Egg yolk antioxidant deposition as a function of parental ornamentation,age, and environment in great tits Parus major

Females can modify phenotype of their offspring through the deposition of biologically active compounds into eggs, including carotenoids, vitamins and other antioxidants. Understanding patterns of deposition is critical for better insight into the significance of maternal effects. Here we investigated how egg yolk antioxidants (lutein, zeaxanthin, β‐carotene, vitamin A and E) related to environmental conditions and parental characteristics in great tits Parus major using data from three breeding seasons. Male and female traits included condition, age and multiple feather ornaments, both carotenoid‐ and melanin‐based (carotenoid and UV chroma of yellow breast feathers, area of black breast band, white cheek immaculateness). Yolk mass increased with ambient temperature during laying, laying date, and the area of male black breast band. Lutein, zeaxanthin, and vitamin E increased with laying date. Total antioxidants increased with female age, immaculateness of female white cheek patch, and UV chroma of carotenoid‐based yellow breast feathers of the social mate. These patterns were thus consistent with 1) environmental effects on yolk mass and composition, 2) higher quality females depositing more antioxidants, and 3) differential allocation of resources in females in relation to male ornamentation. Overall, environmental factors, female traits, and male traits all had an influence on egg yolk characteristics in this socially monogamous songbird.     

Maternal dietary carotenoids mitigate detrimental effects of maternal GnRH on offspring immune function in Japanese quail Coturnix japonica

Maternal resources deposited in eggs can affect the development of several offspring phenotypic traits and result in trade‐offs among them. For example, maternal androgens in eggs may be beneficial to offspring growth and competitive ability, but detrimental to immunocompetence and oxidative stress. In contrast, maternal antioxidants in eggs may be beneficial if they mitigate oxidative stress and immunosuppressive effects of androgens. We investigated possible interactive effects of maternal steroids and carotenoids on aspects of offspring physiology and phenotype, by simultaneously manipulating levels of androgens (via gonadotropin‐releasing hormone, GnRH‐challenges) and carotenoids (via diet supplementation) in captive female Japanese quail Coturnix japonica during egg laying. Carotenoid supplementation of hens, which elevates yolk concentrations of carotenoid and vitamins A and E, enhanced egg hatching success, offspring survival to age 15 d, and size of the bursa of Fabricius in offspring. In contrast, repeated maternal GnRH challenges, which elevated yolk testosterone concentrations, enhanced offspring neonatal size, but negatively affected bursa size. However, interaction among the treatments suggests that the positive effect of maternal carotenoid supplementation on plasma bactericidal capacity was mediated by maternal GnRH challenges. Chicks originating from carotenoid‐supplemented hens were less immunosuppressed than those originating from carotenoid‐supplemented + GnRH‐challenged hens, which were less immunosuppressed than chicks from GnRH‐challenged females not supplemented with carotenoids. Females availability of carotenoid enriched diets allows them to enhance the development of offspring immune system via carotenoids and vitamins deposited in egg yolks and offset detrimental effects of androgens deposited by GnRH‐challenged females.     

Geographical variation in egg mass and egg content in a passerine bird

Reproductive, phenotypic and life-history traits in many animal and plant taxa show geographic variation, indicating spatial variation in selection regimes. Maternal deposition to avian eggs, such as hormones, antibodies and antioxidants, critically affect development of the offspring, with long-lasting effects on the phenotype and fitness. Little is however known about large-scale geographical patterns of variation in maternal deposition to eggs. We studied geographical variation in egg components of a passerine bird, the pied flycatcher (Ficedula hypoleuca), by collecting samples from 16 populations and measuring egg and yolk mass, albumen lysozyme activity, yolk immunoglobulins, yolk androgens and yolk total carotenoids. We found significant variation among populations in most egg components, but ca. 90% of the variation was among individuals within populations. Population however explained 40% of the variation in carotenoid levels. In contrast to our hypothesis, we found geographical trends only in carotenoids, but not in any of the other egg components. Our results thus suggest high within-population variation and leave little scope for local adaptation and genetic differentiation in deposition of different egg components. The role of these maternally-derived resources in evolutionary change should be further investigated.     

Maternal effects mediated by egg quality in the Yellow-legged Gull Larus michahellis in relation to laying order and embryo sex

Background

Maternal effects mediated by egg size and quality may profoundly affect offspring development and performance, and mothers may adjust egg traits according to environmental or social influences. In avian species, context-dependency of maternal effects may result in variation in egg composition, as well as in differential patterns of covariation among selected egg components, according to, for example, position in the laying sequence or offspring sex. We investigated variation in major classes of egg yolk components (carotenoids, vitamins and steroid hormones) in relation to egg size, position in the laying sequence and embryo sex in clutches of the Yellow-legged Gull (Larus michahellis). We also investigated their covariation, to highlight mutual adjustments, maternal constraints or trade-offs in egg allocation.

Results

Laying sequence-specific patterns of allocation emerged: concentration of carotenoids and vitamin E decreased, while concentrations of androgens increased. Vitamin A, estradiol and corticosterone did not show any change. There was no evidence of sex-specific allocation or covariation of yolk components. Concentrations of carotenoids and vitamins were positively correlated. Egg mass decreased along the laying sequence, and this decrease was negatively correlated with the mean concentrations of carotenoids in clutches, suggesting that nutritionally constrained females lay low quality clutches in terms of carotenoid content. Finally, clutches with smaller decline in antioxidants between first- and last-laid eggs had a larger increase in yolk corticosterone, suggesting that a smaller antioxidant depletion along the laying sequence may entail a cost for laying females in terms of increased stress levels.

Conclusions

Since some of the analyzed yolk components (e.g. testosterone and lutein) are known to exert sex-specific phenotypic effects on the progeny in this species, the lack of sex-specific egg allocation by mothers may either result from trade-offs between contrasting effects of different egg components on male and female offspring, or indicate that sex-specific traits are controlled primarily by mechanisms of sexual differentiation, including endogenous hormone production or metabolism of exogenous antioxidants, during embryonic development.     

Determinants of within- and among-clutch variation in yolk corticosterone in the European starling

Maternal glucocorticoids are known to affect offspring phenotype in numerous vertebrate taxa. In birds, the maternal transfer of corticosterone to eggs was recently proposed as a hormonal mechanism by which offspring phenotype is matched to the relative quality of the maternal environment. However, current hypotheses lack supporting information on both intra- and inter-clutch variation in yolk corticosterone for wild birds. As such, we examined variation in yolk corticosterone levels in a wild population of European starlings (Sturnus vulgaris). Maternal condition, clutch size and nesting density were all negatively related to yolk corticosterone deposition; females with high condition indices, those laying larger clutches and those nesting in high-density associations deposited lower amounts of the hormone into eggs than those with low condition indices, laying small clutches and nesting in isolation. Alternatively, we found no effects of maternal age or human disturbance on yolk corticosterone deposition. Intra-clutch variation of yolk corticosterone was significant, with levels increasing across the laying sequence in all clutch sizes examined, with the difference between first and last-laid eggs being greater in large versus small clutches. Given the reported effects of yolk corticosterone on offspring size and growth, intra-clutch variation in yolk corticosterone has the potential to alter the competitive environment within a brood. Furthermore, our results indicate that variation in yolk corticosterone can originate from variation in both the mother's quality as well as the quality of her breeding environment. The presence of inter-female variation in particular is an important pre-requisite in testing whether the exposure of offspring to maternally-derived corticosterone is a mechanistic link between offspring phenotypic plasticity and maternal quality.     

Manipulative signals in family conflict? On the function of maternal yolk hormones in birds

Maternal hormones in the yolk of birds' eggs have been a focus of attention in behavioral and evolutionary ecology stimulated by the pioneering work of Hubert Schwabl. Since then, knowledge of both the factors that influence maternal deposition patterns and their consequences for offspring development has accumulated rapidly. To date, the field has been dominated by the idea that mothers use yolk hormones to adaptively adjust offspring development, a view that assigns control over hormone deposition and its effects on the offspring to the mother. This neglects the possibility that the evolutionary interests of the mother and offspring differ. When there is such parent-offspring conflict, the offspring are selected to respond to the hormones in a way that is adaptive for themselves rather than for the mother. Moreover, sexual conflict between the parents over parental investment may shape the evolution of yolk hormone deposition: females may manipulate the male's contribution to parental care through the effect of yolk hormones on offspring begging, competitiveness, and developmental rate. We therefore suggest that for a full understanding of the evolution of hormone-mediated maternal effects, it is essential to study both fitness consequences and physiological mechanisms and constraints from the perspective of all family members.     

Maternal influences on offspring phenotypes and sex ratios in a multi‐clutching lizard with environmental sex determination

Maternal and environmental factors are important sources of phenotypic variation because both factors influence offspring traits in ways that impact offspring and maternal fitness. The present study explored the effects of maternal factors (maternal body size, egg size, yolk‐steroid allocation, and oviposition‐site choice) and seasonally‐variable environmental factors on offspring phenotypes and sex ratios in a multi‐clutching lizard with environmental sex determination (Amphibolurus muricatus). Maternal identity had strong effects on offspring morphology, but the nature of maternal effects differed among successive clutches produced by females throughout the reproductive season (i.e. maternal identity by environment interactions). The among‐female and among‐clutch variation in offspring traits (including sex ratios) was not mediated through maternal body size, egg size, or variation in yolk steroid hormones. This lack of nongenetic maternal effects suggests that phenotypic variation may be generated by gene by environment interactions. These results demonstrate a significant genetic component to variation in offspring phenotypes, including sex ratios, even in species with environmental sex determination. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 256–266.     

Yolk antioxidants vary with male attractiveness and female condition in the house finch (Carpodacus mexicanus)

The manipulation of egg content is one of the few ways by which female birds can alter offspring quality before hatch. Lipid-soluble vitamins and carotenoids are potent antioxidants. Female birds deposit these antioxidants into eggs in variable amounts according to environmental and social conditions, and the quantities deposited into eggs can have effects on offspring health and immunological condition. Allocation theory posits that females will alter the distribution of resources according to mate quality, sometimes allocating resources according to the differential allocation hypothesis (DAH), investing more in offspring sired by better-quality males, and other times allocating resources according to a compensatory strategy, enhancing the quality of offspring sired by lower-quality males. It is unknown, however, whether antioxidants are deposited into eggs according to the DAH or a compensatory strategy. We examined deposition patterns of yolk antioxidants (including vitamin E and three carotenoids) in relation to laying order, mate attractiveness, female condition, and yolk androgen content in the house finch (Carpodacus mexicanus). Female house finches deposited significantly more total antioxidants into eggs sired by less attractive males. Additionally, yolk antioxidant content was significantly positively correlated with female condition, which suggests a cost associated with the deposition of antioxidants into eggs. Finally, concentrations of antioxidants in egg yolks were positively correlated with total yolk androgen content. We suggest that yolk antioxidants are deposited according to a compensatory deposition strategy, enabling females to improve the quality of young produced with less attractive males. Additionally, yolk antioxidants may act to counter some of the detrimental effects associated with high levels of yolk androgens in eggs and, thus, may exert a complementary effect to yolk androgens.     

Maternal allocation strategies and differential effects of yolk carotenoids on the phenotype and viability of yellow-legged gull (Larus michahellis) chicks in relation to sex and laying order

Egg quality may mediate maternal allocation strategies according to progeny sex. In vertebrates, carotenoids have important physiological roles during embryonic and post-natal life, but the consequences of variation in yolk carotenoids for offspring phenotype in oviparous species are largely unknown. In yellow-legged gulls, yolk carotenoids did not vary with embryo sex in combination with egg laying date, order and mass. Yolk lutein supplementation enhanced the growth of sons from first eggs but depressed that of sons from last eggs, enhanced survival of daughters late in the season, and promoted immunity of male chicks and chicks from small eggs. Lack of variation in egg carotenoids in relation to sex and egg features, and the contrasting effects of lutein on sons and daughters, do not support the hypothesis of optimal sex-related egg carotenoid allocation. Carotenoids transferred to the eggs may rather result from a trade-off between opposing effects on sons or daughters.     

Carotenoids and egg quality in the lesser blackbacked gull Larus fuscus: a supplemental feeding study of maternal effects.

Egg quality is a phenotype of, and can profoundly influence fitness in, both mother and offspring. However, the physiological mechanisms that underlie this maternal effect are poorly understood. Carotenoids are hypothesized to enhance antioxidant activity and immune function, and are responsible for the pigmentation of egg yolk. The proximate basis and consequences of this maternal investment, however, have not previously been studied in wild birds. In this supplemental feeding study of lesser black-backed gulls, Larus fuscus, carotenoid-fed females are shown to have increased integument pigmentation, higher plasma concentrations of carotenoids and antioxidant activity, and lower plasma concentrations of immunoglobulins (Igs) in comparison with controls. In turn, carotenoid-fed females produced eggs containing high carotenoid but low Ig concentrations (i.e. passive immunity), whereas control females produced eggs containing low carotenoid but high Ig concentrations. Within-clutch patterns of these resources varied over the laying sequence in a similar manner in both carotenoid-fed and control nests. Our results suggest that carotenoids could be one resource responsible for egg quality maternal effects in birds. We discuss the possible implications of carotenoid-mediated effects on phenotype for fitness in mothers and their offspring.     

Sex-related effects of maternal egg investment on offspring in relation to carotenoid availability in the great tit

1. Maternal carotenoids in the egg yolk have been hypothesized to promote maturation of the immune system and protect against free radical damages. Depending on availability, mothers may thus influence offspring quality by depositing variable amounts of carotenoids into the eggs. Sex allocation theory predicts that in good quality environments, females should invest into offspring of the sex that will provide larger fitness return, generally males. 2. In a field experiment we tested whether female great tits bias their investment towards males when carotenoid availability is increased, and whether male offspring of carotenoid-supplemented mothers show higher body condition. We partially cross-fostered hatchlings to disentangle maternal effects from post-hatching effects, and manipulated hen flea Ceratophyllus gallinae infestation to investigate the relationship between carotenoid availability and resistance to ectoparasites. 3. As predicted, we found that carotenoid-supplemented mothers produced males that were heavier than their sisters at hatching, while the reverse was true for control mothers. This suggests that carotenoid availability during egg production affects male and female hatchlings differentially, possibly via a differential allocation to male and female eggs. 4. A main effect of maternal supplementation became visible 14 days after hatching when nestlings hatched from eggs laid by carotenoid-supplemented mothers had gained significantly more mass than control nestlings. Independently of the carotenoid treatment, fleas impaired mass gain of nestlings during the first 9 days in large broods only and reduced tarsus length of male nestlings at an age of 14 days, suggesting a cost to mount a defence against parasites. 5. Overall, our results suggest that pre-laying availability of carotenoids affects nestling condition in a sex-specific way with potentially longer-lasting effects on offspring fitness.     

Parental investment and egg yolk lipid composition in gulls

1. Within-clutch variation in parental investment of yolk lipid (including essential fatty acid composition and fat-soluble vitamin content) was investigated in whole clutches of three eggs of Lesser Black-backed Gulls, Larus fuscus .
2. Although there were no significant differences between years in egg size or yolk mass among eggs of the same laying order, eggs collected in 1996 had significantly lower lipid content than those collected in 1997.
3. In both years third-laid c-eggs were smaller, had a smaller mass of yolk and had significantly lower yolk lipid mass compared with earlier laid (a- and b-) eggs.
4. Despite this lower lipid mass of c-eggs, of the different lipid fractions only cholesterol content of c-eggs was significantly lower than that of a- or b-eggs.
5. Earlier laid eggs had greater amounts of arachidonic acid in the phospholipid and cholesterol ester fractions of yolk lipid, and considerably greater quantities of vitamin E and carotenoids.
6. Conversely, levels of linoleic acid and vitamin A were equalized among eggs of different laying sequence as both these essential nutrients have primarily systemic roles.
7. The probable effect of increased antioxidant (vitamin E and carotenoid) content of earlier-laid eggs is to enhance embryonic and neonatal growth and development, reduce susceptibility to pathogens and increase the ability of the embryo to withstand the oxidative stress associated with hatching.
8. The results suggest that there is maternal bias of essential nutrients to offspring with the highest reproductive value in order to maximize reproductive success.     

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.     

Yolk carotenoids and stable isotopes reveal links among environment, foraging behavior and seabird breeding success

Nutrients that are limited in availability, such as carotenoids, are potentially involved in trade-offs between homeostasis and reproduction. Despite their importance, factors that affect the capacity of female birds to meet their carotenoid requirements are poorly understood. We used δ¹⁵N stable isotope analysis to relate foraging behavior to yolk carotenoid deposition in two seabirds, Cassin’s auklet (Ptychoramphus aleuticus) and rhinoceros auklet (Cerorhinca monocerata), during each of five years. As expected from their narrower trophic range, Cassin’s auklets produced yolks with fewer carotenoid types than did rhinoceros auklets (one vs. three). Cassin’s auklets also fed on a lower trophic level diet richer in carotenoids, yet had lower total yolk carotenoid levels, which suggests a role for species-specific adaptations for carotenoid uptake and utilization. Within both species, lower trophic-level feeding was linked to higher yolk carotenoid levels, but through different mechanisms. In Cassin’s auklets, it was due to a population-wide response to environmental variation: in warm-water years, all females fed at a low trophic level and produced carotenoid-rich yolks. In rhinoceros auklets, it was due to individual differences similarly expressed in all years: females fed across a wide trophic range, and those that fed at a low trophic level produced carotenoid-rich yolks. Rhinoceros auklets bred more successfully in years when their yolks were rich in carotenoids, probably due to a correlated response to stronger marine primary production. Our results are novel because they link avian yolk carotenoid deposition to behavioral and environmental variations.     

An analysis of pre- and post-hatching maternal effects mediated by carotenoids in the blue tit

Maternal effects increase phenotypic plasticity in offspring traits and may therefore facilitate adaptation to environmental variability. Carotenoids have been hypothesized to mediate costs of reproduction in females as well as maternal effects. However, assessing potential transgenerational and population consequences of environmental availability of carotenoids requires a better understanding of mechanisms of maternal effects mediated by these antioxidant pigments. Manipulating dietary availability of carotenoids to egg-laying female blue tits and subsequently cross-fostering nestlings between female treatments allowed us to specifically investigate the relative importance of maternal effects through egg carotenoids and through post-hatching care mediated by antioxidants in females. Nestling body size and mass and plasma antioxidants were not significantly affected by pre- or post-hatching maternal effects mediated by antioxidants, although both types of maternal effects in interaction explained the variation in growth, as measured by wing length. Development of the ability to mount a cell-mediated immune response as well as its temporal dynamics was influenced by both pre- and post-hatching maternal effects, with an advantage to nestlings originating from, or reared by, carotenoid-supplemented females. In addition, nestlings reared by carotenoid-fed females had a lower blood sedimentation rate, indicating that they may have been less infected than nestlings from controls. Finally, prehatching maternal effects in interaction with nestling plasma carotenoid levels affected the development of carotenoid-based plumage. Maternal effects mediated by carotenoids may thus act as a proximate factor in development and phenotypic plasticity in traits associated with nestling fitness, such as immune response and ability to metabolize and use antioxidants, and ultimately participate in the evolution of phenotypic traits.     

Egg yolk carotenoids in relation to habitat and reproductive investment in the great tit Parus major

Maternal allocation of antioxidants to egg yolk has been shown to affect early embryonic development and nestling survival. In environments with high levels of anthropogenic pollution, antioxidants (such as carotenoids) are important to protect the body from elevated oxidative stress. Thus, female allocation of antioxidants to yolk may be traded off against self-maintenance. Here we investigate maternal reproductive investment with respect to yolk carotenoid content and composition in relation to subsequent female condition and carotenoid status in urban and rural great tits Parus major. We found no differences between the urban and rural populations in total yolk carotenoids, egg mass, clutch size, hatching success, or female carotenoid status. Interestingly, however, rural eggs contained more zeaxanthin, a more potent antioxidant than lutein, which suggests that rural embryos have better antioxidant protection than urban embryos. Whether rural females actively transfer more zeaxanthin to the yolk or whether it passively reflects differences in dietary access or uptake needs to be further investigated. This highlights the importance of carotenoid identity and composition in future studies of carotenoid physiology, ecology, and signaling.     

Pterin-based ornamental coloration predicts yolk antioxidant levels in female striped plateau lizards (Sceloporus virgatus)

1. Maternal investment in egg quality can have important consequences for offspring fitness. For example, yolk antioxidants can affect embryonic development as well as juvenile and adult phenotype. Thus, females may be selected to advertise their yolk antioxidant deposition to discriminatory males via ornamental signals, perhaps depending on the reproductive costs associated with signal production. 2. Female striped plateau lizards (Sceloporus virgatus) develop pterin-based orange colour patches during the reproductive season that influence male behaviour and that are positively associated with the phenotypic quality of the female and her offspring. Here, we assessed one potential developmental mechanism underlying the relationship between offspring quality and female ornamentation in S. virgatus, by examining the relationship between ornament expression and yolk antioxidant levels. 3. As expected, concentrations of the yolk antioxidants vitamin A, vitamin E and carotenoids (lutein and zeaxanthin) were strongly positively intercorrelated. Eggs from larger clutches had fewer antioxidants than eggs from smaller clutches, suggesting that females may be limited in antioxidant availability or use. Fertilized and unfertilized eggs did not differ in yolk antioxidant levels. 4. The size of a female's ornament was positively related to both the concentration and total amount of yolk antioxidants, and ornament colour was positively related to yolk antioxidant concentration. Thus, in S. virgatus, female ornaments may advertise egg quality. In addition, these data suggest that more ornamented females may produce higher-quality offspring, in part because their eggs contain more antioxidants. As the colour ornament of interest is derived from pterins, not carotenoids, direct resource trade-offs between ornaments and eggs may be eliminated, reducing reproductive costs associated with signalling. 5. This is the first example of a positive relationship between female ornamentation and yolk antioxidants in reptiles and may indicate the general importance of these patterns in oviparous vertebrates.     

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.     

Experimental evidence for genetic heritability of maternal hormone transfer to offspring

In many animal species, embryos are exposed to maternal hormones that affect their development. Maternal hormone transfer varies with environmental conditions of the mother and is often interpreted as being shaped by natural selection to adjust the offspring to prevailing environmental conditions. Such hormone transfer requires genetic variability, which has not yet been experimentally demonstrated. Our study reports direct evidence for additive genetic variance of maternal androgens through a bidirectional selection on yolk testosterone (T) levels in Japanese quail. Lines selected for high egg T (HET) and low egg T (LET) concentration differed in yolk levels of this androgen, resulting in high realized heritability (h2 = 0.42)Correlated responses to selection on other gonadal hormones indicated that selection specifically targeted biologically active androgens. Eggs of HET quail contained higher androstenedione and lower estradiol concentrations than did those of LET quail, with no line differences in yolk progesterone concentration. Plasma T concentrations in adult females were not affected by selection, seriously challenging the hypothesis that transfer of maternal hormones to offspring is constrained by hormone levels in a mother's circulation. Our results suggest that transfer of maternal T represents an indirect genetic effect that has important consequences for the evolution of traits in offspring.     

Parasite-mediated growth patterns and nutritional constraints in a cavity-nesting bird

1. Trade-offs between growth and immunity of nestling birds can be influenced by parasites, but the magnitude of these effects may depend on availability of critical dietary nutrients. Owing to their importance for both immune system function and growth, dietary carotenoids have the potential to mediate parasite-induced developmental strategies of avian hosts. 2. The effects of ectoparasitic blow flies Protocalliphora spp. and dietary carotenoids (lutein and zeaxanthin) on immune function and patterns of growth in nestling mountain bluebirds Sialia currucoides were investigated by combining parasite removal and carotenoid supplementation treatments in a 2 x 2 design. 3. Supplemental carotenoids enhanced nestlings' T-cell-mediated immune response following intradermal injection of phytohaemagglutinin. 4. The effect of carotenoid supplementation on rate of mass gain depended on whether broods were exposed to parasites: among parasitized broods, those receiving supplemental carotenoids gained mass more rapidly than nonsupplemented broods, whereas there was no effect of supplemental carotenoids on growth of mass in broods that had parasites removed. This suggests that additional dietary carotenoids allowed nestlings to compensate for the otherwise detrimental effects of parasites on mass gain. For length of the eighth primary feather at fledging, early and late broods differed in their response to parasitism: early broods showed an increase in feather length when parasites were removed, while nestlings in late broods had shorter feathers in the absence of parasites. We suggest that this may reflect within-season variation in parasite-mediated growth strategies of nestlings. 5. Maternal condition was positively associated with mass, condition and rate of feather growth of offspring under all conditions, and also influenced nestling immunocompetence, but only in the absence of parasites. 6. We conclude that dietary carotenoids alleviate some of the detrimental effects of parasites on nestling birds; however, parasites also appear to specifically influence other growth and resource allocation strategies, and possibly constrain maternal or genetic effects on offspring phenotype, irrespective of dietary carotenoid availability.