Publications by authors named "Michaela Hau"

73 Publications

Quantifying Glucocorticoid Plasticity Using Reaction Norm Approaches: There Still is So Much to Discover!

Integr Comp Biol 2021 Oct 19. Epub 2021 Oct 19.

Department of Biology, University of Konstanz, Konstanz, Germany.

Hormones are highly responsive internal signals that help organisms adjust their phenotype to fluctuations in environmental and internal conditions. Our knowledge of the causes and consequences of variations in circulating hormone concentrations has improved greatly in the past. However, this knowledge comes from population-level studies which generally tend to make the flawed assumption that all individuals respond in the same way to environmental changes. Here, we advocate that we can vastly expand our understanding of the ecology and evolution of hormonal traits once we acknowledge the existence of individual differences by quantifying hormonal plasticity at the individual level, where selection acts. In this review, we use glucocorticoid hormones as examples of highly plastic endocrine traits that interact intimately with energy metabolism but also with other organismal traits like behavior and physiology. First, we highlight the insights gained by repeatedly assessing an individual's glucocorticoid concentrations along a gradient of environmental or internal conditions using a 'reaction norm approach'. This study design should be followed by a hierarchical statistical partitioning of the total endocrine variance into the among-individual component (individual differences in average hormone concentrations, i.e. in the intercept of the reaction norm) and the residual (within-individual) component. The latter is ideally further partitioned by estimating more precisely the hormonal plasticity (i.e. the slope of the reaction norm), which allows to test whether individuals differ in the degree of hormonal change along the gradient. Second, we critically review the published evidence for glucocorticoid variation, focusing mostly on among- and within-individual levels, finding only a good handful of studies that used repeated-measures designs and random regression statistics to investigate glucocorticoid plasticity. These studies indicate that individuals can differ in both the intercept and the slope of their glucocorticoid reaction norm to a known gradient. Third, we suggest rewarding avenues for future work on hormonal reaction norms, for example to uncover potential costs and trade-offs associated with glucocorticoid plasticity or to test whether glucocorticoid plasticity varies when an individual's reaction norm is repeatedly assessed along the same gradient, whether reaction norms in glucocorticoids covary with those in other traits like behavior and fitness (generating multivariate plasticity) or to quantify glucocorticoid reaction norms along multiple external and internal gradients that act simultaneously (leading to multidimensional plasticity). Throughout this review we emphasize the power that reaction norm approaches offer for resolving unanswered questions in ecological and evolutionary endocrinology.
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http://dx.doi.org/10.1093/icb/icab196DOI Listing
October 2021

Natural variation in yolk fatty acids, but not androgens, predicts offspring fitness in a wild bird.

Front Zool 2021 Aug 5;18(1):38. Epub 2021 Aug 5.

Max Planck Institute for Ornithology, Eberhard-Gwinner-Str., 82319, Seewiesen, Germany.

Background: In egg-laying animals, mothers can influence the developmental environment and thus the phenotype of their offspring by secreting various substances into the egg yolk. In birds, recent studies have demonstrated that different yolk substances can interactively affect offspring phenotype, but the implications of such effects for offspring fitness and phenotype in natural populations have remained unclear. We measured natural variation in the content of 31 yolk components known to shape offspring phenotypes including steroid hormones, antioxidants and fatty acids in eggs of free-living great tits (Parus major) during two breeding seasons. We tested for relationships between yolk component groupings and offspring fitness and phenotypes.

Results: Variation in hatchling and fledgling numbers was primarily explained by yolk fatty acids (including saturated, mono- and polyunsaturated fatty acids) - but not by androgen hormones and carotenoids, components previously considered to be major determinants of offspring phenotype. Fatty acids were also better predictors of variation in nestling oxidative status and size than androgens and carotenoids.

Conclusions: Our results suggest that fatty acids are important yolk substances that contribute to shaping offspring fitness and phenotype in free-living populations. Since polyunsaturated fatty acids cannot be produced de novo by the mother, but have to be obtained from the diet, these findings highlight potential mechanisms (e.g., weather, habitat quality, foraging ability) through which environmental variation may shape maternal effects and consequences for offspring. Our study represents an important first step towards unraveling interactive effects of multiple yolk substances on offspring fitness and phenotypes in free-living populations. It provides the basis for future experiments that will establish the pathways by which yolk components, singly and/or interactively, mediate maternal effects in natural populations.
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http://dx.doi.org/10.1186/s12983-021-00422-zDOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC8340462PMC
August 2021

Inferring Whole-Organism Metabolic Rate From Red Blood Cells in Birds.

Front Physiol 2021 16;12:691633. Epub 2021 Jul 16.

Research Group for Evolutionary Physiology, Max Planck Institute for Ornithology, Seewiesen, Germany.

Metabolic rate is a key ecological variable that quantifies the energy expenditure needed to fuel almost all biological processes in an organism. Metabolic rates are typically measured at the whole-organism level (woMR) with protocols that can elicit stress responses due to handling and confinement, potentially biasing resulting data. Improved, non-stressful methodology would be especially valuable for measures of field metabolic rate, which quantifies the energy expenditure of free-living individuals. Recently, techniques to measure cellular metabolic rate (cMR) in mitochondria of blood cells have become available, suggesting that blood-based cMR can be a proxy of organismal aerobic performance. Aerobic metabolism actually takes place in the mitochondria. Quantifying cMR from blood samples offers several advantages such as direct estimates of metabolism and minimized disturbance of individuals. To our knowledge, the hypothesis that blood-based cMR correlates with woMR has not yet been directly tested. We measured cMR in red blood cells of captive great tits (), first during their morning activity period and second after subjecting them to a 2.5 h day-time respirometry protocol to quantify woMR. We predicted cMR to decrease as individuals transitioned from an active to a resting state. In the two blood samples we also assessed circulating corticosterone concentrations to determine the perceived disturbance of individuals. From respirometry traces we extracted initial and final woMR measures to test for a predicted positive correlation with cMR measures, while accounting for corticosterone concentrations. Indeed, cMR declined from the first to the second measurement. Furthermore, woMR and cMR were positively related in individuals that had relatively low corticosterone concentrations and displayed little locomotor activity throughout respirometry. By contrast, woMR and cMR covaried negatively in birds that increased corticosterone concentrations and activity levels substantially. Our results show that red blood cell cMR represents a proxy for woMR when birds do not display signs of stress, i.e., either before increases in hormonal or behavioral parameters have occurred or after they have abated. This method represents a valuable tool for obtaining metabolic data repeatedly and in free-living individuals. Our findings also highlight the importance of accounting for individual stress responses when measuring metabolic rate at any level.
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http://dx.doi.org/10.3389/fphys.2021.691633DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC8322697PMC
July 2021

Early nighttime testosterone peaks are correlated with GnRH-induced testosterone in a diurnal songbird.

Gen Comp Endocrinol 2021 Oct 21;312:113861. Epub 2021 Jul 21.

Max Planck Institute for Ornithology, Evolutionary Physiology Research Group, Seewiesen, Germany; University of Konstanz, Department of Biology, Konstanz, Germany.

Experimental manipulation has established testosterone as a potent, pleiotropic regulator coordinating morphology, physiology and behavior. However, the relationship of field-sampled, unmanipulated testosterone concentrations with traits of interest is often equivocal. Circulating testosterone varies over the course of the day, and recent reports indicate that testosterone is higher during the night in diurnal songbirds. Yet, most field studies sample testosterone during the morning. Sampling at times when levels and individual variation are low may be one reason relationships between testosterone and other traits are not always observed. Testosterone is regulated by the hypothalamic-pituitary-gonadal axis, with gonadotropin-releasing hormone (GnRH) initiating the endocrine cascade. Research has examined GnRH-induced testosterone levels with traits of interest, yet the relevance of these induced levels and their relationship with endogenously produced levels are not fully clear. Using photostimulated male great tits (Parus major) we tested the hypotheses that circulating testosterone levels peak during the night and that GnRH-induced testosterone concentrations are positively related to nightly testosterone peaks. Blood was sampled during first, middle or last third of night. One week later, baseline and GnRH-induced testosterone levels were sampled during mid-morning. Morning baseline testosterone levels were low compared with night-sampled levels that peaked during the first third of the night. Further, GnRH-induced testosterone was strongly positively correlated with levels observed during the first third of the night. These data suggest that morning testosterone samples likely do not reflect an individual's endogenous peak. Instead, GnRH-induced testosterone levels do approximate an individual's nightly peak and may be an alternative for birds that cannot easily be sampled at night in the field. These findings are likely to have implications for research aimed at relating traits of interest with natural variation in sex steroid hormone levels.
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http://dx.doi.org/10.1016/j.ygcen.2021.113861DOI Listing
October 2021

Life history and environment predict variation in testosterone across vertebrates.

Evolution 2021 05 2;75(5):1003-1010. Epub 2021 Apr 2.

Department of Biological Sciences, Simon Fraser University, Burnaby, British Columbia, V5A 1S6, Canada.

Endocrine systems act as key intermediaries between organisms and their environments. This interaction leads to high variability in hormone levels, but we know little about the ecological factors that influence this variation within and across major vertebrate groups. We study this topic by assessing how various social and environmental dynamics influence testosterone levels across the entire vertebrate tree of life. Our analyses show that breeding season length and mating system are the strongest predictors of average testosterone concentrations, whereas breeding season length, environmental temperature, and variability in precipitation are the strongest predictors of within-population variation in testosterone. Principles from small-scale comparative studies that stress the importance of mating opportunity and competition on the evolution of species differences in testosterone levels, therefore, likely apply to the entire vertebrate lineage. Meanwhile, climatic factors associated with rainfall and ambient temperature appear to influence variability in plasma testosterone, within a given species. These results, therefore, reveal how unique suites of ecological factors differentially explain scales of variation in circulating testosterone across mammals, birds, reptiles, amphibians, and fishes.
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http://dx.doi.org/10.1111/evo.14216DOI Listing
May 2021

Sex steroids modulate circadian behavioral rhythms in captive animals, but does this matter in the wild?

Horm Behav 2021 02 29;128:104900. Epub 2020 Nov 29.

North Dakota State University, Department of Biological Sciences, Fargo, ND, USA.

Nearly all organisms alter physiological and behavioral activities across the twenty-four-hour day. Endogenous timekeeping mechanisms, which are responsive to environmental and internal cues, allow organisms to anticipate predictable environmental changes and time their daily activities. Among-individual variation in the chronotype, or phenotypic output of these timekeeping mechanisms (i.e. timing of daily behaviors), is often observed in organisms studied under naturalistic environmental conditions. The neuroendocrine system, including sex steroids, has been implicated in the regulation and modulation of endogenous clocks and their behavioral outputs. Numerous studies have found clear evidence that sex steroids modulate circadian and daily timing of activities in captive animals under controlled conditions. However, little is known about how sex steroids influence daily behavioral rhythms in wild organisms or what, if any, implication this may have for survival and reproductive fitness. Here we review the evidence that sex steroids modulate daily timing in vertebrates under controlled conditions. We then discuss how this relationship may be relevant for the reproductive success and fitness of wild organisms and discuss the limited evidence that sex steroids modulate circadian rhythms in wild organisms.
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http://dx.doi.org/10.1016/j.yhbeh.2020.104900DOI Listing
February 2021

Connecting the data landscape of long-term ecological studies: The SPI-Birds data hub.

J Anim Ecol 2021 09 4;90(9):2147-2160. Epub 2020 Dec 4.

Stazione Ornitologica, Monreale, Italy.

The integration and synthesis of the data in different areas of science is drastically slowed and hindered by a lack of standards and networking programmes. Long-term studies of individually marked animals are not an exception. These studies are especially important as instrumental for understanding evolutionary and ecological processes in the wild. Furthermore, their number and global distribution provides a unique opportunity to assess the generality of patterns and to address broad-scale global issues (e.g. climate change). To solve data integration issues and enable a new scale of ecological and evolutionary research based on long-term studies of birds, we have created the SPI-Birds Network and Database (www.spibirds.org)-a large-scale initiative that connects data from, and researchers working on, studies of wild populations of individually recognizable (usually ringed) birds. Within year and a half since the establishment, SPI-Birds has recruited over 120 members, and currently hosts data on almost 1.5 million individual birds collected in 80 populations over 2,000 cumulative years, and counting. SPI-Birds acts as a data hub and a catalogue of studied populations. It prevents data loss, secures easy data finding, use and integration and thus facilitates collaboration and synthesis. We provide community-derived data and meta-data standards and improve data integrity guided by the principles of Findable, Accessible, Interoperable and Reusable (FAIR), and aligned with the existing metadata languages (e.g. ecological meta-data language). The encouraging community involvement stems from SPI-Bird's decentralized approach: research groups retain full control over data use and their way of data management, while SPI-Birds creates tailored pipelines to convert each unique data format into a standard format. We outline the lessons learned, so that other communities (e.g. those working on other taxa) can adapt our successful model. Creating community-specific hubs (such as ours, COMADRE for animal demography, etc.) will aid much-needed large-scale ecological data integration.
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http://dx.doi.org/10.1111/1365-2656.13388DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC8518542PMC
September 2021

Increased glucocorticoid concentrations in early life cause mitochondrial inefficiency and short telomeres.

J Exp Biol 2020 08 4;223(Pt 15). Epub 2020 Aug 4.

Max Planck Institute for Ornithology, Evolutionary Physiology Group, 82319 Seewiesen, Germany.

Telomeres are DNA structures that protect chromosome ends. However, telomeres shorten during cell replication and at critically low lengths can reduce cell replicative potential, induce cell senescence and decrease fitness. Stress exposure, which elevates glucocorticoid hormone concentrations, can exacerbate telomere attrition. This phenomenon has been attributed to increased oxidative stress generated by glucocorticoids ('oxidative stress hypothesis'). We recently suggested that glucocorticoids could increase telomere attrition during stressful periods by reducing the resources available for telomere maintenance through changes in the metabolic machinery ('metabolic telomere attrition hypothesis'). Here, we tested whether experimental increases in glucocorticoid levels affected telomere length and mitochondrial function in wild great tit () nestlings during the energy-demanding early growth period. We monitored resulting corticosterone (Cort) concentrations in plasma and red blood cells, telomere lengths and mitochondrial metabolism (metabolic rate, proton leak, oxidative phosphorylation, maximal mitochondrial capacity and mitochondrial inefficiency). We assessed oxidative damage caused by reactive oxygen species (ROS) metabolites as well as the total non-enzymatic antioxidant protection in plasma. Compared with control nestlings, Cort-nestlings had higher baseline corticosterone, shorter telomeres and higher mitochondrial metabolic rate. Importantly, Cort-nestlings showed increased mitochondrial proton leak, leading to a decreased ATP production efficiency. Treatment groups did not differ in oxidative damage or antioxidants. Hence, glucocorticoid-induced telomere attrition is associated with changes in mitochondrial metabolism, but not with ROS production. These findings support the hypothesis that shortening of telomere length during stressful periods is mediated by glucocorticoids through metabolic rearrangements.
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http://dx.doi.org/10.1242/jeb.222513DOI Listing
August 2020

Baseline and stress-induced corticosterone levels across birds and reptiles do not reflect urbanization levels.

Conserv Physiol 2020 27;8(1):coz110. Epub 2020 Jan 27.

Department of Ecology and Evolutionary Biology, Cornell University, Ithaca NY 14853, USA.

Rates of human-induced environmental change continue increasing with human population size, potentially altering animal physiology and negatively affecting wildlife. Researchers often use glucocorticoid concentrations (hormones that can be associated with stressors) to gauge the impact of anthropogenic factors (e.g. urbanization, noise and light pollution). Yet, no general relationships between human-induced environmental change and glucocorticoids have emerged. Given the number of recent studies reporting baseline and stress-induced corticosterone (the primary glucocorticoid in birds and reptiles) concentrations worldwide, it is now possible to conduct large-scale comparative analyses to test for general associations between disturbance and baseline and stress-induced corticosterone across species. Additionally, we can control for factors that may influence context, such as life history stage, environmental conditions and urban adaptability of a species. Here, we take a phylogenetically informed approach and use data from HormoneBase to test if baseline and stress-induced corticosterone are valid indicators of exposure to human footprint index, human population density, anthropogenic noise and artificial light at night in birds and reptiles. Our results show a negative relationship between anthropogenic noise and baseline corticosterone for birds characterized as urban avoiders. While our results potentially indicate that urban avoiders are more sensitive to noise than other species, overall our study suggests that the relationship between human-induced environmental change and corticosterone varies across species and contexts; we found no general relationship between human impacts and baseline and stress-induced corticosterone in birds, nor baseline corticosterone in reptiles. Therefore, it should not be assumed that high or low levels of exposure to human-induced environmental change are associated with high or low corticosterone levels, respectively, or that closely related species, or even individuals, will respond similarly. Moving forward, measuring alternative physiological traits alongside reproductive success, health and survival may provide context to better understand the potential negative effects of human-induced environmental change.
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http://dx.doi.org/10.1093/conphys/coz110DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6978728PMC
January 2020

Host dispersal shapes the population structure of a tick-borne bacterial pathogen.

Mol Ecol 2020 02 9;29(3):485-501. Epub 2020 Jan 9.

CNRS - Department of Biometry and Evolutionary Biology (LBBE) - University Lyon 1, University of Lyon, Villeurbanne, France.

Birds are hosts for several zoonotic pathogens. Because of their high mobility, especially of longdistance migrants, birds can disperse these pathogens, affecting their distribution and phylogeography. We focused on Borrelia burgdorferi sensu lato, which includes the causative agents of Lyme borreliosis, as an example for tick-borne pathogens, to address the role of birds as propagation hosts of zoonotic agents at a large geographical scale. We collected ticks from passerine birds in 11 European countries. B. burgdorferi s.l. prevalence in Ixodes spp. was 37% and increased with latitude. The fieldfare Turdus pilaris and the blackbird T. merula carried ticks with the highest Borrelia prevalence (92 and 58%, respectively), whereas robin Erithacus rubecula ticks were the least infected (3.8%). Borrelia garinii was the most prevalent genospecies (61%), followed by B. valaisiana (24%), B. afzelii (9%), B. turdi (5%) and B. lusitaniae (0.5%). A novel Borrelia genospecies "Candidatus Borrelia aligera" was also detected. Multilocus sequence typing (MLST) analysis of B. garinii isolates together with the global collection of B. garinii genotypes obtained from the Borrelia MLST public database revealed that: (a) there was little overlap among genotypes from different continents, (b) there was no geographical structuring within Europe, and (c) there was no evident association pattern detectable among B. garinii genotypes from ticks feeding on birds, questing ticks or human isolates. These findings strengthen the hypothesis that the population structure and evolutionary biology of tick-borne pathogens are shaped by their host associations and the movement patterns of these hosts.
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http://dx.doi.org/10.1111/mec.15336DOI Listing
February 2020

Developmental conditions modulate DNA methylation at the glucocorticoid receptor gene with cascading effects on expression and corticosterone levels in zebra finches.

Sci Rep 2019 11 1;9(1):15869. Epub 2019 Nov 1.

Groningen Institute for Evolutionary Life Sciences, University of Groningen, Groningen, The Netherlands.

Developmental conditions can impact the adult phenotype via epigenetic changes that modulate gene expression. In mammals, methylation of the glucocorticoid receptor gene Nr3c1 has been implicated as mediator of long-term effects of developmental conditions, but this evidence is limited to humans and rodents, and few studies have simultaneously tested for associations between DNA methylation, gene expression and phenotype. Adverse environmental conditions during early life (large natal brood size) or adulthood (high foraging costs) exert multiple long-term phenotypic effects in zebra finches, and we here test for effects of these manipulations on DNA methylation and expression of the Nr3c1 gene in blood. Having been reared in a large brood induced higher DNA methylation of the Nr3c1 regulatory region in adulthood, and this effect persisted over years. Nr3c1 expression was negatively correlated with methylation at 2 out of 8 CpG sites, and was lower in hard foraging conditions, despite foraging conditions having no effect on Nr3c1 methylation at our target region. Nr3c1 expression also correlated with glucocorticoid traits: higher expression level was associated with lower plasma baseline corticosterone concentrations and enhanced corticosterone reactivity. Our results suggest that methylation of the Nr3c1 regulatory region can contribute to the mechanisms underlying the emergence of long-term effects of developmental conditions in birds, but in our system current adversity dominated over early life experiences with respect to receptor expression.
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http://dx.doi.org/10.1038/s41598-019-52203-8DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6825131PMC
November 2019

Macroevolutionary Patterning in Glucocorticoids Suggests Different Selective Pressures Shape Baseline and Stress-Induced Levels.

Am Nat 2019 06 17;193(6):866-880. Epub 2019 Apr 17.

Glucocorticoid (GC) hormones are important phenotypic mediators across vertebrates, but their circulating concentrations can vary markedly. Here we investigate macroevolutionary patterning in GC levels across tetrapods by testing seven specific hypotheses about GC variation and evaluating whether the supported hypotheses reveal consistent patterns in GC evolution. If selection generally favors the "supportive" role of GCs in responding effectively to challenges, then baseline and/or stress-induced GCs may be higher in challenging contexts. Alternatively, if selection generally favors "protection" from GC-induced costs, GCs may be lower in environments where challenges are more common or severe. The predictors of baseline GCs were all consistent with supportive effects: levels were higher in smaller organisms and in those inhabiting more energetically demanding environments. During breeding, baseline GCs were also higher in populations and species with fewer lifetime opportunities to reproduce. The predictors of stress-induced GCs were instead more consistent with the protection hypothesis: during breeding, levels were lower in organisms with fewer lifetime reproductive opportunities. Overall, these patterns indicate a surprising degree of consistency in how some selective pressures shape GCs across broad taxonomic scales; at the same time, in challenging environments selection appears to operate on baseline and stress-induced GCs in distinct ways.
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http://dx.doi.org/10.1086/703112DOI Listing
June 2019

Telomere attrition: metabolic regulation and signalling function?

Biol Lett 2019 03;15(3):20180885

1 Research Group Evolutionary Physiology, Max Planck Institute for Ornithology , 82319 Seewiesen , Germany.

Stress exposure can leave long-term footprints within the organism, like in telomeres (TLs), protective chromosome caps that shorten during cell replication and following exposure to stressors. Short TLs are considered to indicate lower fitness prospects, but why TLs shorten under stressful conditions is not understood. Glucocorticoid hormones (GCs) increase upon stress exposure and are thought to promote TL shortening by increasing oxidative damage. However, evidence that GCs are pro-oxidants and oxidative stress is causally linked to TL attrition is mixed . Based on new biochemical findings, we propose the metabolic telomere attrition hypothesis: during times of substantially increased energy demands, TLs are shortened as part of the transition into an organismal 'emergency state', which prioritizes immediate survival functions over processes with longer-term benefits. TL attrition during energy shortages could serve multiple roles including amplified signalling of cellular energy debt to re-direct critical resources to immediately important processes. This new view of TL shortening as a strategy to resolve major energetic trade-offs can improve our understanding of TL dynamics. We suggest that TLs are master regulators of cell homeostasis and propose future research avenues to understand the interactions between energy homeostasis, metabolic regulators and TL.
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http://dx.doi.org/10.1098/rsbl.2018.0885DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6451386PMC
March 2019

Enzymatic antioxidants but not baseline glucocorticoids mediate the reproduction-survival trade-off in a wild bird.

Proc Biol Sci 2018 11 28;285(1892). Epub 2018 Nov 28.

Department of Evolutionary Physiology, Max Planck Institute for Ornithology, Seewiesen, Starnberg, Germany.

The trade-off between reproductive investment and survival is central to life-history theory, but the relative importance and the complex interactions among the physiological mechanisms mediating it are still debated. Here we experimentally tested whether baseline glucocorticoid hormones, the redox system or their interaction mediate reproductive investment-survival trade-offs in wild great tits (). We increased the workload of parental males by clipping three feathers on each wing, and 5 days later determined effects on baseline corticosterone concentrations (Cort), redox state (reactive oxygen metabolites, protein carbonyls, glutathione peroxidase [GPx], total non-enzymatic antioxidants), body mass, body condition, reproductive success and survival. Feather-clipping did not affect fledgling numbers, chick body condition, nest provisioning rates or survival compared with controls. However, feather-clipped males lost mass and increased both Cort and GPx concentrations. Within feather-clipped individuals, GPx increases were positively associated with reproductive investment (i.e. male nest provisioning). Furthermore, within all individuals, males that increased GPx suffered reduced survival rates. Baseline Cort increases were related to mass loss but not to redox state, nest provisioning or male survival. Our findings provide experimental evidence that changes in the redox system are associated with the trade-off between reproductive investment and survival, while baseline Cort may support this trade-off indirectly through a link with body condition. These results also emphasize that plastic changes in individuals, rather than static levels of physiological signals, may mediate life-history trade-offs.
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http://dx.doi.org/10.1098/rspb.2018.2141DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6283945PMC
November 2018

Glucocorticoid-temperature association is shaped by foraging costs in individual zebra finches.

J Exp Biol 2018 11 27;221(Pt 23). Epub 2018 Nov 27.

Groningen Institute for Evolutionary Life Sciences, University of Groningen, 9747 AG Groningen, The Netherlands.

Glucocorticoid (GC) levels vary with environmental conditions, but the functional interpretation of GC variation remains contentious. A primary function is thought to be metabolic, mobilizing body reserves to match energetic demands. This view is supported by temperature-dependent GC levels, although reports of this effect show unexplained heterogeneity. We hypothesized that the temperature effect on GC concentrations will depend on food availability through its effect on the energy spent to gather the food needed for thermoregulation. We tested this hypothesis in zebra finches living in outdoor aviaries with manipulated foraging conditions (i.e. easy versus hard), by relating within-individual differences in baseline GCs between consecutive years to differences in ambient temperature. In agreement with our hypothesis, we found the GC-temperature association to be significantly steeper in the hard foraging environment. This supports the metabolic explanation of GC variation, underlining the importance of accounting for variation in energy expenditure when interpreting GC variation.
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http://dx.doi.org/10.1242/jeb.187880DOI Listing
November 2018

Corticosterone levels reflect variation in metabolic rate, independent of 'stress'.

Sci Rep 2018 08 29;8(1):13020. Epub 2018 Aug 29.

Groningen Institute for Evolutionary Life Sciences, University of Groningen, Groningen, The Netherlands.

Variation in glucocorticoid hormones (GCs) is often interpreted as reflecting 'stress', but this interpretation is subject of intense debate. GCs induce gluconeogenesis, and we hypothesized therefore that GC variation can be explained by changes in current and anticipated metabolic rate (MR). Alternatively, GC levels may respond to psychological 'stress' over and above its effect on metabolic rate. We tested these hypotheses in captive zebra finches, by inducing an increase in MR using a psychological stressor (noise), and compared its effect on corticosterone (CORT, the primary avian GC) with the effect induced by a decrease in ambient temperature increasing MR to a similar extent. We found the increase in CORT induced by the psychological stressor to be indistinguishable from the level expected based on the noise effect on MR. We further found that a handling and restraint stressor that increased CORT levels also resulted in increased blood glucose levels, corroborating a key assumption underlying our hypothesis. Thus, GC variation primarily reflected variation in energy expenditure, independently of psychological stress. GC levels have many downstream effects besides glucose mobilization, and we propose that these effects can be interpreted as adjustments of physiological functions to the metabolic level at which an organism operates.
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http://dx.doi.org/10.1038/s41598-018-31258-zDOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6115469PMC
August 2018

Corticosterone implants produce stress-hyporesponsive birds.

J Exp Biol 2018 10 11;221(Pt 19). Epub 2018 Oct 11.

Departamento de Biología de la Conservación, Estación Biológica de Doñana, Consejo Superior de Investigaciones Científicas (CSIC), 41092 Seville, Spain

In birds, the use of corticosterone (Cort) implants is a frequent tool aimed at simulating systemic elevations of this hormone and studying effects on biological traits (e.g. physiology, morphology, behavior). This manipulation may alter adrenocortical function, potentially changing both baseline (Cort) and stress-induced (Cort) plasma Cort levels. However, implant effects on the latter trait are rarely measured, disregarding downstream consequences of potentially altered stress responses. Here, we analyzed the effects of Cort implants on both Cort and Cort in nestling and adult European white storks, In addition, we performed a review of 50 studies using Cort implants in birds during the last two decades to contextualize stork results, assess researchers' patterns of use and infer current study biases. High and low doses of Cort implants resulted in a decrease of both Cort (31-71% below controls) and Cort (63-79% below controls) in storks. Our literature review revealed that Cort generally increases (72% of experiments) whereas Cort decreases (78% of experiments) following implant treatment in birds. Our results challenge and expand the prevailing assumption that Cort implants increase circulating Cort levels because: (i) Cort levels show a quadratic association with implant dose across bird species, and decreased levels may occur at both high and low implant doses, and (ii) Cort implants also decrease Cort levels, thus producing stress-hyporesponsive phenotypes. It is time to work towards a better understanding of the effects of Cort implants on adrenocortical function, before addressing downstream links to variation in other biological traits.
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http://dx.doi.org/10.1242/jeb.173864DOI Listing
October 2018

IUCN Conservation Status Does Not Predict Glucocorticoid Concentrations in Reptiles and Birds.

Integr Comp Biol 2018 10;58(4):800-813

Biological Sciences Department, California Polytechnic State University, San Luis Obispo, CA 93407, USA.

Circulating glucocorticoids (GCs) are the most commonly used biomarkers of stress in wildlife. However, their utility as a tool for identifying and/or managing at-risk species has varied. Here, we took a very broad approach to conservation physiology, asking whether International Union for the Conservation of Nature (IUCN) listing status (concern versus no obvious concern) and/or location within a geographic range (edge versus non-edge) predicted baseline and post-restraint concentrations of corticosterone (CORT) among many species of birds and reptiles. Even though such an approach can be viewed as coarse, we asked in this analysis whether CORT concentrations might be useful to implicate species at risk. Indeed, our effort, relying on HormoneBase, a repository of data on wildlife steroids, complements several other large-scale efforts in this issue to describe and understand GC variation. Using a phylogenetically informed Bayesian approach, we found little evidence that either IUCN status or edge/non-edge location in a geographic distribution were related to GC levels. However, we did confirm patterns described in previous studies, namely that breeding condition and evolutionary relatedness among species predicted some GC variation. Given the broad scope of our work, we are reluctant to conclude that IUCN status and location within a range are unrelated to GC regulation. We encourage future more targeted efforts on GCs in at-risk populations to reveal how factors leading to IUCN listing or the environmental conditions at range edges impact individual performance and fitness, particularly in the mammals, amphibians, and fish species we could not study here because data are currently unavailable.
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http://dx.doi.org/10.1093/icb/icy102DOI Listing
October 2018

Species-Specific Means and Within-Species Variance in Glucocorticoid Hormones and Speciation Rates in Birds.

Integr Comp Biol 2018 10;58(4):763-776

Department of Biological Sciences, Simon Fraser University, Burnaby, British Columbia, V5A 1S6, Canada.

At macroevolutionary scales, stress physiology may have consequences for species diversification and subspecies richness. Populations that exploit new resources or undergo range expansion should cope with new environmental challenges, which could favor higher mean stress responses. Within-species variation in the stress response may also play a role in mediating the speciation process: in species with broad variation, there will always be some individuals that can tolerate an unpredictable environment, whereas in species with narrow variation there will be fewer individuals that are able to thrive in a new ecological niche. We tested for the evolutionary relationship between stress response, speciation rate, and subspecies richness in birds by relying on the HormoneBase repository, from which we calculated within- and among-species variation in baseline (BL) and stress-induced (SI) corticosterone levels. To estimate speciation rates, we applied Bayesian analysis of macroevolutionary mixtures that can account for variation in diversification rate among clades and through time. Contrary to our predictions, lineages with higher diversification rates were not characterized by higher BL or SI levels of corticosterone either at the tips or at the deeper nodes of the phylogeny. We also found no association between mean hormone levels and subspecies richness. Within-species variance in corticosterone levels showed close to zero repeatability, thus it is highly unlikely that this is a species-specific trait that influences diversification rates. These results imply that stress physiology may play a minor, if any, role in determining speciation rates in birds.
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http://dx.doi.org/10.1093/icb/icy086DOI Listing
October 2018

Metabolic Scaling of Stress Hormones in Vertebrates.

Integr Comp Biol 2018 10;58(4):729-738

Department of Biology, Hamilton College, Clinton, NY 13323, USA.

Glucocorticoids (GCs) are stress hormones that can strongly influence physiology, behavior, and an organism's ability to cope with environmental change. Despite their importance, and the wealth of studies that have sought to understand how and why GC concentrations vary within species, we do not have a clear understanding of how circulating GC levels vary within and across the major vertebrate clades. New research has proposed that much interspecific variation in GC concentrations can be explained by variation in metabolism and body mass. Specifically, GC concentrations should vary proportionally with mass-specific metabolic rates and, given known scaling relationships between body mass and metabolic rate, GC concentrations should scale to the -1/4 power of body mass and to the power of 1 with mass-specific metabolic rate. Here, we use HormoneBase, the newly compiled database that includes plasma GC concentrations from free-living and unmanipulated vertebrates, to evaluate this hypothesis. Specifically, we explored the relationships between body mass or mass-specific metabolic rate and either baseline or stress-induced GC (cortisol or corticosterone) concentrations in tetrapods. Our phylogenetically-informed models suggest that, whereas the relationship between GC concentrations and body mass across tetrapods and among mammals is close to -1/4 power, this relationship does not exist in amphibians, reptiles, and birds. Moreover, with the exception of a positive association between stress-induced GC concentrations and mass-specific metabolic rate in birds, we found little evidence that GC concentrations are linked to metabolic rate, although the number of species sampled was quite limited for amphibians and somewhat so for reptiles and mammals. Nevertheless, these results stand in contrast to the generally accepted association between the two and suggest that our observed positive association between body mass and GC concentrations may not be due to the well-established link between mass and metabolism. Large-scale comparative approaches can come with drawbacks, such as pooling and pairing observations from separate sources. However, these broad analyses provide an important counterbalance to the majority of studies examining variation in GC concentrations at the population or species level, and can be a powerful approach to testing both long-standing and new questions in biology.
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http://dx.doi.org/10.1093/icb/icy063DOI Listing
October 2018

Do Seasonal Glucocorticoid Changes Depend on Reproductive Investment? A Comparative Approach in Birds.

Integr Comp Biol 2018 10;58(4):739-750

Max Planck Institute for Ornithology, Eberhard-Gwinner-Straße, Seewiesen 82319, Germany.

Animals go through different life history stages such as reproduction, moult, or migration, of which some are more energy-demanding than others. Baseline concentrations of glucocorticoid hormones increase during moderate, predictable challenges and thus are expected to be higher when seasonal energy demands increase, such as during reproduction. By contrast, stress-induced glucocorticoids prioritize a survival mode that includes reproductive inhibition. Thus, many species down-regulate stress-induced glucocorticoid concentrations during the breeding season. Interspecific variation in glucocorticoid levels during reproduction has been successfully mapped onto reproductive investment, with species investing strongly in current reproduction (fast pace of life) showing higher baseline and lower stress-induced glucocorticoid concentrations than species that prioritize future reproduction over current attempts (slow pace of life). Here we test the "glucocorticoid seasonal plasticity hypothesis", in which we propose that interspecific variation in seasonal changes in glucocorticoid concentrations from the non-breeding to the breeding season will be related to the degree of reproductive investment (and thus pace of life). We extracted population means for baseline (for 54 species) and stress-induced glucocorticoids (for 32 species) for the breeding and the non-breeding seasons from the database "HormoneBase", also calculating seasonal glucocorticoid changes. We focused on birds because this group offered the largest sample size. Using phylogenetic comparative methods, we first showed that species differed consistently in both average glucocorticoid concentrations and their changes between the two seasons, while controlling for sex, latitude, and hemisphere. Second, as predicted seasonal changes in baseline glucocorticoids were explained by clutch size (our proxy for reproductive investment), with species laying larger clutches showing a greater increase during the breeding season-especially in passerine species. In contrast, changes in seasonal stress-induced levels were not explained by clutch size, but sample sizes were more limited. Our findings highlight that seasonal changes in baseline glucocorticoids are associated with a species' reproductive investment, representing an overlooked physiological trait that may underlie the pace of life.
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http://dx.doi.org/10.1093/icb/icy022DOI Listing
October 2018

HormoneBase, a population-level database of steroid hormone levels across vertebrates.

Sci Data 2018 05 22;5:180097. Epub 2018 May 22.

Department of Biological Sciences, Simon Fraser University, Burnaby, BC V5A 1S6, Canada.

Hormones are central regulators of organismal function and flexibility that mediate a diversity of phenotypic traits from early development through senescence. Yet despite these important roles, basic questions about how and why hormone systems vary within and across species remain unanswered. Here we describe HormoneBase, a database of circulating steroid hormone levels and their variation across vertebrates. This database aims to provide all available data on the mean, variation, and range of plasma glucocorticoids (both baseline and stress-induced) and androgens in free-living and un-manipulated adult vertebrates. HormoneBase (www.HormoneBase.org) currently includes >6,580 entries from 476 species, reported in 648 publications from 1967 to 2015, and unpublished datasets. Entries are associated with data on the species and population, sex, year and month of study, geographic coordinates, life history stage, method and latency of hormone sampling, and analysis technique. This novel resource could be used for analyses of the function and evolution of hormone systems, and the relationships between hormonal variation and a variety of processes including phenotypic variation, fitness, and species distributions.
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http://dx.doi.org/10.1038/sdata.2018.97DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5963335PMC
May 2018

Effects of El Niño and La Niña Southern Oscillation events on the adrenocortical responses to stress in birds of the Galapagos Islands.

Gen Comp Endocrinol 2018 04 26;259:20-33. Epub 2017 Oct 26.

Department of Ecology and Evolutionary Biology, Princeton University, Princeton, NJ, USA; Max-Planck-Institut für Ornithologie, Seewiesen, and Univerzsity of Konstanz, Konstanz, Germany.

El Niño Southern Oscillation events (ENSO) and the subsequent opposite weather patterns in the following months and years (La Niña) have major climatic impacts, especially on oceanic habitats, affecting breeding success of both land and sea birds. We assessed corticosterone concentrations from blood samples during standardized protocols of capture, handling and restraint to simulate acute stress from 12 species of Galapagos Island birds during the ENSO year of 1998 and a La Niña year of 1999. Plasma levels of corticosterone were measured in samples collected at capture (to represent non-stressed baseline) and subsequently up to 1 h post-capture to give maximum corticosterone following acute stress, and total amount of corticosterone that the individual was exposed to during the test period (integrated corticosterone). Seabird species that feed largely offshore conformed to the brood value hypothesis whereas inshore feeding species showed less significant changes. Land birds mostly revealed no differences in the adrenocortical responses to acute stress from year to year with the exception of two small species (<18 g) that had an increase in baseline and stress responses in the ENSO year - contrary to predictions. We suggest that a number of additional variables, including body size and breeding stage may have to be considered as explanations for why patterns in some species deviated from our predictions. Nevertheless, comparative studies like ours are important for improving our understanding of the hormonal and reproductive responses of vertebrates to large scale weather patterns and global climate change in general.
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http://dx.doi.org/10.1016/j.ygcen.2017.10.015DOI Listing
April 2018

Strong association between corticosterone levels and temperature-dependent metabolic rate in individual zebra finches.

J Exp Biol 2017 12 19;220(Pt 23):4426-4431. Epub 2017 Oct 19.

Groningen Institute for Evolutionary Life Sciences, University of Groningen, 9747 AG Groningen, The Netherlands.

Glucocorticoid hormones (GCs) are often assumed to be indicators of stress. At the same time, one of their fundamental roles is to facilitate metabolic processes to accommodate changes in energetic demands. Although the metabolic function of GCs is thought to be ubiquitous across vertebrates, we are not aware of experiments which tested this directly, i.e. in which metabolic rate was manipulated and measured together with GCs. We therefore tested for a relationship between plasma corticosterone (CORT; ln transformed) and metabolic rate (MR; measured using indirect calorimetry) in a between- and within-individual design in captive zebra finches () of both sexes. In each individual, CORT and MR were measured at two different temperature levels: 'warm' (22°C) and 'cold' (12°C). CORT and MR were both increased in colder compared with warmer conditions within individuals, but also across individuals. At the between-individual level, we found a positive relationship between CORT and MR, with an accelerating slope towards higher MR and CORT values. In contrast, the within-individual changes in CORT and MR in response to colder conditions were linearly correlated between individuals. The CORT-MR relationship did not differ between the sexes. Our results illustrate the importance of including variation at different levels to better understand physiological modulation. Furthermore, our findings support the interpretation of CORT variation as an indicator of metabolic needs.
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http://dx.doi.org/10.1242/jeb.166124DOI Listing
December 2017

Flexible clock systems: adjusting the temporal programme.

Philos Trans R Soc Lond B Biol Sci 2017 Nov;372(1734)

Chronobiology Unit, Groningen Institute for Evolutionary Life Sciences, University of Groningen, Groningen, The Netherlands.

Under natural conditions, many aspects of the abiotic and biotic environment vary with time of day, season or even era, while these conditions are typically kept constant in laboratory settings. The timing information contained within the environment serves as critical timing cues for the internal biological timing system, but how this system drives daily rhythms in behaviour and physiology may also depend on the internal state of the animal. The disparity between timing of these cues in natural and laboratory conditions can result in substantial differences in the scheduling of behaviour and physiology under these conditions. In nature, temporal coordination of biological processes is critical to maximize fitness because they optimize the balance between reproduction, foraging and predation risk. Here we focus on the role of peripheral circadian clocks, and the rhythms that they drive, in enabling adaptive phenotypes. We discuss how reproduction, endocrine activity and metabolism interact with peripheral clocks, and outline the complex phenotypes arising from changes in this system. We conclude that peripheral timing is critical to adaptive plasticity of circadian organization in the field, and that we must abandon standard laboratory conditions to understand the mechanisms that underlie this plasticity which maximizes fitness under natural conditions.This article is part of the themed issue 'Wild clocks: integrating chronobiology and ecology to understand timekeeping in free-living animals'.
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http://dx.doi.org/10.1098/rstb.2016.0254DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5647281PMC
November 2017

Timing as a sexually selected trait: the right mate at the right moment.

Philos Trans R Soc Lond B Biol Sci 2017 Nov;372(1734)

Chronobiology unit, Groningen Institute for Evolutionary Life Sciences, University of Groningen, The Netherlands.

Sexual selection favours the expression of traits in one sex that attract members of the opposite sex for mating. The nature of sexually selected traits such as vocalization, colour and ornamentation, their fitness benefits as well as their costs have received ample attention in field and laboratory studies. However, sexually selected traits may not always be expressed: coloration and ornaments often follow a seasonal pattern and behaviours may be displayed only at specific times of the day. Despite the widely recognized differences in the daily and seasonal timing of traits and their consequences for reproductive success, the actions of sexual selection on the temporal organization of traits has received only scant attention. Drawing on selected examples from bird and mammal studies, here we summarize the current evidence for the daily and seasonal timing of traits. We highlight that molecular advances in chronobiology have opened exciting new opportunities for identifying the genetic targets that sexual selection may act on to shape the timing of trait expression. Furthermore, known genetic links between daily and seasonal timing mechanisms lead to the hypothesis that selection on one timescale may simultaneously also affect the other. We emphasize that studies on the timing of sexual displays of both males and females from wild populations will be invaluable for understanding the nature of sexual selection and its potential to act on differences within and between the sexes in timing. Molecular approaches will be important for pinpointing genetic components of biological rhythms that are targeted by sexual selection, and to clarify whether these represent core or peripheral components of endogenous clocks. Finally, we call for a renewed integration of the fields of evolution, behavioural ecology and chronobiology to tackle the exciting question of how sexual selection contributes to the evolution of biological clocks.This article is part of the themed issue 'Wild clocks: integrating chronobiology and ecology to understand timekeeping in free-living animals'.
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http://dx.doi.org/10.1098/rstb.2016.0249DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5647276PMC
November 2017

Temporal dynamics of the HPA axis linked to exploratory behavior in a wild European songbird (Parus major).

Gen Comp Endocrinol 2017 09 19;250:104-112. Epub 2017 Jun 19.

Department of Animal Ecology, Netherlands Institute of Ecology (NIOO-KNAW), Droevendaalsesteeg 10, 6708 PB Wageningen, The Netherlands.

Variation in the reactivity of the endocrine stress axis is thought to underlie aspects of persistent individual differences in behavior (i.e. animal personality). Previous studies, however, have focused largely on estimating baseline or peak levels of glucocorticoids (CORT), often in captive animals. In contrast, the temporal dynamics of the HPA axis-how quickly it turns on and off, for example-may better indicate how an individual copes with stressors. Moreover, these HPA components might be correlated, thereby representing endocrine suites. Using wild-caught great tits (Parus major) we tested birds for exploratory behavior using a standardized novel environment assay that serves as a validated proxy for personality. We then re-captured a subset of these birds (n=85) and characterized four components of HPA physiology: baseline, endogenous stress response, a dexamethasone (DEX) challenge to estimate the strength of negative feedback, and an adrenocorticotropic hormone (ACTH) challenge to estimate adrenal capacity. We predicted that these four HPA responses would be positively correlated and that less exploratory birds would have a more rapid onset of the stress response (a CORT elevation during the baseline bleed) and weaker negative feedback (higher CORT after DEX). We found support for the first two predictions but not the third. All four components were positively correlated with each other and less exploratory birds exhibited an elevation in CORT during the baseline bleed (<3min from capture). Less exploratory birds, however, did not exhibit weaker negative feedback following the DEX challenge, but did exhibit weaker adrenal capacity. Together, our findings provide partial support for the hypothesis that the temporal reactivity of the HPA axis is linked with consistent individual differences in behavior, with more cautious (slower exploring) individuals exhibiting a faster CORT response.
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http://dx.doi.org/10.1016/j.ygcen.2017.06.011DOI Listing
September 2017

Effects of developmental conditions on glucocorticoid concentrations in adulthood depend on sex and foraging conditions.

Horm Behav 2017 07 9;93:175-183. Epub 2017 Jun 9.

Max Planck Institute for Ornithology, Eberhard-Gwinner-Straße, 82319 Starnberg, Germany.

Developmental conditions in early life frequently have long-term consequences on the adult phenotype, but the adult environment can modulate such long-term effects. Glucocorticoid hormones may be instrumental in mediating developmental effects, but the permanency of such endocrine changes is still debated. Here, we manipulated environmental conditions during development (small vs. large brood size, and hence sibling competition) and in adulthood (easy vs. hard foraging conditions) in a full factorial design in zebra finches, and studied effects on baseline (Bas-CORT) and stress-induced (SI-CORT) corticosterone in adulthood. Treatments affected Bas-CORT in females, but not in males. Females reared in small broods had intermediate Bas-CORT levels as adults, regardless of foraging conditions in adulthood, while females reared in large broods showed higher Bas-CORT levels in hard foraging conditions and lower levels in easy foraging conditions. Female Bas-CORT was also more susceptible than male Bas-CORT to non-biological variables, such as ambient temperature. In line with these results, repeatability of Bas-CORT was higher in males (up to 51%) than in females (25%). SI-CORT was not responsive to the experimental manipulations in either sex and its repeatability was high in both sexes. We conclude that Bas-CORT responsiveness to intrinsic and extrinsic conditions is higher in females than in males, and that the expression of developmental conditions may depend on the adult environment. The latter finding illustrates the critical importance of studying of causes and consequences of long-term developmental effects in other environments in addition to standard laboratory conditions.
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http://dx.doi.org/10.1016/j.yhbeh.2017.05.020DOI Listing
July 2017

Risk-averse personalities have a systemically potentiated neuroendocrine stress axis: A multilevel experiment in Parus major.

Horm Behav 2017 07 26;93:99-108. Epub 2017 May 26.

Evolutionary Physiology Group, Max Planck Institute for Ornithology, Eberhard-Gwinner-Str., 82319 Seewiesen, Germany; Department of Biology, Universitätsstrasse 10, University of Konstanz, Konstanz, Germany. Electronic address:

Hormonal pleiotropy-the simultaneous influence of a single hormone on multiple traits-has been hypothesized as an important mechanism underlying personality, and circulating glucocorticoids are central to this idea. A major gap in our understanding is the neural basis for this link. Here we examine the stability and structure of behavioral, endocrine and neuroendocrine traits in a population of songbirds (Parus major). Upon identifying stable and covarying behavioral and endocrine traits, we test the hypothesis that risk-averse personalities exhibit a neuroendocrine stress axis that is systemically potentiated-characterized by stronger glucocorticoid reactivity and weaker negative feedback. We show high among-individual variation and covariation (i.e. personality) in risk-taking behaviors and demonstrate that four aspects of glucocorticoid physiology (baseline, stress response, negative feedback strength and adrenal sensitivity) are also repeatable and covary. Further, we establish that high expression of mineralocorticoid and low expression of glucocorticoid receptor in the brain are linked with systemically elevated plasma glucocorticoid levels and more risk-averse personalities. Our findings support the hypothesis that steroid hormones can exert pleiotropic effects that organize behavioral phenotypes and provide novel evidence that neuroendocrine factors robustly explain a large fraction of endocrine and personality variation.
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http://dx.doi.org/10.1016/j.yhbeh.2017.05.011DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5552616PMC
July 2017

Endocrine mechanisms, behavioral phenotypes and plasticity: known relationships and open questions.

Front Zool 2015 24;12 Suppl 1:S7. Epub 2015 Aug 24.

Max Planck Institute for Ornithology, Eberhard-Gwinner-Str., D-82319 Seewiesen, Germany.

Behavior of wild vertebrate individuals can vary in response to environmental or social factors. Such within-individual behavioral variation is often mediated by hormonal mechanisms. Hormones also serve as a basis for among-individual variations in behavior including animal personalities and the degree of responsiveness to environmental and social stimuli. How do relationships between hormones and behavioral traits evolve to produce such behavioral diversity within and among individuals? Answering questions about evolutionary processes generating among-individual variation requires characterizing how specific hormones are related to variation in specific behavioral traits, whether observed hormonal variation is related to individual fitness and, whether hormonal traits are consistent (repeatable) aspects of an individual's phenotype. With respect to within-individual variation, we need to improve our insight into the nature of the quantitative relationships between hormones and the traits they regulate, which in turn will determine how they may mediate behavioral plasticity of individuals. To address these questions, we review the actions of two steroid hormones, corticosterone and testosterone, in mediating changes in vertebrate behavior, focusing primarily on birds. In the first part, we concentrate on among-individual variation and present examples for how variation in corticosterone concentrations can relate to behaviors such as exploration of novel environments and parental care. We then review studies on correlations between corticosterone variation and fitness, and on the repeatability over time of corticosterone concentrations. At the end of this section, we suggest that further progress in our understanding of evolutionary patterns in the hormonal regulation of behavior may require, as one major tool, reaction norm approaches to characterize hormonal phenotypes as well as their responses to environments. In the second part, we discuss types of quantitative relationships between hormones and behavioral traits within individuals, using testosterone as an example. We review conceptual models for testosterone-behavior relationships and discuss the relevance of these models for within-individual plasticity in behavior. Next, we discuss approaches for testing the nature of quantitative relationships between testosterone and behavior, concluding that again reaction norm approaches might be a fruitful way forward. We propose that an integration of new tools, especially of reaction norm approaches into the field of behavioral endocrinology will allow us to make significant progress in our understanding of the mechanisms, the functional implications and the evolution of hormone-behavior relationships that mediate variation both within and among individuals. This knowledge will be crucial in light of already ongoing habitat alterations due to global change, as it will allow us to evaluate the mechanisms as well as the capacity of wild populations to adjust hormonally-mediated behaviors to altered environmental conditions.
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http://dx.doi.org/10.1186/1742-9994-12-S1-S7DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4722346PMC
February 2016
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