Yolk Testosterone Concentrations Research Articles

Maternal age and maternal environment affect egg composition, yolk testosterone, offspring growth and behaviour in laying hens

Maternal effects have been reported to alter offspring phenotype in laying hens. In this study, we investigated the effects of maternal environment and maternal age on egg traits and offspring development and behaviour. For this, we ran two experiments. First (E1), commercial hybrid hens were reared either in aviary or barren brooding cages, then housed in aviary, conventional cages or furnished (enriched) cages, thus forming different maternal housing treatments. Hens from each treatment were inseminated at three ages, and measures of egg composition, yolk testosterone concentration and offspring’s development, anxiety and fearfulness were assessed. In experiment 2 (E2), maternal age effects on offspring's growth and behaviour were further investigated using fertile eggs from commercial breeder flocks at three different ages. Results from E1 showed that Old hens laid heavier eggs with less yolk testosterone and produced offspring with fewer indicators of anxiety and fearfulness. Maternal rearing and housing affected egg traits, offspring weight and behaviour, but not in a consistent way. Effects of maternal age were not replicated in E2, possibly due to differences in management or higher tolerance to maternal effects in commercial breeders. Overall, our research confirms that maternal age and maternal environment affects egg composition, with maternal age specifically affecting yolk testosterone concentration, which may mediate physical and behavioural effects in offspring.

In ovo yolk carotenoid and testosterone levels interactively influence female transfer of yolk antioxidants to her eggs.

Mothers can influence prenatal conditions by varying the amount of nutrients, hormones or antioxidants they provide to their developing young. Some of these substances even affect the transfer of these compounds in the next generation, but it is less clear how different maternally transmitted compounds interact with each other to shape reproductive resource allocation in their offspring. Here, we found that female Japanese quails (Coturnix japonica) that were exposed to high carotenoid levels during embryonic development transferred lower concentrations of yolk antioxidants to their own eggs later in life. This effect disappeared when both testosterone and carotenoid concentrations were manipulated simultaneously, showing long-term and interactive effects of these maternally derived egg components on a female's own egg composition. Given that exposure to high levels of testosterone during embryo development stimulates the production of reactive oxygen species (ROS) and impairs antioxidant defenses, we propose that carotenoids act as in ovo antioxidants in an oxidatively stressful environment (i.e. when levels of testosterone are high) but might have prooxidant properties in an environment where they are not used to counteract an increased production of ROS. In line with this hypothesis, we previously showed that prenatal exposure to increased concentrations of yolk carotenoids leads to a rise of oxidative damage at adulthood, but only when yolk testosterone concentrations were not experimentally increased as well. As a consequence, antioxidants in the body may be used to limit oxidative damage in females exposed to high levels of carotenoids during development (but not in females exposed to increased levels of both carotenoids and testosterone), resulting in lower amounts of antioxidants being available for deposition into eggs. Since prenatal antioxidant exposure is known to influence fitness-related traits, the effect detected in this study might have transgenerational consequences.

Protoporphyrin‐based eggshell pigmentation predicts hatching success and offspring sex ratio in the barn swallow

Inter‐ and intraspecific variation in eggshell colouration has long fascinated evolutionary biologists. Among species, such variation may accomplish different functions, the most obvious of which is camouflage and background matching. Within species, it has been proposed that inter‐female variation in eggshell pigmentation patterns can reflect egg, maternal or paternal traits and hence may provide cues to conspecifics about egg, maternal or paternal phenotypic quality. However, the relationship between protoporphyrin‐based eggshell pigmentation and egg or maternal/paternal traits appears to be highly variable among species. We investigated patterns of intraspecific variation in Eurasian barn swallow Hirundo r. rustica protoporphyrin‐based eggshell pigmentation, and analysed its association with egg and clutch characteristics, maternal/paternal phenotypic traits and parental feeding effort. Eggshell pigmentation pattern significantly varied between breeding colonies, was significantly repeatable in first clutches laid by the same females in different years (intraclass correlation coefficient ranging between 0.56 and 0.63), but it was not significantly associated with egg traits, such as position in the laying sequence, egg mass, yolk testosterone concentration and antioxidant capacity. It was weakly or non‐significantly associated with female and male traits (sexual ornaments), but females laying darker (higher pigment intensity) first clutches had higher hatching success, suggesting that eggshell pigment intensity may predict fitness. Male nestling feeding effort was not predicted by eggshell pigmentation. In addition, females with darker breast plumage colouration (a melanin‐based trait related to fitness) laid highly protoporphyrin‐covered eggs, suggesting the presence of a previously unappreciated link between protoporphyrin biosynthesis and plumage melanisation. Moreover, the proportion of male offspring increased in clutches originating from highly protoporphyrin‐covered eggs, suggesting that parents could acquire visual cues about their future brood sex composition before egg hatching. Our results support the idea that intraspecific signalling via eggshell pigmentation is a species‐specific rather than a general feature of avian taxa.

Relationship between maternal environment and DNA methylation patterns of estrogen receptor alpha in wild Eastern Bluebird (Sialia sialis) nestlings: a pilot study.

There is mounting evidence that, across taxa, females breeding in competitive environments tend to allocate more testosterone to their offspring prenatally and these offspring typically have more aggressive and faster‐growing phenotypes. To date, no study has determined the mechanisms mediating this maternal effect's influence on offspring phenotype. However, levels of estrogen receptor alpha (ER α) gene expression are linked to differences in early growth and aggression; thus, maternal hormones may alter gene regulation, perhaps via DNA methylation, of ER α in offspring during prenatal development. We performed a pilot study to examine natural variation in testosterone allocation to offspring through egg yolks in wild Eastern Bluebirds (Sialia sialis) in varying breeding densities and percent DNA methylation of CG dinucleotides in the ER α promoter in offspring brain regions associated with growth and behavior. We hypothesized that breeding density would be positively correlated with yolk testosterone, and prenatal exposure to maternal‐derived yolk testosterone would be associated with greater offspring growth and decreased ER α promoter methylation. Yolk testosterone concentration was positively correlated with breeding density, nestling growth rate, and percent DNA methylation of one out of five investigated CpG sites (site 3) in the diencephalon ER α promoter, but none in the telencephalon (n = 10). Percent DNA methylation of diencephalon CpG site 3 was positively correlated with growth rate. These data suggest a possible role for epigenetics in mediating the effects of the maternal environment on offspring phenotype. Experimentally examining this mechanism with a larger sample size in future studies may help elucidate a prominent way in which animals respond to their environment. Further, by determining the mechanisms that mediate maternal effects, we can begin to understand the potential for the heritability of these mechanisms and the impact that maternal effects are capable of producing at an evolutionary scale.

Maternal transfer of androgens in eggs is affected by food supplementation but not by predation risk

Mothers may affect the future success of their offspring by varying allocation to eggs and embryos. Allocation may be adaptive based on the environmental conditions perceived during early breeding. We investigated the effects of food supplementation and predation risk on yolk hormone transfer in the pied flycatcher Ficedula hypoleuca. In a food supplementation experiment, females were food‐supplemented prior to and during egg‐laying and androgen concentrations were measured throughout the laying order. Predation risk was investigated in three different studies combining both correlative data, where flycatchers bred in close proximity to two different predator species that prey upon adult flycatchers (either Tengmalm's owl Aegolius funereus or pygmy owl Glaucidium passerinum), and an experimental manipulation, where flycatchers were exposed to cues of a nest predator (least weasel Mustela nivalis). Females receiving food supplementation laid eggs with lower concentrations of androstenedione (A4) than females not receiving food supplements. Yolk testosterone (T) concentration showed the same pattern but the difference was not statically significant. Testosterone (but not A4) concentration increased within clutches, from the first to the last egg, independently of the food supplementation. Females breeding under high predation risk did not differ from control females in their yolk androgen levels (A4, T or progesterone). However, concentrations of A4 tended to be lower in the proximity of pygmy owls, which could indirectly increase offspring survival after fledging. Food supplementation during egg‐laying seems to have a stronger impact on maternal transfer of androgens than predation risk. Food availability and predation risk could differentially affect the trade‐offs of androgen allocation for the offspring when raised in good vs. dangerous environments.

Yolk hormones influence in ovo chemosensory learning, growth, and feeding behavior in domestic chicks.

In this study, we assessed whether prenatal exposure to elevated yolk steroid hormones can influence in ovo chemosensory learning and the behavior of domestic chicks. We simulated a maternal environmental challenge by experimentally enhancing yolk progesterone, testosterone, and estradiol concentrations in hen eggs prior to incubation. The embryos from these hormones-treated eggs (HO) as well as sham embryos (O) that had received the vehicle-only were exposed to the odor of fish oil (menhaden) between embryonic Days 11 and 20. An additional group of control embryos (C) was not exposed to the odor. All chicks were tested following hatching for their feeding preferences between foods that were or were not odorized with the menhaden odor. In the 3-min choice tests, the behavior of O chicks differed significantly according to the type of food whereas C and HO chicks showed no preference between odorized and non-odorized food. Our result suggests weaker response in HO chicks. In addition, HO chicks showed impaired growth and reduced intake of an unfamiliar food on the 24-h time scale compared to controls. Our data suggest that embryonic exposure to increased yolk hormone levels can alter growth, chemosensory learning, and the development of feeding behaviors.

Differential Effects of Maternal Yolk Androgens on Male and Female Offspring: A Role for Sex-Specific Selection?

Maternal hormones are important mediators of prenatal maternal effects in animals. Although their effects on offspring phenotype are often sex-specific, the reason why sometimes sons are more sensitive to prenatal hormone exposure and sometimes daughters is not well understood. Here I combine an experimental manipulation of yolk testosterone concentration in the egg and quantification of selection acting on yolk androgen-sensitive traits in a natural population of great tits (Parus major) with a literature review to test the hypothesis that sex-specific selection on traits affected by yolk androgens determines which sex is more sensitive to prenatal hormone exposure. An experimental increase of the testosterone content in the egg boosted the post-hatching growth of male, but not female great tit nestlings. However, I found no evidence that survival selection on body mass or size is acting differently in the two sexes. A literature review revealed that yolk androgen manipulations affect the growth of males and females differently across species. Interestingly, in studies performed in the wild a significant association between the strength and direction of sexual size dimorphism and sex-specific sensitivities to yolk androgens was observed. In studies performed in captivity, no such relationship was found. Thus, across species there is some evidence that sex-specific selection on body size influences how strongly growth trajectories of males and females are affected by maternally-derived yolk androgens.

Reproductive effort of both male and female Bar‐throated Apalis Apalis thoracica is predicted by ornamentation of self and mate

Melanin‐based plumage ornaments have been shown to play an important role in male–male competition, but also influence inter‐sexual communication. Consequently, ornaments may be associated with reproductive effort of both males and females. Females mated to males with larger melanin ornaments may acquire access to better territories or benefit from increased paternal care. Here we investigated whether the melanin‐based breast‐band of male and female Bar‐throated Apalis Apalis thoracica is a signal of information about its bearer and is associated with male and female reproductive effort. Breast‐band size was a highly variable morphometric trait in both sexes, but only in males was it associated with body mass. We then assessed whether male and female breast‐band size predicted maternal and paternal investment. Egg mass increased with male breast‐band size, but decreased with female breast‐band size. Whether females adjust maternal hormone allocation in response to their partner's ornamentation remains a contentious issue. We found that yolk testosterone and androstenedione concentrations were not predicted by male ornamentation or body mass. Finally, males with larger breast‐bands provided their mates with more food, allowing those females to spend more time incubating. Reproductive effort of both parents is therefore predicted by their own and their mate's ornamentation in Bar‐throated Apalis, and thus breast‐band size potentially acts as a signal of reproductive performance in both sexes. These results highlight the need for more comprehensive analyses of a relationship between melanin‐based ornaments and fitness, incorporating multiple behavioural variables associated with reproductive effort.

High heritability of egg testosterone and consequences on performance of offspring in Japanese quail. A review

DOI: 10.15414/afz.2014.17.04.130–135 Received 4. September 2014 ǀ Accepted 2. October 2014 ǀ Available online 24. November 2014 Individual phenotype is a result of an interacting genotype and environment with the most sensitive periods occurring within the late embryonic and early postnatal development. In this time, most of the signals come from the mother and such influences are termed maternal effects. In our review, we focus on hormone-mediated maternal effects in birds because the avian embryo develops without interactions with neuroendocrine milieu of the mother and the egg yolk contains biologically active substances including hormones. Thus, it is expected that the mother can adjust phenotypic development of her progeny to actual environmental conditions through differential deposition of hormones in the egg. The adaptive value of such hormone mediated maternal effects can have significant consequences for poultry performance but is still generally neglected. In the first part of our review, we summarise the data from wild living birds and urge the need to study maternal egg hormones also in poultry species. In the second part of this review, we present our experimental approach that is based on two genetic lines of Japanese quail divergently selected for yolk testosterone concentrations. We estimated relatively high heritability of yolk testosterone concentrations and proved that natural selection can shape hormone-mediated maternal effects. Our selected lines provided us with a unique model to explore mechanisms of maternal hormone transfer into the egg and epigenetic effects of maternal hormones on offspring. We expect that this approach will provide relevant data in the field of trans-generational maternal effects with potential application for poultry breeding and welfare. Keywords : maternal effects, hormones, androgens, genetic lines

Immune responsiveness of Japanese quail selected for egg yolk testosterone content under severe protein restriction

Yolk testosterone concentrations vary in response to environmental conditions and different testosterone contents can subsequently modify the phenotypic traits of offspring. Apart from effects on growth, proactive behaviour and secondary sexual characteristics, the possible negative impacts of maternal testosterone on the immune system are often considered a limitation for its deposition. The effects of maternal testosterone can be modulated by postnatal environmental conditions, such as the availability of food resources. However, the majority of studies considering the effects of maternal testosterone on the immune system have been conducted under optimum conditions. We evaluated the influence of genetic selection for high (HET) and low (LET) egg testosterone content in Japanese quail on immune responsiveness of offspring to phytohaemagglutinin (PHA) and lipopolysaccharide (LPS) stimulation under severe protein restriction. Protein restriction negatively influenced body weight and performance in the PHA-test. We observed an increase in Cort (corticosterone) and He/Ly (heterophil/lymphocyte ratio) after LPS, while no changes occurred in total IgY levels in the protein-restricted group. HET quails showed higher body mass and total IgY levels and lower He/Ly ratio than LET quails, while the PHA index and Cort concentration did not differ between lines. No interactions were found between protein restriction and genetic line. In conclusion, the immune response was not compromised under conditions of severe protein restriction in the faster growing HET line compared with the LET line. We hypothesise that the immune responsiveness of birds with higher yolk testosterone may be linked with other maternally-derived substances in a context-dependent manner.

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

Maternal resources deposited in eggs can aff ect the development of several off spring phenotypic traits and result in tradeoff s among them. For example, maternal androgens in eggs may be benefi cial to off spring growth and competitive ability, but detrimental to immunocompetence and oxidative stress. In contrast, maternal antioxidants in eggs may be benefi cial if they mitigate oxidative stress and immunosuppressive eff ects of androgens. We investigated possible interactive eff ects of maternal steroids and carotenoids on aspects of off spring 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, off spring survival to age 15 d, and size of the bursa of Fabricius in off spring. In contrast, repeated maternal GnRH challenges, which elevated yolk testosterone concentrations, enhanced off spring neonatal size, but negatively aff ected bursa size. However, interaction among the treatments suggests that the positive eff ect 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 off spring immune system via carotenoids and vitamins deposited in egg yolks and off set detrimental eff ects of androgens deposited by GnRHchallenged females.

Do females invest more into eggs when males sing more attractively? Postmating sexual selection strategies in a monogamous reed passerine.

Maternal investment can play an important role for offspring fitness, especially in birds, as females have to provide their eggs with all the necessary nutrients for the development of the embryo. It is known that this type of maternal investment can be influenced by the quality of the male partner. In this study, we first verify that male song is important in the mate choice of female Eurasian reed warblers, as males mate faster when their singing is more complex. Furthermore, female egg investment varies in relation to male song characteristics. Interestingly, clutch size, egg weight, or size, which can be considered as an high-cost investment, is not influenced by male song characteristics, whereas comparably low-cost investment types like investment into diverse egg components are adjusted to male song characteristics. In line with this, our results suggest that female allocation rules depend on investment type as well as song characteristics. For example, egg white lysozyme is positively correlated with male song complexity. In contrast, a negative correlation exists between-song speed and syllable repetitiveness and egg yolk weight as well as egg yolk testosterone concentration. Thus, our results suggest that female egg investment is related to male song performance in several aspects, but female investment patterns regarding various egg compounds are not simply correlated.

Nest Composition and Yolk Hormones: Do Female European Starlings Adjust Yolk Androgens to Nest Quality?

The nest is a key element of avian reproductive fitness. It provides the developmental environment for the embryo and nestling thereby affecting their quality and survival. Nests are often constructed by the male and nest quality, elaboration, and ornamentation vary among males in a species. Therefore, male nest-building behaviour is likely under sexual selection and females may use male nest-building behaviour and nest features as cues in mate choice and their own reproductive investment. We tested this hypothesis in European starlings ( Sturnus vulgaris). Male starlings present green plants to females during their courtship display and furbish their nests with variable quantities of fresh green plants. The highest frequency of plant incorporation into the nest occurs during pair formation and oogenesis. We determined nest composition and measured yolk testosterone concentrations in eggs. Yolk testosterone concentrations were positively correlated with the amount of green plant material in nests. Because males use green nest material in courtship and green nest material positively affects nestlings’ development, we suggest that female European starlings use male nest-building effort, ritualised courtship display of green plants, and nest characteristics in mate choice, and adjust yolk testosterone levels in the eggs to the quality of the nest to optimise offspring growth.

Between-female variation in house sparrow yolk testosterone concentration is negatively associated with CYP19A1 (aromatase) mRNA expression in ovarian follicles

Maternally-derived yolk androgens influence the development and long-term phenotype of offspring in oviparous species. Between-female variation in the amounts of these yolk androgens has been associated with a number of social and environmental factors, suggesting that the variation is adaptive, but the mechanisms behind it are unknown. Using two different approaches, we tested the hypothesis that variation in yolk androgen levels across individuals is associated with variation in their capacity to synthesize androgens. First, we injected female house sparrows with exogenous gonadotropin-releasing hormone (GnRH) to maximally stimulate ovarian steroidogenesis. Second, we collected pre-ovulatory follicle tissue and quantified the mRNA expression of four key enzymes of the steroid synthesis pathway: steroidogenic acute regulatory protein (StAR), cytochrome P450-side chain cleavage enzyme (CYP11A1), 17β-hydroxysteroid dehydrogenase (HSD17B1), and aromatase (CYP19A1). Thirty minutes after GnRH injection, androgen concentrations in both the plasma and in the yolks of pre-ovulatory follicles were significantly elevated compared to controls. However, this measure of steroidogenic capacity did not explain variation in yolk testosterone levels, although physiological differences between house sparrows and more widely studied poultry models were revealed by this approach. Steroidogenic enzyme mRNA levels were detectable in all samples and were significantly lower in the most mature pre-ovulatory follicles. Of the four measured genes, CYP19A1 expression exhibited a significant negative relationship with yolk testosterone concentrations in laid eggs, revealing a key mechanism for between-female variation in yolk testosterone. Furthermore, this suggests that any factors which alter the expression of CYP19A1 within an individual female could have dramatic effects on offspring phenotype.

Patterns of yolk testosterone deposition in two populations of Arctic-breeding Redpolls

The levels of maternal androgens in egg yolk vary considerably within and among clutches. In most species studied to date, high levels have been found to increase offspring competitive ability. We measured yolk testosterone in two populations of Redpolls (Carduelis flammea/hornemanni) nesting around Toolik Lake Field Station and in Barrow, Alaska to test three predictions about the patterns of yolk testosterone concentrations. First, we predicted yolk testosterone to increase with laying sequence so as to minimize the deleterious effects of hatching asynchrony on chicks from later hatched eggs in a species without delayed incubation. Second, we predicted yolk testosterone to be higher in Barrow where food is more abundant and Redpoll body condition is higher. Third, we predicted yolk testosterone to correlate positively with egg mass and yolk mass, which are two other indicators of maternal investment. An initial analysis that did not consider egg and yolk mass lent support to the first of these predictions. However, more thorough statistical analysis revealed that relationships between yolk testosterone and laying sequence and between yolk testosterone and site were not direct but rather driven by associations among yolk testosterone, egg mass, and yolk mass. While yolk testosterone concentration itself did not differ between sites, the relationship between yolk testosterone and egg and yolk mass did. Yolk testosterone correlated positively with egg mass at Barrow and showed a significant quadratic association with yolk mass in Toolik. While laying sequence and yolk testosterone appeared to be correlated when egg and yolk mass were not considered, the full statistical model suggests that egg and yolk mass are directly associated with laying sequence while yolk testosterone concentrations are not. These results suggest complicated relationships among egg mass, yolk mass, yolk testosterone, site, and laying sequence that should be taken into account in future studies. They also suggest that deposition of yolk testosterone may be a regulated rather than being a purely passive process.

Sources of variation in yolk hormone deposition: Consistency, inheritance and developmental effects

Maternal effects occur when the phenotype of the mother affects the phenotype of their offspring. They are thought to have evolved to translate the environmental conditions experienced by the mother into adaptive phenotypic variation of the offspring. However, the integration of environmental cues allowing adaptive responses requires some form of plasticity that depends on the interaction of the maternal phenotype and her environment. In birds, maternal yolk hormones represent such a pathway for maternal effects, and their adaptive significance depends thus on the plasticity in maternal yolk hormone deposition. We studied sources of variation in yolk testosterone deposition, focusing on the often neglected contribution of the (partly heritable) maternal phenotype. We investigated consistency and heritability of yolk testosterone deposition in captive canaries of which the F 1 generation was raised in foster nests and analyzed the potential effects of the early developmental conditions. We found significant female consistency across years in egg mass, yolk mass and total amount of yolk testosterone but not in yolk testosterone concentrations. Females varied the yolk testosterone concentrations of their eggs across years mainly via changes in yolk mass. The heritable variation in egg mass, yolk mass and amount of yolk testosterone but not yolk testosterone concentrations was within the range of previous studies, but not significantly different from zero. Finally, the growth of the daughters as nestling had a significant effect on their yolk testosterone deposition at adulthood indicating the transgenerational potential for environmental effects – via the effects of yolk hormones on offspring development.

Negative effects of yolk testosterone and ticks on growth in canaries

Maternal yolk hormones in bird eggs are thought to adjust the offspring to the post-hatching environment. This implies that the effects of maternal yolk hormones should vary with the post-hatching environment, but to date such context-dependency has largely been ignored. We experimentally increased yolk testosterone concentrations in canary eggs and simultaneously manipulated the post-hatching context via an experimental tick-infestation of the chicks. This allows us to evaluate the context-dependency of hormone-mediated maternal effects, as it has previously been shown that ectoparasites alter the maternal yolk androgen deposition. The experimental tick infestation reduced growth in chicks from sham-treated eggs, indicating harmful effects of this ectoparasite in canaries. Chicks from testosterone-treated eggs were not affected in their development by ticks, suggesting lower ectoparasite vulnerability. But this may also be due to the fact that experimentally elevated yolk testosterone levels impaired growth even under parasite-free conditions. This contrasts previous studies, but these studies often manipulated first laid eggs, while we used eggs of subsequent laying positions. Later laid eggs are presumably of lower quality and contain higher yolk testosterone concentrations. Thus, the effects of elevated yolk testosterone on growth may be dose-dependent or vary with the egg quality, suggesting prenatal context-dependency.

Eavesdropping on Male Singing Interactions Leads to Differential Allocation in Eggs

AbstractFemales invest differently in their eggs depending on the quality of their mates. In oscines, female investment is influenced by the quality of male song. In domestic canaries Serinus canaria, as well as in black‐capped chickadees Poecile atricapillus, females pay attention not only to the intrinsic quality of male song but can also gather information, by eavesdropping on male–male singing interactions, on the relative quality of males. During these interactions, overlapping the song of the rival is more threatening than alternating. Moreover eavesdropping female canaries have been shown to prefer the overlapping song rather than the overlapped song. The present study was designed to assess the effect of the information gathered by eavesdropping on female investment in eggs. First, we broadcasted overlapping interactions to female canaries. Then, we broadcasted to each female one of the two songs previously heard and collected eggs. Females exposed to overlapping songs laid eggs with greater yolk ratio than females exposed to overlapped songs. In contrast, yolk testosterone quantity and concentration were not affected by the treatment. Moreover, we found a variation between eggs with regard to the testosterone deposited in yolk: both quantity and concentration increased with laying order. Our results suggest that female canaries use information gathered by eavesdropping to differentially allocate resources into the eggs. They suggest that singing interactions could influence chick quality via female investment.

Influence of mating preferences on yolk testosterone in the grey partridge

In precocial bird species, the eggs constitute most of the maternal investment because parents do not feed the young after they hatch. Maternal testosterone in egg yolk influences the embryo’s and chick’s development. Females deposit testosterone in the eggs as a response to the environment experienced during the laying period, including the quality of their mate. To assess the relevance of the female’ sm ate selection on egg characteristics in the grey partridge, Perdix perdix, we tested breeding females in a choice trial where they were allowed to choose between two males. After the choice trials, females were mated either with their preferred male (P group) or with the nonpreferred one (NP group). Although eggs laid by females of the two groups did not differ significantly in mass, females of the P group laid eggs with a higher yolk testosterone concentration than females of the NP group. This study agrees with previous work pointing out that partner attractiveness may play an important role in the transfer of maternally derived egg components. 2010 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. Maternal effects imply that the environment and the phenotype of the mother affect the progeny’s phenotype (Mousseau & Fox 1998). In oviparous vertebrates, all the resources needed by the embryo to develop must be present in the egg laid by the female. After the eggs are laid, no further adjustments to their components

Maternal yolk testosterone in canary eggs: toward a better understanding of mechanisms and function

Maternal yolk androgens in avian eggs have been shown to affect numerous offspring traits. These changes in offspring phenotype represent examples of maternal effects and are thought to adjust offspring development to the posthatching environment. When studying the functional consequences of yolk hormones it is, therefore, crucial to manipulate the yolk androgen concentrations as well as the environmental conditions under which the study is performed. However, so far context-dependent effects of maternal yolk hormones have not sufficiently been taken into account, which might contribute to the current level of inconsistency in yolk androgen-mediated effects. We experimentally elevated the yolk testosterone concentrations and manipulated the sibling size hierarchy. We focused on the effects of yolk testosterone on growth and monitored begging behavior and parental feeding preferences in search of the underlying mechanisms of changes in growth. Experimental changes in the yolk testosterone concentrations significantly affected offspring phenotype. However, elevated yolk testosterone concentrations only improved the growth of chicks that were at a competitive disadvantage, whereas it benefited all chicks when placed in staged competition for food as juvenile. This emphasizes the compensatory role of yolk androgens in the context of hatching asynchrony and its context dependency. Enhanced growth did not coincide with intensified begging. Neither males nor females preferentially fed chicks hatching from eggs with elevated yolk testosterone concentrations. Enhanced growth rather resulted from yolk testosterone induced changes in physiology, of which the detailed mechanisms are yet unknown.