Crosstalk between growth and somatic maintenance in young animals

Growing animals face allocation problems whenever receiving suboptimal amounts of resources in very stochastic natural environments, possibly through a trade‐off between growth and somatic maintenance. However, the extent to which such a trade‐off exists has remained an open question. We used an insulin‐like growth factor 1 (IGF‐1) injection treatment in free‐living pied flycatcher Ficedula hypoleuca nestlings to see how IGF‐1 levels mediate the development of an antioxidant phenotype via glutathione peroxidase (GPx). Our study showed that IGF‐1 levels underlie variation in GPx activity and provides important information for understanding the mechanisms behind the growth variation of passerines.     

Mediation of vertebrate life histories via insulin‐like growth factor‐1

Life‐history traits describe parameters associated with growth, size, survival, and reproduction. Life‐history variation is a hallmark of biological diversity, yet researchers commonly observe that one of the major axes of life‐history variation after controlling for body size involves trade‐offs among growth, reproduction, and longevity. This persistent pattern of covariation among these specific traits has engendered a search for shared mechanisms that could constrain or facilitate production of variation in life‐history strategies. Endocrine traits are one candidate mechanism that may underlie the integration of life history and other phenotypic traits. However, the vast majority of this research has been on the effects of steroid hormones such as glucocorticoids and androgens on life‐history trade‐offs. Here we propose an expansion of the focus on glucocorticoids and gonadal hormones and review the potential role of insulin‐like growth factor‐1 (IGF‐1) in shaping the adaptive integration of multiple life‐history traits. IGF‐1 is a polypeptide metabolic hormone largely produced by the liver. We summarize a vast array of research demonstrating that IGF‐1 levels are susceptible to environmental variation and that IGF‐1 can have potent stimulatory effects on somatic growth and reproduction but decrease lifespan. We review the few studies in natural populations that have measured plasma IGF‐1 concentrations and its associations with life‐history traits or other characteristics of the organism or its environment. We focus on two case studies that found support for the hypothesis that IGF‐1 mediates adaptive divergence in suites of life‐history traits in response to varying ecological conditions or artificial selection. We also examine what we view as potentially fruitful avenues of research on this topic, which until now has been rarely investigated by evolutionary ecologists. We discuss how IGF‐1 may facilitate adaptive plasticity in life‐history strategies in response to early environmental conditions and also how selection on loci controlling IGF‐1 signaling may mediate population divergence and eventual speciation. After consideration of the interactions among androgens, glucocorticoids, and IGF‐1 we suggest that IGF‐1 be considered a suitable candidate mechanism for mediating life‐history traits. Finally, we discuss what we can learn about IGF‐1 from studies in free‐ranging animals. The voluminous literature in laboratory and domesticated animals documenting relationships among IGF‐1, growth, reproduction, and lifespan demonstrates the potential for a number of new research questions to be asked in free‐ranging animals. Examining how IGF‐1 mediates life‐history traits in free‐ranging animals could lead to great insight into the mechanisms that influence life‐history variation.     

Food availability modulates temperature‐dependent effects on growth,reproduction, and survival in Daphnia magna

Reduced body size and accelerated life cycle due to warming are considered major ecological responses to climate change with fitness costs at the individual level. Surprisingly, we know little about how relevant ecological factors can alter these life history trade‐offs and their consequences for individual fitness. Here, we show that food modulates temperature‐dependent effects on body size in the water flea Daphnia magna and interacts with temperature to affect life history parameters. We exposed 412 individuals to a factorial manipulation of food abundance and temperature, tracked each reproductive event, and took daily measurements of body size from each individual. High temperature caused a reduction in maximum body size in both food treatments, but this effect was mediated by food abundance, such that low food conditions resulted in a reduction of 20% in maximum body size, compared with a reduction of 4% under high food conditions. High temperature resulted in an accelerated life cycle, with pronounced fitness cost at low levels of food where only a few individuals produced a clutch. These results suggest that the mechanisms affecting the trade‐off between fast growth and final body size are food‐dependent, and that the combination of low levels of food and high temperature could potentially threaten viability of ectotherms.     

A genotypic trade‐off between the number and size of clonal offspring in the stoloniferous herb Potentilla reptans

A negative, genetic correlation between the total number and average size of progeny is a classical life‐history trade‐off that can greatly affect the fitness of organisms in their natural environments. This trade‐off has been investigated for animals and for sexually reproducing plants. However, evidence for a genetical size‐number trade‐off for clonal progeny in plants is still scarce. This study provides experimental evidence for such a trade‐off in the stoloniferous herb Potentilla reptans, and it studies phenotypic plasticity to light availability for the involved traits. Genotypes of P. reptans were collected from distinctively different environments, clonally replicated and exposed to high light and to shaded conditions. We found a significant negative correlation between the average size and the total number of offspring across genotypes for both light environments. Shading reduced ramet numbers, but hardly affected average ramet size.     

Malaria infection negatively affects feather growth rate in the house sparrow Passer domesticus

Birds often face various stressors during feather renewing, for example, enduring infection with blood parasites. Because nutritional resources are typically limited, especially for wild animals, when an individual allocates energy to one physiological system, there is subsequently less for other processes, thereby requiring a trade‐off. Surprisingly, potential trade‐offs between malaria infection and feather growth rate have not been experimentally considered yet. Here, we conducted three studies to investigate whether a trade‐off occurs among feather growth rate, malaria infection and host health conditions. First, we explored whether naturally infected and uninfected house sparrows differed in feather growth rate in the wild. Second, we asked whether experimental inoculation of malaria parasites and/or forcing the renewal of a tail feather. Lastly, we evaluated whether individual condition was affected by experimentally‐induced feather regrowth and/or malaria experimental infection. Our findings showed that feather growth rate was negatively affected by natural malaria infection status in free‐living birds and by experimental infection in captive birds. Furthermore, birds that did not increase body mass or hematocrit during the experimental study had slower feather growth. Together our results suggest that infection with blood parasites has more negative health effects than the growth of tail feathers and that these two processes (response to blood parasite infection and renewal of feathers) are traded‐off against each other. As such, our results highlight the role of malaria parasites as a potential mechanism driving other trade‐offs in wild passerines.     

The effect of Helicobacter pylori and economic status on growth parameters and leptin, ghrelin, and insulin-like growth factor (IGF)-I concentrations in children

Background: It was suggested that gastric colonization with Helicobacter pylori (H. pylori) was associated with suboptimal nutrition and growth in childhood. Furthermore, several studies indicated a relationship between H. pylori colonization and alterations in the circulating levels of growth‐related molecules (GRM). Accordingly, in this study, we investigate the effect of H. pylori infection on GRMs and on the growth of healthy school children, taking into consideration the effect of their economic status (ES) and anthropometric indices of their parents. Methods: To acquire sociodemographic and anthropometric nutritional parameters and to detect H. pylori‐specific serum IgG antibodies and growth‐related molecules, we evaluated a total of 473 children attending four different primary and secondary schools in Istanbul. Subsequently, we assessed the effect of H. pylori on growth‐related parameters (weight for age SDS, height for age SDS, BMI SDS, TSF, and waist‐to‐hip ratio) and on GRMs (leptin, ghrelin, and insulin‐like growth factor‐1 (IGF‐1)), controlling for age, gender, family income, household crowding (HC), breastfeeding, maternal and paternal BMI SDS, and midparental height SDS with complex statistical models. Results: Of the 473 children (275 F/198 M, age 6–15 years; mean: 10.3 ± 0.1 years), 161 (34%) were H. pylori‐positive. The prevalence of H. pylori was significantly higher in lower economic status (ES) groups, in children living in crowded houses, and in older age groups. Using simple statistical models, we did not find any significant associations between H. pylori infection and the growth parameters. However, in complex models for height for age SDS and for weight for age SDS, there was a significant interaction between H. pylori infection status and ES. Whereas in H. pylori‐positive subjects, mid‐income family children were both taller and heavier than the low‐income group, there was no such an association in H. pylori‐negative subjects. Among biochemical parameters, only ghrelin levels were associated with H. pylori infection in all models. Leptin levels were associated with HC in girls, whereas none of the parameters was significantly associated with leptin levels in boys. For IGF‐1 levels, for boys, age and maternal BMI, and for girls, age and HC were significantly associated with IGF‐1 levels. Conclusion: We suggest that H. pylori may impair growth significantly only in susceptible children where unfavorable socioeconomic conditions facilitate its action, probably through mechanisms, at least in part, involving growth‐related molecules.     

Relationship of plasma adiponectin with sex hormone and insulin-like growth factor levels

Objective: Recent studies have suggested that a relationship between adiponectin and sex hormone, prolactin, and insulin‐like growth factor levels could be important for breast cancer risk and insulin sensitivity. Therefore, we assessed the relationship of adiponectin with plasma concentrations of estrone; estradiol; estrone sulfate; testosterone; androstenedione; dehydroepiandrosterone (DHEA); dehydroepiandrosterone sulfate (DHEAS); sex hormone binding globulin (SHBG); prolactin; insulin‐like growth factor (IGF‐1); its binding protein, IGF binding protein 3 (IGFBP‐3); c‐peptide; and IGF binding protein 1 (IGFBP‐1) among 360 postmenopausal women not taking postmenopausal hormones from the Nurses’ Health Study. Research Methods and Procedures: Multivariate models were adjusted for physical activity, alcohol consumption, age at blood draw, age at first birth/parity, fasting status, and time of day of blood draw; a separate model was additionally adjusted for BMI at blood draw. Results: Estrogens were inversely associated with adiponectin levels; however, except for free estradiol, these associations were substantially attenuated after adjustment for BMI. Free estradiol levels were 27% lower among women in the top vs. bottom quartile of adiponectin levels. No consistent associations were observed for the androgens, prolactin, IGF‐1, and IGFBP‐3. However, SHBG, c‐peptide, and IGFBP‐1 were strongly and independently associated with adiponectin levels (r = 0.29, ?0.30, 0.24, respectively). Conclusion: With the exceptions of SHBG, c‐peptide, and IGFBP‐1, the studied analytes were modestly associated with adiponectin and the associations were, in large part, mediated by body fat.     

Stress promotes changes in resource allocation to growth and reproduction in a simultaneous hermaphrodite with indeterminate growth

In iteroparous animals, investment in growth is compromised by investment in reproduction, especially in species with indeterminate growth. Life‐history theory predicts that growth should be favoured over reproduction, assuming size‐related fecundity or survival. Hence, increase body condition represents an increase in reproductive potential. Simultaneous hermaphrodites should adjust their resource allocation to each sex function in response to current conditions but, recently, it has been suggested that, in hermaphrodites, gender allocation should be considered as a three‐way trade‐off, including the investment in somatic growth. Due to the higher costs involved, the female function is affected to a greater extent by environmentally stressful conditions rather than the male function. To examine this, we induced stress in the hermaphroditic earthworm Eisenia fetida (Savigny, 1826) and looked for changes in resource allocation in nonreproductive and reproductive individuals. Experimental stress was induced by using tweezers to elicit contractile escape movements. We predicted that stressed earthworms would preferentially allocate resources to growth. In nonreproductive individuals, however, stress had a negative effect on growth, although weight recovery was rapid once manipulation ceased, indicating the importance of body condition, as well as the existence of mechanisms of compensatory growth for growth trajectories in this earthworm species. The response of reproductive individuals was consistent with our expectation: (1) stressed worms maintained their growth rate at the expense of current reproduction and (2) stressed earthworms laid 25% fewer cocoons, which were 30% lighter than cocoons laid by control earthworms. The present results suggest that E. fetida regulates its reproductive effort and that future reproduction has more impact on its fitness than current reproduction. The trade‐off between current and future reproduction should be taken into consideration in models of sex allocation in simultaneous hermaphrodites. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 91 , 593–600.     

Finite element modeling of shell shape in the freshwater turtle Pseudemys concinna reveals a trade‐off between mechanical strength and hydrodynamic efficiency

Aquatic species can experience different selective pressures on morphology in different flow regimes. Species inhabiting lotic regimes often adapt to these conditions by evolving low‐drag (i.e., streamlined) morphologies that reduce the likelihood of dislodgment or displacement. However, hydrodynamic factors are not the only selective pressures influencing organismal morphology and shapes well suited to flow conditions may compromise performance in other roles. We investigated the possibility of morphological trade‐offs in the turtle Pseudemys concinna. Individuals living in lotic environments have flatter, more streamlined shells than those living in lentic environments; however, this flatter shape may also make the shells less capable of resisting predator‐induced loads. We tested the idea that “lotic” shell shapes are weaker than “lentic” shell shapes, concomitantly examining effects of sex. Geometric morphometric data were used to transform an existing finite element shell model into a series of models corresponding to the shapes of individual turtles. Models were assigned identical material properties and loaded under identical conditions, and the stresses produced by a series of eight loads were extracted to describe the strength of the shells. “Lotic” shell shapes produced significantly higher stresses than “lentic” shell shapes, indicating that the former is weaker than the latter. Females had significantly stronger shell shapes than males, although these differences were less consistent than differences between flow regimes. We conclude that, despite the potential for many‐to‐one mapping of shell shape onto strength, P. concinna experiences a trade‐off in shell shape between hydrodynamic and mechanical performance. This trade‐off may be evident in many other turtle species or any other aquatic species that also depend on a shell for defense. However, evolution of body size may provide an avenue of escape from this trade‐off in some cases, as changes in size can drastically affect mechanical performance while having little effect on hydrodynamic performance. J. Morphol. 2011. © 2011 Wiley‐Liss, Inc.     

Ecophysiology of cognition: How do environmentally induced changes in physiology affect cognitive performance?

Cognitive performance is based on brain functions, which have energetic demands and are modulated by physiological parameters such as metabolic hormones. As both environmental demands and environmental energy availability change seasonally, we propose that cognitive performance in free‐living animals might also change seasonally due to phenotypic plasticity. This is part of an emerging research field, the ‘ecophysiology of cognition’: environmentally induced changes in physiological traits, such as blood glucose and hormone levels, are predicted to influence cognitive performance in free‐living animals. Energy availability for the brain might change, and as such cognition, with changing energetic demands (e.g. reproduction) and changes of energy availability in the environment (e.g. winter, drought). Individuals spending more energy than they can currently obtain from their environment (allostatic overload type I) are expected to trade off energy investment between cognition and other life‐sustaining processes or even reproduction. Environmental changes reducing energy availability might thus impair cognition. However, selection pressures such as predation risk, mate choice or social demands may act on the trade‐off between energy saving and cognition. We assume that different environmental conditions can lead to three different trade‐off outcomes: cognitive impairment, resilience or enhancement. Currently we cannot understand these trade‐offs, because we lack information about changes in cognitive performance due to seasonal changes in energy availability and both the resulting changes in homeostasis (for example, blood glucose levels) and the associated changes in the mechanisms of allostasis (for example, hormone levels). Additionally, so far we know little about the fitness consequences of individual variation in cognitive performance. General cognitive abilities, such as attention and associative learning, might be more important in determining fitness than complex and specialized cognitive abilities, and easier to use for comparative study in a large number of species. We propose to study seasonal changes in cognitive performance depending on energy availability in populations facing different predation risks, and the resulting fitness consequences.     

Effect of weight loss on free insulin-like growth factor-I in obese women with hyposomatotropism

Objective: It has been hypothesized that increased free insulin‐like growth factor (IGF)‐I levels generated from an increase in IGF‐binding protein (IGFBP) protease activity could be the inhibitory mechanism for the decreased growth hormone (GH) secretion observed in obese subjects. Research Methods and Procedures: In this study, we determined basal and 24‐hour levels of free IGF‐I and ‐II, total IGF‐I and ‐II, IGFBP‐1, as well as basal IGFBP‐2, ?3, and ?4, acid‐labile subunit (ALS), IGFBP‐1, ?2, and ?3 protease activity, and 24‐hour GH release in obese women before and after a diet‐induced weight loss. Sixteen obese women (age, 29.5 ± 1.4 years) participated in a weight loss program and 16 age‐matched non‐obese women served as controls. Results: Circulating free IGF‐I and 24‐hour GH release were significantly decreased in obese women at before weight loss compared with non‐obese women (1.29 ± 0.12 vs. 0.60 ± 0.09 μg/L; p < 0.001 and 862 ± 90 vs. 404 ± 77 mU/24 hours; p < 0.001, respectively). Free IGF‐I and 24‐hour GH release were not inversely correlated to each other. IGFBP‐1 and ?2 levels were decreased, whereas ALS, IGFBP‐3 and ?4, and IGFBP‐1, ?2, and ?3 protease activity were similar in obese and non‐obese women. Eight of the 16 obese women achieved an average weight loss of 30 ± 5 kg during 26 to 60 weeks of dieting. After the considerable weight loss, significant differences in free IGF‐I, GH release, and IGFBP‐1 and ?2 levels were no longer present between previously obese and non‐obese women. Discussion: We showed that circulating free IGF‐I is markedly decreased in severely obese women and does not per se mediate the concomitant hyposomatotropism. The decreased levels of free IGF‐I seem to be transient and restored to normal levels after weight loss.     

Maternal investment increases with altitude in a frog on the Tibetan Plateau

Reproducing females can allocate energy between the production of eggs or offspring of different size or number, both of which can strongly influence fitness. The physical capacity to store developing offspring imposes constraints on maximum clutch volume, but individual females and populations can trade off whether more or fewer eggs or offspring are produced, and their relative sizes. Harsh environments are likely to select for larger egg or offspring size, and many vertebrate populations compensate for this reproductive investment through an increase in female body size. We report a different trade‐off in a frog endemic to the Tibetan Plateau, Rana kukunoris. Females living at higher altitudes (n = 11 populations, 2000–3500 m) produce larger eggs, but without a concomitant increase in female body size or clutch size. The reduced diel and seasonal activity at high altitudes may impose constraints on the maximum body size of adult frogs, by limiting the opportunity for energy accumulation. Simultaneously, producing larger eggs likely helps to increase the rate of embryonic development, causing tadpoles to hatch earlier. The gelatinous matrix surrounding eggs, more of which is produced by large females, may help buffer developing embryos from temperature fluctuations or offer protection from ultraviolet radiation. High‐altitude frogs on the Tibetan Plateau employ a reproductive strategy that favours large egg size independent of body size, which is unusual in amphibians. The harsh and unpredictable environmental conditions at high altitudes can thus impose strong and opposing selection pressures on adult and embryonic life stages, both of which can simultaneously influence fitness.     

Optimal growth strategies under divergent predation pressure

The conditions leading to gigantism in nine‐spined sticklebacks Pungitius pungitius were analysed by modelling fish growth with the von Bertalanffy model searching for the optimal strategy when the model's growth constant and asymptotic fish size parameters are negatively related to each other. Predator‐related mortality was modelled through the increased risk of death during active foraging. The model was parameterized with empirical growth data of fish from four different populations and analysed for optimal growth strategy at different mortality levels. The growth constant and asymptotic fish size were negatively related in most populations. Optimal fish size, fitness and life span decreased with predator‐induced mortality. At low mortality, the fitness of pond populations was higher than that of sea populations. The differences disappeared at intermediate mortalities, and sea populations had slightly higher fitness at extremely high mortalities. In the scenario where all populations mature at the same age, the pond populations perform better at low mortalities and the sea populations at high mortalities. It is concluded that a trade‐off between growth constant and asymptotic fish size, together with different mortality rates, can explain a significant proportion of body size differentiation between populations. In the present case, it is a sufficient explanation of gigantism in pond P. pungitius.     

Impacts of environmental variability on desiccation rate,plastic responses and population dynamics of Glossina pallidipes

Physiological responses to transient conditions may result in costly responses with little fitness benefits, and therefore, a trade‐off must exist between the speed of response and the duration of exposure to new conditions. Here, using the puparia of an important insect disease vector, Glossina pallidipes, we examine this potential trade‐off using a novel combination of an experimental approach and a population dynamics model. Specifically, we explore and dissect the interactions between plastic physiological responses, treatment‐duration and ‐intensity using an experimental approach. We then integrate these experimental results from organismal water‐balance data and their plastic responses into a population dynamics model to examine the potential relative fitness effects of simulated transient weather conditions on population growth rates. The results show evidence for the predicted trade‐off for plasticity of water loss rate (WLR) and the duration of new environmental conditions. When altered environmental conditions lasted for longer durations, physiological responses could match the new environmental conditions, and this resulted in a lower WLR and lower rates of population decline. At shorter time‐scales however, a mismatch between acclimation duration and physiological responses was reflected by reduced overall population growth rates. This may indicate a potential fitness cost due to insufficient time for physiological adjustments to take place. The outcomes of this work therefore suggest plastic water balance responses have both costs and benefits, and these depend on the time‐scale and magnitude of variation in environmental conditions. These results are significant for understanding the evolution of plastic physiological responses and changes in population abundance in the context of environmental variability.     

Life History theory hypotheses on child growth: Potential implications for short and long‐term child growth,development and health

Life history theory integrates ecological, physiological, and molecular layers within an evolutionary framework to understand organisms’ strategies to optimize survival and reproduction. Two life history hypotheses and their implications for child growth, development, and health (illustrated in the South African context) are reviewed here. One hypothesis suggests that there is an energy trade‐off between linear growth and brain growth. Undernutrition in infancy and childhood may trigger adaptive physiological mechanisms prioritizing the brain at the expense of body growth. Another hypothesis is that the period from conception to infancy is a critical window of developmental plasticity of linear growth, the duration of which may vary between and within populations. The transition from infancy to childhood may mark the end of a critical window of opportunity for improving child growth. Both hypotheses emphasize the developmental plasticity of linear growth and the potential determinants of growth variability (including the role of parent–offspring conflict in maternal resources allocation). Implications of these hypotheses in populations with high burdens of undernutrition and infections are discussed. In South Africa, HIV/AIDS during pregnancy (associated with adverse birth outcomes, short duration of breastfeeding, and social consequences) may lead to a shortened window of developmental plasticity of growth. Furthermore, undernutrition and infectious diseases in children living in South Africa, a country undergoing a rapid nutrition transition, may have adverse consequences on individuals’ cognitive abilities and risks of cardio‐metabolic diseases. Studies are needed to identify physiological mechanisms underlying energy allocation between biological functions and their potential impacts on health.     

Trade‐offs between growth and reproduction: an analysis of the quantitative genetic evidence

The assumption of a trade‐off between development time and fecundity, resulting from a positive correlation between body size and fecundity and between body size and development time, is a common feature of life history models. The present paper examines the evidence for such a trade‐off as indicated by genetic correlations between traits. The genetic covariances between traits are derived using a model in which maturation occurs when the organism achieves a genetically variable size threshold, and fecundity is an allometric function of body size with one genetically variable parameter (excluding body size itself). This model predicts that the heritabilities of the life history traits (growth rate, development time, fecundity) will not necessarily be less than the heritability of adult size (i.e. morphological traits). It is shown that if growth rate is genetically correlated with adult size then it is not possible, in general, to predict the sign of the genetic correlation between development time and fecundity. For particular cases the signs of the covariances between traits can be predicted. These predictions are tested using data drawn from the literature.     

Photosynthesis,growth, and decay traits in Sphagnum – a multispecies comparison

Peat mosses (Sphagnum) largely govern carbon sequestration in Northern Hemisphere peatlands. We investigated functional traits related to growth and decomposition in Sphagnum species. We tested the importance of environment and phylogeny in driving species traits and investigated trade‐offs among them. We selected 15 globally important Sphagnum species, representing four sections (subgenera) and a range of peatland habitats. We measured rates of photosynthesis and decomposition in standard laboratory conditions as measures of innate growth and decay potential, and related this to realized growth, production, and decomposition in their natural habitats. In general, we found support for a trade‐off between measures of growth and decomposition. However, the relationships are not strong, with r ranging between 0.24 and 0.45 for different measures of growth versus decomposition. Using photosynthetic rate to predict decomposition in standard conditions yielded R² = 0.20. Habitat and section (phylogeny) affected the traits and the trade‐offs. In a wet year, species from sections Cuspidata and Sphagnum had the highest production, but in a dry year, differences among species, sections, and habitats evened out. Cuspidata species in general produced easily decomposable litter, but their decay in the field was hampered, probably due to near‐surface anoxia in their wet habitats. In a principal components analysis, PCA, photosynthetic capacity, production, and laboratory decomposition acted in the same direction. The species were imperfectly clustered according to vegetation type and phylogeny, so that some species clustered with others in the same section, whereas others clustered more clearly with others from similar vegetation types. Our study includes a wider range of species and habitats than previous trait analyses in Sphagnum and shows that while the previously described growth–decay trade‐off exists, it is far from perfect. We therefore suggest that our species‐specific trait measures offer opportunities for improvements of peatland ecosystem models. Innate qualities measured in laboratory conditions translate differently to field responses. Most dramatically, fast‐growing species could only realize their potential in a wet year. The same species decompose fast in laboratory, but their decomposition was more retarded in the field than that of other species. These relationships are crucial for understanding the long‐term dynamics of peatland communities.     

Physiological stress links parasites to carotenoid‐based colour signals

Vertebrates commonly use carotenoid‐based traits as social signals. These can reliably advertise current nutritional status and health because carotenoids must be acquired through the diet and their allocation to ornaments is traded‐off against other self‐maintenance needs. We propose that the coloration more generally reveals an individual’s ability to cope with stressful conditions. We tested this idea by manipulating the nematode parasite infection in free‐living red grouse (Lagopus lagopus scoticus) and examining the effects on body mass, carotenoid‐based coloration of a main social signal and the amount of corticosterone deposited in feathers grown during the experiment. We show that parasites increase stress and reduce carotenoid‐based coloration, and that the impact of parasites on coloration was associated with changes in corticosterone, more than changes in body mass. Carotenoid‐based coloration appears linked to physiological stress and could therefore reveal an individual’s ability to cope with stressors.     

Age‐dependent responses to alarm calls depend on foraging activity in Willow Tits Poecile montanus

Survivorship in animals depends on both foraging activities and avoidance of predation, and thus behavioural decisions often reflect a trade‐off between predation risk and foraging efficiency. In this experimental study, we compared behavioural responses of free‐living adult and juvenile Willow Tits Poecile montanus to a conspecific alarm call in two treatments. The alarm call was played back when a focal bird was either not feeding, or feeding on a sunflower seed on the middle part of a spruce branch. When feeding at the time of the alarm call, juveniles more often stayed motionless or moved shorter distances than adults. Our results suggest that in hierarchical groups, juveniles are forced to take greater risks to maintain access to food or lack experience to optimize between food and safety.