A Dynamic Energy Budget model based on partitioning of net production

We formulate a Dynamic Energy Budget (DEB) model for the growth and reproduction of individual organisms based on partitioning of net production (i.e. energy acquisition rate minus maintenance rate) between growth and energy reserves. Reproduction uses energy from reserves. The model describes both feeding and non-feeding stages, and hence is applicable to embryos (which neither feed nor reproduce), juveniles (which feed but do not reproduce), and adults (which commonly both feed and reproduce). Embryonic growth can have two forms depending on the assumptions for acquisition of energy from yolk. By default, when the energy acquisition rate exceeds the maintenance rate, a fixed proportion of the resulting net production is spent on growth (increase in structural biomass), and the remaining portion is channelled to the reserves. Feeding organisms, however, modulate their allocation of net production energy in response to their total energy content (energy in the reserves plus energy bounded to structural biomass). In variable food environment an organism alternates between periods of growth, no-growth, and balanced-growth. In the latter case the organism adopts an allocation strategy that keeps its total energy constant. Under constant environmental conditions, the growth of a juvenile is always of von Bertalanffy type. Depending on the values of model parameters there are two long-time possibilities for adults: (a) von Bertalanffy growth accompanied by reproduction at a rate that approaches zero as the organism approaches asymptotic size, or (b) abrupt cessation of growth at some finite time, following which, the rate of reproduction is constant. We illustrate the model's applicability in life history theory by studying the optimum values of the energy allocation parameters for constant environment and for each of the dynamic regimes described above. Received: 11 May 1998 / Revised version: 18 February 2000 / Published online: 4 October 2000     

Magellanic penguin telomeres do not shorten with age with increased reproductive effort,investment, and basal corticosterone

All species should invest in systems that enhance longevity; however, a fundamental adult life‐history trade‐off exists between the metabolic resources allocated to maintenance and those allocated to reproduction. Long‐lived species will invest more in reproduction than in somatic maintenance as they age. We investigated this trade‐off by analyzing correlations among telomere length, reproductive effort and output, and basal corticosterone in Magellanic penguins (Spheniscus magellanicus). Telomeres shorten with age in most species studied to date, and may affect adult survival. High basal corticosterone is indicative of stressful conditions. Corticosterone, and stress, has been linked to telomere shortening in other species. Magellanic penguins are a particularly good model organism for this question as they are an unusually long‐lived species, exceeding their mass‐adjusted predicted lifespan by 26%. Contrary to our hypothesis, we found adults aged 5 years to over 24 years of age had similar telomere lengths. Telomeres of adults did not shorten over a 3‐year period, regardless of the age of the individual. Neither telomere length, nor the rate at which the telomeres changed over these 3 years, correlated with breeding frequency or investment. Older females also produced larger volume clutches until approximately 15 years old and larger eggs produced heavier fledglings. Furthermore, reproductive success (chicks fledged/eggs laid) is maintained as females aged. Basal corticosterone, however, was not correlated with telomere length in adults and suggests that low basal corticosterone may play a role in the telomere maintenance we observed. Basal corticosterone also declined during the breeding season and was positively correlated with the age of adult penguins. This higher basal corticosterone in older individuals, and consistent reproductive success, supports the prediction that Magellanic penguins invest more in reproduction as they age. Our results demonstrate that telomere maintenance may be a component of longevity even with increased reproductive effort, investment, and basal corticosterone.     

Developmental energetics of zebrafish, Danio rerio

Using zebrafish (Danio rerio) as a case study, we show that the maturity concept of Dynamic Energy Budget (DEB) theory is a useful metric for developmental state. Maturity does not depend on food or temperature contrary to age and to some extent length. We compile the maturity levels for each developmental milestone recorded in staging atlases. The analysis of feeding, growth, reproduction and aging patterns throughout the embryo, juvenile and adult life stages are well-captured by a simple extension of the standard DEB model and reveals that embryo development is slow relative to adults. A threefold acceleration of development occurs during the larval period. Moreover we demonstrate that growth and reproduction depend on food in predictable ways and their simultaneous observation is necessary to estimate parameters. We used data on diverse aspects of the energy budget simultaneously for parameter estimation using the covariation method. The lowest mean food intake level to initiate reproduction was found to be as high as 0.6 times the maximum level. The digestion efficiency for Tetramin? was around 0.5, growth efficiency was just 0.7 and the value for the allocation fraction to soma (0.44) was close to the one that maximizes ultimate reproduction.     

Trade‐offs between reproduction and self‐maintenance (immune function and body mass) in a small seabird,the little auk

Breeding season is the most energetically and physiologically demanding phase in the avian annual cycle, challenging adults' physiology and survival. However, the timing and extent that self‐maintenance of breeding adults is compromised during the breeding season is poorly understood. We investigated the trade‐off between reproduction and self‐maintenance in relation to breeding phase (prelaying, incubation, chick rearing) and sex in a small Arctic seabird, the little auk Alle alle. To measure a bird's allocation of time for self‐maintenance, we examined size‐adjusted body mass and immunocompetence expressed by bacteria (Escherichia coli) killing capacity (BKC) of blood plasma, heterophils/lymphocyte ratio (H/L) and their numbers of particular leucocytes per 10 000 red blood cells (RBC). We found that size‐adjusted body mass decreased as the breeding season progressed. BKC, number of heterophils and H/L values were all was significantly higher at prelaying when compared to all other phases. Interestingly, we found that heavier individuals had higher BKC and number heterophils at the prelaying and chick rearing phases than light individuals. There were no differences by sex in any studied variables. Our results indicate that immunocompetence and body mass of breeding adults decreases over the course of breeding season. The efficiency of the immune system appears to be dependent on the bird's body reserves. Our results suggest that little auks allocation of resources into reproduction negatively affects their self‐maintenance.     

Experimental testing of dynamic energy budget models

1. Dynamic energy budget (DEB) models describing the allocation of assimilate to the competing processes of growth, reproduction and maintenance in individual organisms have been applied to a variety of species with some success. There are two contrasting model formulations based on dynamic allocation rules that have been widely used (net production and net assimilation formulations). However, the predictions of these two classes of DEB models are not easily distinguished on the basis of simple growth and fecundity data.
2. It is shown that different assumptions incorporated in the rules determining allocation to growth and reproduction in two classes of commonly applied DEB models predict qualitatively distinct patterns for an easily measured variable, cumulative reproduction by the time an individual reaches an arbitrary size.
3. A comparison with experimental data from Daphnia pulex reveals that, in their simplest form, neither model predicts the observed qualitative pattern of reproduction, despite the fact that both formulations capture basic growth features.
4. An examination of more elaborate versions of the two models, in which the allocation rules are modified to account for brief periods of starvation experienced in the laboratory cultures, reveals that a version of the net production model can predict the qualitative pattern seen for cumulative eggs as a function of mass in D. pulex . The analysis leads to new predictions which can be easily tested with further laboratory experiments.     

Parameter Estimations of Dynamic Energy Budget (DEB) Model over the Life History of a Key Antarctic Species: The Antarctic Sea Star Odontaster validus Koehler, 1906

Marine organisms in Antarctica are adapted to an extreme ecosystem including extremely stable temperatures and strong seasonality due to changes in day length. It is now largely accepted that Southern Ocean organisms are particularly vulnerable to global warming with some regions already being challenged by a rapid increase of temperature. Climate change affects both the physical and biotic components of marine ecosystems and will have an impact on the distribution and population dynamics of Antarctic marine organisms. To predict and assess the effect of climate change on marine ecosystems a more comprehensive knowledge of the life history and physiology of key species is urgently needed. In this study we estimate the Dynamic Energy Budget (DEB) model parameters for key benthic Antarctic species the sea star Odontaster validus using available information from literature and experiments. The DEB theory is unique in capturing the metabolic processes of an organism through its entire life cycle as a function of temperature and food availability. The DEB model allows for the inclusion of the different life history stages, and thus, becomes a tool that can be used to model lifetime feeding, growth, reproduction, and their responses to changes in biotic and abiotic conditions. The DEB model presented here includes the estimation of reproduction handling rules for the development of simultaneous oocyte cohorts within the gonad. Additionally it links the DEB model reserves to the pyloric caeca an organ whose function has long been ascribed to energy storage. Model parameters described a slowed down metabolism of long living animals that mature slowly. O. validus has a large reserve that—matching low maintenance costs- allow withstanding long periods of starvation. Gonad development is continuous and individual cohorts developed within the gonads grow in biomass following a power function of the age of the cohort. The DEB model developed here for O. validus allowed us to increase our knowledge on the ecophysiology of this species, providing new insights on the role of food availability and temperature on its life cycle and reproduction strategy.     

Explaining sex differences in lifespan in terms of optimal energy allocation in the baboon

We provide a quantitative test of the hypothesis that sex role specialization may account for sex differences in lifespan in baboons if such specialization causes the dependency of fitness upon longevity, and consequently the optimal resolution to an energetic trade‐off between somatic maintenance and other physiological functions, to differ between males and females. We present a model in which females provide all offspring care and males compete for access to reproductive females and in which the partitioning of available energy between the competing fitness‐enhancing functions of growth, maintenance, and reproduction is modeled as a dynamic behavioral game, with the optimal decision for each individual depending upon his/her state and the behavior of other members of the population. Our model replicates the sexual dimorphism in body size and sex differences in longevity and reproductive scheduling seen in natural populations of baboons. We show that this outcome is generally robust to perturbations in model parameters, an important finding given that the same behavior is seen across multiple populations and species in the wild. This supports the idea that sex differences in longevity result from differences in the value of somatic maintenance relative to other fitness‐enhancing functions in keeping with the disposable soma theory.     

Acorns,insects, and the diet of adult versus nestling Acorn Woodpeckers

ABSTRACT The diet of Acorn Woodpeckers (Melanerpes formicivorus) in central coastal California consists of a mixture of high‐quality (insects) and low‐quality (stored acorns) food during the spring breeding season. We used stable isotope ratios obtained from blood samples to estimate possible differences in the proportion of these items being fed to nestlings and being consumed by breeding adults, as well as the extent to which the diet of nestlings shifts between insects and acorns during the nestling period. Based on both feeding observations and items recovered from nestlings, older nestlings were fed 28% acorns. Using this value as a baseline for the isotope analyses, the diet of breeding adults was estimated to consist of 90% acorns. Based on repeated samples from the same nestlings over time, the estimated proportion of acorns in their diet increased from 19% in 9‐ to 12‐d‐old nestlings to 42% in 23‐ to 26‐d‐old nestlings. There was a significant correlation between the isotope values of adults and the nestlings they fed, indicating that different groups of Acorn Woodpeckers have significantly different diets. Although important for successful reproduction in this species, stored acorns are not the primary food resource used to feed young. Instead, the availability of stored acorns allows adult Acorn Woodpeckers to provide nestlings with more protein‐rich insects while maintaining themselves on relatively protein‐poor, low‐quality acorns. Differences in the diets of adults and their nestlings are likely widespread among species that feed, at least in part, on low‐quality foods, such as fruit and seeds, during the breeding season.     

Cold stress and acclimation – what is important for metabolic adjustment?

As sessile organisms, plants are unable to escape from the many abiotic and biotic factors that cause a departure from optimal conditions of growth and development. Low temperature represents one of the most harmful abiotic stresses affecting temperate plants. These species have adapted to seasonal variations in temperature by adjusting their metabolism during autumn, increasing their content of a range of cryo‐protective compounds to maximise their cold tolerance. Some of these molecules are synthesised de novo. The down‐regulation of some gene products represents an additional important regulatory mechanism. Ways in which plants cope with cold stress are described, and the current state of the art with respect to both the model plant Arabidopsis thaliana and crop plants in the area of gene expression and metabolic pathways during low‐temperature stress are discussed.     

Quantifying and understanding reproductive allocation schedules in plants

A plant's reproductive allocation (RA) schedule describes the fraction of surplus energy allocated to reproduction as it increases in size. While theorists use RA schedules as the connection between life history and energy allocation, little is known about RA schedules in real vegetation. Here we review what is known about RA schedules for perennial plants using studies either directly quantifying RA or that collected data from which the shape of an RA schedule can be inferred. We also briefly review theoretical models describing factors by which variation in RA may arise. We identified 34 studies from which aspects of an RA schedule could be inferred. Within those, RA schedules varied considerably across species: some species abruptly shift all resources from growth to reproduction; most others gradually shift resources into reproduction, but under a variety of graded schedules. Available data indicate the maximum fraction of energy allocated to production ranges from 0.1 to 1 and that shorter lived species tend to have higher initial RA and increase their RA more quickly than do longer‐lived species. Overall, our findings indicate, little data exist about RA schedules in perennial plants. Available data suggest a wide range of schedules across species. Collection of more data on RA schedules would enable a tighter integration between observation and a variety of models predicting optimal energy allocation, plant growth rates, and biogeochemical cycles.     

Resource allocation in Wilson's storm-petrels<Emphasis Type="Italic"> Oceanites oceanicus</Emphasis> determined by measurement of glucocorticoid excretion

When resources are limited, life-history theory predicts that long-lived animals should allocate available resources to body maintenance rather than to reproduction in order to maximise their lifetime reproductive success. In the present study, we estimated physiological stress in a small procellariiform seabird, the Wilson's storm-petrel Oceanites oceanicus, as a means of understanding how limited resources are partitioned between provisioning parents and their chicks. We analysed adrenocortical activity of Wilson's storm-petrels during the breeding season by measuring glucocorticoid (GC) excretion, using an enzyme immunoassay measuring tetrahydrocorticosterone concentrations in extracts of faeces and urine of chicks and adults. Faecal GC measures were negatively correlated with chick body condition, suggesting that measures of tetrahydrocorticosterone in faeces and urine can be used to assess adrenal activity characteristic for physiological stress in Wilson's storm-petrels. In the breeding season of 1999, the colony was subject to low food availability, and the faecal and urine GC levels of chicks were elevated during these months of chronic starvation. In contrast, adults did not show elevated GC levels. The data thus suggest that Wilson's storm-petrels respond to unfavourable conditions by maintaining their own body condition and reducing provisioning of food to their chicks. Communicated by R.F. Oliviera     

Cannabis, cannabinoids and reproduction

In most countries Cannabis is the most widely used illegal drug. Its use during pregnancy in developed nations is estimated to be approximately 10%. Recent evidence suggests that the endogenous cannabinoid system, now consisting of two receptors and multiple endocannabinoid ligands, may also play an important role in the maintenance and regulation of early pregnancy and fertility. The purpose of this review is therefore twofold, to examine the impact that cannabis use may have on fertility and reproduction, and to review the potential role of the endocannabinoid system in hormonal regulation, embryo implantation and maintenance of pregnancy.     

Habitat‐driven life history variation in an amphibian metapopulation

Life‐history theory states that, during the lifetime of an individual, resources are allocated to either somatic maintenance or reproduction. Resource allocation tradeoffs determine the evolution and ecology of life‐history strategies and determine an organisms’ position along the fast–slow continuum. Theory predicts that environmental stochasticity is an important driver of resource allocation and therefore life‐history evolution. Highly stochastic environments are expected to increase uncertainty in reproductive success and select for iteroparity and a slowing down of the life history. To date, most empirical studies have used comparisons among species to examine these theoretical predictions. By contrast, few have investigated how environmental stochasticity affects life‐history strategies at the intraspecific level. In this study, we examined how variation in breeding site stochasticity (among‐year variability in pond volume and hydroperiod) promotes the co‐occurrence of different life‐history strategies in a spatially structured population, and determines life‐history position along the fast–slow continuum in the yellow‐bellied toad Bombina variegata. We collected mark–recapture data from a metapopulation and used multievent capture–recapture models to estimate survival, recruitment and breeding probabilities. We found higher survival and longer lifespans in populations inhabiting variable sites compared to those breeding in stable ones. In addition, probabilities of recruitment and skipping a breeding event were higher in variable sites. The temporal variance of survival and recruitment probabilities, as well as the probability to skip breeding, was higher in variable sites. Taken together, these findings indicate that populations breeding in variable sites experienced a slowing down of the life‐history. Our study thus revealed similarities in the macroevolutionary and microevolutionary processes shaping life‐history evolution.     

Twelve fundamental life histories evolving through allocation‐dependent fecundity and survival

An organism's life history is closely interlinked with its allocation of energy between growth and reproduction at different life stages. Theoretical models have established that diminishing returns from reproductive investment promote strategies with simultaneous investment into growth and reproduction (indeterminate growth) over strategies with distinct phases of growth and reproduction (determinate growth). We extend this traditional, binary classification by showing that allocation‐dependent fecundity and mortality rates allow for a large diversity of optimal allocation schedules. By analyzing a model of organisms that allocate energy between growth and reproduction, we find twelve types of optimal allocation schedules, differing qualitatively in how reproductive allocation increases with body mass. These twelve optimal allocation schedules include types with different combinations of continuous and discontinuous increase in reproduction allocation, in which phases of continuous increase can be decelerating or accelerating. We furthermore investigate how this variation influences growth curves and the expected maximum life span and body size. Our study thus reveals new links between eco‐physiological constraints and life‐history evolution and underscores how allocation‐dependent fitness components may underlie biological diversity.     

Survival and production in variable resource environments

A dynamic energy budget (DEB) model describes the rates at which organisms assimilate and utilize energy from food for maintenance, growth, reproduction and development. We study the dynamic behavior of one particular DEB model, Kooijman’s κ rule model, whose key assumption is that somatic and reproductive tissues are competing for energy. We assume an environment in which the food density fluctuates either periodically or stochastically (pink noise). Both types of fluctuations stimulate growth; the magnitude of the (average) increase in size depends on both the strength and duration of the fluctuations. In a stochastic environment, the risk of mortality due to starvation increases with increasing fluctuation intensity. The mean lifespan is also a function of the model parameter κ characterizing the partitioning of energy between somatic and reproductive tissues. Organisms committing a large fraction of resources to reproduction endure periods of food shortage relatively well. The effects of food fluctuations on reproduction are complex. With stochastic food, reproduction in survivors increases with increasing fluctuation intensities, but lifetime reproduction decreases. Periodic fluctuations may enhance reproduction, depending on the value of κ. Thus, a variable food supply stimulates growth, increases mortality and may enhance reproduction, depending on life history.     

Best be(e) on low fat: linking nutrient perception,regulation and fitness

Preventing malnutrition through consuming nutritionally appropriate resources represents a challenge for foraging animals. This is due to often high variation in the nutritional quality of available resources. Foragers consequently need to evaluate different food sources. However, even the same food source can provide a plethora of nutritional and non‐nutritional cues, which could serve for quality assessment. We show that bumblebees, Bombus terrestris, overcome this challenge by relying on lipids as nutritional cue when selecting pollen. The bees ‘prioritised’ lipid perception in learning experiments and avoided lipid consumption in feeding experiments, which supported survival and reproduction. In contrast, survival and reproduction were severely reduced by increased lipid contents. Our study highlights the importance of fat regulation for pollen foraging bumblebees. It also reveals that nutrient perception, nutrient regulation and reproductive fitness can be linked, which represents an effective strategy enabling quick foraging decisions that prevent malnutrition and maximise fitness.     

Potential applications of mussel modelling

Mussels are extensively cultivated worldwide and are of growing economic importance. However, constraints on the exploitation of wild mussel resources have necessitated the need for tools to improve the management of mussel cultivation towards increased production. Ecological models are increasingly being used as a management tool, and therefore the existing approaches to modelling mussels have been reviewed with respect to their possible application to the improvement of shellfish management strategies. We suggest that dynamic energy budget (DEB) models have the greatest potential in this area, and discuss the mussel DEB models that have been developed to date in terms of their physiological complexity, accuracy of prediction of individual mussel growth and ability to predict mussel population production. Individual mussel production has been predicted; however, the focus of many of the models has been on the growth and reproduction of a single mussel and therefore population effects generally have not been included. Other models at the population level have included only limited population effects, and this has reduced the capacity of many of the models to accurately predict mussel production at the population level. Interactions at the population level (self-thinning and predation) are discussed and the models that describe the consequences of these processes are examined. In future DEB models will need to include the ability to parameterise population level processes if we are to have greater confidence in their application to shellfish management. Electronic Publication     

DNA polymerase κ‐dependent DNA synthesis at stalled replication forks is important for CHK1 activation

Formation of primed single‐stranded DNA at stalled replication forks triggers activation of the replication checkpoint signalling cascade resulting in the ATR‐mediated phosphorylation of the Chk1 protein kinase, thus preventing genomic instability. By using siRNA‐mediated depletion in human cells and immunodepletion and reconstitution experiments in Xenopus egg extracts, we report that the Y‐family translesion (TLS) DNA polymerase kappa (Pol κ) contributes to the replication checkpoint response and is required for recovery after replication stress. We found that Pol κ is implicated in the synthesis of short DNA intermediates at stalled forks, facilitating the recruitment of the 9‐1‐1 checkpoint clamp. Furthermore, we show that Pol κ interacts with the Rad9 subunit of the 9‐1‐1 complex. Finally, we show that this novel checkpoint function of Pol κ is required for the maintenance of genomic stability and cell proliferation in unstressed human cells.     

On the sustainability of a reintroduced Crested Ibis population in Qinling Mountains,Shaanxi, Central China

Reintroduction projects aim to reestablish a self‐sustaining population of an endangered species within its historical range. Adequate post‐release monitoring by gathering demographic data is important to evaluate the success of a reintroduction. Survival and reproduction rates of a reintroduced population can be compared with a self‐sustaining wild population to evaluate the success of a reintroduction. In early 2007, Nipponia nippon (Crested Ibis) was reintroduced into the Qinling Mountains (Shaanxi, Central China). In this study, we attempt to evaluate the demographic status of the reintroduced population. Age‐specific survival rates of 56 released adults and 77 wild‐born fledglings were estimated using mark‐recapture data obtained from 2007 to 2014. Survival rates for the yearlings (0.599, with 95% confidence interval [CI]: 0.467–0.719) were lower than the estimates from a wild population in Yangxian County, but the survival rates of the adults (0.678, with 95% CI: 0.603–0.745) were similar. The number of breeding pairs gradually increased since 2008, although breeding success (52.5%) was somewhat less than that of the wild population (67.6%). The stochastic estimation of population growth rate (1.084 with 95% CI: 1.069–1.098) and population size (5‐fold increase) estimated from an age‐classified Leslie matrix indicate that the reintroduced population of the Crested Ibis is more likely in regulation phase over the next 25 years. We conclude that the reintroduction of the Crested Ibis in Qinling Mountains has great promise, and progress toward a self‐sustaining population has been made under some interventions. Governments, local communities, and scientists need to facilitate habitat restoration for the long‐term survival of this endangered species.