Records |
Author |
Ohtsuki, H.; Iwasa, Y.; Nowak, M.A. |
Title |
Indirect reciprocity provides only a narrow margin of efficiency for costly punishment |
Type |
Journal Article |
Year |
2009 |
Publication |
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Abbreviated Journal |
Nature |
Volume |
457 |
Issue |
7225 |
Pages |
79-82 |
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Abstract |
Indirect reciprocity1, 2, 3, 4, 5 is a key mechanism for the evolution of human cooperation. Our behaviour towards other people depends not only on what they have done to us but also on what they have done to others. Indirect reciprocity works through reputation5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 15, 16, 17. The standard model of indirect reciprocity offers a binary choice: people can either cooperate or defect. Cooperation implies a cost for the donor and a benefit for the recipient. Defection has no cost and yields no benefit. Currently there is considerable interest in studying the effect of costly (or altruistic) punishment on human behaviour18, 19, 20, 21, 22, 23, 24, 25. Punishment implies a cost for the punished person. Costly punishment means that the punisher also pays a cost. It has been suggested that costly punishment between individuals can promote cooperation. Here we study the role of costly punishment in an explicit model of indirect reciprocity. We analyse all social norms, which depend on the action of the donor and the reputation of the recipient. We allow errors in assigning reputation and study gossip as a mechanism for establishing coherence. We characterize all strategies that allow the evolutionary stability of cooperation. Some of those strategies use costly punishment; others do not. We find that punishment strategies typically reduce the average payoff of the population. Consequently, there is only a small parameter region where costly punishment leads to an efficient equilibrium. In most cases the population does better by not using costly punishment. The efficient strategy for indirect reciprocity is to withhold help for defectors rather than punishing them. |
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Macmillan Publishers Limited. All rights reserved |
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0028-0836 |
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10.1038/nature07601 |
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no |
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Equine Behaviour @ team @ |
Serial |
4705 |
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Author |
Harrison, S.A.; Tong, F. |
Title |
Decoding reveals the contents of visual working memory in early visual areas |
Type |
Journal Article |
Year |
2009 |
Publication |
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Abbreviated Journal |
Nature |
Volume |
458 |
Issue |
7238 |
Pages |
632-635 |
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Abstract |
Visual working memory provides an essential link between perception and higher cognitive functions, allowing for the active maintenance of information about stimuli no longer in view1, 2. Research suggests that sustained activity in higher-order prefrontal, parietal, inferotemporal and lateral occipital areas supports visual maintenance3, 4, 5, 6, 7, 8, 9, 10, 11, and may account for the limited capacity of working memory to hold up to 3–4 items9, 10, 11. Because higher-order areas lack the visual selectivity of early sensory areas, it has remained unclear how observers can remember specific visual features, such as the precise orientation of a grating, with minimal decay in performance over delays of many seconds12. One proposal is that sensory areas serve to maintain fine-tuned feature information13, but early visual areas show little to no sustained activity over prolonged delays14, 15, 16. Here we show that orientations held in working memory can be decoded from activity patterns in the human visual cortex, even when overall levels of activity are low. Using functional magnetic resonance imaging and pattern classification methods, we found that activity patterns in visual areas V1–V4 could predict which of two oriented gratings was held in memory with mean accuracy levels upwards of 80%, even in participants whose activity fell to baseline levels after a prolonged delay. These orientation-selective activity patterns were sustained throughout the delay period, evident in individual visual areas, and similar to the responses evoked by unattended, task-irrelevant gratings. Our results demonstrate that early visual areas can retain specific information about visual features held in working memory, over periods of many seconds when no physical stimulus is present. |
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Macmillan Publishers Limited. All rights reserved |
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0028-0836 |
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10.1038/nature07832 |
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Equine Behaviour @ team @ |
Serial |
4944 |
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Author |
Clutton-Brock, T. |
Title |
Cooperation between non-kin in animal societies |
Type |
Journal Article |
Year |
2009 |
Publication |
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Abbreviated Journal |
Nature |
Volume |
462 |
Issue |
7269 |
Pages |
51-57 |
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Abstract |
Explanations of cooperation between non-kin in animal societies often suggest that individuals exchange resources or services and that cooperation is maintained by reciprocity. But do cooperative interactions between unrelated individuals in non-human animals really resemble exchanges or are they a consequence of simpler mechanisms? Firm evidence of reciprocity in animal societies is rare and many examples of cooperation between non-kin probably represent cases of intra-specific mutualism or manipulation. |
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Macmillan Publishers Limited. All rights reserved |
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0028-0836 |
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Notes |
10.1038/nature08366 |
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no |
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Equine Behaviour @ team @ |
Serial |
5270 |
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Author |
Zharkikh, T.L.; Andersen, L. |
Title |
Behaviour of Bachelor Males of the Przewalski Horse (Equus ferus przewalskii) at the Reserve Askania Nova |
Type |
Journal Article |
Year |
2009 |
Publication |
Der Zoologische Garten |
Abbreviated Journal |
Zoologische Garten |
Volume |
78 |
Issue |
5-6 |
Pages |
282-299 |
Keywords |
Equid; Przewalski horse (Equus ferus przewalskii); Bachelor group; Social behaviour |
Abstract |
The aim of this study was to investigate social relationships between Przewalski horses at a high density in a bachelor group housed in a 3.5-ha enclosure. The group consisted of 16 males aged 5 to 16. Behavioural data were collected during 18 days, total 216 h. Fifteen minute focal animal sampling was used; each horse was observed three times a day for a total of 45 min. The occurrence of 25 behaviours was recorded, and group spacing behaviour was studied using nearest neighbour recordings. The group divided into four subgroups; this supports earlier findings of bachelor groups (n>=10) dividing into two or more subgroups if they included several males aged >5 years. The total frequency of social interactions was 14.6±1.1 h-1. Although the density of the group in this study was higher than in other zoos, the males interacted agonistically only 3.6 h-1. The most frequently observed social behaviour categories were friendly interactions. This study shows possibilities to use some investigative behaviours (marking, flehmen, olfactory investigation, etc.) as indicators of social status of animals in a group. |
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0044-5169 |
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Equine Behaviour @ team @ |
Serial |
5098 |
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Author |
Whishaw, I.Q.; Sacrey, L.-A.R.; Gorny, B. |
Title |
Hind limb stepping over obstacles in the horse guided by place-object memory |
Type |
Journal Article |
Year |
2009 |
Publication |
Behavioural Brain Research |
Abbreviated Journal |
Behav. Brain. Res. |
Volume |
198 |
Issue |
2 |
Pages |
372-379 |
Keywords |
Horse place memory; Memory and hind limb stepping; Obstacle memory in horse; Obstacle stepping in horse; Place memory for stepping; Stepping in the horse; Working memory for stepping |
Abstract |
An animal that has stepped over an obstacle with its forelimbs uses a memory of the obstacle to guide the hind limbs so that they also clear the obstacle, even in situations in which long pauses are introduced between forelimb and hind limb stepping. To further clarify the features of hind limb obstacle clearance memory, the present study examined hind limb obstacle clearance in the horse. A rider guided horses over obstacles and paused the horse over obstacles in tests that examined the relationship between forelimb and hind limb stepping, with the following results. First, the horses displayed memory for an obstacle as measured by hind limb lifting over the obstacle for durations lasting as long as 15Â min. The response was not dependent upon ongoing visualization of the obstacle, as limb lifting was unaffected by visual occlusion with blinders, a blindfold, or by removing the obstacle during the pause. Second, previous experience of stepping over an obstacle led to pause-related hind limb lifting at the object's previous location even on trials for which there was no obstacle and so no preceding forelimb lifting. Third, whereas a horse would lift its hind limbs to clear two successively presented obstacles, replacing an obstacle before the horse after the forelimbs had cleared the obstacle prevented subsequent hind limb lifting at the obstacle's previous location. Taken together the results show that hind limb obstacle clearance is guided by a place-object memory. The results are discussed in relation to the differential sensory and memonic control of forelimb and hind limb stepping with the suggestion that place-object memory can guide hind stepping as well as overshadow working memory from front leg stepping. |
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0166-4328 |
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Equine Behaviour @ team @ |
Serial |
4754 |
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Author |
Whistance, L.K.; Sinclair, L.A.; Arney, D.R.; Phillips, C.J.C. |
Title |
Trainability of eliminative behaviour in dairy heifers using a secondary reinforcer |
Type |
Journal Article |
Year |
2009 |
Publication |
Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
Volume |
117 |
Issue |
3-4 |
Pages |
128-136 |
Keywords |
Cattle; Eliminative behaviour; Learning; Clicker training; Clean bedding |
Abstract |
Soiled bedding influences cleanliness and disease levels in dairy cows and there is no evidence of an inherent latrine behaviour in cattle. If cows were trained to use a concrete area of the housing system as a latrine, a cleaner bed could be maintained. Thirteen group-housed, 14-16-month-old Holstein-Friesian heifers, were clicker trained with heifer-rearing concentrate pellets as a reward. Training was carried out in four phases. (Phase 1) Association of feed reward with clicker, criterion: 34/40 correct responses. (Phase 2) Simple task (nose-butting a disc) to reinforce phase 1 association, criterion: 17/20 correct responses. (Phase 3) Association of eliminative behaviour with reward where criterion was four sessions with only one incorrect response: criteria for each heifer in phases 1-3 were set using binomial tests. (Phase 4) Shaping eliminative behaviour to occur on concrete. Possible responses were, eliminating on concrete (C) or straw (S), or moving from one substrate to another immediately before eliminating: C --> S, S --> C. Heifers were rewarded for the desired behaviours C and S --> C and ignored when S and C --> S occurred. If learning was achieved, C should increase as C --> S decreased and S --> C should increase as S decreased: tested with Spearman rank correlations. All heifers achieved criterion by day 4 of phase 1 (P = 0.001); day 1 of phase 2 (P = 0.001) and day 10 of phase 3 (P < 0.009). Responses changed throughout phase 3 beginning with (i) looking at the trainer whilst voiding then moving to trainer after the click, and later including (ii) moving to trainer immediately before- or (iii) during voiding. No relationship was found between S and S --> C (rs = -0.14; P = 0.63) or C and C --> S (rs = -0.33; P = 0.25). All group members eliminated more often on concrete (580) than on straw (141) but four heifers with consistently longer lying bouts also showed more C --> S before lying down (Mann-Whitney, P = 0.007). The present study is believed to be the first reported work to show that cattle can be trained to show an awareness of their own eliminative behaviour. This was not successfully shaped to latrine behaviour, however, and it is suggested that floor type may not have been a sufficiently salient cue. Voiding on straw occurred largely with response C --> S (0.73) and general behaviour suggested that this was strongly linked to lying patterns of individual heifers. |
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0168-1591 |
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Equine Behaviour @ team @ |
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4765 |
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Author |
VanDierendonck, M.C.; de Vries, H.; Schilder, M.B.H.; Colenbrander, B.; Þorhallsdóttir, A.G. and Sigurjónsdóttir, H. |
Title |
Interventions in social behaviour in a herd of mares and geldings |
Type |
Journal Article |
Year |
2009 |
Publication |
Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
Volume |
116 |
Issue |
1 |
Pages |
67-73 |
Keywords |
Horses; Social relationships; Interventions; Mares; Geldings; Play; Allogrooming; Social network |
Abstract |
Social dynamics and maintenance of social cohesion were studied by analysing social interventions in two groups of horses consisting of adult mares, their offspring, adult geldings and sub-adults. The animals were observed for a total of 1316 h. All relevant dyadic and triadic social interactions, including initial behaviour, possible intervention and outcome, were recorded. The main question was: do horses use interventions in affiliative interactions to safeguard their social network? Horses were significantly more likely to intervene in allogrooming or play interactions that involved a preferred partner. The stronger the preferred association in allogrooming, the higher the likelihood the intervener took over allogrooming with an initial dyad member. Interveners originating from two newly introduced groups (n = 3 and 5), intervened significantly more often when a member of their own group allogroomed with an unfamiliar horse. In play, no correlation with unfamiliarity was found. Overall, the intervening horses stopped more than half of the initial allogrooming interactions, and a third of all interactions. Therefore, social facilitation cannot sufficiently explain interference behaviour. This study shows that maintaining relationships with preferred partners is important to horses and has implications for equine husbandry and management. |
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0168-1591 |
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Equine Behaviour @ team @ |
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4766 |
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Author |
Hartmann, E.; Christensen, J.W.; Keeling, L.J. |
Title |
Social interactions of unfamiliar horses during paired encounters: Effect of pre-exposure on aggression level and so risk of injury |
Type |
Journal Article |
Year |
2009 |
Publication |
Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
Volume |
121 |
Issue |
3-4 |
Pages |
214-221 |
Keywords |
Equine; Behaviour; Welfare; Mixing; Housing; Wounds |
Abstract |
Group housing of horses is not widely applied in practice despite the welfare advantages of keeping animals socially rather than individually. In particular, concerns have been raised about the possible increased risk of injury and how to introduce a new horse into an established group. This study investigated two hypotheses: (1) pre-exposure of young horses in neighbouring boxes reduces the frequency of aggressive interactions when the same horses are subsequently put together in a paddock compared to horses without this previous box experience, (2) the occurrence of aggressive behaviour, in particular contact aggression in the paddock can be predicted after observing the horses' social interactions in neighbouring boxes. Danish Warmblood mares (n = 20), 2 years old, were kept in two groups of 10 horses. In total, 60 encounters were arranged whereby each horse was confronted pair-wise with six horses from the other group, three according to each treatment: treatment I--box (B) and subsequent paddock meeting (BP), and treatment II--only paddock meeting (P). Horses met in neighbouring boxes for 5 min and together in the same paddock for 10 min. The frequencies of aggressive and non-aggressive interactions were analysed from video recordings. Total aggression levels between BP and P did not differ, but [`]contact aggression', i.e. bite, kick, strike, push, tended to be lower in BP compared to P (median BP = 1, P = 2; p = 0.083) and there were less bites in BP than P (median BP = 0, P = 1; p = 0.050). Frequencies of [`]non-aggressive' interactions, e.g. friendly approach, nasal sniff, were lower in BP than P (median BP = 2.5, P = 10; p < 0.01). Results further revealed that [`]bite threat' performed in boxes correlated with [`]contact aggression' in the paddock (r = 0.46, p = 0.011). In conclusion, pre-exposure of young horses in neighbouring boxes may reduce [`]contact aggression', especially biting, in the paddock and [`]bite threat' shown in boxes may help to predict contact aggression when horses are later turned out together. The reduced non-aggressive interactions in the paddock in the BP test were probably a consequence of horses having exchanged these behaviours in the preceding B test. Exposing young horses in boxes next to each other may be a helpful tool before mixing them because horses meet in a safe environment that could assist in reducing the type of aggression where horses are most at risk of being injured. |
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0168-1591 |
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Equine Behaviour @ team @ |
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5089 |
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Author |
Nagy, K.; Bodó, G.; Bárdos, G.; Harnos, A.; Kabai, P. |
Title |
The effect of a feeding stress-test on the behaviour and heart rate variability of control and crib-biting horses (with or without inhibition) |
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Journal Article |
Year |
2009 |
Publication |
Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
Volume |
121 |
Issue |
2 |
Pages |
140-147 |
Keywords |
Stereotypic behaviour; Crib-biting; Modified Forssell's operation; Cribbing collar; Equine welfare; Stress coping |
Abstract |
Crib-biting is a form of oral stereotypy affecting 4-5% of horses. Once fixed, crib-biting is difficult to eliminate by behaviour therapy, however, its performance can be inhibited by collar or surgery treatment (modified Forssell's procedure). Although surgical intervention is widespread, the effects on stress coping in horses have not been studied. In the present study we evaluated changes in behaviour response and heart rate variability in 9 control, 10 crib-biting, 10 collar and 11 surgically treated horses in a feeding stress-test, in which a feeding-bowl was placed in front but out of the reach of the horses, from which tidbits were given 3 times. We found that stress triggers high oral activity, mainly cribbing in crib-biting horses, elevates other forms of oral activities in the inhibited groups and does not affect oral activities of controls. Instead of performing oral activities, control horses tended to target an unavailable feeding-bowl by pawing or head-tossing. Changes in stress level were indistinguishable in controls and crib-biters as heart rate variability returned to baseline values in both groups. In contrast, horses inhibited to perform crib-biting showed elevated stress level throughout the test period. Our results suggest that crib-biting may develop to cope with stress, and such coping function diminishes when inhibited. |
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0168-1591 |
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Equine Behaviour @ team @ |
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5091 |
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Jørgensen, G.H.M.; Borsheim, L.; Mejdell, C.M.; Søndergaard, E.; Bøe, K.E. |
Title |
Grouping horses according to gender--Effects on aggression, spacing and injuries |
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Journal Article |
Year |
2009 |
Publication |
Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
Volume |
120 |
Issue |
1-2 |
Pages |
94-99 |
Keywords |
Horse; Group housing; Gender; Injuries; Social behaviour |
Abstract |
Many horse owners tend to group horses according to gender, in an attempt to reduce aggressive interactions and the risk of injuries. The aim of our experiment was to test the effects of such gender separation on injuries, social interactions and individual distance in domestic horses. A total of 66 horses were recruited from 4 different farms in Norway and Denmark and divided into six batches. Within each batch, horses were allotted into one mare group, one gelding group and one mixed gender group, with most groups consisting of three or four animals. After 4-6 weeks of acclimatisation, a trained observer recorded all social interactions using direct, continuous observation 1 h in the morning and 1 h in the afternoon for three consecutive days. Recordings of the nearest neighbour of each horse were performed using instantaneous sampling every 10 min. The horses were inspected for injuries before grouping, day 1 after grouping and after 4-6 weeks. No significant effect of gender composition was found on social interactions (P > 0.05), spacing (P > 0.07) or injuries (P > 0.23). Eighty percent of all aggressive interactions recorded were threats, not involving physical contact. Horses with the smallest space allowance showed the highest mean number of aggressive interactions (28.6 ± 6.1 interactions per 6 h) compared to the mean of all the other batches (8.3 ± 1.0 interactions per 6 h). Very few injuries were found and most were superficial. In conclusion, gender composition does not seem to have any effect on aggression level, spacing or injuries. However, the early social experience of horses, management of feeding and space allowance probably represents more important factors for successful group housing of domestic horses. |
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0168-1591 |
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Equine Behaviour @ team @ |
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5093 |
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