any stage of the trial. Eye temperature increased significantly in ALL horses during SC, peaking at +10min into the procedure (p<0.05) and then decreased substantially when SC had ceased (p<0.01). Although no significant differences were found between C and NC per se, there was a significant interaction between group and phase of trial (p<0.05) with the NC group showing a greater decrease in eye temperature post SC. There was a significant positive correlation between changes in salivary cortisol concentration and eye temperature (p<0.01) but no correlation between any of the other measures. Although the behavioral response of C and NC to this procedure was significantly different the physiological responses indicated that ALL horses found the procedure aversive. Eye temperature could be used as an objective and immediate measure of how an animal is responding to a specific situation in order to evaluate management procedures and adapt them where appropriate to reduce the negative impact on animal health and welfare.
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Lucidi, P., Bacco, G., Sticco, M., Mazzoleni, G., Benvenuti, M., Bernabò, N., et al. (2013). Assessment of motor laterality in foals and young horses (Equus caballus) through an analysis of derailment at trot. Physiol. Behav., 109, 8–13.
Abstract: The conflicting results regarding the study of motor laterality in horses may indicate that there does not exist a proper method to assess the degree and the direction of motor bias in these animals. Unfortunately, even less is known about the development of laterality in horses, and to what extent early manipulations can still exert their effects in adulthood. We propose a new method that can be easily applied at a very early age thus avoiding testing adult horses eventually biased by human handling and/or training. Forty-six horses (29 nine-month-old foals and 17 two-year old horses) were handled since birth bilaterally and housed in groups in wide areas. At the time of the analysis, in order to minimize environmental and sensorial disturbances, each horse was tested in a round pen individually or as dyad mother-foal. The ability/inability to properly execute a circle at trot was then recorded, assuming the direction of derailment, i.e. the cutting of the circle, as an indicator of motor bias. From the results of the study it is arguable that motor laterality in horses is acquired over time: in fact foals tested while their mothers were being subjected to longeing showed a higher percentage of ambidextrous animals, while two-year-old horses appeared biased toward the right (p<0.05). Results are discussed in the light of the scientific knowledge about equine biomechanics, taking into account horses' locomotion that leads to the advancement of the body mass through the activation of a kinetic chain that originates from the hindquarters.
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Stahlbaum, C. C., & Houpt, K. A. (1989). The role of the Flehmen response in the behavioral repertoire of the stallion. Physiol. Behav., 45(6), 1207–1214.
Abstract: The role of the Flehmen response in equine behavior was investigated under field and laboratory conditions. In Experiment 1, a field study made of five stallions on pasture with between three and eighteen mares each during the season indicated the following: 1) The Flehmen response was most frequently preceded by nasal, rather than oral, investigation of substances; 2) The stallions' rate of Flehmen varied with the estrous cycles of the mares; 3) The rate of Flehmen response did not show a variation with time of day; and 4) The Flehmen response was most frequently followed by marking behaviors rather than courtship behaviors. The results suggest that the Flehmen response is not an immediate component of sexual behavior, e.g., courtship of the stallion but may be involved in the overall monitoring of the mare's estrous cycle. Therefore the Flehmen response may contribute to the chemosensory priming of the stallion for reproduction. In Experiment 2 stallions were presented with urine or feces of mares in various stages of the reproductive cycle as well as with their own or other males' urine or feces. The occurrence of sniffing and Flehmen was used to determine the discriminatory ability of the stallions. Stallions can differentiate the sex of a horse on the basis of its feces alone, but cannot differentiate on the basis of urine. This ability may explain the function of fecal marking behavior of stallions.
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Houpt, K. A., Thornton, S. N., & Allen, W. R. (1989). Vasopressin in dehydrated and rehydrated ponies. Physiol. Behav., 45(3), 659–661.
Abstract: Six pony mares deprived of water for 24 hours showed significant increases in plasma vasopressin (2.8 pg/ml) and osmolality (9 mosmol/kg). When water was made available the ponies drank rapidly (5 of 6 drank to satiety within 90 seconds) and corrected their fluid deficits precisely. Vasopressin did not return to predehydration levels until osmolality did after 15 minutes of access to water. The horse differs from rodents and humans, but is similar to pigs in that vasopressin levels do not fall before osmolality returns to normal. Oropharyngeal factors, therefore, may not be as important in vasopressin release in horses as in other species.
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Houpt, K. A., Perry, P. J., Hintz, H. F., & Houpt, T. R. (1988). Effect of meal frequency on fluid balance and behavior of ponies. Physiol. Behav., 42(5), 401–407.
Abstract: Twelve ponies were fed their total daily ration either as one large meal or divided into six small meals. Pre- and post-feeding behavior was recorded six times a day. Blood samples were taken for 30 min before and two hr after the meal. Plasma protein increased from 7.0 to a peak of 7.3 g/dl with small meals and from 7.3 to 8.1 g/dl with large meals, and returned to pre-feeding levels by 90 min post-feeding. Hematocrit rose from 33.3 to 34.1% with small meals and from 33.0 to 36.0% with large meals. These rapid and short-lived increases indicate a decrease in plasma volume. Plasma osmolality rose with feeding from 283 to 285 mosmoles/kg with small meals and from 281 to 288 mosmoles/kg with large meals. Water availability had no significant effect on blood changes. Digestibility and rate of passage were measured with chromic oxide, but there were no differences. Vocalizing (neighing) and walking occurred more often before than after feeding, while eating bedding and engaging in other oral behaviors were more frequent after feeding.
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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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