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	Author	Nicol, C.J.
	Title	The social transmission of information and behaviour			Type	Journal Article
	Year	1995	Publication	Applied Animal Behaviour Science	Abbreviated Journal	Appl. Anim. Behav. Sci.
	Volume	44	Issue	2-4	Pages	79-98
	Keywords	Social learning; Imitation; Social facilitation; Cultural transmission; Stereotypies
	Abstract	Social influences on established behaviour and on the acquisition of new information and behaviour are reviewed. Distinctions between social facilitation and contagious behaviour are drawn and suggestions for further research on contagious behaviour are made. Socially derived visual, olfactory and auditory cues are considered as important influences on behaviour and subsequent learning. The evidence supporting two potential mechanisms of social learning, i.e. stimulus enhancement followed by individual learning, and imitation, is reviewed in detail. It is argued that the functions of social learning are similarly heterogeneous and include motor skill acquisition, gathering of environmental information, and social conformity. Factors affecting the spread of socially acquired skills, including the social relationship between demonstrator and observer, are highlighted. Lastly, the few studies of social learning that have been conducted with domestic species are reviewed and potential applied goals that could stimulate further research in this area are suggested.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN		ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	refbase @ user @			Serial	577
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	Author	Nicol, C.J.; Badnell-Waters, A.J.; Bice, R.; Kelland, A.; Wilson, A.D.; Harris, P.A.
	Title	The effects of diet and weaning method on the behaviour of young horses			Type	Journal Article
	Year	2005	Publication	Applied Animal Behaviour Science	Abbreviated Journal	Appl. Anim. Behav. Sci.
	Volume	95	Issue	3-4	Pages	205-221
	Keywords	Horse; Diet; Weaning; Temperament test
	Abstract	The effects of diet on horse behaviour have not previously been quantified in detail. In this study, we examined the behaviour of 17 foals from the age of 2 to 40 weeks. Each foal received either a starch and sugar (SS) diet or a fat and fibre (FF) diet. The two diets contained similar digestible energy, crude protein and micronutrients, but differed in the fat and non-structural carbohydrate balance. The baseline behaviour of the foals was observed every 2 weeks by focal animal sampling. Additional behavioural observations were conducted when the foals were weaned by one of two methods. Approximately 2 months after weaning, the temperament and tractability of the young horses was assessed using standardised tests. Responses to a novel object, to a novel person, and during a handling test were observed and quantified. Horses grew well on both diets with no apparent effects of diet on growth rate or baseline behaviour. Immediately after weaning, horses receiving the FF diet cantered less frequently (F = 5.10; p < 0.05), for a shorter duration (F = 7.23; p < 0.05) and appeared to be more settled. Foals that were barn-weaned appeared more stressed than foals that were paddock-weaned. In the temperament tests, horses receiving the FF diet spent significantly more time investigating (F = 6.78; p < 0.05), and less time looking at (F = 7.93; p < 0.05), the novel object than horses receiving the SS diet. They also spent less time walking away from the novel person (F = 5.16; p < 0.05) and their time taken to complete the handling test was significantly lower (F = 8.72; p = 0.01). Overall, the horses that received the FF diet appeared less distressed immediately after weaning and seemed calmer and more inquisitive during a range of temperament tests.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN		ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	3642
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	Author	Hothersall, B.; Harris, P.; Sörtoft, L.; Nicol, C.J.
	Title	Discrimination between conspecific odour samples in the horse (Equus caballus)			Type	Journal Article
	Year	2010	Publication	Applied Animal Behaviour Science	Abbreviated Journal	Appl. Anim. Behav. Sci.
	Volume	126	Issue	1-2	Pages	37-44
	Keywords	Horse; Olfaction; Social discrimination; Urine; Faeces
	Abstract	Behavioural observations suggest that smell is important in social discriminations between horses but balanced studies of this capacity are lacking. We used a habituation-discrimination procedure to investigate the ability of horses to distinguish between pairs of odour samples from different individuals. In Study 1, separate tests were conducted for urine, faeces or fleece fabric previously rubbed on the coat (to pick up body odour samples (BOS)) and donor pairs differed in sex, and age. 10 pregnant mares each underwent three tests, one per sample type. A test consisted of three successive 2-min presentations of a sample from Individual A with a simultaneous presentation of a sample from Individual B during the final presentation. Doubly repeated measures ANOVA indicated a main effect of sample type on investigative response (df = 2, f = 7.98, P = 0.004): durations were longer for BOS than for urine or faeces but habituation across trials was most consistent for urine. In the final presentation, mares demonstrated discrimination by investigating the novel urine sample (B) more than the repeated sample (novel: median 8.0 s, IQR = 10; repeated: median 2.5 s, IQR = 6; z = -2.558, P = 0.008). In Study 2, urine samples from castrated male donors were used and neither mares nor their 4-month-old foals discriminated between samples from different individuals in the final presentation. The findings suggest that urine odour may contain some information that horses can use to discriminate between conspecifics. This may be limited to the level of broad categories such as sex or reproductive status; further investigation is needed to reveal what functional information can be transmitted and what compounds are involved.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0168-1591	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5134
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	Author	Søndergaard, E.; Jensen, M.B.; Nicol, C.J.
	Title	Motivation for social contact in horses measured by operant conditioning			Type	Journal Article
	Year		Publication	Applied Animal Behaviour Science	Abbreviated Journal	Appl. Anim. Behav. Sci.
	Volume	In Press, Corrected Proof	Issue		Pages
	Keywords	Horse; Housing; Social behaviour; Operant conditioning; Motivation
	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.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0168-1591	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5388
Permanent link to this record



	Author	Søndergaard, E.; Jensen, M.B.; Nicol, C.J.
	Title	Motivation for social contact in horses measured by operant conditioning			Type	Journal Article
	Year	2011	Publication	Applied Animal Behaviour Science	Abbreviated Journal	Appl. Anim. Behav. Sci.
	Volume	132	Issue	3-4	Pages	131-137
	Keywords	Horse; Housing; Social behaviour; Operant conditioning; Motivation
	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.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0168-1591	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5410
Permanent link to this record



	Author	Ahrendt, L.P.; Labouriau, R.; Malmkvist, J.; Nicol, C.J.; Christensen, J.W.
	Title	Development of a standard test to assess negative reinforcement learning in horses			Type	Journal Article
	Year	2015	Publication	Applied Animal Behaviour Science	Abbreviated Journal	Appl. Anim. Behav. Sci.
	Volume	169	Issue		Pages	38-42
	Keywords	Algometry; Horse behaviour; Learning performance; Operant conditioning; Pressure-release; Horse training
	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.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0168-1591	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	6650
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	Author	Nicol, C.J.; Pope, S.J.
	Title	The effects of demonstrator social status and prior foraging success on social learning in laying hens			Type	Journal Article
	Year	1999	Publication	Animal Behaviour.	Abbreviated Journal	Anim. Behav.
	Volume	57	Issue	1	Pages	163-171
	Keywords
	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.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN		ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	refbase @ user @			Serial	715
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	Author	Abeyesinghe, S.M.; Nicol, C.J.; Hartnell, S.J.; Wathes, C.M.
	Title	Can domestic fowl, Gallus gallus domesticus, show self-control?			Type	Journal Article
	Year	2005	Publication	Animal Behaviour.	Abbreviated Journal	Anim. Behav.
	Volume	70	Issue	1	Pages	1-11
	Keywords
	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.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
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	ISSN		ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	2897
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	Author	Nicol, C.J.; Badnell-Waters, A.J.
	Title	Suckling behaviour in domestic foals and the development of abnormal oral behaviour			Type	Journal Article
	Year	2005	Publication	Animal Behaviour.	Abbreviated Journal	Anim. Behav.
	Volume	70	Issue	1	Pages	21-29
	Keywords
	Abstract	We investigated how the behaviour of domestic foals, Equus caballus, living at pasture with their dams was associated with foal gender, mare rank and the development of abnormal oral behaviour, both during the preweaning period, and over a period of up to 4 years postweaning. A population of 186 foals belonging to private owners and commercial studs was studied. The behaviour of male and female foals hardly differed, but mare rank affected patterns of foal social interaction and suckling behaviour, with foals of subordinate mares involved in more affiliative interactions. These foals also spent more time in perisuckling activities such as teat nuzzling than foals of other mares. During the study, 18 foals developed abnormal oral behaviour before weaning and 42 foals developed abnormal oral behaviour after weaning. The development of abnormal oral behaviour was associated with suckling behaviour in a variety of ways. Foals that had already developed abnormal oral behaviour at the time of the preweaning observations were involved in more suckling terminations within bouts than normal foals or foals that developed future abnormal behaviour, and pushing the udder with the muzzle was most frequent in these foals. Foals that had no current abnormal oral behaviour, but that would develop this in the future, spent more time suckling and twice as much time teat nuzzling as other foals. The results add to the growing evidence of associations between digestive function and abnormal oral behaviour in horses.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN		ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	3631
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	Author	Nicol, C.J.; Adachi, M.; Akiyama, T.E.; Gonzalez, F.J.
	Title	PPARgamma in endothelial cells influences high fat diet-induced hypertension			Type	Journal Article
	Year	2005	Publication	American journal of hypertension : journal of the American Society of Hypertension	Abbreviated Journal	Am J Hypertens
	Volume	18	Issue	4 Pt 1	Pages	549-556
	Keywords	Administration, Oral; Animals; Antihypertensive Agents/pharmacology; Blood Pressure/drug effects; Diabetes Mellitus, Type 2/physiopathology; Dietary Fats/administration & dosage/pharmacology; Dose-Response Relationship, Drug; Endothelial Cells/metabolism; Female; Heart Rate/drug effects; Hypertension/etiology; Ligands; Male; Mice; Mice, Knockout; PPAR gamma/metabolism; Sodium Chloride/administration & dosage/pharmacology; Thiazolidinediones/pharmacology
	Abstract	BACKGROUND: Peroxisome proliferator-activated receptor gamma (PPARgamma) ligands improve human hypertension. However, the mechanism and site of this effect remains unknown, confounded by PPARgamma expression in many cell types, including endothelial cells (ECs). METHODS: To evaluate the vascular role of PPARgamma we used a conditional null mouse model. Specific disruption of PPARgamma in ECs was created by crossing Tie2-Cre+ transgenic (T2T+) and PPARgamma-floxed (fl/fl) mice to generate PPARgamma (fl/fl)T2T+ (PPARgamma E-null) mice. Conscious 8- to 12-week-old congenic PPARgamma (fl/fl)Cre- (wild type) and PPARgamma E-null mice were examined for changes in systolic blood pressure (BP) and heart rate (HR), untreated, after 2 months of salt-loading (drinking water), and after treatment for 3 months with high fat (HF) diet alone or supplemented during the last 2 weeks with rosiglitazone (3 mg/kg/d). RESULTS: Untreated PPARgamma E-nulls were phenotypically indistinguishable from wild-type littermates. However, compared to similarly treated wild types, HF-treated PPARgamma E-nulls had significantly elevated systolic BP not seen after normal diet or salt-loading. Despite sex-dependent baseline differences, salt-loaded and HF-treated PPARgamma E-nulls of either sex had significantly elevated HR versus wild types. Interestingly, rosiglitazone improved serum insulin levels, but not HF diet-induced hypertension, in PPARgamma E-null mice. CONCLUSIONS: These results suggest that PPARgamma in ECs not only is an important regulator of hypertension and HR under stressed conditions mimicking those arising in type 2 diabetics, but also mediates the antihypertensive effects of rosiglitazone. These data add evidence supporting a beneficial role for PPARgamma-specific ligands in the treatment of hypertension, and suggest therapeutic strategies targeting ECs may prove useful.
	Address	Laboratory of Metabolism, Center for Cancer Research, National Cancer Institute, National Institutes of Health, Bethesda, Maryland 20892, USA
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0895-7061	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:15831367			Approved	no
	Call Number	refbase @ user @			Serial	69
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