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T Garland

Publications and source records attributed to T Garland.

At least 37 records · Page 2Linked to original sources

Diet, phylogeny, and basal metabolic rate in phyllostomid bats.

Aside from the pervasive effects of body mass, much controversy exists as to what factors account for interspecific variation in basal metabolic rates (BMR) of mammals; however, both diet and phylogeny have been strongly implicated. We examined variation in BMR within the New World bat family Phyllostomidae, which shows the largest diversity of food habits among mammalian families, including frugivorous, nectarivorous, insectivorous, carnivorous and blood-eating species. For 27 species, diet was taken from the literature and BMR was either measured on animals captured in Brazil or extracted from the literature. Conventional (nonphylogenetic) analysis of covariance (ANCOVA), with body mass as the covariate, was first used to test the effects of diet on BMR. In this analysis, which assumes that all species evolved simultaneously from a single ancestor (i.e., a "star" phylogeny), diet exerted a strong effect on mass-independent BMR: nectarivorous bats showed higher mass-independent BMR than other bats feeding on fruits, insects or blood. In phylogenetic ANCOVAs via Monte Carlo computer simulation, which assume that species are part of a branching hierarchical phylogeny, no statistically significant effect of diet on BMR was observed. Hence, results of the nonphylogenetic analysis were misleading because the critical values for testing the effect of diet were underestimated. However, in this sample of bats, diet is perfectly confounded with phylogeny, because the four dietary categories represent four separate subclades, which greatly reduces statistical power to detect a diet (= subclade) effect. But even if diet did appear to exert an influence on BMR in this sample of bats, it would not be logically possible to separate this effect from the possibility that the dietary categories differ for some other reason (i.e., another synapomorphy of one or more of the subclades). Examples such as this highlight the importance of considering phylogenetic relationships when designing new comparative studies, as well as when analyzing existing data sets. We also discuss some possible reasons why BMR may not coadapt with diet.

Journal Article↗

The quantitative genetics of maximal and basal rates of oxygen consumption in mice.

A positive genetic correlation between basal metabolic rate (BMR) and maximal (VO(2)max) rate of oxygen consumption is a key assumption of the aerobic capacity model for the evolution of endothermy. We estimated the genetic (V(A), additive, and V(D), dominance), prenatal (V(N)), and postnatal common environmental (V(C)) contributions to individual differences in metabolic rates and body mass for a genetically heterogeneous laboratory strain of house mice (Mus domesticus). Our breeding design did not allow the simultaneous estimation of V(D) and V(N). Regardless of whether V(D) or V(N) was assumed, estimates of V(A) were negative under the full models. Hence, we fitted reduced models (e.g., V(A) + V(N) + V(E) or V(A) + V(E)) and obtained new variance estimates. For reduced models, narrow-sense heritability (h(2)(N)) for BMR was <0.1, but estimates of h(2)(N) for VO(2)max were higher. When estimated with the V(A) + V(E) model, the additive genetic covariance between VO(2)max and BMR was positive and statistically different from zero. This result offers tentative support for the aerobic capacity model for the evolution of vertebrate energetics. However, constraints imposed on the genetic model may cause our estimates of additive variance and covariance to be biased, so our results should be interpreted with caution and tested via selection experiments.

Animals↗

Genetic selection of mice for high voluntary wheel running: effect on skeletal muscle glucose uptake.

Effects of genetic selection for high wheel-running activity (17th generation) and access to running wheels on skeletal muscle glucose uptake were studied in mice with the following treatments for 8 wk: 1) access to unlocked wheels; 2) same as 1, but wheels locked 48 h before glucose uptake measurement; or 3) wheels always locked. Selected mice ran more than random-bred (nonselected) mice (8-wk mean +/- SE = 8,243 +/- 711 vs. 3,719 +/- 233 revolutions/day). Body weight was 5-13% lower for selected vs. nonselected groups. Fat pad/body weight was ~40% lower for selected vs. nonselected and unlocked vs. locked groups. Insulin-stimulated glucose uptake and fat pad/body weight were inversely correlated for isolated soleus (r = -0.333; P < 0.005) but not extensor digitorum longus (EDL) or epitrochlearis muscles. Insulin-stimulated glucose uptake was higher in EDL (P < 0.02) for selected vs. nonselected mice. Glucose uptake did not differ by wheel group, and amount of running did not correlate with glucose uptake for any muscle. Wheel running by mice did not enhance subsequent glucose uptake by isolated muscles.

Animals↗

Selection for high voluntary wheel-running increases speed and intermittency in house mice (Mus domesticus).

In nature, many animals use intermittent rather than continuous locomotion. In laboratory studies, intermittent exercise regimens have been shown to increase endurance compared with continuous exercise. We hypothesized that increased intermittency has evolved in lines of house mice (Mus domesticus) that have been selectively bred for high voluntary wheel-running (wheel diameter 1.12 m) activity. After 23 generations, female mice from four replicate selection lines ran 2.7 times more revolutions per day than individuals from four random-bred control lines. To measure instantaneous running speeds and to quantify intermittency, we videotaped mice (N=41) during a 5-min period of peak activity on night 6 of a 6-day exposure to wheels. Compared with controls (20 revs min(-1) while actually running), selection-line females (41 revs min(-1)) ran significantly faster. These instantaneous speeds closely matched the computer-recorded speeds over the same 5-min period. Selection-line females also ran more intermittently, with shorter (10.0 s bout(-1)) and more frequent (7.8 bouts min(-1)) bouts than controls (16.8 s bout(-1), 3.4 bouts min(-1)). Inter-bout pauses were also significantly shorter in selection-line (2.7 s) than in control-line (7.4 s) females. We hypothesize that intermittency of locomotion is a key feature allowing the increased wheel-running performance at high running speeds in selection-line mice.

Aging↗

Maximum cold-induced food consumption in mice selected for high locomotor activity: implications for the evolution of endotherm energy budgets.

We studied house mice (Mus domesticus) that had been artificially selected for high activity to test the hypothesis that a high capacity for energy assimilation in cold-exposed endotherms could evolve as a correlated response to selection for increased locomotor activity. After 10 generations of selection for increased voluntary wheel-running, mice from four selected lines ran 75 % more wheel revolutions per day than did mice from four random-bred, control lines. The maximum cold-induced rates of food consumption (C(max); mean 10.6 g day(-1)) and energy assimilation (A(max); mean 141 kJ day(-1)) were not significantly higher in the selected than in the control mice. However, in cold-exposure trials, mice from the selected lines maintained body mass better than did mice from the control lines. C(max) and A(max) were positively correlated with the amount of wheel-running activity measured before cold-exposure and also with the rates of food consumption measured when the mice had access to running wheels. In females at least, the correlation was significant not only among individuals but also among adjusted means of the replicate lines, which suggests the presence of a positive genetic correlation between the traits. Thus, despite the lack of a significant difference between the selected and control lines in maximum rate of food consumption, the remaining results conform to the hypothesis that a selection for increased locomotor activity could be a factor behind the evolution of the ability to sustain activity and maintain energy balance during prolonged cold-exposure, as occurred during the evolution of mammalian and avian endothermy.

Animals↗

Phylogenetic comparison and artificial selection. Two approaches in evolutionary physiology.

Interspecific comparison has a long and productive history in physiology. Conceptual and statistical advances over the last 15 years have demonstrated several ways in which comparisons can be enhanced by consideration of phylogenetic information, i.e., empirical estimates of the ways in which organisms are related (evolutionary trees). Choice of species to be compared should be informed by phylogenetic information. For example, a comparison of three species that inhabit high altitude with three that live at low altitude would be suspect if each of the two groups were composed of closely-related species (e.g., within single genera). To avoid such "phylogenetic pseudoreplication," one might instead study species from three different genera, each containing one high-altitude and one low-altitude inhabitant. Unfortunately, many studies have not been so carefully designed, sometimes because organisms were not accessible or because the studies incorporated data from the literature. Fortunately, several new statistical methods correct for problems caused by phylogenetic relatedness and descent with modification, the most common being phylogenetically independent contrasts. Another tool that can be used in comparative physiology is selective breeding, which has been practiced for millennia and applied in scientific contexts for over a century. In the last 20 years, ecological and evolutionary physiologists have begun using selection experiments to study processes of genetic adaptation in physiological and behavioral traits. For example, house mice have been maintained in the cold for multiple generations to see what adaptations may occur naturally in response to reduced ambient temperature ("laboratory natural selection"). Our own laboratory has used selective breeding to create four replicate lines of mice that exhibit high levels of voluntary wheel-running behavior, as well as various morphological and physiological characteristics that cause or allow the elevated locomotor activity. Similar experiments could be used to study adaptation to hypoxia.

Animals↗

Body temperatures of house mice artificially selected for high voluntary wheel-running behavior: repeatability and effect of genetic selection.

We studied rectal body temperatures of house mice (Mus domesticus) that had been artificially selected for high voluntary wheel running.1. At generation 17, mice from the four replicate selected lines ran, on average, 2.5-times as many revolutions/day as did mice from the four random-bred control lines.2. During the day, repeatability of individual differences in body temperature measured 4 days apart was low; at night, repeatability was statistically significant across three time scales (1 day, 1 week, 2 weeks).3. During the day, body temperatures of selected and control animals did not differ; at night, mice from selected lines had higher body temperatures. However, when amount of wheel running immediately prior to measurement was included as a covariate, the difference was no longer statistically significant.Higher body temperatures, associated with increased activity, might enhance locomotor abilities through Q10 effects, increase metabolic rate and food requirements, affect sleep patterns, and alter expression of heat-shock proteins.

Journal Article↗

Toxicologic disease of the digestive tract.

There is a diverse and long list of toxicants that can affect the digestive system of food-producing animals. The plants and other natural toxicants discussed in this article are those primarily affecting the GI system. A number of other plants may also affect the digestive tract, but the effects from these are considered secondary and less pronounced. Often, plant poisonings affecting the digestive tract present with similar clinical signs, and a good thorough history is necessary to help differentiate between them. Moreover, a careful walk through the pasture with a keen eye to note plants that have been browsed or grazed may greatly assist the history. In cases where toxins are suspected as the cause of a GI disorder, consultation with a veterinary toxicologist at a diagnostic laboratory may be indicated. These professionals are knowledgeable about a wide variety of natural and other toxicants that may be present in your area. They can help with developing a differential diagnosis and the selection of appropriate samples to confirm the diagnosis.

Animals↗

Nesting behavior of house mice (Mus domesticus) selected for increased wheel-running activity.

Nest building was measured in "active" (housed with access to running wheels) and "sedentary" (without wheel access) mice (Mus domesticus) from four replicate lines selected for 10 generations for high voluntary wheel-running behavior, and from four randombred control lines. Based on previous studies of mice bidirectionally selected for thermoregulatory nest building, it was hypothesized that nest building would show a negative correlated response to selection on wheel-running. Such a response could constrain the evolution of high voluntary activity because nesting has also been shown to be positively genetically correlated with successful production of weaned pups. With wheel access, selected mice of both sexes built significantly smaller nests than did control mice. Without wheel access, selected females also built significantly smaller nests than did control females, but only when body mass was excluded from the statistical model, suggesting that body mass mediated this correlated response to selection. Total distance run and mean running speed on wheels was significantly higher in selected mice than in controls, but no differences in amount of time spent running were measured, indicating a complex cause of the response of nesting to selection for voluntary wheel running.

Animals↗

Effects of voluntary activity and genetic selection on muscle metabolic capacities in house mice Mus domesticus.

Selective breeding is an important tool in behavioral genetics and evolutionary physiology, but it has rarely been applied to the study of exercise physiology. We are using artificial selection for increased wheel-running behavior to study the correlated evolution of locomotor activity and physiological determinants of exercise capacity in house mice. We studied enzyme activities and their response to voluntary wheel running in mixed hindlimb muscles of mice from generation 14, at which time individuals from selected lines ran more than twice as many revolutions per day as those from control (unselected) lines. Beginning at weaning and for 8 wk, we housed mice from each of four replicate selected lines and four replicate control lines with access to wheels that were free to rotate (wheel-access group) or locked (sedentary group). Among sedentary animals, mice from selected lines did not exhibit a general increase in aerobic capacities: no mitochondrial [except pyruvate dehydrogenase (PDH)] or glycolytic enzyme activity was significantly (P < 0.05) higher than in control mice. Sedentary mice from the selected lines exhibited a trend for higher muscle aerobic capacities, as indicated by higher levels of mitochondrial (cytochrome-c oxidase, carnitine palmitoyltransferase, citrate synthase, and PDH) and glycolytic (hexokinase and phosphofructokinase) enzymes, with concomitant lower anaerobic capacities, as indicated by lactate dehydrogenase (especially in male mice). Consistent with previous studies of endurance training in rats via voluntary wheel running or forced treadmill exercise, cytochrome-c oxidase, citrate synthase, and carnitine palmitoyltransferase activity increased in the wheel-access groups for both genders; hexokinase also increased in both genders. Some enzymes showed gender-specific responses: PDH and lactate dehydrogenase increased in wheel-access male but not female mice, and glycogen phosphorylase decreased in female but not in male mice. Two-way analysis of covariance revealed significant interactions between line type and activity group; for several enzymes, activities showed greater changes in mice from selected lines, presumably because such mice ran more revolutions per day and at greater velocities. Thus genetic selection for increased voluntary wheel running did not reduce the capability of muscle aerobic capacity to respond to training.

Animals↗

Laboratory endurance capacity predicts variation in field locomotor behaviour among lizard species.

I measured locomotor endurance capacities of lizards on a motorized treadmill in the laboratory and compared average values for different species with quantitative measures of their movement in the field (percentage of time moving, N=15 species; moves/min, N=13; daily movement distance, N=11). I hypothesized that endurance would be positively related to all three movement indices. Relationships between log endurance and log movement were computed as conventional Pearson product-moment correlations and as the equivalent with phylogenetically independent contrasts. Endurance was significantly positively related to both the percentage of time moving and the daily movement distance. This is the first study to demonstrate such relationships with phylogenetically based statistical methods. These results suggest that endurance capacities of lizards are coadapted with their typical locomotor behaviour. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗

Behaviour of house mice artificially selected for high levels of voluntary wheel running.

We have developed a novel model to study the correlated evolution of behavioural and morphophysiological traits in response to selection for increased locomotor activity. We used selective breeding to increase levels of voluntary wheel running in four replicate lines of laboratory house mice, Mus domesticus, with four random-bred lines maintained as controls. The experiment presented here tested for correlated behavioural responses in the wheel-cage complex, with wheels either free to rotate or locked (environmental factor). After 13 generations, mice from selected lines ran 2.2 times as many revolutions/day as controls on days 5 and 6 of initial exposure to wheels (10 826 versus 4890 revolutions/day, corresponding to 12.1 and 5.5 km/day, respectively). This increase was caused primarily by mice from selected lines running faster, not more minutes per day. Focal-animal observations confirmed that the increase in revolutions/day involved more actual running (or climbing in locked wheels), not an increase in coasting (or hanging). Not surprisingly, access to free versus locked wheels had several effects on behaviour, including total time spent in wheels, sniffing and biting. However, few behaviours showed statistically significant differences between the selected and control lines. Selection did not increase the total time spent in wheels (either free or locked), the frequency of nonlocomotor activities performed in the wheels, nor the amount of locomotor activity in cages attached to the wheels; as well, selection did not decrease the amount of time spent sleeping. Thus, wheel running is, at the genetic level, a largely independent axis of behaviour. Moreover, the genetic architecture of overall wheel running and its components seem conducive to increasing total distance moved without unduly increasing energy or time-related costs. The selection experiment also offers a new approach to study the proximate mechanisms of wheel-running behaviour itself. For example, frequencies of sniffing and wire biting were reduced in selected females but not males. This result suggests that motivation or function of wheel running may differ between the sexes. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗

Genetic variation and correlations between genotype and locomotor physiology in outbred laboratory house mice (Mus domesticus).

Laboratory strains of house mice (Mus domesticus) are increasingly used as model organisms in evolutionary physiology, so information on levels of genetic variation is important. For example, are levels of genetic variation comparable to those found in populations of wild house mice? We studied allozymes to estimate genetic variation in outbred Hsd:ICR mice, which have been used in several studies with evolutionary emphasis. The physiological significance of allozyme variation remains obscure. Several workers have reported relationships between multi-locus heterozygosity and metabolic traits, but endotherms have not been studied. Therefore, we also measured mice for basal metabolic rate (BMR), maximal oxygen consumption during forced treadmill exercise (VO2max), and 12 other traits related to locomotor physiology, before genotyping them for 10 allozyme loci. Four of these loci were polymorphic, all were in Hardy-Weinberg equilibrium, and inbreeding coefficients were not significantly different from zero. Average heterozygosities were 11%, similar to values reported for wild populations of house mice. Fourteen percent of the associations between single-locus genotype and physiological traits were statistically significant. Multi-locus heterozygosity was not significantly related to VO2max, but was positively correlated with BMR, a result opposite to the negative correlation between standard metabolic rate and heterozygosity reported in many ectotherms. Therefore, the proposed mechanisms for the effect of multi-locus heterozygosity on metabolic rate in ectotherms may not apply to endotherms.

Animals↗

Polytomies and phylogenetically independent contrasts: examination of the bounded degrees of freedom approach.

We examined the effect of soft polytomies on the performance (Type I error rate and bias) of Felsenstein's (1985; Am. Nat. 125:1-15) method of phylogenetically independent contrasts for estimating a bivariate correlation. We specifically tested the adequacy of bounding degrees of freedom, as suggested by Purvis and Garland (1993; Syst. Biol. 42:569-575). We simulated bivariate character evolution under Brownian motion (assumed by independent contrasts) and eight other models on five phylogenetic trees. For non-Brownian motion simulations, the adequacy of branch-length standardization was checked with a simple diagnostic (Garland et al., 1992; Syst. Biol. 41:18-32), and transformations were applied as indicated. Surprisingly, soft polytomies tended to have negligible effects on Type I error rates when models other than Brownian motion were used. Overall, and irrespective of evolutionary model, degrees of freedom were appropriately bounded for hypothesis testing, and unbiased estimates of the correlation coefficient were obtained. Our results, along with those of previous simulation studies, suggest that independent contrasts can reliably be applied to real data, even with phylogenetic uncertainty.

Models, Genetic↗

Effects of genetic selection and voluntary activity on the medial gastrocnemius muscle in house mice.

In a previous study, we found that in house mice both genetic selection (10 generations of artificial selection for high voluntary activity on running wheels) and access to running wheels (7-8 weeks) elicited a modest increase in maximal oxygen consumption. Based on these results, we hypothesized that genetic selection would affect the changes in endurance and oxidative capacity of the medial gastrocnemius (MG) muscle induced by wheel access (training response). Wheel access increased the isotonic endurance of the MG in both genetically selected and random-bred (control) mice. However, this exercise-induced improvement in isotonic endurance of the MG was similar between genetically selected and control mice. Wheel access also increased the succinate dehydrogenase activity of MG muscle fibers in both selected and control lines. However, this exercise-induced increase in succinate dehydrogenase activity was comparable between genetically selected and control animals. Taken together, these results indicate that the modest increase in maximal oxygen consumption associated with genetic selection is not reflected by the training-induced changes in oxidative capacity and endurance of MG muscle fibers.

Animals↗

Artificial selection for increased wheel-running activity in house mice results in decreased body mass at maturity.

To test the hypothesis that body size and activity levels are negatively genetically correlated, we conducted an artificial selection experiment for increased voluntary wheel-running activity in house mice (Mus domesticus). Here, we compare body masses of mice from control and selected lines after 14 generations of selection. In both groups, beginning at weaning and then for 8 weeks, we housed half of the individuals with access to running wheels that were free to rotate and the other half with wheels that were locked to prevent rotation. Mice from selected lines were more active than controls at weaning (21 days) and across the experiment (total revolutions during last week: females 2.5-fold higher, males 2.1-fold higher). At weaning, mice from selected and control lines did not differ significantly in body mass. At 79 days of age, mice from selected lines weighed 13.6 % less than mice from control lines, whereas mice with access to free wheels weighed 4.5 % less than 'sedentary' individuals; both effects were statistically significant and additive. Within the free-wheel-access group, individual variation in body mass of males was negatively correlated with amount of wheel-running during the last week (P<0.01); for females, the relationship was also negative but not statistically significant (P>0.40). The narrow-sense genetic correlation between wheel-running and body mass after 8 weeks of wheel access was estimated to be -0. 50. A negative genetic correlation could account for the negative relationship between voluntary wheel-running and body mass that has been reported across 13 species of muroid rodents.

Animals↗

Glucocorticoid response to forced exercise in laboratory house mice (Mus domesticus).

We examined the time course and sex differences of the glucocorticoid response to forced, moderate-intensity treadmill exercise in outbred laboratory house mice. Mice (n = 64 total) were divided into eight groups, each of four males and four females, which were run on a motorized treadmill at 1.0 km/h for either 0, 2, 5, 10, 15, 25, 40, or 60 min. Serum samples were taken immediately after exercise and corticosterone (CORT) concentration was determined by radioimmunoassay. Resting CORT levels ranged between 11.6 and 29.5 ng/mL for both sexes. CORT levels increased with length of exercise and then exhibited a plateau by 25 min in females and by 40 min in males. Females displayed a significantly more rapid increase in serum CORT levels and attained higher maximal CORT levels than males. Females also had significantly larger adrenal glands, both in absolute terms and relative to body mass.

Adrenal Glands↗

Artificial selection for increased wheel-running behavior in house mice.

Replicated within-family selection for increased voluntary wheel running in outbred house mice (Mus domesticus; Hsd:ICR strain) was applied with four high-selected and four control lines (10 families/line). Mice were housed individually with access to activity wheels for a period of 6 days, and selection was based on the mean number of revolutions run on days 5 and 6. Prior to selection, heritabilities of mean revolutions run per day (rev/day), average running velocity (rpm), and number of minutes during which any activity occurred (min/day) were estimated by midparent-offspring regression. Heritabilities were 0.18, 0.28, and 0.14, respectively; the estimate for min/day did not differ significantly from zero. Ten generations of selection for increased rev/day resulted in an average 75% increase in activity in the four selected lines, as compared with control lines. Realized heritability averaged 0.19 (range, 0.12-0.24 for the high-activity lines), or 0.28 when adjusted for within-family selection. Rev/day increased mainly through changes in rpm rather than min/day. These lines will be studied for correlated responses in exercise physiology capacities and will be made available to other researchers on request.

Analysis of Variance↗