Growth of yellow-headed blackbird Xanthocephalus xanthocephalus nestlings in relation to maternal body condition, egg mass, and yolk carotenoids concentrations

Condition‐dependent resource allocation to eggs can affect offspring growth and survival, with potentially different effects on male and female offspring, particularly in sexually dimorphic species. We investigated the influence of maternal body condition (i.e., mass‐tarsus residuals) and two measures of female resource allocation (i.e., egg mass, yolk carotenoid concentrations) on nestling mass and growth rates in the polygynous and highly size dimorphic yellow‐headed blackbird Xanthocephalus xanthocephalus. Egg characteristics and carotenoid concentrations were obtained from the third‐laid egg of each clutch and were correlated with the mass and growth rates of the first two asynchronously hatched nestlings. Maternal body condition was associated with the growth of first‐hatched, but not second‐hatched nestlings. Specifically, females in better body condition produced larger and faster growing first‐hatched nestlings than females in poorer body condition. As predicted for a polygynous, size‐dimorphic species, females that fledged first‐hatched sons were in better body condition than females that fledged first‐hatched daughters. Associations between egg mass, yolk carotenoid content, and nestling growth were also specific to hatching‐order. Egg mass was positively correlated with the mass and growth rates of second‐hatched nestlings, and yolk concentrations of β‐carotene were positively correlated with second‐hatched nestling mass. Surprisingly, the relationship between yolk lutein and hatchling growth differed between the sexes. Females with high concentrations of yolk lutein produced larger and faster growing first‐hatched sons, but smaller first‐hatched daughters than females with lower lutein concentrations. Mass and growth rates did not differ between first‐ and second‐hatched nestlings of the same sex, despite asynchronous hatching in the species. Results from this study suggest that maternal body condition and the allocation of resources to eggs have carotenoid‐, sex‐, and/or hatch‐order‐specific effects on yellow‐headed blackbird nestlings.     

Sex-specific effects of yolk testosterone on survival, begging and growth of zebra finches

Yolk androgens affect offspring hatching, begging, growth and survival in many bird species. If these effects are sex-specific, yolk androgen deposition may constitute a mechanism for differential investment in male and female offspring. We tested this hypothesis in zebra finches. In this species, females increase yolk-testosterone levels and produce male-biased sex ratios when paired to more attractive males. We therefore predicted that especially sons benefit from elevated yolk androgens. Eggs were injected with testosterone or sesame oil (controls) after 2 days of incubation. Testosterone had no clear effect on sex-specific embryonic mortality and changed the pattern of early nestling mortality independent of offspring sex. Testosterone-treated eggs took longer to hatch than control eggs. Control males begged significantly longer than females during the first days after hatching and grew significantly faster. These sex differences were reduced in offspring from testosterone-treated eggs due to prolonged begging durations of daughters, enhanced growth of daughters and reduced growth of sons. The results show that variation in maternal testosterone can play an important role in avian sex allocation due to its sex-specific effects on offspring begging and growth.     

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.     

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.     

Experimental manipulation of egg carotenoids affects immunity of barn swallow nestlings

The yolk of bird eggs contains maternal carotenoids that may act as antioxidants thus influencing offspring performance and survival. However, to our knowledge, this hypothesis has not been subjected to experimental tests and the function of transmission of carotenoids to the egg is largely unknown. We directly manipulated the concentration of the main carotenoid (lutein) in the eggs of barn swallows (Hirundo rustica) and analysed the effect of experimental manipulation on growth of nestlings and two fundamental components of their acquired immunity. Nestlings hatched from lutein-inoculated eggs had larger T-cell-mediated immune response compared with those of two control groups. T-cell-mediated immune response predicted nestling survival until fledging. However, lutein inoculation did not affect antibody response to an immunogen, body mass, tarsus length or plumage development. Nestling body mass and plumage development declined with egg laying order, but the effects of lutein inoculation were independent of egg laying order for all traits. Our results show that maternal yolk carotenoids can have a major effect in promoting a fundamental component of immunity that predicts offspring survival and suggests that adaptive early maternal effects can be mediated by transmission of antioxidants to eggs.     

Differential maternal testosterone allocation among siblings benefits both mother and offspring in the zebra finch Taeniopygia guttata

Parents are selected to preferentially invest in the offspring with highest reproductive value. One mechanism for achieving this is the modification of competitive asymmetries between siblings by maternal hormones. In many organisms, offspring value varies according to birth position in the brood, which determines survival chances and competitive advantage over access to resources. In birds, variation in yolk androgen allocation over the laying sequence is thought to modulate dominance of senior chicks over junior brood mates. We tested this hypothesis in zebra finches, which show a naturally decreasing pattern of within-clutch testosterone allocation. We abolished these within-clutch differences by experimentally elevating yolk testosterone levels in eggs 2-6 to the level of egg 1, and we assessed fitness measures for junior offspring (eggs 2-6), senior offspring (egg 1), and their mothers. Testosterone-injected eggs hatched later than control eggs. Junior, but not senior, chicks in testosterone-treated broods attained poorer phenotypic quality compared to control broods, which was not compensated for by positive effects on seniors. Mothers were generally unaffected by clutch treatment. Thus, naturally decreasing within-clutch yolk testosterone allocation appears to benefit all family members and does not generally enhance brood reduction by favoring senior chicks, in contrast to the widely held assumption.     

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.     

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.     

Maternal allocation of androgens and antagonistic effects of yolk androgens on sons and daughters

Mothers can influence the phenotype of their offspring by adjustingthe quality of their eggs in relation to sex and reproductivevalue of the progeny. Maternal androgens in the eggs of vertebratesmay mediate such adaptive early maternal effects. However, theevolution of early maternal effects mediated by egg androgensmay be constrained by the inability of mothers to differentiallyallocate androgens to eggs with a male or a female if androgenshave different effects on sons and daughters. In this study,we increased the concentration of androgens in the eggs of barnswallows (Hirundo rustica) within the physiological range ofvariation and analyzed the effect on nestling growth and beggingbehavior. Egg androgens increased body size and mass of sonsbut reduced these characters in daughters when compared to twocontrol groups in a repeated-measures analysis of variance ofdata collected at different ages. However, the differentialeffect of androgen on the two sexes was no longer significantwhen the analysis was restricted to the age of 12 days, whenfinal body size is attained. In a second experiment, we testedwhether mothers differentially allocated androgens to eggs withsons rather than daughters while manipulating a paternal secondarysexual character. Androgen concentration did not vary in relationto paternal ornamentation or embryo sex. Hence, antagonisticeffects of egg androgens on sons and daughters may exist inthe very early posthatching life and may constrain the evolutionof adaptive maternal effects because mothers do not differentiallyallocate androgens in relation to embryo sex.     

Maternal antibodies but not carotenoids in barn swallow eggs covary with embryo sex

Mothers influence their offspring phenotype by varying egg quality. Such maternal effects may be mediated by transmission of antibodies and antioxidants. Mothers should adjust allocation of maternal substances depending on embryonic sex because of differences in reproductive value, potentially dependent on paternal genetic effects as reflected by secondary sexual characters. We manipulated sexual attractiveness of male barn swallows (Hirundo rustica) and investigated maternal investment in eggs in relation to offspring sex. Mothers allocated more antibodies against a pathogen to eggs with a daughter than a son. However, concentration of antioxidants was independent of embryonic sex. Sex-dependent allocation was independent of paternal attractiveness. Thus, mothers adjusted allocation of substances to offspring in a complex manner, that may be part of a strategy of favouritism of daughters, which have larger mortality than sons. Such effects may have important consequences for secondary and tertiary sex ratios, but also for ontogeny of adult phenotype.     

Evolution of sex-biased maternal effects in birds: III. Adjustment of ovulation order can enable sex-specific allocation of hormones, carotenoids, and vitamins

Overlap in growth of offspring should constrain the opportunity for sex-biased maternal effects, yet sex-specific allocation of maternal resources among simultaneously growing ova is often observed in vertebrates. In birds, such allocation can be accomplished either by temporal clustering of ova that become the same sex, resulting in sex-biased egg-laying order, or by follicle-specific delivery of maternal resources. Two house finch populations at the northern and southern boundaries of the species range have opposite ovulation sequences of male and female eggs, and thus, in the absence of sex differences in ova growth or sex-specific maternal strategies, would be expected to have opposite sex-specific accumulation of maternal products. We found that the populations had strong and similar gradients of steroid distribution in relation to ovulation order, whereas distribution of carotenoids and vitamins correlated with each follicle's accumulation of steroids. In both populations, temporal bias in production of sons and daughters within a clutch enabled strongly sex-specific acquisition of maternal products, and oocytes of the same sex were highly interdependent in their accumulation of steroids. Moreover, in nests where the sex-bias in relation to ovulation order deviated from population-specific patterns, eggs had highly distinct concentrations of steroids, carotenoids and vitamins. These results and previous findings of sex-specific yolk partitioning among oocytes suggest that oocytes that become males and females are temporally or spatially clustered during their ovarian growth. We discuss the implication of these findings for the evolution of sex-specific maternal resource allocation.     

Laying order and offspring sex in blue tits Parus caeruleus

According to sex allocation theory, females may benefit from a differential investment in sons and daughters, where fitness returns from male and female offspring vary. One way in which investment may be differentiated is by assigning male and female offspring to eggs of different sizes, and/or eggs laid early or late in the laying order. Here, we investigate whether such sex-specific adjustments occur in blue tits Parus caeruleus . We found that the proportion of males changed non-linearly with laying order showing the increase in early-laid eggs. Egg size increased with laying order, but this pattern differed between female and male eggs. Female eggs increased initially in size and became smaller later in the laying sequence, while male eggs exhibited constant increase in mass. However, egg size did not differ between genders. Egg size and laying order was found to interact in moulding sex of the embryo. Thus, females seem to simultaneously adjust the sex of the offspring in relation to their investments into eggs and the position of the egg in the laying sequence.     

Different optimal offspring sizes for sons versus daughters may favor the evolution of temperature-dependent sex determination in viviparous lizards

Temperature-dependent sex determination (TSD) has evolved independently in at least two lineages of viviparous Australian scincid lizards, but its adaptive significance remains unclear. We studied a montane lizard species (Eulamprus heatwolei) with TSD. Our data suggest that mothers can modify the body sizes of their offspring by selecting specific thermal regimes during pregnancy (mothers with higher and more stable temperatures produced smaller offspring), but cannot influence sons versus daughters differentially in this way. A field mark-recapture study shows that optimal offspring size differs between the sexes: larger body size at birth enhanced the survival of sons but reduced the survival of daughters. Thus, a pregnant female can optimize the fitness of either her sons or her daughters (via yolk allocation and thermoregulation), but cannot simultaneously optimize both. One evolutionary solution to reduce this fitness cost is to modify the sex-determining mechanism so that a single litter consists entirely of either sons or daughters; TSD provides such a mechanism. Previous work has implicated a sex difference in optimal offspring size as a selective force for TSD in turtles. Hence, opposing fitness determinants of sons and daughters may have favored evolutionary transitions from genetic sex determination to TSD in both oviparous turtles and viviparous lizards.     

Does the differential cost of sons and daughters lead to sex ratio adjustment in great frigatebirds Fregataminor?

Sex allocation theory predicts that if benefits of producing sons and daughters differ and outweigh the costs of sex ratio adjustment, parents should produce more of the offspring that provide them with greater fitness. Potential benefits may be more likely to outweigh costs where sexual size dimorphism and, in birds, single‐egg clutches exist. Great frigatebirds Fregataminor are seabirds in which females are larger than males and clutch size is one egg. In our study population, sexual size dimorphism develops primarily during the period of complete juvenile dependence on parental care, consistent with a higher cost of producing daughters than sons. Over the course of the 1998 breeding season there was a shift from early season prevalence of daughters to late‐season prevalence of sons. Variation in food availability at time of egg laying, as indexed by sea surface temperature (SST), was a strong predictor of offspring sex in 1998. In contrast, SST in 2003 was not a predictor of offspring sex, nor was there a seasonal shift in the hatching sex ratio, despite a seasonal shift in SST. Besides food availability, we tested two additional factors in 2003 that could explain sex ratio adjustment in relation to the cost of reproduction. Offspring sex in 2003 was not related to natural or experimentally induced variation in maternal body condition; pre‐laying food supplements raised the body condition of females at the time of egg laying but did not affect offspring sex or egg mass. In addition, offspring sex was not predicted by the length of maternal telomere restriction fragments (TRFs), an index of age and possibly of reproductive experience. Broad confidence intervals on effect size suggest that undetected effects of maternal condition on offspring sex ratio could easily exist, but confidence intervals were narrower on the non‐significant effects of SST and TRF length on offspring sex ratio. The cause of different seasonal patterns of hatching sex ratio and different SST effects in 1998 and 2003 is unclear.     

Within‐female plasticity in sex allocation is associated with a behavioural polyphenism in house wrens

Sex allocation theory assumes individual plasticity in maternal strategies, but few studies have investigated within‐individual changes across environments. In house wrens, differences between nests in the degree of hatching synchrony of eggs represent a behavioural polyphenism in females, and its expression varies with seasonal changes in the environment. Between‐nest differences in hatching asynchrony also create different environments for offspring, and sons are more strongly affected than daughters by sibling competition when hatching occurs asynchronously over several days. Here, we examined variation in hatching asynchrony and sex allocation, and its consequences for offspring fitness. The number and condition of fledglings declined seasonally, and the frequency of asynchronous hatching increased. In broods hatched asynchronously, sons, which are over‐represented in the earlier‐laid eggs, were in better condition than daughters, which are over‐represented in the later‐laid eggs. Nonetheless, asynchronous broods were more productive later within seasons. The proportion of sons in asynchronous broods increased seasonally, whereas there was a seasonal increase in the production of daughters by mothers hatching their eggs synchronously, which was characterized by within‐female changes in offspring sex and not by sex‐biased mortality. As adults, sons from asynchronous broods were in better condition and produced more broods of their own than males from synchronous broods, and both males and females from asynchronous broods had higher lifetime reproductive success than those from synchronous broods. In conclusion, hatching patterns are under maternal control, representing distinct strategies for allocating offspring within broods, and are associated with offspring sex ratios and differences in offspring 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.     

No sexual differences in embryonic period in jackdaws Corvus monedula and black-headed gulls Larus ridibundus

Offspring survival probability usually decreases with hatching order, especially in species with brood reduction. Brood reduction in combination with a sex difference in embryonic period (the time between laying and hatching of an egg) can potentially have a profound effect on sex allocation, with higher investment in chicks of the early hatching sex because they are more likely to survive to fledge. Two recent studies reported sex differences in the embryonic period, but compared embryonic period between, rather than within, clutches, which does not control for possible environmental effects on both clutch sex ratio and embryonic period. We compared the embryonic period of sons and daughters within clutches in jackdaws Corvus monedula and black-headed gulls Larus ridibundus , two species with frequent brood reduction, and found no sexual difference in embryonic period. This suggests that sex allocation is not affected by sex differences in embryonic period in these species, but more studies are required to verify whether this is a general pattern.     

Flaxseed and dried tomato waste used together in laying hens diet

The aim of this study was to investigate the effects of simultaneous supplementation of laying hens with dietary sources of n-3 polyunsaturated fatty acids (PUFA) and carotenoids on egg quality, fatty acids and carotenoid profile of the egg yolk and on feed and yolk lipid peroxidation. A 6-week experiment was carried out with 53-week old laying hens (96 Tetra SL) assigned to a control and three treatment groups supplemented with 5% flaxseeds and different levels of dried tomato waste (DTW, 2.5%, 5.0% and 10.0%). Hens from the groups supplemented with 5% and 7.5% DTW had a significantly lower average daily feed intake and laying percentage as compared to the control. Increased doses of dietary DTW enhanced yolk Roche colour score in direct correlation with the enrichment of egg yolk in carotenoids but decreased their transfer efficiency from feed to egg. After 4 weeks, egg yolk from hens fed with 5% flaxseeds and 7.5% DTW had increased lutein and zeaxanthin levels (by 29% and 24%, respectively) and the colour score was 3.5 fold higher compared to the control group. As a result of the dietary supplementation with flaxseed, the n-3 fatty acid content was 3.1–3.7-fold higher in egg yolk compared with the control and the n-6/n-3 ratio decreased from 18.3 (control) to 4.1–5.4 in supplemented diets. Dietary supplementation with 5% DTW effectively prevented lipid oxidation of eggs enriched with n-3 PUFA, but the increase in DTW content depressed the absorption and deposition of n-3 PUFA in egg yolk.     

Maternal nutrition affects reproductive output and sex allocation in a lizard with environmental sex determination

Life-history traits such as offspring size, number and sex ratio are affected by maternal feeding rates in many kinds of animals, but the consequences of variation in maternal diet quality (rather than quantity) are poorly understood. We manipulated dietary quality of reproducing female lizards (Amphibolurus muricatus; Agamidae), a species with temperature-dependent sex determination, to examine strategies of reproductive allocation. Females maintained on a poor-quality diet produced fewer clutches but massively (twofold) larger eggs with lower concentrations of yolk testosterone than did conspecific females given a high-quality diet. Although all eggs were incubated at the same temperature, and yolk steroid hormone levels were not correlated with offspring sex, the nutrient-deprived females produced highly male-biased sex ratios among their offspring. These responses to maternal nutrition generate a link between sex and offspring size, in a direction likely to enhance maternal fitness if large body size enhances reproductive success more in sons than in daughters (as seems plausible, given the mating system of this species). Overall, our results show that sex determination in these animals is more complex, and responsive to a wider range of environmental cues, than that suggested by the classification of 'environmental sex determination'.     

Do mothers regulate facultative and obligate siblicide by differentially provisioning eggs with hormones?

We examined androgens in clutches of two booby species that differ in their sibling conflict. Blue-footed booby Sula nebouxii chicks show an aggression-submission relationship, aggression is normally moderate and siblicide is facultative. Brown booby Sula leucogaster chicks show an aggression-aggression relationship, aggression of both chicks can be relentless and siblicide is obligate. The parental favoritism hypothesis predicts that egg mass, yolk mass and yolk androgens should decline with laying order less in the blue-footed booby than in the brown booby, to promote the survival of the former's junior chick. The eggs of the blue-footed booby had higher yolk concentrations of 5α-dihydrotestosterone (DHT) and lower concentrations of testosterone (T); both species had similarly high yolk concentrations of androstenedione (A). Intra-clutch variation in yolk DHT, T and A failed to support our predictions. In both species, first and second eggs showed similar concentrations of all three hormones and were of similar size, although in the blue-footed booby (only) yolk masses declined with laying order. Brown booby mothers make junior chicks vulnerable to siblicide by hatching them 5 d after their broodmates, but not by differential allocation of egg androgens or nutrients. Blue-footed booby mothers appear to prepare junior chicks for thriving in a subordinate non-provocative role by hatching them 4 d after their broodmates and providing them with 10% less yolk. To orchestrate agonism within the brood, these boobies may rely more on hatch intervals and yolk provision than on androgens.