An experimental test of the contributions and condition dependence of microstructure and carotenoids in yellow plumage coloration

A combination of structural and pigmentary components is responsible for many of the colour displays of animals. Despite the ubiquity of this type of coloration, neither the relative contribution of structures and pigments to variation in such colour displays nor the relative effects of extrinsic factors on the structural and pigment-based components of such colour has been determined. Understanding the sources of colour variation is important because structures and pigments may convey different information to conspecifics. In an experiment on captive American goldfinches Carduelis tristis, we manipulated two parameters, carotenoid availability and food availability, known to affect the expression of carotenoid pigments in a full-factorial design. Yellow feathers from these birds were then analysed in two ways. First, we used full-spectrum spectrometry and high-performance liquid chromatography to examine the extent to which variation in white structural colour and total carotenoid content was associated with variation in colour properties of feathers. The carotenoid content of yellow feathers predicted two colour parameters (principal component 1--representing high values of ultraviolet and yellow chroma and low values of violet-blue chroma-and hue). Two different colour parameters (violet-blue and yellow chroma) from white de-pigmented feathers, as well as carotenoid content, predicted reflectance measurements from yellow feathers. Second, we determined the relative effects of our experimental manipulations on white structural colour and yellow colour. Carotenoid availability directly affected yellow colour, while food availability affected it only in combination with carotenoid availability. None of our manipulations had significant effects on the expression of white structural colour. Our results suggest that the contribution of microstructures to variation in the expression of yellow coloration is less than the contribution of carotenoid content, and that carotenoid deposition is more dependent on extrinsic variability than is the production of white structural colour.     

Colourful parrot feathers resist bacterial degradation

The brilliant red, orange and yellow colours of parrot feathers are the product of psittacofulvins, which are synthetic pigments known only from parrots. Recent evidence suggests that some pigments in bird feathers function not just as colour generators, but also preserve plumage integrity by increasing the resistance of feather keratin to bacterial degradation. We exposed a variety of colourful parrot feathers to feather-degrading Bacillus licheniformis and found that feathers with red psittacofulvins degraded at about the same rate as those with melanin and more slowly than white feathers, which lack pigments. Blue feathers, in which colour is based on the microstructural arrangement of keratin, air and melanin granules, and green feathers, which combine structural blue with yellow psittacofulvins, degraded at a rate similar to that of red and black feathers. These differences in resistance to bacterial degradation of differently coloured feathers suggest that colour patterns within the Psittaciformes may have evolved to resist bacterial degradation, in addition to their role in communication and camouflage.     

Nanostructure predicts intraspecific variation in ultraviolet-blue plumage colour

Evidence suggests that structural plumage colour can be an honest signal of individual quality, but the mechanisms responsible for the variation in expression of structural coloration within a species have not been identified. We used full-spectrum spectrometry and transmission electron microscopy to investigate the effect of variation in the nanostructure of the spongy layer on expression of structural ultraviolet (UV)-blue coloration in eastern bluebird (Sialia sialis) feathers. Fourier analysis revealed that feather nanostructure was highly organized but did not accurately predict variation in hue. Within the spongy layer of feather barbs, the number of circular keratin rods significantly predicted UV-violet chroma, whereas the standard error of the diameter of these rods significantly predicted spectral saturation. These observations show that the precision of nanostructural arrangement determines some colour variation in feathers.     

Comparing plumage colour measurements obtained directly from live birds and from collected feathers: the case of the great tit Parus major

Birds frequently display a colourful plumage which is important both in inter and intraespecific communication, and either in sexual and social contexts. In last years some methodologies have been developed to, analyse plumage coloration, but the use of the spectrometers has been particularly important for UV range. Measurement of plumage coloration with the spectrometer may be taken directly on the bird or, alternatively by collecting some feathers and measuring them later in the laboratory. However, few is known about the reliability of measures obtained from feathers and whether these are really representative of plumage coloration. We tested this assumption analysing measurements of carotenoids-based coloration components (lightness, chroma and hue) and lutein peak of the yellow breast of the great tit Parus major. We used two spectrometers (Ocean optics and Minolta) which calculate differently the colour components. Our results showed that direct measurement of bird was highly repeatable to determine lightness, chroma and hue for both spectrometers. Similar results we found for collected feathers procedure for both devices. Collected feathers provided high representative measurements of colour values with Minolta spectrometer. Lightness was highly repeatable when we used Ocean optic spectrometer, but chroma and hue were moderate. Lutein peak was also highly repeatable in all cases. The number of feathers used to measure plumage coloration in collected feathers procedure strongly influenced values of colour plumage variables. In general, values of lightness, chroma and hue stabilised when more than 10–15 feathers were used although we found slight differences between spectrometers. However, only four feathers were needed for lutein peak. Thus, our results stress the need to use a minimum number of feathers in measuring plumage coloration from collected feathers.     

Dramatic colour changes in a bird of paradise caused by uniquely structured breast feather barbules

The breast-plate plumage of male Lawes' parotia (Parotia lawesii) produces dramatic colour changes when this bird of paradise displays on its forest-floor lek. We show that this effect is achieved not solely by the iridescence--that is an angular-dependent spectral shift of the reflected light--which is inherent in structural coloration, but is based on a unique anatomical modification of the breast-feather barbule. The barbules have a segmental structure, and in common with many other iridescent feathers, they contain stacked melanin rodlets surrounded by a keratin film. The unique property of the parotia barbules is their boomerang-like cross section. This allows each barbule to work as three coloured mirrors: a yellow-orange reflector in the plane of the feather, and two symmetrically positioned bluish reflectors at respective angles of about 30°. Movement during the parotia's courtship displays thereby achieves much larger and more abrupt colour changes than is possible with ordinary iridescent plumage. To our knowledge, this is the first example of multiple thin film or multi-layer reflectors incorporated in a single structure (engineered or biological). It nicely illustrates how subtle modification of the basic feather structure can achieve novel visual effects. The fact that the parotia's breast feathers seem to be specifically adapted to give much stronger colour changes than normal structural coloration implies that colour change is important in their courtship display.     

Plumage colour in nestling blue tits: sexual dichromatism,condition dependence and genetic effects

Sexual-selection theory assumes that there are costs associated with ornamental plumage coloration. While pigment-based ornaments have repeatedly been shown to be condition dependent, this has been more difficult to demonstrate for structural colours. We present evidence for condition dependence of both types of plumage colour in nestling blue tits (Parus caeruleus). Using reflectance spectrometry, we show that blue tit nestlings are sexually dichromatic, with males having more chromatic (more 'saturated') and ultraviolet (UV)-shifted tail coloration and more chromatic yellow breast coloration. The sexual dimorphism in nestling tail coloration is qualitatively similar to that of chick-feeding adults from the same population. By contrast, the breast plumage of adult birds is not sexually dichromatic in terms of chroma. In nestlings, the chroma of both tail and breast feathers is positively associated with condition (body mass on day 14). The UV/blue hue of the tail feathers is influenced by paternally inherited genes, as indicated by a maternal half-sibling comparison. We conclude that the expression of both carotenoid-based and structural coloration seems to be condition dependent in blue tit nestlings, and that there are additional genetic effects on the hue of the UV/blue tail feathers. The signalling or other functions of sexual dichromatism in nestlings remain obscure. Our study shows that nestling blue tits are suitable model organisms for the study of ontogenetic costs and heritability of both carotenoid-based and structural colour in birds.     

Molecular and functional studies of tilapia (Oreochromis mossambicus) NMDA receptor NR1 subunits

The Pin-tailed Manakin (Ilicura militaris) is a small, sexually dimorphic, frugivorous suboscine songbird (Pipridae; Passeriformes; Aves) endemic to the Atlantic Forest of Brazil. A variant individual of this species was recently described in which the red patches that characterise the male's Definitive plumage were replaced by orange-yellow ones. We show here that the pigments in the feathers of the colour variant are common dietary carotenoids (zeaxanthin, beta-cryptoxanthin), not carotenoids synthesised by birds, lending support to the suggestion that the individual is a colour mutant lacking the capability to transform yellow dietary pigments into the red pigments normally present in these feathers. By comparison, the yellow crown feathers of a close relative, the Golden-winged Manakin (Masius chrysopterus), contained predominantly endogenously produced epsilon-caroten-3'-ones. Surprisingly, the normal-coloured feathers of the male Pin-tailed Manakin owe their red hue to rhodoxanthin, an unusual carotenoid more commonly found in plants, rather than 4-keto-carotenoids typically found in red plumages and found lacking in previously characterised bird colour variants. The implication is that birds, like the tilapia fish, may be able to synthesise this unusual pigment endogenously from dietary precursors. A newly described carotenoid, 6-hydroxy-epsilon,epsilon-carotene-3,3'-dione, here named piprixanthin, present in the red feathers of the Pin-tailed Manakin, provides a plausible intermediate between epsilon,epsilon-carotene-3,3'-dione (canary-xanthophyll B), a bright yellow pigment found in this and other songbirds, and rhodoxanthin. It is apparent that pigeons (Columbidae, Columbiformes) also have the capability to produce rhodoxanthin, and a structurally related pigment, endogenously. The ability to synthesise rhodoxanthin might have arisen at least twice in birds.     

Condition-dependent variation in the blue-ultraviolet coloration of a structurally based plumage ornament

After years of investigation into the function of sexually dimorphic ornamental traits, researchers are beginning to understand how bright plumage colour in birds acts as an intraspecific signal. This work has focused primarily on pigment-based ornaments because they are highly variable in patch size, hue and brightness for some species. In contrast, structurally based ornaments have been little studied, in part because they do not appear to be as variable as pigment-based ornaments. We investigated a structurally based plumage ornament in a wild population of blue grosbeaks (Guiraca caerulea), a sexually dimorphic passerine. We report plumage variation that extends into the ultraviolet region of the spectrum. The pattern of covariation between four out of five elements of plumage variation suggests that structurally based ornamentation is pushed towards extreme expression of the trait as predicted by the sexual selection theory. The ''bluest'' birds have the highest percentage of blue feathers on the body. These ornamental feathers reflect light maximally at the shortest wavelengths (ultraviolet), with the greatest intensity and the greatest contrast. Age may have some effect on expression of blueness. In addition, plumage variables are correlated with growth bars in tail feathers (a record of nutritional condition during moult in a non-ornamental trait). This suggests that the ornament is partially condition dependent. Thus, blue plumage in male grosbeaks may serve as an honest indicator of age and quality.     

The reliability of achromatic displays is island-dependent in nocturnal Storm Petrels

Nocturnal birds rely on achromatic visual signals to assess rivals and potential mates, but variation in the expression of these displays has been understudied. Here we use UV-visible reflectance spectrometry to study colour variation and the potential signalling function of the dark brown chest and white rump plumage – a colour pattern conspicuously exhibited during twilight courtship displays – in nocturnal Mediterranean Storm Petrels Hydrobates pelagicus melitensis from three islands with different exposures to predation and human disturbance. Human activity may increase perceived predation risk and thus affect activity budgets and the physiological stress response, with possible consequences on the relationship between individual body condition and plumage coloration. We found that chest and especially rump feathers reflected in the ultraviolet (UV) but there was no evidence of sexual dichromatism. However, we found support for a male-specific signalling role of plumage coloration. Indeed, chest yellow–red chroma and rump UV chroma were positively related to body condition in males, but only in the two islands representing the poorer environmental conditions. Therefore, environmental variability can potentially modify the reliability of achromatic displays, with possible consequences on sexual selection patterns in a nocturnal bird.     

ON THE NATURE AND ORIGIN OF THE FEATHER COLOURATION IN THE GREAT WHITE PELICAN PELECANUS ONOCROTALUS ROSEUS IN ETHIOPIA

The yellow or orange-brown colour on the breast feathers and to a lesser extent on other feathers of the Great White Pelican in a breeding colony on an island in Lake Shala, Ethiopia, is due to the presence of ferric oxide.
The feathers most probably become stained when the birds are in the water, and the source of the ferric oxide may be iron–rich silt carried into the lake by the Gidu River.     

Oil droplets of bird eyes: microlenses acting as spectral filters

An important component of the cone photoreceptors of bird eyes is the oil droplets located in front of the visual-pigment-containing outer segments. The droplets vary in colour and are transparent, clear, pale or rather intensely yellow or red owing to various concentrations of carotenoid pigments. Quantitative modelling of the filter characteristics using known carotenoid pigment spectra indicates that the pigments’ absorption spectra are modified by the high concentrations that are present in the yellow and red droplets. The high carotenoid concentrations not only cause strong spectral filtering but also a distinctly increased refractive index at longer wavelengths. The oil droplets therefore act as powerful spherical microlenses, effectively channelling the spectrally filtered light into the photoreceptor''s outer segment, possibly thereby compensating for the light loss caused by the spectral filtering. The spectral filtering causes narrow-band photoreceptor spectral sensitivities, which are well suited for spectral discrimination, especially in birds that have feathers coloured by carotenoid pigments.     

Continent-wide variation in feather colour of a migratory songbird in relation to body condition and moulting locality

Understanding the causes of variation in feather colour in free-living migratory birds has been challenging owing to our inability to track individuals during the moulting period when colours are acquired. Using stable-hydrogen isotopes to estimate moulting locality, we show that the carotenoid-based yellow-orange colour of American redstart (Setophaga ruticilla) tail feathers sampled on the wintering grounds in Central America and the Caribbean is related to the location where feathers were grown the previous season across North America. Males that moulted at southerly latitudes were more likely to grow yellowish feathers compared with males that moulted more orange-red feathers further north. Independent samples obtained on both the breeding and the wintering grounds showed that red chroma-an index of carotenoid content-was not related to the mean daily feather growth rate, suggesting that condition during moult did not influence feather colour. Thus, our results support the hypothesis that feather colour is influenced by ecological conditions at the locations where the birds moulted. We suggest that these colour signals may be influenced by geographical variation in diet related to the availability of carotenoids.     

The plumage and colouration of an enantiornithine bird from the early cretaceous of china

Brilliant colour displays and diverse feather morphologies that are often sexual ornaments are common throughout much of extant Avialae. Here we describe a new basal enantiornithine bird specimen recovered from the Early Cretaceous Jiufotang Formation of Liaoning Province in northeastern China. We present new information on the plumage of Bohaiornithidae as well as the first detailed colour reconstruction of an enantiornithine bird. The new specimen retains subadult skeletal characteristics, including periosteal pitting of the long bone epiphyses and unfused elements, while also preserving plumage evidence consistent with sexual maturity at the time of death. Exceptionally‐preserved feathers cover the body, including elongate crown feathers, body contour feathers, asymmetrically‐veined wing primaries, an alula and two elongate rachis‐dominated rectrices that may have been sexual ornaments. The crown, neck, and body contour feathers retain elongate melanosome morphologies associated with weakly iridescent colouration in extant feathers. We provide additional evidence of preserved melanin using Raman spectroscopy; a rapid, non‐destructive chemical technique. The new specimen provides data on skeletal ontogeny in the Bohaiornithidae as well as evidence for intraspecific communication functions of plumage.     

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.     

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.     

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.     

Mechanical and structural adaptations to migration in the flight feathers of a Palaearctic passerine

Current avian migration patterns in temperate regions have been developed during the glacial retreat and subsequent colonization of the ice‐free areas during the Holocene. This process resulted in a geographic gradient of greater seasonality as latitude increased that favoured migration‐related morphological and physiological (co)adaptations. Most evidence of avian morphological adaptations to migration comes from the analysis of variation in the length and shape of the wings, but the existence of intra‐feather structural adjustments has been greatly overlooked despite their potential to be under natural selection. To shed some light on this question, we used data from European robins Erithacus rubecula overwintering in Campo de Gibraltar (Southern Iberia), where sedentary robins coexist during winter with conspecifics showing a broad range of breeding origins and, hence, migration distances. We explicitly explored how wing length and shape, as well as several functional (bending stiffness), developmental (feather growth rate) and structural (size and complexity of feather components) characteristics of flight feathers, varied in relation to migration distance, which was estimated from the hydrogen stable isotope ratios of the summer‐produced tail feathers. Our results revealed that migration distance not only favoured longer and more concave wings, but also promoted primaries with a thicker dorsoventral rachis and shorter barb lengths, which, in turn, conferred more bending stiffness to these feathers. We suggest that these intra‐feather structural adjustments could be an additional, largely unnoticed, adaptation within the avian migratory syndrome that might have the potential to evolve relatively quickly to facilitate the occupation of seasonal environments.     

棉蚜体色变化的生态遗传学研究

调查了不同寄主上棉蚜刀Aphis gossypll自受精卵孵化出的自然种群、室内混合饲养以及单个饲养蚜虫的体色变化。结果表明：不论是自然还是实验种群,是群体还是个体饲养,不论寄主、栽培条件、生育期营养相同与否,棉蚜体色在世代内稳定不变,即出生时是什么颜色保持终生不变;在世代间则随温度升高体色渐变为黄色,温度降低体色逐渐转绿。伏蚜由苗蚜而来。X²检验证实：棉蚜体色变化与营养、寄主种类、光照、光质、栽培条件等无关,仅与温度密切相关,属于同一基因型在不同环境条件下的反应规范。但在太槿上还发现有个别深黄色棉蚜,从卵孵化到迁飞体色不随温度变化,表明棉蚜体色变化中还存在遗传多态现象。胚胎学观察与染色体校型分析结果证实了上述结论与观点。     

Bill colour and correlates of male quality in blackbirds: an analysis using canonical ordination

Carotenoid-dependent plumage displays are widely assumed to be honest indicators of individual health or quality, which are used as cues during mate choice and/or agonistic signalling. Despite the fact that red, yellow and orange pigmentation of bills is common, and also variable between individuals, comparatively little is known about bill colouration as a condition-dependent trait. Furthermore, many studies of avian colouration are confounded by the lack of objective colour quantification and the use of overly simplistic univariate techniques for analysis of the relationship between the condition-dependent trait and individual quality variables. In this study, we correlated male blackbird bill colour (a likely carotenoid-dependent sexually selected trait) with body/condition variables that reflect male quality. We measured bill colour using photometric techniques, thus ensuring objectivity. The data were analysed using the multivariate statistical techniques of canonical ordination. Analyses based on reflectance spectra of male blackbird bill samples and colour components (i.e. hue, chroma and brightness) derived from the reflectance spectra were very similar. Analysing the entire reflectance spectra of blackbird bill samples with Redundancy Analysis (RDA) allowed examination of individual wavelengths and their specific associations with the body/condition variables. However, hue, chroma and brightness values also provided useful information to explain colour variation, and the two approaches may be complimentary. We did not find any significant associations between male blackbird bill colour and percent incidence of ectoparasites or cloaca size. However, both the colour component and full spectral analyses showed that culmen length explained a significant amount of variation in male blackbird bill colour. Culmen length was positively associated with greater reflectance from the bill samples at longer wavelengths and a higher hue value (i.e. more orange-pigmented bills). Larger males may have larger territories or be better at defending territories during male-male interactions, ensuring access to carotenoid food sources. Future studies should elucidate the relationship between bill colour and behavioural measures such as aggressiveness, territory size, song rate and nest attendance.     

Regulation of integumentary colour and plasma carotenoids in American Kestrels consistent with sexual selection theory

1. Sexually selected traits are expected to vary seasonally, with the maximal expression of the character being evident during mate choice; however, the mechanisms controlling or regulating such traits are generally poorly known.
2. Carotenoid pigments responsible for bright red or yellow coloration in the feathers, skin or other integumentary structures of birds are generally believed to vary seasonally because of diet.
3. Variation in carotenoid-dependent skin colour between winter and spring (mating season) was investigated, as was variation in plasma carotenoids across the breeding season in captive American Kestrels, Falco sparverius , fed a uniform diet.
4. Kestrels were more brightly coloured in the mating period than in winter, and plasma carotenoid concentrations declined from the time of mating to the rearing of young.
5. Although carotenoid levels were highly sexually dimorphic during mating and laying, males and both breeding and non-breeding females all had similar levels by the incubation period, and the pattern of variation over time suggests rheostatic regulation.
6. These results suggest kestrels may have the ability to regulate (rather than merely control) their colour physiologically, the variation in colour and carotenoids is consistent with that expected of a sexually selected trait, and the loss of colour after breeding may suggest a trade-off between the show and health functions of carotenoids.