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Telomere length (TL) predicts health and survival across taxa. Variation in TL between individuals is thought to be largely of genetic origin, but telomere inheritance is unusual, because zygotes already express a TL phenotype, the TL of the parental gametes. Offspring TL changes with paternal age in many species including humans, presumably through age-related TL changes in sperm, suggesting an epigenetic inheritance mechanism. However, present evidence is based on cross-sectional analyses, and age at reproduction is confounded with between-father variation in TL. Furthermore, the quantitative importance of epigenetic TL inheritance is unknown. Using longitudinal data of free-living jackdaws Corvus monedula, we show that erythrocyte TL of subsequent offspring decreases with parental age within individual fathers, but not mothers. By cross-fostering eggs, we confirmed the paternal age effect to be independent of paternal age dependent care. Epigenetic inheritance accounted for a minimum of 34% of the variance in offspring TL that was explained by paternal TL. This is a minimum estimate, because it ignores the epigenetic component in paternal TL variation and sperm TL heterogeneity within ejaculates. Our results indicate an important epigenetic component in the heritability of TL with potential consequences for offspring fitness prospects., Author summary Telomeres are DNA-protein structures at chromosome ends and a short telomere length predicts reduced survival in humans, birds and other organisms. Variation in telomere length between individuals is thought to be largely of genetic origin, but telomere inheritance may be unusual because not only genes regulating telomere length are inherited, but a fertilised cell already has a telomere length (from the parental gametes). Using long-term individual-based data of jackdaw families (a small corvid species), we found that as fathers aged, they produced chicks with shorter telomeres. This suggests that telomere length inheritance has an epigenetic component. To investigate to what extent telomere length in the fertilised cell affects telomere length after birth, we compared telomere length over years within fathers with the telomere length of their consecutive offspring. This epigenetic component explained a substantial part (≥ one third) of the telomere length inheritance; whereas there was no such effect of maternal telomere length. The sex difference fits the idea that lifelong sperm formation leads to change in telomere length of the sperm cells, whereas female gametes are all formed before birth and their telomere length does not change over time.

International audience; Leukocyte telomere length (LTL) shortens with age. Longitudinal studies have reported accelerated LTL attrition when baseline LTL is longer. However, the dependency of LTL attrition on baseline LTL might stem from a statistical artifact known as regression to the mean (RTM). To our knowledge no published study of LTL dynamics (LTL and its attrition rate) has corrected for this phenomenon. We illustrate the RTM effect using replicate LTL measurements, and show, using simulated data, how the RTM effect increases with a rise in stochastic measurement variation (representing LTL measurement error), resulting in spurious increasingly elevated dependencies of attrition on baseline values. In addition, we re-analyzed longitudinal LTL data collected from four study populations to test the hypothesis that LTL attrition depends on baseline LTL. We observed that the rate of LTL attrition was proportional to baseline LTL, but correction for the RTM effect reduced the slope of the relationship by 57% when measurement error was low (coefficient of variation ~2%). A modest but statistically significant effect remained however, indicating that high baseline LTL is associated with higher LTL attrition even when correcting for the RTM effect. Baseline LTL explained 1.3% of the variation in LTL attrition, but this effect, which differed significantly between the study samples, appeared to be primarily attributable to the association in men (3.7%).

Mutual mate choice is prevalent in humans, where both males and females have a say in their choice of partner. How the choices made by one sex constrain the choice of the other remains poorly understood, however, because human studies have mostly limited themselves to measuring preferences. We used a sample of 5782 speed-daters making 128 104 choices to link preferences for partner height to actual choice and the formation of a match (the mutual expression of interest to meet again). We show that sexual conflict at the level of preferences is translated into choice: women were most likely to choose a speed-dater 25 cm taller than themselves, whereas men were most likely to choose women only 7 cm shorter than themselves. As a consequence, matches were most likely at an intermediate height difference (19 cm) that differed significantly from the preferred height difference of both sexes. Thus, our study reveals how mutual mate choice can result in suboptimal pair formation for both sexes, highlighting the importance of assessing the mate choice process in its entirety. (C) 2013 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Dietary restriction (DR) extends lifespan in multiple species from various taxa. This effect can arise via two distinct but not mutually exclusive ways: a change in aging rate and/or vulnerability to the aging process (i.e. initial mortality rate). When DR affects vulnerability, this lowers mortality instantly, whereas a change in aging rate will gradually lower mortality risk over time. Unraveling how DR extends lifespan is of interest because it may guide toward understanding the mechanism(s) mediating lifespan extension and also has practical implications for the application of DR. We reanalyzed published survival data from 82 pairs of survival curves from DR experiments in rats and mice by fitting Gompertz and also GompertzMakeham models. The addition of the Makeham parameter has been reported to improve the estimation of Gompertz parameters. Both models separate initial mortality rate (vulnerability) from an age-dependent increase in mortality (aging rate). We subjected the obtained Gompertz parameters to a meta-analysis. We find that DR reduced aging rate without affecting vulnerability. The latter contrasts with the conclusion of a recent analysis of a largely overlapping data set, and we show how the earlier finding is due to a statistical artifact. Our analysis indicates that the biology underlying the life-extending effect of DR in rodents likely involves attenuated accumulation of damage, which contrasts with the acute effect of DR on mortality reported for Drosophila. Moreover, our findings show that the often-reported correlation between aging rate and vulnerability does not constrain changing aging rate without affecting vulnerability simultaneously.

According to both the scientific literature and popular media, all one needs to win a US presidential election is to be taller than one's opponent. Yet, such claims are often based on an arbitrary selection of elections, and inadequate statistical analysis. Using data on all presidential elections, we show that height is indeed an important factor in the US presidential elections. Candidates that were taller than their opponents received more popular votes, although they were not significantly more likely to win the actual election. Taller presidents were also more likely to be reelected. In addition, presidents were, on average, much taller than men from the same birth cohort. The advantage of taller candidates is potentially explained by perceptions associated with height: taller presidents are rated by experts as 'greater', and having more leadership and communication skills. We conclude that height is an important characteristic in choosing and evaluating political leaders. (C) 2012 Elsevier Inc. All rights reserved.

Estimating the number of genome copies in a tissue sample can serve various purposes. For example, such an estimate serves as a scaling variable when measuring telomeres with quantitative PCR. We describe the primer development and evaluation for the glyceraldehyde-3-phosphate dehydrogenase (GAPDH) gene in the Great Tit Parus major, as a control gene to estimate genome copy number. We demonstrate specific amplification with negligible variation in 48 Great Tits and cross-amplification in 53 Blue Tits Cyanistes caeruleus. We conclude this primer set to be reliable for amplification of GAPDH as a reference gene for quantitative PCR analysis in Great and Blue Tits.

Iteroparous organisms face a trade-off between reproduction and survival, but knowledge of whether how and when costs of long-term increases in workload are paid is scant. We increased locomotion costs for a whole year by equipping male great tits with a backpack during breeding, removing the backpacks 1 year later. We applied 3 different treatments: control (without backpack), light ("empty" backpack, 0.1g), and heavy ("full" backpack, 0.9g, similar to 5% of body mass). Backpacks were administered in 3 cohorts, and we monitored effects on mass of nestlings and the male, wing length, reproduction, and survival. Added mass had a negative effect on nestling mass in both the starting year of the experiment and 1 year later, but not on production of fledglings or recruits. In winter and the next breeding season, males equipped with heavy backpacks had a higher (net) body mass and had shorter third primary feathers than the other 2 groups. Heavy backpack males were less likely to sleep in a nest box in winter. Nest boxes are optimal roosting sites, and we interpret this finding as a treatment effect on success in competition over this resource. However, there was no effect of the manipulation on survival. Overall, we found no long-term fitness consequences, and we discuss possible explanations and implications for the "starvation-predation theory" of optimal body mass. However, we found short-term effects of carrying extra weight suggesting that behavioral studies using small devices should consider the effects of equipping small non-migratory passerines with devices such as transmitters.

Bird populations provide excellent systems to investigate variation in longevity in the wild since individuals can often be monitored over their lifetime. A number of recent studies suggest that the dynamics of protective telomere chromosome caps (telomere length and rate of loss) are indicative of biological state and potentially useful as indicators of future longevity. Currently, Terminal Restriction Fragment (TRF) analysis and relative quantitative PCR (qPCR) are used to measure telomeres in birds, but with limitations. TRF analysis is time consuming, while relative qPCR gives a within-study relative value making it difficult to compare across experiments. Utilising an approach first developed in humans of using synthetic oligomer telomeric (TTAGGG)n and normaliser gene standards of known length to calibrate qPCR values, we describe a methodological adaptation to the avian qPCR telomere assay to make results comparable within, and potentially between, bird species. We evaluate this absolute qPCR method in the Seychelles warbler Acrocephalus sechellensis against relative qPCR measurements on the same samples. Telomere estimates from both methods showed an age-related decline in telomere length, and were highly correlated (r = 0.99). Absolute qPCR avian telomere analysis may prove a useful means of estimating telomere lengths in a calibrated, sensitive, and efficient way using small amounts of archived bird sample.

Human male height is associated with mate choice and intra-sexual competition, and therefore potentially with reproductive success. A literature review (n = 18) on the relationship between male height and reproductive success revealed a variety of relationships ranging from negative to curvilinear to positive. Some of the variation in results may stem from methodological issues, such as low power, including men in the sample who have not yet ended their reproductive career, or not controlling for important potential confounders (e.g. education and income). We investigated the associations between height, education, income and the number of surviving children in a large longitudinal sample of men (n = 3,578; Wisconsin Longitudinal Study), who likely had ended their reproductive careers (e.g. > 64 years). There was a curvilinear association between height and number of children, with men of average height attaining the highest reproductive success. This curvilinear relationship remained after controlling for education and income, which were associated with both reproductive success and height. Average height men also married at a younger age than shorter and taller men, and the effect of height diminished after controlling for this association. Thus, average height men partly achieved higher reproductive success by marrying at a younger age. On the basis of our literature review and our data, we conclude that men of average height most likely have higher reproductive success than either short or tall men. Electronic supplementary material The online version of this article (doi:10.1007/s00265-011-1283-2) contains supplementary material, which is available to authorized users.

Long-term effects of unfavourable conditions during development can be expected to depend on the quality of the environment experienced by the same individuals during adulthood. Yet, in the majority of studies, long-term effects of early developmental conditions have been assessed under favourable adult conditions only. The immune system might be particularly vulnerable to early environmental conditions as its development, maintenance and use are thought to be energetically costly. Here, we studied the interactive effects of favourable and unfavourable conditions during nestling and adult stages on innate immunity (lysis and agglutination scores) of captive male and female zebra finches (Taeniopygia guttata). Nestling environmental conditions were manipulated by a brood size experiment, while a foraging cost treatment was imposed on the same individuals during adulthood. This combined treatment showed that innate immunity of adult zebra finches is affected by their early developmental conditions and varies between both sexes. Lysis scores, but not agglutination scores, were higher in individuals raised in small broods and in males. However, these effects were only present in birds that experienced low foraging costs. This study shows that the quality of the adult environment may shape the long-term consequences of early developmental conditions on innate immunity, as long-term effects of nestling environment were only evident under favourable adult conditions.

1. Between-individual variation in rates of senescence has recently been found to relate to natal and early-life conditions in several natural populations. Mechanistic theories of senescence have predicted between-individual variation in basal metabolic rate (BMR) to also underlie such variation in rates of senescence. The question whether variation in BMR is linked to natal and early-life conditions with effects on senescence has, however, not yet been addressed. 2. Using cross-sectional data on winter BMR of nearly 700 individual great tits Parus major, we tested whether factors associated with individual variation in the rate of senescence were also associated with differences in BMR. 3. We found that winter BMR was a repeatable trait (36%), and that variation in winter BMR was partly explained by body mass, and interactive effects of sex, age and seasonal date with ambient temperature. 4. Our data, however, revealed that neither the level of BMR, nor its age-specific decline, was explained by three parameters previously shown to underlie variation in rates of reproductive senescence in our study population: maternal age, immigrant status and early-life reproductive performance. 5. These results therefore did not support the suggestion that variation in BMR underlies variation in rates of reproductive senescence in relation to maternal age, immigrant status and early-life reproductive performance. 6. We further found a low repeatability of BMR from winter to the subsequent breeding season (1%, n = 55), which suggested that a relationship between BMR and proxies for rates of senescence may have been absent in our data set because winter BMR did not represent year-round metabolic strategies. 7. Further study, preferably using longitudinal data, is required to resolve the link between BMR at different times of year and senescence.

P>1. Evidence suggests that developing and maintaining an effective immune system may be costly and that an organism has to make a trade-off between immune function and other fitness-enhancing traits. To test for a trade-off between growth and immune function we carried out a meta-analysis of data from lines of poultry that had been divergently selected for either growth (body mass) or an aspect of immune function. This is relevant to our understanding of the evolution of immune function, but also because the increased prevalence of antibiotic-resistant bacteria and calls to restrict the use of antibiotics in the agricultural industry has made immune function of livestock an important theme. Has the selection of animals for rapid growth unintentionally resulted in reduced immune function?2. The lines selected for increased growth all showed a strong and significant decrease in immune function (standard difference in means = 0 center dot 8; P 3. Testing for publication bias revealed that the effect of selection for body mass on immune function was robust. However, there was considerable heterogeneity in both body mass and immune function data. The heterogeneity in the growth-selected lines cannot be accounted for by gender or species: the only turkey line had an effect size between that of the two chicken lines.4. In conclusion, we found that selection for growth does indeed compromise immune function, but selection for immune function did not consistently affect growth. This is in agreement with the supposition that the costs of growth are large relative to the costs of immune function, and on a practical level this suggests that it may be possible to breed animals for increased growth without loss of immune function.

Male zebra finches display multiple secondary sexual traits such as song and red bill coloration. This color is dependent on carotenoids, which enhance immune function and are antioxidants. A red bill may thus function as an indicator signal. The zebra finch is extensively used in the study of carotenoid-dependent signaling. However, studies of female mate preferences for redder bills show mixed results. Here, we report a meta-analysis of mate-choice studies that reveals that female zebra finches do prefer males with redder bills (r = 0.61), except when there was reduced opportunity for imprinting or when bill color was experimentally manipulated, which both reduced preference for red bills to approximately zero. The latter may either be due to aspects of the experimental design or due to bill color being correlated with another trait such as song rate as was previously suggested. We show, however, in a separate meta-analysis on a different set of studies that the correlation between bill coloration and song rate (r = 0.14) was significantly lower than the r = 0.61 between bill color and attractiveness. We conclude, therefore, that the role of bill coloration in mate choice cannot be solely due to an association with song rate. Thus, we conclude that females do prefer males with redder bills when there was sufficient opportunity for sexual imprinting, but to what extent this is causally related to the bill color remains to be established. Copyright 2011, Oxford University Press.

1. Longitudinal studies of various vertebrate populations have recently demonstrated senescent declines in reproductive performance and/or survival probability with age to be almost ubiquitous in nature. Little is known, however, about the extent to which rates of senescence vary between individuals, and about causes or consequences of such variation. Quantifying these links in natural populations is important for understanding the constraints and adaptive processes underlying the evolution of senescence. 2. Here, we analyse breeding data from 1029 female great tits Parus major to quantify the effect of natal conditions and early life reproduction on rates of reproductive senescence, reproductive life span and lifetime reproductive success. 3. Although for locally born females we find no evidence that natal conditions influence rates of reproductive senescence, we show that immigrant females suffer from faster rates of senescence than locally born females, and that this difference contributes to immigrants having a lower lifetime reproductive success. 4. Additionally, and independently, we find rates of reproductive senescence to increase with rates of early life reproduction, as predicted by the disposable soma hypothesis. Despite accelerated senescence late in life, high early life reproduction is, however, positively associated with lifetime reproductive success across individuals. 5. Female immigrant status and early life reproduction do not relate to reproductive life span. 6. We thus show that both immigration into our population, and high levels of early life reproduction, are associated with reduced late life reproductive performance in female great tits, but that fitness can be increased by high levels of early life reproduction at the expense of accelerated reproductive senescence. These results suggest disposable soma to be a likely mechanism underlying senescence in these birds, and encourage further study of the genetic basis (i.e. antagonistic pleiotropy) of such an early vs. late life trade-off.

Ageing, long thought to be too infrequent to study effectively in natural populations, has recently been shown to be ubiquitous, even in the wild. A major challenge now is to explain variation in the rates of ageing within populations. Here, using 49 years of data from a population of great tits (Parus major), we show that offspring life-history trajectories vary with maternal age. Offspring hatched from older mothers perform better early in life, but suffer from an earlier onset, and stronger rate, of reproductive senescence later in life. Offspring reproductive lifespan is, however, unaffected by maternal age, and the different life-history trajectories result in a similar fitness payoff, measured as lifetime reproductive success. This study therefore identifies maternal age as a new factor underlying variation in rates of ageing, and, given the delayed trans-generational nature of this effect, poses the question as to proximate mechanisms linking age-effects across generations.

Eurasian Oystercatchers Haematopus ostralegus ostralegus are a classic example of individual feeding specialization. Feeding specialization causes morphological differentiation in Oystercatchers' bill shapes due to varying degrees of abrasion associated with specific handling techniques for each prey species. Consequently, the Oystercatcher's bill shape has been used as a proxy for diet specialization, as it provides a quick and easy way to assess the diet choice of this marine top-predator. However, bill shapes of Oystercatchers are categorized visually in distinct types, while it has been argued that the relevant variation is continuous. Also, it is unclear how comparable the bill-shape classification is among studies and between the sexes and how universal bill shape-diet relationships are. Here we investigate the usefulness of bill-shape types as a proxy for diet choice in Oystercatchers, using four new and two published datasets. We show that quantitative bill-morphometrics provide no evidence that b...

Deterioration of reproductive traits with age is observed in an increasing number of species. Although such deterioration is often attributed to reproductive senescence, a within-individual decline in reproductive success with age, few studies on wild animals have focused on direct fitness measures while accounting for selective disappearance and terminal effects, and to our knowledge none have determined how senescence effects arise from underlying reproductive traits. We show for female great tits that such an approach helps understanding of the onset, impact and architecture of senescence. Cross-sectional analysis of 49 years of breeding data shows annual recruit production to decline from 3.5 years of age, this decline affecting 9 per cent of females each year. Longitudinal analyses, however, show that selective disappearance of poor-quality breeders partly masks senescence, which in fact starts at 2.8 years and affects 21 per cent of females each year. There is no evidence for abrupt terminal effects. Analyses of underlying traits show no deterioration in clutch size, but significant declines in brood size and fledgling number. Furthermore, these traits contribute −9, 12 and 39 per cent to the senescent decline in recruit production, respectively. Besides providing detailed knowledge of the patterns and architecture of senescence in a natural population, these results illustrate the importance of modelling individual variation, and facilitate study of the underlying mechanisms of senescence.

Molecular sexing of birds has been possible for over a decade, but for practical reasons many studies still use biometric data for sex discrimination. In some species, the sexes are easy to distinguish but sexual dimorphism is often more subtle, requiring the use of statistical analyses of biometric measurements to discriminate sexes. These models are usually parameterized and validated using data from a limited number of sites and years. However, the resulting discriminant functions are often applied to other populations and periods. A crucial, but usually untested, assumption is that sexual dimorphism does not vary in time and space. Here we illustrate the consequences of violation of this assumption in Eurasian Oystercatchers Haematopus ostralegus, a species for which most studies have relied on biometric sexing. Using biometric data from captures of known-sex birds, we show that sexual dimorphism varied substantially in time and even reversed in some months and years. Furthermore, some biometric traits used in sexing changed gradually over time, causing a reduction in sexual dimorphism. We show that the consequences of this variation on sex discrimination in Oystercatchers are subtle and easily overlooked, but can result in inaccurate and strongly male- or female-biased sex-ratio estimates. We recommend that biometric sexing should be avoided in Oystercatchers unless specific calibration for each month, year and area is carried out. This recommendation also applies to other species where biometric traits may depend on environmental conditions. We argue that this condition might apply to many bird species and therefore advise caution when interpreting results based on biometric sex discrimination.

Females often select mates on the basis of sexual signals, which can be reliable indicators of male quality when the costliness of these signals prevents cheating. The immunocompetence handicap hypothesis (ICHH) provides a mechanistic explanation of these costs, by proposing a trade-off between immune function and sexual displays. This trade-off arises because testosterone enhances sexual signals, but suppresses immune function. Many studies have investigated the ICHH by administrating testosterone, and a recent meta-analysis found little evidence that testosterone suppressed immune function. However, another component of the ICHH, which has received less empirical interest, suggests that there may also be an interaction in the other direction, with immune activation suppressing testosterone levels. We present a meta-analysis to test for this effect. Overall, there was a strong suppressive effect of experimental immune activation on testosterone levels ( r =−0.52), regardless of whether live pathogens or non-pathogenic antigens were used to challenge the immune system. The latter is important because it shows that immune activation per se suppresses testosterone levels. Thus, a trade-off between immunocompetence and sexual displays may primarily be generated by the effect of immune activation on testosterone, rather than the opposite effect that has received most attention.

Post-natal stress has been shown to have important short and long term effects on many adult traits in birds. During stress, metabolic alterations often result in the mobilization of energy away from energy-sensitive functions such as growth, which could have significant implications for developing animals. However, little is known about the implications of stress hormones for energy consumption in growing individuals. We experimentally increased corticosterone (CORT) levels in nestling zebra finches via oral administration, between the ages of 7 and 18 days. The standard metabolic rate (SMR) of birds was measured twice overnight when birds were between 11-13 and 55-65 days of age. Developmental CORT administration significantly elevated overnight variability in SMR (sd) in nestling birds (during the treatment period), but not at 55-65 days (5-6 weeks after the treatment period). The effect on variability was seen more prominently in birds from larger brood sizes and in females. We found no effects of our treatments on mean SMR overnight. However, brood size and sex had interactive effects, with males from larger brood sizes having higher SIVIR at 55-65 days of age. These results suggest that stress hormones can have significant effects on energy metabolism and possibly nocturnal arousal and sleep fragmentation. However, there were no detectable long term effects of our treatments on SIVIR, suggesting that these effects are only short-lived, in order to maintain homeostasis in the short term. (c) 2008 Elsevier Inc. All rights reserved.

We studied primary sex ratio of clutches in relation to social dominance for 6 years in a colony of free-living jackdaws, a small corvid. Social dominance was strongly associated with clutch sex ratio, with the difference in clutch sex ratio between the most and least dominant pairs being 30--40%. To our knowledge, this is the first demonstration of an association between social dominance and sex allocation in birds. However, the direction of this effect varied between years. Dominant jackdaws produced more sons during the first years of the study but fewer sons during the last years. Offspring sex was not related to laying order within a clutch, and the effect of social dominance on sex ratio was similar on eggs laid first, middle, or last. We investigated the effect of 2 factors (laying date and parental condition) that could have mediated the shift in the effect of social dominance on sex allocation in the course of the study. Laying date was positively associated with the proportion of males, but this effect was independent of social dominance. Maternal condition (residual mass over tarsus and egg volume) was related to social dominance but not to clutch sex ratio. Paternal condition (residual mass over tarsus) was not related to clutch sex ratio. We discuss how spatial or temporal variation in effects of variables such as social dominance on sex allocation can contribute to our understanding of the evolution of sex allocation in species with complex life histories. Copyright 2008, Oxford University Press.

Climate variability is changing on multiple temporal scales and little is known of the consequences of increases in short-term variability, particularly in endotherms. Using mortality data with high temporal resolution of zebra finches living in large outdoor aviaries (5 years, 359.220 bird-days), we show that mortality rate increases almost two-fold per 1°C increase in diurnal temperature range (DTR). Interestingly, the DTR effect differed between two groups with low versus high experimentally manipulated foraging costs, reflecting a typical laboratory ‘easy’ foraging environment and a ‘hard’ semi-natural environment respectively. DTR increased mortality on days with low minimum temperature in the easy foraging environment, but on days with high minimum temperature in the semi-natural environment. Thus, in a natural environment DTR effects will become increasingly important in a warming world, something not detectable in an ‘easy’ laboratory environment. These effects were particularly apparent at young ages. Critical time window analyses showed that the effect of DTR on mortality is delayed up to three months, while effects of minimum temperature occurred within a week. These results show that daily temperature variability can substantially impact the population viability of endothermic species.

Many migrating birds undertake extraordinary long flights. How birds are able to perform such endurance flights of over 100-hour durations is still poorly understood. We examined energy expenditure and physiological changes in Northern Bald Ibis Geronticus eremite during natural flights using birds trained to follow an ultra-light aircraft. Because these birds were tame, with foster parents, we were able to bleed them immediately prior to and after each flight. Flight duration was experimentally designed ranging between one and almost four hours continuous flights. Energy expenditure during flight was estimated using doubly-labelled-water while physiological properties were assessed through blood chemistry including plasma metabolites, enzymes, electrolytes, blood gases, and reactive oxygen compounds. Instantaneous energy expenditure decreased with flight duration, and the birds appeared to balance aerobic and anaerobic metabolism, using fat, carbohydrate and protein as fuel. This made flight both economic and tolerable. The observed effects resemble classical exercise adaptations that can limit duration of exercise while reducing energetic output. There were also in-flight benefits that enable power output variation from cruising to manoeuvring. These adaptations share characteristics with physiological processes that have facilitated other athletic feats in nature and might enable the extraordinary long flights of migratory birds as well.

Long-lived mutants of model organisms have brought remarkable progress in our understanding of aging mechanisms. However, long-lived mutants are usually maintained in optimal standardized laboratory environments (SLEs), and it is not obvious to what extent insights from long-lived mutants in SLEs can be generalized to more natural environments. To address this question, we reviewed experiments that compared the fitness and lifespan advantage of long-lived mutants relative to wild type controls in SLEs and more challenging environments in various model organisms such as yeast Saccharomyces cerevisiae, the nematode worm Caenorhabditis elegans, the fruitfly Drosophila melanogaster and the mouse Mus musculus. In competition experiments over multiple generations, the long-lived mutants had a lower fitness relative to wild type controls, and this disadvantage was the clearest when the environment included natural challenges such as limited food (N = 6 studies). It is well known that most long-lived mutants have impaired reproduction, which provides one reason for the fitness disadvantage. However, based on 12 experiments, we found that the lifespan advantage of long-lived mutants is diminished in more challenging environments, often to the extent that the wild type controls outlive the long lived mutants. Thus, it appears that information on aging mechanisms obtained from long-lived mutants in SLEs may be specific to such environments, because those same mechanisms do not extend lifespan in more natural environments. This suggests that different mechanisms cause variation in aging and lifespan in SLEs compared to natural populations. (C) 2015 Elsevier Inc. All rights reserved.

Telomere length predicts survival in birds, and many stressors that presumably reduce fitness have also been linked to telomere length. The response to selection of telomere length will be largely determined by the heritability of this trait; however, little is known about the genetic component of telomere length variation in animals other than humans. Moreover, published heritability estimates of telomere length are based on telomere measurements with techniques that do not distinguish between terminal telomeres, which are susceptible to age and stress, and the interstitial telomeric repeats, which are relatively inert. Heritability estimates that combine interstitial and terminal telomeres are difficult to interpret in species such as birds, where interstitial telomeres are often numerous. We estimated the heritability of terminal telomere length in a captive Zebra Finch population of cross-fostered (half-)siblings using data obtained with an electrophoresis technique that excludes the interstitial repeats from the measurements. We used both a Bayesian quantitative genetic 'animal' model and a frequentist sibling regression approach to estimate heritability. With the animal model, we estimated a high heritability of telomere length (h (2) = 0.99, 95 % credible interval = 0.87-1), but had insufficient statistical power to separate parental and permanent environment effects. The frequentist approach yielded similar heritability estimates, although with large confidence intervals. We used general linear mixed models to disentangle variance components of telomere length. The relative contributions of the individual, mother and father to telomere length variation were statistically indistinguishable at 23-31 %. Chicks were cross-fostered 4-days after hatching, and no effect of rearing nest was found, indicating that any undetected environmental effects exerted their influence prior to, or soon after, hatching. Thus, we conclude that telomere length resemblance between relatives is high and proportional to their relatedness, but we cannot conclusively distinguish between genetic and other forms of inheritance.

Across cultures, taller stature is linked to increased social status, but the potential reasons why this should be are unclear. One potential explanation is that taller individuals are more likely to win a dyadic confrontation with a competitor (i.e., they are more dominant), which leads to higher social rank. Although some previous studies have shown that perceptions of status or dominance are related to height, and are therefore consistent with such an explanation, there is surprisingly little research testing whether height actually has any influence on the behavioural outcomes in real-life social interactions. Here, we present three naturalistic observational studies demonstrating that height predicts interpersonal dominance during brief dyadic interactions. Study 1 investigated the likelihood of giving way in a narrow passage (N = 92); Study 2 investigated giving way in a busy shopping street, plus the likelihood of colliding with another individual (N = 1,108); and Study 3 investigated the likelihood of maintaining a linear path while walking, and potentially entering another individual's personal space (N = 1,056). We conclude that human height is positively related to interpersonal dominance, and may well contribute to the widely observed positive association between height and social status.

1. Long-term effects of conditions during early development on fitness are important for life history evolution and population ecology. Using multistrata mark-recapture models on 20 years of data, we quantified the relation between rearing conditions and lifetime fitness in a long-lived shorebird, the oystercatcher (Haematopus ostralegus). We addressed specifically the relative contribution of short- and long-term effects of rearing conditions to overall fitness consequences. 2. Rearing conditions were defined by differences in natal habitat quality, in which there is a clear dichotomy in our study population. In the first year of life, fledglings from high-quality natal origin had a 1.3 times higher juvenile survival. Later in life (age 3-11), individuals of high-quality natal origin had a 1.6 times higher adult prebreeder survival. The most striking effect of natal habitat quality was that birds that were reared on high-quality territories had a higher probability of settling in high-quality habitat (44% vs. 6%). Lifetime reproductive success of individuals born in high-quality habitat was 2.2 times higher than that of individuals born in low-quality habitat. This difference increased further when fitness was calculated over several generations, due to a correlation between the quality of rearing conditions of parents and their offspring. 3. Long-term effects of early conditions contributed more to overall fitness differences as short-term consequences, contrary to common conceptions on this issue. 4. This study illustrates that investigating only short-term effects of early conditions can lead to the large underestimation of fitness consequences. We discuss how long-term consequences of early conditions may affect settlement decisions and source-sink population interactions.

Variation in rearing conditions, due either to parental or to environmental quality, can result in offspring of different quality (e.g. body condition, immune function). However, evidence is accumulating that egg size and composition can also affect offspring quality. In Oystercatchers Haematopus ostralegus, high-quality rearing conditions result in a higher quantity as well as quality of offspring. This is thought to be caused by increased parental food provisioning to the chicks in high-quality environments. However, variation in egg quality between rearing conditions could also affect the quantity and quality of offspring. Determining the mechanism and ontogeny of quality differences is important in unravelling the causes of variation in reproductive success. To disentangle the effects of egg quality, and quality of the rearing conditions, on the future survival of offspring, we cross-fostered complete clutches between nests. When reared under conditions of similar environmental quality, chicks originating from eggs laid in low-quality environments survived as well as chicks originating from eggs laid in a high-quality environment. However, chicks reared in high-quality environments survived twice as long as chicks reared in low-quality environments, independent of the environmental quality in which the eggs were laid. This suggests that variation in the future survival of offspring is primarily caused by differences in environmental and/or parental quality, with no clear effect of egg quality (size).

Many studies have suggested that reproductive performance improves during the pair-bond, which might explain why individuals remate with the same partner in many species. However, discussion exists about whether the association between reproductive performance and pair-bond duration that is reported in these studies reflects a causal relationship. Usually it is unclear whether a positive association is caused by pairs improving during their pair-bond or by high-quality pairs staying together for longer. Furthermore, reproductive performance often also depends on the age or breeding experience of parents, which all covary with pair-bond duration. A much needed experimental approach is lacking so far. We investigated the effect of pair-bond duration on reproductive performance in a long-lived monogamous bird species based on natural as well as experimental variation. The duration of oystercatcher (Haematopus ostralegus) pair-bonds, which were followed for 21 years, strongly affected reproductive output, even after controlling for effects of age and breeding experience. Pairs improved during their pair-bond, and there were no indications of selective disappearance of low-quality pairs; however, pairs that stayed together for very long performed badly. Experimental removal of one partner showed that the reproductive cost of divorce depended on the pair-bond duration with the old partner. In addition, after remating, the newly formed pairs strongly improved again, independent of the age and breeding experience of the remated pair members. As such, this study provides the first experimental evidence of a causal effect of pairbond duration on reproductive performance. Key words: age-dependent reproduction, constraint, divorce, intrapair cooperation, long-term monogamy, restraint. [Behav Ecol]

Intake rate maximization alone is not always sufficient in explaining prey size selection in predators. For example, bivalve-feeding oystercatchers regularly select smaller prey than expected if they aimed to maximize their intake rate. It has been proposed that to these birds large prey are ‘‘risky,’’ in the sense that birds may damage their bills when feeding on large bivalves. Large bivalves yield more energy, but according to this hypothesis this is achieved at the expense of energy yield in the long term when (1) the risk of bill damage increases with prey size and (2) foraging with a damaged bill is less effective. In accordance with this hypothesis, we show that captive oystercatchers feeding on large cockles experienced a high probability of bill tip damage, while bill damage was absent when cockles were small. Moreover, among free-living oystercatchers the prevalence of bill damage was correlated with mean cockle size near the capture site, and the data on captive birds fit in this pattern. Food intake of captive oystercatchers feeding exclusively on cockles was reduced by 23% after bill damage, and free-living birds with damaged bills had 14 g lower mass. Because lower body mass was associated with higher mortality probability, these results indicate long-term costs associated with feeding on large cockles. We conclude that the risk of bill damage can potentially explain why oystercatchers avoid large bivalves and that oystercatchers may maximize long-term intake rate by selecting prey sizes that are ‘‘suboptimal’’ from a short-term rate-maximizing point of view. Key words: prey choice, prey profitability, prey size selection, risky prey. [Behav Ecol]

Background: In eutherian mammals and in humans, the female fetus may be masculinized while sharing the intra-uterine environment with a male fetus. Telomere length (TL), as expressed in leukocytes, is heritable and is longer in women than in men. The main determinant of leukocyte TL (LTL) is LTL at birth. However, LTL is modified by age-dependent attrition.Methods: We studied LTL dynamics (LTL and its attrition) in adult same-sex (monozygotic, n = 268; dizygotic, n = 308) twins and opposite-sex (n = 144) twins. LTL was measured by Southern blots of the terminal restriction fragments.Results: We observed that in same-sex (both monozygotic and dizygotic) twins, as reported in singletons, LTL was longer in females than in males [estimate +/- standard error (SE): 163 +/- 63 bp, PConclusions: These findings are compatible with masculinization of the female fetus in opposite-sex twins. They suggest that the sex difference in LTL, seen in the general population, is largely determined in utero, perhaps by the intrauterine hormonal environment. Further studies in newborn twins are warranted to test this thesis.

Sexual selection has resulted in a wide array of ornaments used in mate choice, and such indicator traits signal quality honestly when they bear costs, precluding cheating. Carotenoid-dependent coloration has attracted considerable attention in this context, because investing carotenoids in coloration has to be traded off against its physiological functions; carotenoids are antioxidants and increase immunocompetence. This trade-off is hypothesized to underlie the honesty of carotenoid-dependent coloration, signaling the handicap of allocating carotenoids away from somatic maintenance toward sexual display. Utilizing recent advances in modeling adaptive evolution, we used a comparative approach to investigate the evolution of plasma carotenoid levels using a species-level phylogeny of 178 bird species. We find that the evolutionary optimum for carotenoid levels is higher in lineages that evolved carotenoid-dependent coloration, with strong attraction toward this optimum. Hence, carotenoids do not appear to be limiting, given that higher carotenoid levels readily evolve in response to the evolution of carotenoid-dependent coloration. These findings challenge the assumption that carotenoids are a scarce resource and thus also challenge the hypothesis that physiological resource value of carotenoids underlies honesty of carotenoid-dependent traits. Therefore, the comparative evidence suggests that other factors, such as the acquisition and incorporation of carotenoids, are involved in maintaining signal honesty.

SUMMARY Knowledge of the physiological consequences of variation in food availability may be essential for understanding behavioural and life history responses to such variation. To study the physiological consequences of food availability animals are generally subjected to caloric restriction or starvation, thereby reducing the upper limit to the energy budget. The relevance of this approach to free-living animals is questionable, however,because under natural conditions low food availability often results in higher foraging costs, and everything else remaining equal this results in a higher energy budget. We manipulated food availability by varying the foraging costs and studied effects on daily energy expenditure (DEE) and energy allocation of captive starlings Sturnus vulgaris. Birds in a closed economy earned their food by flying between two perches 5 m apart. The probability of a reward was set at three different levels, thereby creating a `poor',`intermediate' and `rich' environment. Compared with the rich environment,birds flew 4 times more (2.3 h per day) in the poor environment, and increased DEE by 43% to 220 kJ day-1 (3.7×BMR), within the range of free-living parents rearing young. To our knowledge this is the first study to show an increase in DEE with decreasing food availability. Body mass, basal metabolic rate (BMR) and pectoral muscle size were reduced in the poor environment. Nocturnal energy expenditure was further reduced by reaching BMR earlier in the night. Calculations show that the energy demands in the poor environment could not be met with the flight costs of 20.5 W that we measured previously in a rich environment. Flight costs derived indirectly from the energy budget were lower, at 17.5 W, probably due to lower body mass. By reducing body mass by 20%, and economising during sleep, the birds achieved savings of 37% in their DEE. Without these savings, a DEE substantially higher than measured in free-living parents rearing young would be required to remain in energy balance. Surprisingly little data exist to verify whether free-living animals use the same tactics to survive periods with low food availability.

Birds rearing experimentally enlarged broods have lower antibody responses to a novel antigen, and we tested three hypotheses that could explain this result. We used zebra finches Taeniopygia guttata inoculated with sheep red blood cells (SRBC) as a study system, for which this trade-off was previously demonstrated. 1. Compensatory cellular immunity: The humoral immune response is slow, and removal of SRBC through up-regulated cellular immunity could pre-empt an antibody response. However, cellular immune response to PHA decreased with increasing brood size, allowing rejection of this hypothesis. 2. Costs of anti body-production: Chicks in large broods grow less well, and birds with large broods may allocate resources to chicks instead of antibodies when these are costly. Compared to saline controls, SRBC suppressed metabolic rate in the hours following immunisation, but there was no effect in the following night, or at any time 4 and 8 days later. Fitness costs were measured by repeatedly immunising parents with SRBC while rearing young. Chick growth, parental condition, and subsequent reproduction of the parents were not affected by SRBC. We conclude that the costs of antibody formation cannot explain the trade-off between brood size and antibody responsiveness. 3. Costs of immune system maintenance: Maintaining a system enabling antibody-formation may be very costly, and birds rearing large broods may have down-regulated this system. Based on this hypothesis we predicted that antibody formation would still be reduced in parents rearing large broods when immunised after rearing the chicks. Our results confirmed this prediction, and we suggest that birds rearing large broods have lower antibody responses because they economised on the maintenance costs of the immune system.

Biomarkers of aging are essential to predict mortality and aging-related diseases. Paradoxically, age itself imposes a limitation on the use of known biomarkers of aging because their associations with mortality generally diminish with age. How this pattern arises is, however, not understood. With meta-analysis we show that human leucocyte telomere length (TL) predicts mortality, and that this mortality association diminishes with age, as found for other biomarkers of aging. Subsequently, we demonstrate with simulation models that this observation cannot be reconciled with the popular hypothesis that TL is proportional to biological age. Using the reliability theory of aging, we instead propose that TL is a biomarker of somatic redundancy, the body's capacity to absorb damage, which fits the observed pattern well. We discuss to what extent diminishing redundancy with age may also explain the observed diminishing mortality modulation with age of other biomarkers of aging. Considering diminishing somatic redundancy as the causal agent of aging may critically advance our understanding of the aging process, and improve predictions of life expectancy and vulnerability to aging-related diseases.

SUMMARYAlthough most birds are accustomed to making short flights, particularly during foraging, the flight patterns during these short periods of activity differ between species. Nectarivorous birds, in particular, often spend time hovering, while non-nectarivorous birds do not. The cost of short flights is likely therefore to differ between nectarivorous and non-nectarivorous birds because of the different energetic contributions of different flight types to the behaviour. The 13C-labelled bicarbonate technique was used to measure the energy cost of short flights in the nectarivorous Palestine sunbird Nectarinia osea (mean mass 6.17±0.16 g, N=8)and the non-nectarivorous starling Sturnus vulgaris (mean mass 70.11±1.11 g, N=9). The technique was initially calibrated in five individuals for each species at temperatures ranging from 1 to 35°C,by comparing the isotope elimination rate to the metabolic rate measured simultaneously by indirect calorimetry. The cost for short intermittent flight was then measured by encouraging birds to fly between two perches at either end of a narrow corridor (perch distance for sunbirds, 6 m; for starlings, 5 m), and measuring the amount of isotope eliminated during the flight. The isotope elimination rate was interpolated onto the calibration equation to predict flight cost, as a direct calibration could not be performed during flight. Mean energy expenditure during flight was 1.64±0.32 W in sunbirds, while in starlings the flight costs averaged 20.6±0.78 W. Energy cost of flight relative to basal metabolic rate was substantially greater in the starling than the sunbird. Phylogenetic analysis of different modes of flight in these and additional species suggests that differences in flight behaviour may cause these elevated costs in slow flying non-nectarivores such as starlings, compared to birds that are more prone to short intermittent flights like the sunbirds.

Social dominance is intuitively assumed to be associated with higher fitness, because social dominance implies better access to resources. We found that, in a colony of jackdaws, the dominant males consistently produced fewer fledglings, which had lower chances of survival to 1 year of age. Laying date and clutch size were independent of dominance, but females that mated with dominant males were in poorer condition and laid smaller eggs. Parental survival was independent of social dominance, and the frequency of extrapair fertilizations in jackdaws is negligible. Dominance was a stable trait of individuals, and not a state that all individuals eventually attained. We conclude that, in this colony, dominant jackdaws had lower fitness. To our knowledge, this is the first example of such a pattern in a free-living species. We hypothesize that the high density of our colony resulted in high testosterone titres, which suppressed paternal care of mate and offspring to the extent that it outweighed the benefits of higher resource access. (C) 2004 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Oxidative metabolism has reactive oxygen species (ROS) as unavoidable by-products, and the damage ROS inflicts on DNA, proteins and lipids is considered to be a major agent of senescence. Increasing reproductive effort accelerates senescence, but whether reproductive effort is increased at the expense of protection against oxidative damage has not yet been tested. We manipulated reproductive effort in zebra finches through brood size manipulation and measured the activity of two major antioxidant enzymes (superoxide dismutase (SOD) and glutathione peroxidase (GPx)) in the pectoral muscle after 19-20 days of brood rearing. Oxidative stress is reflected by the balance between oxidative protection and ROS exposure, and we therefore scaled SOD and GPx activity to daily energy expenditure (DEE) as an index of ROS production. SOD and GPx activity decreased with increasing brood size by 28% and 24%, respectively. This effect was identical in the two sexes, but arose in different ways: males did not change their DEE, but had lower absolute enzyme activity, and females increased their DEE, but did not change absolute enzyme activity. This result suggests that senescence acceleration by increased reproductive effort is at least in part mediated by oxidative stress.

Sexual conflict occurs when the optimal solution regarding e.g. a life history trait differs between co-operating individuals of different sex. When deciding a conflict is not instantaneous, some form of negotiation can be expected to evolve. In great tits, Parus major, a sexual conflict exists over the number of clutches that are reared, because the fitness costs of a second clutch are greater for females. A conflict is also likely to exist over investment in the first brood - each parent benefiting from a greater investment by the partner. Male great tits sing when rearing the first brood, and if acoustic signals play a role in the negotiation of a sexual conflict, a positive association between male song rate and maternal investment is predicted. In agreement with this hypothesis, maternal effort (in kJ/day) relative to paternal effort was positively correlated with male song rate. Furthermore, females were more likely to start a second clutch when their male had a high song rate, and high song rate was associated with shorter inter-clutch intervals. Song rate was higher when brood size was experimentally reduced and, independent of brood size manipulation, males with high song rate produced higher quality fledglings. These results indicate that song rate reflects the males' state, suggesting it may function as a handicap signal. Although song rate seems too low (

Food availability modulates survival in interaction with (for example) competition, disease and predators, but to what extent food availability in natural populations affects survival independent of these factors is not well known. We tested the effect of food availability on lifespan and actuarial senescence in a large population of captive zebra finches by increasing the effort required to obtain food, reflecting natural contrasts in food availability. Food availability may not affect all individuals equally and we therefore created heterogeneity in phenotypic quality by raising birds with different numbers of siblings. Low food availability had no effect on lifespan for individuals from benign developmental conditions (raised in small broods), but shortened lifespan for individuals from harsh developmental conditions. The lifespan difference arose through higher baseline mortality rate of individuals from harsh developmental conditions, despite a decrease in the rate of actuarial senescence. We found no evidence for sex-specific environmental sensitivity, but females lived shorter than males due to increased actuarial senescence. Thus, low food availability by itself shortens lifespan, but only in individuals from harsh developmental conditions. Our food availability manipulation resembles dietary restriction as applied to invertebrates, where it extends lifespan in model organisms and we discuss possible reasons for the contrasting results.

The vitamin A-redox hypothesis provides an explanation for honest signaling of phenotypic quality by carotenoid-dependent traits. A key aspect of the vitamin A-redox hypothesis, applicable to both yellow and red coloration, is the hypothesized negative feedback of tightly regulated Vitamin A plasma levels on the enzyme responsible for sequestering both Vitamin A and carotenoids from the gut. We performed a meta-analysis and find that vitamin A levels are positively related to carotenoid plasma levels (r = 0.50, P = 0.0002). On the basis of this finding and further theoretical considerations, we propose that the vitamin A-redox hypothesis is unlikely to explain carotenoid-dependent honest signaling.

Personality and metabolic rate are predicted to show covariance on methodological and functional grounds, but empirical studies at the individual level are rare, especially in natural populations. Here we assess the relationship between exploration behaviour, an important axis of personality, and basal metabolic rate (BMR) for 680 free-living great tits Parus major, studied over three years. We find that exploration behaviour is weakly negatively related to BMR among female, but not male, birds. Moreover, we find exploration behaviour to be independent of methodological aspects of BMR measurements (e.g. activity levels, time to acclimatize) which have been suggested to be indicative of personality-related activity or stress levels during measurement. This suggests that the weak link between exploration behaviour and BMR found here is functional rather than methodological. We therefore test the hypothesis that selection favours covariance between exploration behaviour and metabolic rate, but find no evidence for correlational survival or fecundity selection. Our data therefore provide at best only very weak evidence for a functional link between personality and metabolic rate, and we suggest that studies of personality and metabolic strategies, or personality and daily energy expenditure, are required to further resolve the link between personality and metabolic rate. © 2013 The Authors.

Sexual selection theory posits that females should choose mates in a way that maximises their reproductive success. But what exactly is the optimal choice? Most empirical research is based on the assumption that females seek a male of the highest possible quality (in terms of the genes or resources he can provide), and hence show directional preferences for indicators of male quality. This implies that attractiveness and quality should be highly correlated. However, females frequently differ in what they find attractive. New theoretical and empirical insights provide mounting evidence that a female's own quality biases her judgement of male attractiveness, such that male quality and attractiveness do not always coincide. A recent experiment in songbirds demonstrated for the first time that manipulation of female condition can lead to divergent female preferences, with low-quality females actively preferring low-quality males over high-quality males. This result is in line with theory on state-dependent mate choice and is reminiscent of assortative mating preferences in humans. Here we discuss the implications of this work for the study of mate preferences.

1. To elucidate the links between avian brood size, parental effort and parental investment, we measured daily energy expenditure (DEEfem), condition (residuals of mass on tarsus) and feeding rate in female great tits Parus major L. rearing broods in which the number of young was either reduced, unmanipulated or enlarged. 2. Female condition was negatively correlated with manipulation when measured at the nestling age of 8 days (measured during the day), which suggests a shift in allocation from self-feeding to chick-feeding. However, there was no detectable manipulation effect on condition measured at the nestling age of 12 days (measured during the night). Either female condition was only affected by manipulation in the early nestling phase or the females adjusted their diurnal mass trajectory in response to brood size manipulation. More detailed data are required to verify this point. There were no indications of a fitness cost associated with the condition during the day, but condition at night was positively related to winter survival. Since manipulation only affected condition during the day, there was no link between manipulation and winter survival. 3. The duration of the working day was not affected by manipulation and female feeding rate tended to flatten off with manipulated brood size. Similarly, brood reduction resulted in a lower DEEfem, whilst brood enlargement had no effect. This suggests that females worked at an energetic ceiling when rearing an unmanipulated brood. However, the level of this 'ceiling' in DEEfem was not fixed: it differed between years. This leads us to conclude that the observed ceiling was imposed by extrinsic factors (e.g. available foraging time) and not by an intrinsic factor such as maximum energy assimilation rate. We hypothesize that time limitation was the cause for the observed ceiling in energy expenditure and that the annual variation in the level of this ceiling was due to annual variation in ambient temperature. 4. A cost of reproduction was previously demonstrated in this population: brood enlargement caused a reduction in the incidence of second clutches. However, since DEEfem did not differ between control and enlarged broods, we judge it unlikely that daily energy expenditure is a general predictor for parental investment. [KEYWORDS: clutch size; condition; energy expenditure; cost of reproduction; parental effort; Parus major L. Starlings sturnus-vulgaris; kestrel falco-tinnunculus; parus-major; brood size; clutch size; blue tits; food availability; feeding frequency; field metabolism; trade-offs]

Western scrub-jays ( Aphelocoma californica ) engage in a variety of cache-protection strategies to reduce the chances of cache theft by conspecifics. Many of these strategies revolve around reducing visual information to potential thieves. This study aimed to determine whether the jays also reduce auditory information during caching. Each jay was given the opportunity to cache food in two trays, one of which was filled with small pebbles that made considerable noise when cached in (‘noisy’ tray), whereas the other one contained soil that made little detectable noise when cached in (‘quiet’ tray). When the jays could be heard, but not seen, by a competitor, they cached proportionally less food items in the ‘noisy’ substrate than when they cached alone in the room, or when they could be seen and heard by competitors. These results suggest that western scrub-jays know when to conceal auditory information, namely when a competitor cannot see but can hear the caching event.

Females often select their mates on the basis of the size or intensity of sexual ornaments, and it is thought that such traits are reliable indicators of male quality because the costliness of these traits prevents cheating. The immunocompetence handicap hypothesis is a recently proposed mechanistic explanation of these costs and states that males carry ornaments at the expense of their resistance to disease and parasites. The tradeoff between immunocompetence and sexual ornamentation was hypothesized to arise as a consequence of the dual effect of androgens on ornamentation (+) and immune function (−). To test this hypothesis, we compared comb size between male domestic chickens Gallus domesticus of lines divergently selected for antibody responses to sheep erythrocytes (three lines: selected for low response or high response and a control line). The importance of comb size in inter- and intrasexual selection is well established, and comb size is strongly dependent on testosterone level. Comb size was larger in the males of the low line than in the high line, and comb size of control males was intermediate, indicating a tradeoff between ornamentation and immunocompetence. Testosterone (T) levels varied in a similar fashion (T Low > T Control > T High ), suggesting that this hormone could mediate the tradeoff between ornamentation and immunocompetence. These results support the idea that a tradeoff with immune function may constrain the expression of secondary sexual ornaments.