Simon Verhulst

University of Groningen | RUG · Groningen Institute for Evolutionary Life Sciences

Sexual conflict occurs when the optimal solution regarding e.g. a life history trait differs between co-operating individuals of different sex. When deciding a conflict is not instantaneous, some form of negotiation can be expected to evolve. In great tits, Parus major, a sexual conflict exists over the number of clutches that are reared, because t...

Biomarkers of aging are essential to predict mortality and aging related diseases. Paradoxically, age itself imposes a limitation on the use of known biomarkers of aging, because their associations with mortality generally diminish with age. How this pattern arises is however not understood. With meta-analysis we show that human leucocyte telomere...

Developmental stressors often have long-term fitness consequences, but linking offspring traits to fitness prospects has remained a challenge. Telomere length predicts mortality in adult birds, and may provide a link between developmental conditions and fitness prospects. Here, we examine the effects of manipulated brood size on growth, telomere dy...

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 's...

Telomere length (TL) predicts health and survival across taxa. Variation in TL between individuals is thought to be largely of genetic origin, but telomere inheritance is unusual, because zygotes already express a TL phenotype, the TL of the parental gametes. Offspring TL changes with paternal age in many species including humans, presumably throug...

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 fu...

Growth trajectories of young animals are intimately connected to their fitness prospects, but we have little knowledge of growth regulation mechanisms, particularly in the wild. Insulin-like growth factor 1 (IGF-1) is a central hormone in regulating resource allocation, with higher IGF-1 levels resulting in more growth. IGF-1 levels generally incre...

Exposure to rising sublethal temperatures can affect development and somatic condition, and thereby Darwinian fitness. In the context of climate warming, these changes could have implications for population viability, but they can be subtle and consequently difficult to quantify. Using telomere length (TL) as a known biomarker of somatic condition...

Early-life environment can affect organisms for life on many levels. The glucocorticoid receptor (GR) gene has a pivotal role mediating organismal physiological and behavioral responses to environmental change, and is sensitive to early-life environmental conditions and epigenetic programming. Longitudinal studies require non-lethal sampling of per...

Early-life conditions impact fitness, but whether the combined effect of extrinsic stressors is additive or synergistic is not well known. This is a major knowledge gap, because exposure to multiple stressors is likely to be frequent. Telomere dynamics may be instrumental when testing whether combined stressor effects are additive or synergistic, b...

Inferring the chronological and biological age of individuals is fundamental to population ecology and our understanding of ageing itself, its evolution, and the biological processes that affect or even cause ageing. Epigenetic clocks based on DNA methylation (DNAm) at specific CpG sites show a strong correlation with chronological age in humans, a...

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 fu...

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 fu...

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 fu...

Young animals need to grow to a large body size fast to maximise their survival prospects until sexual maturity. However, body size varies substantially in wild populations, and neither the selection pressures maintaining this variation, nor the regulatory mechanisms are well understood. IGF-1 administration has been shown to accelerate growth, but...

Variation in developmental conditions is known to affect fitness in later life, but the mechanisms underlying this relationship remain elusive. We previously found in jackdaws Corvus monedula that larger eggs resulted in larger nestlings up to fledging. Through a cross‐foster experiment of complete clutches we tested whether this association can be...

Suboptimal conditions during development can shorten telomeres, the protective DNA caps on the end of chromosomes. Shorter early-life telomere length (TL) can indicate reduced somatic maintenance, leading to lower survival and shorter lifespan. However, despite some clear evidence, not all studies show a relationship between early-life TL and survi...

An accurate inference of the chronological and biological age of individuals is fundamental to population ecology and our understanding of ageing itself, its evolution and the biological processes that affect or even cause ageing. In humans, epigenetic clocks based on the DNA methylation (DNAm) at selected CpG sites correlate highly with chronologi...

Collisions between birds and airplanes can damage aircrafts, resulting in delays and cancellation of flights, costing the international civil aviation industry more than 1.4 billion US dollars annually. Driving away birds is therefore crucial, but the effectiveness of current deterrence methods is limited. Live avian predators can be an effective d...

The innate immune system is essential for survival, yet many immune traits are highly variable between and within individuals. In recent years, attention has shifted to the role of environmental factors in modulating this variation. A key environmental factor is food availability, which plays a major role in shaping life histories, and may affect r...

Variation in developmental conditions is known to affect fitness in later life, but the mechanisms underlying this relationship remain elusive. We previously found in jackdaws Corvus monedula that larger eggs resulted in larger nestlings up to fledging. Through a cross-foster experiment of complete clutches we tested whether this association can be...

Climate warming is increasingly exposing wildlife to sublethal high temperatures, which may lead to chronic impacts and reduced fitness. Telomere length (TL) may link heat exposure to fitness, particularly at early-life stages, because developing organisms are especially vulnerable to adverse conditions, adversity can shorten telomeres, and TL pred...

Food availability modulates survival, reproduction and thereby population size. In addition to direct effects, food availability has indirect effects through density of conspecifics and predators. We tested the prediction that food availability in isolation affects reproductive success by experimentally manipulating food availability continuously f...

Collisions between birds and airplanes, bird strikes, can damage aircrafts, resulting in delays and cancellation of flights, costing the international civil aviation industry more than 1.4 billion U.S. dollars annually. Bird deterrence is therefore crucial, but the effectiveness of all available deterrence methods is limited. For example, live avia...

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) in zebra finches, we tested for a link between the capacity to regulate glucose levels and survival. We also inve...

Mercury is a heavy metal, which is pervasive and persistent in the marine environment. It bioaccumulates within organisms and biomagnifies in the marine food chain. Due to its high toxicity, mercury contamination is a major concern for wildlife and human health. Telomere length is a biomarker of aging and health, because it predicts survival, makin...

Telomere length (TL) shortens with age but telomere dynamics can relate to fitness components independent of age. Immune function often relates to such fitness components and can also interact with telomeres. Studying the link between TL and immune function may therefore help us understand telomere–fitness associations. We assessed the relationship...

Background Severe COVID-19 T-cell lymphopenia is more common among older adults and entails poor prognosis. Offsetting the decline in T-cell count during COVID-19 demands fast and massive T-cell clonal expansion, which is telomere length (TL)-dependent. Methods We developed a model of TL-dependent T-cell clonal expansion capacity with age and virt...

Quantifying an individual’s state as fitness proxy has proven challenging, but accumulating evidence suggests that telomere length and attrition may indicate individual somatic state and success at self‐maintenance, respectively. Sexual ornamentation is also thought to signal phenotypic quality, but links between telomeres and sexual ornamentation...

Costs of reproduction shape the life-history evolution of investment in current and future reproduction and thereby aging. Androgens have been proposed to regulate the physiology governing these investments. Furthermore, androgens are hypothesized to play a central role in carotenoid-dependent sexual signaling, regulating how much carotenoids are d...

Telomeres are protective, nucleoprotein structures at the end of chromosomes that have been associated with lifespan across taxa. However, the extent to which these associations can be attributed to absolute length versus the rate of telomere shortening prior to sampling remains unresolved. In a longitudinal study, we examined the relationship betw...

Recent technological advancements allow researchers to measure electrophysiological parameters of animals, such as sleep, in remote locations by using miniature dataloggers. Yet, continuous recording of sleep might be constrained by the memory and battery capacity of the recording devices. These limitations can be alleviated by recording intermitte...

Aging is associated with declines in physiological performance; declining immune defenses particularly could have consequences for age-related fitness and survival. In aging vertebrates, adaptive (memory-based) immune responses typically become impaired, innate (nonspecific) responses undergo lesser declines, and inflammation increases. Longitudina...

Birds exposed to food insecurity—defined as temporally variable access to food—respond adaptively by storing more energy. To do this, they may reduce energy allocation to other functions such as somatic maintenance and repair. To investigate this trade-off, we exposed juvenile European starlings ( Sturnus vulgaris , n = 69) to 19 weeks of either un...

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 gram backpack for a full year. In addition, we ana...

Telomere length and telomere shortening predict survival in many organisms. This raises the question of the contribution of genetic and environmental effects to variation in these traits, which is still poorly known, particularly for telomere shortening. We used experimental (cross‐fostering) and statistical (quantitative genetic ‘animal’ models) m...

Body condition is an important concept in behaviour, evolution and conservation, commonly used as a proxy of an individual's performance, for example in the assessment of environmental impacts. Although body condition potentially encompasses a wide range of health state dimensions (nutritional, immune or hormonal status), in practice most studies o...

Telomeres are protective, nucleoprotein structures at the end of chromosomes that have been associated with lifespan across taxa. However, the extent to which these associations can be attributed to absolute length versus the rate of telomere shortening prior to sampling remains unresolved. In a longitudinal study, we examined the relationship betw...

Quantifying an individual’s state as fitness proxy has proven challenging, but accumulating evidence suggests that telomere length and attrition may indicate individual somatic state and success at self-maintenance, respectively. Sexual ornamentation is also thought to signal phenotypic quality, but links between telomeres and sexual ornamentation...

The slow pace of global vaccination and the rapid emergence of SARS-CoV-2 variants suggest recurrent waves of COVID-19 in coming years. Therefore, understanding why deaths from COVID-19 are highly concentrated among older adults is essential for global health. Severe COVID-19 T-cell lymphopenia is more common among older adults, and it entails poor...

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 gram backpack for a full year. In addition, we ana...

Early life conditions can impact individuals for life, with harsh developmental conditions resulting in lower fitness, but the underlying mechanisms are not well understood. We hypothesised that immune function may be part of the underlying mechanism, when harsh developmental conditions result in less effective immune function. We tested this hypot...

Recent hypotheses propose that the human placenta and chorioamniotic membranes (CAMs) experience telomere length (TL)-mediated senescence. These hypotheses are based on mean TL (mTL) measurements, but replicative senescence is triggered by short and dysfunctional telomeres, not mTL. We measured short telomeres by a vanguard method, the Telomere sho...

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. Here we investigated metabolic ageing in outdoor‐living captive zebra finches experiencing foraging costs. We longitudinally monitored 407 individuals over 6 years and...

Many organisms are capable of growing faster than they do. Restrained growth rate has functionally been explained by negative effects on lifespan of accelerated growth. However, the underlying mechanisms remain elusive. Telomere attrition has been proposed as a causal agent and has been mostly studied in endothermic vertebrates. We established that...

Profound T-cell lymphopenia is the hallmark of severe coronavirus disease 2019 (COVID-19). T-cell proliferation is telomere length (TL) dependent and telomeres shorten with age. Older COVID-19 patients, we hypothesize, are, therefore, at a higher risk of having TL-dependent lymphopenia. We measured TL by the novel Telomere Shortest Length Assay (Te...

Use of telomere length (TL) as a biomarker for various environmental exposures and diseases has increased in recent years. Various methods have been developed to measure telomere length. Polymerase chain reaction (PCR)-based methods remain wide-spread for population-based studies due to the high-throughput capability. While several studies have eva...

Telomeres are protective caps at the end of eukaryotic chromosomes that shorten with age and in response to stressful or resource‐demanding conditions. Their length predicts individual health and lifespan across a wide range of animals, but whether the observed positive association between telomere length and lifespan is environmentally induced, or...

Telomere length (TL) and shortening rate predict survival in many organisms. Evolutionary dynamics of TL in response to survival selection depend on the presence of genetic variation that selection can act upon. However, the amount of standing genetic variation is poorly known for both TL and TL shortening rate, and has not been studied for both tr...

1. 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 w...

Background: Lymphopenia due to a plummeting T-cell count is a major feature of severe COVID-19. T-cell proliferation is telomere length (TL)-dependent and TL shortens with age. Older persons are disproportionally affected by severe COVID-19, and we hypothesized that those with short TL have less capacity to mount an adequate T-cell proliferative r...

Use of telomere length (TL) as a biomarker for various environmental exposures and diseases has increased in recent years. Various methods have been developed to measure telomere length. PCR-based methods remain wide-spread for population-based studies due to the high-throughput capability. While several studies have evaluated TL measurement method...

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...

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...

The capacity to adequately respond to (physiological) perturbations is a fundamental aspect of physiology, and may affect health and thereby Darwinian fitness. However, little is known of the degree of individual variation in this capacity in non-model organisms. The glucose tolerance test evaluates the individual’s ability to regulate circulating...

Many organisms are capable of growing faster than they do. Restrained growth rate has functionally been explained by negative effects on lifespan of accelerated growth. However, the underlying mechanisms remain elusive. Telomere attrition has been proposed as a causal agent and has been studied in endothermic vertebrates. We established that telome...

Individuals in free‐living animal populations generally differ substantially in reproductive success, lifespan and other fitness‐related traits and the molecular mechanisms underlying this variation are poorly understood. Telomere length and dynamics are candidate traits explaining this variation, as long telomeres predict a higher survival probabi...

Sleep is considered to be of crucial importance for performance and health, yet much of what we know about sleep is based on studies in a few mammalian model species under strictly controlled laboratory conditions. Data on sleep in different species under more natural conditions may yield new insights in the regulation and functions of sleep. We th...

Importance Leukocyte telomere length (LTL) is a trait associated with risk of cardiovascular disease and cancer, the 2 major disease categories that largely define longevity in the United States. However, it remains unclear whether LTL is associated with the human life span. Objective To examine whether LTL is associated with the life span of cont...

The assumption that reproductive effort decreases somatic state, accelerating ageing, is central to our understanding of life‐history variation. Maximal reproductive effort early in life is predicted to be maladaptive by accelerating ageing disproportionally, decreasing fitness. Optimality theory predicts that reproductive effort is restrained earl...

Telomere length (TL) and shortening is increasingly shown to predict variation in survival and lifespan, raising the question of what causes variation in these traits. Oxidative stress is well known to accelerate telomere attrition in vitro, but its importance in vivo is largely hypothetical. We tested this hypothesis experimentally by supplementin...

Telomeres are protective caps at the end of chromosomes, and their length is positively correlated with individual health and lifespan across taxa. Longitudinal studies have provided mixed results regarding the within-individual repeatability of telomere length. While some studies suggest telomere length to be highly dynamic and sensitive to resour...

Most of our knowledge about the regulation and function of sleep is based on studies in a restricted number of mammalian species, particularly nocturnal rodents. Hence, there is still much to learn from comparative studies in other species. Birds are interesting because they appear to share key aspects of sleep with mammals, including the presence...

Crucial to our understanding of the ageing process is identifying how traits change with age, which variables alter their ageing process and whether these traits associate with lifespan. We here investigated metabolic ageing in zebra finches. We longitudinally monitored 407 individuals during six years and collected 3213 measurements of two indepen...

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 test...

Telomeres, the complex of the repeated DNA sequence TTAGGGn and associated proteins at the end of linear chromosomes, are attracting increasing attention because of associations of telomere length with morbidity and mortality (e.g. Boonekamp, Simons, Hemerik & Verhulst, 2013; Verhulst et al., 2016). There are multiple techniques to measure telomere...