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Burns, T. E., & Clayton, H. M. (1997). Comparison of the temporal kinematics of the canter pirouette and collected canter. Equine Vet J Suppl, (23), 58–61.
Abstract: The objectives were to compare the temporal characteristics of canter pirouette strides with collected canter strides in elite dressage horses, and to determine whether the stride kinematics of the canter pirouettes fulfilled the requirements specified in the Federation Equestre Internationale Rules for Dressage Events. Eleven horses were videotaped (60 fields/s) during the individual medal competition at the 1992 Olympic Games. Temporal variables were extracted from the videotapes using standard methods. Two strides were analysed on each of the left and right leads and these were pooled to give mean values for the collected canter and the pirouettes. The pirouettes were completed in 4-9 strides, (mean of 6.4). In the collected canter strides, mean duration of the suspension was 0.013 s. There was no suspension in any of the pirouette strides, instead the stance phases of the leading forelimb and trailing hindlimb overlapped by a mean of 0.163 s. In 9 horses the trailing forelimb contacted the ground before the diagonal leading hindlimb in the collected canter, whereas in the pirouettes the leading hindlimb always made contact before the trailing forelimb (mean dissociation 0.164 s), giving the strides a distinct 4 beat rhythm. Due to increases in advanced placement between the diagonal limb pair and between the 2 forelimbs, the stride duration was longer in the pirouette (0.879 s) than the collected canter (0.629 s). It is concluded that the canter pirouette strides did not maintain the rhythm and timing of the the collected canter strides in any of the 11 horses.
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Cameron, E. Z. (2004). Facultative adjustment of mammalian sex ratios in support of the Trivers-Willard hypothesis: evidence for a mechanism. Proc Biol Sci, 271(1549), 1723–1728.
Abstract: Evolutionary theory predicts that mothers of different condition should adjust the birth sex ratio of their offspring in relation to future reproductive benefits. Published studies addressing variation in mammalian sex ratios have produced surprisingly contradictory results. Explaining the source of such variation has been a challenge for sex-ratio theory, not least because no mechanism for sex-ratio adjustment is known. I conducted a meta-analysis of previous mammalian sex-ratio studies to determine if there are any overall patterns in sex-ratio variation. The contradictory nature of previous results was confirmed. However, studies that investigated indices of condition around conception show almost unanimous support for the prediction that mothers in good condition bias their litters towards sons. Recent research on the role of glucose in reproductive functioning have shown that excess glucose favours the development of male blastocysts, providing a potential mechanism for sex-ratio variation in relation to maternal condition around conception. Furthermore, many of the conflicting results from studies on sex-ratio adjustment would be explained if glucose levels in utero during early cell division contributed to the determination of offspring sex ratios.
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Carroll, G. L., Matthews, N. S., Hartsfield, S. M., Slater, M. R., Champney, T. H., & Erickson, S. W. (1997). The effect of detomidine and its antagonism with tolazoline on stress-related hormones, metabolites, physiologic responses, and behavior in awake ponies. Vet Surg, 26(1), 69–77.
Abstract: Six ponies were used to investigate the effect of tolazoline antagonism of detomidine on physiological responses, behavior, epinephrine, norepinephrine, cortisol, glucose, and free fatty acids in awake ponies. Each pony had a catheter inserted into a jugular vein 1 hour before beginning the study. Awake ponies were administered detomidine (0.04 mg/kg intravenously [i.v.]) followed 20 minutes later by either tolazoline (4.0 mg/kg i.v.) or saline. Blood samples were drawn from the catheter 5 minutes before detomidine administration (baseline), 5 minutes after detomidine administration, 20 minutes before detomidine administration which was immediately before the administration of tolazoline or saline (time [T] = 0), and at 5, 30, and 60 minutes after injections of tolazoline or saline (T = 5, 30, and 60 minutes, respectively). Compared with heart rate at T = 0, tolazoline antagonism increased heart rate 45% at 5 minutes. There was no difference in heart rate between treatments at 30 minutes. Blood pressure remained stable after tolazoline, while it decreased over time after saline. Compared with concentrations at T = 0, tolazoline antagonism of detomidine in awake ponies resulted in a 55% increase in cortisol at 30 minutes and a 52% increase in glucose at 5 minutes. The change in free fatty acids was different for tolazoline and saline over time. Free fatty acids decreased after detomidine administration. Free fatty acids did not change after saline administration. After tolazoline administration, free fatty acids increased transiently. Tolazoline tended to decrease sedation and analgesia at 15 and 60 minutes postantagonism. Antagonism of detomidine-induced physiological and behavioral effects with tolazoline in awake ponies that were not experiencing pain appears to precipitate a stress response as measured by cortisol, glucose, and free fatty acids. If antagonism of an alpha-agonist is contemplated, the potential effect on hormones and metabolites should be considered.
Keywords: Adrenergic alpha-Antagonists/administration & dosage/*pharmacology; Animals; Behavior, Animal/drug effects/physiology; Blood Glucose/metabolism; Blood Pressure/drug effects/physiology; Consciousness/physiology; Dose-Response Relationship, Drug; Drug Interactions; Epinephrine/blood; Fatty Acids, Nonesterified/blood; Female; Heart Rate/drug effects/physiology; Horse Diseases/metabolism/physiopathology/psychology; Horses/blood/metabolism/*physiology; Hydrocortisone/blood; Hypnotics and Sedatives/administration & dosage/*pharmacology; Imidazoles/administration & dosage/*pharmacology; Injections, Intravenous; Male; Norepinephrine/blood; Receptors, Adrenergic, alpha/drug effects/*physiology; Stress/metabolism/physiopathology/veterinary; Time Factors; Tolazoline/administration & dosage/*pharmacology
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Cerutti, D. T., & Staddon, J. E. R. (2004). Immediacy versus anticipated delay in the time-left experiment: a test of the cognitive hypothesis. J Exp Psychol Anim Behav Process, 30(1), 45–57.
Abstract: In the time-left experiment (J. Gibbon & R. M. Church, 1981), animals are said to compare an expectation of a fixed delay to food, for one choice, with a decreasing delay expectation for the other, mentally representing both upcoming time to food and the difference between current time and upcoming time (the cognitive hypothesis). The results of 2 experiments support a simpler view: that animals choose according to the immediacies of reinforcement for each response at a time signaled by available time markers (the temporal control hypothesis). It is not necessary to assume that animals can either represent or subtract representations of times to food to explain the results of the time-left experiment.
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Christensen, J. W., Rundgren, M., & Olsson, K. (2006). Training methods for horses: habituation to a frightening stimulus. Equine Vet J, 38(5), 439–443.
Abstract: REASONS FOR PERFORMING STUDY: Responses of horses in frightening situations are important for both equine and human safety. Considerable scientific interest has been shown in development of reactivity tests, but little effort has been dedicated to the development of appropriate training methods for reducing fearfulness. OBJECTIVES: To investigate which of 3 different training methods (habituation, desensitisation and counter-conditioning) was most effective in teaching horses to react calmly in a potentially frightening situation. HYPOTHESES: 1) Horses are able to generalise about the test stimulus such that, once familiar with the test stimulus in one situation, it appears less frightening and elicits a reduced response even when the stimulus intensity is increased or the stimulus is presented differently; and 2) alternative methods such as desensitisation and counter-conditioning would be more efficient than a classic habituation approach. METHODS: Twenty-seven naive 2-year-old Danish Warmblood stallions were trained according to 3 different methods, based on classical learning theory: 1) horses (n = 9) were exposed to the full stimulus (a moving, white nylon bag, 1.2 x 0.75 m) in 5 daily training sessions until they met a predefined habituation criterion (habituation); 2) horses (n = 9) were introduced gradually to the stimulus and habituated to each step before the full stimulus was applied (desensitisation); 3) horses (n = 9) were trained to associate the stimulus with a positive reward before being exposed to the full stimulus (counter-conditioning). Each horse received 5 training sessions of 3 min per day. Heart rate and behavioural responses were recorded. RESULTS: Horses trained with the desensitisation method showed fewer flight responses in total and needed fewer training sessions to learn to react calmly to test stimuli. Variations in heart rate persisted even when behavioural responses had ceased. In addition, all horses on the desensitisation method eventually habituated to the test stimulus whereas some horses on the other methods did not. CONCLUSIONS AND POTENTIAL RELEVANCE: Desensitisation appeared to be the most effective training method for horses in frightening situations. Further research is needed in order to investigate the role of positive reinforcement, such as offering food, in the training of horses.
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Clayton, H. M. (1993). Development of conditioning programs for dressage horses based on time-motion analysis of competitions. J Appl Physiol, 74(5), 2325–2329.
Abstract: The time-motion characteristics of Canadian basic- and medium-level dressage competitions are described, and the results are applied in formulating sport-specific conditioning programs. One competition was analyzed at the six levels from basic 1 to medium 3. Each test was divided into a series of sequences based on the type and speed of activity. The durations of the sequences were measured from videotapes. The basic-level tests had fewer sequences, and they were shorter in distance and duration than the medium tests (P < 0.10), but the average speed did not differ between the two levels. It is recommended that horses competing at the basic levels be conditioned using 5-min exercise periods, with short (10-s) bursts of lengthened trot and canter included at basic 2 and above. In preparation for medium-level competitions, the duration of the work periods increases to 7 min, 10- to 12-s bursts of medium or extended trot and canter are included, and transitions are performed frequently to simulate the energy expenditure in overcoming inertia.
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Clayton, H. M. (1997). Classification of collected trot, passage and piaffe based on temporal variables. Equine Vet J Suppl, (23), 54–57.
Abstract: The objective was to determine whether collected trot, passage and piaffe could be distinguished as separate gaits on the basis of temporal variables. Sagittal plane, 60 Hz videotapes of 10 finalists in the dressage competitions at the 1992 Olympic Games were analysed to measure the temporal variables in absolute terms and as percentages of stride duration. Classification was based on analysis of variance, a graphical method and discriminant analysis. Stride duration was sufficient to distinguish collected trot from passage and piaffe in all horses. The analysis of variance showed that the mean values of most variables differed significantly between passage and piaffe. When hindlimb stance percentage was plotted against diagonal advanced placement percentage, some overlap was found between all 3 movements indicating that individual horses could not be classified reliably in this manner. Using hindlimb stance percentage and diagonal advanced placement percentage as input in a discriminant analysis, 80% of the cases were classified correctly, but at least one horse was misclassified in each movement. When the absolute, rather than percentage, values of the 2 variables were used as input in the discriminant analysis, 90% of the cases were correctly classified and the only misclassifications were between passage and piaffe. However, the 2 horses in which piaffe was misclassified as passage were the gold and silver medallists. In general, higher placed horses tended toward longer diagonal advanced placements, especially in collected trot and passage, and shorter hindlimb stance percentages in passage and piaffe.
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Clement, T. S., & Zentall, T. R. (2003). Choice based on exclusion in pigeons. Psychon Bull Rev, 10(4), 959–964.
Abstract: When humans acquire a conditional discrimination and are given a novel-sample-comparison choice, they often reject a comparison known to be associated with a different sample and choose the alternative comparison by default (or by exclusion). In Experiment 1, we found that if, following matching training, we replaced both of the samples, acquisition took five times longer than if we replaced only one of the samples. Apparently, the opportunity to reject one of the comparisons facilitated the association of the other sample with the remaining comparison. In Experiment 2, we first trained pigeons to treat two samples differently (to associate Sample A with Comparison 1 and Sample B with Comparison 2) and then trained them to associate one of those samples with a new comparison (e.g., Sample A with Comparison 3) and to associate a novel sample (Sample C) with a different, new comparison (Comparison 4). When Sample B then replaced Sample C, the pigeons showed a significant tendency to choose Comparison 4 over Comparison 3. Thus, when given the opportunity, pigeons will choose by exclusion.
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Clement, T. S., Feltus, J. R., Kaiser, D. H., & Zentall, T. R. (2000). “Work ethic” in pigeons: reward value is directly related to the effort or time required to obtain the reward. Psychon Bull Rev, 7(1), 100–106.
Abstract: Stimuli associated with less effort or with shorter delays to reinforcement are generally preferred over those associated with greater effort or longer delays to reinforcement. However, the opposite appears to be true of stimuli that follow greater effort or longer delays. In training, a simple simultaneous discrimination followed a single peck to an initial stimulus (S+FR1 S-FR1) and a different simple simultaneous discrimination followed 20 pecks to the initial stimulus (S+FR20 S-FR20). On test trials, pigeons preferred S+FR20 over S+FR1 and S-FR20 over S-FR1. These data support the view that the state of the animal immediately prior to presentation of the discrimination affects the value of the reinforcement that follows it. This contrast effect is analogous to effects that when they occur in humans have been attributed to more complex cognitive and social factors.
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Cottin, F., Barrey, E., Lopes, P., & Billat, V. (2006). Effect of repeated exercise and recovery on heart rate variability in elite trotting horses during high intensity interval training. Equine Vet J Suppl, (36), 204–209.
Abstract: REASONS FOR PERFORMING STUDY: Interval training is a commonly used training method for trotting horses. In addition, trainers are provided with efficient and inexpensive heart rate monitor devices for the management of training. HYPOTHESIS: Since the high frequency (HF) frequency peak (fHF) of heart rate variability (HRV) corresponds to the breathing frequency in combination with stride frequency during trotting, it is hypothesised that modifications of breathing and stride frequencies induced by repeated exercise could be detected from fHF. METHODS: RR interval time series of 7 trotting horses were recorded during an interval training session. Interval training was made up of 5 successive 800 m high-velocity trotting runs (H1, H2...H5) separated by 1 min recovery bouts at low speed (R1, R2...R5). Fast Fourier transform (FFT) and Poincare plot analysis techniques were applied to RR series. RESULTS: Repeated exercise had significant effects on HRV components during interval training. Despite constant trotting velocities during high-speed and recovery, repetition induced a decrease in mean RR interval (H1: 295 +/- 19 vs. H5: 283 +/- 15 msec, P<0.05) and in the root mean square of successive differences in RR series (RMSSD; H1: 6.31 +/- 1.28 vs. H5: 5.31 +/- 1.31 msec, P<0.05). Furthermore, high-speed and recovery repetitions induced an increase in fHF (H1: 1.37 +/- 0.35 vs. H5: 1.62 +/- 0.40 Hz and R1: 0.22 +/- 0.02 vs. R4: 0.64 +/- 0.38 Hz, P<0.05). Hence, recovery induced a decrease in the s.d. of the successive RR series (SDRR; R3: 10.5 +/- 3.96 vs. R5: 6.17 +/- 2.65 msecs, P>0.05) and in the long term index of Poincare plot (SD2; R1: 43.29 +/- 28.90 vs. R5: 18.19 +/- 9.35 msecs, P<0.05). CONCLUSIONS: The observed increase in fHF during the interval training could be induced by alterations of the coupling between breathing and stride frequency linked to the emergence of fatigue. The decrease in SD2 and SDRR during successive recovery bouts could be linked with a deterioration of the recovery pattern. POTENTIAL RELEVANCE: HRV can provide breathing frequency data of Standardbreds during training without any respiratory device. Furthermore, HRV could provide useful makers of the emergence of fatigue states during training.
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