Phenotypic correlates of yolk and plasma carotenoid concentration in yellow-legged gull chicks

Abstract Carotenoids perform important biological actions in animal tissues, including contributing antioxidant protection. However, the function of transmission of maternal carotenoids to bird eggs is still largely unknown. We made a yolk biopsy of yellow-legged gull (Larus michahellis) eggs and found that the concentration of lutein declined with laying date and across the laying order and increased with egg mass. The concentration of all the main carotenoids (lutein, zeaxanthin, and dehydrolutein) pooled also declined with date and increased with egg mass. We also performed a partial reciprocal cross-fostering of eggs between clutches and investigated the covariation between morphology, T cell-mediated immunity, and plasma carotenoid concentrations of the chicks and carotenoid concentrations in their original eggs. Absolute plasma carotenoid concentrations did not covary with those in the yolk, whereas a positive covariation was found for relative concentrations. Yolk and absolute plasma carotenoid concentrations positively predicted chick body mass and size but not the intensity of the cell-mediated immune response. Thus, yolk carotenoid concentrations may affect chick carotenoid profile and growth, possibly mediating early maternal effects. However, rearing conditions also contributed to determining relative concentrations of circulating carotenoids. Since yolk or plasma antioxidant capacity did not correlate with carotenoid concentrations, future studies of maternal effects mediated by antioxidants should integrate information on carotenoids with information on other components of the antioxidant systems.     

Yolk carotenoids increase fledging success in great tit nestlings

Avian mothers can influence offspring phenotype through the deposition of different compounds into eggs, such as antibodies, hormones and antioxidants. The concentration of carotenoids in yolk is larger than in maternal plasma, suggesting an important role of these compounds for offspring development. Since carotenoids have to be acquired from the diet, they may be available in limiting amounts to the mothers. Here, we investigated the role of egg carotenoids for offspring growth by experimentally increasing the concentration of yolk lutein, the main carotenoid in great tit (Parus major) yolk. We subsequently measured body condition, oxidative stress, immune response, plumage colouration and fledging success. Lutein increased body mass soon after hatching and fledging success, but did not affect tarsus length, oxidative stress, immune response and plumage colouration. The higher content of yolk lutein could have increased body mass by reducing oxidative stress caused by high metabolic rates of rapidly growing embryos or by promoting cell differentiation and proliferation. The positive effect of lutein on fledging success seems to be mediated by its influence on body mass 3 days post-hatch, since these two traits were correlated. The finding that our treatment did not affect traits measured later in the nestling period, except for fledging success, suggests that yolk lutein has short-term effects that are essential to increase survival until fledging. Our study shows the positive effect of yolk lutein on offspring survival in the great tit, and therefore suggests an important role of carotenoid-mediated maternal effects.     

Effects of carotenoid availability during laying on reproduction in the blue tit

Carotenoids are antioxidant pigments involved in several physiological processes and signalling in animals that cannot synthesise them and therefore must acquire them from food. We experimentally investigated the effects of carotenoid availability in the diet during egg laying on antioxidant deposition in egg yolk and the related effects on nestling condition, female body condition and parental investment in the blue tit (Parus caeruleus). Carotenoid supplementation of egg-laying females resulted in a significant increase in carotenoid concentration in egg yolk, but not in vitamin E or A concentration. There was no relationship between yellow plumage colour of adult females and carotenoid deposition in eggs, and no differential effect of feeding treatment depending on female colour. Nestlings from eggs laid by carotenoid supplemented females had longer tarsi, had faster development of the immune system as reflected by leukocyte concentration in blood, and grew brighter yellow feathers than nestlings from control females. However, nestlings from the two groups did not differ significantly in body mass, plasma antioxidants or plumage colour hue. At the time of chick rearing, carotenoid-fed females had increased plasma vitamin E levels compared to controls. However, females from the two treatment groups did not differ significantly in body condition or feeding rate. These results suggest that carotenoid availability is limiting during egg laying, and that females may have to balance the benefits of investing in egg quality against the potential costs of impairing their own future antioxidant protection. In addition, there may be considerable variation in carotenoid availability not only across seasons, but also among different stages of the breeding season.     

No effect of carotenoid supplementation on phytohemagglutinin response or body condition of nestling house wrens

Carotenoids are an essential and often limiting resource in animals and play important roles in immune system function. In birds, the period shortly after hatching is an energetically demanding stage characterized by rapid growth in body size and organ systems, including the immune system. Availability of carotenoids for the growing nestlings may be of particular importance and potentially limiting at this stage of development. We tested the hypothesis that the availability of carotenoids for the embryo in the egg and in the diet of nestlings limits the condition and immune responses of nestling house wrens (Troglodytes aedon Vieillot 1809), a species with melanin-based plumage pigments. In one experiment, nestlings within females' second broods were randomly assigned to receive either a control or a lutein supplement (2008); in a second experiment, females, before their first broods, were either induced to lay additional eggs or not induced, and nestlings within both kinds of broods were supplemented as in the first experiment (2009). There were no significant effects of lutein supplementation on nestling condition or phytohemagglutinin response. There was a significant effect of lutein supplementation on nestling mass in 2008, but the difference was opposite to that predicted. Moreover, even when breeding females were stressed by inducing them to lay supernumerary eggs, lutein supplementation of nestlings had no effect on the size or condition of nestlings hatching from these eggs. These results suggest that maternally derived lutein in the egg and that provided in the diet of nestlings are not limiting to normal development and to the components of the immune system involved in the phytohemagglutinin response of nestling house wrens.     

Effects of carotenoids, immune activation and immune suppression on the intensity of chronic coccidiosis in greenfinches

Allocation trade-offs of carotenoids between their use in the immune system and production of integumentary colouration have been suggested as a proximate mechanism maintaining honesty of signal traits. We tested how dietary carotenoid supplementation, immune activation and immune suppression affect intensity of coccidian infection in captive greenfinches Carduelis chloris, a passerine with carotenoid-based plumage. Immune activation with phytohaemagglutinin (PHA) decreased body mass among birds not supplemented with lutein, while among the carotenoid-fed birds, PHA had no effect on mass dynamics. Immune suppression with dexamethasone (DEX) induced loss of body mass and reduced the swelling response to PHA. DEX and PHA increased the concentration of circulating heterophils. Lutein supplementation increased plasma carotenoid levels but had no effect on the swelling response induced by PHA. PHA and DEX treatments did not affect plasma carotenoids. Immune stimulation by PHA suppressed the infection, but only among carotenoid-supplemented birds. Priming of the immune system can thus aid in suppressing chronic infection but only when sufficient amount of carotenoids is available. Our experiment shows the importance of carotenoids in immune response, but also the complicated nature of this impact, which could be the reason for inconsistent results in studies investigating the immunomodulatory effects of carotenoids. The findings about involvement of carotenoids in modulation of an immune response against coccidiosis suggest that carotenoid-based ornaments may honestly signal individuals’ ability to manage chronic infections.     

The effects of diet quality and quantity on plumage colour and growth of great tit Parus major nestlings: a food manipulation experiment along a pollution gradient

The yellow carotenoid-based plumage coloration of great tit Parus major nestlings is found to be paler in polluted and urban environments. Because carotenoid pigmentation is often considered to be a condition dependent trait in birds we wanted to find out whether food-limitation and poor nestling condition could explain the pale plumage colour in a polluted area. P. major nestlings were supplemented with variable diets along a well known heavy metal pollution gradient around a copper smelter: two food treatments with carotenoids, one food treatment with little carotenoid and one unsupplemented control. Our field experiment showed that nestlings in the polluted area grew better with carotenoid rich diets, while such effect was not found in the unpolluted area. Nestlings showed higher plasma carotenoid (lutein) levels and higher plumage carotenoid chroma values in the unpolluted area than in the polluted area. However, plasma lutein levels or plumage colour were not associated with heavy metal levels in nestling faeces (a proxy for dietary exposure). Our results provide only weak evidence for carotenoid-based colouration to be condition-dependent in great tit nestlings as we found a positive relationship between body mass and carotenoid chroma in the non-supplemented control group only. The positive relationship between body mass and plumage colour intensity is more likely to be produced by the fact that good availability of caterpillars, an important food source for P. major, also means a good availability of carotenoids to nestlings. Our results suggest that main reason for pale nestling plumage in the polluted area is lower quality invertebrate food, and not nutrition-related oxidative stress.     

Dietary carotenoid supplementation facilitates egg laying in a wild passerine

During egg laying, females face a trade‐off between self‐maintenance and investment into current reproduction, since providing eggs with resources is energetically demanding, in particular if females lay one egg per day. However, the costs of egg laying not only relate to energetic requirements, but also depend on the availability of specific resources that are vital for egg production and embryonic development. One of these compounds are carotenoids, pigments with immuno‐stimulatory properties, which are crucial during embryonic development. In this study, we explore how carotenoid availability alleviates this trade‐off and facilitates egg laying in a small bird species, the blue tit (Cyanistes caeruleus). Blue tits have among the largest clutch size of all European passerines and they usually lay one egg per day, although laying interruptions are frequent. We performed a lutein supplementation experiment and measured potential consequences for egg laying capacity and egg quality. We found that lutein‐supplemented females had less laying interruptions and thus completed their clutch faster than control females. No effects of treatment were found on the onset of egg laying or clutch size. Experimentally enhanced carotenoid availability did not elevate yolk carotenoid levels or egg mass, but negatively affected eggshell thickness. Our results provide hence evidence on the limiting role of carotenoids during egg laying. However, the benefits of laying faster following lutein supplementation were counterbalanced by a lower accumulation of calcium in the eggshell. Thus, even though single components may constrain egg laying, it is the combined availability of a range of different resources which ultimately determines egg quality and thus embryonic development.     

The effect of an acute phase response on tissue carotenoid levels of growing chickens (Gallus gallus domesticus)

Plasma, liver and skin carotenoids decrease following infectious disease challenges. Since these challenges often involve substantial host pathology and chronic immune responses, the mechanism underlying altered carotenoid deposition is unclear. Therefore, changes in tissue carotenoid levels were examined during an acute phase response induced by lipopolysaccharide (LPS) or interleukin-1 (IL-1). In two experiments, chicks were hatched from carotenoid-deplete eggs (n=28, n=64, respectively) and fed 0, 8 or 38 mg carotenoids (lutein+canthaxanthin)/kg diet. For chicks fed 38 mg carotenoids, but not those fed 0 or 8 mg, LPS generally reduced plasma lutein, canthaxanthin and total carotenoids (P<0.05), and liver lutein, zeaxanthin, canthaxanthin and total carotenoids (P<0.05). Additionally, LPS reduced thymic total carotenoids (P=0.05) and increased thymocyte lutein (P=0.07), zeaxanthin (P=0.07) and total carotenoids (P=0.07). Finally, LPS increased bursal canthaxanthin (P<0.01), but had no effect on shank carotenoids (P>0.5). In chicks hatched from carotenoid-replete eggs (n=36) and fed dietary lutein (38 mg/kg diet), LPS reduced plasma and liver zeaxanthin and liver total carotenoids (P<0.05); IL-1 reduced plasma and liver lutein, zeaxanthin and total carotenoids (P<0.05). Therefore, an acute phase response plays a role in reduced tissue carotenoids during infectious disease.     

Dietary carotenoid pigments and immune function in a songbird with extensive carotenoid-based plumage coloration

Carotenoid pigments can directly enhance the immune responsesof vertebrates, and they are used by many animals to createornamental color displays. It has been hypothesized that thesetwo functions of carotenoid pigments are linked: animals musttrade off use of carotenoid pigments for immune function versusornamental display. We tested two key predictions of this hypothesiswith captive American goldfinches, Carduelis tristis, a specieswith extensive carotenoid-based plumage coloration. First, wetested whether the immune systems of male goldfinches are carotenoidlimited during molt by supplying treatment groups with low,approximately normal, or high dietary access to lutein and zeaxanthin.Dietary treatment had a significant effect on plumage and billcolor but not on immunocompetence. We compared the cell-mediatedand humoral immune responses and the course of disease afterinfection for males in the different treatments. We observedno significant effect of the carotenoid content of diet on immuneresponse or disease resistance. Second, we tested whether therewas a positive relationship between immune function and expressionof ornamental coloration by comparing both the pre- and posttreatmentplumage coloration of males to their immune responses. We failedto find the predicted trade-off between ornament display andimmune function. These findings do not support the hypothesisthat songbirds with extensive carotenoid-based plumage displaystrade off the use of carotenoids for ornamentation versus immunefunction.     

Plasma carotenoid concentrations of incubating American kestrels (Falco sparverius) show annual,seasonal, and individual variation and explain reproductive outcome

In wild birds, the proximate and ultimate factors that affect circulating carotenoid concentrations remain poorly understood. We studied variation in plasma carotenoid concentrations across several scales: annual, seasonal, pair, territory and individual, and evaluated whether plasma carotenoid concentrations explained reproductive outcome of wild American kestrels (Falco sparverius). We sampled plasma carotenoid concentrations of 99 female and 80 male incubating kestrels from April to June in 2008 to 2012. Plasma carotenoid concentrations were explained by an interaction between year and sex, date, and random effects for pair and individual identity. In general, plasma carotenoid concentrations of males were significantly higher than females, but this depended on year. Within a breeding season, earlier nesting kestrels had higher carotenoid concentrations than later nesting kestrels, a pattern that is coincident with seasonal trends in local fitness. Pair and individual identity explained variation in carotenoid concentrations suggesting that carotenoid concentrations of mated birds were correlated, and some individuals consistently maintained higher carotenoid levels than others. Male carotenoid concentrations were positively associated with number of young fledged per pair. These results are consistent with the hypothesis that higher quality individuals have higher carotenoid levels compared to lower quality individuals, despite annual variations in carotenoid availability.     

Geographical trends in the yolk carotenoid composition of the pied flycatcher (Ficedula hypoleuca)

Carotenoids in the egg yolks of birds are considered to be important antioxidants and immune stimulants during the rapid growth of embryos. Yolk carotenoid composition is strongly affected by the carotenoid composition of the female??s diet at the time of egg formation. Spatial and temporal differences in carotenoid availability may thus be reflected in yolk concentrations. To assess whether yolk carotenoid concentrations or carotenoid profiles show any large-scale geographical trends or differences among habitats, we collected yolk samples from 16 European populations of the pied flycatcher, Ficedula hypoleuca. We found that the concentrations and proportions of lutein and some other xanthophylls in the egg yolks decreased from Central Europe northwards. The most southern population (which is also the one found at the highest altitude) also showed relatively low carotenoid levels. Concentrations of ??-carotene and zeaxanthin did not show any obvious geographical gradients. Egg yolks also contained proportionally more lutein and other xanthophylls in deciduous than in mixed or coniferous habitats. We suggest that latitudinal gradients in lutein and xanthophylls reflect the lower availability of lutein-rich food items in the northern F. hypoleuca populations and in montane southern populations, which start egg-laying earlier relative to tree phenology than the Central European populations. Similarly, among-habitat variation is likely to reflect the better availability of lutein-rich food in deciduous forests. Our study is the first to indicate that the concentration and profile of yolk carotenoids may show large-scale spatial variation among populations in different parts of the species?? geographical range. Further studies are needed to test the fitness effects of this geographical variation.     

Carotenoid coloration in great black‐backed gull Larus marinus reflects individual quality

Carotenoids are a large group of biochemicals, with similar properties, synthesised by bacteria, fungi, algae and plants. Vertebrates obtain these biologically active pigments through the diet, and they are a disproportionately common component of animal colour signals and play important roles in immune functions and as antioxidants. Carotenoids are believed to be a limited resource and because of the trade‐off between allocation of carotenoids to signals and to other functions, carotenoid based signals are often thought to be handicap signals. The purpose of this study was to investigate the signalling potential of carotenoid‐based tissue coloration in the great black‐backed gull Larus marinus. The intensity of carotenoid‐based coloration in bill, gape and eye‐ring coloration was investigated in relation to body condition, reproductive parameters, levels of immune activity, and sexual dimorphism. In males there was a positive relationship between colour intensity and body condition, but in females no such relationship was found. However, females with high colour intensity had larger eggs and clutches. Additionally, females with high red scores tended to have high density of circulating lymphocytes. There was no sexual dimorphism in coloration and there was a negative relationship between colour intensity and sampling time, which indicates that this coloration is most intensely expressed early in the breeding season. The results in this study suggest that carotenoid‐based coloration in great black‐backed gull are partly condition dependent and reveal information about individual quality in both males an females. Hence, it might have evolved as an important signal for assessing the quality of potential mates.     

Patterns of serum carotenoid accumulation and skin colour variation in kestrel nestlings in relation to breeding conditions and different terms of carotenoid supplementation

Carotenoids are pigments synthesised by autotrophic organisms. For nestlings of raptorial species, which obtain carotenoids from the consumption of other heterotrophic species, the access to these pigments can be crucial. Carotenoids, indeed, have fundamental health maintenance functions, especially important in developing individuals as nestling kestrels. The aim of this study was to investigate how body carotenoid levels and skin pigmentation vary in kestrel nestlings (Falco tinnunculus) in relation to nesting parameters. Furthermore, we experimentally altered carotenoid availability (short- medium- and long-term) for nestlings and investigated skin and serum variance. The skin colour variance of 151 nestlings was explained by nest of origin, age and by the body condition (body mass corrected by age), older nestlings with higher body condition being redder. No difference in skin colour was detected between sexes. Differences in hue (skin “redness”) between treatments did not emerge during the first week, but did occur 15 days after administration between long-term supplemented and control chicks. In contrast, the serum carotenoid concentration showed a treatment-dependent increase after 5 days from the first carotenoid administration and at least after two supplemented feedings. In general, hue but not serum carotenoids, was correlated with the body condition of nestlings. Based on the increased skin pigmentation of nestling kestrels in the long-term experimental group, we suggest carotenoid availability to be limited for colour expression. The small increase of serum carotenoids due to supplementation is consistent with the hypothesis that there is a physiological constraint on these pigments, as well as an environmental limitation. The presented results are useful for the understanding of carotenoid uptake and accumulation by a wild raptorial species, located at the top of the food web, highlighting that carotenoids are a limited resource for kestrel nestlings.     

Carotenoid supplementation and GnRH challenges influence female endocrine physiology, immune function, and egg-yolk characteristics in Japanese quail (Coturnix japonica)

Androgens and carotenoids circulating in plasma affect the physiology and behavior of vertebrates. Much is known about control mechanisms and functions of each of these substances, yet their interactive effects are not well understood. Here we examine possible additive, multiplicative, and interactive effects of testosterone and carotenoids on female endocrine physiology, immunocompetence, and investment in eggs by simultaneously manipulating levels of testosterone [via gonadotropin releasing hormone (GnRH) challenges] and carotenoids (via diet supplementation) in captive female Japanese quail (Coturnix japonica). Females were randomly assigned to one of four treatments: carotenoid supplementation, GnRH challenge, GnRH challenge?+?carotenoid supplementation, or control. Carotenoid supplementation significantly increased circulating plasma carotenoid levels and acquired immune system performance, but not innate immunity. GnRH challenges elevated circulating testosterone and carotenoid levels, and induced immunosuppression in females. However, females in the GnRH challenge?+?carotenoid supplementation treatment had higher cell-mediated immune responses than control females and similar responses to those of carotenoid-supplemented females. Hence, availability of carotenoids in female quail seemed to counteract immunosuppressive effects of GnRH challenges. Our results provide further evidence for synergistic effects of carotenoids and testosterone on endocrine physiology and immune function in female birds. Elevated plasma testosterone or carotenoids levels resulted in increased deposition of those compounds to eggs, respectively. Furthermore, because we found that concentrations of testosterone and carotenoids in yolks were correlated within each treatment group, differential deposition of hormones and carotenoids in eggs may not only respond to surrounding social and environmental conditions, but also to other components of the egg.     

Anhydrolutein in the zebra finch: a new,metabolically derived carotenoid in birds

Many birds acquire carotenoid pigments from the diet that they deposit into feathers and bare parts to develop extravagant sexual coloration. Although biologists have shown interest in both the mechanisms and function of these colorful displays, the carotenoids ingested and processed by these birds are poorly described. Here we document the carotenoid-pigment profile in the diet, blood and tissue of captive male and female zebra finches (Taeniopygia guttata). Dietary carotenoids including: lutein; zeaxanthin; and β-cryptoxanthin were also present in the plasma, liver, adipose tissue and egg-yolk. These were accompanied in the blood and tissues by a fourth pigment, 2′,3′-anhydrolutein, that was absent from the diet. To our knowledge, this is the first reported documentation of anhydrolutein in any avian species; among animals, it has been previously described only in human skin and serum and in fish liver. We also identified anhydrolutein in the plasma of two closely related estrildid finch species (Estrilda astrild and Sporaeginthus subflavus). Anhydrolutein was the major carotenoid found in zebra finch serum and liver, but did not exceed the concentration of lutein and zeaxanthin in adipose tissue or egg yolk. Whereas the percent composition of zeaxanthin and β-cryptoxanthin were similar between diet and plasma, lutein was comparatively less abundant in plasma than in the diet. Lutein also was proportionally deficient in plasma from birds that circulated a higher percentage of anhydrolutein. These results suggest that zebra finches metabolically derive anhydrolutein from dietary sources of lutein. The production site and physiological function of anhydrolutein have yet to be determined.     

Carotenoids, immunocompetence, and the information content of sexual colors: an experimental test

Many male birds use carotenoid pigments to acquire brilliant colors that advertise their health and condition to prospective mates. The direct means by which the most colorful males achieve superior health has been debated, however. One hypothesis, based on studies of carotenoids as antioxidants in humans and other animals, is that carotenoids directly boost the immune system of colorful birds. We studied the relationship between carotenoid pigments, immune function, and sexual coloration in zebra finches (Taeniopygia guttata), a species in which males incorporate carotenoid pigments into their beak to attract mates. We tested the hypotheses that increased dietary carotenoid intake enhances immunocompetence in male zebra finches and that levels of carotenoids circulating in blood, which also determine beak coloration, directly predict the immune response of individuals. We experimentally supplemented captive finches with two common dietary carotenoid pigments (lutein and zeaxanthin) and measured cell-mediated and humoral immunity a month later. Supplemented males showed elevated blood-carotenoid levels, brighter beak coloration, and increased cell-mediated and humoral immune responses than did controls. Cell-mediated responses were predicted directly by changes in beak color and plasma carotenoid concentration of individual birds. These experimental findings suggest that carotenoid-based color signals in birds may directly signal male health via the immunostimulatory action of ingested and circulated carotenoid pigments.     

Die rolle der Carotinoide und des Gallenfarbstoffs bei der Farbmodifikation der Puppen von Pieris brassicae

The carotenoids and the bile pigment in larvae and pupae of Pieris brassicae were analysed. Their rôle in the morphological colour adaptation of the pupae was studied by quantitative measurements.The carotenoids are β-carotene, lutein mono-ester, free lutein, and zeaxanthin. Metabolized carotenoids were not found. There are no differences between pupae showing different grades of melanization in the quality of the carotenoids, or in the total amounts, or in the relative portions of each carotenoid fraction. However, the carotenoid content of the integument alone is twofold in the light pupae as compared to dark ones. The integumental carotenoids are deposited mainly in the epidermis. β-Carotene, lutein, and zeaxanthin are selectively absorbed by the larvae from the diet. β-Carotene and lutein ester are localized mainly in the fat body, whereas lutein is predominant in the haemolymph and in the integument.The pupal bile pigment is protobiliverdin-IXγ (pterobilin), which is also known to be the larval pigment. The bile pigment is synthesized mainly during the last larval instar up to the pharate pupal stage. In the pupae the bile pigment content is related to the melanization: pupae exposed to the same light conditions contain less bile pigment the more melanized they are (negative correlation). On the whole there is a strong enhancement by blue light of the bile pigment content besides the known stimulation of melanization (positive correlation). But within such a sample the negative correlation between the amounts of bile pigment and melanin is maintained.     

Condition dependence of nestling mouth colour and the effect of supplementing carotenoids on parental behaviour in the hihi (Notiomystis cincta)

Carotenoids are integument pigments that often reflect foraging efficiency, disease resistance and body condition. In contrast to the widespread attention this relationship has received in adult birds, the condition dependence of nestling colouration remains an understudied component of animal communication. Here we assess the condition dependence of carotenoid pigmentation in nestling hihi (Notiomystis cincta, an endangered New Zealand bird) and examine the influence of carotenoid supplementation on nestling quality and parental visitation rates. Our results show that carotenoids provided to breeding adult hihi were transferred to their offspring and resulted in an intensified orange-yellow flange colour. After accounting for carotenoid supplementation the parameter that most consistently explained variation in nestling flange colour was nestling tarsus length at 23 days, indicating condition dependence of this trait. We did not, however, detect direct effects of carotenoid supplementation on nestling mass or immune response (or any other fitness parameter measured). Carotenoid supplementation did, however, result in an increased paternal provisioning rate.     

Selective absorption of carotenoids in the common green iguana (Iguana iguana)

In the animal kingdom, species-specific differences with regard to the absorption of intact carotenoids are observed. The causes of these differences are not entirely understood. To investigate the absorption of selected carotenoids, 20 juvenile green iguanas (Iguana iguana) were fed a carotenoid deficient basal diet for 56 days. Thereafter, the iguanas were assigned to receive a basal diet supplemented with different carotenoids (80 mg/kg diet) such as beta-carotene, canthaxanthin and apo-8'-carotenoic acid ethyl ester for 28 days. Changes in plasma carotinoid concentrations associated with the individual diets were used as indicators of carotenoid absorption. In both the experimental and control groups, only the oxygenated carotenoids (xanthophylls), lutein, zeaxanthin and canthaxanthin, were found in the plasma. Canthaxanthin and apo-8'-carotenoic acid ethyl ester were readily absorbed and recovered from the plasma. However, the supplementation of beta-carotene caused no increase in plasma beta-carotene concentration. Additionally, beta-carotene, canthaxanthin or apo-8'-carotenoic acid ethyl ester did not affect the concentrations of retinol and alpha-tocopherol in plasma. In conclusion, the study demonstrates that iguanas appear to be selective accumulators of polar xanthophylls. The iguana might, therefore, be a valuable model to investigate the selectiveness of carotenoid absorption as well as the function of xanthophylls in animals.     

Phenotypic variation in nestlings of a bird of prey under contrasting breeding and diet conditions

Environmental conditions often vary in space and time, and this may explain variation in the expression of phenotypic traits related to individual quality, such as ornamental coloration. Furthermore, the direction and strength of the relationship between coloured trait expression and individual quality might vary under contrasting conditions. These issues have been explored in adult birds but much less so in nestlings, which are more likely to experience different selective pressures and different physiological trade‐offs than adults. Here, we empirically investigated the effects of contrasting breeding and diet conditions on the expression of carotenoid‐based colour traits displayed by marsh harrier (Circus aeruginosus) nestlings. We studied the variation in coloration, body condition, and immune responsiveness of nestlings in four populations over a 5‐year period. We characterized spatiotemporal differences in rearing conditions experienced by C. aeruginosus nestlings in terms of breeding (laying date, clutch size, and number of nestlings hatched and fledged) and diet (percentage of mammal in diet and prey diversity) conditions. We found that breeding conditions influenced the co‐variation between coloration and immune responsiveness in female nestlings, and that diet conditions influenced the condition‐dependence of nestling coloration in later‐hatched nestlings. In addition, breeding conditions influenced nestling body condition and immune responsiveness, whereas diet conditions influenced nestling coloration and body condition. Our study highlights that nestling phenotype (levels of signalling, circulating carotenoids, and immunity) varies both spatially and temporally, and that some of this variation is related to differences in breeding and diet conditions. Moreover, under contrasting conditions, the direction of the relationships between nestling carotenoid‐based coloration and nestling quality may also vary. In order to fully understand the evolution and maintenance of colour traits in nestling birds, studies and experiments should ideally be replicated under contrasting rearing conditions. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.