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Author Malmgren, L.; Andresen, Ø.; Dalin, A.-M. url  doi
openurl 
  Title (down) Effect of GnRH immunisation on hormonal levels, sexual behaviour, semen quality and testicular morphology in mature stallions Type Journal Article
  Year 2001 Publication Equine Veterinary Journal Abbreviated Journal Equine vet. J.  
  Volume 33 Issue 1 Pages 75-83  
  Keywords horse; GnRH; immunisation; stallion; testosterone; oestrone sulphate; sexual behaviour; semen quality  
  Abstract Summary The aim of this study was to investigate the effect of gonadotrophin-releasing hormone (GnRH) immunisation on mature stallions that had been used for breeding. Four Standardbred stallions were used in the study: 3 experimental animals and 1 control animal. Semen was collected regularly, i.e. twice/week, during the 4 months prior to the experimental period. The stallions were immunised against GnRH with a GnRH-BSA conjugate. Equimune was used as the adjuvant. The stallions were immunised on 5 occasions, 4 at 2 week intervals, and the fifth 4 weeks after the fourth. Blood samples were taken once a week for analysis of GnRH antibody titre and every third week for testosterone and oestrone sulphate analyses. Semen was collected once a week, and libido and sexual behaviour were observed. Ejaculate volume, sperm concentration, total number of sperm in the ejaculate, sperm motility and sperm morphology were evaluated. Testicular size was measured once a week. At the end of the study, the stallions were castrated, and a histological examination of the testes performed. All immunised stallions produced antibodies against GnRH, and plasma testosterone concentration decreased. However, the effect of immunisation varied between stallions. In 2 of the stallions, high levels of antibodies were found, while in the third, the level was moderate. Four weeks after the first immunisation, a decrease in libido was observed. Two months after the first immunisation, marked changes in semen quality were observed in the 2 stallions with high antibody titres. Fourteen weeks after the first immunisation, the total number of sperm/ejaculate had decreased from >8.6×109 to<2.7×109, sperm motility from >59 to<10% and the frequency of morphological normal spermatozoa had decreased from >60 to<14%. The dominating abnormalities were abnormal head shapes, proximal cytoplasmic droplets and detached heads. In the third stallion, only slight changes in semen quality were found. No changes were observed in the control stallion. Decreases in testicular size were noted in all of the experimental stallions. Pronounced histological alterations in the testes were observed in 2 of the stallions. It is concluded that the vaccine was effective in stimulating production of GnRH antibodies and in suppressing testicular function and androgen secretion. However, there was an individual variation in the responses among the stallions and, further, libido was not totally suppressed.  
  Address  
  Corporate Author Thesis  
  Publisher Blackwell Publishing Ltd Place of Publication Editor  
  Language Summary Language Original Title  
  Series Editor Series Title Abbreviated Series Title  
  Series Volume Series Issue Edition  
  ISSN 2042-3306 ISBN Medium  
  Area Expedition Conference  
  Notes Approved no  
  Call Number Equine Behaviour @ team @ Serial 5312  
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Author Falewee, C.; Gaultier, E.; Lafont, C.; Bougrat, L.; Pageat, P. url  openurl
  Title (down) Effect of a synthetic equine maternal pheromone during a controlled fear-eliciting situation Type Journal Article
  Year 2006 Publication Applied Animal Behaviour Science Abbreviated Journal Appl. Anim. Behav. Sci.  
  Volume 101 Issue 1-2 Pages 144-153  
  Keywords Horses; Behaviour; Fear; Pheromone; Heart rate; Performance  
  Abstract Horses are known to show fear reactions when confronted with novelty and this can be a considerable hindrance in the context of working situations such as riding, dressage or racing. The aim of the present study was to measure the potential effects of a synthetic analogue of the Equine Appeasing Pheromone on saddled horses when subjected to a stressful situation using a double-blinded, placebo controlled study design. A group of 40 horses was analyzed during this study and horses were divided by sex, breed and reactivity into two homogenized groups. The test, which consisted of walking the horse through a fringed curtain, was selected from a range of tests which are used to assess behaviour for the selection of French breeding stock. Horses that could have been subjected to the test on a previous occasion, and therefore be familiar with it, were not included. Behavioural and physiological parameters were both taken into account with measures of time to go through the curtain, fear related typical behavioural patterns, based on available literature detailed in the bibliography, and heart rate being recorded. Parameters were analyzed by means of Mann-Whitney U-test. Significant differences were noticed between the two groups concerning heart rate data during the test (UMeanHR = 100.5, pMeanHR = 0.02; UMaxHR = 75, pMaxHR = 0.001) and during the whole measured period (UMeanHR = 67, pMeanHR = 0.005; UMaxHR = 58, pMaxHR = 0.002). Observation of the animals also revealed less behavioural items characteristic of fear within the treated group. As a result, horses performed the test with a better time performance when they received the pheromone analogue (U = 62, p = 0.002). The main parameter, area under the HR graph, is based on heart rate measure and performance. Differences noticed (U = 74, p = 0.002) for this parameter lead to the conclusion that horses who received EAP underwent less stress related consequences in terms of their cardiac physiology. As horses are subjected to a number of foreseeable stressful events this study suggests that the use of Equine Appeasing Pheromone could be a significant factor in improving the welfare of this species.  
  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 3992  
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Author Giles, S.L.; Nicol, C.J.; Harris, P.A.; Rands, S.A. url  doi
openurl 
  Title (down) Dominance rank is associated with body condition in outdoor-living domestic horses (Equus caballus) Type Journal Article
  Year 2015 Publication Applied Animal Behaviour Science Abbreviated Journal  
  Volume Issue Pages  
  Keywords Equine; fatness; obesity; social behaviour; displacement  
  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.  
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  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 @ Giles2015 Serial 5864  
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Author Yarnell, K.; Hall, C.; Royle, C.; Walker, S.L. url  doi
openurl 
  Title (down) Domesticated horses differ in their behavioural and physiological responses to isolated and group housing Type Journal Article
  Year 2015 Publication Physiology & Behavior Abbreviated Journal Physiol.Behav.  
  Volume 143 Issue Pages 51-57  
  Keywords Equine; Behaviour; Corticosterone; Housing  
  Abstract Abstract The predominant housing system used for domestic horses is individual stabling; however, housing that limits social interaction and requires the horse to live in semi-isolation has been reported to be a concern for equine welfare. The aim of the current study was to compare behavioural and physiological responses of domestic horses in different types of housing design that provided varying levels of social contact. Horses (n = 16) were divided equally into four groups and exposed to each of four housing treatments for a period of five days per treatment in a randomized block design. The four housing treatments used were single housed no physical contact (SHNC), single housed semi-contact (SHSC), paired housed full contact (PHFC) and group housed full contact (GHFC). During each housing treatment, adrenal activity was recorded using non-invasive faecal corticosterone metabolite analysis (fGC). Thermal images of the eye were captured and eye temperature was assessed as a non-invasive measure of the stress response. Behavioural analysis of time budget was carried out and an ease of handling score was assigned to each horse in each treatment using video footage. SHNC horses had significantly higher (p = 0.01) concentrations of fGC and were significantly (p = 0.003) more difficult to handle compared to the other housing types. GHFC horses, although not significantly different, had numerically lower concentrations of fGC and were more compliant to handling when compared to all other housing treatments. Eye temperature was significantly (p = 0.0001) lower in the group housed treatment when compared to all other treatments. These results indicate that based on physiological and behavioural measures incorporating social contact into the housing design of domestic horses could improve the standard of domestic equine welfare.  
  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 0031-9384 ISBN Medium  
  Area Expedition Conference  
  Notes Approved no  
  Call Number Equine Behaviour @ team @ Serial 5920  
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Author Siniscalchi, M.; Sasso, R.; Pepe, A.M.; Vallortigara, G.; Quaranta, A. url  doi
openurl 
  Title (down) Dogs turn left to emotional stimuli Type Journal Article
  Year 2010 Publication Behavioural Brain Research Abbreviated Journal Behav. Brain. Res.  
  Volume 208 Issue 2 Pages 516-521  
  Keywords Dog; Laterality; Vision; Behaviour; Physiology; Cognition; Emotion; Animal welfare  
  Abstract During feeding behaviour, dogs were suddenly presented with 2D stimuli depicting the silhouette of a dog, a cat or a snake simultaneously into the left and right visual hemifields. A bias to turn the head towards the left rather than the right side was observed with the cat and snake stimulus but not with the dog stimulus. Latencies to react following stimulus presentation were lower for left than for right head turning, whereas times needed to resume feeding behaviour were higher after left rather than after right head turning. When stimuli were presented only to the left or right visual hemifields, dogs proved to be more responsive to left side presentation, irrespective of the type of stimulus. However, cat and snake stimuli produced shorter latencies to react and longer times to resume feeding following left rather than right monocular visual hemifield presentation. Results demonstrate striking lateralization in dogs, with the right side of the brain more responsive to threatening and alarming stimuli. Possible implications for animal welfare are discussed.  
  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 0166-4328 ISBN Medium  
  Area Expedition Conference  
  Notes Approved no  
  Call Number Equine Behaviour @ team @ Serial 5080  
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Author Stachurska, A.; Pieta, M.; Ussing, A.P.; Kapron, A.; Kwiecinska, N. url  doi
openurl 
  Title (down) Difficulty of cross-country obstacles for horses competing in Three Day Events Type Journal Article
  Year 2010 Publication Applied Animal Behaviour Science Abbreviated Journal Appl. Anim. Behav. Sci.  
  Volume 123 Issue 3-4 Pages 101-107  
  Keywords Cross-country; Obstacles; Eventing horse; Behaviour  
  Abstract The objective of the study was to determine which cross-country obstacles are more difficult for eventing horses. Jumping scores were considered in terms of the horse's reaction to novelty and to the fearfulness of novel objects which are the fences situated in novel terrain. The data concerned 11 classes of One to Four Star level (stars showing the difficulty of the class) held at the Olympic Games and three international Three Day Events. A total of 400 entries, in which 259 horses jumped 372 obstacles were considered. Scores of 11,341 jumps at particular fences were categorized either as faulty jumps or non-faulty jumps. Factors describing the fences versus the jumping scores were studied with least square analysis of variance, with respect to the interaction between the star level and the fence traits. The overall frequency of faults at the cross-country amounted to 4.33 ± 0.57%. Among the effects analyzed, the difficulty of cross-country fences for the horses depends upon whether an obstacle is single or is an element of a combination, whether it is straight or requiring an effort in both height and spread, is broad or narrow, has a solid top or a brush, has an alternative or not and whether it is a water crossing or not. At One Star level, the less experienced horses react differently to certain fence traits compared to horses participating in Three or Four Star levels. It is concluded that the equine visionary system, being less developed towards identification of stationary objects than the human visionary system may be a key towards explaining the horse's behaviour while jumping the obstacle. The height-spread obstacles, those of the narrow front, with the brush and with the alternative seem to involve more faults because of the equine low-acuity vision. The same reason may justify the similar frequency of faults at the single obstacles and the first elements in combinations. Accurate methods of measuring equine vision would complement behavioural tests and should both be introduced into the selection of eventing 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 5076  
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Author Nagy, K.; Bodó, G.; Bárdos, G.; Bánszky, N.; Kabai, P. url  doi
openurl 
  Title (down) Differences in temperament traits between crib-biting and control horses Type Journal Article
  Year 2010 Publication Applied Animal Behaviour Science Abbreviated Journal Appl. Anim. Behav. Sci.  
  Volume 122 Issue 1 Pages 41-47  
  Keywords Crib-biting; Stereotypic behaviour; Equine temperament; Trainability; Coping strategies  
  Abstract Recent studies have suggested that crib-biting in horses is associated with diminished capacity of learning or coping with stress. Such findings raise the question whether trainability, which is fundamentally important in practice, could also be affected by stereotypic behaviour. Trainability of a horse is difficult to assess in simple tests, however, it is reliably estimated by experienced riders. To assess trainability and other characteristics related to that, a questionnaire survey was conducted with the owners of 50 crib-biting and 50 control horses. Where possible, control horses were selected from the same establishment as crib-biters. Groups did not differ significantly regarding age, breed, gender, training level or usage. Principal component analysis revealed three main factors which can be labelled as [`]Anxiety', [`]Affability' and [`]Trainability'. The [`]Anxiety' factor consisted of the items [`]Nervousness', [`]Excitability', [`]Panic', [`]Inconsistent emotionality', [`]Vigilance', [`]Skittishness', and [`]Timidity'. [`]Affability' consisted of [`]Friendliness toward people', [`]Cooperation', [`]Docility' and [`]Friendliness toward horses'. [`]Trainability' involved [`]Concentration', [`]Trainability', [`]Memory', and [`]Perseverance'. Temperament traits were not affected by age, gender, breed or training level, but the usage of the horse and the presence of crib-biting behaviour had significant effects. Competition horses had lower level of [`]Anxiety' (p = 0.032) and higher level of [`]Trainability' (p = 0.068) than leisure horses. Crib-biting horses had significantly lower level of [`]Anxiety' than control horses (p < 0.001), while [`]Trainability' and [`]Affability' did not differ between groups (p = 0.823 and p = 0.543, respectively). Competition horses are more often exposed to novel environment and to frightening stimuli (e.g. colourful obstacles) than leisure horses and therefore might have also become more habituated to these types of stimuli. Coping with novel situation may be enhanced by defusing nervous behaviour by the more experienced riders of competition. Previous studies indicated crib-biting horses to be less reactive when challenged as compared to control horses. We suggest that the virtual calmness and lower nervousness of the crib-biting horses might be due to the passive coping style of these animals. [`]Affability' of horses might be more related to housing and management conditions than to crib-biting. Contrary to expectations, scores on [`]Trainability' had not coincided with the impaired learning of crib-biting horses reported in laboratory tests. However, previous behavioural tests on equine learning rarely had a direct relevance to the training abilities of the horses. Our results do not support crib-biting stereotypy to affect performance in training, which is a complex learning process involving cooperation and docility in the social environment.  
  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 5078  
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Author VanDierendonck, M.C. url  openurl
  Title (down) Diff erences in social behaviour between late pregnant, post-partum and barren mares in a herd of Icelandic horses Type Manuscript
  Year 2006 Publication The Importance of Social Relationships in Horses Abbreviated Journal  
  Volume Chapter 5 Issue Pages  
  Keywords Feral horses are social animals, which have to rely on survival strategies centered on the formation of cohesive social bonds within their bands. Many problems in the husbandry of social animals such as horses, are due to the fact that the limits of their adaptive abilities are exceeded. Evidence suggests that the fundamental social characteristics of domestic horses have remained relatively unchanged. The social structure, social strategies and social interactions were investigated (3 non-consecutive years, 24 hr per day for several weeks) in long term established groups of domestic horses (mares and geldings of all ages) and a few small introduced groups, kept in (semi)natural environments. The general aim was to investigate the social needs of domestic horses. The social life of domestic horses was characterised by long lasting bonds with preferred partners which were established and maintained by allogrooming, play, proximity and dominance behaviours. Bonding partners were mainly found within the same sex-age group, but adult geldings also bonded with sub-adult mares and geldings. Adult mares were clustered in a group, while the other animals formed a second group. Among the adult mares, subgroups according to reproductive state were formed. Individuals regulated their social network by interfering with interactions between other members of the herd, which in itself is complex. An intervention is a behavioural action of one animal that actively interferes with an ongoing interaction between a dyad with the apparent aim of altering that interaction. This was verified by post-hoc analyses of disturbed and undisturbed interactions. Interventions in allogrooming or play were performed significantly more often when at least one member of the initial dyad was a preferred partner of, or familiar to (within the small introduced bands) the intervener. The stronger the preferred association in allogrooming between the intervener and member(s) of the initial dyad, the higher the probability the intervener would displace one initial member and continue allogrooming with the other. Just five behaviours were extracted which reliably reflected the dominance relations among horses. Aggression with the hind quarters was used both offensively and defensively and therefore not suitable as a reliable parameter. Individual dominance relationships were related to social experience. The implications of these findings for horse husbandry were assessed. It is argued that the execution of affiliative behaviours may be rewarding in itself, and therefore always will be a highly motivated behaviour. It is shown that social positive physical interactions (allogrooming, play) with other horses is an ethological need and therefore indispensable in modern husbandry systems. Ethological needs are so important for the animal that husbandry systems that lack the possibilities to execute such behaviours will cause chronic stress. It is concluded that all horses need physical social contact, and that horses, which lack appropriate social learning experiences during ontogeny, may be hampered in their social functioning later in life. Solutions for problems, including dominance problems, in individual social housing and group housing are presented.  
  Abstract Feral horses are social animals, which have to rely on survival strategies centered on the formation of cohesive social bonds within their bands. Many problems in the husbandry of social animals such as horses, are due to the fact that the limits of their adaptive abilities are exceeded. Evidence suggests that the fundamental social characteristics of domestic horses have remained relatively unchanged. The social structure, social strategies and social interactions were investigated (3 non-consecutive years, 24 hr per day for several weeks) in long term established groups of domestic horses (mares and geldings of all ages) and a few small introduced groups, kept in (semi)natural environments. The general aim was to investigate the social needs of domestic horses. The social life of domestic horses was characterised by long lasting bonds with preferred partners which were established and maintained by allogrooming, play, proximity and dominance behaviours. Bonding partners were mainly found within the same sex-age group, but adult geldings also bonded with sub-adult mares and geldings. Adult mares were clustered in a group, while the other animals formed a second group. Among the adult mares, subgroups according to reproductive state were formed. Individuals regulated their social network by interfering with interactions between other members of the herd, which in itself is complex. An intervention is a behavioural action of one animal that actively interferes with an ongoing interaction between a dyad with the apparent aim of altering that interaction. This was verified by post-hoc analyses of disturbed and undisturbed interactions. Interventions in allogrooming or play were performed significantly more often when at least one member of the initial dyad was a preferred partner of, or familiar to (within the small introduced bands) the intervener. The stronger the preferred association in allogrooming between the intervener and member(s) of the initial dyad, the higher the probability the intervener would displace one initial member and continue allogrooming with the other. Just five behaviours were extracted which reliably reflected the dominance relations among horses. Aggression with the hind quarters was used both offensively and defensively and therefore not suitable as a reliable parameter. Individual dominance relationships were related to social experience. The implications of these findings for horse husbandry were assessed. It is argued that the execution of affiliative behaviours may be rewarding in itself, and therefore always will be a highly motivated behaviour. It is shown that social positive physical interactions (allogrooming, play) with other horses is an ethological need and therefore indispensable in modern husbandry systems. Ethological needs are so important for the animal that husbandry systems that lack the possibilities to execute such behaviours will cause chronic stress. It is concluded that all horses need physical social contact, and that horses, which lack appropriate social learning experiences during ontogeny, may be hampered in their social functioning later in life. Solutions for problems, including dominance problems, in individual social housing and group housing are presented.  
  Address  
  Corporate Author Thesis Ph.D. thesis  
  Publisher Place of Publication Universiteit Utrecht 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 2370  
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Author Ahrendt, L.P.; Labouriau, R.; Malmkvist, J.; Nicol, C.J.; Christensen, J.W. url  doi
openurl 
  Title (down) 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 Valderrabano-Ibarra, C.; Brumon, I.; Drummond, H. url  doi
openurl 
  Title (down) Development of a linear dominance hierarchy in nestling birds Type Journal Article
  Year 2007 Publication Animal Behaviour. Abbreviated Journal Anim. Behav.  
  Volume 74 Issue 6 Pages 1705-1714  
  Keywords agonistic behaviour; blue-footed booby; dominance; hatch asynchrony; hierarchy; Sula nebouxii; trained winning  
  Abstract Theoreticians propose that trained winning and losing are important processes in creating linear animal dominance hierarchies, and experiments have shown that both processes can occur in animals, but their actual roles in creating natural hierarchies are unknown. We described agonism in 18 broods of three blue-footed boobies, Sula nebouxii, a species for which trained winning and losing have been demonstrated, to infer how these processes generate and maintain a natural hierarchy. Ranks in the linear hierarchy that emerged in every brood were initially assigned by asymmetries in age, size and maturity, which led to differences between broodmates in levels of expressed and received aggression and, consequently, to differences in the training of their aggressiveness and submissiveness. Later, ranks appeared to be maintained by the chicks' acquired aggressive and submissive tendencies combined with ongoing effects of persisting differences in size and maturity. Our results suggest that trained winning and trained losing are important in the construction of booby hierarchies but that these two axes of learning are largely independent. Increase in submissiveness occurs over a period of about 10-20 days, and the level of submissiveness reached varies with the amount of aggression received. After training, submissiveness is apparently maintained by a lower level of aggression and increasing use of threats. Threats become increasingly effective as chicks age, but are never as effective as attacks.  
  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 4318  
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