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1.
Feather pecking is an abnormal behaviour where laying hens peck the feathers of conspecifics, damaging the plumage or even injuring the skin. If it occurs in a flock, more and more birds show it within a short period of time. A possible mechanism is social transmission. Several studies have shown that laying hen chicks, Gallus gallus domesticus, are able to modify their own behaviour when observing the behaviour of other chicks, for example, when feeding and foraging. As there is good experimental evidence that feather pecking originates from foraging behaviour, we hypothesized that feather pecking could also be socially transmitted. To test this, we reared 16 groups of 30 chicks. After week 4, the birds were regrouped into 16 groups of 20 chicks into each of which we introduced either five chicks that showed high frequencies of feather pecking or, as controls, five chicks that had not developed feather pecking. We then determined the feather-pecking rate and the frequency of foraging, dustbathing, feeding, drinking, preening, moving, standing and resting of all birds in a group. Data from the introduced birds were analysed separately and excluded from the group data. Chicks in groups with introduced feather-pecking chicks had a significantly higher feather-pecking rate than chicks in the control groups. In addition, birds in groups with introduced feather peckers showed significantly lower foraging frequencies than those in the control groups, although the housing conditions were identical and there were no differences in either the number or the quality of the stimuli relevant to foraging behaviour. The study therefore suggests that feather pecking is socially transmitted in groups of laying hen chicks. Copyright 2000 The Association for the Study of Animal Behaviour.  相似文献   

2.
The objective of this experiment was to investigate the effect of rearing density on pecking behaviour and plumage during rearing and throughout the laying period in aviaries. Chicks were reared on sand at high (H; 13 m−2) or low (L; 6.5 m−2) density, in four rearing pens of 390 chicks and eight pens of 195 chicks, respectively, each pen measuring 30 m2. Proportions of chicks per pen performing various types of pecking behaviour were recorded by scan sampling during 16 observation bouts in each rearing pen at 6 weeks of age and during 24 observation bouts at 12 weeks. Individual body weights and plumage condition were recorded. Later, these pullets were housed at 17 hens m−2 in Tiered Wire Floor (TWF; 3 H and 3 L pens of 275 hens) and Laco-Volétage (2 H and 2 L pens of 275 hens) aviaries. At 35 weeks, two samples of eight hens from each aviary pen were observed for pecking behaviour in a test pen. Throughout the laying period, additional records were collected on pecking behaviour, body weight, plumage condition, egg production, and mortality. The L birds had better plumage condition at 6 weeks of age and throughout the laying period. These birds also ground pecked more frequently than H birds during rearing and the laying period. At 12 weeks, L birds feather pecked less than H birds, but no relationship was found between rearing density and feather-pecking behaviour during the laying period. Although TWF hens feather pecked more frequently than Volétage hens, there was no interaction between rearing density and type of aviary for the various pecking behaviours.  相似文献   

3.
Feather pecking is a behavioural disorder in laying hens which consists of pecking the feathers of conspecifics, causing feather damage or even injuries to the skin. Its development can be explained by redirection of foraging behaviour. While the occurrence of feather pecking strongly depends on the kind of housing condition, it is also known that there are strain differences in the tendency to feather peck. From the inverse relation between feather pecking and foraging behaviour found earlier, we hypothesised that genetically determined differences in foraging behaviour could be responsible for the observed differences in feather pecking between strains.In a first experiment we tested whether there are differences in the foraging behaviour of two hybrids. As hybrids, we used Lohman selected leghorn (LSL) and Dekalb; eight groups of 20 1-day old chicks each. They were kept in enriched pens (265cmx90cm) with a litter area (200cmx90cm) consisting of wood-shavings, chaff, straw, polystyrene blocks, sand area (65cmx90cm) and elevated perches. Behavioural observations were carried out in week 4. In a subsequent experiment with the same birds we tested how the foraging behaviour of the two hybrids differed when housing conditions were changed from enriched to restricted and to what extent they developed feather pecking. A 2x2 factorial design with hybrid (LSL, Dekalb) and housing condition (restricted, enriched) as factors and with four replicates of each factor combination was used. Half of the pens of each hybrid were changed from enriched to restricted housing conditions by covering the litter area with slats. Behavioural observations were carried out in weeks 5 and 6.In experiment 1, LSL and Dekalb spent the same amount of time foraging, but Dekalb spent significantly more of that time with pecking and hacking at the polystyrene blocks. On the other hand, LSL spent significantly more time at the feeding troughs and rested significantly less than Dekalb. In the restricted environment of experiment 2, again, the total foraging time did not differ between hybrids, but LSL chicks spent significantly less time scratching, while Dekalb spent significantly more time moving. Both hybrids developed feather pecking but LSL showed significantly higher rates than Dekalb.Our results demonstrate genetic differences in the foraging behaviour and in the way hybrids cope with the change in housing condition from enriched to an environment that is restricted in relation to foraging possibilities. We conclude that the results support the hypothesis put forward that genetic differences in foraging behaviour could be the basis for the genetic influence in the development of feather pecking.  相似文献   

4.
It has been proposed that chicks acquire substrate preferences during an early 'sensitive' period. If a suitable substrate is absent during this period birds may develop alternative preferences for pecking at feathers. The aim of this study was to examine whether early substrate exposure has durable effects on the subsequent behaviour of adult hens. The effects of duration of substrate exposure, substrate change, age at exposure and time since exposure on adult bird behaviour were examined. From days 1 to 210, 144 laying strain birds were housed in pairs in pens with wire floors. The floors were replaced with solid floors covered in wood shavings at different ages and for different durations by allocation to 1 of 12 treatments. Adult birds that had never experienced shavings performed significantly more feather pecking than birds in any other treatment group. Thus, exposure to shavings, even for the minimum exposure duration of 10 days, was protective. However, current substrate was of great importance and adult birds housed on shavings performed significantly more ground pecking and less feather pecking than birds on wire, regardless of previous experience. From day 211 all hens were given shavings or straw, presented alternately for five 24h sessions over 10 consecutive days. Birds foraged on both substrates and their foraging behaviour was not influenced by previous experience. Dustbathing occurred primarily on shavings and was significantly influenced by the age at which birds had previously been exposed to shavings. Dustbathing on shavings was fairly constant throughout the 10-day test period in all groups, suggesting that relatively stable preferences had developed. A secondary 'sensitive period' for the formation of adult dustbathing substrate preference may have superseded the early 'imprinting' process. However, adult behaviour was generally flexible and strongly influenced by current substrate.  相似文献   

5.
Feather pecking is a problem in commercial laying hens, particularly in loose-housing systems, where many hens can be affected by only a few feather peckers. In addition, feather pecking can become an even larger problem if it spreads throughout the flock. There are several possible ways that feather pecking may spread. The simplest way is that one hen may damage the feathers of a hen, and another hen may find the damaged feathers an attractive pecking target. The aim of this experiment was to determine if damaged feathers were feather-pecked more than undamaged feathers on the same body area, and to determine whether some types of feather-body area manipulations were preferred over others as pecking stimuli. Manipulations involved damaging the feathers on the rump, tail or belly of different hens, with two or three levels of severity of manipulation at each body area. Sixteen groups of 11 Lohmann Brown hens between 26 and 28 weeks were observed with the recipient, the feather pecker and the body area that was pecked all being recorded. The feather pecks were classified separately as either gentle or severe. Damaged feathers received significantly more severe feather pecks than undamaged feathers. There were also more gentle feather pecks to damaged feathers, although this did not reach statistical significance. The feather-body area manipulations that received the greatest number of severe feather pecks were the tail feathers when they were cut very short, the rump feathers when they were trimmed, and the rump when feathers were removed. These results support the suggestion that feather pecking does indeed spread through flocks by damaged feathers becoming an attractive target for feather-pecking behaviour. An unexpected result of performing the feather manipulations was an outbreak of cannibalism in half of the experimental groups. Even though there was no visible damage to the skin of the hens after having the feathers manipulated, 13 of the 16 attacked hens were wounded on the part of the body where the feathers had been damaged in some way.  相似文献   

6.
Reaction to frustration of high (HFP) and low feather pecking (LFP) laying hens was investigated. From a HFP- and a LFP-line five birds with a HFP- and five birds with a LFP-phenotype were selected. Birds from the HFP-line were expected to show more key pecking and covered feeder pecking during frustration than birds from the LFP-line. When a bunch of feathers was presented, birds with a HFP-phenotype were expected to redirect their pecks at the bunch. Birds were trained to peck a key for a food reward in an automated Skinnerbox and subjected to two sessions: a control session, where food was available, and a frustration session, where the feeder was covered with Perspex. These two sessions were repeated in the presence of a bunch of feathers. Unexpectedly, birds from the LFP-line had a stronger reaction to frustration than birds from the HFP-line, expressed in pecking behaviour. When a bunch of feathers was offered, birds with a HFP-phenotype did not show more bunch pecking during frustration than birds with a LFP-phenotype.  相似文献   

7.
New housing systems for commercial egg production, furnished cages and non-cage systems, should improve the welfare of laying hens. In particular, thanks to the presence of a litter area, these new housing systems are thought to satisfy the dust-bathing motivation of hens more than in conventional cages, in which no litter area is present. However, although apparently obvious, there is no concrete evidence that non-cage systems, particularly aviaries, satisfy hens' motivation to dust-bathe and thus improve hens' welfare in terms of dust-bathing behaviour. The aim of this study was to compare hens' dust-bathing motivation when housed for a long time under similar conditions to commercial conditions in laying aviaries (with litter) and in conventional cages (without litter). Three treatments were compared: hens reared in floor pens then housed in conventional cages, hens reared in furnished floor pens then housed in a laying aviary, and hens reared in rearing aviaries then housed in a laying aviary. All three treatments provided access to litter during the rearing period. After transfer to the laying systems, access to litter was maintained for the aviary hens but stopped for the cage hens. Twelve groups of four hens per treatment were tested 36 to 43 weeks after transfer. The hens were placed in sawdust-filled testing arenas, and latency to dust-bathe, duration and number of dust baths, and number of hens dust-bathing were recorded. Latency to dust-bathe was shorter, dust baths were longer and more numerous and more hens dust-bathed among cage hens than among aviary hens. Our results indicate that hens' motivation to dust-bathe was more satisfied in laying aviaries than in conventional cages. Thus, laying aviaries improve hens' welfare in term of dust-bathing behaviour compared with conventional cages.  相似文献   

8.
Feather eating has been associated with feather pecking, which continues to pose economic and welfare problems in egg production. Knowledge on feather eating is limited and studies of feather eating in commercial flocks of laying hens have not been performed previously. Therefore, the main objective was to investigate feather eating and its association with plumage damage and floor feather characteristics in commercial flocks of layers in barn and organic production systems. The study was performed in 13 flocks of barn layers and 17 flocks of organic layers. Each flock was visited at around 32 and 62 weeks of age. During both visits, the plumage condition was assessed and the density of floor feathers recorded. In week 62, droppings and floor feathers were collected. Droppings were examined for presence of feather content, whereas length, downiness and pecking damage were recorded for each floor feather. In week 62, a higher prevalence of hens with poor plumage condition was found in barn (22.2%) compared with organic production systems (7.4%; P<0.001), but the prevalence of droppings with feather content did not differ between the two production systems (8.5% in barn v. 4.3% in organic; P=0.99). Our hypothesis about a positive correlation between feather eating and plumage damage was not supported as no correlation was found between the prevalence of poor plumage condition and the prevalence of droppings with feather content. However, the prevalence of pecking damaged floor feathers was positively correlated both with prevalence of droppings with feather content (P<0.05) and poor plumage condition (P<0.01), indicating a possible association between feather eating and feather pecking. In conclusion, it was confirmed that feather eating occurs on-farm, but feather eating was only found to be positively correlated to the number of floor feathers with pecking damage and not as expected to the prevalence of plumage damage. More research is needed into the sources from where feathers are selected for ingestion, that is, whether they are picked from the floor litter, plucked directly from other hens or dislodged during preening of own feathers.  相似文献   

9.
The effect of rearing conditions on feather pecking and reaction to frustration was studied in two lines of laying hens. From commercial rearing conditions (large group, no mother hen), seven birds from a high feather pecking line (HC birds) and eight birds from a low feather pecking line (LC birds) were used. From semi-natural rearing conditions (small group, mother hen present) seven birds from the high feather pecking line (HN birds) were used. Feather pecking behaviour of HC, LC, and HN groups was recorded for 30 min. After that, each bird was food deprived and trained to peck a key for a food reward in a Skinnerbox. After training, each bird was subjected to a frustration session in a Skinnerbox, where the feeder was covered with Perspex. Three HC birds showed severe feather pecking, compared with one HN bird and zero LC birds. Differences in reaction to frustration were found between birds from different lines, but not in birds from different rearing conditions. LC birds tended to put their head in the feeder more frequently than HC birds over all sessions. Although limited, this study indicates that rearing conditions influence feather pecking, but not reaction to frustration.  相似文献   

10.
The aim of this experiment was to study the relationship between feather pecking and ground pecking in laying hens and the effect of group size on feather pecking behaviour. Hisex White hens were kept in floor pens in group sizes of 15, 30, 60 and 120 birds, each with four replicates. Behavioural observations were performed at four different ages and focused on the number of feather pecks and aggressive pecks, both given and received. The part of the body pecked and the location of the bird was recorded as well as the number of pecks made to the floor, feeder and drinker.The results showed that most feather pecking activity occurred in the largest group size (120 birds) and there was some evidence of an increasing frequency of aggressive pecks with increasing group size. The parts of the body which were targets for feather pecking varied depending on the location of the bird giving the peck and the bird receiving it. When looking at the behaviour of individuals, birds doing a lot of feather pecking also showed more ground pecking.  相似文献   

11.
The aim of this review is to discuss the effects of selection method and early-life history on the behavioural development of laying hens. Especially in larger groups, laying hens often develop damaging behaviours, such as feather pecking and cannibalism, leading to impaired animal welfare. We hypothesise that the propensity to develop feather pecking and cannibalism is affected by a bird's genetic background and by its early-life history. The genetic background can be influenced by genetic selection. Laying hens are traditionally selected on individual performance, which may lead to co-selection of feather pecking and cannibalism. For hens kept in small groups, it has recently been demonstrated that a novel group selection method, focusing on group performance, can help to reduce cannibalism. However, the biological background behind the success of group selection is unknown. It is also not known whether these results from small groups can be translated to larger groups of laying hens. Regarding early-life history, laying, brooding and rearing conditions have been shown to have major effects on behavioural development and on feather pecking and cannibalism. The presence of a hen during rearing has been shown to improve foraging- and social behaviour, to decrease feather pecking and to decrease fearfulness in chicks. Applying group selection and rearing laying hens in a more natural environment may be key factors in solving the problems caused by feather pecking and cannibalism, especially if the promising results of group selection from small groups in experimental settings can be translated to large-group housing systems.  相似文献   

12.
Stressful environmental conditions affect the adrenocortical function of developing animals, which can have consequences for their fitness. Discovery of the avian stress hormone corticosterone (CORT) in feathers has the potential to broaden the application of endocrine research in ecological and evolutionary studies of wild birds by providing a long‐term measure of CORT secretion. Mechanisms of CORT deposition in feathers are not well known and few studies have related feather CORT to circulating plasma CORT during feather growth. Our objective was to experimentally test the validity of using feather CORT as a measure of CORT secretion in developing birds experiencing nutritional stress. Caspian tern Hydroprogne caspia chicks were fed ad libitum or restricted (35% less than ad libitum) diets for four weeks. We measured CORT in feathers from these chicks to examine the relationship between feather CORT concentrations and nutritional limitation, circulating plasma CORT, and feather development. We found that feather CORT was higher in controls fed ad libitum than in restricted individuals, despite higher levels of plasma CORT in restricted chicks compared to controls. Feather mass and growth rates were strongly and positively related to feather CORT concentrations in both treatments. This is the first experimental study to show that feather CORT concentrations can be lower in response to nutritional stress, even when plasma CORT concentrations are elevated. Our results indicate that CORT deposition in feathers may be confounded when feather mass and growth rates are compromised by nutritional stress. We conclude that feather CORT can be used for assessing nutritional stress in growing birds, but the direction of response depends on how strongly stress affects feather development.  相似文献   

13.
14.
B Riedstra 《Animal behaviour》2004,67(6):1037-1042
Recently we proposed that early feather pecking is a form of social exploration. Social recognition, important for exploration, is a lateralized function in the domestic chick. Lateralization of functions can be influenced by light exposure late in embryonic development. Therefore, we investigated whether this light exposure affected early posthatching feather-pecking behaviour in domestic chicks, Gallus gallus domesticus. White leghorn embryos either were exposed to light or remained in darkness in the last week of incubation. After hatching, they were housed in groups of two light-exposed and two dark-incubated chicks. Light-exposed chicks showed more feather pecking than did their dark-incubated cagemates. Dark-incubated chicks preferred to direct feather pecks to unfamiliar peers than to familiar peers; light-exposed chicks showed no preference. These effects were present in the first week after hatching and remained at least another 3 weeks. These results support the hypothesis that early gentle feather pecking is part of the normal behavioural repertoire of young chicks and influences social exploration. We discuss a possible mechanism underlying these results. We also suggest that it may be worthwhile not to expose embryos to light during the last week of incubation when housing hatchlings in commercial conditions, where feather pecking is a serious problem.  相似文献   

15.
Two experiments examined the responses of 16 individually caged laying hens to the presentation of feathers plucked from dead birds of the same genetic line. In the first experiment, hens known from a previous experiment to be either feather 'peckers' or 'non-peckers' (8 of each) were tested for their propensity to eat feathers in four 10min trials, in which they were offered fresh semiplumes measuring 4-6cm (length), one at a time, in front of their cage. Wide variation between birds was observed in numbers of feathers eaten, pecked, picked-up and manipulated. Fourteen out of 16 birds readily ate presented feathers on one or more occasion and both birds that ate no feathers were non-peckers. Peckers ate, picked-up and manipulated feathers significantly more often than did non-peckers (P<0.05, P<0.01 and P<0.01, respectively). A second experiment investigated the possibility that presence of preen (uropygial) oil might contribute to the attractiveness of feathers to eat. The same group of 16 pecker and non-pecker hens were offered a choice between 20 washed and 20 unwashed semiplumes, presented simultaneously in separate containers, in two 10min trials. Unwashed feathers were eaten, pecked and picked-up in preference to washed feathers by both peckers and non-peckers (P<0.05, P<0.01, and P<0.01, respectively), indicating an attraction towards unwashed feathers, or an avoidance of washed feathers for some reason. Peckers and non-peckers did not differ significantly in their preferences. These results provide evidence of a relationship between feather eating and feather pecking at an individual level. The finding that hens could distinguish between normal feathers and those treated in such a way as to alter their olfactory (but not visual) properties suggests olfactory cues may be of importance in determining the attractiveness of conspecific feathers.  相似文献   

16.
With a view to alleviate the feeling of hunger in broiler breeders, different types of fibre sources were used in high-fibre diets to increase feed quantity while limiting growth to industry recommended levels. Using scatter feeding, three diets (C1: commercial control diet, 1 × fibre content, 80% insoluble fibre (ISF); H2: 2 × fibre content, 89% ISF; and L2: 2 × fibre content, 71% ISF) were each fed to 10 groups of 16 broiler breeder chickens. Similar growth rates were obtained on different quantities of food with all birds reaching commercial target weight at 15 weeks of age. In a hunger test, birds fed C1 ate significantly faster and showed a higher compensatory feed intake than birds on diets H2 and L2, indicating that the two high-fibre diets did reduce the level of hunger experienced by the birds. Behavioural observations carried out at 14 weeks of age showed high levels of tail pecking in birds fed C1 and almost none in birds fed L2, whereas birds fed H2 were intermediate. Stereotypic pecking at fixtures was seen twice as frequently in birds fed C1. Birds on diet L2 displayed behavioural signs indicative of discomfort, and the high water usage on this diet created problems with litter quality. Birds on diet H2 continued to show foraging behaviour throughout the day, and were more frequently engaged in dust bathing and other comfort behaviour. This experiment indicates that high-fibre diets can alleviate the feeling of hunger currently experienced by broiler breeders, and a high ratio of ISF may improve the well-being of the birds.  相似文献   

17.
Carotenoid-based plumage coloration of birds has been hypothesized to honestly reflect individual quality, either because carotenoids are difficult to acquire via food or because of a trade-off in allocation of carotenoids between maintenance and signaling functions. We tested whether differential foraging ability is a necessary precondition for maintaining individual differences in carotenoid-based plumage coloration in male greenfinches (Carduelis chloris). Wild-caught birds were brought into captivity, where half of them were supplemented with carotenoids while the other half was maintained on a carotenoid-poor diet. Color of the yellow parts of tail feathers, grown under natural conditions, was compared with that of the replacement feathers, grown in captivity. Carotenoid supplementation increased feather chroma (saturation). Color of wild-grown feathers significantly correlated with the color of lab-grown feathers. This result demonstrates the existence of a significant component of variation in carotenoid coloration, which reflects physiological qualities or genetic differences among individuals independent of foraging ability. Among both experimental groups, plasma carotenoid concentration during feather growth strongly correlated with chroma of the feathers grown in captivity. This indicates that carotenoid-based plumage coloration can reveal circulating carotenoid levels over a very wide range of concentrations, suggesting the ample signaling potential of such a mechanism.  相似文献   

18.
Feather pecking is a serious economic and welfare problem in laying hens. Feather damage occurs mainly through severe feather pecking (SFP). Selection experiments have proved that this behavior is heritable and lines have been divergently selected for high (HFP) and low feather pecking (LFP). The number of bouts of SFP per hen follows a Poisson distribution with a maximum nearby 0. A few studies indicate that the distribution within flocks is not homogenous but contains sub-groups of birds showing extremely high levels of feather pecking (EFP). It was the aim of the current study to re-analyze data on SFP of lines selected for HFP/LFP and their F2 cross so as to uncover hidden sub-populations of EFP birds. Data of seven selection generations of HFP and LFP selection lines as well as their F2 cross have been used. We fitted a two-component mixture of Poisson distributions in order to separate the sub-group of EFP from the remaining birds. HFP and LFP lines differed mainly in mean bouts per bird. The proportion of EFP was only marginal in the LFP as compared with the HFP and the F2 population. Selection for LFP did not result in total elimination of EFP. The presence of even small proportions of EFP may play an important role in initiating outbreaks of feather pecking in large flocks. Further studies on feather pecking should pay special attention to the occurrence of EFP sub-groups.  相似文献   

19.
Feather and skin pecking leading to feather loss and tissue damage is a welfare problem in commercial turkeys. A factorial experiment was designed to compare a line of unselected turkeys that previously did not exhibit this behaviour with a genetically selected male line with a propensity for damaging pecking. We also compared the time budgets of male and female turkeys from the two lines at 3, 6 and 9 weeks of age and at different times of the day. Damaging feather pecking occurred in 32% of male and 15% of female male line turkeys but was not observed in the traditional line. Time budgets of males and females were similar. Preening increased and resting, feeding and general pecking declined with age. Resting was higher in the afternoon than in the morning and male line turkeys were observed resting more often than traditional turkeys at 3 and 6 weeks. Traditional turkeys displayed more gentle feather pecks than male line poults at 9 weeks. Male turkeys of both lines showed more strong feather pecks and pulls at 3 weeks whereas female turkeys showed more at 9 weeks of age. The frequency of strong feather pulls recorded electronically increased with age but was not related to mortality. It is suggested that damaging feather pecking in turkeys may be the result of vigorous investigative pecking.  相似文献   

20.
Growing feathers and mounting immune responses are both energetically costly for birds. According to the life history trade‐off hypothesis, it has been posited that the costs of feather growth lead to temporal isolation between molt and other expensive activities, reproduction for example. In contrast to life cycle events, the need to mount an immune response can occur at any time, including during feather growth. Thus, we hypothesized that mounting an immune response during feather growth may divert energy and resources from feather growth and impair feather renewal. To test this hypothesis, we clipped or plucked the same feathers of male house sparrows Passer domesticus biblicus. In the clipped group (n = 16), the feathers were absent with no regrowth; in the plucked group (n = 14), feathers were absent and regrowth was initiated. We also had an intact control group of 15 sparrows. We then initiated an inflammatory immune response by subcutaneous injection over the left breast muscle of the birds with a lipopolysaccharide (LPS) and quantified behavioral and physiological responses. We predicted that sparrows with plucked feathers would incur the highest energetic costs while mounting an immune response, and would increase their foraging effort to offset this cost. We found no difference in body mass and resting metabolic rates among sparrows subjected to the different feather and immune treatments. However, we did find that while sparrows with plucked feathers increased foraging efficiency following the immune challenge by paying fewer but longer visits to the food tray, allowing them to maintain food consumption. Foraging efficiency in sparrows with clipped feathers was reduced. We also found that quality of newly grown feathers after the immune challenge was poorer in the plucked group, suggesting that mounting an immune response competes with feather growth for resources.  相似文献   

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