Declining body size: a third universal response to warming?

A recently documented correlate of anthropogenic climate change involves reductions in body size, the nature and scale of the pattern leading to suggestions of a third universal response to climate warming. Because body size affects thermoregulation and energetics, changing body size has implications for resilience in the face of climate change. A review of recent studies shows heterogeneity in the magnitude and direction of size responses, exposing a need for large-scale phylogenetically controlled comparative analyses of temporal size change. Integrative analyses of museum data combined with new theoretical models of size-dependent thermoregulatory and metabolic responses will increase both understanding of the underlying mechanisms and physiological consequences of size shifts and, therefore, the ability to predict the sensitivities of species to climate change.     

Life-history responses of the rice stem borer Chilo suppressalis to temperature change: Breaking the temperature–size rule

Temperature is a key environmental factor for ectotherms and affects a large number of life history traits. In the present study, development time from hatching to pupation and adult eclosion, pupal and adult weights of the rice stem borer, Chilo suppressalis were examined at 22, 25, 28 and 31 °C under L18:D 6. Larval and pupal times were significantly decreased with increasing rearing temperature and growth rate was positively correlated with temperature. Larval and pupal developmental times were not significantly different between females and males. The relationship between body weight and rearing temperature in C. suppressalis did not follow the temperature–size rule (TSR), both males and females gained the highest body weight at 31 °C. Females were significantly larger than males at all temperatures, showing a female biased sex size dimorphism (SSD). Contrary to Rensch's rule, SSD and body weight in C. suppressalis tended to increase with rising temperature. Male pupae lost significantly more weight at metamorphosis compared to females. We discuss the adaptive significance of the reverse-TSR in the moth's life history.     

Cope's rule in cryptodiran turtles: do the body sizes of extant species reflect a trend of phyletic size increase?

Cope's rule of phyletic size increase is questioned as a general pattern of body size evolution. Most studies of Cope's rule have examined trends in the paleontological record. However, neontological approaches are now possible due to the development of model-based comparative methods, as well as the availability of an abundance of phylogenetic data. I examined whether the phylogenetic distribution of body sizes in extant cryptodiran turtles is consistent with Cope's rule. To do this, I examined body size evolution in each of six major clades of cryptodiran turtles and also across the whole tree of cryptodirans (n = 201 taxa). Extant cryptodiran turtles do not appear to follow Cope's rule, as no clade showed a significant phyletic body size trend. Previous analyses in other extant vertebrates have also found no evidence for phyletic size increase, which is in contrast to the paleontological data that support the rule in a number of extinct vertebrate taxa.     

Phenotypic plasticity of body size in a temperate population ofDrosophila melanogaster: When the temperature—size rule does not apply

A natural population ofDrosophila melanogaster in southern France was sampled in three different years and 10 isofemale lines were investigated from each sample. Two size-related traits, wing and thorax length, were measured and the wing/thorax ratio was also calculated. Phenotypic plasticity was analysed after development at seven different constant temperatures, ranging from 12‡C to 31‡C. The three year samples exhibited similar reaction norms, suggesting a stable genetic architecture in the natural population. The whole sample (30 lines) was used to determine precisely the shape of each reaction norm, using a derivative analysis. The practical conclusion was that polynomial adjustments could be used in all cases, but with different degrees: linear for the wing/thorax ratio, quadratic for thorax length, and cubic for wing length. Both wing and thorax length exhibited concave reaction norms, with a maximum within the viable thermal range. The temperatures of the maxima were, however, quite different, around 15‡C for the wing and 19.5‡C for the thorax. Assuming that thorax length is a better estimate of body size, it is not possible to state that increasing the temperature results in monotonically decreasing size (the temperature-size rule), although this is often seen to be the case for genetic variations in latitudinal clines. The variability of the traits was investigated at two levels—within and between lines—and expressed as a coefficient of variation. The within-line (environmental) variability revealed a regular, quadratic convex reaction norm for the three traits, with a minimum around 21‡C. This temperature of minimum variability may be considered as a physiological optimum, while extreme temperatures are stressful. The between-line (genetic) variability could also be adjusted to quadratic polynomials, but the curvature parameters were not significant. Our results show that the mean values of the traits and their variance are both plastic, but react in different ways along a temperature gradient. Extreme low or high temperatures decrease the size but increase the variability. These effects may be considered as a functional response to environmental stress.     

Metabolic responses of the body to breathing with hypoxic argon–oxygen and nitrogen–oxygen mixtures

We have studied metabolic responses of six male volunteers who were exposed to hypoxic nitrogen–oxygen and argon–oxygen respiratory mixtures under hermetic chamber conditions for a long time. We measured the values of 44 biochemical parameters of venous blood, which reflected the state of different parts of metabolism, as well as the state of organs and tissues. Under the conditions of argon–oxygen respiratory mixture with the level of oxygen of 12.8% within the first five days and 12.0% from six to ten days of the experiment, the activation of processes of anaerobic glycolysis and lipolysis with the development of metabolic acidosis was observed. Against this background, biochemical signs of unfavorable changes in the myocardium were recorded. When we used the nitrogen–oxygen respiratory mixture with the level of oxygen of 13.0% during the first five days of the experiment and 12.1% from the sixth to the tenth day, these changes were expressed much more significantly: the activation of lipolysis and a decrease in the level of serum iron were observed and the signs of damage of mitochondria, depression of the kidney function, and development of hypodynamia were recorded. Complete stabilization of metabolic processes was reached only after seven to eight days of the recovery period for both mixtures. These findings lead to the conclusion that it is physiologically more preferable to use oxygen–argon respiratory mixtures at small immersion depths.     

Evolution of body size in food webs: does the energetic equivalence rule hold?

The energetic equivalence rule (EER), which is derived from empirical observations linking population density and body size and from the allometric law linking metabolism and body size, predicts that the amount of energy used by the various species should be independent of body size. Here, we examine this hypothesis using a model that allows entire food webs to emerge from coevolution of interacting species. Body size influences both individual metabolism and interactions among species in the model. Overall, population density does decrease with body size roughly following a power law whose exponent is variable. We discuss this variability in the light of empirical data sets. The emerging relationship between the flux of resources exploited by the various species and their body size follows a decreasing power law, thus contradicting the EER. Our model emphasizes the importance of considering the influence of body size on species interactions in attempting to explain large-scale patterns related to body size.     

Effect of temperature on the dose–response curves for auxin-induced elongation growth in maize coleoptile segments

The dose–response curves for IAA-induced growth in maize coleoptile segments were studied as a function of time and temperature. In addition, the kinetics of growth rate responses at some auxin concentrations and temperatures was also compared. It was found that the dose–response curves for IAA-induced elongation growth were, independently of time and temperature, bell-shaped with an optimal concentration at 10⁻⁵ M IAA. The kinetics of IAA-induced growth rate responses depended on IAA concentration and temperature, and could be separated into two phases (biphasic reaction). The first phase (very rapid) was followed by a long lasting one (second phase), which began about 30 min after auxin addition. For coleoptile segments incubated at 30°C, the amplitudes of the first and second phase were significantly higher, when compared with 25°C, at all IAA concentrations studied. However, when coleoptile segments were incubated at 20°C, the elongation growth of coleoptile segments treated with suboptimal IAA concentrations was diminished, mainly as a result of both phases reduction. In conclusion, we propose that the shape of the dose–response curves for IAA-induced growth in maize coleoptile segments is connected with biphasic kinetic of growth rate response.     

Egg size in relation to body size in rotifers: an indication of reproductive strategy?

Egg sizes and body sizes of 43 egg-bearing rotifer species of numerous, mostly tropical, general have been recorded. Larger absolute egg volumes have been found for larger rotifer species, but the increase was lower than expected in proportion to body size, i.e. the relative egg volume decreased with increasing body size. Obviously the relative investment per offspring is smaller in larger rotifer species.     

Does body size affect the response to exercise in shortnose sturgeon (Acipenser brevirostrum)?

The effect of body size on various hematological variables was examined in juvenile shortnose sturgeon (Acipenser brevirostrum) ranging in mass from 38 to 730 g. The blood was examined for differences in plasma ionic composition (Na⁺, K⁺, Cl^?), blood oxygen carrying capacities (hemoglobin, hematocrit), and plasma metabolite concentrations (lactate, glucose), before and following a standard 5‐min chasing stress. All measured resting hematological variables were size independent in shortnose sturgeon. After exercise, levels of plasma lactate, potassium, and hemoglobin increased in all fish. Only post‐exercise levels of chloride and hemoglobin changed in a size‐dependent manner; however, the relationships were weak. The general lack of a relationship between body size and hematological variables might reflect the narrow range of fish sizes used in the present study. From a practical perspective, the results suggest that when examining the hematological stress response in juvenile shortnose sturgeon, a range of fish sizes could be used. This is important considering the variability in the growth rates of juvenile shortnose sturgeon under laboratory conditions.     

Global warming and Bergmann’s rule: do central European passerines adjust their body size to rising temperatures?

Recent climate change has caused diverse ecological responses in plants and animals. However, relatively little is known about homeothermic animals’ ability to adapt to changing temperature regimes through changes in body size, in accordance with Bergmann’s rule. We used fluctuations in mean annual temperatures in south-west Germany since 1972 in order to look for direct links between temperature and two aspects of body size: body mass and flight feather length. Data from regionally born juveniles of 12 passerine bird species were analysed. Body mass and feather length varied significantly among years in eight and nine species, respectively. Typically the inter-annual changes in morphology were complexly non-linear, as was inter-annual variation in temperature. For six (body mass) and seven species (feather length), these inter-annual fluctuations were significantly correlated with temperature fluctuations. However, negative correlations consistent with Bergmann’s rule were only found for five species, either for body mass or feather length. In several of the species for which body mass and feather length was significantly associated with temperature, morphological responses were better predicted by temperature data that were smoothed across multiple years than by the actual mean breeding season temperatures of the year of birth. This was found in five species for body mass and three species for feather length. These results suggest that changes in body size may not merely be the result of phenotypic plasticity but may hint at genetically based microevolutionary adaptations.     

The temperature–size rule in a rotifer is determined by the mother and at the egg stage

In most ectotherms, compared with development at low temperatures, development at high temperatures results in the acceleration of maturation, which in turn results in a smaller size (temperature–size rule, TSR). It is not known at which developmental stages this thermal response is determined. We exposed different life stages of the rotifer Lecane inermis to 15, 20, or 25 °C to determine whether the TSR in the F1 generation is governed by the thermal conditions experienced by the mothers (F0 generation) during their development, during egg production, or during the development of the eggs or hatchlings. We found that the adult size was affected by the thermal conditions experienced by the mothers and embryos, but not by the conditions during post-hatching growth. We suggest that the thermal plasticity producing the TSR in rotifers may reflect the joint impacts of a maternal effect and a direct effect of the environment during egg development. The two-point control of the TSR resembles the thermal determination of other biological phenomena, similar to the thermally determined sex determination in ectotherms. Our results contribute not only to better understanding the proximate mechanisms of TSR, but also to comprehending the general biological mechanisms of response to temperature, which is one of the most important ecological factors.     

Is bigger really bigger? Differential responses to temperature in measures of body size of the mosquito,Aedes albopictus

When confronted with variation in temperature, most ectotherms conform to a growth rule that "hotter is smaller". This phenomenon can have important implications on population dynamics, interactions with other species, and adaptation to new environments for arthropods. However, the impact of other environmental factors and genetics may affect that general rule. Furthermore, most studies measure a single body part, and do not examine how temperature and other factors alter the allometric relationship between measurements of growth. In this study, we test the hypothesis that temperature and nutrition, while strongly affecting growth, do not change the allometric relationship between body mass and wing length in the mosquito Aedes albopictus. We tested this hypothesis by growing larval mosquitoes from two populations at five temperatures with three food levels. Contrary to our hypothesis, we find that temperature has a profound effect on allometry, with higher temperatures resulting in mosquitoes with shorter wings and greater body mass, and that the effects of temperature are dependent upon available food and population origin. We therefore reject our hypothesis and propose several testable mechanisms that will provide greater insight into the relationship between temperature, food, and measures of growth.     

The effect of body temperature on the dynamic respiratory system compliance–breathing frequency relationship in the rat

The mechanical inhomogeneity of the respiratory system is frequently investigated by measuring the frequency dependence of dynamic compliance, but no data are currently available describing the effects of body temperature variations. The aim of the present report was to study those effects in vivo. Peak airway pressure was measured during positive pressure ventilation in eight anesthetized rats while breathing frequency (but not tidal volume) was altered. Dynamic compliance was calculated as the tidal volume/peak airway pressure, and measurements were taken in basal conditions (mean rectal temperature 37.3 °C) as well as after total body warming (mean rectal temperature 39.7 °C). Due to parenchymal mechanical inhomogeneity and stress relaxation-linked effects, the normal rat respiratory system exhibited frequency dependence of dynamic lung compliance. Even moderate body temperature increments significantly reduced the decrements in dynamic compliance linked to breathing rate increments. The results were analyzed using Student’s and Wilcoxon’s tests, which yielded the same results (p < 0.05). Body temperature variations are known to influence respiratory mechanics. The frequency dependence of dynamic compliance was found, in the experiments described, to be temperature-dependent as temperature variations affected parenchymal mechanical inhomogeneity and stress relaxation. These results suggest that body temperature variations should be taken into consideration when the dynamic compliance–breathing frequency relationship is being examined during clinical assessment of inhomogeneity of lung parenchyma in patients.     

Scaling of species diversity and body mass in mammals: Cope’s rule and the evolutionary cost of large size

Abstract

Equations are constructed describing the inverse correlation of species diversity and body mass in extant and Cenozoic mammals. Cope’s rule, the tendency for many mammal clades to increase in body size through time, through phyletic change in single lineages or turnover within species groups, is interpreted as a probability function reducing diversity potential as a tradeoff for ecological/evolutionary gains. The inverse rule predicts that large species in clades will be less diverse than smaller species and, unless origination rates remain high among smaller clade members, clades conforming to Cope’s rule will decline in diversity, moving towards extinction. This proposition is evaluated in the Cenozoic histories of five North American mammal clades; cotton rats, felids, canids, hyaenodontids, and equids. Diversity potential of different size classes within the 3.75 million year phyletic history of the muskrat, Ondatra zibethicus, is also examined. A corollary prediction of the inverse rule, that large species should have longer durations (species lifespans) than small species, is unresolved. Successful clades maintain small size or a significant number of smaller species relative to clade average size. The potential loss of unique extant large mammal species justifies the conservation effort to protect them. The similarity of scaling exponents of species diversity to mass around a slope of -1.0 suggests that species diversity is correlated with home range size, the latter related to the probability of population fragmentation.     

Exceptions to the temperature–size rule: no Lilliput Effect in end-Permian ostracods (Crustacea) from Aras Valley (northwest Iran)

The body size of marine ectotherms is often negatively correlated with ambient water temperature, as seen in many clades during the hyperthermal crisis of the end-Permian mass extinction (c. 252 Ma). However, in the case of ostracods, size changes during ancient hyperthermal events are rarely quantified. In this study, we evaluate the body size changes of ostracods in the Aras Valley section (northwest Iran) in response to the drastic warming during the end-Permian mass extinction at three taxonomic levels: class, order, species. At the assemblage level, the warming triggers a complete species turnover in the Aras Valley section, with larger, newly emerging species dominating the immediate post-extinction assemblage for a short time. Individual ostracod species and instars do not show dwarfing or a change in body size as an adaptation to the temperature stress during the end-Permian crisis. This may indicate that the ostracods in the Aras Valley section might have been exceptions to the temperature–size rule (TSR), using an adaptation mechanism that does not involve a decrease in body size. This adaptation might be similar to the accelerated development despite constant instar body sizes that can be observed in some recent experimental studies of ostracod responses to thermal stress.     

Resistance to oomycetes: a general role for the hypersensitive response?

Oomycete plant pathogens, such as Phytophthora, downy mildews and Pythium, have devastating disease effects on numerous crop and ornamental plants. Various types of genetic resistance to oomycetes occur in plants, and can be determined at the subspecific or varietal level (race or cultivar-specific resistance), or at the species or genus level (nonhost resistance). In addition, resistance might be a quantitative phenotype (partial resistance). Resistance reactions are often associated with the hypersensitive response – a programed cell death pathway. Recent advances in the genetic, biochemical and cytological characterization of disease resistance suggests that the hypersensitive response is associated with all forms of resistance to Phytophthora and downy mildews. Identification of the resistance genes involved in nonhost and partial resistance to oomycetes remains an important challenge.     

Effect of CO₂ on the ventilatory sensitivity to rising body temperature during exercise

We examined the degree to which ventilatory sensitivity to rising body temperature (the slope of the regression line relating ventilation and body temperature) is altered by restoration of arterial PCO(2) to the eucapnic level during prolonged exercise in the heat. Thirteen subjects exercised for ~60 min on a cycle ergometer at 50% of peak O(2) uptake with and without inhalation of CO(2)-enriched air. Subjects began breathing CO(2)-enriched air at the point that end-tidal Pco(2) started to decline. Esophageal temperature (T(es)), minute ventilation (V(E)), tidal volume (V(T)), respiratory frequency (f(R)), respiratory gases, middle cerebral artery blood velocity, and arterial blood pressure were recorded continuously. When V(E), V(T), f(R), and ventilatory equivalents for O(2) uptake (V(E)/VO(2)) and CO(2) output (V(E)/VCO(2)) were plotted against changes in T(es) from the start of the CO(2)-enriched air inhalation (ΔT(es)), the slopes of the regression lines relating V(E), V(T), V(E)/VO(2), and V(E)/VCO(2) to ΔT(es) (ventilatory sensitivity to rising body temperature) were significantly greater when subjects breathed CO(2)-enriched air than when they breathed room air (V(E): 19.8 ± 10.3 vs. 8.9 ± 6.7 l·min(-1)·°C(-1), V(T): 18 ± 120 vs. -81 ± 92 ml/°C; V(E)/VO(2): 7.4 ± 5.5 vs. 2.6 ± 2.3 units/°C, and V(E)/VCO(2): 7.6 ± 6.6 vs. 3.4 ± 2.8 units/°C). The increase in Ve was accompanied by increases in V(T) and f(R). These results suggest that restoration of arterial PCO(2) to nearly eucapnic levels increases ventilatory sensitivity to rising body temperature by around threefold.     

Hydration of serine–metal cation complexes: implication for the role of water in the origin of homochirality on the Earth

Amino Acids - It may be taken for granted that the processes that occurred in water were of crucial importance for the origin of life. Therefore, it is quite likely that water was also important...