Carotenoid-based plumage pigmentation and concentration as a function of sex and habitat type in the Yellow-breasted Boubou Laniarius atroflavus

The study of avian integumentary colouration can offer insight into dietary and metabolic processes as well as fitness in focal species. Yet, we know relatively less about the system of feather colouration in African birds in comparison to Europe, North America and the neotropics. In this study, we biochemically characterised and quantified the pigmentary basis for breast plumage colouration in the Yellow-breasted Boubou Laniarius atroflavus, a little-known Afromontane species restricted to the Nigerian–Cameroon Highlands. We also measured differences in carotenoid concentration and feather reflectance between sexes, and between birds inhabiting edge and riparian habitats. Six carotenoid pigments were recovered from the yellow feathers – canary xanthophyll A and B, a cis isomer of each, isoastaxanthin and an unidentified carotenoid. We determined that the yellow colour of the breast feathers is carotenoid-based, with the greater proportion as canary xanthophylls. The presence of the ketocarotenoid, isoastaxanthin, provides the basis for further studies into red, orange and yellow coloured congenerics. Males appeared to have higher feather pigment concentrations than females, and birds resident in the edge habitat appeared to have slightly higher feather pigment concentrations than those in the degraded riparian habitat. There was little indication of differences in feather reflectance between sexes and habitat types. However, low samples size restricted further differentiation. There is also the need for further studies on the dietary and metabolic pathways of feather colouration to better understand how ecological variation may shape pigment uptake, transport, synthesis and deposition in feathers.     

Energetic constraints on expression of carotenoid-based plumage coloration

Carotenoid pigments are used by many bird species as feather colorants, creating brilliant yellow, orange, and red plumage displays. Such carotenoid-based plumage coloration has been shown to function as an honest signal that is used in female mate choice. Despite recent interest in carotenoid-based ornamental traits, the basis for individual variation in expression of carotenoid-based plumage coloration remains incompletely understood. I tested the hypothesis that, independent of carotenoid access, food stress during molt would cause reduced expression of carotenoid pigmentation. I fed molting male House Finches Carpodacus mexicanus seed diets supplemented with either the red carotenoid pigment canthaxanthin or the yellow/orange carotenoid pigment β-cryptoxanthin (in the form of tangerine juice). Within each diet treatment, one group of males was given restricted food access and the other group was given unrestricted food access. Carotenoid supplements were placed in water so carotenoid access was controlled independent of food access. The results indicated a strong effect of both carotenoid access and food access on color display. Some males in the β-cryptoxanthin-supplemented group grew red plumage, suggesting that they can metabolically modify yellow pigments into red pigments, but no bird supplemented with β-cryptoxanthin grew plumage as red as birds supplemented with canthaxanthin. Males in the unrestricted food groups grew redder and more intensely pigmented plumage than males in the restricted food groups. These observations provide the best evidence to date of an energetic cost of carotenoid utilization in the generation of colorful plumage.     

基于类胡萝卜素着色的鸟类羽色多样性形成机制

人们对动物体色的研究由来已久。作为一类让生物呈现出多变色彩的重要色素, 类胡萝卜素可以在鸟类的羽毛、鸟喙和皮肤等体表组织中沉积, 产生红、橙、黄、粉、紫等颜色。类胡萝卜素不能在鸟类体内合成, 需从食物中摄取, 进而在体内完成吸收、运输、代谢和沉积等一系列过程, 才能用于羽毛着色。与类胡萝卜素着色相关的生理及遗传调控机制一直备受关注, BCO2、SCARB1和CYP2J19等影响类胡萝卜素在鸟类羽毛中着色的关键基因, 推动了对羽色遗传调控机制的深入认识。本文介绍了鸟类可利用类胡萝卜素的主要类型和基本特征, 综述了类胡萝卜素着色相关的生理过程以及调控基因研究的最新进展, 旨在增加对鸟类羽毛中类胡萝卜素着色过程和相关遗传机制的理解。     

Carotenoid coloration in greenfinches is individually consistent irrespective of foraging ability

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.     

Pigment patterns in neural crest chimeras constructed from quail and guinea fowl embryos

The pattern of pigmentation in bird embryos is determined by the spatial organization of melanocyte differentiation. Some of the results from recent, neural crest transplantation experiments support a model based on a prepattern in the feathers; others could be interpreted in terms of a nonspecific pattern resulting from a failure of the crest cells to read the positional values in another species. To distinguish between these possibilities, the crucial test is to construct chimeras from two species with different pigment patterns. We have examined the wing plumage of quail and guinea fowl embryos. The quail has a characteristic pattern of pigmented and unpigmented feather papillae, whereas the guinea fowl shows uniform pigmentation. Chimeras were constructed by grafting wing buds isotopically between embryos. The wing buds were transplanted before they had become invaded by neural crest cells. Quail wing buds grafted to the guinea fowl developed, in most cases, a pigment pattern resembling that of the quail and not that of the guinea fowl. A few cases became uniformly pigmented and appeared to represent nonspecific patterns. The reciprocal grafts (guinea fowl wing buds grafted to the quail) became pigmented all over. We found evidence that the timing of melanocyte differentiation is controlled by cues in the feather papillae. Some cases developed a severe inflammatory response. The model which best accounts for these findings--and which can account for inconsistencies in previous reports--is the following. A prepattern is present in the feathers and this can control the differentiation of melanoblasts, even if they come from a different species. The local cues which constitute the prepattern are not positional values. In some chimeras melanoblasts fail to respond to the prepattern and so a nonspecific pattern of uniform pigmentation is produced.     

Differential accumulation and pigmenting ability of dietary carotenoids in colorful finches

Many animals develop bright red, orange, or yellow carotenoid pigmentation that they use to attract mates. Colorful carotenoid pigments are acquired from the diet and are either directly incorporated as integumentary colorants or metabolized into other forms before deposition. Because animals often obtain several different carotenoids from plant and animal food sources, it is possible that these pigments are accumulated at different levels in the body and may play unique roles in shaping the ultimate color expression of individuals. We studied patterns of carotenoid accumulation and integumentary pigmentation in two colorful finch species--the American goldfinch (Carduelis tristis) and the zebra finch (Taeniopygia guttata). Both species acquire two main hydroxycarotenoids, lutein and zeaxanthin, from their seed diet but transform these into a series of metabolites that are used as colorful pigments in the plumage (goldfinches only) and beak (both species). We conducted a series of carotenoid-supplementation experiments to investigate the relative extent to which lutein and zeaxanthin are accumulated in blood and increase carotenoid coloration in feathers and bare parts. First, we supplemented the diets of both species with either lutein or zeaxanthin and measured plasma pigment status, feather carotenoid concentration (goldfinches only), and integumentary color. Zeaxanthin-supplemented males grew more colorful feathers and beaks than lutein-supplemented males, and in goldfinches incorporated a different ratio of carotenoids in feathers (favoring the accumulation of canary xanthophyll B). We also fed goldfinches different concentrations of a standard lutein-zeaxanthin mix and found that at physiologically normal and high concentrations, birds circulated proportionally more zeaxanthin over lutein than occurred in the diet. Collectively, these results demonstrate that zeaxanthin is preferentially accumulated in the body and serves as a more potent substrate for pigmentation than lutein in these finches.     

Theory of the growth and evolution of feather shape

We present the first explicit theory of the growth of feather shape, defined as the outline of a pennaceous feather vane. Based on a reanalysis of data from the literature, we propose that the absolute growth rate of the barbs and rachis ridges, not the vertical growth rate, is uniform throughout the follicle. The growth of feathers is simulated with a mathematical model based on six growth parameters: (1) absolute barb and rachis ridge growth rate, (2) angle of helical growth of barb ridges, (3) initial barb ridge number, (4) new barb ridge addition rate, (5) barb ridge diameter, and (6) the angle of barb ramus expansion following emergence from the sheath. The model simulates growth by cell division in the follicle collar and, except for the sixth parameter, does not account for growth by differentiation in cell size and shape during later keratinization. The model can simulate a diversity of feather shapes that correspond closely in shape to real feathers, including various contour feathers, asymmetrical feathers, and even emarginate primaries. Simulations of feather growth under different parameter values demonstrate that each parameter can have substantial, independent effects on feather shape. Many parameters also have complex and redundant effects on feather shape through their influence on the diameter of the follicle, the barb ridge fusion rate, and the internodal distance. Simulated isochrones-the loci, or sets, of feather cells of the same age-have the same oblique chevron-shaped position in the mature feather as fault bars, which are isochronic defects in the barbules created by a disruptions during development. Accurate simulation of fault bar shape and position confirms the uniform absolute growth rate hypothesis and the general realism of the model. The theory defines a six-parameter feather morphospace, and provides many predictions about the developmental determination of feather shape that can be tested with detailed observations and experiments on developing feathers. This theory also provides testable predictions about the changes in developmental mechanisms required to evolve different feather shapes to accomplish various functions.     

Carotenoid‐based plumage coloration in golden‐crowned kinglets Regulus satrapa: pigment characterization and relationships with migratory timing and condition

Carotenoid‐based ornamental coloration has long been proposed to honestly signal quality due to its dependence on individual condition. Because migration can be one of the most stressful periods of an animal's annual cycle, developing colourful plumage may be particularly challenging for species in which migration and moult periods overlap or occur sequentially. The purpose of this study was to investigate pigmentary and condition‐dependent bases of carotenoid colour variation in a small migratory passerine, the golden‐crowned kinglet Regulus satrapa (Family Regulidae). We captured 186 male and female kinglets of various ages during fall migration in southwestern Ontario, Canada and recorded arrival date, body condition index, fat and pectoral muscle scores, wing mite infestation, and feather growth rate as measures of condition. We quantified crown coloration using reflectance spectrometry and analyzed feather carotenoids using high‐performance liquid chromatography. Yellow crown feathers of female kinglets contained only yellow hydroxycarotenoids, whereas orange feathers of males harboured a suite of eight carotenoid pigments. Males with longer wavelength orange crown hues deposited greater concentrations of ketocarotenoids, especially canthaxanthin. Female kinglets with longer wavelength crown hues and males with longer wavelength crown hues and more saturated crown coloration left for migration earlier in the year. Females with longer wavelength crown hues had fewer feather mites and tended to be in better condition. However, male kinglets with more saturated coloration possessed smaller pectoral muscles. This is the first study to identify plumage carotenoids in this North American bird family and to determine the pigmentary basis for both inter‐ and intrasexual colour variation. Our results provide further support for the condition‐dependence of carotenoid coloration and suggest that ornamental elaboration in both sexes may encode information about fall condition and migratory performance.     

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.     

Shh-Bmp2 signaling module and the evolutionary origin and diversification of feathers

To examine the role of development in the origin of evolutionary novelties, we investigated the developmental mechanisms involved in the formation of a complex morphological novelty-branched feathers. We demonstrate that the anterior-posterior expression polarity of Sonic hedgehog (Shh) and Bone morphogenetic protein 2 (Bmp2) in the primordia of feathers, avian scales, and alligator scales is conserved and phylogenetically primitive to archosaurian integumentary appendages. In feather development, derived patterns of Shh-Bmp2 signaling are associated with the development of evolutionarily novel feather structures. Longitudinal Shh-Bmp2 expression domains in the marginal plate epithelium between barb ridges provide a prepattern of the barbs and rachis. Thus, control of Shh-Bmp2 signaling is a fundamental component of the mechanism determining feather form (i.e., plumulaceous vs. pennaceous structure). We show that Shh signaling is necessary for the formation and proper differentiation of a barb ridge and that it is mediated by Bmp signaling. BMP signaling is necessary and sufficient to negatively regulate Shh expression within forming feather germs and this epistatic relationship is conserved in scale morphogenesis. Ectopic SHH and BMP2 signaling leads to opposing effects on proliferation and differentiation within the feather germ, suggesting that the integrative signaling between Shh and Bmp2 is a means to regulate controlled growth and differentiation of forming skin appendages. We conclude that Shh and Bmp signaling is necessary for the formation of barb ridges in feathers and that Shh and Bmp2 signaling constitutes a functionally conserved developmental signaling module in archosaur epidermal appendage development. We propose a model in which branched feather form evolved by repeated, evolutionary re-utilization of a Shh-Bmp2 signaling module in new developmental contexts. Feather animation Quicktime movies can be viewed at http://fallon.anatomy.wisc.edu/feather.html.     

Sexual Dimorphism in Melanin Pigmentation,Feather Coloration and Its Heritability in the Barn Swallow (Hirundo rustica)

Melanin is the main pigment in animal coloration and considerable variation in the concentrations of the two melanin forms (pheo- and eumlanin) in pigmented tissues exists among populations and individuals. Melanin-based coloration is receiving increasing attention particularly in socio-sexual communication contexts because the melanocortin system has been hypothesized to provide a mechanistic basis for covariation between coloration and fitness traits. However, with few notable exceptions, little detailed information is available on inter-individual and inter-population variation in melanin pigmentation and on its environmental, genetic and ontogenetic components. Here, we investigate melanin-based coloration in an Italian population of a passerine bird, the barn swallow (Hirundo rustica rustica), its sex- and age-related variation, and heritability. The concentrations of eu- and pheomelanin in the throat (brown) and belly (white-to-brownish) feathers differed between sexes but not according to age. The relative concentration of either melanin (Pheo:Eu) differed between sexes in throat but not in belly feathers, and the concentrations in males compared to females were larger in belly than in throat feathers. There were weak correlations between the concentrations of melanins within as well as among plumage regions. Coloration of belly feathers was predicted by the concentration of both melanins whereas coloration of throat feathers was only predicted by pheomelanin in females. In addition, Pheo:Eu predicted coloration of throat feathers in females and that of belly feathers in males. Finally, we found high heritability of color of throat feathers. Melanization was found to differ from that recorded in Hirundo rustica rustica from Scotland or from H. r. erythrogaster from North America. Hence, present results show that pigmentation strategies vary in a complex manner according to sex and plumage region, and also among geographical populations, potentially reflecting adaptation to different natural and sexual selection regimes, and that some coloration components seem to be highly heritable.     

Which strategy is better for linkage analysis: single-nucleotide polymorphisms or microsatellites? Evaluation by identity-by-state – identity-by-descent transformation affected sib-pair method on GAW14 data

Background

The genetics of plumage of Japanese quail is of interest both from a biological standpoint, for comparative studies between avian species, and from a zootechnical standpoint, for identifying commercial selection lines or crosses. There are only few plumage mutations reported in quail, and the present work describes a new color variant "rusty" and a new feather structure "curly", and their heredity from an F1 and F2 segregation experiment.

Results

Curly feathers result from abnormal early growth caused by transient joining of follicle walls of adjacent feathers around 10 days of age, but the expression of the trait is variable. Rusty plumage color results from the replacement of the wild-type plumage pattern on the tip of the feather by a reddish coloration, but the pigmentation of the bottom part of the feather is not affected. Two lines breeding true for the curly or the rusty phenotype were developed. Both characters are determined by autosomal recessive mutations which are independent. The curly mutation has also a positive effect on body weight at 5 weeks of age.

Conclusion

The curly line is a new model which may be used for further work on the growth of the feather, and the rusty mutation is a new addition to the panel of plumage mutations available for comparative studies in poultry, and more generally among avian species.     

Molecular biology of feather morphogenesis: a testable model for evo-devo research

Darwin's theory describes the principles that are responsible for evolutionary change of organisms and their attributes. The actual mechanisms, however, need to be studied for each species and each organ separately. Here we have investigated the mechanisms underlying these principles in the avian feather. Feathers comprise one of the most complex and diverse epidermal organs as demonstrated by their shape, size, patterned arrangement and pigmentation. Variations can occur at several steps along each level of organization, leading to highly diverse forms and functions. Feathers develop gradually during ontogeny through a series of steps that may correspond to the evolutionary steps that were taken during the phylogeny from a reptilian ancestor to birds. These developmental steps include 1) the formation of feather tract fields on the skin surfaces; 2) periodic patterning of the individual feather primordia within the feather tract fields; 3) feather bud morphogenesis establishing anterio-posterior (along the cranio-caudal axis) and proximo-distal axes; 4) branching morphogenesis to create the rachis, barbs and barbules within a feather bud; and 5) gradual modulations of these basic morphological parameters within a single feather or across a feather tract. Thus, possibilities for variation in form and function of feathers occur at every developmental step. In this paper, principles guiding feather tract formation, distributions of individual feathers within the tracts and variations in feather forms are discussed at a cellular and molecular level.     

The evolutionary origin and diversification of feathers

Progress on the evolutionary origin and diversification of feathers has been hampered by conceptual problems and by the lack of plesiomorphic feather fossils. Recently, both of these limitations have been overcome by the proposal of the developmental theory of the origin of feathers, and the discovery of primitive feather fossils on nonavian theropod dinosaurs. The conceptual problems of previous theories of the origin of feathers are reviewed, and the alternative developmental theory is presented and discussed. The developmental theory proposes that feathers evolved through a series of evolutionary novelties in developmental mechanisms of the follicle and feather germ. The discovery of primitive and derived fossil feathers on a diversity of coelurosaurian theropod dinosaurs documents that feathers evolved and diversified in nonavian theropods before the origin of birds and before the origin of flight. The morphologies of these primitive feathers are congruent with the predictions of the developmental theory. Alternatives to the theropod origin of feathers are critique and rejected. Hypotheses for the initial function of feathers are reviewed. The aerodynamic theory of feather origins is falsified, but many other functions remain developmentally and phylogenetically plausible. Whatever their function, feathers evolved by selection for a follicle that would grow an emergent tubular appendage. Feathers are inherently tubular structures. The homology of feathers and scales is weakly supported. Feathers are composed of a suite of evolutionary novelties that evolved by the duplication, hierarchical organization, interaction, dissociation, and differentiation of morphological modules. The unique capacity for modular subdivision of the tubular feather follicle and germ has fostered the evolution of numerous innovations that characterize feathers. The evolution of feather keratin and the molecular basis of feather development are also discussed.     

Ancient origins and multiple appearances of carotenoid-pigmented feathers in birds

The broad palette of feather colours displayed by birds serves diverse biological functions, including communication and camouflage. Fossil feathers provide evidence that some avian colours, like black and brown melanins, have existed for at least 160 million years (Myr), but no traces of bright carotenoid pigments in ancient feathers have been reported. Insight into the evolutionary history of plumage carotenoids may instead be gained from living species. We visually surveyed modern birds for carotenoid-consistent plumage colours (present in 2956 of 9993 species). We then used high-performance liquid chromatography and Raman spectroscopy to chemically assess the family-level distribution of plumage carotenoids, confirming their presence in 95 of 236 extant bird families (only 36 family-level occurrences had been confirmed previously). Using our data for all modern birds, we modelled the evolutionary history of carotenoid-consistent plumage colours on recent supertrees. Results support multiple independent origins of carotenoid plumage pigmentation in 13 orders, including six orders without previous reports of plumage carotenoids. Based on time calibrations from the supertree, the number of avian families displaying plumage carotenoids increased throughout the Cenozoic, and most plumage carotenoid originations occurred after the Miocene Epoch (23 Myr). The earliest origination of plumage carotenoids was reconstructed within Passeriformes, during the Palaeocene Epoch (66–56 Myr), and not at the base of crown-lineage birds.     

Growth and pigmentation in Sphingobacterium antarcticus,a psychrotrophic bacterium from Antarctica

Sphingobacterium antarcticus, a yellow pigmented psychrotrophic bacterium from Antarctica exhibited enhanced pigmentation with increasing temperatures of incubation. This behavior was opposite to mesophilic Sphingobacterium in which pigmentation was reduced upon raising temperatures. The UV-visible spectrum of the crude pigment was characteristic of carotenoids and the pigment gave negative tests for flexirubins. Diphenylamine (DPA), a standard biochemical blocker of carotenoid biosynthesis reduced the growth of broth cultures when grown at extremes of the optimum temperature. Mutants defective in pigmentation were capable of growing between 1–31°C suggesting that pigmentation does not play any role in adapting the bacterium to the psychrotrophic growth temperature. On the other hand, our results suggest that DPA, which is known to block desaturation reactions in the carotenoid biosynthesis pathway may be affecting other desaturation reactions within the bacterium thereby causing reduced growth at extreme temperatures.     

Red-winged blackbirds Agelaius phoeniceus use carotenoid and melanin pigments to color their epaulets

Over the past three decades, the red‐winged blackbird Agelaius phoeniceus has served as a model species for studies of sexual selection and the evolution of ornamental traits. Particular attention has been paid to the role of the colorful red‐and‐yellow epaulets that are striking in males but reduced in females and juveniles. It has been assumed that carotenoid pigments bestow the brilliant red and yellow colors on epaulet feathers, but this has never been tested biochemically. Here, we use high‐performance liquid chromatography (HPLC) to describe the pigments present in these colorful feathers. Two red ketocarotenoids (astaxanthin and canthaxanthin) are responsible for the bright red hue of epaulets. Two yellow dietary precursors pigments (lutein and zeaxanthin) are also present in moderately high concentrations in red feathers. After extracting carotenoids, however, red feathers remained deep brown in color. HPLC tests show that melanin pigments (primarily eumelanin) are also found in the red‐pigmented barbules of epaulet feathers, at an approximately equal concentration to carotenoids. This appears to be an uncommon feature of carotenoid‐based ornamental plumage in birds, as was shown by comparable analyses of melanin in the yellow feathers of male American goldfinches Carduelis tristis and the red feathers of northern cardinals Cardinalis cardinalis, in which we detected virtually no melanins. Furthermore, the yellow bordering feathers of male epaulets are devoid of carotenoids (except when tinged with a carotenoid‐derived pink coloration on occasion) and instead are comprised of a high concentration of primarily phaeomelanin pigments. The dual pigment composition of red epaulet feathers and the melanin‐only basis for yellow coloration may have important implications for the honesty‐reinforcing mechanisms underlying ornamental epaulets in red‐winged blackbirds, and shed light on the difficulties researchers have had to date in characterizing the signaling function of this trait. As in several other birds, the melanic nature of feathers may explain why epaulets are used largely to settle aggressive contests rather than to attract mates.     

Sex and age differences in reflectance and biochemistry of carotenoid‐based colour variation in the great tit Parus major

The plumage coloration in great tits (Parus major) is the subject of much behavioural and ecophysiological research, yet there is a lack of analyses of the natural colour variation and its mechanisms. We used reflectance spectrometry and high‐performance liquid chromatography to explore individual, sexual and age‐related variation in carotenoid coloration and pigmentation, paramount to the often presumed, but rarely substantiated, costs and ‘honesty’ of carotenoid displays. In adults, we found that sex was the strongest predictor of ‘brightness’ (higher in males) and of ‘hue’ (longer wavelength in females). There was no sex difference in ‘carotenoid chroma’ or carotenoid content of feathers which also was unrelated to adult age (1 or 2+ years) and condition. Similar patterns were revealed for nestlings. Regarding the biochemical ‘signal content’, ‘carotenoid chroma’, but not ‘hue’, was significantly related to the carotenoid content (lutein and zeaxanthin) of feathers. These results refute the previously assumed exaggeration of carotenoid pigmentation in male great tits, and question the condition‐dependence of carotenoid coloration in this species. However, the sexual dimorphism in total reflectance or ‘brightness’, most likely due to melanins rather than carotenoids, may have implications for signalling or other adaptive explanations that need to be explored. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 758–765.     

Preen waxes do not protect carotenoid plumage from bleaching by sunlight

The plumage coloration of wild birds often changes during the breeding season. One of the possible reasons for this is that sunlight, and particularly ultraviolet (UV) wavelengths, degrades the pigments responsible for plumage coloration. It has been suggested that birds may apply preen wax to feathers to protect feathers from bleaching. This hypothesis is tested by exposing carotenoid-based breast feathers of Great Tits to ambient light, light filtered to exclude UV and darkness. Preen waxes were experimentally removed from feather samples and the effect of light on coloration of treatment and control feathers compared. Ambient light had an effect on feather colour but preen wax did not. Feathers exposed to sun gradually became less saturated and hues shifted towards shorter wavelengths. This was not apparent in control feathers kept in darkness. Feathers exposed to full-spectra sunlight faded more than those that were kept in light with UV wavelengths removed. There was a decrease in brightness of feathers in both experimental and control groups, which was assumed to be an effect of dirt accumulation. This experiment confirmed earlier suspicions regarding the detrimental effects of UV irradiation on carotenoid-based coloration of avian feathers but failed to show any protective function of preen waxes. The possible consequences of these mechanisms of colour change for birds with regard to mating strategies are discussed.