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Theodore Garland

Publications and source records attributed to Theodore Garland.

At least 37 records · Page 2Linked to original sources

Number of arginine-vasopressin neurons in the suprachiasmatic nuclei is not related to level or circadian characteristics of wheel-running activity in house mice.

House mouse lines bidirectionally selected for nest-building behavior show a correlation between number of AVP cells in the suprachiasmatic nuclei (SCN), the master circadian clock in mammals, and level of nest-building behavior as well as a correlation between wheel-running activity and SCN AVP content. Similar genetic correlations between wheel-running activity and nest-building behavior have been reported in house mouse lines selected for increased voluntary wheel-running behavior. These similarities in genetic correlation structure in independently selected mouse lines allowed us to test whether AVP in the SCN and wheel running activity are truly correlated traits under identical testing procedures. In the mouse lines selected for voluntary wheel-running, no difference was found between the lines selected for high levels of voluntary wheel-running and randomly-bred control lines in the number of AVP immunoreactive neurons in the SCN ( F1,6 = 0.09, NS; replicate line effect: F1,22 = 0.05, NS). This finding was confirmed at the level of individual variation, which revealed no relationship between number of AVP neurons in the SCN and total daily activity ( R = -0.086, NS, n = 24), or circadian organization (i.e., the chi-squared periodogram waveform amplitude; R = -0.071, NS). Therefore our data do not support the hypothesis that differences in activity level and the circadian expression of activity in young adult mice are related to differences in the number of AVP-immunoreactive cells in the SCN.

Animals↗

Climatic adaptation and the evolution of basal and maximum rates of metabolism in rodents.

Metabolic rate is a key aspect of organismal biology and the identification of selective factors that have led to species differences is a major goal of evolutionary physiology. We tested whether environmental characteristics and/or diet were significant predictors of interspecific variation in rodent metabolic rates. Mass-specific basal metabolic rates (BMR) and maximum metabolic rates (MMR, measured during cold exposure in a He-O2 atmosphere) were compiled from the literature. Maximum (Tmax) and minimum (Tmin) annual mean temperatures, latitude, altitude, and precipitation were obtained from field stations close to the capture sites reported for each population (N = 57). Diet and all continuous-valued traits showed statistically significant phylogenetic signal, with the exception of mass-corrected MMR and altitude. Therefore, results of phylogenetic analyses are emphasized. Body mass was not correlated with absolute latitude, but was positively correlated with precipitation in analyses with phylogenetically independent contrasts. Conventional multiple regressions that included body mass indicated that Tmax (best), Tmin, latitude, and diet were significant additional predictors of BMR. However, phylogenetic analyses indicated that latitude was the only significant predictor of mass-adjusted BMR (positive partial regression coefficient, one-tailed P = 0.0465). Conventional analyses indicated that Tmax, Tmin (best), and altitude explained significant amounts of the variation in mass-adjusted MMR. With body mass and Tmin in the model, no additional variables were significant predictors. Phylogenetic contrasts yielded similar results. Both conventional and phylogenetic analyses indicated a highly significant positive correlation between residual BMR and MMR (as has also been reported for birds), which is consistent with a key assumption of the aerobic capacity model for the evolution of vertebrate energetics (assuming that MMR and exercise-induced maximal oxygen consumption are positively functionally related). Our results support the hypothesis that variation in environmental factors leads to variation in the selective regime for metabolic rates of rodents. However, the causes of a positive association between BMR and latitude remain obscure. Moreover, an important area for future research will be experiments in all taxa are raised under common conditions to allow definitive tests of climatic adaptation in endotherm metabolic rates and to elucidate the extent of adaptive phenotypic plasticity.

Animals↗

Voluntary running in deer mice: speed, distance, energy costs and temperature effects.

The energetics of terrestrial locomotion are of considerable interest to ecologists and physiologists, but nearly all of our current knowledge comes from animals undergoing forced exercise. To explore patterns of energy use and behavior during voluntary exercise, we developed methods allowing nearly continuous measurements of metabolic rates in freely behaving small mammals, with high temporal resolution over periods of several days. We used this approach to examine relationships between ambient temperature (Ta), locomotor behavior and energy costs in the deer mouse, a small mammal that routinely encounters a large range of temperatures in its natural habitat. We tested for individual consistency in running behavior and metabolic traits, and determined how locomotor costs vary with speed and Ta. Because of the importance of thermoregulatory costs in small mammals, we checked for substitution of exercise heat for thermostatic heat production at Ta below the thermal neutral zone and determined the fraction of the daily energy budget comprising exercise costs. Locomotor behavior was highly variable among individuals but had high repeatability, at least over short intervals. We found few temperature-related changes in speed or distance run, but Ta strongly affected energy costs. Partial substitution of exercise heat for thermogenic heat occurred at low Ta. This reduced energy expenditure during low-temperature running by 23-37%, but running costs comprised a fairly minor fraction of the energy budget, so the daily energy savings via substitution were much smaller. Deer mice did not adjust running speed to maximize metabolic economy, as they seldom used the high speeds that provide the lowest cost of transport. The highest voluntary speeds (4-5 km h(-1)) were almost always below the predicted maximal aerobic speed, and were much less than the species' maximal sprint speed. Maximum voluntarily attained rates of oxygen consumption (VO2) were highest at low Ta, but rarely approached maximal VO2 during forced treadmill exercise. Mean respiratory exchange ratios coincident with maximal voluntary VO2 increased slightly as Ta declined, but were always below 1.0 (another indication that metabolic rate was less than the aerobic maximum). Individuals with high running performance (cumulative distance and running time) had high resting metabolism, which suggests a cost of having high capacity or propensity for activity.

Animals↗

Effects of voluntary exercise and genetic selection for high activity levels on HSP72 expression in house mice.

We studied expression of heat shock protein 72 (HSP72) in female mice from four replicate lines that had been selectively bred for high voluntary wheel running (S) and from four random-bred control lines (C). Mice from generation 23 were sampled after 6 days of wheel access, and those from generation 14 were sampled after 8 wk of access to wheels either free to rotate or locked. Mice from S lines ran approximately 2.6 times as many revolutions per day as did those from C lines. Western blotting of tissues from generation 23 mice indicated that S mice had elevated HSP72 expression in triceps surae muscle, but levels in spleen, kidney, heart, and lung were similar in S and C mice. HSP72 expression in triceps surae from generation 14 mice was measured by ELISA and analyzed with a two-way analysis of covariance. The interaction between wheel type and line type (S vs. C) was statistically significant, and subsequent analyses indicated that S mice had significantly elevated HSP72 expression only when housed with free wheels. Mice with the previously described mini-muscle phenotype (Houle-Leroy P, Guderley H, Swallow JG, and Garland T Jr. Am J Physiol Regul Integr Comp Physiol 284: R433-R443, 2003) occurred in both generations and had elevated HSP72 expression in triceps surae. For the generation 23 sample, wheel running as a covariate had a significant negative association with HSP72 expression, and the effect of line type was still statistically significant. Therefore, the increased HSP72 expression of S mice is not a simple proximate effect of their increased wheel running.

Animals↗

Food wasting by house mice: variation among individuals, families, and genetic lines.

Under ad libitum conditions, laboratory house mice (Mus domesticus) fragment considerable amounts of pelleted food and leave it scattered in their cages. The proportion of food thus wasted (in relation to food eaten) varies remarkably among individuals, from 2% to 40%, but is highly consistent in consecutive trials, even when the mice were moved from 22 to -10 degrees C and food consumption doubled. Food wasting did not differ either between the sexes or between genetic lines that had been selected (10 generations) for high voluntary wheel-running behavior (n=4) and their unselected control lines (n=4). However, it varied significantly among replicate lines within the selection groups and among families within the lines (coefficient of intraclass correlation for full sibs, rhof=0.41 in room temperature trials and rhof=0.34 in cold trials). Moreover, the percent of food wasted was negatively correlated with food consumption in the cold trials (males: r=-.36, females: r=-.20) and with total litter mass at weaning (the litters into which they were born; r=-.24), two traits that may affect Darwinian fitness. We conclude that food wastage should not be ignored without justification in calculations of food consumption. In addition, "table manners" can convey reliable information about family origin of an individual and its quality, and therefore could potentially play a role in establishment of social status.

Animals↗

Predatory aggression, but not maternal or intermale aggression, is associated with high voluntary wheel-running behavior in mice.

Predatory (towards crickets), intermale, and maternal aggression were examined in four replicate lines of mice that had been selectively bred for high wheel-running (S) and in four random-bred control lines (C). In generation 18, individual differences in both predatory and intermale aggression were significantly consistent across four trial days, but predatory and intermale aggression were uncorrelated both at the individual level and among the eight line means. Latencies to attack crickets were significantly lower in S lines as a group. Intermale aggression, however, did not differ between S and C lines. S lines were significantly smaller in body mass, but did not differ in either testes mass or plasma testosterone. In generations 28 and 30, respectively, S and C lines did not differ in either maternal or intermale aggression. However, significant differences among the individual lines were found for maternal aggression, and one S line exhibited an extremely high mean time of aggression (>120 sec for a 5-min test). Maternal and intermale aggression were not correlated among the eight line means or at the level of individual variation. Overall, our results suggest: (1) predatory aggression and voluntary wheel-running are positively related at the genetic level; (2) predatory and intermale aggression are unrelated at a genetic level; and (3) maternal and intermale aggression are not tightly related at the genetic level. Possible relationships between predatory aggression, dopamine, and wheel-running behavior are discussed.

Aggression↗

Exercise increases hippocampal neurogenesis to high levels but does not improve spatial learning in mice bred for increased voluntary wheel running.

The hippocampus is important for the acquisition of new memories. It is also one of the few regions in the adult mammalian brain that can generate new nerve cells. The authors tested the hypothesis that voluntary exercise increases neurogenesis and enhances spatial learning in mice selectively bred for high levels of wheel running (S mice). Female S mice and outbred control (C) mice were housed with and without running wheels for 40 days. 5-Bromodeoxyuridine was used to label dividing cells. The Morris water maze was used to measure spatial learning. C runners showed a strong positive correlation between running distance and new cell number, as well as improved learning. In S runners, neurogenesis increased to high levels that reached a plateau, but no improvement in learning occurred. This is the first evidence that neurogenesis can occur without learning enhancement. The authors propose an alternative function of neurogenesis in the control of motor behavior.

Animals↗

Patterns of brain activity associated with variation in voluntary wheel-running behavior.

Rodents spontaneously run on wheels, but what underlies variation within and between species is unknown. This study used Fos immunoreactivity to compare brain activity in mice selectively bred for high wheel running (S) versus control (C) mice. Mice ran for 6 days, but on Day 7, half the mice were prevented from running. A strong positive correlation was found between running distance and Fos in the dentate gyrus of C runners that was lost in S runners. In mice prevented from running, Fos was higher in S than in C in the lateral hypothalamus, medial frontal cortex, and striatum. Results implicate specific brain regions in motivation to run and others in control of the intensity of the locomotor behavior itself.

Analysis of Variance↗

Do precocial mammals develop at a faster rate? A comparison of rates of skull development in Sigmodon fulviventer and Mus musculus domesticus.

Variation in neonatal maturity among mammals is often explained by variation in gestation length, but species may also differ in developmental rate, a quantity that is difficult to measure because the conventional formalism makes two important and potentially unrealistic assumptions: (1) ontogeny of form can be described by a single line, and (2) species have the same ontogeny of form. We examine two species, one precocial (Sigmodon fulviventer), the other altricial (Mus musculus domesticus), and find that neither assumption is met. Therefore, we introduce an alternative metric, the rate of shape differentiation away from the average neonate. We find that S. fulviventer has a lower developmental rate than M. m. domesticus; consequently, while more mature at birth, S. fulviventer loses ground to M. m. domesticus over time. Surprisingly, despite differences in gestation length and developmental rate, these species reach developmental and life-history milestones at nearly identical degrees of skull shape maturity.

Adaptation, Physiological↗

Testing for phylogenetic signal in comparative data: behavioral traits are more labile.

The primary rationale for the use of phylogenetically based statistical methods is that phylogenetic signal, the tendency for related species to resemble each other, is ubiquitous. Whether this assertion is true for a given trait in a given lineage is an empirical question, but general tools for detecting and quantifying phylogenetic signal are inadequately developed. We present new methods for continuous-valued characters that can be implemented with either phylogenetically independent contrasts or generalized least-squares models. First, a simple randomization procedure allows one to test the null hypothesis of no pattern of similarity among relatives. The test demonstrates correct Type I error rate at a nominal alpha = 0.05 and good power (0.8) for simulated datasets with 20 or more species. Second, we derive a descriptive statistic, K, which allows valid comparisons of the amount of phylogenetic signal across traits and trees. Third, we provide two biologically motivated branch-length transformations, one based on the Ornstein-Uhlenbeck (OU) model of stabilizing selection, the other based on a new model in which character evolution can accelerate or decelerate (ACDC) in rate (e.g., as may occur during or after an adaptive radiation). Maximum likelihood estimation of the OU (d) and ACDC (g) parameters can serve as tests for phylogenetic signal because an estimate of d or g near zero implies that a phylogeny with little hierarchical structure (a star) offers a good fit to the data. Transformations that improve the fit of a tree to comparative data will increase power to detect phylogenetic signal and may also be preferable for further comparative analyses, such as of correlated character evolution. Application of the methods to data from the literature revealed that, for trees with 20 or more species, 92% of traits exhibited significant phylogenetic signal (randomization test), including behavioral and ecological ones that are thought to be relatively evolutionarily malleable (e.g., highly adaptive) and/or subject to relatively strong environmental (nongenetic) effects or high levels of measurement error. Irrespective of sample size, most traits (but not body size, on average) showed less signal than expected given the topology, branch lengths, and a Brownian motion model of evolution (i.e., K was less than one), which may be attributed to adaptation and/or measurement error in the broad sense (including errors in estimates of phenotypes, branch lengths, and topology). Analysis of variance of log K for all 121 traits (from 35 trees) indicated that behavioral traits exhibit lower signal than body size, morphological, life-history, or physiological traits. In addition, physiological traits (corrected for body size) showed less signal than did body size itself. For trees with 20 or more species, the estimated OU (25% of traits) and/or ACDC (40%) transformation parameter differed significantly from both zero and unity, indicating that a hierarchical tree with less (or occasionally more) structure than the original better fit the data and so could be preferred for comparative analyses.

Animals↗

Ontogenies in mice selected for high voluntary wheel-running activity. I. Mean ontogenies.

The evolutionary importance of postnatal ontogenies has long been recognized, but most studies of ontogenetic trajectories have focused exclusively on morphological traits. For animals, this represents a major omission because behavioral traits and their ontogenies often have relatively direct relationships to fitness. Here four replicate lines of house mice artificially selected for high early-age wheel running and their four replicate control lines were used to evaluate the effects of early-age directional selection, genetic drift, and activity environment (presence or absence of a running wheel) on variation in the ontogenies of three traits known to be genetically correlated: voluntary wheel running, body mass, and food consumption. Early-age selection significantly changed both the shape and position of the wheel-running and food-consumption ontogenies while influencing the position, but not the shape, of the body mass ontogeny. Genetic drift (as indicated by variation among replicate lines) produced significant changes in both the position and shape of all three ontogenies; however, its effect differed between the selection and control groups. For wheel running and food consumption, genetic drift only influenced the control ontogenies, whereas for body mass, genetic drift had a significant effect in both selection groups. Both body-mass and food-consumption ontogenies were significantly altered by activity environment, with the environment causing significant changes in the shape and position of both ontogenies. Overall the results demonstrate strong effects of early-age selection, genetic drift, and environmental variation on the evolution and expression of behavioral and morphological ontogenies, with selection changing only the position of the morphological ontogeny but both the position and shape of the behavioral ontogenies.

Analysis of Variance↗

Artificial selection for high activity favors mighty mini-muscles in house mice.

After 14 generations of selection for voluntary wheel running, mice from the four replicate selected lines ran, on average, twice as many revolutions per day as those from the four unselected control lines. To examine whether the selected lines followed distinct strategies in the correlated responses of the size and metabolic capacities of the hindlimb muscles, we examined mice from selected lines, housed for 8 wk in cages with access to running wheels that were either free to rotate ("wheel access" group) or locked ("sedentary"). Thirteen of twenty individuals in one selected line (line 6) and two of twenty in another (line 3) showed a marked reduction ( approximately 50%) in total hindlimb muscle mass, consistent with the previously described expression of a small-muscle phenotype. Individuals with these "mini-muscles" were not significantly smaller in total body mass compared with line-mates with normal-sized muscles. Access to free wheels did not affect the relative mass of the mini-muscles, but did result in typical mammalian training effects for mitochondrial enzyme activities. Individuals with mini-muscles showed a higher mass-specific muscle aerobic capacity as revealed by the maximal in vitro rates of citrate synthase and cytochrome c oxidase. Moreover, these mice showed the highest activities of hexokinase and carnitine palmitoyl transferase. Females with mini-muscles showed the highest levels of phosphofructokinase, and males with mini-muscles the highest levels of pyruvate dehydrogenase. As shown by total muscle enzyme contents, the increase in mass-specific aerobic capacity almost completely compensated for the reduction caused by the "loss" of muscle mass. Moreover, the mini-muscle mice exhibited the lowest contents of lactate dehydrogenase and glycogen phosphorylase. Interestingly, metabolic capacities of mini-muscled mice resemble those of muscles after endurance training. Overall, our results demonstrate that during selection for voluntary wheel running, distinct adaptive paths that differentially exploit the genetic variation in morphological and physiological traits have been followed.

Animals↗

Different effects of intensity and duration of locomotor activity on circadian period.

An outstanding unresolved issue in chronobiology is how the level of locomotor activity influences length of the free-running, endogenous circadian period (tau). To address this issue, the authors studied a novel model, 4 replicate lines of laboratory house mice (Mus domesticus) that had been selectively bred for high wheel-running activity (S) and their 4 unselected control (C) lines. Previous work indicates that S mice run approximately twice as many revolutions/day and exhibit an altered dopaminergic function as compared with C mice. The authors report that S mice have a tau shorter by about 0.5 h as compared with C mice. The difference in tau was significant both under constant light (control lines: tau = 25.5 h; selected: tau = 24.9 h) and under constant dark (control lines: 23.7 h; selected: 23.4 h). Moreover, the difference remained statistically significant even when the effects of running speed and time spent running were controlled in ANCOVA. Thus, something more fundamental than just intensity or duration of wheel-running activity per se must underlie the difference in tau between the S and C lines. However, despite significant difference in total wheel-running activity between females and males, tau did not differ between the sexes. Similarly, among individuals within lines, tau was not correlated with wheel-running activity measured as total revolutions per day. Instead, tau tended to decrease with average running speed but increase with time spent running. Finally, within individuals, an increase in time spent running resulted in decreased tau in the next few days, but changes in running speed had no statistically significant effect. The distinctions between effects of duration versus intensity of an activity, as well as between the among- versus within-individual correlations, are critical to understanding the relation between locomotor activity and pace of the circadian clock.

Animals↗

Reduction in muscle fibre number during the adaptive radiation of notothenioid fishes: a phylogenetic perspective.

The fish fauna of the continental shelf of the Southern Ocean is dominated by a single sub-order of Perciformes, the Notothenioidei, which have unusually large diameter skeletal muscle fibres. We tested the hypothesis that in fast myotomal muscle a high maximum fibre diameter (FD(max)) was related to a reduction in the number of muscle fibres present at the end of the recruitment phase of growth. We also hypothesized that the maximum fibre number (FN(max)) would be negatively related to body size, and that both body size and size-corrected FN(max) would show phylogenetic signal (tendency for related species to resemble each other). Finally, we estimated ancestral values for body size and FN(max). A molecular phylogeny was constructed using 12S mitochondrial rRNA sequences. A total of 16 species were studied from the Beagle Channel, Tierra del Fuego (5-11 degrees C), Shag Rocks, South Georgia (0.5-4 degrees C), and Adelaide Island, Antarctic Peninsula (-1.5 to 0.5 degrees C). The absence of muscle fibres of less than 10 micro m diameter was used as the criterion for the cessation of fibre recruitment. FD(max) increased linearly with standard length (SL), reaching 500-650 micro m in most species. Maximum body size was a highly significant predictor of species variation in FN(max), and both body size and size-corrected FN(max) showed highly significant phylogenetic signal (P<0.001). Estimates of trait values at nodes of the maximum likelihood phylogenetic tree were consistent with a progressive reduction in fibre number during part of the notothenioid radiation, perhaps serving to reduce basal energy requirements to compensate for the additional energetic costs of antifreeze production. For example, FN(max) in Chaenocephalus aceratus (12 700+/-300, mean +/- S.E.M., N=18) was only 7.7% of the value found in Eleginops maclovinus (164 000+/-4100, N=17), which reaches a similar maximum length (85 cm). Postembryonic muscle fibre recruitment in teleost fish normally involves stratified followed by mosaic hyperplasia. No evidence for this final phase of growth was found in two of the most derived families (Channichthyidae and Harpagiferidae). The divergence of the notothenioids in Antarctica after the formation of the Antarctic Polar Front and more recent dispersal north would explain the high maximum diameter and low fibre number in the derived sub-Antarctic notothenioids. These characteristics of notothenioids may well restrict their upper thermal tolerance, particularly for Champsocephalus esox and similar Channichthyids that lack respiratory pigments.

Adaptation, Biological↗

Dominance, plasma testosterone levels, and testis size in house mice artificially selected for high activity levels.

Male house mice (Mus domesticus) from four replicate lines selectively bred for high voluntary wheel-running behavior were compared with four random-bred control lines with respect to dominance, testis size, and plasma testosterone level. Behavior was measured with a tube apparatus in which focal mice encountered a standard opponent from an inbred strain, and positions of mice were scored over a 10-min period; the test was replicated the following day. Blood samples were taken from undisturbed mice 1 week prior to testing (baseline condition) and immediately after the first tube test; plasma testosterone was measured by enzyme immunoassay with chromatography. As compared with control lines, mice from selected lines tended to be smaller in body mass, to have larger testes, and were significantly less likely to advance towards their opponent during the second tube-test encounter. However, no significant differences in either baseline or post-encounter testosterone levels were detected. Significant differences in body mass, relative testis size, position during the first tube-test encounter, and baseline testosterone were found among the replicate lines within linetype, which indicates founder effects, random genetic drift, unique mutations, and/or multiple responses to selection. At the level of individual variation (residuals from nested analysis of covariance models), an inverse relationship between baseline testosterone and advancing in the tube test was observed, and the relationship was stronger during the second test day. This unexpected result may reflect an alternate coping strategy.

Animals↗

Antioxidant gene expression in active and sedentary house mice (Mus domesticus) selected for high voluntary wheel-running behavior.

We present liver mRNA levels of the two antioxidant enzymes catalase (CAT) and Mn-superoxide dismutase (SOD2) in four treatment groups of house mice assayed by RNase protection at 20 months of age. These groups were mice from four replicate selection and four replicate control lines from the sixteenth generation of selective breeding for high voluntary wheel running, housed with or without running wheels from age 3 weeks through 20 months. Exercising control females had induced CAT expression; SOD2 exhibited a similar pattern in females from two of the four control lines. Exercising male mice had induced CAT expression, but not SOD2 expression, irrespective of genetic background. We discuss these results with respect to both evolutionary (genetic) and training (exercise-induced) adaptations and explore predictions of these results in relation to the oxidative-damage theory of senescence.

Aging↗

Evolution of a small-muscle polymorphism in lines of house mice selected for high activity levels.

To study the correlated evolution of locomotor behavior and exercise physiology, we conducted an artificial selection experiment. From the outbred Hsd:ICR strain of Mus domesticus, we began eight separate lines, each consisting of 10 breeding pairs. In four of the lines, we used within-family selection to increase voluntary wheel running. The remaining four lines were random-bred (within lines) to serve as controls. Various traits have been monitored to test for correlated responses. Here, we report on organ masses, with emphasis on the triceps surae muscle complex, an important extensor of the ankle. Mice from the selected lines exhibit reduced total body mass, increased relative (mass-corrected) kidney mass, and reduced relative triceps surae mass. In addition, a discrete muscle-mass polymorphism was observed: some individuals had triceps surae that were almost 50% lighter than normal for their body mass. This small-muscle phenotype was observed in only three of the eight lines: in one control line, it has fluctuated in frequency between zero and 10%, whereas in two of the selected lines it has increased in frequency to approximately 50% by generation 22. Data from a set of parents and offspring (generations 23 and 24) are consistent with inheritance as a single autosomal recessive allele. Evidence for the adaptive significance of the small-muscle allele was obtained by fitting multiple-generation data to hierarchical models that include effects of genetic drift and/or selection. The small-muscle allele is estimated to have been present at low frequency (approximately 7%) in the base population, and analysis indicates that strong selection favors the allele in the selected but not control lines. We hypothesize that the small muscles possess functional characteristics and/or that the underlying allele causes pleiotropic effects (e.g., reduced total body mass; increased relative heart, liver, and kidney mass) that facilitate high levels of wheel running. Nevertheless, at generation 22, wheel running of affected individuals did not differ significantly from those with normal-sized muscles, and the magnitude of response to selection has been similar in all four selected lines, indicating that multiple genetic "solutions" are possible in response to selection for high activity levels.

Animals↗

A comparative analysis of brain size in relation to foraging ecology and phylogeny in the Chiroptera.

Variations in total brain mass and in the mass of three brain regions (main olfactory bulb, hippocampus, auditory nuclei) were examined using a data set for 63 species of bats (Chiroptera). Using both conventional and phylogenetically based analysis of covariance (log body mass as covariate), we tested several hypotheses that relate total brain mass or the size of the components to variation in foraging ecology, categorized as phytophagous, gleaner, and aerial insectivore. In some analyses, the category phytophagous was split into phytophagous pteropodid and phytophagous phyllostomid to examine differences between two distinct clades of bats. Because the Megachiroptera orient primarily by vision and olfaction, whereas all other bats rely on laryngeal echolocation to locate their prey, we hypothesized that the former would differ in size of the main olfactory bulb, as compared with all other bats. This hypothesis was supported by our analyses. Our more general prediction was that insectivorous bats, which rely heavily on echolocation for the pursuit and capture of their prey, would have larger auditory nuclei than do phytophagous species. This, too, was supported. We also compared phytophagous (fruit or nectar consuming) bats in two families, the Pteropodidae and the Phyllostomidae. We hypothesized that the phyllostomids, which use echolocation while foraging, would have larger auditory nuclei. Although statistical power is low in phylogenetically informed comparisons of the two clades, we did find weak evidence in support of this hypothesis. We conclude that bat brains show evidence of adaptation to foraging ecology.

Adaptation, Physiological↗