Coccidial infection does not influence preening behavior in American goldfinches

Preening behavior in birds is important for the maintenance of thermoregulatory and ornamental functions of plumage. It has been repeatedly demonstrated that birds trade off time between plumage maintenance and other activities. However, the condition-dependent constraints of preening remain virtually unstudied. Here, we present the first experimental test of the hypothesis that intestinal parasite infection impairs preening activity. We studied male American goldfinches (Spinus tristis), a species with carotenoid-based plumage coloration. Following pre-alternate (spring) molt, we manipulated the health of males by infecting some birds with Isospora spp. coccidia and keeping others free of the infection. Although the goldfinches increased preening throughout the captive period, we found no significant effect of coccidial treatment on preening behavior. The effect of coccidia on plumage maintenance may be more pronounced under natural conditions where birds have limited access to food and engage in more activities that might limit time available for preening.     

Experimental exposure to trace metals affects plumage bacterial community in the feral pigeon

Bacteria are fundamental associates of animals, and recent studies have highlighted their major role in host behaviour, immunity or reproductive investment. Thus, any environmental factor modifying bacterial community may affect host fitness. In birds, trace metals emitted by anthropogenic activities accumulate onto the plumage where they may alter bacterial community and ultimately affect bird fitness. Although trace metals are current major environmental issues in urban habitats, their effects on feather bacterial community have never been investigated. Here, we supplemented feral pigeons Columba livia, an emblematic urban species, with zinc and/or lead in drinking and bath water. As expected, lead and zinc supplementations modified plumage bacterial community composition. Zinc decreased bacterial load, while lead decreased bacterial richness and the frequency of preening behaviour in birds, known to regulate feather bacteria. Our results demonstrate for the first time the effects of common urban trace metals on plumage bacterial community and shed light on one of the mechanisms by which trace metals can affect bird fitness. Further studies are now needed to investigate how this effect modulates avian life history traits known to depend on plumage bacterial community.     

Feather eating and its associations with plumage damage and feathers on the floor in commercial farms of laying hens

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.     

Metabolism of carotenoid pigments in birds

Carotenoid pigments are an important component in the plumage of birds. The metabolic precursors are dietary in origin but many species have the capacity to chemically modify and selectively deposit the pigments. The ensuing plumage patterns are important in communication and identification. The bright yellows, oranges, and reds are due mostly to xanthophylls; keto and hydroxy carotenes. Some are deposited unmodified (e.g., lutein) whereas others are modified chemically (canthaxanthin, astaxanthin). Early workers concentrated on demonstrating that feather carotenoids were derived from the diet and deposited selectively. Progress in defining and solving biological problems depended on advances in chemical and analytical techniques. Subsequent investigation showed that various plumage colormorphs, seasonal plumage changes or colors in common mutant, were due to relatively simple chemical changes in carotenoids but had profound biological consequences. Equally important was the realization that many of these processes were under genetic control. Validation came from feeding studies of flamingos and finches. Recent studies have employed the plumage carotenoids to test hypotheses of genetic divergence, to relate plumage color to environmental process, and to demonstrate the influence of synthetic changes on color. Understanding the processes has advanced with the introduction of high-resolution separation techniques and the ability to determine both conformation and absolute configuration. The next steps will be in the direction of understanding the enzymatic modification, transport, and tissue selectivity of feather carotenoids.     

Genetic,environmental, and neighbor effects on the severity of stereotypies and feather picking in Orange-winged Amazon parrots (Amazona amazonica): An epidemiological study

Stereotypies and psychogenic feather picking represent two distinct forms of abnormal behavior in parrots, with implications for welfare and owner relinquishment. While both behaviors are common in captive parrots, relatively little is known about the factors that may predispose a parrot to their development. To assess the effects of potential social, environmental, and genetic factors, we performed an epidemiological study of a colony of 64 Orange-winged Amazon parrots. We first used mixed model methods to assess the narrow heritability, or genetic determination, of stereotypy, which was −0.08 ± 0.14, and feather picking, which was 1.14 ± 0.27. We then focused our study on one room of 27 birds that had always been housed in the same cage location. We examined the effects of neighbor behavior, number of neighbors, sex, family, age, cage row (beside an aisle or beside a wall), and distance from the door, on each behavior. Stereotypy was negatively correlated with the number of neighbors, but was unaffected by any other factor. In contrast, feather picking was affected by family, was more severe in females, and was progressively more severe in birds housed closer to the door when they had a direct line of sight of the door. Feather picking was spatially clustered in the room, but siblings from particular families were also more likely to be housed as neighbors than were non-siblings. Given the strong effect of family on feather picking and the effect of proximity to the door, these factors together explained the spatial clustering of feather picking. In addition, the feather picking of neighbors was uncorrelated. Thus, there was no evidence for social transmission of feather picking. Our results indicate that cages should be located away from potential stressors, and suggest that there is a potential for breeding against feather picking behavior in parrots. Finally, the markedly different genetic and environmental effects on stereotypies versus feather picking emphasizes that these are two distinct classes of Abnormal Repetitive Behavior in parrots.     

Cohabitation with farm animals rather than breeding effort increases the infection with feather‐associated bacteria in the barn swallow Hirundo rustica

Feather‐associated bacteria are widespread inhabitants of avian plumage. However, the determinants of the between‐individual variation in plumage bacterial loads are less well understood. Infection intensities can be determined by ecological factors, such as breeding habitat, and can be actively regulated by hosts via preening. Preening, yet, is a resource intensive activity, and thus might be traded‐off against reproductive investment in breeding birds. Here, we studied barn swallows Hirundo rustica to assess the bacterial cost of reproduction in relation to nesting site micro‐habitats. Barn swallows prefer to breed in the company of large‐sized farm animals, although the presence of mammalian livestock in barns assures a warm and humid micro‐climate that favours bacterial proliferation. Thus, we experimentally manipulated brood sizes of birds breeding in barns with, or without, farm animals and measured total cultivable bacteria (TCB) and feather‐degrading bacteria (FDB) from the plumage. We found that the abundance of feather‐associated bacteria (i.e. both TCB and FDB) in females, but not males, breeding in barns with livestock were significantly higher than in conspecifics breeding in empty barns. Plumage bacterial loads, however, were not affected by brood size manipulations in either sex. In addition, we report a negative relationship between both TCB and FDB and hatching date in females, and several sex and seasonal differences in plumage bacterial abundances. Our study is the first to show that breeding micro‐habitat (i.e. livestock co‐tenancy) has consequences for the abundance of feather‐associated bacteria.     

Feather mass and winter moult extent are heritable but not associated with fitness-related traits in a long-distance migratory bird

In birds, the allocation of resources to plumage production may have important fitness consequences. However, we have only a limited understanding of how plumage traits respond to natural selection, making it difficult to predict how variation in plumage traits may contribute to the adaptation of birds to environmental change. In this study, we collected plumage-related data in a pedigreed population of a long-distance migratory bird (the Pied Flycatcher Ficedula hypoleuca) to estimate the heritability of two plumage traits: feather mass (as a proxy of feather quality) and the extent of winter moult. We further explored whether these plumage features were associated with some fitness-related traits. Variation in plumage characteristics could be explained by differences in sex, age and year, which indicates a high degree of plasticity in these traits. After controlling for these effects, however, feather mass and winter moult extent were highly repeatable (r = 0.58–0.82) and heritable (h² = 0.59–0.65), suggesting that additive genetic variation accounts for a significant proportion of the residual phenotypic variation of plumage traits in this population. Although the studied characteristics showed evolutionary potential, we did not find any relationship between plumage features and fitness-related traits like spring arrival date, egg-laying date, mating success or mating-time. We conclude that current selection on feather mass and moult extent, if existing, is weak, and that these moult-related traits are currently of minor importance for the adaptation of our study population to global warming.     

Migratory behaviour affects the trade-off between feather growth rate and feather quality in a passerine bird

Migratory birds have less time for moulting than sedentary birds, which may force them to produce their feathers faster at the expense of reducing feather quality. However, the effects of migration on the trade-off between moult speed and plumage quality remain to be studied in natural populations. We analysed the relationship between growth rate and quality of individual feathers, taking advantage of natural variation between migratory and sedentary populations of blackcaps Sylvia atricapilla . As predicted by life-history theory, individual blackcaps showed variable individual quality, which was revealed by positive correlations between feather growth rate and feather mass within populations. However, migrants grew up their feathers faster, producing lighter feathers than sedentary blackcaps. These results support the idea that feather growth rate and feather quality are traded against each other in blackcaps. Such a trade-off is apparently caused by different selection associated to migratory and sedentary life styles, which opens new insights into the diversification of moult patterns in birds.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 98–105.     

A natural experiment on the condition-dependence of achromatic plumage reflectance in black-capped chickadees

Honest advertisement models posit that only individuals in good health can produce and/or maintain ornamental traits. Even though disease has profound effects on condition, few studies have experimentally tested its effects on trait expression and even fewer have identified a mechanistic basis for these effects. Recent evidence suggests that black and white, but not grey, plumage colors of black-capped chickadees (Poecile atricapillus) are sexually selected. We therefore hypothesized that birds afflicted with avian keratin disorder, a condition that affects the beak and other keratinized tissues, would show reduced expression of black and white, but not grey, color. UV-vis spectrometry of black-capped chickadees affected and unaffected by avian keratin disorder revealed spectral differences between them consistent with this hypothesis. To elucidate the mechanistic bases of these differences, we used scanning electron microscopy (SEM), electron-dispersive x-ray spectroscopy (EDX) and a feather cleaning experiment. SEM showed extreme feather soiling in affected birds, and EDX revealed that this was most likely from external sources. Experimentally cleaning the feathers increased color expression of ornamental feathers of affected, but not unaffected, birds. These data provide strong evidence that black and white color is an honest indicator in chickadees, and that variation in feather dirtiness, likely due to differences in preening behavior is a mechanism for this association.     

Adaptive significance of avian beak morphology for ectoparasite control

The beaks of Darwin's finches and other birds are among the best known examples of adaptive evolution. Beak morphology is usually interpreted in relation to its critical role in feeding. However, the beak also plays an important role in preening, which is the first line of defence against harmful ectoparasites such as feather lice, fleas, bugs, flies, ticks and feather mites. Here, we show a feature of the beak specifically adapted for ectoparasite control. Experimental trimming of the tiny (1-2 mm) maxillary overhang of rock pigeons (Columba livia) had no effect on feeding efficiency, yet triggered a dramatic increase in feather lice and the feather damage they cause. The overhang functions by generating a shearing force against the tip of the lower mandible, which moves forward remarkably quickly during preening, at up to 31 timesper second. This force damages parasite exoskeletons, significantly enhancing the efficiency of preening for parasite control. Overhangs longer than the natural mean of 1.6mm break significantly more often than short overhangs. Hence, stabilizing selection will favour overhangs of intermediate length. The adaptive radiation of beak morphology should be re-assessed with both feeding and preening in mind.     

Bh (black at hatch) gene appears to be expressed in melanocytes of feather germs in the Japanese quail (Coturnix coturnix japonica)

The Bh (black at hatch) gene was examined to determine whether it is expressed in plumage melanocytes by analyzing pigmentation patterns of Bh melanocytes placed in the micro-environment of the feather germs of quail embryos with pink eyes. These host quails genetically lack a large part of plumage melanin. The Bh locus in these almost white quails is wild-type. When Bh neural crest cells were transplanted orthotopically into the host embryos, wild-type and Bh /+ melanocytes, which differentiated from the transplanted neural crest cells, formed plumage pigmentation patterns characteristic of each genotype in the micro-environment of the host feather germs. Brown plumage pigmentation, which was very similar to that of 10-day Bh / Bh embryos, was also observed in the feather germs of host embryos that received Bh neural crest cells, although the genotype of the donors could not be determined. These donors died before pigmentation of their feather germs occurred. The results demonstrate that pigmentation patterns of Bh menalocytes are not altered in the micro-environment of the host germs, suggesting that the Bh gene is autonomous in Bh melanocytes and is expressed in melanocytes of both Bh and the host feather germs, and that it causes the normal pigmentation pattern to be altered.     

Environmental Pollution Affects the Plumage Color of Great Tit Nestlings through Carotenoid Availability

Birds need to acquire carotenoids for their feather pigmentation from their diet, which means that their plumage color may change as a consequence of human impact on their environment. For example, the carotenoid-based plumage coloration of Great tit, Parus major, nestlings is associated with the degree of environmental pollution. Breast feathers of birds in territories exposed to heavy metals are less yellow than those in unpolluted environments. Here we tested two hypotheses that could explain the observed pattern: (I) deficiency of carotenoids in diet, and (II) pollution-related changes in transfer of carotenoids to feathers. We manipulated dietary carotenoid levels of nestlings and measured the responses in plumage color and tissue concentrations. Our carotenoid supplementation produced the same response in tissue carotenoid concentrations and plumage color in polluted and unpolluted environments. Variation in heavy metal levels did not explain the variation in tissue (yolk, plasma, and feathers) carotenoid concentrations and was not related to plumage coloration. Instead, the variation in plumage yellowness was associated with the availability of carotenoid-rich caterpillars in territories. Our results support the hypothesis that the primary reason for pollution-related variation in plumage color is carotenoid deficiency in the diet.     

From preen secretions to plumage: the chemical trajectory of blue petrels' Halobaena caerulea social scent

Petrel seabirds heavily rely on their olfactory sense and are thus appropriate models for the study of avian chemical communication. The uropygial secretions of blue petrels Halobaena caerulea, for instance, have been shown to encapsulate elaborate sociochemical information including species, gender and identity. Yet, it is the plumage, and not preening secretions per se, which acts as the final substrate of avian scents. Importantly, the chemical relationship between secretions from the uropygial gland, located at the dorsal base of the tail, and plumage lipids has been considered in only a handful of studies which reported large qualitative differences. The emission process of avian scents, and the possible participation of the uropygial gland in particular, are therefore not elucidated. In the present study, we examine the early chemical trajectory of blue petrels' social chemosignals by comparing secretion and feather samples using Gas‐Chromatography Mass‐Spectrometry (GC/MS) and recently developed multivariate statistics. Our results indicate that (1) 85% of the feather lipids come from the uropygial secretions, (2) chemical differentiation between secretions and feather lipids includes qualitative and quantitative variations, which both have interesting implications for scent production, (3) the sociochemical information contained within the secretions (i.e. a sex‐specific signal and individual chemical signatures) are present in very conserved forms on the plumage. In the light of these results, it is apparent that the uropygial gland plays a critical role for chemical communication in petrels and possibly other avian groups.     

Is Preening Behaviour Sexually Selected? An Experimental Approach

Elaborate or colourful feathers are important traits in female mate choice in birds but little attention has been given to potential costs of maintaining these traits in good condition with preening behaviour. Recent studies indicate that the time and energy required to maintain ornamental plumage in good condition reinforces the honesty of plumage trait. It has been proposed that some behaviours, whose primary function is not to transfer information, can also evolve as signalling components. Here we investigate whether the preening behaviour intensity has a signalling component: we hypothesized that if only high quality males can invest a lot of time in preening, this behaviour may be used by females as a quality signal (attractive preening hypothesis). We tested this hypothesis by using female budgerigars in mate‐choice tests in captivity. We tried to experimentally manipulate the preening behaviour of two groups of budgerigar males (treatment and control group). The proportion of time in which treated males preened in front of females was statistically higher than for control males, however, females spent similar amounts of time with treated males and control males. Moreover, males did not show significant quantitative changes in preening (for both groups) when females were present, suggesting that male budgerigars did not use this behaviour to convey information. These results are inconsistent with the ‘attractive preening’ hypothesis which predicts that preening behaviour itself provides information on condition and is used in female choice.     

Population differences in the structure and coloration of great tit contour feathers

Contour feathers cover most of the avian body and play critical roles in insulation, social communication, aerodynamics, and water repellency. Feather production is costly and the development of the optimum characteristics for each function may be constrained by limited resources or time, and possibly also lead to trade‐offs among the different characteristics. Populations exposed to different environmental conditions may face different selective pressures, resulting in differences in feather structure and coloration, particularly in species with large geographical distributions. Three resident populations of great tit Parus major L. from different latitudes differed in feather structure and coloration. Individuals from the central population exhibited less dense and longer contour feathers, with a higher proportion of plumulaceous barbs than either northern or southern birds, which did not differ in their feather structure. Ultraviolet reflectance and brightness of the yellow of the contour feathers of the breast was higher for the southern than for the northern population. Birds with greener plumage (higher hue) had less dense but longer feathers, independently of the population of origin. Differences in feather structure across populations appear to be unrelated to the contour feather colour characteristics except for hue. Nutritional and time constraints during molt might explain the pattern of feather structure, whereas varying sexual selection pressure might underlie the coloration patterns observed. Our results suggest that different selective pressures or constraints shape contour feather traits in populations exposed to varying environmental conditions. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 114 , 82–91.     

Plumage maintenance affects ultraviolet colour and female preference in the budgerigar

Elaborate or colourful feathers are important traits in female-mate choice in birds but little attention has been given to the potential costs of maintaining these traits in good condition via preening behaviour. While preening is known to be an important component of plumage maintenance, it has received little attention with respect to colouration. We investigated whether preening can influence plumage reflectance and whether females show a preference for plumage cleanliness in captive-bred, wild-type budgerigars, Melopsittacus undulatus. To do this, we compared the spectral colour of birds that were allowed to preen their plumage and individuals that were prevented from preening. The plumage of birds that were prevented from preening showed a significant lower reflectance in the UV range (300-400 nm). Subsequently, we measured females’ preferences for preened and unpreened males using a two-choice test. In a second experiment we allowed females to choose between an unpreened male and a male smeared with UV-absorbing chemicals (UV-blocked male). The proportion of time that females stayed near preened males was statistically higher than for unpreened males, but females spent similar amounts of time with unpreened males and UV-blocked males. These results are consistent with the idea that female budgerigars are able to discriminate between preened and unpreened males, and that UV colours, mediated by preening, can convey information about a bird's current condition.     

The Relationship between Personality Dimensions and Resiliency to Environmental Stress in Orange-Winged Amazon Parrots (Amazona amazonica), as Indicated by the Development of Abnormal Behaviors

Parrots are popular companion animals, but are frequently relinquished because of behavioral problems, including abnormal repetitive behaviors like feather damaging behavior and stereotypy. In addition to contributing to pet relinquishment, these behaviors are important as potential indicators of diminished psychological well-being. While abnormal behaviors are common in captive animals, their presence and/or severity varies between animals of the same species that are experiencing the same environmental conditions. Personality differences could contribute to this observed individual variation, as they are known risk factors for stress sensitivity and affective disorders in humans. The goal of this study was to assess the relationship between personality and the development and severity of abnormal behaviors in captive-bred orange-winged Amazon parrots (Amazona amazonica). We monitored between-individual behavioral differences in enrichment-reared parrots of known personality types before, during, and after enrichment deprivation. We predicted that parrots with higher scores for neurotic-like personality traits would be more susceptible to enrichment deprivation and develop more abnormal behaviors. Our results partially supported this hypothesis, but also showed that distinct personality dimensions were related to different forms of abnormal behavior. While neuroticism-like traits were linked to feather damaging behavior, extraversion-like traits were negatively related to stereotypic behavior. More extraverted birds showed resiliency to environmental stress, developing fewer stereotypies during enrichment deprivation and showing lower levels of these behaviors following re-enrichment. Our data, together with the results of the few studies conducted on other species, suggest that, as in humans, certain personality types render individual animals more susceptible or resilient to environmental stress. Further, this susceptibility/resiliency can have a long-term effect on behavior, as evidenced by behavioral changes that persisted despite re-enrichment. Ours is the first study evaluating the relationship between personality dimensions, environment, and abnormal behaviors in an avian species.     

A PHYLOGENETIC PERSPECTIVE ON THE EVOLUTION OF SEXUAL DICHROMATISM IN TANAGERS (THRAUPIDAE): THE ROLE OF FEMALE VERSUS MALE PLUMAGE

The evolution of sexual dichromatism in tanagers (family Thraupidae) was studied from a phylogenetic perspective using a molecular-based phylogeny. Mapping patterns of sexual dimorphism in plumage onto the phylogeny reveals that changes in female plumage occur more frequently than changes in male plumage. Possible explanations for this pattern include sexual selection acting on female plumage and natural selection for background matching. The results of this study and other recent phylogenetic and comparative studies suggest that factors affecting female plumage are important in shaping patterns of sexual dimorphism.     

An analysis of displacement preening in the domestic fowl

The preening behaviour of the domestic fowl was filmed under both normal and thwarting conditions. The film was analysed frame by frame and the duration of individual preening movements was measured. It was found that preening movements in the thwarting situation were of shorter duration than those in the normal situation and it was thought that this accounted for the ‘frantic’ appearance of the displacement preening. The pattern of preening also differed in the thwarting situation with plumage areas which were easy to reach receiving more attention than in the control situation.     

Concordant evolution of plumage colour, feather microstructure and a melanocortin receptor gene between mainland and island populations of a fairy-wren

Studies of the patterns of diversification of birds on islands have contributed a great deal to the development of evolutionary theory. In white-winged fairy-wrens, Malurus leucopterus, mainland males develop a striking blue nuptial plumage whereas those on nearby islands develop black nuptial plumage. We explore the proximate basis for this divergence by combining microstructural feather analysis with an investigation of genetic variation at the melanocortin-1 receptor locus (MC1R). Fourier analysis revealed that the medullary keratin matrix (spongy layer) of the feather barbs of blue males was ordered at the appropriate nanoscale to produce the observed blue colour by coherent light scattering. Surprisingly, the feather barbs of black males also contained a spongy layer that could produce a similar blue colour. However, black males had more melanin in their barbs than blue males, and this melanin may effectively mask any structural colour produced by the spongy layer. Moreover, the presence of this spongy layer suggests that black island males evolved from a blue-plumaged ancestor. We also document concordant patterns of variation at the MC1R locus, as five amino acid substitutions were perfectly associated with the divergent blue and black plumage phenotypes. Thus, with the possible involvement of a melanocortin receptor locus, increased melanin density may mask the blue-producing microstructure in black island males, resulting in the divergence of plumage coloration between mainland and island white-winged fairy-wrens. Such mechanisms may also be responsible for plumage colour diversity across broader geographical and evolutionary scales.