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Wilson, M. T., Silvestrini, M. C., Morpurgo, L., & Brunori, M. (1979). Electron transfer kinetics between Rhus vernicifera stellacyanin and cytochrome c (horse heart cytochrome c and Pseudomonas cytochrome c551). J Inorg Biochem, 11(2), 95–100.
Abstract: The electron transfer reactions between Rhus vernicifera stellacyanin and either horse heart cytochrome c or Pseudomonas aeruginosa cytochrome c551 were investigated by rapid reaction techniques. The time course of electron transfer is monophasic under all conditions, and thus consistent with a simple formulation of the reaction. Both stopped-flow and temperature-jump experiments yield equilibrium constants in reasonable agreement with values calculated from the redox potentials. The differences in reaction rate between the two cytochromes and stellacyanin are discussed in terms of the Marcus theory.
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Waran, N. K., Robertson, V., Cuddeford, D., Kokoszko, A., & Marlin, D. J. (1996). Effects of transporting horses facing either forwards or backwards on their behaviour and heart rate. Vet. Rec., 139(1), 7–11.
Abstract: The effects of transporting horses facing either forwards or backwards were compared by transporting six thoroughbred horses in pairs in a lorry on one journey facing in the direction of travel, and on another journey facing away from the direction of travel, over a standard one-hour route. Heart rate monitors were used to record their heart rate before, during and after the journey and the horses' behaviour was recorded by scan sampling each horse every other minute. The average heart rate was significantly lower (P < 0.05) when the horses were transported facing backwards, and they also tended to rest on their rumps more (P = 0.059). In the forward-facing position, the horses moved more frequently (P < 0.05) and tended to hold their necks in a higher than normal position and to vocalise more frequently (P = 0.059). During loading the average peak heart rate was 38 bpm lower (P < 0.05) when the horses were backed into the horse box for rear-facing transport than when they were loaded facing forwards. However, there was no difference between transport facing forwards or backwards in terms of the peak unloading heart rate, or the average heart rate during loading or unloading. The horses seemed to find being transported less physically stressful when they were facing backwards than when they were facing forwards.
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Stull, C. L., Spier, S. J., Aldridge, B. M., Blanchard, M., & Stott, J. L. (2004). Immunological response to long-term transport stress in mature horses and effects of adaptogenic dietary supplementation as an immunomodulator. Equine Veterinary Journal, 36(7), 583–589.
Abstract: Reasons for performing study: Little information exists on the immunological effects of transport or the use of supplements to minimise transport stress. Objectives: To establish baseline ranges and evaluate immunophenotypic and functional changes associated with transport and a nutritional ‘adaptogen’ supplement. Methods: Horses received either supplement (n = 10) or placebos (n = 9) during the 30 day study. After 28 days in stalls, 12 horses (6 supplement; 6 placebo) were transported for 24 h, then unloaded and recovered. Venous blood samples were collected on Days 1, 14 and 28 to establish baselines, and on Days 28, 29 and 30 to examine changes during transport and recovery. Results: Transport prompted elevations (P<0.05) in cortisol concentration, neutrophil count and white blood cell counts, while lymphocyte subpopulation counts (CD3+, CD4+, CD8+, CD21+) decreased (P<0.05). Normal phenotypic lymphocyte profiles returned within 24 h of recovery. Supplement effects on immunophenotype (CD21+ and CD8+) were observed in stabled horses (P<0.05), but not in transported horses. Conclusions: These results provide insights into the immunological mechanisms associated with long-term transport. Potential relevance: The existence of a small window of immunological uncertainty follows long-term transportation, enhancing the potential risk of infectious disease in susceptible individuals.
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Strand, S. C., Tiefenbacher, S., Haskell, M., Hosmer, T., McDonnell, S. M., & Freeman, D. A. (2002). Behavior and physiologic responses of mares to short-term isolation. Appl. Anim. Behav. Sci., 78(2-4), 145–157.
Abstract: The aim of this study was to evaluate the behavior and physiologic responses of mares to removal from an established pasture herd and to isolation in a pasture setting for 6 h (Group I, n=5). Responses of mares in Group I were compared to mares that were transported and returned to the herd (Group T, n=5) and to mares moved to the isolation pasture with a companion (Group C, n=5). Behavior was recorded continuously for 6 h on the day before the isolation procedures (baseline, Day 0) and again on the day of the procedure (test, Day 1). Plasma cortisol, white blood cell count (WBC), neutrophil:lymphocyte ratio (N:L), and hematocrit (HCT) were measured once on Day 0 (a.m.) and twice on Day 1 (a.m. and p.m.). Heart rate (HR) was monitored continuously during Day 0 and Day 1. Intradermal response to phytohemagglutinin (PHA) injection was measured 18 h following injection, which was administered at the end of Day 1. Average time spent standing alert increased (P<0.05) in Groups I and C and average time spent grazing decreased (P<0.05) in Group C from Day 0 to Day 1. Also, there was a significant difference between groups (based on a calculated χ2-square value) in the proportion of mares that autogroomed, defecated, urinated, rolled, and whinnied on Day 1. Activity shift rate (ASR) and temperament scores increased significantly in Groups I and C from Day 0 to Day 1 (P<0.05). Plasma cortisol increased significantly in all groups from Day 0 to Day 1, a.m. (P<0.05) and decreased significantly from Day 1, a.m. to Day 1, p.m. (P<0.05). HCT significantly increased in all three groups from Day 0 to Day 1, a.m. (P<0.05). WBC significantly increased in Group T from Day 0 to Day 1, a.m. (P<0.05). N:L ratio significantly increased in Groups I and C from Day 0 and Day 1, a.m. to Day 1, p.m. (P<0.05). A variety of measures did indicate a response to removal from the pasture group, however, the overall, short-term response was minimal. Since the responses of Groups I and C were similar, the effects of isolation versus a novel environment or separation from the established herd could not be differentiated.
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Shanahan, S. (2003). Trailer loading stress in horses: behavioral and physiological effects of nonaversive training (TTEAM). J Appl Anim Welf Sci, 6(4), 263–274.
Abstract: Resistance in the horse to trailer loading is a common source of stress and injury to horses and their handlers. The objective of this study was to determine whether nonaversive training based on the Tellington-Touch Equine Awareness Method (TTEAM; Tellington-Jones &Bruns, 1988) would decrease loading time and reduce stress during loading for horses with a history of reluctance to load. Ten horses described by their owners as “problem loaders” were subjected to pretraining and posttraining assessments of loading. Each assessment involved two 7-min loading attempts during which heart rate and saliva cortisol were measured. The training consisted of six 30-min sessions over a 2-week period during which the horse and owner participated in basic leading exercises with obstacles simulating aspects of trailering. Assessment showed heart rate and saliva cortisol increased significantly during loading as compared to baseline (p <.001 and p <.05, respectively). Reassessment after training showed a decrease in loading time (p <.02), reduced heart rate during loading (p <.002), and reduced saliva cortisol as compared to pretraining assessments. Seven “good loaders” also were subject to loading assessment for physiological comparison. Increases in heart rate during loading were significantly higher in the good loaders (p <.001). Nonaversive training simulating aspects of loading may effectively reduce loading time and stress during loading for horses with a history of resistance to trailer loading.
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Schmidt, A., Möstl, E., Wehnert, C., Aurich, J., Müller, J., & Aurich, C. (2010). Cortisol release and heart rate variability in horses during road transport. Horm. Behav., 57(2), 209–215.
Abstract: Based on plasma cortisol concentrations it is widely accepted that transport is stressful to horses. So far, cortisol release during transport has not been evaluated in depth by non-invasive techniques such as analysis of salivary cortisol and faecal cortisol metabolites. Transport also causes changes in heart rate and heart rate variability (HRV). In this study, salivary cortisol, faecal cortisol metabolites, heart rate and HRV in horses transported by road for short (one and 3.5 h) and medium duration (8 h) were determined. With the onset of transport, salivary cortisol increased immediately but highest concentrations were measured towards the end of transport (4.1 ± 1.6, 4.5 ± 2.6, 6.5 ± 1.8 ng/ml in horses transported for one, 3.5 and 8 h, respectively). Faecal cortisol metabolite concentrations did not change during transport, but 1 day after transport for 3.5 and 8 h had increased significantly (p < 0.01), reflecting intestinal passage time. Compared to salivary cortisol, changes in faecal cortisol metabolites were less pronounced. Heart rate increased and beat-to-beat (RR) interval decreased (p < 0.05) with the onset of transport. Standard deviation of heart rate increased while root mean square of successive RR differences (RMSSD) decreased in horses transported for 3.5 (from 74 ± 5 to 45 ± 6 ms) and 8 h (from 89.7 ± 7 to 59 ± 7 ms), indicating a reduction in vagal tone. In conclusion, transport of horses over short and medium distances leads to increased cortisol release and changes in heart rate and HRV indicative of stress. The degree of these changes is related to the duration of transport. Salivary cortisol is a sensitive parameter to detect transient changes in cortisol release.
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Schmidt, A., Biau, S., Möstl, E., Becker-Birck, M., Morillon, B., Aurich, J., et al. (2010). Changes in cortisol release and heart rate variability in sport horses during long-distance road transport. Domest Anim Endocrinol, 38(3), 179–189.
Abstract: It is widely accepted that transport is stressful for horses, but only a few studies are available involving horses that are transported regularly and are accustomed to transport. We determined salivary cortisol immunoreactivity (IR), fecal cortisol metabolites, beat-to-beat (RR) interval, and heart rate variability (HRV) in transport-experienced horses (N = 7) in response to a 2-d outbound road transport over 1370 km and 2-d return transport 8 d later. Salivary cortisol IR was low until 60 min before transport but had increased (P < 0.05) 30 min before loading. Transport caused a further marked increase (P < 0.001), but the response tended to decrease with each day of transport. Concentrations of fecal cortisol metabolites increased on the second day of both outbound and return transports and reached a maximum the following day (P < 0.001). During the first 90 min on Day 1 of outbound transport, mean RR interval decreased (P < 0.001). Standard deviations of RR interval (SDRR) decreased transiently (P < 0.01). The root mean square of successive RR differences (RMSSD) decreased at the beginning of the outbound and return transports (P < 0.01), reflecting reduced parasympathetic tone. On the first day of both outbound and return transports, a transient rise in geometric HRV variable standard deviation 2 (SD2) occurred (P < 0.01), indicating increased sympathetic activity. In conclusion, transport of experienced horses leads to increased cortisol release and changes in heart rate and HRV, which is indicative of stress. The degree of these changes tended to be most pronounced on the first day of both outbound and return transport.
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Marlin, D. J., Schroter, R. C., White, S. L., Maykuth, P., Matthesen, G., Mills, P. C., et al. (2001). Recovery from transport and acclimatisation of competition horses in a hot humid environment. Equine Vet J, 33(4), 371–379.
Abstract: The aims of the present field-based study were to investigate changes in fit horses undergoing acclimatisation to a hot humid environment and to provide data on which to base recommendations for safe transport and acclimatisation. Six horses (age 7-12 years) were flown from Europe to Atlanta and underwent a 16 day period of acclimatisation. Exercise conditions during acclimatisation (wet bulb globe temperature index 27.6+/-0.0 [mean +/- s.e.]) were more thermally stressful compared with the European climate from which the horses had come (22.0+/-1.8, P<0.001). Following the flight, weight loss was 4.1+/-0.8% bodyweight and took around 7 days to recover. Water intake during the day was significantly increased (P<0.05) compared with night during acclimatisation. Daily mean exercise duration was 72+/-12 min and the majority of work was performed with a heart rate below 120 beats/min. Respiratory rate (fR) was increased (P<0.05) throughout acclimatisation compared with in Europe, but resting morning (AM) and evening (PM) rectal temperature (TREC), heart rate (fC) and plasma volume were unchanged. White blood cell (WBC) count was significantly increased at AM compared with in Europe on Days 4 and 10 of acclimatisation (P<0.01), but was not different by Day 16. In conclusion, horses exposed to hot humid environmental conditions without prior acclimatisation are able to accommodate these stresses and, with appropriate management, remain fit and clinically healthy, without significant risk of heat illness or heat-related disorders, provided they are allowed sufficient time to recover from transport, acclimatisation is undertaken gradually and they are monitored appropriately.
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Kay, R., & Hall, C. (2009). The use of a mirror reduces isolation stress in horses being transported by trailer. Appl. Anim. Behav. Sci., 116(2-4), 237–243.
Abstract: Horse trailers are a common form of transportation for horses and ponies and often require the animal to travel alone or with a single companion. The current study investigated the effect of transporting horses alone, in company or with an acrylic safety mirror (measuring 81cm61.5cm) that provided surrogate companionship. The behavioural and physiological responses of 12 mature horses during a 30-min journey by trailer under the three treatments were compared. Behaviours (vocalisation, eating, head-tossing, pawing, and head-turning) were recorded. In order to assess circulatory changes that occur as part of the response to transport, heart rate (HR), rectal (Tr) and ear-pinna (Tp) temperatures were recorded. When travelling with a live companion significantly less time was spent vocalising (p<0.001), head-turning (p<0.001), head-tossing (p<0.01) and pawing (p<0.01); eating behaviour increased (p<0.05). Physiological responses (increases in HR and Tr and decreases in Tp) were also significantly reduced when travelling with a live companion (p<0.01). Travelling with the mirror did not significantly affect physiological responses compared with travelling alone, but the rise in Tr and fall in Tp was reduced (p=0.052 and p=0.051, respectively) and can be considered a trend. When travelling with a mirror significantly less time was spent turning the head (p<0.01), vocalising (p<0.05) and head-tossing (p<0.05); eating behaviour increased (p<0.05). The only significant difference between travelling with a live companion and a mirror was that the time spent turning the head round was less with a live companion (p<0.05). The provision of surrogate companionship in the form of a mirror was found to be preferable to travel alone, but where possible a live companion is recommended. Isolation during transportation was found to suppress feeding behaviour. Although peripheral blood flow (Tp) has been used to assess transport stress in other species it has not previously been used in the horse. Further evaluation of this non-invasive measure is now required.
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Kalin, N. H., & Shelton, S. E. (2003). Nonhuman primate models to study anxiety, emotion regulation, and psychopathology. Ann N Y Acad Sci, 1008, 189–200.
Abstract: This paper demonstrates that the rhesus monkey provides an excellent model to study mechanisms underlying human anxiety and fear and emotion regulation. In previous studies with rhesus monkeys, stable, brain, endocrine, and behavioral characteristics related to individual differences in anxiety were found. It was suggested that, when extreme, these features characterize an anxious endophenotype and that these findings in the monkey are particularly relevant to understanding adaptive and maladaptive anxiety responses in humans. The monkey model is also relevant to understanding the development of human psychopathology. For example, children with extremely inhibited temperament are at increased risk to develop anxiety disorders, and these children have behavioral and biological alterations that are similar to those described in the monkey anxious endophenotype. It is likely that different aspects of the anxious endophenotype are mediated by the interactions of limbic, brain stem, and cortical regions. To understand the brain mechanisms underlying adaptive anxiety responses and their physiological concomitants, a series of studies in monkeys lesioning components of the neural circuitry (amygdala, central nucleus of the amygdala and orbitofrontal cortex) hypothesized to play a role are currently being performed. Initial findings suggest that the central nucleus of the amygdala modulates the expression of behavioral inhibition, a key feature of the endophenotype. In preliminary FDG positron emission tomography (PET) studies, functional linkages were established between the amygdala and prefrontal cortical regions that are associated with the activation of anxiety.
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