Effects of elevated yolk androgens on perinatal begging behavior in yellow-legged gull (Larus michahellis) chicks

Maternal hormones may represent an important pathway by which mothers can adaptively adjust offspring traits and performance to suit the prevailing environmental conditions. Earlier studies of birds have shown that egg androgens of maternal origin may enhance post-natal offspring 'begging' displays, functioning to solicit parental care. Here we investigate the effects of elevated egg androgen levels on the prenatal begging behavior of yellow-legged gull (Larus michahellis) chicks. At laying, we experimentally increased the concentration of yolk testosterone (T) within the natural range of variation, and, shortly before hatching, we compared the structural properties, rate, and loudness of vocalizations of embryos developing in T- and oil-injected (control) eggs. In addition, we compared the early post-hatch begging rate (measured as the pecking rate towards a dummy gull head) in chicks of the two experimental groups. We found that T embryos produced louder embryonic vocalizations than controls, whereas structural properties and the calling rate did not differ between T and control embryos. The post-hatch begging rate was unaffected by T treatment, but strongly decreased with increasing chick body mass, suggesting that intensity of the begging display was sensitive to chick state and may therefore reliably indicate the need of food in this species. Therefore, the results of this study show for the first time that prenatal T exposure modulates the quality of embryonic vocalizations, but are not in accordance with previous findings reporting increased post-hatching begging intensity following increased prenatal exposure to androgens.     

Darwin's Finch Begging Intensity Does Not Honestly Signal Need in Parasitised Nests

Parental care should be selected to respond to honest cues that increase offspring survival. When offspring are parasitised, the parental food compensation hypothesis predicts that parents can provision extra food to compensate for energy loss due to parasitism. Chick begging behaviour is a possible mechanism to solicit increased feeding from attending parents. We experimentally manipulated parasite intensity from Philornis downsi in nests of Darwin's small ground finch (Geospiza fuliginosa) to test its effects on chick begging intensity and parental food provisioning. We used in‐nest video recordings of individually marked chicks to quantify nocturnal parasite feeding on chicks, subsequent diurnal chick begging intensity and parental feeding care. Our video analysis showed that one chick per brood had the highest parasite intensity during the night (supporting the tasty chick hypothesis) and weakest begging intensity during the day, which correlated with low parental care and rapid death. We observed sequential chick death on different days rather than total brood loss on a given day. Our within‐nest video images showed that (1) high nocturnal larval feeding correlated with low diurnal begging intensity and (2) parent birds ignored weakly begging chicks and provisioned strongly begging chicks. Excluding predation, all parasite‐free chicks survived (100% survival) and all parasitised chicks died in the nest (100% mortality). Weak begging intensity in parasitised chicks, which honestly signalled recent parasite attack, was not used as a cue for parental provisioning. Parents consistently responded to the strongest chick in both parasitised and parasite‐free nests.     

Equivocal Responses of Feeding Parents to Experimental Brood Sizes in a Hawk–Cuckoo Host: Brood Size as a Reference for Parental Provisioning Decisions?

The number of offspring surviving until independence is the fundamental drive in the evolution of parental care. Because of the related costs, parental investment must be balanced with essential resources for parents themselves, among the resources available in the environment under the current parental condition. It is advantageous for parents to adjust their level of investment to the number of offspring; however, there is little evidence that parents employ numerical competence in adjusting their investment level. We investigated how parents respond to experimentally manipulated brood sizes in a passerine species, known as a host of a brood parasitic cuckoo whose chicks presumably deceive their hosts numerically. Parents reduced their provisioning to broods of reduced sizes, whereas parents did not increase provisioning to enlarged broods compared to that in the control condition. These parental responses can be attributed to the response of chicks to the experimental treatments compared to that in the control: chicks lowered begging intensity in the reduced condition, while they did not intensify being in the enlarged condition. Further analyses revealed that eagerness of parents to respond to chick begging intensity differed between the experimental treatments: strong parental response was detected toward begging chicks only in the reduced condition. We propose that the detected equivocality of parental responses might be related to the difference in the number of chicks between the unmanipulated and experimentally manipulated broods, the former reflecting the initial parental decision on the amount of resources to allocate to the brood.     

Hatching Asynchrony Decreases the Magnitude of Parental Care in Domesticated Zebra Finches: Empirical Support for the Peak Load Reduction Hypothesis

Parent–offspring conflict over the supply of parental care results in offspring attempting to exert control using begging behaviours and parents attempting to exert control by manipulating brood sizes and hatching patterns. The peak load reduction hypothesis proposes that parents can exert control via hatching asynchrony, as the level of competition amongst siblings is determined by their age differences and not by their growth rates. Theoretically, this benefits the parents by reducing both the peak load of the offspring's demand and their overall demand for food and benefits the offspring by reducing the amplification of their competition. However, the peak load reduction hypothesis has only received mixed support. Here, we describe an experiment where we manipulated the hatching patterns of domesticated zebra finch Taeniopygia guttata broods and quantified patterns of nestling begging and parental feeding effort. There was no difference in the begging intensity of nestlings raised in asynchronous or experimentally synchronous broods, yet parental feeding effort was lower when provisioning asynchronous broods and particularly so when levels of nestling begging were low. Further, both parents acted in unison, as there was no evidence of parentally biased favouritism in relation to hatching pattern. Therefore, our study provided empirical support for the prediction that hatching asynchrony reduces the feeding effort of parents, thereby providing empirical support for the peak load reduction hypothesis.     

Coadaptation of offspring begging and parental provisioning: A role for prenatal maternal effects?

Recent studies on birds have shown that offspring begging and parental provisioning covary at the phenotypic level, which is thought to reflect genetic correlations. However, prenatal maternal factors, like yolk testosterone, may also facilitate parent-offspring coadaptation via their effects on offspring begging and development. In fact, maternal effects are thought to adjust offspring phenotype to the environmental conditions they will experience after birth, which are in turn strongly dependent on the levels of parental provisioning. Using cross-fostering experiments in canaries, we tested the role of maternal effects on parent-offspring coadaptation from two different approaches. First, we analyzed whether females deposit yolk testosterone in relation to their own or their partner's prospective parental provisioning, measured as the rate of parental feeding to foster nestlings. Second, we investigated whether females deposit yolk testosterone in relation to costs they incurred when raising a previous brood, as this likely impinges on their capacity to provide parental care in the near future. However, from the results of both experiments we have no evidence that canary females deposit yolk testosterone in order to match offspring begging to the levels of care they and/or their partners provide. We therefore found no evidence that yolk testosterone facilitates parent-offspring coadaptation. In addition, our results suggest that the functional consequences of yolk testosterone deposition may relate to hatching asynchrony since it primarily varied with egg laying order.     

Sex roles in nest keeping – how information asymmetry contributes to parent‐offspring co‐adaptation

Parental food provisioning and offspring begging influence each other reciprocally. This makes both traits agents and targets of selection, which may ultimately lead to co‐adaptation. The latter may reflect co‐adapted parent and offspring genotypes or could be due to maternal effects. Maternal effects are in turn likely to facilitate in particular mother‐offspring co‐adaptation, further emphasized by the possibility that mothers are sometimes found to be more responsive to offspring need. However, parents may not only differ in their sensitivity, but often play different roles in postnatal care. This potentially impinges on the access to information about offspring need. We here manipulated the information on offspring need as perceived by parents by playing back begging calls at a constant frequency in the nest‐box of blue tits (Cyanistes caeruleus). We measured the parental response in provisioning to our treatment, paying particular attention to sex differences in parental roles and whether such differences alter the perception of the intensity of our manipulation. This enabled us to investigate whether an information asymmetry about offspring need exists between parents and how such an asymmetry relates to co‐adaptation between parental provisioning and offspring begging. Our results show that parents indeed differed in the frequency how often they perceived the playback due to the fact that females spent more time with their offspring in the nest box. Correcting for the effective exposure of an adult to the playback, the parental response in provisioning covaried more strongly (positive) with offspring begging intensity, independent of the parental sex, indicating coadaptation on the phenotypic level. Females were not more sensitive to experimentally increased offspring need than males, but they were exposed to more broadcasted begging calls. Therefore, sex differences in access to information about offspring need, due to different parental roles, have the potential to impinge on family conflicts and their resolution.     

European starling chicks benefit from high yolk testosterone levels during a drought year

Avian egg yolk contains androgenic hormones, such as testosterone, of maternal origin. Experimental elevation of yolk testosterone levels enhances growth of canary chicks. Success in sibling competition, due to increased begging, is presumed to underlie this growth enhancement, because canary hatchlings from testosterone-treated eggs beg longer in response to vibrational stimuli than controls. Furthermore, experimental elevation of both yolk androstenedione and testosterone increased chick growth and begging in black-headed gulls. We measured daily growth of European starling (Sturnus vulgaris) chicks hatching from testosterone-treated or vehicle-treated (control) eggs until 14 days of age, and measured begging behavior at hatching and at 5 days of age. A temporary drought caused relatively high levels of early brood reduction for this population; 2- and 3-day-old chicks were most likely to starve. We found that chicks from testosterone-treated eggs were less likely to starve than control chicks, and were heavier on the days when most brood reduction occurred. However, chicks from testosterone-treated eggs begged less than control chicks on the day of hatching, and begged similarly at 5 days of age. Thus, while yolk testosterone did increase growth during periods of (presumably) high competition, increased begging does not appear to mediate this effect. Instead, testosterone may induce more efficient energy use, for example, by decreasing ineffective begging. While our results indicate that elevated yolk testosterone enhances survival, and thus offspring and parental fitness, further evidence regarding the fitness consequences of yolk androgens are vital to understanding their role in avian life history.     

Coadaptation of Offspring Begging and Parental Provisioning - An Evolutionary Ecological Perspective on Avian Family Life

Offspring begging and parental provisioning are the two central social behaviours expressed during the period of parental care. Both behaviours influence each other and it is, therefore, hypothesized that they should ultimately become (genetically) correlated, stabilized by fitness costs to parents and/or offspring. By reciprocally exchanging entire clutches in canaries (Serinus canaria), we tested (1) whether there is covariation between these behaviours and (2) whether a mismatch - as introduced by cross-fostering - entails costs. Begging was scored in a standardized begging test and parental provisioning was measured via (a) the actual feeding rate and (b) using the growth rate of the foster nestlings as a proxy. Costs were established in terms of future reproductive investment in subsequent clutches and offspring growth. We found a positive and significant phenotypic covariation between offspring begging and parental feeding when using the growth rate as a proxy and, to a lesser extent, in case of the parental feeding rate. Female parents suffered no future reproductive costs when feeding foster nestlings that were more demanding than their own nestlings. Neither growth measured amongst all offspring nor the reproductive investment measured amongst the female offspring as adults was influenced by their begging behaviour. However, the reproductive investment of female offspring tended to depend on the parental qualities of their foster parents. Thus, offspring may only be able to extract resources within the limit of generosity of their foster parents. This suggests parental control of feeding, which is also supported by the positive covariation between offspring begging and parental feeding.     

Food Availability and Maternal Immunization Affect Transfer and Persistence of Maternal Antibodies in Nestling Pigeons

The ability of mothers to transfer antibodies (Abs) to their young and the temporal persistence of maternal Abs in offspring constitute important life-history traits that can impact the evolution of host-parasite interactions. Here, we examined the effects of food availability and parental immunization on the transfer and persistence of maternal antibodies in nestling pigeons (Columba livia). This species can transmit maternal Abs to offspring before hatching through the egg yolk and potentially after hatching through crop milk. However, the role of this postnatal substance in immunity remains elusive. We used a full cross-fostering design to disentangle the effects of food limitation and parental immunization both before and after hatching on the levels and persistence of maternal Abs in chicks. Parents were immunized via injection with keyhole limpet hemocyanin antigens. Using an immunoassay that specifically detected the IgY antibodies that are known to be transmitted via the yolk, we found that the levels of anti-KLH Abs in newly hatched chicks were positively correlated with the levels of anti-KLH Abs in the blood of their biological mothers. However, this correlation was not present between chicks and their foster parents, suggesting limited IgY transfer via crop milk to the chick’s bloodstream. Interestingly, biological mothers subjected to food limitation during egg laying transferred significantly fewer specific maternal Abs, which suggests that the transfer of antibodies might be costly for them. In addition, the persistence of maternal Abs in a chick’s bloodstream was not affected by food limitation or the foster parents’ anti-KLH Ab levels; it was only affected by the initial level of maternal anti-KLH Abs that were present in newly hatched chicks. These results suggest that the maternal transfer of Abs could be costly but that their persistence in an offspring’s bloodstream may not necessarily be affected by environmental conditions.     

Maternal investment tactics in superb fairy-wrens

In cooperatively breeding species, parents often use helper contributions to offspring care to cut their own costs of investment (i.e. load-lightening). Understanding the process of load-lightening is essential to understanding both the rules governing parental investment and the adaptive value of helping behaviour, but little experimental work has been conducted. Here we report the results of field experiments to determine maternal provisioning rules in cooperatively breeding superb fairy-wrens (Malurus cyaneus). By manipulating carer: offspring ratios, we demonstrate that helpers allow females to reduce the rate at which they provision their brood. Female reductions, however, were less than that provided by helpers, so that chicks still received food at a faster rate in the presence of helpers. Despite this, chicks fed by parents and helpers were not heavier than those provisioned by parents alone. This is because maternal load-lightening not only occurs during the chick provisioning stage, but also at the egg investment stage. Theoretically, complete load-lightening is predicted when parents value themselves more highly than their offspring. We tested this idea by 'presenting' mothers with a 'choice' between reducing their own levels of care and increasing investment in their offspring. We found that mothers preferred to cut their contributions to brood care, just as predicted. Our experiments help to explain why helper effects on offspring success have been difficult to detect in superb fairy-wrens, and suggest that the accuracy with which theoretical predictions of parental provisioning rules are matched in cooperative birds depends on measuring maternal responses to helper presence at both the egg and chick stages.     

Structural (UV) and carotenoid‐based plumage coloration – signals for parental investment?

Parental care increases parental fitness through improved offspring condition and survival but comes at a cost for the caretaker(s). To increase life‐time fitness, caring parents are, therefore, expected to adjust their reproductive investment to current environmental conditions and parental capacities. The latter is thought to be signaled via ornamental traits of the bearer. We here investigated whether pre‐ and/or posthatching investment of blue tit (Cyanistes caeruleus) parents was related to ornamental plumage traits (UV crown coloration and carotenoid‐based plumage coloration) expressed by either the individual itself (i.e. “good parent hypothesis”) or its partner (i.e. “differential allocation hypothesis”). Our results show that neither prehatching (that is clutch size and offspring begging intensity) nor posthatching parental investment (provisioning rate, offspring body condition at fledging) was related to an individual's UV crown coloration or to that of its partner. Similar observations were made for carotenoid‐based plumage coloration, except for a consistent positive relationship between offspring begging intensity and maternal carotenoid‐based plumage coloration. This sex‐specific pattern likely reflects a maternal effect mediated via maternally derived egg substances, given that the relationship persisted when offspring were cross‐fostered. This suggests that females adjust their offspring's phenotype toward own phenotype, which may facilitate in particular mother‐offspring co‐adaptation. Overall, our results contribute to the current state of evidence that structural or pigment‐based plumage coloration of blue tits are inconsistently correlated with central life‐history traits.     

Maternal yolk androgens stimulate territorial behaviour in black-headed gull chicks

Avian eggs contain substantial amounts of maternal androgens. The concentrations of these yolk androgens are affected by the maternal environment, such as the level of social competition, parasite exposure or food conditions. Since yolk androgens have been shown to affect a wide array of offspring traits, they may adjust the chicks to the expected post-hatching environment, but experimental evidence is still scarce. We investigate in colonial breeding black-headed gulls whether high concentrations of yolk androgens, such as those found in environments with high numbers of social interactions, facilitate aggressiveness and territorial behaviour of the chicks. Black-headed gulls are highly suitable for this, as the semi-precocial chicks defend the natal territory and food against intruders. We manipulated yolk androgen concentrations and investigated their role in both within-nest and between-nest aggression. We found that chicks hatching from androgen-treated eggs defended the natal territory more often than their nest mates from control eggs, without increasing sibling aggression. This suggests that variation in yolk androgen concentrations in relation to the social environment of the mother may indeed allow adjustment of the offspring''s behaviour to the expected frequency of territorial interactions with conspecifics post-hatching.     

Lesser double‐collared sunbirds Nectarinia chalybea do not compensate for hatching asynchrony by adjusting egg mass or yolk androgens

Substantial amounts of maternal androgens are found in birds’ eggs and have been shown to benefit offspring development. Within‐clutch patterns of increasing androgen concentrations over the laying sequence are often hypothesized to compensate for the negative effects of hatching asynchrony. However, detrimental effects to offspring fitness of exposure to high yolk androgen levels have also been demonstrated. This suggests that mothers should forego these costs to their offspring when the need for compensation for hatching asynchrony is low or when alternative compensatory strategies, e.g. in terms of increasing egg mass, are available. Here we show that in the south‐temperate lesser double‐collared sunbird Nectarinia chalybea, a species with hatching asynchrony but also with high survival of last‐hatched chicks, mothers do not deposit resources differentially in terms of either yolk androgen concentration or egg mass across the laying sequence. We discuss to what extend this challenges the original explanation of within‐clutch variation in these egg parameters and offer some explanation for their between‐clutch variation which was related to female body mass.     

Does testosterone mediate the trade-off between nestling begging and growth in the canary (Serinus canaria)?

Nestling birds solicit food from their parents with vigorous begging displays, involving posturing, jostling and calling. In some species, such as canaries, begging is especially costly because it causes a trade off against nestling growth. Fitness costs of begging like this are predicted by evolutionary theory because they function to resolve conflicts of interest within the family over the provision of parental investment. However, the mechanism that links these costs with nestling behaviour remains unclear. In the present study, we determine if the relationships between nestling androgen levels, nestling begging intensities and nestling growth rates are consistent with the hypothesis that testosterone is responsible for the trade-off between begging and growth. We test this idea with a correlational study, using fecal androgens as a non-invasive method for assaying nestling androgen levels. Our results show that fecal androgen levels are positively correlated with nestling begging intensity, and reveal marked family differences in each trait. Furthermore, changes in fecal androgen levels between 5 and 8 days after hatching are positively associated with changes in nestling begging intensity, and negatively associated with nestling growth during this time. Although these correlational results support our predictions, we suggest that that experimental manipulations are now required to test the direct or indirect role of testosterone in mediating the trade-off between begging and growth.     

Flexible communication within bird families—The consequences of behavioral plasticity for parent–offspring coadaptation

Offspring are selected to demand more resources than what is optimal for their parents to provide, which results in a complex and dynamic interplay during parental care. Parent–offspring communication often involves conspicuous begging by the offspring which triggers a parental response, typically the transfer of food. So begging and parental provisioning reciprocally influence each other and are therefore expected to coevolve. There is indeed empirical evidence for covariation of offspring begging and parental provisioning at the phenotypic level. However, whether this reflects genetic correlations of mean levels of behaviors or a covariation of the slopes of offspring demand and parental supply functions (= behavioral plasticity) is not known. The latter has gone rather unnoticed—despite the obvious dynamics of parent–offspring communication. In this study, we measured parental provisioning and begging behavior at two different hunger levels using canaries (Serinus canaria) as a model species. This enabled us to simultaneously study the plastic responses of the parents and the offspring to changes in offspring need. We first tested whether parent and offspring behaviors covary phenotypically. Then, using a covariance partitioning approach, we estimated whether the covariance predominantly occurred at a between‐nest level (i.e., indicating a fixed strategy) or at a within‐nest level (i.e., reflecting a flexible strategy). We found positive phenotypic covariation of offspring begging and parental provisioning, confirming previous evidence. Yet, this phenotypic covariation was mainly driven by a covariance at the within‐nest level. That is parental and offspring behaviors covary because of a plastic behavioral coadjustment, indicating that behavioral plasticity could be a main driver of parent–offspring coadaptation.     

The quantitative genetic basis of offspring solicitation and parental response in a passerine bird with biparental care.

The coevolution of parental investment and offspring solicitation is driven by partly different evolutionary interests of genes expressed in parents and their offspring. In species with biparental care, the outcome of this conflict may be influenced by the sexual conflict over parental investment. Models for the resolution of such family conflicts have made so far untested assumptions about genetic variation and covariation in the parental resource provisioning response and the level of offspring solicitation. Using a combination of cross-fostering and begging playback experiments, we show that, in the great tit (Parus major), (i) the begging call intensity of nestlings depends on their common origin, suggesting genetic variation for this begging display, (ii) only mothers respond to begging calls by increased food provisioning, and (iii) the size of the parental response is positively related to the begging call intensity of nestlings in the maternal but not paternal line. This study indicates that genetic covariation, its differential expression in the maternal and paternal lines and/or early environmental and parental effects need to be taken into account when predicting the phenotypic outcome of the conflict over investment between genes expressed in each parent and the offspring.     

Parent-offspring interactions in food provisioning of Manx shearwaters: implications for nestling obesity

Procellariiform seabirds such as the Manx shearwater Puffinus puffinus, rear only one chick at a time but may breed many times in their lives; parents should thus limit food delivery to the chick in keeping with the balance between current and future reproductive output. Yet procellariiform chicks accumulate large quantities of lipid, which may provide a buffer against pronounced and unpredictable variation in food provisioning, resulting in part from an inability of parents to regulate food supply to the nest. We switched chicks between nests to examine the roles of parents and offspring in controlling food delivery. The serial autocorrelation in age-specific body masses for unmanipulated chicks decreased from 0.61 (P< 0.01) to 0.35 (NS) over a period of 15 days and remained nonsignificant thereafter. By contrast, the serial autocorrelation for switched chicks increased from 0.64 (P< 0.01) to 0.83 (P< 0.001) and the serial cross-correlation rose from 0.23 (NS) to 0.50 (P< 0.05). These results supported both chick determination and parental determination models of food provisioning, indicating that chicks conveyed information about their nutritional status, which parents acted upon by adjusting their rate of food delivery. We discuss these results in relation to the optimization of nestling lipid reserves and parental foraging effort. We suggest that information conveyed by the chick's begging intensity serves to reduce the provisioning rate to well-fed chicks, but parents cannot or do not increase food provisioning to poorly fed chicks. Such adjustment of food provisioning does not refute the hypothesis that nestling obesity provides a buffer against highly variable food delivery. Copyright 1999 The Association for the Study of Animal Behaviour.     

Experimental evidence for offspring learning in parent-offspring communication

The offspring of birds and mammals solicit food from their parents by a combination of movements and vocalizations that have come to be known collectively as 'begging'. Recently, begging has most often been viewed as an honest signal of offspring need. Yet, if offspring learn to adjust their begging efforts to the level that rewards them most, begging intensities may also reflect offsprings' past experience rather than their precise current needs. Here we show that bird nestlings with equal levels of need can learn to beg at remarkably different levels. These experiments with hand-raised house sparrows (Passer domesticus) indicated that chicks learn to modify begging levels within a few hours. Moreover, we found that the begging postures of hungry chicks in natural nests are correlated with the average postures that had previously yielded them parental feedings. Such learning challenges parental ability to assess offspring needs and may require that, in response, parents somehow filter out learned differences in offspring signals.     

Elevated yolk androgen levels benefit offspring development in a between-clutch context

The field of androgen deposition in avian eggs and its consequencesfor offspring development has received a lot of attention inrecent research. However, although variation within clutchesin yolk androgens is relatively well understood, the adaptivesignificance of patterns of variation between clutches remainsrather unclear. Furthermore, it has been hypothesized that yolkandrogens act as a means of an adaptive maternal effect to adjustoffspring to a given posthatching environment. Thus, the consequencesof maternal yolk androgens for offspring development are likelyto depend on the specific environment of a given brood. We experimentallymanipulated yolk androgen concentrations in spotless starlingeggs, using a between-brood design in which full broods weremanipulated applying either an androgen or sham treatment, inorder to test the effects of between-brood variation in yolkandrogen levels. We also included in the analysis several femalecharacters that have been shown to affect androgen deposition.Androgen-chicks tended to gain more mass, a similar effect tothat shown in previous studies where direct competition betweenchicks belonging to different treatments was allowed, but didnot gain a survival benefit. Androgen-chicks had wider beakflanges, an effect that has not been described previously andcould play an important role in food acquisition. In addition,androgen-chicks had higher endogenous plasma levels of androgens,which could induce higher begging intensity. We hypothesizethat these effects are an important mechanistic link for ourunderstanding of how yolk androgens exert their effects on offspringdevelopment after hatching. Contrary to our predictions, wefound no evidence that the observed effects depended on theenvironment under which it was investigated. Because high yolkandrogen levels seem to be beneficial for nestlings in thisspecies, we hypothesize that yolk hormone deposition could becostly for females or alternatively that potential negativeeffects later in adulthood may constrain maternal hormone allocation.     

Yolk androgens do not appear to mediate sexual conflict over parental investment in the collared flycatcher Ficedula albicollis

Males and females are in conflict over parental care, as it would be favourable for one parent to shift labour to the other. Yolk hormones may offer a mechanism through which female birds could influence offspring traits in ways that increase the relative investment by the male. We studied the role of yolk androgens in mediating sexual conflict over parental care in the collared flycatcher (Ficedula albicollis). In a cross-fostering experiment, the male's proportion of total feeding visits increased with increasing androgen levels in the foster eggs. This could suggest that sexual conflict over parental care may be influenced by the female's differential allocation of yolk androgens or a maternal effect associated with yolk androgens. However, when we experimentally elevated yolk androgen levels, male feeding rates did not differ between control and androgen-manipulated nests. This suggests that other egg components correlated with yolk androgen levels, rather than yolk androgen levels per se, may influence male parental effort. In conclusion, yolk androgens per se do not appear to mediate sexual conflict over parental investment in the collared flycatcher.