Pigmentation In Animals Research Articles

The level of putative carotenoid-binding proteins determines the body color in two species of endemic Lake Baikal amphipods.

Color is an essential clue for intra- and interspecies communication, playing a role in selection and speciation. Coloration can be based on nanostructures and pigments; carotenoids and carotenoproteins are among the most widespread pigments in animals. Over 350 species and subspecies of amphipods (Crustacea: Amphipoda) endemic to Lake Baikal exhibit an impressive variability of colors and coloration patterns, including intraspecific color morphs. However, the mechanisms forming this diversity are underexplored, as while the carotenoid composition of several transparent, green, and red species was investigated, there have been no reports on the corresponding carotenoid-binding proteins. In this work, we analyze the coloration of two brightly colored Baikal amphipods characterized by intraspecific color variability, Eulimnogammarus cyaneus and E. vittatus. We showed that the color of either species is defined by the level of putative carotenoid-binding proteins similar to the pheromone/odorant-binding protein family, as the concentration of these putative crustacyanin analogs was higher in blue or teal-colored animals than in the orange- or yellow-colored ones. At the same time, the color did not depend on the total carotenoid content, as it was similar between animals of contrasting color morphs. By exploring the diversity of these sequences within a larger phylogeny of invertebrate crustacyanins, we show that amphipods lack orthologs of the well-studied crustacyanins A and C, even though they possess some crustacyanin-like sequences. The analysis of expression levels in E. cyaneus showed that the transcripts encoding crustacyanin analogs had much higher expression than the crustacyanin-like sequences, suggesting that the former indeed contribute to the color of these brightly colored animals. The crustacyanin analogs seem to act in a similar way to the well-studied crustacyanins in body color formation, but the details of their action are still to be revealed.

The dark side of coloration: Ecogeographical evidence supports Gloger's rule in American marsupials.

Geographical distribution of color phenotypes and associations with ecological predictors remains poorly understood. An important geographic pattern concerning this topic is Gloger's rule, which predicts the increase of pigmentation in endothermic animals from cold and dry to warm and wet environments. Didelphid marsupials exhibit a variety of color patterns, ranging from light and dark uniform to more complex colorations. However, surprisingly little is known about the adaptive significance of dark coloration in this singular group of mammals. Using a phylogenetic comparative approach, we investigated whether coloration in different body regions of didelphids (i.e., dorsum and face) is associated with variables representing heat and humidity of the environment, as predicted by Gloger's rule. We demonstrated that Gloger's rule explains the interspecific color variation in American marsupials, especially when considering the facial region. Thus, dark coloration was more frequent among didelphid species occupying warm and wet environments than cold and dry environments. We also discuss the selective forces that can potentially explain coat color variation in didelphid marsupials, including camouflage, pathogen resistance, and pleiotropy hypotheses.

Exposure to UV radiance predicts repeated evolution of concealed black skin in birds

Plumage is among the most well-studied components of integumentary colouration. However, plumage conceals most skin in birds, and as a result the presence, evolution and function of skin colour remains unexplored. Here we show, using a database of 2259 species encompassing >99% of bird genera, that melanin-rich, black skin is found in a small but sizeable percentage (~5%) of birds, and that it evolved over 100 times. The spatial distribution of black skin follows Gloger’s rule, which states that pigmentation of endothermic animals increases towards the equator. Furthermore, most black-skinned birds inhabit high irradiation regions, and tend to be bald and/or have white feathers. Thus, taken together, our results suggest that melanin-rich, black skin helps to protect birds against ultraviolet irradiation. More generally, our results illustrate that feathered skin colour varies taxonomically, ontogenetically and temporally, providing an additional dimension for avian colour research.

Genome and Transcriptome Sequencing of casper and roy Zebrafish Mutants Provides Novel Genetic Clues for Iridophore Loss.

casper has been a widely used transparent mutant of zebrafish. It possesses a combined loss of reflective iridophores and light-absorbing melanophores, which gives rise to its almost transparent trunk throughout larval and adult stages. Nevertheless, genomic causal mutations of this transparent phenotype are poorly defined. To identify the potential genetic basis of this fascinating morphological phenotype, we constructed genome maps by performing genome sequencing of 28 zebrafish individuals including wild-type AB strain, roy orbison (roy), and casper mutants. A total of 4.3 million high-quality and high-confidence homozygous single nucleotide polymorphisms (SNPs) were detected in the present study. We also identified a 6.0-Mb linkage disequilibrium block specifically in both roy and casper that was composed of 39 functional genes, of which the mpv17 gene was potentially involved in the regulation of iridophore formation and maintenance. This is the first report of high-confidence genomic mutations in the mpv17 gene of roy and casper that potentially leads to defective splicing as one major molecular clue for the iridophore loss. Additionally, comparative transcriptomic analyses of skin tissues from the AB, roy and casper groups revealed detailed transcriptional changes of several core genes that may be involved in melanophore and iridophore degeneration. In summary, our updated genome and transcriptome sequencing of the casper and roy mutants provides novel genetic clues for the iridophore loss. These new genomic variation maps will offer a solid genetic basis for expanding the zebrafish mutant database and in-depth investigation into pigmentation of animals.

Pigmentation formation and expression analysis of tyrosinase in Siniperca chuatsi.

Animal pigmentation primarily depends on the presence and mixing ratio of chromatophores, functioning in animal survival and communication. For the benthic and carnivorous Siniperca chuatsi, pigmentation pattern is key to concealment and predation. In this study, the formation, distribution, and main pattern of chromatophores were observed in the embryos, larvae, skins, and visceral tissues from S. chuatsi. Melanophores were firstly visualized in the yolk sac at segmentation stage, and then they were migrated to the whole body and further clustered into the black stripes, bands, and patches. In adult S. chuatsi, the head, black band, and body side skins mainly contained melanophores, showing as deep or light black. The abdomen skin mainly contained iridophores, showing as silvery. In the eye, the pigment layers were located in the epithelial layers of iris and retina and shown as black. Then, the pigmentation-related gene, tyrosinase gene from S. chuatsi (Sc-tyr) was analyzed by bioinformatics and quantitative methods. The Sc-tyr gene encoded a protein with 540 amino acids (Sc-TYR). The Sc-TYR contained two copper ion binding sites, which were coordinated by six conserved histidines (H182, H205, H214, H366, H370, H393) and necessary for catalytic activity. The Sc-TYR was well conserved compared with TYR of various species with higher degree of sequence similarity with other fishes (77.6-98.3%). The qRT-PCR test showed that the Sc-tyr mRNA reached the peak value at segmentation stage in the embryo development, the black skins displayed a higher expression level than that in silvery skin, and the eye had the highest expression level compared with other tissues. Further research on enzyme activity showed that the expression patterns of tyrosinase activity were similar to that of the Sc-tyr mRNA. Comparing with the results of molecular and phenotype, it was found that the temporal and spatial distributions of tyrosinase corresponded well with changes in pigmentation patterns and the intensity of skin melanization. This study initially explored the pigmentation formation and tyrosinase expression, which served as a foundation for further insight into the genetics mechanism of body color formation in S. chuatsi.

Impairment of mixed melanin-based pigmentation in parrots.

Parrots and allies (Order Psittaciformes) have evolved an exclusive capacity to synthesize polyene pigments called psittacofulvins at feather follicles, which allows them to produce a striking diversity of pigmentation phenotypes. Melanins are polymers constituting the most abundant pigments in animals, and the sulphurated form (pheomelanin) produces colors that are similar to those produced by psittacofulvins. However, the differential contribution of these pigments to psittaciform phenotypic diversity has not been investigated. Given the color redundancy, and physiological limitations associated to pheomelanin synthesis, we hypothesized that the latter would be avoided by psittaciform birds. Here we test this by using Raman spectroscopy to identify pigments in feathers exhibiting colors suspicious of being produced by pheomelanin (i.e., dull red, yellow and grey- and green-brownish) in 26 species from the three main lineages of Psittaciformes. We detected the non-sulphurated melanin form (eumelanin) in black, grey and brown plumage patches, and psittacofulvins in red, yellow and green patches, but no evidence of pheomelanin. As natural melanins are assumed to be composed of eumelanin and pheomelanin in varying ratios, our results represent the first report of impairment of mixed melanin-based pigmentation in animals. Given that psittaciforms also avoid the uptake of circulating carotenoid pigments, these birds seem to have evolved a capacity to avoid functional redundancy between pigments, likely by regulating follicular gene expression. Ours study provides the first vibrational characterization of different psittacofulvin-based colors and thus helps to determine the relative polyene chain length in these pigments, which is related to their antireductant protection activity.

A comparative microstructural study of the animal pigment, bilirubin, in seed arils of Strelitzia nicolai

The unexpected finding of an animal pigment, bilirubin, in the arils of the plant Strelitzia nicolai, has resulted in interest regarding bilirubin's formation, its ultimate function as well as its ultrastructure both in the humans and in the plant. The main objective of this study was to identify the structural components of seed arils of S. nicolai at a cellular level. Furthermore, this study compared the five different stages of aril maturation and tracked bilirubin formation within the cells. This was accomplished by stereomicroscopic, light microscopic and electron microscopical analysis. These included arils stained with toluidine blue and the modified Hall's staining to detect the presence of bilirubin. The results from the light, scanning electron microscopy (SEM) and transmission electron microscopy (TEM) showed the different stages of aril development containing a large number of cellular contents. However, as the aril matures, the contents of the cell became denser. Additionally, arils became a more structured and rigid as they matured. The most noticeable feature of the arils was that of the colour change as they mature. It was clarified that the colour change of the arils was attributed to the production and accumulation of bilirubin within the cells.

Convergent evolution of cytochrome P450s underlies independent origins of keto-carotenoid pigmentation in animals.

Keto-carotenoids contribute to many important traits in animals, including vision and coloration. In a great number of animal species, keto-carotenoids are endogenously produced from carotenoids by carotenoid ketolases. Despite the ubiquity and functional importance of keto-carotenoids in animals, the underlying genetic architectures of their production have remained enigmatic. The body and eye colorations of spider mites (Arthropoda: Chelicerata) are determined by β-carotene and keto-carotenoid derivatives. Here, we focus on a carotenoid pigment mutant of the spider mite Tetranychus kanzawai that, as shown by chromatography, lost the ability to produce keto-carotenoids. We employed bulked segregant analysis and linked the causal locus to a single narrow genomic interval. The causal mutation was fine-mapped to a minimal candidate region that held only one complete gene, the cytochrome P450 monooxygenase CYP384A1, of the CYP3 clan. Using a number of genomic approaches, we revealed that an inactivating deletion in the fourth exon of CYP384A1 caused the aberrant pigmentation. Phylogenetic analysis indicated that CYP384A1 is orthologous across mite species of the ancient Trombidiformes order where carotenoids typify eye and body coloration, suggesting a deeply conserved function of CYP384A1 as a carotenoid ketolase. Previously, CYP2J19, a cytochrome P450 of the CYP2 clan, has been identified as a carotenoid ketolase in birds and turtles. Our study shows that selection for endogenous production of keto-carotenoids led to convergent evolution, whereby cytochrome P450s were independently co-opted in vertebrate and invertebrate animal lineages.

Pheomelanin synthesis varies with protein food abundance in developing goshawks.

The accumulation of the amino acid cysteine in lysosomes produces toxic substances, which are avoided by a gene (CTNS) coding for a transporter that pumps cystine out of lysosomes. Melanosomes are lysosome-related organelles that synthesize melanins, the most widespread pigments in animals. The synthesis of the orange melanin, termed pheomelanin, depends on cysteine levels because the sulfhydryl group is used to form the pigment. Pheomelanin synthesis may, therefore, be affected by cysteine homeostasis, although this has never been explored in a natural system. As diet is an important source of cysteine, here we indirectly tested for such an effect by searching for an association between food abundance and pheomelanin content of feathers in a wild population of Northern goshawk Accipiter gentilis. As predicted on the basis that CTNS expression may inhibit pheomelanin synthesis and increase with food abundance as previously found in other strictly carnivorous birds, we found that the feather pheomelanin content in nestling goshawks, but not in adults, decreased as the abundance of prey available to them increased. In contrast, variation in the feather content of the non-sulphurated melanin form (eumelanin) was only explained by sex in both nestlings and adults. We also found that the feather pheomelanin content of nestlings was negatively related to that of their mothers, suggesting a relevant environmental influence on pheomelanin synthesis. Overall, our findings suggest that variation in pheomelanin synthesis may be a side effect of the maintenance of cysteine homeostasis. This may help explaining variability in the expression of pigmented phenotypes.

Метод электронной спектроскопии диффузного отражения в исследовании мышечной ткани диких и домашних животных

The aim of the work was to study the application of diffuse reflection electron spectroscopy (DRES) method for obtaining the optical surface characteristics of slices and chopped muscle tissue, muscle fibre and sarcoplasmic protein extracts of wild (wild boar, elk, deer, freshwater and marine fish) and domestic (pork, beef, lamb) animal tissue depending on the following processing conditions: freezing, water extraction, lyophilisation and enzyme-microbial exposure. Electronic spectra of transverse sections of whole-piece material, ground muscle tissue, muscle fibre and aqueous extracts were recorded using a Specord M-200 spectrophotometer (AIZ Engineering GmbH, Germany) having a vertical pathway relative to the Spectralon standard in the wavelength range of 200-700 nm. During testing of biological tissues of animal origin in solid and liquid state, the high sensitivity, resolution and analytical features of the express DRES method was shown to allow the acquisition of factual information concerning changes in the spectral characteristics of their main components under experimental conditions. Regardless of the species and type of animal organism, proteins, mucopolysaccharides, lipids, pigment protein and various oxidised forms demonstrate absorption bands in four clearly differentiated regions of the electromagnetic spectrum: 200-300, 400-425, 300-380, 540-580 and 635 nm, respectively. A comparative study of the pork and beef muscle tissue demonstrated significant differences in the nature of changes in the spectral characteristics of the main components and components of muscle tissue and muscle fibre, depending on the degree of oxidation, mechanical destruction, strength of myofibrils and pigment protein myoglobin associated with them.

Non-Synonymous variants in premelanosome protein (PMEL) cause ocular pigment dispersion and pigmentary glaucoma.

Pigmentary glaucoma (PG) is a common glaucoma subtype that results from release of pigment from the iris, called pigment dispersion syndrome (PDS), and its deposition throughout the anterior chamber of the eye. Although PG has a substantial heritable component, no causative genes have yet been identified. We used whole exome sequencing of two independent pedigrees to identify two premelanosome protein (PMEL) variants associated with heritable PDS/PG. PMEL encodes a key component of the melanosome, the organelle essential for melanin synthesis, storage and transport. Targeted screening of PMEL in three independent cohorts (n=394) identified seven additional PDS/PG-associated non-synonymous variants. Five of the nine variants exhibited defective processing of the PMEL protein. In addition, analysis of PDS/PG-associated PMEL variants expressed in HeLa cells revealed structural changes to pseudomelanosomes indicating altered amyloid fibril formation in five of the nine variants. Introduction of 11-base pair deletions to the homologous pmela in zebrafish by the clustered regularly interspaced short palindromic repeats (CRISPR)-Cas9 method caused profound pigmentation defects and enlarged anterior segments in the eye, further supporting PMEL's role in ocular pigmentation and function. Taken together, these data support a model in which missense PMEL variants represent dominant negative mutations that impair the ability of PMEL to form functional amyloid fibrils. While PMEL mutations have previously been shown to cause pigmentation and ocular defects in animals, this research is the first report of mutations in PMEL causing human disease.

Transcriptional regulation of Rab32/38, a specific marker of pigment cell formation in Ciona robusta

Through a myriad of pigments stored in different cells, animal pigmentation represents a crucial process to face disparate environmental and ecological challenges. In vertebrates, the small GTPase Rab32 and Rab38 have a conserved role in the transport of key melanogenic enzymes, as tyrosinase (tyr) and tyrosinase-related protein (tyrp), to the melanosomes in formation. We provide a survey on Rab32/38 evolution and its regulatory logics during pigment cell formation in Ciona robusta. Our phylogeny supports the existence of a single Rab32/38 gene in tunicates, which is probably the unique transporter for tyrosinase family members in this clade. Different deletions allow us to identify the minimal cis-regulatory element able to recapitulate the endogenous gene expression during pigment cell development in C. robusta. In this conserved region, we identified two putative binding sites for the transcription factor Mitf, which is known for its role as regulator of pigmentation in vertebrates. Mutational analysis revealed that both Mitf binding sites are essential for the activity of this regulatory region and we demonstrated that Mitf misexpression is able to induce ectopic activation of the Rab32/38 regulatory region in vivo.Our results strongly indicate that Mitf is involved in the regulation of Rab32/38 activity during Ciona pigment cell development.

Differential predation alters pigmentation in threespine stickleback (Gasterosteus aculeatus).

Animal pigmentation plays a key role in many biological interactions, including courtship and predator avoidance. Sympatric benthic and limnetic ecotypes of threespine stickleback (Gasterosteus aculeatus) exhibit divergent pigment patterns. To test whether differential predation by cutthroat trout contributes to the differences in pigmentation seen between the ecotypes, we used a within-generation selection experiment on F2 benthic-limnetic hybrids. After 10months of differential selection, we compared the pigmentation of fish under trout predation to control fish not exposed to trout predation. We found that stickleback exhibited more lateral barring in ponds with trout predation. Ponds with trout were also less turbid, and a greater degree of barring was negatively correlated with the magnitude of turbidity across pond replicates. A more benthic diet, a proxy for habitat use, was also correlated with greater lateral barring and green dorsal pigmentation. These patterns suggest that differential exposure to cutthroat trout predation may explain the divergence in body pigmentation between benthic and limnetic ecotypes.

Solar and terrestrial radiations explain continental-scale variation in bird pigmentation.

Animals living on the earth's surface are protected from the damaging effects of solar ultraviolet (UV) radiation by melanin pigments that color their integument. UV levels that reach the earth's surface vary spatially, but the role of UV exposure in shaping clinal variations in animal pigmentation has never been tested. Here, we show at a continental scale in Europe that golden eagles Aquila chrysaetos reared in territories with a high solar UV-B radiation exposure deposit lower amounts of the sulphurated form of melanin (pheomelanin) in feathers and consequently develop darker plumage phenotypes than eagles from territories with lower radiation exposure. This clinal variation in pigmentation is also explained by terrestrial γ radiation levels in the rearing territories by a similar effect on the pheomelanin content of feathers, unveiling natural radioactivity as a previously unsuspected factor shaping animal pigmentation. These findings show for the first time the potential of solar and terrestrial radiations to explain pigmentation phenotype diversity in animals, including humans, at large spatial scales.

Molecular vibration as a novel explanatory mechanism for the expression of animal colouration.

Animal colouration is characterized by the concentration of pigments in integumentary structures and by the nanoscale arrangement of constitutive elements. However, the influence of molecular vibration on colour expression has been overlooked in biology. Molecular vibration occurs in the infrared spectral region, but vibrational and electronic properties can influence each other. Thus, the vibration of pigment molecules may also affect their absorption properties and the resulting colours. We calculated for the first time the relative contribution of molecular vibration (by means of Raman spectroscopy) and concentration (by means of HPLC) of melanin polymers, the most common animal pigments, to generate diversity in plumage colour in 47 species of birds. Vibrational characteristics explained >9 times more variance in colour expression than the concentration of melanins. Additionally, we modelled melanin Raman spectra on the basis of the chemical structure of their constituent monomers and calculated the Huang-Rhys factors for each vibrational mode, which indicate the contribution of these modes to the electronic spectra responsible for the resulting colours. High Huang-Rhys factors frequently coincided with the vibrational modes of melanin monomers. Our results can be explained by the influence of molecular vibration on the absorption properties of melanins. The colour of organisms may thus mainly result from the vibrational properties of their molecules and only residually from their concentration. As a given melanin concentration can give rise to different colours because different structural melanin conformations can present different vibrational characteristics, vibrational effects may favour phenotypic plasticity and thus constitute an important evolutionary force.

Dark pigmentation limits thermal niche position in birds

Abstract Animal pigmentation has evolved because of several adaptive functions. In the case of pigmentation produced by melanins, the most common pigments in animals, the main function is protection against UV radiation. However, pigmentation also affects animal surface's ability to absorb solar radiation and gain heat, which may represent a thermal constraint for endotherms due to their relatively high and constant body temperatures. As darker colours absorb more radiation than lighter colours, dark‐pigmented endotherm animals may exhibit limited performance under high ambient temperatures and thus be constrained at occupying hot environments. While the influence of pigmentation on the determination of the thermal niches of ectotherms, particularly reptiles, has been the focus of several studies, this remains an unexplored issue for endotherm animals. Here, we made a detailed quantification of the expression of pigmentation phenotypes produced by melanins in 96 species of birds inhabiting the Spanish sector of the Iberian Peninsula and estimated their climate niche position by calculating the effects of ambient temperature, insolation and precipitation on bird occurrence at a fine 10 × 10 km spatial scale. After controlling for the body size and the nocturnal condition of species, and for phylogenetic and spatial effects, we found a positive association between plumage reflectance and the functional response of bird distribution to spring–summer ambient temperature and insolation but not to precipitation. Thus, bird species preferentially occupying the hotter and sunnier areas of Spain exhibit lighter plumage pigmentation. Our findings suggest that darker birds are limited from occupying environments with high temperatures, unveiling a constraint in endotherms imposed by their pigmentation phenotype. A plain language summary is available for this article.

Reproductive melanization may protect sperm from harmful solar radiation

Animal pigmentation is incredibly diverse, serving a variety of functions. However, the function of the pigmentation that surrounds the testes of some vertebrates is unknown. Why are the tissues surrounding the testes (scrotum, tunica vaginalis and/or tunica albuginea) melanized in some species but not others? We examined this question in bats, where there is extensive species-specific variation in the presence of darkly pigmented (melanized) tissues surrounding the male gonads, as well as diversity in their ecologies, mating systems, and physiologies. Using data from 136 species of bats, we found that melanin surrounding the testes and epididymis is associated with more exposed roosting sites (presumably with greater sun exposure- and UV radiation), and with the occurrence of long-term male sperm storage. These findings suggest that scrotal melanin may protect mature sperm from UV damage, and from oxidative damage in species with male sperm storage. We found no evidence of an association between group size or mating system with reproductive melanin, or that phylogeny explains the distribution of pigmentation. Although our study suggests that scrotal melanin may protect sperm, the mechanism remains unknown. We outline several avenues for future work on reproductive pigmentation aimed at investigating additional roles of reproductive melanization in male bats.

Oxygen consumption and swimming performance in Arctic charr with different pigmentation patterns

Pigmentation in animals often reflects behavioural and physiological traits, such as health status, stress responsiveness, and dominance. Individual variation in stress response and behaviour has earlier been proposed to be connected to differences in metabolism. The aim of this study was to investigate the connection between pigmentation, behaviour, and oxygen consumption in Arctic charr (Salvelinus alpinus). In this species, we have earlier found an association between number of carotenoid spots on the skin and physiological response to stress (stress coping style). Swimming endurance and respiration rates were estimated from video recorded behaviour and oxygen consumption in a swim tunnel. Flow velocity was kept as 1.6 BL (body lengths) s−2 for 60 min, and oxygen consumption (mg min−1 g−1) during that time was used as a proxy for metabolism. Oxygen consumption was negatively correlated with number of spots. Also behaviour scores from a principal component analysis varied with pigmentation, with a negative correlation between number of spots and behaviour scores connected with endurance. Fish with fewer spots rested more against the rear of the chamber, and fish with more spots were sooner, and more often, pressed to the rear grid by the water current. The variation in oxygen consumption in the swim tunnel indicates a relation between respiration rate, swimming endurance, and pigmentation in the Arctic charr. With earlier findings on covariations between spot numbers and stress coping in this species, there seems to be connections between stress coping style, behaviour, and metabolism. Thus, it would be possible to identify individual expression of these features, based on the pigmentation patterns of the fish.

Melanin in a changing world: brown trout coloration reflects alternative reproductive strategies in variable environments

Melanins are the most widespread pigments in animals but their adaptive significance remains elusive. Recent studies suggest that intraspecific variation in melanin-based coloration reflects individual genetic-based alternative strategies to cope with environment variability, which could be crucial for their responses to climate changes. However, empirical evidence is still scarce. In this study, we tested how skin coloration in natural populations of brown trout Salmo trutta fario would reflect alternative reproductive strategies in different environments. We experimentally manipulated the flow regime (constant vs. variable) in artificial streams and compared the reproductive investment (body mass and plasma triglyceride variations), innate immunity (variations in plasma peroxidase and lysozyme activity) and reproductive success (number of mates and offspring) of differently colored brown trout over 2 reproductive seasons. Results show that darker males had a higher reproductive investment, but similar immune variations during reproduction compared to paler males. In addition, this reproductive investment was higher in variable environments. However, this did not translate into a higher reproductive success in variable environments, as darker males had a similar number of mates and offspring compared to their paler counterparts under a variable water flow. Since climate change will likely lead to an increased flow variability in the next decades, this suggests that darker brown trout could incur a higher energetic cost of reproduction and could be more impacted by climate changes than their paler counterparts. This highlights the need to take into account intraspecific variability to better forecast the response of natural populations to climate changes.

Gyrfalcons Falco rusticolus adjust CTNS expression to food abundance: a possible contribution to cysteine homeostasis.

Melanins form the basis of animal pigmentation. When the sulphurated form of melanin, termed pheomelanin, is synthesized, the sulfhydryl group of cysteine is incorporated to the pigment structure. This may constrain physiological performance because it consumes the most important intracellular antioxidant (i.e., glutathione, GSH), of which cysteine is a constitutive amino acid. However, this may also help avoid excess cysteine, which is toxic. Pheomelanin synthesis is regulated by several genes, some of them exerting this regulation by controlling the transport of cysteine in melanocytes. We investigated the possibility that these genes are epigenetically labile regarding protein intake and thus contribute to cysteine homeostasis. We found in the Icelandic population of gyrfalcon Falco rusticolus, a species that pigments its plumage with pheomelanin, that the expression of a gene regulating the export of cystine out of melanosomes (CTNS) in feather melanocytes of developing nestlings increases with food abundance in the breeding territories where they were reared. The expression of other genes regulating pheomelanin synthesis by different mechanisms of influence on cysteine availability (Slc7a11 and Slc45a2) or by other processes (MC1R and AGRP) was not affected by food abundance. As the gyrfalcon is a strict carnivore and variation in food abundance mainly reflects variation in protein intake, we suggest that epigenetic lability in CTNS has evolved in some species because of its potential benefits contributing to cysteine homeostasis. Potential applications of our results should now be investigated in the context of renal failure and other disorders associated with cystinosis caused by CTNS dysfunction.