GENETIC CORRELATIONS IN A WILD RODENT: GRASS-EATERS AND FAST-GROWERS EVOLVE HIGH BASAL METABOLIC RATES

Basal metabolic rate (BMR), commonly used as a measure of the cost of living, is highly variable among species, and sources of the variation are subject to an enduring debate among comparative biologists. One of the hypotheses links the variation in BMR with diversity of food habits and life-history traits. We test this hypothesis by asking how BMR of a particular species, the bank vole Myodes (= Clethrionomys ) glareolus , would change under selection for high growth rate (measured as a postweaning body mass change; MD_PW ) and the ability to cope with a low-quality herbivorous diet (measured as body mass change during a four-day test; MD_LQD ). We show that both of the traits are heritable in the narrow sense ( MD_PW : h ²= 0.30; MD_LQD : h ²= 0.19), and are genetically correlated with mass-independent BMR (additive genetic correlation, r_A = 0.28 for MD_PW and 0.37 for MD_LQD ). Thus, both of the traits could change in response to a selection, and the selection would also result in a correlated evolution of the level of metabolism. The results are consistent with the hypothesis that a part of the interspecific variation in BMR evolved in response to selection for life-history and ecological traits such as food habits.     

Basal metabolic rate, body weight and diet in primates: an evaluation of the evidence.

The relationship between basal metabolic rate (BMR), body weight and diet is examined for primates. Contrary to the results reported in several recent works, there is no strong evidence that diet is directly linked to BMR, although a low BMR, relative to body weight, may be found in species with folivorous diets. There is some evidence that nocturnal haplorhine species have a relatively low BMR, but strepsirhines appear to have a uniformly low relative BMR regardless of their primary activity period. The evolution of BMR in primates is discussed in the light of these findings. Some predictions are made about the relative BMRs that should be found for other species whose BMR is, as yet, unknown.     

Heritability of energetics in a wild mammal,the leaf-eared mouse (Phyllotis darwini)

Abstract.— As a first examination of the additive genetic variance of thermoregulatory traits in a natural population of endotherms, we studied the quantitative genetics of key physiological ecology traits in the leaf-eared mouse, Phyllotis darwini. We measured basal metabolic rate (BMR), nonshivering thermogenesis (NST), maximum metabolic rate for thermoregulation (MMR), thermal conductance (C_T), body temperature (T_b), and factorial aerobic scope (FAS) in individuals acclimated to cold and warm conditions. For comparability with previous studies, we included the following morphological traits: foot length (F_L), total length (T_L), body mass (mb, at birth, sexual maturity, 6 months, and 8 months). Variance components were obtained from two different procedures: the expected variance component in an ANOVA Type III sum of squares and an animal model approach using restricted maximum likelihood. Results suggest the presence of additive genetic variance in F_L (h²= 0.47, P = 0.045), C_T of cold-acclimated animals (h²= 0.66, P = 0.041), and night body temperature, measured in cold-acclimated animals (h²= 0.68, P = 0.080). Heritabilities of m_b were near zero at all ages, but maternal effects and common environment effects were high and significant. We found no evidence of additive genetic variance in BMR, NST, MMR, or FAS (i.e., estimates were not significantly different from zero for all tests). Our results are in general agreement with previous studies of mammals that reported low heritability for: (1) BMR and MMR; (2) daytime body temperature; and (3) body mass for wild, but not laboratory or domestic, populations.     

Field metabolic rates of black-browed albatrosses Thalassarche melanophrys during the incubation stage

Field metabolic rates (FMR) and activity patterns of black-browed albatrosses Thalassarche melanophrys were measured while at sea and on nest during the incubation stage at Kerguelen Island, southwestern Indian Ocean. Activity-specific metabolic rates of five albatrosses at sea (FMR_at-sea) were measured using doubly labeled water (DLW), and by equipping birds with wet-dry activity data loggers that determined when birds were in flight or on the water. The metabolic rates of four birds incubating their eggs (FMR_on-nest) were also measured using DLW. The mean±SD FMR_at-sea of albatrosses was 611±96 kJ kg⁻¹ d⁻¹ compared to FMR_on-nest of 196±52 kJ kg⁻¹ d⁻¹. While at sea, albatrosses spent 52.9±8.2% (N=3) of their time in flight and they landed on the water 41.2±13.9 times per day. The FMR of black-browed albatrosses appear to be intermediate to that of three other albatross species. Based on at-sea activity, the power requirement of flight was estimated to be 8.7 W kg⁻¹ (or 4.0×predicted BMR), which is high compared to other albatross species, but may be explained by the high activity levels of the birds when at sea. The FMR_at-sea of albatrosses, when scaled with body mass, are lower than other seabirds of similar body size, which probably reflects the economical nature of their soaring flight.     

Influence of seasonal changes in relative liver size, condition, relative gonad size and variability in ovarian development in multiple spawning fish species used in environmental monitoring programmes

Basic biological information was collected to assist in understanding the biology of selected small-bodied fishes for use in environmental monitoring programmes. Gonado-somatic index ( I _G) and hepato-somatic index ( I _H) profiles of female blacknose dace Rhinichthys atratulus , mummichog Fundulus heteroclitus and golden shiner Notemigonus crysoleucas were similar, decreasing gradually during the spawning season. I _G and I _H profiles of female northern redbelly dace Phoxinus eos peaked twice suggesting oocyte recruitment was continuous throughout the spawning season. Regressions of ovary mass to adjusted body mass ( M _A) were more variable in dace relative to the other fish species. For female blacknose dace, variability in the relationship between ovary mass and M _A was minimized by selecting either 2 year-old fish and fish that weighed 2–4 g.     

Distribution of C3 and C4 grasses along an altitudinal gradient in Central Argentina

The distribution pattern of C₃ and C₄ grasses was studied in eight sites located between 350 m and 2100 m along an altitudinal gradient in Central Argentina. Of 139 taxa fifty-nine are C₃ and eighty C₄. Species of the C₃ tribes (Stipeae, Poeae, Meliceae, Aveneae, Bromeae and Triticeae) and C₃ Paniceae species increase in number at higher elevations; only one C₃ species was found below 650 m. C₄ Aristideae, Pappophoreae, Eragrostideae, Cynodonteae, Andropogoneae and Paniceae increase at lower altitudes. The floristic crossover point is at about 1500 m; the ground cover cross-over point is at about 1000 m. Analysis of the relationships between % C₄ species along the gradient and nine climatic and environmental variables showed the highest correlation with July mean temperature, but all temperature variables show highly significant correlations with % C₄. Correlation with annual rainfall is lower but also significant. These results are consistent with previous research showing the relative importance of C₄ grasses as temperature increases. C₃ species make a high contribution to relative grass coverage below the C₃/C₄ floristic crossover point but are rare below 1000 m.     

Evaporative water loss in two sympatric species of vespertilionid bat, Plecotus auritus and Myotis daubentoni: relation to foraging mode and implications for roost site selection

Simultaneous measures of oxygen consumption and evaporative water loss (EWL) were made in two species of temperate-zone vespertilionid bat ( Plecotus auritus and Myotis daubentoni ; mean body mass 9.12 and 10.12g, respectively) at ambient temperatures (T_a) of 5, 15 and 25°C and variable vapour pressure deficit. EWL was directly dependent on vapour pressure deficit and oxygen consumption and inversely dependent on T_a. EWL was significantly greater in P. auritus than in M. daubentoni. A model for EWL in P. auritus under a variety of environmental conditions (5–25°C and 20–80% relative humidity) suggested that EWL from bats in shallow summer torpor will be lowest at low T_a, and that, except at low (> 50%) relative humidity, EWL from euthermic bats will be lowest at high T_a. At low relative humidity (< 20%), resting bats could lose over 30% of body mass per day (24 h) through evaporation. At high T_a (> 25°C), EWL from euthermic bats could be over 65% lower at high (> 80%) compared to low (< 20%) relative humidity. In bats in shallow summer torpor at low (5°C) T_a the equivalent saving was > 96%. At low relative humidity predicted EWL from bats in shallow summer torpor was 34 to 81% of that from euthermic bats, and at low T_a and high relative humidity was only 2%. In the wild, M. daubentoni has freer access to drinking water than does P. auritus and yet EWL at rest was higher in the latter species. We suggest that post-prandial dumping of urinary water by M. daubentoni leads to a limit in the amount of body water available to this species to cover evaporative losses once within the day roost, which in turn has led to an adaptation of physiology towards the minimization of EWL when at rest.     

Quantitative observations on the pulmonary anatomy of the domestic Muscovy duck (Cairina moschata)

The lungs of five female domestic Muscovy ducks, mean body weight 1.627 kg, total lung volume 48.07 cm³, were analysed by standard morphometric methods. Principal results obtained are: lung volume per unit body weight, 30.17 cm³/g; volume densities of exchange tissue relative to lung volume, 49.24%, blood capillaries relative to exchange tissue, 29.63%, tissue of the blood gas (tissue) barrier relative to exchange tissue, 5.88%; surface area of the blood-gas (tissue) barrier per unit body weight, 30.04 cm²/g; ratios of the surface area of the blood-gas (tissue) barrier per unit volume of the lung and per unit volume of exchange area, 979 cm²/cm³ and 200.06 mm²/mm³, respectively; harmonic and arithmetic mean thicknesses of the tissue barrier, 0.199 μm and 0.303 μm, respectively. The anatomical diffusing capacity of the tissue barrier for oxygen ( DtO₂ ) and the total pulmonary diffusing capacity ( DLO₂ ), 49.58 ml O₂/min/mmHg/kg and 4.55 ml O₂/min/mm Hg/kg, respectively. The lungs of the domestic Muscovy duck appear to be about as well adapted anatomically for gas exchange as the lungs of wild anatid species, and there is no clear evidence that domestication has been associated with any deterioration in the anatomical capacity for oxygen uptake. The weight-specific anatomical diffusing capacity of the lung for oxygen ( DLO₂/W ) was about 3.6 times greater than the weight-specific physiological value, a factor which falls within the expected range.     

Empirical models of the conductance of leaves in apple orchards

Abstract. Seasonal data on leaf conductance (g_l) for three different apple cultivars grown in four separate orchards with different aged trees was studied between 1979 and 1985. A number of empirical models for predicting leaf conductance from environmental measurements were compared using this data and a general method for adapting such models for the prediction of different data sets is proposed. Although stepwise multiple regression identified relative humidity or vapour pressure as important variables, it frequently did not identify the optimal set of independent variables, which often did not include either of these. There was no advantage in regressing g_l against principal components of the environment, rather than against the raw environmental variables. A simple model involving air vapour pressure deficit, air temperature and a hyperbolic function of irradiance was found to explain between 32 and 62% of the variance in g_l for the different data sets. Parameters fitted for one data set led to the effective prediction of g_l in other years or plots. The model fit could generally be improved significantly by including soil moisture deficit among the independent variables.     

Roles of endothermy in niche differentiation for ball-rolling dung beetles (Coleoptera: Scarabaeidae) along an altitudinal gradient

Abstract.  1. An analysis of whether niche differentiation in ball-rolling dung beetles can be explained by the way in which they regulate their body temperature was conducted.
2.  A priori assumptions were: (i) if thermoregulation affects niche partitioning, sympatric species must have different endothermic strategies that minimise encounters; or, alternatively (ii) if two co-occurring species show the same thermoregulation pattern and their flight periods overlap, they might be avoiding competition by exhibiting different resource preferences or different food relocation behaviour.
3. The ball-rolling dung beetles studied showed a hierarchical structure based on the species' endothermic capacity, measured as temperature excess [ T _ex= difference between body temperature ( T _b) and ambient temperature ( T _a)]. Those with a high T _ex (10–15 °C) were located exclusively at altitudes >1000 m a.s.l. On the coastal plains, species with a high T _ex were restricted to flying at night when the T _a was lower. Species with a lower T _ex (less than 10 °C higher than T _a) were found in the coastal plains zone.
4. Where there was sympatry with similar trophic habits, the species involved showed very different thermal niches, and where there was significant overlap of thermal niches between sympatric species, trophic habits of species were very different.
5. The results suggest that it is possible to use the concept of the thermal niche as a tool to explain interspecific interactions and the spatial distribution of species.     

N-3 polyunsaturated fatty acids impair lifespan but have no role for metabolism

Although generally considered as beneficial components of dietary fats, polyunsaturated fatty acids (PUFA) have been suspected to compromise maximum lifespan (MLSP) in mammals. Specifically, high amounts of phospholipid PUFAs are thought to impair lifespan due to an increase in the susceptibility of membranes to lipid peroxidation and its damaging effect on cellular molecules. Also, there is evidence from in vitro studies suggesting that highly unsaturated PUFAs elevate basal metabolic rate (BMR). Previous comparative studies in this context were based on small sample sizes, however, and, except for one study, failed to address possible confounding influences of body weight and taxonomic relations between species. Therefore, we determined phospholipid membrane composition in skeletal muscle from 42 mammalian species to test for a relation with published data on MLSP, and with literature data on BMR (30 species). Using statistical models that adjust for the effects of body weight and phylogeny, we found that among mammals, MLSP indeed decreases as the ratio of n-3 to n-6 PUFAs increases. In contrast to previous studies, we found, however, no relation between MLSP and either membrane unsaturation (i.e. PUFA content or number of double bonds) or to the very long-chain, highly unsaturated docosahexaenoic acid (DHA). Similarly, our data set gave no evidence for any notable relation between muscle phospholipid fatty acid composition and BMR, or MLSP and BMR in mammals. These results contradict the 'membrane pacemaker theory of aging', that is, the concept of a direct link between high amounts of membrane PUFAs, elevated BMR, and thus, impaired longevity.     

Bioenergetic Constraints on Primate Abundance

Explaining variation in primate population densities is central to understanding primate ecology, evolution, and conservation. Yet no researchers to date have successfully explained variation in primate population density across dietary class and phylogeny. Most previous work has focused on measures of food availability, as access to food energy likely constrains the number of individuals supported in a given area. However, energy output may provide a measure of energy constraints on population density that does not require detailed data on food availability for a given taxon. Across mammals, many studies have shown that population densities generally scale with body mass^−0.75. Because individual energy expenditures scale with body mass^0.75, population energy use (the product of population density and individual energy use) does not change with body mass, which suggests the existence of energy constraints on population density across body sizes, i.e., taxa are limited to a given amount of energy use, constraining larger taxa to lower densities. We examined population energy use and individual energy expenditure in primates and tested this energy equivalence across body mass. We also used a residual analysis to remove the effects of body mass on primate population densities and energy expenditures using basal metabolic rates (BMR; kcal/d) as a proxy for total daily energy expenditure. After taking into account phylogeny, population energy use did not significantly correlate with body mass. Larger primates, which use more energy per day, live at lower population densities than smaller primates. In addition, we found a significant negative correlation between residuals of BMR from body mass and residuals of population density from body mass after taking phylogeny into account. Thus, energy costs constrain population density across a diverse sample of primates at a given body mass, and primate species that have relatively low BMRs exist at relatively high densities. A better understanding of the determinants of primate energy costs across geography and phylogeny will ultimately help us explain and predict primate population densities.     

Amphipod gigantism dictated by oxygen availability?

A recent study of gigantism in amphipods proposes a novel mechanism which could be used to explain Bergmann's rule, namely differences in environmental O₂ concentration. Consideration of the respiratory biology of this group indicates significant flaws in this proposal. Nonetheless, it is not unreasonable to predict a relationship between ectotherm body size and O₂ partial pressure ( p O₂) in some systems, where the partial pressure gradient is steep, either in time (comparison of Carboniferous and Permian atmospheres) or space (animals inhabiting oceanic O₂ minimum layers, and, particularly in freshwaters, living at high altitude). Animal gigantism can be correlated with periods of very high environmental p O₂ over geological time. Unfortunately, data for present day spatial patterns are scant.     

Response of beech (Fagus sylvatica) to elevated CO2 and N: Influence on larval performance of the gypsy moth Lymantria dispar (Lep., Lymantriidae)

Two-year-old beech seedlings were kept from germination to bioassays with Lymantria dispar under the following conditions: ambient CO₂/low N, elevated CO₂/low N, ambient CO₂/elevated N, and elevated CO₂/elevated N. The effect of these growing conditions of the trees on the performance of the defoliator L. dispar was studied 2 years after initiating the tree cultivation. The developmental success of third-instar larvae of L. dispar was characterized by the weight gained, percentage of weight gain, relative growth rate (RGR), relative consumption rate (RCR), and efficiency of conversion of ingested food into body substance (ECI). Contrary to our expectations, additional N-fertilization did not increase and elevated CO₂ did not delay larval growth rate. However, the environmental treatments of the beech seedlings were found to affect the larval performance. Larvae consumed significantly higher amounts of foliage (RCR) on beech trees under controlled conditions (ambient CO₂ and low N) compared to those under elevated CO₂ and enhanced N. The opposite was true for ECI. The lowest efficacy to convert consumed food to body substance was observed under control conditions and the highest when the larvae were kept on beech trees grown under elevated CO₂ and additional N-fertilization. These opposite effects resulted in the weight gain-based parameters (absolute growth, percentage of growth, and RGR) of the gypsy moth larvae remaining unaffected. The results indicate that the gypsy moth larvae are able to change their ECI and RCR to obtain a specific growth rate. This is discussed as an adaptation to specific food qualities.     

Correlation between digestion rate and feeding frequency in the stomachless teleost, Blennius pholis L.

Foregut emptying in Blennius pholis at 16–18°C is exponential ( S_t = S_o e^-Rt; R=0.006 when t is in min and s in mg dry weight). In consequence, the time for the foregut to empty completely varies in proportion to ln (meal size) in a chosen size of fish or to ln (body weight) in different sized fish fed to the same relative amount (% body weight). Appetite returns rapidly in comparison with other species (4–15 h for fish of 0.5−30 g) and, because of the tidal constraints imposed on this littoral fish, voluntarily ingested meals are large (14–9% body weight depending on fish size). Studies on digestion in small, stomachless fish are not adequately made using X-radiography and must be supported by sequential gut samples after a meal.     

Basal energy metabolism of mustelids

Summary Oxygen consumption was measured during sleep in seven species of mustelids. Their body weight ranged from 50 g to 15 kg. When basal metabolic rate (BMR) was plotted against weight on logarithmic coordinates (Fig. 1), a break in the linearity appeared at a weight level of about one kg. In species with a body weight below one kg, the regression line of BMR against weight is best represented by the equation M = 95.8W^0.55 (±0.03; standard error of estimate) where M is basal metabolic rate in kcal/day and W is body weight in kg. The equation M = 84.6 W^0.78 (±0.15) describes the relationship of animals weighing one kg or more, indicating that the BMR is proportional to almost the same fractional power of body weight, 0.75, as that of other mammals. The high BMR observed in weasels and stoats, suggests that a metabolic adjustment has occurred in the smaller species of the mustelid family.     

Postcanine tooth size in female primates

The results of many allometric studies of postcanine tooth size in mammals have not corresponded to expectations of tooth size based on energy requirements and dental function. The purpose of this study is to investigate the relationship between postcanine occlusal surface area, body size, and the metabolic demands of pregnancy and lactation in female primates. Tooth and body sizes from 38 primate species were taken from the literature to test two hypotheses: 1) females should have relatively larger teeth than males in order to masticate additional food for the energetic costs of reproduction; 2) taxa with the largest neonatal size (a measure of average metabolic costs of pregnancy and lactation) should have females with a greater degree of relative dental enlargement. The results show that relatively large female teeth are not found consistently in primate species. Females have less occlusal surface area than expected on the basis of the male tooth and body size regression in 21% of the species, and there is no correlation between relative female tooth size and relative newborn size across higher primate taxa. The degree of female dental enlargement is most closely related to degree of sexual dimorphism in body weight. The correlation between degree of body weight dimorphism and relatively larger postcanine teeth in females than in males is 0.87 in the 38 species. Species that are monomorphic in weight tend to be monomorphic in tooth size even though females apparently require more food than males.(ABSTRACT TRUNCATED AT 250 WORDS)     

The photosynthetic drought physiology of a diverse group of southern hemisphere conifer species is correlated with minimum seasonal rainfall

1. The instantaneous and integrated leaf gas exchange of 13 species of southern hemisphere conifers grown under identical glasshouse conditions were examined to determine whether there was any correlation between the characteristics of water use at the leaf level and environmental water availability.
2. In the conifer species examined, the minimum ratio of internal to ambient CO₂ measured in leaves during artificially imposed drought [( c _i/ c _a)_min] was strongly correlated with the minimum rainfall observed within the natural range of each species. This suggests that the distributions of these species are constrained by the drought tolerance of their photosynthetic apparatus.
3. A good correlation was found between the ratio of internal to ambient CO₂ measured in leaves under optimal conditions ( c _i/ c _a)_max and leaf δ¹³C (and hence inferred ∫[ c _i/ c _a]). Neither of these, however, correlated with the environmental parameters considered most likely to be limiting species distribution, i.e. precipitation and altitude.
4. These data suggest that decreasing water availability may have been the major factor responsible for the restriction and extinction of conifers in the southern hemisphere.     

Phenotypic flexibility in basal metabolic rate is associated with rainfall variability among populations of rufous-collared sparrow

Phenotypic flexibility in metabolic rates allows organisms to reversibly adjust their energy flow to meet challenges imposed by a variable environment. In turn, the food habits hypothesis (FHH) predicts that species or populations adjust their basal metabolic rate (BMR) according to the diet attributes such as food abundance or predictability. Desert ecosystems represent a temporally heterogeneous environment because of low rain pulse predictability, which is also associated with temporal variation in food resources. In the present study, we investigated the relationship between the magnitude of BMR flexibility in response to dietary acclimation and the inter-annual rainfall variability in three populations of rufous-collared sparrows. Specifically we addressed the question of whether birds from a desert environment are more flexible in BMR than those from non-desert habitats. We found a positive trend between BMR flexibility and the inter-annual rainfall variability. In fact, dietary treatments had a significant effect only in desert birds, a result that also supported the FHH. Our study confirms the existence of phenotypic variation in response to environmental conditions among populations, and also highlights the importance of considering the circumstances in which phenotypic flexibility evolves and the specific environmental cues that induce their expression.     

Effect of body weight and temperature on the maximum daily food consumption of Gasterosteus aculeatus L. and Phoxinus phoxinus (L.): selecting an appropriate model

The relationships between the maximum, daily food consumption, body weight and water temperature for the stickleback, Gasterosteus aculeatus L. , and the minnow, Phoxinus phoxinus (L.), were studied. The two species occupy the same lakes in mid-Wales, but differ in that G. aculeatus has a true stomach whereas P. phoxinus lacks a true stomach.
For both species, the relationship between food consumption (F) and body weight ( W ) could be described by the model: log_eF=a + b log_e W . The effect of temperature (T) could be incorporated in the form: log_ef= a + b, log_e W+b₂log_e T. These models were selected because they accounted for a high proportion of the variance in the dependent variable, and because an analysis of the residuals obtained after fitting a model showed no trends which would have signalled the inadequacy of the model.