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Søndergaard, E., Jensen, M. B., & Nicol, C. J. (). Motivation for social contact in horses measured by operant conditioning. Appl. Anim. Behav. Sci., In Press, Corrected Proof.
Abstract: Although horses are social animals they are often housed individually with limited social contact to other horses and this may compromise their welfare. The present study included eight young female horses and investigated the strength of motivation for access to full social contact, head contact and muzzle contact, respectively, to a familiar companion horse. Horses were housed individually next to their companion horse and separations between pens prevented physical contact. During daily test sessions horses were brought to a test area where they could access an arena allowing social contact. Arena access during 3 min was given after completion of a predetermined number of responses on a panel. Fixed ratios (FR) of 8, 16, 24, 32 and 40 responses per arena access were applied in a random order, one per daily test session, within each test week (Monday to Friday), and the number of rewards per daily test session was recorded. All horses could access all three types of social contact in a cross-over design, and an empty arena was used as control. Motivational strength was assessed using elasticity of demand functions, which were estimated based on the number of rewards earned and FR. Elasticities of demand for the three types of social contact were low (-0.20), and not significantly different, although increasing FR still resulted in a decrease in rewards obtained for all three types of social contact (P < 0.001). Across FR-levels horses earned more rewards for social contact than for an empty arena, as shown by much higher intercept values (2.51 vs. 0.99; P < 0.001). However, the elasticity of demand for infrequent access to an empty arena (-0.08) was lower than for social contact (P < 0.01) and not significantly different from zero (P = 0.07). Horses performed more social behaviour the lesser the restriction on social contact (full > head > muzzle). However, the finding that horses showed a similar and high motivation for all three types of social contact suggests that they are valued equally highly in a situation where the alternative is no social contact.
Keywords: Horse; Housing; Social behaviour; Operant conditioning; Motivation
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Ahrendt, L. P., Labouriau, R., Malmkvist, J., Nicol, C. J., & Christensen, J. W. (2015). Development of a standard test to assess negative reinforcement learning in horses. Appl. Anim. Behav. Sci., 169, 38–42.
Abstract: Most horses are trained by negative reinforcement. Currently, however, no standardised test for evaluating horses' negative reinforcement learning ability is available. The aim of this study was to develop an objective test to investigate negative reinforcement learning in horses. Twenty-four Icelandic horses (3 years old) were included in this study. The horses were tested in a pressure-release task on three separate days with 10, 7 and 5 trials on each side, respectively. Each trial consisted of pressure being applied on the hindquarter with an algometer. The force of the pressure was increased until the horse moved laterally away from the point of pressure. There was a significant decrease in required force over trials on the first test day (P<0.001), but not the second and third day. The intercepts on days 2 and 3 differed significantly from day 1 (P<0.001), but not each other. Significantly stronger force was required on the right side compared to the left (P<0.001), but there was no difference between first and second side tested (P=0.56). Individual performance was evaluated by median-force and the change in force over trials on the first test day. These two measures may explain different characteristics of negative reinforcement learning. In conclusion, this study presents a novel, standardised test for evaluating negative reinforcement learning ability in horses.
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Giles, S. L., Nicol, C. J., Harris, P. A., & Rands, S. A. (2015). Dominance rank is associated with body condition in outdoor-living domestic horses (Equus caballus). Applied Animal Behaviour Science, .
Abstract: Abstract The aim of our study was to explore the association between dominance rank and body condition in outdoor group-living domestic horses, Equus caballus. Social interactions were recorded using a video camera during a feeding test, applied to 203 horses in 42 herds. Dominance rank was assigned to 194 individuals. The outcome variable body condition score (BCS) was recorded using a 9-point scale. The variables age and height were recorded and considered as potential confounders or effect modifiers. Results were analysed using multivariable linear and logistic regression techniques, controlling for herd group as a random effect. More dominant (p = 0.001) individuals generally had a higher body condition score (p = 0.001) and this association was entirely independent of age and height. In addition, a greater proportion of dominant individuals fell into the obese category (BCS >= 7/9, p = 0.005). There were more displacement encounters and a greater level of interactivity in herds that had less variation in age and height, lending strength to the hypothesis that phenotypic variation may aid cohesion in group-living species. In addition there was a strong quadratic relationship between age and dominance rank (p < 0.001), where middle-aged individuals were most likely to be dominant. These results are the first to link behavioural predictors to body condition and obesity status in horses and should prompt the future consideration of behavioural and social factors when evaluating clinical disease risk in group-living animals.
Keywords: Equine; fatness; obesity; social behaviour; displacement
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Dixon, G., Green, L. E., & Nicol, C. J. (2006). Effect of diet change on the behavior of chicks of an egg-laying strain. J Appl Anim Welf Sci, 9(1), 41–58.
Abstract: Injurious pecking has serious welfare consequences in flocks of hens kept for egg laying, especially when loose-housed. Frequent diet change is a significant risk for injurious pecking; how the mechanics of diet change influence pecking behavior is unknown. This study investigated the effect of diet change on the behavior of chicks from a laying strain. The study included a 3-week familiarity phase: 18 chick pairs received unflavored feed (Experiment 1); 18 pairs received orange oil-flavored (Experiment 2). All chicks participated in a dietary preference test (P); a diet change (DC); or a control group (C), 6 scenarios. All P chicks preferred unflavored feed. In Experiment 1, DC involved change from unflavored to orange-flavored; Experiment 2, orange- flavored to unflavored. Compared with controls, Experiment 2 DC chicks exhibited few behavioral differences; Experiment 1 DC chicks exhibited increased behavioral event rates on Days 1 and 7. They pecked significantly longer at their environment; by Day 7, they showed significantly more beak activity. There was little evidence of dietary neophobia. Change from more preferred to less preferred feed led to increased activity and redirected pecking behavior.
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Haslam, S. M., Brown, S. N., Wilkins, L. J., Kestin, S. C., Warriss, P. D., & Nicol, C. J. (2006). Preliminary study to examine the utility of using foot burn or hock burn to assess aspects of housing conditions for broiler chicken. Br Poult Sci, 47(1), 13–18.
Abstract: 1. Eleven broiler chicken farms, representing 4 production system types, were visited during the last 5 d of the flock cycle: bird and flock details were recorded. Litter friability was assessed at 9 sites within the house, atmospheric ammonia was measured at three sites and bird cleanliness was assessed on a numerical rating scale. 2. For these flocks, hock burn, foot burn and breast burn were measured at the processing plant by standardised assessors. 3. Significant correlations were identified between the percentage of birds with foot burn and average litter score, average house ammonia concentrations and feather score. 4. No correlation was found between the percentage of birds with hock burn or breast burn and average litter scores, average ammonia concentrations or feather score. 5. No correlation was found between stocking density and foot burn, hock burn or breast burn.6. If confirmed, these findings may have implications for the draft EU Broiler Directive, for which it is proposed that permitted stocking density on farm may be determined by the incidence and severity of contact dermatitis measured on plant.
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Crosby, M. B., Svenson, J. L., Zhang, J., Nicol, C. J., Gonzalez, F. J., & Gilkeson, G. S. (2005). Peroxisome proliferation-activated receptor (PPAR)gamma is not necessary for synthetic PPARgamma agonist inhibition of inducible nitric-oxide synthase and nitric oxide. J Pharmacol Exp Ther, 312(1), 69–76.
Abstract: Peroxisome proliferation-activated receptor (PPAR)gamma agonists inhibit inducible nitric-oxide synthase (iNOS), tumor necrosis factor-alpha, and interleukin-6. Because of these effects, synthetic PPARgamma agonists, including thiazolidinediones, are being studied for their impact on inflammatory disease. The anti-inflammatory concentrations of synthetic PPARgamma agonists range from 10 to 50 microM, whereas their binding affinity for PPARgamma is in the nanomolar range. The specificity of synthetic PPARgamma agonists for PPARgamma at the concentrations necessary for anti-inflammatory effects is thus in question. We report that PPARgamma is not necessary for the inhibition of iNOS by synthetic PPARgamma agonists. RAW 264.7 macrophages possess little PPARgamma, yet lipopolysaccharide (LPS)/interferon (IFN)gamma-induced iNOS was inhibited by synthetic PPARgamma agonists at 20 microM. Endogenous PPARgamma was inhibited by the transfection of a dominant-negative PPARgamma construct into murine mesangial cells. In the transfected cells, synthetic PPARgamma agonists inhibited iNOS production at 10 microM, similar to nontransfected cells. Using cells from PPARgamma Cre/lox conditional knockout mice, baseline and LPS/IFNgamma-induced nitric oxide levels were higher in macrophages lacking PPARgamma versus controls. However, synthetic PPARgamma agonists inhibited iNOS at 10 microM in the PPARgamma-deficient cells, similar to macrophages from wild-type mice. These results indicate that PPARgamma is not necessary for inhibition of iNOS expression by synthetic PPARgamma agonists at concentrations over 10 microM. Intrinsic PPARgamma function, in the absence of synthetic agonists, however, may play a role in inflammatory modulation.
Keywords: Animals; Cell Line; Gene Expression/drug effects; Macrophages/drug effects/metabolism; Mice; Mice, Inbred C57BL; Nitric Oxide/*metabolism; Nitric Oxide Synthase/*metabolism; Nitric Oxide Synthase Type II; PPAR delta/metabolism; PPAR gamma/*agonists/deficiency; Thiazolidinediones/pharmacology
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Nicol, C. J. (2004). Development, direction, and damage limitation: social learning in domestic fowl. Learn Behav, 32(1), 72–81.
Abstract: This review highlights two areas of particular interest in the study of social learning in fowl. First, the role of social learning in the development of feeding and foraging behavior in young chicks and older birds is described. The role of the hen as a demonstrator and possible teacher is considered, and the subsequent social influence of brood mates and other companions on food avoidance and food preference learning is discussed. Second, the way in which work on domestic fowl has contributed to an understanding of the importance of directed social learning is examined. The well-characterized hierarchical social organization of small chicken flocks has been used to design studies which demonstrate that the probability of social transmission is strongly influenced by social relationships between birds. The practical implications of understanding the role of social learning in the spread of injurious behaviors in this economically important species are briefly considered.
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Freire, R., Wilkins, L. J., Short, F., & Nicol, C. J. (2003). Behaviour and welfare of individual laying hens in a non-cage system. Br Poult Sci, 44(1), 22–29.
Abstract: 1. A leg band containing a transponder was fitted to 80 birds in a perchery containing 1,000 birds. 2. The transponder emitted a unique identification number when a bird walked on one of 8 flat antennae on the floor. The recording apparatus was used to measure the amount of time that each of the tagged birds spent on the slatted and littered areas in a 6-week period. 3. Some birds spent long periods of time on the slats, possibly as a means of avoiding repeated attacks. Duration on the slats was greatest in birds with the worst (as opposed to better) feather scores of the head, back and tail regions. 4. Birds that spent long periods on the slats were lighter than other birds at both 39 weeks of age and 72 weeks of age and had greater back, head and tail feather damage, consistent with these birds being victims of pecking. 5. Tagged birds received a social avoidance test outside the perchery at 39 weeks of age, which suggested that birds retreated to the slats in response to pecks rather than just to close proximity to other birds. 6. The failure to find that duration on the slats was related to anatomical indicators of stress (liver, spleen and bursa of Fabricius) suggests that retreating to the slats following pecking attenuates physiological stress responses. 7. We conclude that the provision of areas where birds in a large group can avoid pecking may improve the welfare of a minority of victimised birds.
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Nicol, C. J., & Pope, S. J. (1999). The effects of demonstrator social status and prior foraging success on social learning in laying hens. Anim. Behav., 57(1), 163–171.
Abstract: Opportunities for social learning within a group of animals are likely to be influenced by the social dynamics of that group. Some individuals may be more influential demonstrators than others even when there are no differences in their skill level or performance. In this study of domestic hens,Gallus gallus domesticus, differences in demonstrator salience were examined. From 24 separate flocks we selected as demonstrators a dominant cockerel, a dominant hen, a mid-ranking hen or a subordinate hen. Demonstrators were pretrained to perform an operant discrimination task to obtain food. Six observers from each flock individually watched the demonstrator perform the task for four 5-min sessions held on consecutive days. On the fifth day observers were tested individually in the operant chamber. We analysed data from 19 flocks, where there were no quantitative differences in demonstrator performance. Observer hens of relatively high social status performed more correct operant pecks than observer hens of relatively low social status. Demonstrator category also had a significant effect on subsequent observer behaviour. Hens that had observed cockerels performed very few general pecks or operant pecks. Hens that had observed dominant hens performed more operant pecks, but hens that had observed sub-ordinate hens performed more general pecks in the chamber. The results suggested either that there was an interaction between dominance and gender in demonstrator salience or that dominant hens might have been influential because of some factor imperfectly associated with their dominance status. A possible candidate was the foraging ability of the dominant hens. In a second experiment using the same protocol, we manipulated the prior foraging success of dominant hens from four additional flocks but this had no significant effect on their subsequent influence as demonstrators.
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Abeyesinghe, S. M., Nicol, C. J., Hartnell, S. J., & Wathes, C. M. (2005). Can domestic fowl, Gallus gallus domesticus, show self-control? Anim. Behav., 70(1), 1–11.
Abstract: An important aspect of cognition is whether animals live exclusively in the present or can anticipate the future. Defined as self-control, the ability to choose a large, remote reinforcer over a small, proximate reinforcer available at the same frequency has been examined in a number of species, often proving difficult to demonstrate. We investigated self-control for food in domestic fowl using a standard two-key operant task and an equivalent two-choice return maze (TCRM) task. When hens chose between a 2-s delay to a 3-s feed access (impulsive) and a 6-s delay to a 7-s feed access (self-control), they appeared unable to discriminate in the TCRM but were impulsive in the operant task. We explored reasons for not choosing self-control in the operant task, first by examining the relation between feed access time and actual feed intake. A second operant experiment examined whether failure to show self-control could be attributed to an inability to combine the delay and access (quantity) reward information associated with choices to reach overall predictions of value. New hens chose between a 2-s delay to a 3-s feed access (impulsive) and either a 22-s delay to a 22-s feed access (standard self-control) or a 6-s delay to a 22-s feed access (jackpot self-control). While hens were impulsive in the standard condition, they showed significant and pronounced self-control in the jackpot condition, eliminating the possibility of an absolute cognitive constraint. Impulsive behaviour can instead be explained by temporal discounting: perceived depreciation of reward value as a function of the uncertainty associated with delay. Implications for welfare are discussed.
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