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Mothers who produce multiple offspring within one reproductive attempt often allocate resources differentially; some maternally derived substances are preferentially allocated to last-produced offspring and others to first-produced offspring. The combined effect of these different allocation regimes on the overall fitness of offspring produced early or late in the sequence is not well understood, partly because production order is often coupled with birth order, making it difficult-to-separate effects of pre-natal maternal allocation from those of post-natal social environments. In addition, very little is known about the influence of laying order on fitness in later life. In this study, we used a semi-natural captive colony of black-headed gulls to test whether an offspring's position in the laying order affected its early-life survival and later-life reproductive success, independent of its hatching order. Later-laid eggs were less likely to hatch, but among those that did, survival to adulthood was greater than that of first-laid eggs. In adulthood, the laying order of females did not affect their likelihood of breeding in the colony, but male offspring hatched from last-laid eggs were significantly less likely to gain a breeding position than earlier-laid males. In contrast, later-laid female parents hatched lower proportions of their clutches than first-laid females, but hatching success was unrelated to the laying order of male parents. Our results indicate that gull mothers induce complex and sex-specific effects on both the early survival of their offspring and on long-term reproductive success through laying order effects among eggs of the same breeding attempt.

Title: "Pathway for personality development: juvenile red knots vary more in diet and exploratory behaviour than adults" Authors: Selin Ersoy, Christine E. Beardsworth, Elif Duran, Marcel T.J. van der Meer, Theunis Piersma, Ton G. G. Groothuis, Allert I. Bijleveld GENERAL INFORMATION Titles of the Dataset: Juv-Adult_Expl_Allyears (Exploration speed (mean speed cm/sec) in mobile arena), isotope data (plasma and red blood cell) Date of the data collection: Catching between 14 August 2018 - 19 October 2021 3. Geographic location of data collection: Western Dutch Wadden Sea (53°15'N, 5°15'E) 4. Corresponding author information: Selin Ersoy, NIOZ Royal Netherlands Institute for Sea Research, Department of Coastal Systems, PO Box 59, 1790 AB Den Burg, Texel, The Netherlands, selin.ersoy@gmail.com METHODS Isotope data: Red blood cell and plasma d13C and d15N isotopes collected from the blood samples of red knots. Exploration speed: Movement tracjectories were collected from the recordings of the top camera during experiments in the mobile arena. We used the distance between estimated positions to calculate speed. Errors in the positioning algorithm were filtered by excluding speeds higher than 200 cm/s. An individual’s exploration speed was calculated as the average speed during each 20 min trial. The dataset includes two repeats of exploration speed. DATA & FILE OVERVIEW Juv-Adult_Expl_Allyears : Exploration speed repeated measures from Juvenile and Adult red knots Column 1: FB: Unique metal ring number of red knots. Ring were put on the legs Column 2: Exp: Repeat number of the exploration assay trials Column 3: Age: Adult (3) or juvenile (1) age Column 4: Mean: Exploratory score of individual measured in mobile arena (mean speed cm/sec) measured through automated tracking Column 5: Year: Year of the testing Isotope : Red blood cell and plasma d13C and d15N isotopes collected from the blood samples of juvenile and adult red knots Column 1: RingNr: Unique metal ring number of red knots. Ring were put on the legs Column 2: Type: Blood type plasma or red blood cell Column 3: Repeat: If there is a repeated samples of an individual Column 4: d15N: Nitrogen 15 Isotope value Column 5: d13C: Carbon 13 Isotope value Column 6: ProcessDate:Process date of blood samples Column 7: Comments: important commments about samples Column 8: AGE: Adult (3) or juvenile (1) age Column 9: CatchDate: Cathing date of the individuals R file includes code for all the statistics and visualization of the data.&nbsp

Maternal effects, or the influence of maternal environment and phenotype on offspring phenotype, may allow mothers to fine‐tune their offspring's developmental trajectory and resulting phenotype sometimes long after the offspring has reached independence. However, maternal effects on offspring phenotype do not evolve in isolation, but rather within the context of a family unit, where the separate and often conflicting evolutionary interests of mothers, fathers and offspring are all at play. While intrafamilial conflicts are routinely invoked to explain other components of reproductive strategy, remarkably little is known about how intrafamilial conflicts influence maternal effects. We argue that much of the considerable variation in the relationship between maternally derived hormones, nutrients and other compounds and the resulting offspring phenotype might be explained by the presence of conflicting selection pressures on different family members. In this review, we examine the existing literature on maternal hormone allocation as a case study for maternal effects more broadly, and explore new hypotheses that arise when we consider current findings within a framework that explicitly incorporates the different evolutionary interests of the mother, her offspring and other family members. Specifically, we hypothesise that the relationship between maternal hormone allocation and offspring phenotype depends on a mother's ability to manipulate the signals she sends to offspring, the ability of family members to be plastic in their response to those signals and the capacity for the phenotypes and strategies of various family members to interact and influence one another on both behavioural and evolutionary timescales. We also provide suggestions for experimental, comparative and theoretical work that may be instrumental in testing these hypotheses. In particular, we highlight that manipulating the level of information available to different family members may reveal important insights into when and to what extent maternal hormones influence offspring development. We conclude that the evolution of maternal hormone allocation is likely to be shaped by the conflicting fitness optima of mothers, fathers and offspring, and that the outcome of this conflict depends on the relative balance of power between family members. Extending our hypotheses to incorporate interactions between family members, as well as more complex social groups and a wider range of taxa, may provide exciting new developments in the fields of endocrinology and maternal effects.

Hormones transferred from mothers to their offspring are thought to be a tool for mothers to prepare their progeny for expected environmental conditions, thus increasing fitness. Thyroid hormones (THs) are crucial across vertebrates for embryonic and postnatal development and metabolism. Yet yolk THs have mostly been ignored in the context of hormone-mediated maternal effects. In addition, the few studies on maternal THs have yielded contrasting results that could be attributed to either species or environmental differences. In this study, we experimentally elevated yolk THs (within the natural range) in a wild population of a migratory passerine, the European pied flycatcher (Ficedula hypoleuca), and assessed the effects on hatching success, nestling survival, growth, and oxidative status (lipid peroxidation, antioxidant enzyme activity, and oxidative balance). We also sought to compare our results with those of a closely related species, the collared flycatcher (Ficedula albicolis), that has strong ecological and life-history similarities with our species. We found no effects of yolk THs on any of the responses measured. We could detect only a weak trend on growth: elevated yolk THs tended to increase growth during the second week after hatching. Our results contradict the findings of previous studies, including those of the collared flycatcher. However, differences in fledging success and nestling growth between both species in the same year suggest a context-dependent influence of the treatment. This study should stimulate more research on maternal effects mediated by THs and their potential context-dependent effects.

Maternal hormones constitute a key signalling pathway for mothers to shape offspring phenotype and fitness. Thyroid hormones (THs; triiodothyronine, T3; and thyroxine, T4) are metabolic hormones known to play crucial roles in embryonic development and survival in all vertebrates. During early developmental stages, embryos exclusively rely on exposure to maternal THs, and maternal hypothyroidism can cause severe embryonic maldevelopment. The TH molecule includes iodine, an element that cannot be synthesised by the organism. Therefore, TH production may become costly when environmental iodine availability is low. This may yield a trade-off for breeding females between allocating the hormones to self or to their eggs, potentially to the extent that it even influences the number of laid eggs. In this study, we investigated whether low dietary iodine may limit TH production and transfer to the eggs in a captive population of rock pigeons (Columba livia). We provided breeding females with an iodine-restricted (I−) diet or iodine-supplemented (I+) diet and measured the resulting circulating and yolk iodine and TH concentrations and the number of eggs laid. Our iodine-restricted diet successfully decreased both circulating and yolk iodine concentrations compared with the supplemented diet, but not circulating or yolk THs. This indicates that mothers may not be able to independently regulate hormone exposure for self and their embryos. However, egg production was clearly reduced in the I− group, with fewer females laying eggs. This result shows that restricted availability of iodine does induce a cost in terms of egg production. Whether females reduced egg production to preserve THs for themselves or to prevent embryos from exposure to low iodine and/or THs is as yet unclear., Summary: Restricted dietary iodine in captive rock pigeons reduces egg production in some females, thus inducing a trade-off between offspring quality and offspring quantity.

1. In vertebrates, thyroid hormones (THs) play an important role in the regulation of growth, development, metabolism, photoperiodic responses and migration. Maternally transferred THs are important for normal early-phase embryonic development when embryos are not able to produce endogenous THs. Previous studies have shown that variation in maternal THs within the physiological range can influence offspring phenotype. 2. Given the essential functions of maternal THs in development and metabolism, THs may be a mediator of life-history variation across species. 3. We tested the hypothesis that differences in life histories are associated with differences in maternal TH transfer across species. Using birds as a model, we specifically tested whether maternally transferred yolk THs co-vary with migratory status, developmental mode, and traits related to pace-of-life (e.g. basal metabolic rate, maximum lifespan). 4. We collected un-incubated eggs (n = 1-21 eggs per species, median = 7) from 34 wild and captive bird species across 17 families and 6 orders to measure yolk THs (both triiodothyronine, T3 and thyroxine, T4), compiled life-history trait data from the literature, and used Bayesian phylogenetic mixed models to test our hypotheses. 5. Our models indicated that both concentrations and total amounts of the two main forms of THs (T3 and T4) were higher in the eggs of migratory species compared to resident species, and total amounts were higher in the eggs of precocial species, which have longer prenatal developmental periods, than in those of altricial species. However, maternal yolk THs did not show clear associations with pace-of-life related traits, such as fecundity, basal metabolic rate, or maximum lifespan. 6. We quantified interspecific variation in maternal yolk THs in birds and our findings suggest higher maternal TH transfer is associated with the precocial mode of development and migratory status. Whether maternal THs represent a part of the mechanism underlying the evolution of precocial development and migration or a consequence of such life histories is currently unclear. We therefore encourage further studies to explore the physiological mechanisms and evolutionary processes underlying these patterns.

Brain lateralization is a fundamental aspect of the organization of brain and behavior in the animal kingdom, begging the question about its Darwinian function. We tested the possibility that lateralization enhances cognitive performance in single- and dual-tasks. Previous studies reported mixed results on this topic and only a handful of studies have measured functional brain lateralization and performance independently and simultaneously. We therefore examined a possible positive effect of the strength and direction of lateralization on two demanding cognitive tasks: A visuospatial task (mental rotation MR), and a language task (word generation WG), executed either as a singletask or as dual-task. Participants (n = 72) performed these tasks while their single-task brain lateralization was assessed with functional Transcranial Doppler for both tasks. From these measurements we determined strength and direction of lateralization for both tasks and the individual pattern of lateralization (contralateral or ipsilateral) was derived. These factors, along with sex, were used in a GLM analysis to determine if they predicted the respective performance measure of the tasks. We found that for MR there was a significant medium effect of direction of lateralization on performance with better performance in left-lateralized (atypical) participants (partial eta squared 0.061; p = .039). After correction for outliers, there was a significant effect for strength (p = .049). For the dual-task, there was a significant positive medium effect of strength of lateralization on performance (partial eta squared 0.062; p = .038, respectively) No other association between direction or strength in either tests were found. We conclude that there is no evidence for hemispheric crowding, and that strength of lateralization may be a factor that contributes to the evolutionary selection of functional brain lateralization. Pattern of lateralization does not, explaining the large inter-individual variation in these traits.

A remarkable feature of human handedness at the population level is specialization of the hands, the right hand performing usually better than the left. This specialization might have an evolutionary advantage, because it provides the individual and population with a wider range of skill. We therefore investigated the relationships between hand skill asymmetry and potential markers of Darwinian fitness that have been hypothesized to explain the bias in hand preference: creativity, aggression and developmental and health problems. Over twenty thousand participants (56% left-handers) completed an online survey, including a finger-tapping task to measure hand skill asymmetry. Left-skilled individuals were overall more aggressive than right-skilled individuals and rated themselves as more artistically creative. However, when assessed with a questionnaire, they were less creative on problem solving and equally artistically creative compared to right-skilled individuals, who reported more health problems. Conclusion: we found some evidence for current selection on the direction of lateralization of hand skill although the effect sizes were rather low. Strength of lateralization of hand skill showed only a few associations with fitness proxies. We suggest that Darwinian selection on hand preference (Zickert, Feen, van der, Geuze, & Groothuis, 2018. Fitness costs and benefits associated with hand preference in humans: A large internet study in a Dutch sample. Evolution and Human Behavior, 39, 235-248) and hand skill asymmetry (present study) may be attenuated in modern society.

Ocklenburg et al. (2020, Laterality 2020: Entering the next decade. Laterality) provided the field of laterality research with a stimulating research perspective for the coming decade, based on the current state of the art in both animal and human laterality research. Although this is paper takes many different approaches of laterality into account, we emphasize that the eco-evolutionary approach needs more attention. This concerns the question why organisms are lateralized in the first place, in other words, how does lateralization enhance the Darwinian fitness of the individual. We argue that laterality can be distinguished along four dimensions, and that each of them requires different ultimate explanations. Studying these functional and evolutionary explanations requires the development of ecologically relevant tests, adapted to the species at hand. It also requires experimental manipulation of laterality, testing the effect in (semi)-natural conditions on fitness parameters. Tools for such manipulation of laterality urgently require a better understanding of the developmental plasticity of lateralization, extending the field of evo-devo to that of eco-evo-devo. We also warn against seeing the minority in the distribution of direction or strength of lateralization as being a pathology as such minorities in biology can often be explained as adaptations by natural frequency dependent selection.

Competing theories have posited roles for foetal androgen exposure in the development of human handedness. However, due to practical and ethical considerations, few studies have used hormonal measures to examine this possibility. The current paper reviews this literature and reveals a generally inconsistent pattern of results. We also present data from a longitudinal study of prenatal sex hormone exposure and subsequent handedness. More specifically, we examine correlations between testosterone and estradiol measured from second trimester amniotic fluid and hand preference (Dutch language version of the Edinburgh Handedness Inventory) and hand skill asymmetry (pegboard task) measured at 15 years of age. Prenatal sex hormone exposure was not associated with the direction of hand preference in either males or females. However, in females, high levels of prenatal testosterone were associated with weaker lateralisation of hand skill, and high levels of prenatal estradiol were associated with weaker hand preference. In addition, high levels of prenatal testosterone were associated with increased task duration (i.e., slow hand speed) for the right and left hands of males. The pattern of results observed here is not entirely consistent with any of the main theories linking sex hormones with handedness, suggesting that an association between these variables may be more complex than initially thought.

Maternal hormones constitute a key signalling pathway for mothers to shape offspring phenotype and fitness. Thyroid hormones (THs; triiodothyronine, T3 and thyroxine, T4) are metabolic hormones known to play crucial roles in embryonic development and survival in all vertebrates. During early developmental stages, embryos exclusively rely on the exposure to maternal THs, and maternal hypothyroidism can cause severe embryonic maldevelopment. The TH molecule includes iodine, an element that cannot be synthesised by the organism. Therefore, TH production may become costly when environmental iodine availability is low. This may yield a trade-off for breeding females between allocating the hormones to self or to their eggs, potentially to the extent that it even influences the number of laid eggs. In this study, we investigated whether low dietary iodine may limit TH production and transfer to the eggs in a captive population of Rock pigeons (Columba livia). We provided breeding females with an iodine-restricted (I- diet) or iodine-supplemented diet (I+ diet) and measured the resulting circulating and yolk iodine and TH concentrations and the number of eggs laid. Our iodine-restricted diet successfully decreased both circulating and yolk iodine concentrations compared to the supplemented diet, but not circulating or yolk THs. This indicates that mothers may not be able to independently regulate hormone exposure for self and their embryos. However, egg production was clearly reduced in the I- group, with fewer females laying eggs. This result shows that restricted availability of iodine does induce a cost in terms of egg production. Whether females reduced egg production to preserve THs for themselves or to prevent embryos from exposure to low iodine and/or THs is as yet unclear.

Many plants and animals adaptively downsize the number of already-produced propagules if resources become insufficient to raise all of them. In birds, mothers often induce hatching asynchrony by incubating first eggs before last eggs are laid, creating an age/size hierarchy within broods which selectively eliminates the smallest chicks in poor food conditions. However, mothers also deposit more testosterone into late-laid eggs, which boosts competitive abilities of younger chicks, counteracts the competitive hierarchy, and ostensibly creates a paradox. Since testosterone also carries costs, we hypothesized that benefits of maternally deposited testosterone outweigh its costs in good food conditions, but that testosterone has a net detrimental effect in poor food conditions. We found experimental evidence that elevated maternal testosterone in the egg caused higher chick mortality in poor food conditions but better chick growth in good food conditions. These context-dependent effects resolve the paradox, suggesting co-evolution of two maternal effects, and explain inconsistent results of egg hormone manipulations in the literature.

Hormones transferred from mothers to their offspring are considered a maternal tool to prepare progeny for expected environmental conditions, increasing maternal fitness. To flexibly influence offspring, mothers should be able to transmit the hormonal signals independent of their own hormonal status. However, the ability to regulate hormone transfer to the next generation is under debate. We studied the transfer of thyroid hormones (THs) to eggs in a bird model. We elevated thyroxine (T4, the prohormone for the biological active triiodothyronine, T3) during egg-laying using T4 implants on females of a wild population of pied flycatchers (Ficedula hypoleuca), and measured plasma and yolk T4 and T3 as a response. To our knowledge, studies that manipulated a prohormone and measured the change in its active metabolites have rarely been conducted. We found an increase in plasma and yolk T4 and no change in T3 concentrations leading to a similar decrease in yolk T3/T4 ratio in response to the T4 treatment in plasma and yolk. This suggests that mothers are able to regulate the conversion of T4 in T3 to avoid potential costs of elevated exposure to the active hormone to herself and to her progeny. Finally, contrary to our predictions, we found no evidence of regulatory mechanisms at the follicle level, which is essential for independent regulation of yolk hormone transfer.Summary statementThyroid hormones have been overlooked in the context of hormone-mediated maternal effects. We found that mothers may regulate yolk thyroid hormone transfer by regulating the conversion of the active form of the hormone.

Maternal thyroid hormones (THs) are known to be crucial in embryonic development in humans, but their influence on other, especially wild, animals remains poorly understood. So far, the studies that experimentally investigated the consequences of maternal THs focused on short-term effects, while early organisational effects with long-term consequences, as shown for other prenatal hormones, could also be expected. In this study, we aimed at investigating both the short- and long-term effects of prenatal THs in a bird species, the Japanese quail Coturnix japonica. We experimentally elevated yolk TH content (the prohormone T4, and its active metabolite T3, as well as a combination of both hormones). We analysed hatching success, embryonic development, offspring growth and oxidative stress as well as their potential organisational effects on reproduction, moult and oxidative stress in adulthood. We found that eggs injected with T4 had a higher hatching success compared with control eggs, suggesting conversion of T4 into T3 by the embryo. We detected no evidence for other short-term or long-term effects of yolk THs. These results suggest that yolk THs are important in the embryonic stage of precocial birds, but other short- and long-term consequences remain unclear. Research on maternal THs will greatly benefit from studies investigating how embryos use and respond to this maternal signalling. Long-term studies on prenatal THs in other taxa in the wild are needed for a better understanding of this hormone-mediated maternal pathway.

After decades of research, the influence of prenatal testosterone on brain lateralization is still elusive, whereas the influence of pubertal testosterone on functional brain lateralization has not been investigated, although there is increasing evidence that testosterone affects the brain in puberty. We performed a longitudinal study, investigating the relationship between prenatal testosterone concentrations in amniotic fluid, pubertal testosterone concentrations in saliva, and brain lateralization (measured with functional Transcranial Doppler ultrasonography (fTCD)) of the Mental Rotation, Chimeric Faces and Word Generation tasks. Thirty boys and 30 girls participated in this study at the age of 15 years. For boys, we found a significant interaction effect between prenatal and pubertal testosterone on lateralization of Mental Rotation and Chimeric Faces. In the boys with low prenatal testosterone levels, pubertal testosterone was positively related to the strength of lateralization in the right hemisphere, while in the boys with high prenatal testosterone levels, pubertal testosterone was negatively related to the strength of lateralization. For Word Generation, pubertal testosterone was negatively related to the strength of lateralization in the left hemisphere in boys. For girls, we did not find any significant effects, possibly because their pubertal testosterone levels were in many cases below quantification limit. To conclude, prenatal and pubertal testosterone affect lateralization in a task-specific way. Our findings cannot be explained by simple models of prenatal testosterone affecting brain lateralization in a similar way for all tasks. We discuss alternative models involving age dependent effects of testosterone, with a role for androgen receptor distribution and efficiency.

Lateral preference is a widespread organizational principle in human and nonhuman animals. In humans, the most apparent lateralized trait (handedness) is unique in the animal kingdom because of a very pronounced bias towards right-handedness on a population level. In this study, based on previous experiments, we test the hypothesis that this bias was-among other factors-shaped by evolution through the facilitation of social learning. We exposed 134 subjects to footage of right- or left-handed knot making and analyzed whether concordant handedness between instructor and student facilitated quicker and more successful imitation. We used a set of nautical knots of different difficulty levels in order to test whether the potential effect of concordance became stronger with increasing knot difficulty. For all three performance measures (time until correct completion, number of attempts needed and correct imitation), we found hand congruency and difficulty level to be significant predictors but not the interaction of the two. We conclude that concordance of handedness between teacher and student of a motor skill enhances the speed and accuracy of imitation, which may have been a beneficial trait for selection to act upon, thereby shaping the human population bias in handedness.

Vertebrates evolved in concert with bacteria and have developed essential mutualistic relationships. Gut bacteria are vital for the postnatal development of most organs and the immune and metabolic systems and may likewise play a role during prenatal development. Prenatal transfer of gut bacteria is shown in four mammalian species, including humans. For the 92% of the vertebrates that are oviparous, prenatal transfer is debated, but it has been demonstrated in domestic chicken. We hypothesize that also non-domestic birds can prenatally transmit gut bacteria. We investigated this in medium-sized Rock pigeon (Columba livia), ensuring neonates producing fair-sized first faeces. The first faeces of 21 neonate rock pigeons hatched in an incubator, contained a microbiome (bacterial community) the composition of which resembled the cloacal microbiome of females sampled from the same population (N = 5) as indicated by multiple shared phyla, orders, families, and genera. Neonates and females shared 16.1% of the total number of OTUs present (2881), and neonates shared 45.5% of their core microbiome with females. In contrast, the five females shared only 0.3% of the 1030 female OTUs present. These findings suggest that prenatal gut bacterial transfer may occur in birds. Our results support the hypothesis that gut bacteria may be important for prenatal development and present a heritability pathway of gut bacteria in vertebrates.

Several studies show that avian females prefer males based on their secondary sexual ornaments and dominance status. We tested in red junglefowl (Goth's gallus) how comb size affected the result of fighting and how the dominance status related to testosterone concentrations in their circulation and ejaculates. We subsequently tested how social status was related to female reproductive investment, including yolk hormone transfer. We found that after a fight 1) winners increased plasma T and decreased ejaculates T whereas losers' T remained unchanged, and 2) plasma T of winners was higher but ejaculates T was lower than those of losers. We argued those are consistent with the different reproductive strategies of dominant and subordinate males. Furthermore, in line with offspring sex-dependent growth patterns females transferred significantly more androstenedione to female than male embryos when mated with winners, while doing the opposite when mated with losers. We concluded therefore that female reproductive investment was affected by both partner quality and embryo sex. The results indicate that male quality influences sex-specific maternal investment, which could be mediated by ejaculate testosterone concentration.

Exposure of yolk androgens can positively stimulate chick growth and competitive ability, but may negatively affect immunity. It has been hypothesized that only chicks from immunologically superior fathers can bear the cost of prenatal exposure to high androgen levels. To test this hypothesis, we paired roosters from two selection lines, one up- and one down-selected for natural antibodies (NAbs), with hens from a control line. We measured yolk testosterone and androstenedione levels, and we injected the treatment group of eggs of each female with testosterone suspended in sesame oil and the control group with sesame oil only. We then measured hatching success and growth, and characterized the humoral and cellular immune responses using three different challenges: a phyto-hemagglutinin, a lipopolysaccharide and a sheep red blood cell challenge. We found that the hatching success, body mass, initial levels of natural antibodies and the chicks’ immunological responses to the three different challenges and development were affected neither by paternal immunocompetence nor by treatment. These results do not support the hypothesis that chicks from low-NAb line fathers are more sensitive to testosterone exposure during embryonic development than chicks from high-NAb line fathers., Summary: Our study shows that there were no effects of paternal natural antibody line, increased embryonic testosterone exposure or the interaction of both on immunocompetence of chicks.

Maternal thyroid hormones (THs) are known to be crucial in embryonic development in humans, but their influence on other, especially wild, animals remains poorly understood. So far, the studies that experimentally investigated the consequences of maternal THs focused on short-term effects, while long-term organisational effects, as shown for other prenatal hormones, could also be expected. In this study, we aimed at investigating both the short- and long-term effects of prenatal THs in a bird species, the Japanese quail Coturnix japonica . We experimentally elevated yolk TH content (the prohormone T 4 , and its active metabolite T 3 , as well as a combination of both hormone) We analysed hatching success, embryonic development, offspring growth and oxidative stress as well as their potential organisational effects on reproduction, moult, and oxidative stress in adulthood. We found that eggs injected with both hormones had a higher hatching success compared with control eggs, suggesting conversion of T 4 into T 3 by the embryo. We detected no other short-term or organisational effects of yolk THs. Unfortunately, sex-specific responses could not be properly tested due to low sample sizes. These results suggest that yolk thyroid hormones are important in the embryonic stage of precocial birds, but may not have other short and long-term consequences, at least not in captivity. Research on maternal thyroid hormones will greatly benefit from studies investigating how embryos use and respond to this maternal signalling. Long-term studies on prenatal THs in other taxa in the wild are needed for a better understanding of this hormone-mediated maternal pathway. Summary statement Thyroid hormones are important hormones in all vertebrates, although overlooked in the context of maternal effect. We found short-term effects of prenatal THs but no evidence of programming effects.

Exposure of the vertebrate embryo to maternal hormones can have long-lasting effects on its phenotype, which has been studied extensively by experimentally manipulating maternal steroids, mostly androgens, in bird eggs. Yet, there is a severe lack of understanding of how and when these effects are actually mediated, hampering both underlying proximate and ultimate explanations. Here we report a novel finding that the embryo expresses androgen receptor (AR) and estrogen receptor (ERα) mRNA in its extraembryonic membranes (EMs) as early as before its own hormone production starts, suggesting a novel substrate for action of maternal hormones on the offspring. We also report the first experimental evidence for steroid receptor regulation in the avian embryo in response to yolk steroid levels: the level of AR is dependent on yolk androgen levels only in the EMs but not in body tissues, suggesting embryonic adaptation to maternal hormones. The results also solve the problem of uptake of lipophilic steroids from the yolk, why they affect multiple traits, and how they could mediate maternal effects without affecting embryonic sexual differentiation.

Substantial concentrations of testosterone are not only present in a male’s circulation, but also in its ejaculate. Surprisingly, the regulation of ejaculate T and its effects on females and their offspring, potentially a cryptic paternal trait, are not known. We found lower circulating and higher ejaculate T concentrations in subordinate red junglefowl (Gallus gallus gallus) males compared to dominant males, suggestive of an adaptive trade-off in T allocation to circulation and their ejaculate. Subsequently, we artificially inseminated females with either testosterone enriched (TE) or control ejaculates (CE) in a cross-over design. TE females produced heavier eggs than CE females. Offspring growth and tonic immobility were affected in a sex-specific way by TE. TE sons were heavier with shorter TI duration than CE sons, and TE daughters were lighter than CE daughters but daughters did not differ in TI score. However, the chicks competitiveness was not influenced by the TE nor CE. This indicates a previously unknown function of ejaculate testosterone as well as a new form of interaction between a cryptic paternal trait and a maternal effect that may be widespread in the animal kingdom.

Parasites may have strong eco‐evolutionary interactions with their hosts. Consequently, they may contribute to host diversification. The radiation of cichlid fish in Lake Victoria provides a good model to study the role of parasites in the early stages of speciation. We investigated patterns of macroparasite infection in a community of 17 sympatric cichlids from a recent radiation and 2 older species from 2 nonradiating lineages, to explore the opportunity for parasite‐mediated speciation. Host species had different parasite infection profiles, which were only partially explained by ecological factors (diet, water depth). This may indicate that differences in infection are not simply the result of differences in exposure, but that hosts evolved species‐specific resistance, consistent with parasite‐mediated divergent selection. Infection was similar between sampling years, indicating that the direction of parasite‐mediated selection is stable through time. We morphologically identified 6 Cichlidogyrus species, a gill parasite that is considered a good candidate for driving parasite‐mediated speciation, because it is host species‐specific and has radiated elsewhere in Africa. Species composition of Cichlidogyrus infection was similar among the most closely related host species (members of the Lake Victoria radiation), but two more distantly related species (belonging to nonradiating sister lineages) showed distinct infection profiles. This is inconsistent with a role for Cichlidogyrus in the early stages of divergence. To conclude, we find significant interspecific variation in parasite infection profiles, which is temporally consistent. We found no evidence that Cichlidogyrus‐mediated selection contributes to the early stages of speciation. Instead, our findings indicate that species differences in infection accumulate after speciation., Species composition of Cichlidogyrus infection was similar among the closely related cichlid species of the Lake Victoria radiation (in orange), but differed between this lineage and two distantly related species (belonging to nonradiating sister lineages, in blue).

Ecological speciation is facilitated when divergent adaptation has direct effects on selective mating. Divergent sensory adaptation could generate such direct effects, by mediating both ecological performance and mate selection. In aquatic environments, light attenuation creates distinct photic environments, generating divergent selection on visual systems. Consequently, divergent sensory drive has been implicated in the diversification of several fish species. Here, we experimentally test whether divergent visual adaptation explains the divergence of mate preferences in Haplochromine cichlids. Blue and red Pundamilia co‐occur across south‐eastern Lake Victoria. They inhabit different photic conditions and have distinct visual system properties. Previously, we documented that rearing fish under different light conditions influences female preference for blue versus red males. Here, we examine to what extent variation in female mate preference can be explained by variation in visual system properties, testing the causal link between visual perception and preference. We find that our experimental light manipulations influence opsin expression, suggesting a potential role for phenotypic plasticity in optimizing visual performance. However, variation in opsin expression does not explain species differences in female preference. Instead, female preference covaries with allelic variation in the long‐wavelength‐sensitive opsin gene (LWS), when assessed under broad‐spectrum light. Taken together, our study presents evidence for environmental plasticity in opsin expression and confirms the important role of colour perception in shaping female mate preferences in Pundamilia. However, it does not constitute unequivocal evidence for the direct effects of visual adaptation on assortative mating., Pundamilia cichlids exhibit light‐induced opsin expression plasticity, which correlates with female mate preference, but does not explain species differences in preference. Instead, female preference covaries with allelic variation in the LWS opsin in broad‐spectrum light, confirming the important role of colour perception in shaping Pundamilia mate preferences.

Maternal effects can adaptively modulate offspring developmental trajectories in variable but predictable environments. Hormone synthesis is sensitive to environmental factors, and maternal hormones are thus a powerful mechanism to transfer environmental cues to the next generation. Birds have become a key model for the study of hormone-mediated maternal effects because the embryo develops outside the mother's body, facilitating the measurement and manipulation of prenatal hormone exposure. At the same time, birds are excellent models for the integration of both proximate and ultimate approaches, which is key to a better understanding of the evolution of hormone-mediated maternal effects. Over the past two decades, a surge of studies on hormone-mediated maternal effects has revealed an increasing number of discrepancies. In this review, we discuss the role of the environment, genetic factors and social interactions in causing these discrepancies and provide a framework to resolve them. We also explore the largely neglected role of the embryo in modulating the maternal signal, as well as costs and benefits of hormone transfer and expression for the different family members. We conclude by highlighting fruitful avenues for future research that have opened up thanks to new theoretical insights and technical advances in the field. This article is part of the theme issue ‘Developing differences: early-life effects and evolutionary medicine’.

Hormones transferred from mothers to their offspring are thought to be a maternal tool for mothers to prepare their progeny for expected environmental conditions, thus increasing fitness. Thyroid hormones (THs) are crucial across vertebrates for embryonic and post-natal development and metabolism. Nevertheless, the studies that investigated the consequences of maternal hormones have mostly focused on steroid hormones and ignored maternally-derived thyroid hormones. In this study, we experimentally elevated yolk thyroid hormones in a wild population of a migratory passerine, the European Pied flycatcher Ficedula hypoleuca. We injected eggs with a mixture of T4 and T3 within the natural range of the species to assess its effects on hatching success, nestling survival, growth and oxidative status (antioxidant enzyme activity, lipid peroxidation and oxidative balance). We found no effects of yolk THs on egg hatchability or nestling survival. Yolk THs increased nestling growth during the second week post hatching, but this potentially beneficial effect did not incur any costs in terms of oxidative stress. The results should stimulate more research on thyroid hormone mediated maternal effects, further studies into the underlying mechanistic pathways for these effects and how they translate into adulthood and fitness.Summary statementThyroid hormones have been overlooked in the context of hormone-mediated maternal effects. We found that yolk thyroid hormones in a wild bird species increase growth without incurring oxidative stress.

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.

Global warming has substantially changed the environment, but the mechanisms to cope with these changes in animals, including the role of maternal effects, are poorly understood. Maternal effects via hormones deposited in eggs, have important environment-dependent effects on offspring development and fitness: thus females are expected to adjust these hormones to the environment, such as the ambient temperature. Longer-term temperature variation could function as a cue, predicting chick rearing conditions to which yolk hormone levels are adjusted, while short-term temperature variation during egg formation may causally affect hormone transfer to eggs. We studied the effects of ambient temperature on yolk androgens (testosterone and androstenedione) and thyroid hormones (thyroxine and triiodothyronine) in great tits (Parus major) using data from unmanipulated clutches from a wild population and from aviary birds (ad libitum food) exposed to different experimental temperature treatments during five years. Both in the wild and in captivity, longer-term pre-laying ambient temperature was not associated with clutch mean yolk hormone levels, while the way androstenedione and thyroxine levels varied across the laying sequence did associate with pre-laying temperature in the wild. Yolk testosterone levels were positively correlated with short-term temperature (during yolk formation) changes within clutches in both wild and captivity. We also report, for the first time in a wild bird, that yolk thyroxine levels correlated with a key environmental factor: thyroxine levels were negatively correlated with ambient temperature during egg formation. Thus, yolk hormone levels, especially testosterone, seem to be causally affected by ambient temperature. These short-term effects might reflect physiological changes in females with changes in ambient temperature. The adaptive value of the variation with ambient temperatures pre-laying or during egg formation should be studied with hormone manipulations in different thermal environments. (C) 2016 Published by Elsevier Inc.

De linker- en de rechterhemisfeer van de hersenen zijn grotendeels symmetrisch, maar er zijn subtiele verschillen in anatomie en functie. Ze communiceren met elkaar, onder andere via de hersenbalk, het corpus callosum. Algemeen bekend is dat bij de meeste mensen de linkerhemisfeer gespecialiseerd is voor taalverwerking en de rechterhemisfeer voor de verwerking van visuoruimtelijke processen. De term lateralisatie verwijst naar hoe de specialisatie voor neuropsychologische functies is verdeeld over de hersenhelften.Naast lateralisatie op het niveau van de hersenen, onderscheidt men ook lateralisatie op gedragsniveau. Voorbeelden daarvan zijn handvoorkeur, waarbij rechterhandvoorkeur duidt op dominantie van de linkerhemisfeer voor fijne motoriek, en het rechteroorvoordeel bij dichotisch luisteren, wat duidt op een linksdominantie voor taal. Lateralisatie op gedragsniveau wordt vaak gebruikt als indicator voor lateralisatie op het niveau van de hersenen. De vraag is of dat terecht is. In dit artikel bespreken we twee vragen: 1) Wat zijn voordelen van hersenlateralisatie? en 2) Zijn linkshandigen anders gelateraliseerd dan rechtshandigen?, De linker en de rechter hemisfeer van de hersenen zijn grotendeels symmetrisch, maar er zijn subtiele verschillen in anatomie en functie. Ze communiceren met elkaar onder andere via de hersenbalk, het Corpus Callosum. Algemeen bekend is dat bij de meeste mensen de linker hemisfeer gespecialiseerd is voor taalverwerking en de rechter hemisfeer voor verwerking van visuo-ruimtelijke processen. De term lateralisatie verwijst naar hoe de specialisatie voor neuropsychologische functies is verdeeld over de hersenhelften.Naast lateralisatie op het niveau van de hersenen onderscheidt men ook lateralisatie op gedragsniveau. Voorbeelden daarvan zijn handvoorkeur waarbij rechterhandvoorkeur duidt op dominantie van de linker hemisfeer voor fijne motoriek, en het rechteroorvoordeel bij dichotisch luisteren wat duidt op een linker dominantie voor taal (zie bijvoorbeeld [1]). Lateralisatie op gedragsniveau wordt vaak gebruikt als indicator voor lateralisatie op het niveau van de hersenen. De vraag is of dat terecht is. In dit artikel bespreken we twee vragen: 1) wat zijn voordelen van hersenlateralisatie? 2) zijn linkshandigen anders gelateraliseerd dan rechtshandigen?

Maternal reproductive investment can critically influence offspring phenotype, and thus these maternal effects are expected to be under strong natural selection. Knowledge on the extent of heritable variation in the physiological mechanisms underlying maternal effects is however limited. In birds, resource allocation to eggs is a key mechanism for mothers to affect their offspring and different components of the egg may or may not be independently adjusted. We studied the heritability of egg components and their genetic and phenotypic covariation in great tits (Parus major), using captive-bred full siblings of wild origin. Egg mass, testosterone (T) and androstenedione (A4) hormone concentrations showed moderate heritability, in agreement with earlier findings. Interestingly, yolk triiodothyronine hormone (T3), but not its precursor, thyroxine hormone (T4), concentration was heritable. An immune factor, albumen lysozyme, showed moderate heritability, but yolk immunoglobulins (IgY) did not. The genetic correlation estimates were moderate but statistically nonsignificant; a trend for a positive genetic correlation was found between A4 and egg mass, T and lysozyme and IgY and lysozyme, respectively. Interestingly, phenotypic correlations were found only between A4 and T, and T4 and T3, respectively. Given that these egg components are associated with fitness-related traits in the offspring (and mother), and that we show that some components are heritable, it opens the possibility that natural selection may shape the rate and direction of phenotypic change via egg composition.Heredity advance online publication, 6 July 2016; doi:10.1038/hdy.2016.49.

In many animal species, mothers expose their embryos to maternal hormones. This provides a powerful pathway for affecting offspring phenotype and a potential mechanism for adaptive nongenomic inheritance. This has been studied extensively in birds because the embryo develops outside the mother's body, facilitating experimental studies. Some studies have demonstrated that maternal yolk testosterone (T) can have long-lasting effects into the period after fledging. However, these results are based on offspring in the juvenile stage or their first breeding season only. The lack of data further into adulthood limits a better assessment of the fitness consequences of prenatal T. In a series of previous experiments in the rock pigeon, Columba livia, we manipulated yolk T levels using oil-injected eggs as controls. In the present study, we examined the reproductive behaviour of birds from these eggs when they were 1-3 years old and housed together in a large aviary. Our results showed that males from T-injected eggs were less aggressive and females from T-injected eggs laid lighter eggs than controls. Moreover, the experimental birds showed strongly disassortative mating with respect to hormone treatment. There were no indications of treatment effects on the process of sexual differentiation. Our results indicate that the effects of prenatal T do not diminish over age and may last far into adulthood, which should be taken into account in assessing the fitness consequences of maternal hormones. They may also stimulate new physiological and neurobiological studies on the mechanisms underlying such long-lasting effects of prenatal T exposure. (C) 2016 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Several studies show effects of yolk androgens in avian eggs on the phenotype of the offspring. Yolk hormone concentrations decline strongly already in the first few days of incubation. Although early embryonic uptake of yolk androgens is suggested by the presence of radioactivity in the embryo when eggs are injected with radiolabelled androgens, these studies do not verify the chemical identity of radioactive compound(s), while it is known that these androgens can be metabolized substantially. By using stable isotope labelled testosterone and androstenedione in combination with mass spectrometry, enabling verification of the exact molecular identity of labelled compounds in the embryo, we found that after five days of incubation the androgens are not taken up by the embryo. Yet their concentrations in the entire yolk-albumen homogenates decline strongly, even when corrected for dilution by albumen and water. Our results indicate metabolism of maternal androgens, very likely to 5β-androstane-3α,17β-diol, etiocholanolone, and their conjugated forms. The results imply that the effects of increased exposure of the embryo to maternal androgens either take place before this early conversion or are mediated by these metabolites with a so far unknown function, opening new avenues for understanding hormone mediated maternal effects in vertebrates.

Electronic supplementary material is available online at https://dx.doi.org/10.6084/m9.figshare. c.4440182 When different genotypes choose different habitats to better match their phenotypes, genetic differentiation within a population may be promoted. Mating within those habitats may subsequently contribute to reproductive isolation. In cichlid fish, visual adaptation to alternative visual environments is hypothesized to contribute to speciation. Here, we investigated whether variation in visual sensitivity causes different visual habitat preferences, using two closely related cichlid species that occur at different but overlapping water depths in Lake Victoria and that differ in visual perception (Pundamilia spp.). In addition to species differences, we explored potential effects of visual plasticity, by rearing fish in two different light conditions: broad-spectrum (mimicking shallow water) and red-shifted (mimicking deeper waters). Contrary to expectations, fish did not prefer the light environment that mimicked their typical natural habitat. Instead, we found an overall preference for the broad-spectrum environment. We also found a transient influence of the rearing condition, indicating that the assessment of microhabitat preference requires repeated testing to control for familiarity effects. Together, our results show that cichlid fish exert visual habitat preference but do not support straightforward visual habitat matching info:eu-repo/semantics/publishedVersion

Maternal hormones deposited in the egg can provide a powerful model for the study of maternal effects. The differential amount of maternal hormones in the yolk of freshly laid eggs is assumed to represent differential maternal allocation. However, some evidence suggests that these amounts do not reflect maternal allocation that in fact takes place before ovulation. We compared the amounts of a wide array of gonadal steroids and their metabolites in the yolk of pre-ovulatory follicles with those of freshly laid eggs of rock pigeons using mass spectrometry. We found that between the follicle and egg stages the levels of progesterone increase whereas androstenedione and testosterone decrease in which the strength of decrease was dependent on the laying order of the egg. For conjugated estrone the change between follicle and egg differed in direction for first and second laying position yielding a significant interaction effect. For conjugated testosterone the interaction did not reach but was close to significance. This extremely early steroid metabolism was not due to maternal enzymes in the yolk as indicated by incubation of pre-ovulatory yolks treated with proteinase-K, a protein digesting enzyme. The results have significant consequences for the functional and evolutionary interpretation as well as experimental manipulation of hormone-mediated maternal effects.

Mothers may modulate the phenotype of their offspring by affecting their development based on her own environment. In changing environments, these maternal effects are thought to adjust offspring physiology and development and thus produce offspring better prepared to the environment experienced by the mother. However, evidence for this is scarce. Here we test the consequences of a match or mismatch between mother and offspring temperature conditions on growth, adult morphology and reproduction into the grandchildren generation in the fruit fly Drosophila melanogaster. This experimental design tests the relative contribution of maternal effects and offspring intrinsic plasticity to the phenotypic response to temperature conditions. We manipulated maternal temperature conditions by exposing mothers to either 18°C or 29°C conditions. Their eggs developed at a temperature that was either matched or mismatched with the maternal one. Survival from egg to adult was higher when the maternal and offspring environments matched, showing maternal effects affecting a trait that is a close proxy for fitness. However developmental speed, adult size and fecundity responded to temperature mostly through offspring phenotypic plasticity and maternal effects only had a small contribution. The results provide experimental evidence for maternal effects in influencing a potentially adaptive offspring response to temperature in the model organismDrosophila melanogaster.These effects appear to modulate early embryonic phenotypes such as survival, more than the adult phenotypes of the offspring.

Vertebrate embryos are exposed to maternal hormones that can profoundly affect their later phenotype. Although it is known that the embryo can metabolize these maternal hormones, the metabolic outcomes, their quantitative dynamics and timing are poorly understood. Moreover, it is unknown whether embryos can adjust their metabolic activity to, for example, hormones or other maternal signals. We studied the dynamics of maternal steroids in fertilized and unfertilized rock pigeon eggs during early incubation. Embryos of this species are naturally exposed to different amounts of maternal steroids in the egg according to their laying position, which provides a natural context to study differential embryonic regulation of the maternal signals. We used mass spectrometric analyses to map changes in the androgen and estrogen pathways of conversion. We show that the active hormones are heavily metabolized only in fertilized eggs, with a corresponding increase in supposedly less potent metabolites already within one-fourth of total incubation period. Interestingly, the rate of androgen metabolism was different between embryos in different laying positions. The results also warrant a re-interpretation of the timing of hormone mediated maternal effects and the role of the supposedly biologically inactive metabolites. Furthermore, the results also provide a potential solution as to how the embryo can prevent maternal steroids in the egg from interfering with its sexual differentiation processes as we show that the embryo can metabolize most of the maternal steroids before sexual differentiation starts.

In many seasonally breeding birds, female and male testosterone (T) levels peak at the start of the breeding season, coinciding with pair bonding and nesting activities. Shortly after the onset of egg laying, T levels slowly decline to baseline levels in both sexes, but more rapidly so in females. During this period, T in males may still function to facilitate territorial behaviour, mate guarding and extra pair copulations, either via short lasting peaks or elevated basal levels of the hormone. In some species, however, males become insensitive to increased T after the onset of egg laying. It has been postulated that in these species bi-parental care is essential for offspring survival, as T is known to inhibit paternal care. However, only very few studies have analysed this for females. As females are heavily involved in parental care, they too might become insensitive to T after egg laying. Alternatively, because territorial defence, mate guarding and extra pair copulations are expected to be less important for females than for males, they may not have had the need to evolve a mechanism to become insensitive to T during the period of maternal care, because their natural T levels are never elevated during this part of the breeding season anyway. We tested these alternative hypotheses in female great tits (Parus major). Male great tits have previously been shown to be insensitive to T after egg laying with regard to nestling feeding behaviour (but not song rate). When females had started nest building, we experimentally elevated their T levels up to the nestling feeding phase, and measured incubation behaviour (only females incubate) and reproductive success. T did not significantly affect nest building or egg laying behaviour, although egg laying tended to be delayed in T females. Females with experimentally enhanced T maintained lower temperature during incubation but did not spend less time incubating. This might explain the reduced hatching success of their eggs, smaller brood size and lower number of fledglings we found in this study. As in this species T-dependent behaviour by females during the phase of parental care is not needed, the results support the hypothesis that in this species the need for selection in favour of T-insensitivity did not occur. (C) 2016 Elsevier Inc. All rights reserved.

In birds, mothers can affect their offspring's phenotype and thereby survival via egg composition. It is not well known to what extent and time-scales environmental variation in resource availability, either via resource constrains or adaptive adjustment to predicted rearing conditions, influences maternal effects. We experimentally studied whether egg and yolk mass and yolk hormone levels respond to short-term changes in food availability during laying in wild great tits (Parus major). Our treatment groups were: 1) food supplementation (mealworms) from the 1st until the last egg; 2) food supplementation from the 1st first until the 5th egg, where the effect of cessation of the supplementary food treatment could also be studied; 3) no food supplementation (controls). We analysed both nutritional resources (egg, yolk and albumen mass), and the important developmental signals, yolk androgens (testosterone and androstenedione), and for the first time in a wild population, yolk thyroid hormones (thyroxine and 3,5,3’-triiodothyronine). Egg mass is a costly resource for females, androgens most likely non-costly signals, whereas thyroid hormones may be costly signals, requiring environmental iodine. In the food supplemented group egg, yolk and albumen mass increased rapidly relative to controls and when food supplementation was halted, egg and albumen mass decreased, indicating rapid responses to resource availability. Yolk androgen and thyroid hormone levels were not affected by food supplementation during laying. Thyroxine showed an increase over the laying sequence and its biological meaning needs further study. The rapid changes in egg mass to variation in within-clutch food availability suggest energetic/protein/nutrient constrains on egg formation. The lack of a response in yolk hormones suggest that perhaps in this species the short-term changes in resource availability during egg laying do not predict offspring rearing conditions, or (for thyroid hormones) do not cause systemic changes in circulating hormones, and hence do not affect maternal signaling.

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.

Animal personality traits emerge developmentally from the interaction of genetic and early environmental factors. Maternal hormones, such as androgens (testosterone, T and androstenedione, A4), transferred to embryos and egg yolks may simultaneously organize multiple behavioural and physiological traits. Whereas previous studies demonstrated an association between the mother's personality and yolk androgen levels, the independent effects of the male partner's personality and pair combination are unknown. We test this association using an ecological model species for personality research, the great tit (Parus major) using multiple approaches: 1) a wild population, 2) a randomly-mated captive population, and 3) an experimental study with (dis)assortatively mated pairs from lines selected for fast exploration/boldness or slow exploration/shyness. Egg androgen concentrations were associated with variation in female personality traits, and the experimental data suggested that this is independent of male personality: experimental females from slow-shy line tended to have higher egg T concentrations than females from the fast-bold line, with no effect of male personality. Shy females from the wild population had higher egg A4 concentration than bold females. However, in the correlative data yolk hormones were linked with male personality, as well as the interaction between female and male traits and yolk androgen levels: Male handling responsiveness correlated negatively with egg A4 concentration in wild birds. In randomly-mated birds, pairs that were mated assortatively for personality had lower egg T concentrations than disassortatively mated pairs. Given that egg androgens are known mediators of avian personality, our results suggest that maternal hormones might contribute to the heritability of personality, may be sensitive to the social context of mating, and act as key drivers of individual differences. This article is protected by copyright. All rights reserved.

Differences in the deposition of limited maternal resources to eggs can reflect the optimal allocation to manipulate offspring phenotype, or constraints caused by maternal condition. We examined multiple maternal substances transferred to eggs in the magpie Pica pica to test the hypothesis that certain substances represent constraint and some optimal allocation. We did this by supplementary feeding magpies prior to egg-laying and then comparing the effect of food on maternal substances in conjunction with laying order relative to a control group. Certain substances such as carotenoids, immunoglobulins and avidin responded positively to food supplementation, whereas others, such as testosterone and total protein content of eggs, did not. Immunoglobulin allocation increased with laying order in fed but not in unfed groups. Our results suggest that patterns of optimal allocation and constraint in maternally-provisioned substances co-occur within clutches. Furthermore, constraint and optimal allocation may co-occur within the same substance i.e. immunoglobulins, highlighting the complexities of maternal resource allocation.

Early-life food conditions can have profound impact on adult behavioural performance. In song birds, early-life food conditions affect adult physiology and cognitive performance such as song learning and spatial learning. However, effects on reproductive behaviour other than song, such as visual courtship display, pair formation, and egg laying, which are all important determinants of fitness, received hardly any attention. In this study, rock pigeons Columba livia were raised either in a food ad-libitum or at impoverished (quality) and restricted (quantity) food conditions, in which the latter was applied for either 26 or 8 d after hatching and in slightly different severity. Their growth and reproductive behaviour in adulthood was analysed under semi-natural conditions. Impoverished and restricted food (IRF) negatively affected adult body mass, pair-bonding behaviour, courtship (males) and being courted (females), and induced incomplete compensatory growth. Further analyses suggest that the effects of IRF on behaviour could result from the strong effects on body mass, and perhaps also from detrimental effects of compensatory growth. These results provide evidence that adverse early-life condition can negatively impact on reproductive success, as suggested by the significant effect on egg production, which will very likely lead to a lower life-time fitness. The detrimental effects on male courtship indicate that IRF can affect sexual displays via other pathways than by impaired cognition as has been suggested for bird song, since courtship in the pigeons is not learned.