Melanin-based Plumage Coloration Research Articles

Melanin-based plumage coloration and melanin content in organs in the barn owl

Although the evolutionary ecology of melanin pigments and melanin-based coloration has been studied in great details, particularly in birds, little is known about the function of melanin stored inside the body. In the barn owl Tyto alba, in which individuals vary in the degree of reddish pheomelanin-based coloration and in the size of black eumelanic feather spots, we measured the concentration in melanin pigments in seven organs. The eyes had by far the most melanin then the skin, pectoral muscle, heart, liver, trachea, and uropygial gland. The concentration in eumelanin was not necessarily correlated with the concentration in pheomelanin suggesting that their production can be regulated independently from each other. Redder barn owls had more pheomelanin in the skin and uropygial gland than white owls, while owls displaying larger black feather spots had more eumelanin in the skin than small-spotted owls. More data are required to evaluate whether melanin-based traits can evolve as an indirect response to selection exerted on melanin deposition in organs.

Melanin-based plumage coloration is associated with exposure in tawny owls under novel conditions

Intraspecific color variation is often associated with camouflage or protection, but melanin-based color variation is also linked to behavioral and physiological aspects including boldness. In the melanin-based plumage color polymorphic tawny owl (Strix aluco), the grey morph is known to be more cryptic than the brown morph. Using 19 captive tawny owls (11 grey and 8 brown), we tested if these two color morphs that differ in camouflage tend to differently use exposed perches in a familiar environment (home aviary) and a novel environment (experimental aviary), as well as whether their response to predation risk and mobbing cues differs. The two color morphs did not differ in their exposure under known conditions (in their home aviary), but brown tawny owls were more likely to use exposed perches than grey tawny owls after release in a novel environment.Significance statementMelanin-based coloration can be associated with several behavioral traits. However, it is still unknown how predators with genetically-based color polymorphism vary in their efficiency of active background choice and risk-prone behavior while facing predation risk or mobbing harassment. Using captive tawny owls, we investigated predators’ behavior (attention towards the stimulus and use of space) while confronted to mobbing events and predation risk. We showed that tawny owls were using their space differently according to their color morph in a novel environment (experimental aviary) but not in a familiar environment (home aviary).

Does melanin-based plumage coloration reflect health status of free-living birds in urban environments?

BackgroundEcological functions and processes in urban ecosystems are governed by various human activities. City-adapted and city-exploiting animal species are expected to present certain specific behavioral and physiological traits in comparison to city-avoiders or conspecific individual frequenting less urbanized or rural environments. A trait of high importance, the plumage color polymorphism has been selected as the main study model and was correlated with different morphological and physiological parameters to highlight its importance in determining the possible health status of urban Feral Pigeons (Columba livia) in North African urban habitats.MethodsDifferent body morphometrics, hematological and hemoparasitic parameters were quantified on free-living Feral Pigeons in urban environments of northern Algeria. Moreover, plumage melanin-based coloration (MBC) was measured and the data collected at the individual scale was correlated with the previous parameters using linear and non-linear modeling approaches.ResultsPlumage MBC scores of the sampled Feral Pigeons ranged between 0.3% and 74.8%. Among the 12 morphological traits measured, body weight, tail length and total length were deemed to be positively correlated with MBC. Darker morphs appeared to have more hemoparasites compared to lighter pigeons. Quite the same observation goes with the immunity but with non-linear trends. The number of monocytes and granulocytes increased with the increase in MBC levels in lighter morphs, while pigeons with high MBC scores exhibited negative relationships between MBC levels and the number of white blood cells.ConclusionsDespite the existence of a number of studies demonstrating phenotypic directional selection, further studies are undoubtedly necessary to understand in detail the underlying mechanisms in species life-history strategies between differently colored individuals. Findings of this correlative study open exciting perspectives revealing that MBC can be considered a good indicator of and health status and adaptation strategies to changes in urban environments.

Carotenoid- but not melanin-based plumage coloration is negatively related to metal exposure and proximity to the road in an urban songbird

Rapid urbanization is a global phenomenon that is increasingly exposing organisms to novel stressors. These novel stressors can affect diverse aspects of organismal function, including development of condition-dependent ornaments, which play critical roles in social and sexual selection. We investigated the relationship between metal pollution, proximity to roads, and carotenoid- and melanin-based plumage coloration in a common songbird, the great tit (Parus major). We studied populations located across a well-characterized metal pollution gradient and surrounded by roadway networks. Metal exposure and road-associated pollution could reduce carotenoid-based pigmentation by inducing oxidative stress or affecting habitat quality, but metals could also enhance melanin-based pigmentation, through effects on melanogenesis and testosterone concentrations. Using a large sample size (N > 500), we found that birds residing close to a point source for metals had reduced ultraviolet chroma, a component of carotenoid-based pigmentation. Moreover, birds with high feather metal concentrations had lower carotenoid chroma, hue, and ultraviolet chroma, with effects modified by age class. Birds residing closer to roads also had lower carotenoid chroma and hue. Melanin-based pigmentation showed high between-year repeatability, and no association with anthropogenic pollution. Results suggest that carotenoid-, but not melanin-, based pigmentation is negatively affected by multiple anthropogenic stressors. We are the first to demonstrate a negative association between roads and a plumage-based signaling trait, which could have important implications for sexual signaling dynamics in urban landscapes.

Brightness of melanin-based plumage coloration is a cue to oxidative stress in Himalayan Black Bulbuls (Hypsipetes leucocephalus nigerrimus)

Melanin-based coloration is often found to be associated with variations in physiological and behavioral traits that are essential in avian communication. However, the function of melanin-based traits to their bearers has been investigated less than that of carotenoid-based ones. It is commonly believed that expression of melanin-based pigmentation is genetically controlled. However, whether it could be modulated by physical condition has remained controversial. In this study, we demonstrate that the level of melanin-based plumage pigmentation could be negatively correlated with an individual’s oxidative stress level. In this study, we used two groups of Black Bulbuls (Hypsipetes leucocephalus nigerrimus) obtained from a pet-shop in 2010 and 2011 to examine the correlation between the melanin-based plumage coloration and the ratio of lymphocytes to heterocytes, which is an indicator of the oxidative stress experienced by an individual. We found a negative correlation between oxidative stress levels and expression of melanin-based pigmentation on the breast and scapular feathers, but the correlation only appeared in the 2011 group, individuals of which had higher oxidative stress level and brighter plumage (i.e. more melanins) than those of the 2010 group. Our data suggest that melanin-based plumage could reflect an individual’s physical condition in certain situations. Therefore, melanin-based plumage coloration could have a function in Himalayan Black Bulbuls’ communication.

The adaptive function of melanin-based plumage coloration to trace metals

Trace metals produced by anthropogenic activities are of major importance in urban areas and might constitute a new evolutionary force selecting for the ability to cope with their deleterious effects. Interestingly, melanin pigments are known to bind metal ions, thereby potentially sequestering them in inert body parts such as coat and feathers, and facilitating body detoxification. Thus, a more melanic plumage or coat coloration could bring a selective advantage for animals living in polluted areas. We tested this hypothesis by investigating the link between melanin-based coloration and zinc and lead concentrations in feathers of urban feral pigeons, both at capture time and after one year of captivity in standardized conditions. Results show that differently coloured pigeons had similar metal concentrations at capture time. Metal concentrations strongly decreased after one year in standardized conditions, and more melanic pigeons had higher concentrations of zinc (but not lead) in their feathers. This suggests that more melanic pigeons have a higher ability to store some metals in their feathers compared with their paler counterparts, which could explain their higher success in urbanized areas. Overall, this work suggests that trace metal pollution may exert new selective forces favouring more melanic phenotypes in polluted environments.

Brownish, small and lousy barn swallows have greater natal dispersal propensity

Natal dispersal contributes to population dynamics and genetic structure. Individuals differ in whether or not they disperse and in the distance they travel from their natal site before settling to breed. Differences in natal dispersal are often associated with variation in other traits. These associations may arise because suites of morphological and behavioural traits are ultimately controlled by the same set of genes. The genes that control melanogenesis in vertebrates pleiotropically influence physiology and behaviour, including boldness and exploration. Because these personality traits predict dispersal, we tested the hypothesis that in the barn swallow, Hirundo rustica , melanic coloration predicts natal dispersal, using a solid matched case–control sampling design and a large sample. We found that males but not females with colour traits that reflect relatively more pheomelanic feather pigmentation were more likely to disperse, consistently with observations on the only other species for which dispersal in relation to plumage melanic coloration has so far been studied. To control for any confounding effects, we also analysed the association of dispersal with morphological traits and parasite infestation. Philopatric individuals were larger than dispersers, whereas dispersal strategy did not differ according to tail length, which is a sexually selected trait. Finally, philopatric females had a smaller infestation of a haematophagous louse fly. The present findings corroborate previous evidence that melanic coloration covaries with a suite of traits. In particular, they show that melanin-based plumage coloration predicts natal dispersal, independently of other factors also influencing dispersal. In addition, our results show that philopatric individuals were larger than dispersers possibly because individuals return to a benign natal place or because large body size confers an advantage in competitive interactions. Finally, they are compatible with the idea of host adaptation to local strains of a parasite with presumably small population size and low dispersing capacity. • Individuals of the same species can differ broadly in natal dispersal. • The genes for melanization affect important life history traits including behaviour. • We analyse barn swallow dispersal in relation to melanin colour, size and parasitism. • Darker, smaller and more infested swallows were more likely to disperse from colony. • Melanin-based coloration may reflect different life history strategies.

Sex-specific phenotypic integration: endocrine profiles, coloration, and behavior in fledgling boobies

The intensity of color expression in animals plays a key role in social environments as a mechanism to signal individual capacities in competitive contests. Selective pressures for resource competition differ at different stages of life and between sexes; therefore, coloration is expected to vary between juveniles and adults and between males and females. Exploring the covariance between coloration and other traits may help to understand the functional significance of color and the action of natural selection on multivariate phenotypes. Melanin-based plumage coloration was investigated in the masked booby Sula dactylatra in relation to melanin concentration, sex, hormone levels, and shy–bold behavior of chicks close to fledging. Darker brown boobies showed higher levels of both eumelanin and pheomelanin concentration and lower body mass. Males behaved bolder than females and showed on average 8% larger brown patches. Bolder females had smaller brown patches. Bolder individuals also had lower levels of circulating testosterone, but no differences in corticosterone levels were found. Stronger phenotypic integration was observed in females than males. Our study suggests that juvenile melanic coloration may reflect behavioral strategies by sex, endocrine profiles, and body mass indicating the convergence of different adaptive functions in a given phenotype, this being more evident in females. Direction of correlations differed from those predicted under the pleiotropic idea for color-related traits. These results suggest the possibility that juvenile plumage acts as a signaling system in a social context within the age class and suggest that plumage coloration may indicate different behavioral strategies.

Signaling stress? An analysis of phaeomelanin-based plumage color and individual corticosterone levels at two temporal scales in North American barn swallows, Hirundo rustica erythrogaster

Sexually selected traits confer greater reproductive benefits to individuals with more elaborate forms of the signal. However, whether these signals convey reliable information about the physiology underlying trait development remains unknown in many species. The steroid hormone corticosterone (CORT) mediates important physiological and behavioral processes during the vertebrate stress response, and CORT secretion itself can be modulated by melanocortins. Thus, sexually selected melanin-based plumage coloration could function as an honest signal of an individual's ability to respond to stressors. This hypothesis was tested in North American barn swallows, Hirundo rustica erythrogaster, where males with darker ventral plumage color exhibit higher phaeomelanin content and are more successful at reproduction. Because reproductive behavior occurs months after plumage signals are developed, we also addressed the potential temporal disconnect of physiological state during trait development and trait advertisement by analyzing three different measurements of CORT levels in adult males during the breeding season (trait advertisement) and in nestling males while they were growing their feathers (trait development). Variation in adult plumage color did not predict baseline or stress-induced CORT, or stress responsiveness. Likewise, there was no relationship between nestling plumage color and any of the CORT measurements, but heavier nestlings had significantly lower baseline CORT. Our finding that a predominantly phaeomelanin-based trait is unrelated to circulating CORT suggests that phaeomelanin and eumelanin signals may convey different physiological information, and highlights the need for further study on the biochemical links between the hypothalamic–pituitary–adrenal (HPA) axis and the production of different melanin-based pigments.

House Finches Are Not Just What They Eat: A Reply To Hill

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Evolution of sexual dichromatism: contribution of carotenoid- versus melanin-based coloration

In birds, carotenoid-based plumage coloration is more dependent on physical condition and foraging abilities and less constrained developmentally than is melanin-based coloration. Thus, female mate choice for honest signals should result in more intense sexual selection on carotenoid- than on melanin-based plumage coloration. Using variation in sexual dimorphism as an indirect measure of the intensity of sexual selection, we tested the prediction that variation in sexual dimorphism is driven more by change in carotenoid-based coloration between males and females than by change in melanin-based coloration. Examination of historical changes in carotenoid- versus melanin-based pigmentation in 126 extant species of Cardueline finches supported this prediction. We found that carotenoid-derived coloration changed more frequently among congeners than melanin-based coloration. In both sexes, increase in carotenoid-based coloration score, but not in melanin-based coloration score, was strongly associated with increase in sexual dichromatism. In addition, sexual dimorphism in carotenoid-based coloration contributed more to overall dichromatism than dimorphism in melanin-based plumage. Our results supported the hypothesis that melanin-based and carotenoid-based coloration have fundamentally different signal content and suggest that combining melanin-based and carotenoid-based coloration in comparative analyses is not appropriate.

Evolution of sexual dichromatism: contribution of carotenoid- versus melanin-based coloration

In birds, carotenoid-based plumage coloration is more dependent on physical condition and foraging abilities and less constrained developmentally than is melanin-based coloration. Thus, female mate choice for honest signals should result in more intense sexual selection on carotenoid- than on melanin-based plumage coloration. Using variation in sexual dimorphism as an indirect measure of the intensity of sexual selection, we tested the prediction mat variation in sexual dimorphism is driven more by change in carotenoid-based coloration between males and females than by change in melanin-based coloration. Examination of historical changes in carotenoid- versus melanin-based pigmentation in 126 extant species of Cardueline finches supported this prediction. We found that carotenoid-derived coloration changed more frequently among congeners than melanin-based coloration. In both sexes, increase in carotenoid-based coloration score, but not in melanin-based coloration score, was strongly associated with increase in sexual dichromatism. In addition, sexual dimorphism in carotenoid-based coloration contributed more to overall dichromatism than dimorphism in melanin-based plumage. Our results supported the hypothesis that melanin-based and carotenoid-based coloration have fundamentally different signal content and suggest that combining melanin-based and carotenoid-based coloration in comparative analyses is not appropriate.

Melanin–based plumage coloration in the house finch is unaffected by coccidial infection

For most species of birds, ornamental plumage coloration may result from two types of pigments: carotenoids and melanins. Despite the fact that melanin pigments can be synthesized by birds from basic, amino acid precursors, while carotenoids cannot be synthesized by birds and must be ingested, melanin-based plumage coloration and carotenoid-based plumage coloration have often been treated as a single trait in investigations of the function and evolution of plumage coloration. Expression of carotenoid-based coloration is known to be dependent on condition, while the effects of individual condition have not been well-tested for expression of melanin-based coloration. In this study, we experimentally tested the effect of coccidial infection of the intestinal tract of male house finches during moult on expression of melanin-based plumage coloration. Coccidial infection had a significant negative effect on carotenoid-based coloration, but it had no significant effect on melanin-based feather coloration. Unlike carotenoid-based coloration, melanin-based coloration may be cheap to produce, and honesty of melanin-based coloration my require social mediation.