Thermal acclimation of metabolic rate may be seasonally dependent in the subtropical anuran Latouche's frog (Rana latouchii, Boulenger)

We tested the hypothesis that the lack of metabolic thermal acclimation ability in tropical and subtropical amphibians is dependent on season and investigated the effects of body size, sex, time of day, and season on metabolic rates in Rana latouchii. The males were acclimated at 15 degrees, 20 degrees, and 25 degrees C, and their oxygen consumption was measured at 15 degrees, 20 degrees, 25 degrees, and 30 degrees C in all four seasons, with the exception that we did not measure oxygen consumption at 30 degrees C in winter frogs. We also acclimated the males at 30 degrees C in summer for investigating diel variation of metabolic rate. The females were acclimated at 20 degrees and 25 degrees C, and their oxygen consumption was measured at 15 degrees , 20 degrees , 25 degrees , and 30 degrees C in summer. Our results showed that metabolic rates of R. latouchii differed by time of day, season, and acclimation temperature but did not differ by sex if the results were adjusted for differences in body mass. Summer males exhibited a 26%-48% increase in metabolic rates from the lowest values in the seasons. There was a trend of increased oxygen consumption in cold-acclimated males, but it was significant only at 15 degrees and 25 degrees C in summer, autumn, and winter. These results support the hypothesis that thermal acclimation of metabolism is seasonally dependent, which has not been reported in other tropical and subtropical amphibians.     

Corticosterone,testosterone and life-history strategies of birds

Steroid hormones have similar functions across vertebrates, but circulating concentrations can vary dramatically among species. We examined the hypothesis that variation in titres of corticosterone (Cort) and testosterone (T) is related to life-history traits of avian species. We predicted that Cort would reach higher levels under stress in species with higher annual adult survival rates since Cort is thought to promote physiological and behavioural responses that reduce risk to the individual. Conversely, we predicted that peak T during the breeding season would be higher in short-lived species with high mating effort as this hormone is known to promote male fecundity traits. We quantified circulating hormone concentrations and key life-history traits (annual adult survival rate, breeding season length, body mass) in males of free-living bird species during the breeding season at a temperate site (northern USA) and a tropical site (central Panama). We analysed our original data by themselves, and also combined with published data on passerine birds to enhance sample size. In both approaches, variation in baseline Cort (Cort0) among species was inversely related to breeding season length and body mass. Stress-induced corticosterone (MaxCort) also varied inversely with body mass and, as predicted, also varied positively with annual adult survival rates. Furthermore, species from drier and colder environments exhibited lower MaxCort than mesic and tropical species; T was lowest in species from tropical environments. These findings suggest that Cort0, MaxCort and T modulate key vertebrate life-history responses to the environment, with Cort0 supporting energetically demanding processes, MaxCort promoting survival and T being related to mating success.     

Environmental influence on the genetic correlations between life-history traits in Caenorhabditis elegans

Empirical evidence is mounting to suggesting that genetic correlations between life-history traits are environment specific. However, detailed knowledge about the loci underlying genetic correlations in different environments is scant. Here, we studied the influence of temperature (12 degrees C and 24 degrees C) on the genetic correlations between egg size, egg number and body mass in the nematode Caenorhabditis elegans. We used a quantitative trait loci (QTL) approach based on a genetic map with evenly spaced single nucleotide polymorphism markers in an N2 x CB4856 recombinant inbred panel. Significant genetic correlations between various traits were found at both temperatures. We detected pleiotropic or closely linked QTL, which supported the negative correlation between egg size and egg number at 12 degrees C, the positive correlation across temperatures for body mass, and the positive correlation between body mass and egg size at 12 degrees C. The results indicate that specific loci control the covariation in these life-history traits and the locus control is prone to environmental conditions.     

Life history traits associated with body size covary along a latitudinal gradient in a generalist grasshopper

Animal body size often varies systematically along latitudinal gradients, where individuals are either larger or smaller with varying season length. This study examines ecotypic responses by the generalist grasshopper Melanoplus femurrubrum (Orthoptera: Acrididae) in body size and covarying, physiologically based life history traits along a latitudinal gradient with respect to seasonality and energetics. The latitudinal compensation hypothesis predicts that smaller body size occurs in colder sites when populations must compensate for time constraints due to short seasons. Shorter season length requires faster developmental and growth rates to complete life cycles in one season. Using a common garden experimental design under laboratory conditions, we examined how grasshopper body size, consumption, developmental time, growth rate and metabolism varied among populations collected along an extended latitudinal gradient. When reared at the same temperature in the lab, individuals from northern populations were smaller, developed more rapidly, and showed higher growth rates, as expected for adaptations to shorter and generally cooler growing seasons. Temperature-dependent, whole organism metabolic rate scaled positively with body size and was lower at northern sites, but mass-specific standard metabolic rate did not differ among sites. Total food consumption varied positively with body size, but northern populations exhibited a higher mass-specific consumption rate. Overall, compensatory life history responses corresponded with key predictions of the latitudinal compensation hypothesis in response to season length.     

Seasonal metabolism of a small, arboreal monitor lizard, Varanus scalaris, in tropical Australia

The field metabolic rates (FMR) and water fluxes of Varanus scalaris were measured during the wet and dry seasons by the doubly-labelled water technique. Seasonal measurements of standard (night-time) metabolism (SMR) and resting (daytime) metabolism (RMR) were made in the laboratory at 18, 24, 30 and 36°C, and maximal oxygen consumption was measured at 36°C on a motorized treadmill. This population was active throughout the year. In the wet season, the mean FMR was 7.8 kJ day⁻¹ (128.0 kJkg⁻¹ day⁻¹; mean mass = 66.4 g, n = 13), and during the dry season the mean was 5.0 kJ day⁻¹ (67.6 kJ kg⁻¹ day⁻¹; mean mass = 77.4 g, n = 17). The mean water flux rates for these animals were 3.6 and 1.2 ml day⁻¹, respectively (60.4 and 16.6 ml kg⁻¹ day⁻¹). The seasonal means of FMR and water flux were significantly different by ANCOVA ( P < 0.0001). Measurements of SMR and RMR were significantly higher in the wet season (ANCOVA: P < 0.0001), but we found no difference in the maximal oxygen consumption between seasons (ANCOVA: P = 0.6). The maximal oxygen consumption of the lizards on the treadmill (2.9 ml min⁻¹= 1.8 ml g⁻¹ h⁻¹), mean mass = 97.4 g, n = 16) was 20 times that of the SMR at the same temperature during the dry season, and 11 times that of the SMR during the wet season. The seasonal differences in FMR were attributable to: changes in SMR (12.2%) and RMR (16.4%); differences in night-time body temperatures (11.3) and daytime body temperatures (16.4%); and activity (broadly defined to include locomotion, digestion, and reproductive costs (43.7%).     

Inter-sexual differences in resting metabolic rates in the Texas tarantula, Aphonopelma anax

Intra-specific variation in life history and mating strategies can lead to differences in energy allocation and expenditure in males and females. This may, in turn, explain large-scale evolutionary patterns. In this study, I investigated the effects of body mass, temperature and sex on resting metabolic rates (RMRs) in sexually mature male and female tarantulas (Aphonopelma anax (Chamberlin)), a species that exhibits extreme inter-sexual differences in life history after reaching sexual maturity. RMRs were measured as rates of CO(2) production in an open-flow respirometry system at 20, 25, 30 and 35 degrees C. These temperatures are typical to what this species experiences under natural conditions. In addition, a respiratory quotient (RQ) of 0.92 was calculated from rates of CO(2) production and O(2) consumption in a closed, constant-volume respirometry system. As expected, RMRs increased with increasing temperature and body mass. However, after adjusting for the influence of body mass, males had substantially higher metabolic rates than females at each temperature. This higher metabolic rate is proposed as an adaptive strategy to support higher energetic demands for males during their active, locomotory search for females during the mating season.     

Body size response to warming: time of the season matters in a tephritid fly

Whether shrinking body size is a universal response to climate change remains controversial. Moreover, the mechanisms underlying body size shifts are poorly understood. Here, assuming that life history traits evolve to maximize fitness according to life history plasticity theory, we hypothesized that under global warming temperate multivoltine insects should emerge earlier with a smaller body mass in the early growing season, but emerge later with a larger body mass in the late season. We tested this hypothesis by conducting two field artificial warming experiments in an alpine meadow: 1) with one pre‐dispersal seed predator species (tephritid flies, Tephritis femoralis) and its two host‐plant species flowering in early and late growing seasons, respectively, and 2) with the tephritid flies and one host species with a flowering season that occupies parts of both the early and late growing seasons. These experiments were complemented by a microcosm chamber warming experiment, in which increasing and decreasing temperature trends were set to simulate temperature variation pattern in early and late growing seasons, respectively, but photoperiod was held constant. Warming generally advanced the adult emergence and decreased the body size (adult body mass) in the early season but delayed the emergence and increased the size in the late season in both field experiments, indicating that the seasonally different effects of warming on the fly body size was constant regardless of host‐plant identity. The chamber warming resulted in consistent responses of emerging timing and body size to the simulated seasonal warming, demonstrating that the temperature increase per se and its interaction with direction of temperature change, but not other correlated effects, should be primarily responsible for the observed contrasting shifts of body size at different times of the season. Our results indicate that taking into account temperate seasonal patterns of temperature variation could be of general importance for predicting animal body size changes in the warmed future.     

Elevation impacts the balance between growth and oxidative stress in coal tits

The short favorable period of time available for the growth in seasonal environments could constrain the resources allocation between growth and other life-history traits, and the short-term fitness benefits of increased growth rate may prevail over other functions. Accelerated growth rates have been associated with long-term deleterious consequences (e.g., decreased lifespan), and recently oxidative stress (the imbalance between pro-oxidants generation and antioxidant defenses) has been suggested as a mediator of these effects. Here, we examined the impact of elevation on growth rate and self-maintenance parameters (resting metabolism, oxidative damage, and antioxidant defenses) of coal tit chicks (Periparus ater). We predicted that the shorter favorable season at the higher-elevation site could lead to a reallocation of resources towards growth at the expense of self-maintenance processes. We found that chicks at high elevation grew significantly faster in terms of body mass and body size. Chicks from the high-elevation site presented higher resting metabolism, higher oxidative damage level, but similar antioxidant defenses, compared to low-elevation chicks. Interestingly, the chicks exhibiting the better antioxidant defenses at 7 days were also those with the highest resting metabolic rate, and the chicks that grew at the faster rate within the high-elevation site were those with the highest levels of oxidative damage on DNA. Our study supports the idea that increasing elevation leads to a higher growth rate in coal tit chicks, possibly in response to a shorter favorable season. In accordance with life-history theory, a bigger investment in growth was done at the expense of body maintenance, at least in terms of oxidative stress.     

An experimental test of the role of environmental temperature variability on ectotherm molecular, physiological and life-history traits: implications for global warming

Global climate change is one of the greatest threats to biodiversity; one of the most important effects is the increase in the mean earth surface temperature. However, another but poorly studied main characteristic of global change appears to be an increase in temperature variability. Most of the current analyses of global change have focused on mean values, paying less attention to the role of the fluctuations of environmental variables. We experimentally tested the effects of environmental temperature variability on characteristics associated to the fitness (body mass balance, growth rate, and survival), metabolic rate (VCO(2)) and molecular traits (heat shock protein expression, Hsp70), in an ectotherm, the terrestrial woodlouse Porcellio laevis. Our general hypotheses are that higher values of thermal amplitude may directly affect life-history traits, increasing metabolic cost and stress responses. At first, results supported our hypotheses showing a diversity of responses among characters to the experimental thermal treatments. We emphasize that knowledge about the cellular and physiological mechanisms by which animals cope with environmental changes is essential to understand the impact of mean climatic change and variability. Also, we consider that the studies that only incorporate only mean temperatures to predict the life-history, ecological and evolutionary impact of global temperature changes present important problems to predict the diversity of responses of the organism. This is because the analysis ignores the complexity and details of the molecular and physiological processes by which animals cope with environmental variability, as well as the life-history and demographic consequences of such variability.     

Phenotypic plasticity in reproductive traits induced by food availability in a lacertid lizard, Takydromus septentrionalis

Understanding the proximate determinants of phenotypic variations in life-history traits can provide powerful insights into a species' life-history strategies. I experimentally manipulated availability of food (high vs low) to examine plasticity in the reproductive traits of northern grass lizards, Takydromus septentrionalis (Lacertidae), from eastern China. Food availability significantly affected reproductive frequency and thereby seasonal reproductive output, but had little effect on reproductive output per clutch. Low-food females postponed reproduction and produced less clutches in the reproductive season than did high-food females. After producing their second clutches, low-food females were in lower body condition than the high-food counterparts. By the end of the experiment, however, all females exhibited similar body condition. Clutch size and clutch mass differed between the first and second clutches but not between the treatments. Egg size and phenotypic traits of hatchlings (body size, morphology and locomotor performance) in T. septentrionalis did not vary significantly from first to second clutches nor between the two treatments. These results support optimal egg size (offspring) theory. Female T. septentrionali s "decide" whether or not to reproduce largely based on current energy intake; lowered feeding rates thus delay oviposition and reduce reproductive frequency. In contrast, clutch size, egg size and relative clutch mass remain unchanged.     

Plasticity of growth rate and metabolism in Daphnia magna populations from different thermal habitats

The aim of this study was to evaluate the effect of temperature on growth and aerobic metabolism in clones of Daphnia magna from different thermal regimes. Growth rate (increment in size), somatic juvenile growth rate (increment in mass), and oxygen consumption were measured at 15 and 25 degrees C in 21 clones from one northern and two southern sites. There were no significant differences in body size and growth rate (increase in length) at both 15 and 25 degrees C among the three sites. Clones from southern site 2 had a higher mass increment than clones from the other two sites at both temperatures. Clone had a significant effect on growth (body length) and body size at both temperatures. As expected, age at maturity was lower at 25 degrees C (4.5 days) than at 15 degrees C, (11.6 days) and body sizes, after the release of the third clutch, were larger at 15 degrees C than at 25 degrees C. Northern clones had higher oxygen consumption rates and specific dynamic action (SDA) than southern clones at 15 degrees C. By contrast, southern clones from site 1 had a higher oxygen consumption and SDA than subarctic clones at 25 degrees C. Clones from southern site 2 had high oxygen consumption rates at both temperatures. Our results reveal important differences in metabolic rates among Daphnia from different thermal regimes, which were not always reflected in growth rate differences.     

Changes in body composition during breeding: Reproductive strategies of three species of seabirds under poor environmental conditions

Seabirds differ dramatically in life history traits and breeding strategies. For example, gulls have short incubation shifts (several hours) and high metabolic rates, auks have medium-length incubation shifts (12-24h) and high metabolic rates, and petrels have long incubation shifts (days) and low metabolic rates. How these different strategies affect the dynamics of body components is poorly known. We compared body, organ and lipid mass changes among three different seabirds (gull: black-legged kittiwake Rissa tridactyla; auk: thick-billed murre Uria lomvia; petrel: northern fulmar Fulmarus glacialis) at Prince Leopold Island, Nunavut, Canada during 2002 (a year with low reproductive success and poor chick growth across all three species). This study is among the first to compare mass and lipid dynamics among different species foraging in the same food web and at similar trophic levels during the same breeding season (same environmental conditions). In fulmars and murres, most of decreases in body mass reflected decreases in lipid mass while in kittiwakes the increase in body mass reflected an increase in lean mass, especially the muscle. The species with the longest fasting endurance (incubation shift length) had the highest percent body lipids during incubation (fulmars: 13.3%, murres: 7.3%, kittiwakes: 6.9%), the highest variability in body lipids, tended to regulate body mass primarily through lipid stores and tended to regulate exercise and digestive organs separately. In contrast, in the species with the highest metabolic rate, all organ systems were adjusted similarly and in relation to body mass, and in a similar manner between incubation (stress due to heavy ice conditions) and chick-rearing (lower stress due to ice-free conditions). In high metabolic rate species, we suggest that organ size varies in response to environmental stress. We conclude that the organ dynamics of seabirds are set by a combination of key life history traits (like incubation shift and metabolic rate) and environmental conditions.     

Seasonality and the scheduling of life history at different latitudes

For many ectotherms, the annual cycle is partitioned into 'growing' (summer) and 'non-growing' (winter) seasons, and the lengths of these seasons are inversely related across a latitudinal gradient. This pattern of variation has the potential to affect diverse life-history traits profoundly. A key selective agent is size-dependent winter mortality which, with increasing latitude, places an increasing premium on attainment of large body size before the first winter of life. Winter body size is determined primarily by two factors: (1) birth date, which defines the beginning of the first growing season and (2) somatic growth rate within the first season. Using examples drawn from the Atlantic silverside, Menidia menidia (L.), and other fishes, I show how latitudinal variation in the scheduling of the spawning season, countergradient variation in the capacity for growth and reproduction, and mode of sex determination (environmental v. genetic) represent adaptations to seasonality.     

Seasonal changes in the cardiorespiratory responses to hypercarbia and temperature in the bullfrog, Rana catesbeiana

We assessed the seasonal variations in the effects of hypercarbia (3 or 5% inspired CO2) on cardiorespiratory responses in the bullfrog Rana catesbeiana at different temperatures (10, 20 and 30 degrees C). We measured breathing frequency, blood gases, acid-base status, hematocrit, heart rate, blood pressure and oxygen consumption. At 20 and 30 degrees C, the rate of oxygen consumption had a tendency to be lowest during winter and highest during summer. Hypercarbia-induced changes in breathing frequency were proportional to body temperature during summer and spring, but not during winter (20 and 30 degrees C). Moreover, during winter, the effects of CO2 on breathing frequency at 30 degrees C were smaller than during summer and spring. These facts indicate a decreased ventilatory sensitivity during winter. PaO2 and pHa showed no significant change during the year, but PaCO2 was almost twice as high during winter than in summer and spring, indicating increased plasma bicarbonate levels. The hematocrit values showed no significant changes induced by temperature, hypercarbia or season, indicating that the oxygen carrying capacity of blood is kept constant throughout the year. Decreased body temperature was accompanied by a reduction in heart rate during all four seasons, and a reduction in blood pressure during summer and spring. Blood pressure was higher during winter than during any other seasons whereas no seasonal change was observed in heart rate. This may indicate that peripheral resistance and/or stroke volume may be elevated during this season. Taken together, these results suggest that the decreased ventilatory sensitivity to hypercarbia during winter occurs while cardiovascular parameters are kept constant.     

Metabolism, body size and life span: a case study in evolutionarily divergent populations of the garter snake (Thamnophis elegans)

We present a case study of metabolism, life history and aging in the western terrestrial garter snake (Thamnophis elegans). Early research in the field supported the rate-of-living hypothesis as an explanation of aging, which was based on an apparent negative relationship between mass-specific metabolic rate and lifespan in endotherms. This hypothesis in its original form has not withstood additional tests and comparisons between the two main lineages of endotherms-birds and mammals, but there is still much to be discovered of the causative links among rate of oxygen consumption, physiology and life history, particularly in ectothermic reptiles. We present data that show adult short-lived snakes, from naturally occurring ecotypes of garter snakes, have higher mass-specific resting metabolic rates at any given body mass (metabolic intensity) across a series of normal activity temperatures (15-32°C). The short-lived ecotype in this geographic region reaches a larger body size, and has life-history traits that place it at the fast end of a pace-of-life continuum (fast growth, early maturation, high reproductive output) relative to individuals of the small-bodied long-lived ecotype. The difference between ecotypes in metabolic intensity, even after acclimation to identical conditions, may reflect evolutionary divergence and genetic differences between ecotypes. The difference in metabolic intensity is not, however, present at birth, so an alternative is that developmental environment may permanently influence metabolic rate and life history. Such developmental canalization could lead to altered gene expression via environmental influences on the epigenome and result in altered metabolic trajectories in the snakes' natural habitats.     

Trade-offs in the life history and energy budget of the parthenogenetic collembolan Folsomia candida (Willem)

Three clones of Folsomia candida from different locations in Europe were compared in four experiments investigating genetic and phenotypic correlations between life-history traits. The first three experiments focused on the effects of food type, clone and temperature on traits associated with the first clutch. Differences in clutch size between clones and treatments were almost completely attributable to body size. Clones differed in length of the juvenile period, but the difference decreased at low temperatures. Age and weight at first reproduction were negatively correlated in the food type experiment and positively correlated in the temperature experiment, an often-encountered result for which no general explanation is as yet available. In the temperature experiment egg size variation was considerable, and was highest at low temperatures. The fourth experiment, with two clones at two feeding levels, aimed at finding trade-offs, in particular between reproduction and survival. It was hypothesized that higher fecundity led to increased scenescence through a higher metabolic rate. The trade-off was clearly present among the clones: one combined fast growth, late reproduction and high lifetime fecundity with lower survival, while in the other the relation between these traits was opposite. The proposed mechanism, however, was not confirmed, as no difference in metabolic rate was found. The effect of food level was too small to result in significant differences in the life-history traits in either of the clones.     

A Review for Life-history Traits Variation in Frogs Especially for Anurans in China

Environmental variation can promote differentiation in life-history traits in species of anurans. Increased environmental stress usually results in larger age at sexual maturity, older mean age, longer longevity, slower growth, larger body size, and a shift in reproductive allocation from offspring quantity to quality, and a stronger trade-off between offspring size and number. However, previous studies have suggested that there are inconsistent geographical variations in life-history traits among anuran species in China. Hence, we here review the intraspecific patterns and differences in life-history traits(i.e., egg size, clutch size, testes size, sperm length, age at sexual maturity, longevity, body size and sexual size dimorphism) among different populations within species along geographical gradients for anurans in China in recent years. We also provide future directions for studying difference in sperm performance between longer and shorter sperm within a species through transplant experiments and the relationships between metabolic rate and brain size and life-history.     

Life-history correlates of evolution under high and low adult mortality

Abstract. Life-history theory predicts evolutionary changes in reproductive traits and intrinsic mortality rates in response to differences in extrinsic mortality rates. Trade-offs between life- history traits play a pivotal role in these predictions, and such trade-offs are mediated, at least in part, by physiological allocations. To gain insight into these trade-offs, we have been performing a long-term experiment in which we allow fruitflies, Drosophila melanogaster , to evolve in response to high (HAM) and low (LAM) adult mortality rates. Here we analyze the physiological correlates of the life-history trade-offs. In addition to changing development time and early fecundity in the direction predicted, high adult mortality affected three traits expressed early in life–body size, growth rate, and ovariole number–but had little or no effect on body composition (relative fat content), viability, metabolic rate, activity, starvation resistance, or desiccation resistance. Correlations among lines revealed trade-offs between early fecundity, late fecundity, and starvation resistance, which appear to be mediated by differential allocation of lipids.     

Seasonal acclimatization in metabolic rate of the fan-fingered gecko,Ptyodactylus hasselquistii (Reptilia: Gekkonidae)

The resting metabolic rate of the fan-fingered gecko Ptyodactylus hasselquistii of various body masses was determined in relation to ambient temperatures ranging from 20 to 35°C during winter and summer acclimatization. Oxygen consumption (ml g⁻¹ h⁻¹) decreased with increasing mass at each temperature. The intraspecific exponents of body mass in relation to metabolic rate ranged from 0.62 to 0.79. Winter-acclimatized geckos had significantly lower metabolic rates than summer-acclimatized geckos at different temperatures, especially at low temperature (20°C). The pattern of acclimatization exhibited by P. hasselquistii may conserve energy during inactivity in winter and make activity more easily achieved during active seasons.     

Patterns of life-history traits in open-nesting palearctic passerines as a function of the climatic variation of their ranges

Latitude, a surrogate of climatic conditions, is commonly used in the examination of life-history variation. However, the climatic mechanisms underlying latitudinal life-history variation have only rarely been tested. Here, we test whether the number of climates to which species are subjected in their ranges predicts geographical life-history variation. In particular, we examine whether eurytopic species, the range of which covers more climates, show different reproductive effort to stenotopic species, which are distributed over climatically more homogeneous environments. We examined female body mass, egg mass controlled for female body mass, clutch size and the number of breeding attempts per season for 34 sedentary and short-distance migratory passerine species of the Western Palearctic. For each species, we assessed how many climate zones extend over the species' wintering and breeding ranges. We found that avian body mass, and also clutch size, significantly increases with the number of climatic zones extended over the species' wintering range. In turn, species whose breeding ranges span more climates show more breeding attempts per season. Whereas the mass of a single egg declines, clutch size increases with increasing climatic variation in breeding ranges. Our study suggests that the level of climatic variation over species' ranges during and outside the breeding season might be responsible for variation in life-history traits in open-nesting Western Palearctic passerines.