Meta-analysis reveals glucocorticoid levels reflect variation in metabolic rate, not ‘stress’

Glucocorticoid (GC) variation has long been thought to reflect variation in organismal ‘stress,’ but associations between GCs and Darwinian fitness components are diverse in magnitude, direction, and highly context-dependent. This paradox reveals our poor understanding of the causes of GC variation, contrasting with the detailed knowledge of the functional consequences of GC variation. Amongst an array of effects in many physiological systems, GCs orchestrate energy availability to anticipate and recover from predictable and unpredictable environmental fluctuations and challenges. Although this is mechanistically well-known, the extent to which GC levels are quantitatively explained by energy metabolism is unresolved. We investigated this association through meta-analysis, selecting studies of endotherms in which (1) an experiment was performed that affected metabolic rate and (2) metabolic rate and GC levels were measured simultaneously. We found that an increase in metabolic rate was associated with an increase in GC levels in 20 out of 21 studies (32 out of 35 effect sizes). More importantly, there was a strong positive correlation between the increases in metabolic rate and GCs (p=0.003). This pattern was similar in birds and mammals, and independent of the nature of the experimental treatment. We conclude that metabolic rate is a major driver of GC variation within individuals. Stressors often affect metabolic rate, leading us to question whether GC levels provide information on ‘stress’ beyond the stressor’s effect on metabolic rate

Insulin-like growth factor 1 of wild vertebrates in a life-history context

Broad variation in intra- and interspecific life-history traits is largely shaped by resource limitation and the ensuing allocation trade-offs that animals are forced to make. Insulin-like growth factor 1 (IGF-1), a growth-hormone-dependent peptide, may be a key player in the regulation of allocation processes. In laboratory animals, the effects of IGF-1 on growth- and development (positive), reproduction (positive), and longevity (negative) are well established. We here review the evidence on these effects in wild vertebrates, where animals are more likely to face resource limitation and other challenges. We point out the similarities and dissimilarities in patterns of IGF-1 functions obtained in these two different study settings and discuss the knowledge we need to develop a comprehensive picture of the role of IGF-1 in mediating life-history variation of wild vertebrates

Mosaic metabolic ageing:Basal and standard metabolic rates age in opposite directions and independent of environmental quality, sex and life span in a passerine

1. Crucial to our understanding of the ageing process is identifying how traits change with age, which variables alter their ageing process and how these traits associate with fitness. 2. Here we investigated metabolic ageing in outdoor-living captive zebra finches experiencing foraging costs. We longitudinally monitored 407 individuals over 6 years and collected 3,213 measurements of two independent mass-adjusted metabolic traits: basal metabolic rate (BMRm) at thermoneutral temperatures and standard metabolic rate (SMRm), measured as BMRm but at ambient temperatures below thermoneutrality. 3. We define mosaic or asynchronous ageing as the difference in standardized absolute ageing rates between traits, and we estimate the degree of asynchrony using the within-individual correlation of change in trait values with age. 4. BMRm decreased linearly with age, consistent with earlier reports. In contrast, SMRm increased linearly with age. The absolute standardized change with age was significantly faster for BMRm compared to SMRm, and the within-individual correlation of age related change was negligible. To the best of our knowledge, this is the first quantification of SMRm ageing, and the finding that SMRm and BMRm age in opposite directions. 5. Neither metabolic rate nor metabolic ageing rate were associated with variation in life span between individuals. Moreover, experimental manipulations of environmental quality that decreased BMRm and SMRm and shortened life span by 6 months (12%) did not affect the ageing of either metabolic trait. Females lived 2 months (4%) shorter than males, but none of the metabolic traits showed sex-specific differences at any age. 6. Our findings indicate, in contrast to the current view, that baseline energy requirements increase with age, because animals do not generally live in thermoneutral conditions, and illustrate the importance of studying the ageing phenotype in an ecologically realistic setting

Glucose tolerance predicts survival in old zebra finches

The capacity to deal with external and internal challenges is thought to affect fitness, and the age-linked impairment of this capacity defines the ageing process. Using a recently developed intra-peritoneal glucose tolerance test (GTT), we tested for a link between the capacity to regulate glucose levels and survival in zebra finches. We also investigated for the effects of ambient factors, age, sex, and manipulated developmental and adult conditions (i.e. natal brood size and foraging cost, in a full factorial design) on glucose tolerance. Glucose tolerance was quantified using the incremental 'area under the curve' (AUC), with lower values indicating higher tolerance. Glucose tolerance predicted survival probability in old birds, above the median age, with individuals with higher glucose tolerance showing better survival than individuals with low or intermediate glucose tolerance. In young birds there was no association between glucose tolerance and survival. Experimentally induced adverse developmental conditions did not affect glucose tolerance, but low ambient temperature at sampling and hard foraging conditions during adulthood induced a fast return to baseline levels (i.e. high glucose tolerance). These findings can be interpreted as an efficient return to baseline glucose levels when energy requirements are high, with glucose presumably being used for energy metabolism or storage. Glucose tolerance was independent of sex. Our main finding that old birds with higher glucose tolerance had better survival supports the hypothesis that the capacity to efficiently cope with a physiological challenge predicts lifespan, at least in old birds

Improving comparability between qPCR-based telomere studies

Comparability of findings from qPCR-based telomere studies is hampered by such measurement results being assay-specific, precluding a direct quantitative comparisons of observed differences and/or slopes of associations between studies. It is proposed that this can be partially alleviated by expressing qPCR-based telomere data as Z-scores

The influence of the starvation-predation trade-off on the relationship between ambient temperature and body size among endotherms

Aim: The tendency for animals at higher latitudes to be larger (Bergmann's rule) is generally explained by recourse to latitudinal effects on ambient temperature and the food supply, but these receive only mixed support and do not explain observations of the inverse to Bergmann's rule. Our aim was to better understand how ecological variables might influence body size and thereby explain this mixed support. Location: World-wide. Methods: Previous explanations do not allow for the selective pressure exerted by the trade-off between predation and starvation, which we incorporate in a model of optimal body size and energy storage of a generalized homeotherm. In contrast to existing arguments, we concentrate on survival over winter when the food supply is poor and can be interrupted for short periods. Results: We use our model to assess the logical validity of the heat conservation hypothesis and show that it must allow for the roles of both food availability and predation risk. We find that whether the effect of temperature on body size is positive or negative depends on temperature range, predator density, and the likelihood of long interruptions to foraging. Furthermore, changing day length explains differing effects of altitude and latitude on body size, leading to opposite predictions for nocturnal and diurnal endotherms. Food availability and ambient temperature can have counteracting selective pressures on body mass, and can lead to a non-monotonic relationship between latitude and size, as observed in several studies. Main conclusions: Our work provides a theoretical framework for understanding the relationships between the costs and benefits of large body size and eco-geographical patterns among endotherms world-wide

Telomere dynamics in relation to experimentally increased locomotion costs and fitness in great tits

Evidence that telomere length (TL) and dynamics can be interpreted as proxy for 'life stress' experienced by individuals stems largely from correlational studies. We tested for effects of an experimental increase of workload on telomere dynamics by equipping male great tits (Parus major) with a 0.9 g backpack for a full year. In addition, we analysed associations between natural life-history variation, TL and TL dynamics. Carrying 5% extra weight for a year did not significantly accelerate telomere attrition. This agrees with our earlier finding that this experiment did not affect survival or future reproduction. Apparently, great tit males were able to compensate behaviourally or physiologically for the increase in locomotion costs we imposed. We found no cross-sectional association between reproductive success and TL, but individuals with higher reproductive success (number of recruits) lost fewer telomere base pairs in the subsequent year. We used the TRF method to measure TL, which method yields a TL distribution for each sample, and the association between reproductive success and telomere loss was more pronounced in the higher percentiles of the telomere distribution, in agreement with the higher impact of ageing on longer telomeres within individuals. Individuals with longer telomeres and less telomere shortening were more likely to survive to the next breeding season, but these patterns did not reach statistical significance. Whether successful individuals are characterized by losing fewer or more base pairs from their telomeres varies between species, and we discuss aspects of ecology and social organisation that may explain this variation