Brood Size Manipulation Research Articles

Experimental support for partial compensation, not matched, partner response rules in blue tits.

Outcomes of theoretical models on conflict resolution between investing partners in bi-parental care systems typically predict "partial compensation" or "matched" response rules,depending on underlying assumptions. Further, although experimental tests in birds suggest that care levels by pair members are largely associated with partial compensation responses, this outcome partly depends on the type of experiment used and its effects on model assumptions. To elucidate support for both the underlying assumptions and predictions of models predicting partner compensation versus matching, we performed temporary, bi-directional brood size manipulations during late nestling provisioning in blue tits (Cyanistes caeruleus) in the French Pyrenees. We found support for assumptions of both compensation and matching models. For example, females spent more time with the brood than males, leading to potential information asymmetries on brood demand as expected by matching models. Nevertheless, both pair members changed their provisioning comparably in response to brood size manipulations, suggesting that pair members have comparable cost-benefit functions in response to variation in brood demand, as assumed under partial compensation models. Despite support for the underlying assumptions of both models, we primarily found support for predictions of partial compensation models in provisioning responses. Notably, modest changes by one member of the pair on experimental days were met with larger changes by the other, after controlling for brood size and provisioning rates on control days. Our results corroborate previous findings in blue tits using alternative experimental approaches. We suggest that future studies could benefit from understanding when partial compensation responses dominate matched ones, despite apparent asymmetries in information over brood demand.

Early-life environmental effects on mitochondrial aerobic metabolism: a brood size manipulation in wild great tits.

In avian species, the number of chicks in the nest and subsequent sibling competition for food are major components of the offspring's early-life environment. A large brood size is known to affect chick's growth, leading in some cases to long-lasting effects for the offspring, such as a decrease in size at fledgling and in survival after fledging. An important pathway underlying different growth patterns could be the variation in offspring mitochondrial metabolism through its central role in converting energy. Here, we performed a brood size manipulation in great tits (Parus major) to unravel its impact on offspring's mitochondrial metabolism and reactive oxygen species (ROS) production in red blood cells. We investigated the effects of brood size on chicks' growth and survival, and tested for long-lasting effects on juvenile mitochondrial metabolism and phenotype. As expected, chicks raised in reduced broods had a higher body mass compared to enlarged and control groups. However, mitochondrial metabolism and ROS production were not significantly affected by the treatment either at chick or juvenile stages. Interestingly, chicks raised in very small broods were smaller in size and had higher mitochondrial metabolic rates. The nest of rearing had a significant effect on nestling mitochondrial metabolism. The contribution of the rearing environment in determining offspring mitochondrial metabolism emphasizes the plasticity of mitochondrial metabolism in regards to the nest environment. This study opens new avenues regarding the implication of postnatal environmental conditions in shaping the offspring's early-life mitochondrial metabolism.

Plumage color degradation indicates reproductive effort: an experiment

Plumage color has traditionally been regarded as a static ornamental trait, but evidence is accumulating for significant color changes without molt that typically reduce the conspicuousness of ornamentation. In some species, the social partner seems to increase its reproductive investment if the color trait is experimentally enhanced, suggesting that color change could act as a signal. However, the information content of this signal is so far unclear. For example, birds in poor condition or making greater effort may deteriorate more severely. We used brood size manipulations to alter the reproductive effort of male and female collared flycatchers Ficedula albicollis. Both sexes showed less severe decline in some reflectance attribute of their white breast when their brood was experimentally reduced. In each sex, greater deterioration of the reflectance trait affected by the manipulation was accompanied by increased feeding rate by the partner. These feeding patterns do not prove, but are consistent with, a compensatory response by the partner to induced degradation. The manipulation effects on color change we detected confirm for the first time that plumage color deterioration can indicate current reproductive effort, thereby providing a potential fitness advantage to social partners that react to such deterioration.

Ectoparasite presence and brood size manipulation interact to accelerate telomere shortening in nestling jackdaws.

Early-life conditions impact fitness, but whether the combined effect of extrinsic stressors is additive or synergistic is not well known. This is a major knowledge gap because exposure to multiple stressors is frequent. Telomere dynamics may be instrumental when testing how stressors interact because many factors affect telomere shortening, and telomere shortening predicts survival. We evaluated the effect of manipulated brood size and natural infestation by the carnid fly Carnus hemapterus on nestling growth and telomere shortening of wild jackdaws (Corvus monedula). Telomere length, measured in blood using TRF, shortened on average by 264 bp, and on average, Carnus infection induced more telomere shortening. Further analyses showed that in enlarged broods, nestlings' telomeres shortened more when parasitized, while in reduced broods there was no effect of infection on telomere shortening. We conclude that there is a synergistic effect of number of siblings and Carnus infection on telomere shortening rate: blood-sucking parasites may negatively impact telomeres by increasing cell proliferation and/or physiological stress, and coping with infection may be less successful in enlarged broods with increased sibling competition. Larger nestlings had shorter telomeres independent of age, brood manipulation or infection. Growth was independent of infestation but in enlarged broods, nestlings were lighter at fledging. Our findings indicate that (i) evaluating consequences of early-life environmental conditions in isolation may not yield a full picture due to synergistic effects, and (ii) effects of environmental conditions may be cryptic, for example, on telomeres, with fitness consequences expressed beyond the temporal framework of the study.

Variation in oxidative status, but not structural and physiological development, associated with changing ontogenetic environments.

Despite the potential for temporally-dependent relationships between trait values and fitness (e.g., as juveniles approach life-stage transitions such as fledging), how developmental stage affects canalization (a measure of robustness to environmental variation) of morphological and physiological traits is rarely considered. To test the sensitivity of morphological and physiological traits to environmental variation in two developmental stages, we manipulated brood size at hatch in European starlings (Sturnus vulgaris) and cross-fostered chicks between enlarged and reduced broods approaching fledging. We measured body size (mass, tarsus, wing length) and physiological state (aerobic capacity, oxidative status) at asymptotic mass on day 15, then cross-fostered chicks between 'high' and 'low' quality environments and assessed the same traits again on day 20, after five days of pre-fledging mass recession. Chicks in reduced broods were heavier at asymptotic mass and had lower reactive oxygen metabolites than enlarged broods, while structural size, aerobic capacity, and antioxidant capacity were unaffected by experimental brood size. The observed canalization of structural and physiological traits during early development was maintained after cross-fostering, during late development. However, in contrast to early development, antioxidant capacity approaching fledging appeared sensitive to environmental conditions, as trajectories varied by cross-fostering treatment. Elevated reactive oxygen metabolites observed after early development in enlarged brood chicks were maintained after cross-fostering, suggesting canalized development in low-quality environments could produce oxidative costs that carry over between life-stages, even when conditions improve. These data reveal trait-specific relationships between environmental conditions and development and highlight how natal environment effects may vary by developmental stage.

Females with Increased Costs Maintain Reproductive Output: A Field Experiment in a Common Songbird.

Reproduction and self-maintenance are energetically costly activities involved in classic life history trade-offs. However, few studies have measured the responses of wild organisms to simultaneous changes in reproductive and self-maintenance costs, which may have interactive effects. In free-living female Barn Swallows (Hirundo rustica), we simultaneously manipulated reproductive costs (by adding or removing two nestlings) and self-maintenance costs (by attaching a ∼1 g weight in the form of a GPS tag to half of our study birds) and measured mass, immune status, blood glucose, feather growth, and reproductive output (likelihood of a second clutch, number of eggs, and time between clutches). GPS tags allowed us to analyze how movement range size affected response to brood size manipulation. Tagging altered females' immune function as evidenced by an elevated heterophil to lymphocyte (H:L) ratio, but all females were equally likely to lay more eggs. There was no evidence of interactive effects of the tagging and brood size treatment. Range size was highly variable, and birds with large ranges grew feathers more slowly, but analyzing the effect of brood size manipulation while accounting for variation in range size did not result in any physiological response. Our results support the theoretical prediction that short-lived vertebrates do face a trade-off between reproduction and self-maintenance and, when faced with increased costs, tend to preserve investment in reproduction at the expense of parental condition. This experiment also helps us to understand how movement patterns may be relevant to life history trade-offs in wild birds.

When parents play favorites: brood demand shapes parental preference for offspring UV color.

Parents might initially produce more offspring than they might be able to raise. However, when offspring demand exceeds their parents´ rearing capacity, parents might shift care towards the offspring which yield greater fitness returns to achieve their optimal brood size via brood reduction. Such favoritism could rely on offspring signaling traits if these inform parents about offspring quality and hence about the pay-offs of their investment. Here we investigated whether favoritism of blue tit (Cyanistes caeruleus) parents for an offspring signal (i.e., ultraviolet (UV) plumage coloration) varies with brood demand. To test this, we experimentally blocked the UV reflectance of yellow breast feathers in half of the nestlings of each brood, and then we sequentially performed two opposing brood size manipulations to vary nestling demand below or above parental rearing capacity. In reduced broods, nestlings begged overall less intensely and gained more body mass, supporting that parental rearing capacities sufficed to satisfy brood demand. Moreover, in reduced broods, UV-blocked nestlings (i.e., low-quality offspring) were fed and prey-tested more often. Yet, they begged more than control nestlings, suggesting that they were perhaps treated differently by other family members or which they may exploit parental preferences beyond actual need (at least in reduced nests). Parents flexibly shifted their feeding rate and favoritism in response to short-term changes in family size, as there was no parental preference for enlarged broods. Such flexible parental feeding rules may allow parents to gain the upper hand in parent-offspring conflict. However, we did not find evidence that parental favoritism facilitated brood reduction, at least in conditions where demand was temporally enhanced.

Insulin-like growth factor 1 and the hormonal mediation of sibling rivalry

In altricial animals, young are completely dependent on parents for provisioning. The ability to outcompete siblings to receive parental provisioning has clear fitness benefits, and may be mediated by hormones that influence growth. We analyzed the effects of insulin-like growth factor 1 (IGF-1) on body size, growth, and sibling rivalry in eastern bluebirds (Sialia sialis). To determine whether IGF-1 is upregulated in response to the competitive environment, we manipulated brood sizes and examined the effect on IGF-1 levels, nestling body size, growth rate, and behavior. In a separate experiment, we injected nestlings with exogenous IGF-1 to study its impacts on body size, growth rate, and sibling competition. Brood size manipulation did not influence endogenous IGF-1 levels, but male nestlings with higher IGF-1 levels early in the nestling period tended to have greater mass gain than males with lower IGF-1 levels. Nestlings with higher IGF-1 levels also tended to be fed more frequently by parents. In the injection experiment, IGF-1 injected individuals tended to be heavier than vehicle injected young by the end of the nestling period, which suggests that IGF-1 can influence mass gain in bluebirds. IGF-1 has been proposed to be a mediator of life-history strategies and post-hatching behavior. Our results suggest that although bluebird nestlings do not adaptively elevate IGF-1 in response to the presence or number of siblings, IGF-1 may influence growth during the nestling period. These findings shed light on sibling competition, life history strategies, and the hormones that underlie them.

Visitation rate, but not foraging range, responds to brood size manipulation in an aerial insectivore

Life history theory predicts that increased investment in current offspring decreases future fecundity or survival. Avian parental investment decisions have been studied either via brood size manipulation or direct manipulation of parental energetic costs (also known as handicapping). However, we have limited experimental data on the potential interactive effects of these manipulations on parent behavior. Additionally, we know little about how these manipulations affect spatial foraging behavior away from the nest. We simultaneously manipulated brood size and parental costs (via added weight in the form of a GPS tag) in wild female barn swallows (Hirundo rustica). We measured multiple aspects of parent behavior at and away from the nest while controlling for measures of weather conditions. We found no significant interactive effects of manipulated brood size and parental costs. Both sexes increased their visitation rate with brood size, but nestlings in enlarged broods grew significantly less post-brood size manipulation than those in reduced broods. Foraging range area was highly variable among GPS-tagged females but was unaffected by brood size. As such, increased visitation rate in response to brood size may be more energetically costly for far-ranging females. GPS-tagged females did not alter their visitation rate relative to un-tagged birds, but their mates had higher visitation rates. This suggests that GPS tagging may affect some unmeasured aspect of female behavior, such as prey delivery. Our findings indicate that investigation of foraging tactics alongside visitation rate is critical to understanding parental investment and the benefits and costs of reproduction.Significance statementAvian parental investment decisions have been studied by either brood size manipulation or direct manipulation of parental costs, but rarely both simultaneously. We simultaneously manipulated brood size and parental costs (via addition of a GPS tag) in a wild avian system, allowing us to examine interactive effects of these manipulations. Additionally, studies of parental investment often examine behaviors at the nest, but measurements of parental care behavior away from the nest are rare. Our study is unique in that we measured multiple aspects of parental care, including spatial foraging behavior tracked with GPS tags. We found no interactive effects of manipulated brood size and parental costs on visitation rate or nestling growth, and spatial foraging behavior of females was individually variable. Documenting foraging tactics alongside visitation rate is critical to understanding parental investment because the same visitation rate might be more costly for far-ranging females.

Access to resources buffers against effects of current reproduction on future ability to provide care in a burying beetle.

Studies investigating the trade‐off between current and future reproduction often find that increased allocation to current reproduction is associated with a reduction in the number or quality of future offspring. In species that provide parental care, this effect on future offspring may be mediated through a reduced future ability to provide care. Here, we test this idea in the burying beetle Nicrophorus vespilloides, a species in which parents shift the cost of reproduction toward future offspring and provide elaborate parental care. We manipulated brood size to alter the costs females experienced in association with current reproduction and measured the level of parental care during a subsequent breeding attempt. Given that these beetles breed on carcasses of small vertebrates, it is important to consider confounding effects due to benefits associated with resource access during breeding. We, therefore, manipulated access to carrion and measured the level of parental care during a subsequent breeding attempt. We found that females provided the same level of care regardless of previous brood size and resource access, suggesting that neither affected future ability to provide care. This may reflect that parents feed on carrion during breeding, which may buffer against any costs of previous breeding attempts. Our results show that increased allocation to current reproduction is not necessarily associated with a reduction in future ability to provide care. Nevertheless, this may reflect unique aspects of our study system, and we encourage future work on systems where parents do not have access to a rich resource during breeding.

The interactive effect of ambient temperature and brood size manipulation on nestling body mass in blue tits: an exploratory analysis of a long-term study

BackgroundRelatively few studies have examined the interactive effects of ecological factors on physiological responses in wild animals. Nearly all of them have been short-term investigations that did not include experimental manipulations, limiting our ability to understand how climate change will affect natural populations. Using a 10-year brood size manipulation experiment in wild blue tits (Cyanistes caeruleus), we quantified the impact of weather conditions and brood competition on the body mass and structural size (tarsus length) of nestlings just prior to leaving the nest.ResultsWe found that variation in nestling body mass on day 14 after hatching was explained by an interactive effect between average ambient temperature experienced during nestling period and brood size treatment. Specifically, in control broods nestling body mass was correlated with temperature in a non-linear manner (concave) with the vertex point (maximum body mass) at ca. 13 °C. In contrast, in enlarged broods nestling body mass permanently increased (also non-linearly) as temperature advanced.ConclusionsOur results highlight the importance of considering the effects of brood rearing conditions alongside other environmental factors experienced during growth while investigating early-life environmental effects on body condition.

Brood size manipulation reveals cost on the size of the second clutch in the earwig Anisolabis maritime

Abstract There may be a trade-off between the duration of parental care and future reproductive success. Traditionally, studies about the cost of parental care have included the removal of the parent. However, producing a secondary clutch after the failure of the first one is a compensatory behaviour that occurs in cases of brood failure. In this study, attempts were made to detect the cost of maternal care in the earwig, Anisolabis maritima (Dermaptera: Anisolabididae) by either extending the period of care or increasing the brood size to prevent compensation through the brood’s success. The results indicated that manipulation did not change the inter-clutch interval, although my previous study revealed shortening of these intervals after the removal of the clutch in this species. In this study, decreased clutch size manipulation increased the size of the following clutch.

Parental Care System and Brood Size Drive Sex Difference in Reproductive Allocation: An Experimental Study on Burying Beetles

Life-history theory predicts that increased resource allocation in current reproduction comes at the cost of survival and future reproductive fitness. In taxa with biparental care, each parent can adjust investment on current reproduction according to changes in their partner’s effort, but these adjustments may be different for males and females as they may have different reproductive strategies. Numerous theoretical and empirical studies have proposed the mechanism underlying such adjustments. In addition, the value of the brood or litter (brood size) has also been suggested to affect the amount of care through manipulation of brood size. While the two conditions have been studied independently, the impact of their interplay on potential sex-dependent future reproductive performance remains largely unknown. In this study, we simultaneously manipulated both care system (removal of either parent vs. no removal) and brood size in a burying beetle (Nicrophorus vespilloides) to understand their joint effect on reproductive allocation and trade-off between current and future reproduction. Our results show that males compensated for mate loss by significantly increasing the level of care regardless of brood size, while females exhibited such compensation only for small brood size. Additionally, with an increase in allocation to current reproduction, males showed decreased parental investment during the subsequent breeding event as a pair. These findings imply a dual influence of parental care system and brood size on allocation in current reproduction. Moreover, the impact of such adjustments on sex-dependent differences in future reproduction (parental care, larvae number, and average larval mass at dispersal) is also demonstrated. Our findings enhance the understanding of sex roles in parental investment and highlight their importance as drivers of reproductive allocation.

Variation in reproductive investment increases body temperature amplitude in a temperate passerine

Many birds and mammals show substantial circadian variation in body temperature, which has been attributed to fluctuations in ambient temperature and energy reserves. However, to fully understand the variation in body temperature over the course of the day, we also need to consider effects of variation in work rate. We made use of a dataset on body temperature during the resting and active periods in female marsh tits (Poecile palustris) that bred in a temperate area and were subjected to experimental changes in reproductive investment through brood size manipulations. Furthermore, the amplitude increased with daytime, but were unaffected by nighttime, ambient temperature. Amplitudes in females with manipulated broods were 44% above predictions based on inter-specific allometric relationships. In extreme cases, amplitudes were > 100% above predicted values. However, no individual female realised the maximum potential amplitude (8.5 °C, i.e. the difference between the highest and lowest body temperature within the population) but seemed to prioritise either a reduction in body temperature at night or an increase in body temperature in the day. This suggests that body temperature amplitude might be constrained by costs that preclude extensive use of both low nighttime and high daytime body temperatures within the same individual. Amplitudes in the range found here (0.5–6.7 °C) have previously mostly been reported from sub-tropical and/or arid habitats. We show that comparable values can also be found amongst birds in relatively cool, temperate regions, partly due to a pronounced increase in body temperature during periods with high work rate.

Effects of maternal exposure to a bacterial antigen and altered post-hatching rearing conditions on avian offspring behaviour

The early-life environment plays a crucial role in shaping morphological, physiological, and behavioural traits, with potential long-term consequences for fitness. Indeed, a set of factors experienced by offspring during prenatal and early post-natal development has been recognised to affect behavioural trait expression in later life. Several studies have shown that in birds, nutritional and social rearing conditions and maternal and/or neonatal immunisations may profoundly determine the development and establishment of behaviour in offspring. To our knowledge, no research has examined whether and how the interaction between immune-mediated maternal effects and post-hatching rearing conditions affects offspring behaviour. Here, we studied the effects of maternal exposure to a bacterial antigen and altered brood size on docility, breathing rate, and aggression in the offspring of great tit, Parus major. We used a 2 × 2 design to investigate the interactive effects of maternal immunisation and brood size manipulation on offspring behavioural development. We found no such interactive effect on offspring behaviour, although we observed it regarding to offspring body mass and tarsus length. Maternal immunisation itself did not affect offspring behaviour. However, we demonstrated that the offspring breathing rate and level of aggression were affected by brood size manipulation. Both breathing rate and aggression in offspring reared in enlarged broods were lower than those in offspring reared in non-manipulated broods. Our study did not confirm earlier reports that immune-mediated maternal effects modulate offspring behavioural development, but we showed that brood size during rearing might indeed be a factor that affects offspring behaviour.Significance statementThe early environment experienced by offspring constitutes a significant source of developmental plasticity, which may profoundly affect the establishment of their behavioural traits. Food availability, social conditions, and maternal or offspring infection are crucial factors shaping various behavioural traits in birds. However, there remains a lack of studies emphasising the potential interactive effects of early-life conditions on behavioural trait development in natural bird populations. Here, to our knowledge for the first time, we experimentally examined how maternal immunisation and altered post-hatching rearing conditions interact to determine the behaviour of fledged offspring. We found that maternal treatment and brood size manipulation interactively affected offspring body mass and tarsus length, but this interaction had no effect on offspring behaviour. Our findings suggest that different mechanisms may underlie the development of morphological and behavioural traits.

Gull chicks grow faster but lose telomeres when prenatal cues mismatch the real presence of sibling competitors.

During embryonic life, individuals should adjust their phenotype to the conditions that they will encounter after birth, including the social environment, if they have access to (social) cues that allow them to forecast future conditions. In birds, evidence indicates that embryos are sensitive to cues from clutch mates, but whether embryos adjust their development to cope with the expected level of sibling competition has not hitherto been investigated. To tackle this question, we performed a 'match versus mismatch' experimental design where we manipulated the presence of clutch mates (i.e. clutch size manipulation) and the real (postnatal) level of sibling competition (i.e. brood size manipulation) in the yellow-legged gull (Larus michahellis). We provide evidence that the prenatal cues of sibling presence induced developmental changes (such as epigenetic profiles) that had programming effects on chick begging behaviour and growth trajectories after hatching. While receiving mismatching information favoured chick begging and growth, this came at the cost of reduced antioxidant defences and a premature loss of telomeres. Our findings highlight the role of the prenatal social environment in developmental plasticity and suggest that telomere attrition may be an important physiological cost of phenotype-environment mismatch.

Increased allocation to reproduction reduces future competitive ability in a burying beetle.

The existence of a trade-off between current and future reproduction is a fundamental prediction of life history theory. Support for this prediction comes from brood size manipulations, showing that caring for enlarged broods often reduces the parent's future survival or fecundity. However, in many species, individuals must invest in competing for the resources required for future reproduction. Thus, a neglected aspect of this trade-off is that increased allocation to current reproduction may reduce an individual's future competitive ability. We tested this prediction in the burying beetle, Nicrophorus vespilloides, a species where parents care for their offspring and where there is fierce competition for resources used for breeding. We manipulated reproductive effort by providing females with either a small brood of 10 larvae or a large brood of 40 larvae and compared the ability of these females, and virgin females that had no prior access to a carcass, to compete for a second carcass against a virgin competitor. We found that increased allocation to current reproduction reduced future competitive ability, as females that had cared for a small brood were more successful when competing for a second carcass against a virgin competitor than females that had cared for a large brood. In addition, the costs of reproduction were offset by the benefits of feeding from the carcass during an initial breeding attempt, as females that had cared for a small brood were better competitors than virgin females that had no prior access to a carcass, whilst females that had cared for a large brood were similar in competitive ability to virgin females. Our results add to our understanding of the trade-off between current and future reproduction by showing that this trade-off can manifest through differences in future competitive ability and that direct benefits of reproduction can offset some of these costs.

Experimentally increased brood size accelerates actuarial senescence and increases subsequent reproductive effort in a wild bird population.

The assumption that reproductive effort decreases somatic state, accelerating ageing, is central to our understanding of life‐history variation. Maximal reproductive effort early in life is predicted to be maladaptive by accelerating ageing disproportionally, decreasing fitness.Optimality theory predicts that reproductive effort is restrained early in life to balance the fitness contribution of reproduction against the survival cost induced by the reproductive effort. When adaptive, the level of reproductive restraint is predicted to be inversely linked to the remaining life expectancy, potentially resulting in a terminal effort in the last period of reproduction.Experimental tests of the reproductive restraint hypothesis require manipulation of somatic state and subsequent investigation of reproductive effort and residual life span. To our knowledge the available evidence remains inconclusive, and hence reproductive restraint remains to be demonstrated.We modulated somatic state through a lifelong brood size manipulation in wild jackdaws and measured its consequences for age‐dependent mortality and reproductive success.The assumption that lifelong increased brood size reduced somatic state was supported: Birds rearing enlarged broods showed subsequent increased rate of actuarial senescence, resulting in reduced residual life span.The treatment induced a reproductive response in later seasons: Egg volume and nestling survival were higher in subsequent seasons in the increased versus reduced broods' treatment group. We detected these increases in egg volume and nestling survival despite the expectation that in the absence of a change in reproductive effort, the reduced somatic state indicated by the increased mortality rate would result in lower reproductive output. This leads us to conclude that the higher reproductive success we observed was the result of higher reproductive effort.Our findings show that reproductive effort negatively covaries with remaining life expectancy, supporting optimality theory and confirming reproductive restraint as a key factor underpinning life‐history variation.

Differential effects of early growth conditions on colour-producing nanostructures revealed through small angle X-ray scattering and electron microscopy.

The costs associated with the production and maintenance of colour patches is thought to maintain their honesty. Although considerable research on sexual selection has focused on structurally coloured plumage ornaments, the proximate mechanisms of their potential condition dependence, and thus their honesty, is rarely addressed, particularly in an experimental context. Blue tit (Cyanistes caeruleus) nestlings have ultraviolet (UV)-blue structurally coloured tail feathers, providing a unique opportunity for investigation of the causes of variation in their colour. Here, we examined the influence of early growing conditions on the reflectance and structural properties of UV-blue-coloured tail feathers of blue tit nestlings. We applied a two-stage brood size manipulation to determine which stage of development more strongly impacts the quality of tail feather colouration and microstructure. We used small-angle X-ray scattering (SAXS) and electron microscopy to characterise the nanoscale and microscale structure of tail feather barbs. Nestlings from the broods enlarged at a later stage of growth showed a sex-specific rectrix development delay, with males being more sensitive to this manipulation. Contrary to predictions, treatment affected neither the quality of the barbs' nanostructures nor the brightness and UV chroma of feathers. However, at the microscale, barbs' keratin characteristics were impaired in late-enlarged broods. Our results suggest that nanostructure quality, which determines the UV-blue colour in tail feathers, is not sensitive to early rearing conditions. Furthermore, availability of resources during feather growth seems to impact the quality of feather microstructure more than body condition, which is likely to be determined at an earlier stage of nestling growth.

Alternation of nest visits varies with experimentally manipulated workload in brood-provisioning great tits

In species with biparental care, the amount of care devoted to offspring is affected by the negotiation rules that the parents adopt. Recently, turn taking in provisioning visits has been proposed as a negotiation rule by which parents respond to their partner's behaviour, which results in a perfect alternation of the nest visits by the parents. Empirical evidence suggests that parents do not strictly alternate their visits, and, so far, this imperfect alternation has received no experimental investigation. In this study, we tested whether alternation of nest visits might be subject to time constraints affecting the ability of parents to strictly take turns. We manipulated the workload of 15 great tit, Parus major, pairs using a short-term brood size manipulation. Parental nest visits alternated more in reduced than control and enlarged broods. To understand whether this variability could be caused by changes in turn taking, we explored the rate and regularity of the parents' intervisit intervals. Treatment differences in alternation were still present when controlling for the rate and regularity of the visits by each of the two parents, suggesting that workload also affected alternation via the temporal sequence of the intervals (e.g. via turn taking). Our results show that alternation of nest visits varies in response to workload and is not merely a by-product of variation in visit rate or regularity.