Lower-level predictors and behavioral correlates of maximal aerobic capacity and sprint speed among individual lizards.

The standard paradigm of organismal biology views lower-level traits (e.g. aspects of physiology) as determining organismal performance ability (e.g. maximal sprint speed), which in turn constrains behavior (e.g. social interactions). However, few studies have simultaneously examined all three levels of organization. We used focal observations to record movement behaviors and push-up displays in the field for adult male Sceloporus occidentalis lizards during the breeding season. We then captured animals, measured aspects of their physiology, morphology and performance, and counted ectoparasites and endoparasites as potential predictors of sprint speed and maximal oxygen consumption (V̇O2,max). Field behaviors were statistically repeatable, but not strongly so. Sprint speed and V̇O2,max were repeatable using residuals from regressions on body mass (speed: r=0.70; V̇O2,max: r=0.88). Both calf [standardized partial regression (path) coefficient B=0.53] and thigh [B=-0.37] muscle mass (as residuals from regressions on body mass) were significant predictors of sprint speed; hemoglobin concentration (B=0.42) was a predictor of V̇O2,max. In turn, V̇O2,max predicted the maximum number of four-legged push-ups per bout (B=0.39). In path analysis, log likelihood ratio tests indicated no direct paths from lower-level traits to behavior, supporting the idea that morphology, in the broad sense, only affects behavior indirectly through measures of performance. Our results show that inter-individual variation in field behaviors can be related to performance ability, which in turn reflect differences in morphology and physiology, although not parasite load. Given the low repeatability of field behaviors, some of the relationships between behavior and performance may be stronger than suggested by our results.

Effects of early-life voluntary exercise and fructose on adult activity levels, body composition, aerobic capacity, and organ masses in mice bred for high voluntary wheel-running behavior.

Fructose (C6H12O6) is acutely obesogenic and is a risk factor for hypertension, cardiovascular disease, and nonalcoholic fatty liver disease. However, the possible long-lasting effects of early-life fructose consumption have not been studied. We tested for effects of early-life fructose and/or wheel access (voluntary exercise) in a line of selectively bred High Runner (HR) mice and a non-selected Control (C) line. Exposures began at weaning and continued for 3 weeks to sexual maturity, followed by a 23-week "washout" period (equivalent to ∼17 human years). Fructose increased total caloric intake, body mass, and body fat during juvenile exposure, but had no effect on juvenile wheel running and no important lasting effects on adult physical activity or body weight/composition. Interestingly, adult maximal aerobic capacity (VO2max) was reduced in mice that had early-life fructose and wheel access. Consistent with previous studies, early-life exercise promoted adult wheel running. In a 3-way interaction, C mice that had early-life fructose and no wheel access gained body mass in response to 2 weeks of adult wheel access, while all other groups lost mass. Overall, we found some long-lasting positive effects of early-life exercise, but minimal effects of early-life fructose, regardless of the mouse line.

Herpetofauna of protected areas in the Caatinga VIII: An updated checklist for the Serra das Confusões region with new data from Serra Vermelha, Piauí, Brazil / Herpetofauna das áreas protegidas da Caatinga VIII: Um inventário atualizado para a região da Serra das Confusões com novos dados da Serra Vermelha, Piauí, Brazil

Abstract It is repeatedly stressed the need to characterize the extant biodiversity in tropical ecosystems. However, inventory studies are still progressing slowly in dry ecosystems, leading to the underestimation of their true biodiversity and hindering conservation efforts. In this study, we present primary and secondary data, along with an updated list of amphibians and reptiles from two localities in the São Francisco-Gurguéia region in Piauí. Additionally, we compare the species composition between nine areas within the Caatinga, which were sampled using standardized methods over the past ten years, to examine broader spatial patterns of community composition. To survey reptiles and amphibians, we employed similar methods and sampling efforts in two areas within the Serra das Confusões National Park (SCNP) region. Our surveys recorded a total of 73 species of amphibians and reptiles, of which 24 are new distribution records for the SCNP region. Consequently, our findings increase the known herpetofauna in the region to 94 species. Despite their proximity, the two sites in the SCNP region exhibited only 42% similarity in species composition, and they differed significantly from other areas within the Caatinga. Furthermore, even the closer Caatinga areas presented differences in species composition, highlighting the necessity to evaluate biodiversity across the landscape and contribute to understanding biogeographic patterns.

Resumo É repetidamente enfatizada a necessidade de caracterizar a biodiversidade vivente em ecossistemas tropicais. No entanto, os estudos de inventário ainda estão progredindo lentamente em ecossistemas secos, levando à subestimação de sua verdadeira biodiversidade e dificultando os esforços de conservação. Neste estudo, apresentamos dados primários e secundários, juntamente com uma lista atualizada de anfíbios e répteis de duas localidades na região de São Francisco-Gurguéia, do Piauí. Além disso, comparamos a composição de espécies entre nove áreas dentro da Caatinga, que foram amostradas usando métodos padronizados nos últimos dez anos, para examinar padrões espaciais mais amplos de composição da comunidade. Para estudar répteis e anfíbios, utilizamos métodos e esforços de amostragem semelhantes em duas áreas na região do Parque Nacional da Serra das Confusões (PNSC). Nossos levantamentos registraram um total de 73 espécies de anfíbios e répteis, das quais 24 são novos registros de distribuição para a região do PNSC. Consequentemente, nossos resultados aumentam a herpetofauna conhecida na região para 94 espécies. Apesar da proximidade, os dois locais na região do PNSC exibiram apenas 42% de similaridade na composição de espécies e diferiram significativamente de outras áreas dentro da Caatinga. Mesmo áreas mais próximas da Caatinga apresentaram diferenças na composição de espécies, destacando a necessidade de avaliar a biodiversidade em toda a paisagem e contribuir para a compreensão de padrões biogeográficos.

Conditioned place preference for cocaine and methylphenidate in female mice from lines selectively bred for high voluntary wheel-running behavior.

Behavioral addictions can come in many forms, including overeating, gambling and overexercising. All addictions share a common mechanism involving activation of the natural reward circuit and reinforcement learning, but the extent to which motivation for natural and drug rewards share similar neurogenetic mechanisms remains unknown. A unique mouse genetic model in which four replicate lines of female mice were selectively bred (>76 generations) for high voluntary wheel running (High Runner or HR lines) alongside four non-selected control (C) lines were used to test the hypothesis that high motivation for exercise is associated with greater reward for cocaine (20 mg/kg) and methylphenidate (10 mg/kg) using the conditioned place preference (CPP) test. HR mice run ~three times as many revolutions/day as C mice, but the extent to which they have increased motivation for other rewards is unknown. Both HR and C mice displayed significant CPP for cocaine and methylphenidate, but with no statistical difference between linetypes for either drug. Taken together, results suggest that selective breeding for increased voluntary running has modified the reward circuit in the brain in a way that increases motivation for running without affecting cocaine or methylphenidate reward.

Phylogenetic analysis of maximal oxygen consumption during exercise (V̇O2max) and ecological correlates among lizard species.

The maximum amount of oxygen consumed during forced exercise (V̇O2max) sets the upper limit to the effort that can be sustained over relatively long periods and can limit activity levels in nature. Among ectotherms, V̇O2max is primarily affected by body size and body temperature, but it should also coadapt with behavior, ecology, and life history aspects. We compiled published data from 11 different families of lizards, including 58 species and 7 populations (total 65 data points) and tested whether V̇O2max was related to diet (herbivore, insectivore, insectivore/carnivore, carnivore, and omnivore), climate (tropical, temperate, and arid), nocturnality, viviparity, or family. We fitted models that included body mass and measurement temperature as covariates, and all possible combinations of other independent variables using ordinary least-squares (OLS) and phylogenetic regressions assuming an Ornstein-Uhlenbeck model of residual trait evolution (RegOU). The sum of Akaike weights for each independent variable revealed viviparity (∑w i =0.996) and the combined set of dummy variables coding for helodermatids, varanids, and skinks (∑w i =0.996) as the most important predictors. These three families had relatively high V̇O2max. They are composed mainly of active foragers that probably benefit from higher V̇O2max. Viviparity had a negative effect on V̇O2max. Ecological or behavioral factors associated with viviparity (e.g., activity levels), but not included here, may explain this effect. The average allometric slope of V̇O2max from the top eight models (which accounted for 99% of the cumulative evidence) was 0.803, which is similar to that reported previously for lizards and for mammals in general.

Phylogenetic analysis of maximal oxygen consumption during exercise (V̇O2max) and ecological correlates among lizard species.

The maximum amount of oxygen consumed during forced exercise (V̇O2max) sets the upper limit to the effort that can be sustained over relatively long periods and can limit activity levels in nature. Among ectotherms, V̇O2max is primarily affected by body size and body temperature, but it should also coadapt with behavior, ecology, and life history aspects. We compiled published data from 11 different families of lizards, including 58 species and 7 populations (total 65 data points) and tested whether V̇O2max was related to diet (herbivore, insectivore, insectivore/carnivore, carnivore, and omnivore), climate (tropical, temperate, and arid), nocturnality, viviparity, or family. We fitted models that included body mass and measurement temperature as covariates, and all possible combinations of other independent variables using ordinary least-squares (OLS) and phylogenetic regressions assuming an Ornstein-Uhlenbeck model of residual trait evolution (RegOU). The sum of Akaike weights for each independent variable revealed viviparity (∑wi=0.996) and the combined set of dummy variables coding for helodermatids, varanids, and skinks (∑wi=0.996) as the most important predictors. These three families had relatively high V̇O2max. They are composed mainly of active foragers that probably benefit from higher V̇O2max. Viviparity had a negative effect on V̇O2max. Ecological or behavioral factors associated with viviparity (e.g., activity levels), but not included here, may explain this effect. The average allometric slope of V̇O2max from the top eight models (which accounted for 99% of the cumulative evidence) was 0.803, which is similar to that reported previously for lizards and for mammals in general.

Locomotion, Energetics, Performance, and Behavior: A Mammalian Perspective on Lizards, and Vice Versa.

SYNOPSIS: Animals are constrained by their abilities and by interactions with environmental factors, such as low ambient temperatures. These constraints range from physical impossibilities to energetic inefficiencies, and may entail trade-offs. Some of the constraints related to locomotion and activity metabolism can be illustrated through allometric comparisons of mammals and lizards, as representative terrestrial vertebrate endotherms and ectotherms, respectively, because these lineages differ greatly in aerobic metabolic capacities, resting energetic costs, and thermoregulatory patterns. Allometric comparisons are both useful and unavoidable, but "outlier" species (unusual for their clade) can also inform evolutionary scenarios, as they help indicate extremes of possible adaptation within mammalian and saurian levels of organization. We compared mammals and lizards for standard metabolic rate (SMR), maximal oxygen consumption during forced exercise (VO2max), net (incremental) cost of transport (NCT), maximal aerobic speed (MAS), daily movement distance (DMD), daily energy expenditure (DEE) during the active season, and the ecological cost of transport (ECT = percentage of DEE attributable to locomotion). (Snakes were excluded because their limbless locomotion has no counterpart in terrestrial mammals.) We only considered lizard SMR, VO2max, NCT, MAS, and sprint speed data if measured at 35-40 °C. On average, MAS is ∼7.4-fold higher in mammals, whereas SMR and VO2max are ∼6-fold greater, but values for all three of these traits overlap (or almost overlap) between mammals and lizards, a fact that has not previously been appreciated. Previous studies show that sprint speeds are similar for smaller mammals and lizards, but at larger sizes lizards are not as fast as some mammals. Mammals move ∼6-fold further each day than lizards, and DMD is by far the most variable trait considered here, but their NCT is similar. Mammals exceed lizards by ∼11.4-fold for DEE. On average for both lineages, the ECT is surprisingly low, somewhat higher for lizards, and positively allometric. If a lizard and mammal of 100 g body mass were both to move their entire DMD at their MAS, they could do so in ∼21 and 17 min, respectively, thus de-emphasizing the possible importance of time constraints. We conclude that ecological-energetic constraints related to locomotion are relatively more likely to occur in large, carnivorous lizards. Overall, our comparisons support the idea that the (gradual) evolution of mammalian endothermy did not necessarily require major changes in locomotor energetics, performance, or associated behaviors. Instead, we speculate that the evolution of thermoregulatory responses to low temperatures (e.g., shivering) may have been a key and "difficult" step in this transition.

Maternal exposure to Western diet affects adult body composition and voluntary wheel running in a genotype-specific manner in mice.

Some human diseases, including obesity, Type II diabetes, and numerous cancers, are thought to be influenced by environments experienced in early life, including in utero. Maternal diet during the perinatal period may be especially important for adult offspring energy balance, potentially affecting both body composition and physical activity. This effect may be mediated by the genetic background of individuals, including, for example, potential "protective" mechanisms for individuals with inherently high levels of physical activity or high basal metabolic rates. To examine some of the genetic and environmental factors that influence adult activity levels, we used an ongoing selection experiment with 4 replicate lines of mice bred for high voluntary wheel running (HR) and 4 replicate, non-selected control lines (C). Dams (half HR and half C) were fed a "Western" diet (WD, high in fat and sucrose) or a standard diet (SD) from 2weeks prior to mating until their pups could feed on solid food (14days of age). We analyzed dam and litter characteristics from birth to weaning, and offspring mass and physical activity into adulthood. One male offspring from each litter received additional metabolic and behavioral tests. Maternal WD caused pups to eat solid food significantly earlier for C litters, but not for HR litters (interaction of maternal environment and genotype). With dam mass as a covariate, mean pup mass was increased by maternal WD but litter size was unaffected. HR dams had larger litters and tended to have smaller pups than C dams. Home-cage activity of juvenile focal males was increased by maternal WD. Juvenile lean mass, fat mass, and fat percent were also increased by maternal WD, but food consumption (with body mass as a covariate) was unaffected (measured only for focal males). Behavior in an elevated plus maze, often used to indicate anxiety, was unaffected by maternal WD. Maximal aerobic capacity (VO2max) was also unaffected by maternal WD, but HR had higher VO2max than C mice. Adult lean, fat, and total body masses were significantly increased by maternal WD, with greater increase for fat than for lean mass. Overall, no aspect of adult wheel running (total distance, duration, average running speed, maximum speed) or home-cage activity was statistically affected by maternal WD. However, analysis of the 8 individual lines revealed that maternal WD significantly increased wheel running in one of the 4 HR lines. On average, all groups lost fat mass after 6days of voluntary wheel running, but the absolute amount lost was greater for mice with maternal WD resulting in no effect of maternal WD on absolute or % body fat after wheel access. All groups gained lean and total body mass during wheel access, regardless of maternal WD or linetype. Measured after wheel access, circulating leptin, adiponectin, and corticosterone concentrations were unaffected by maternal WD and did not differ between HR and C mice. With body mass as a covariate, heart ventricle mass was increased by maternal WD in both HR and C mice, but fat pads, liver, spleen, and brain masses were unaffected. As found previously, HR mice had larger brains than C mice. Body mass of grand-offspring was unaffected by grand-maternal WD, but grand-offspring wheel running was significantly increased for one HR line and decreased for another HR line by grand-maternal WD. In summary, maternal Western diet had long-lasting and general effects on offspring adult morphology, but effects on adult behavior were limited and contingent on sex and genetic background.

Relationship between Maximal Oxygen Consumption (VO2max) and Home Range Area in Mammals.

Home range is defined as the area traversed during normal daily activities, such as foraging, avoiding predators, and social or antagonistic behaviors. All else being equal, larger home ranges should be associated with longer daily movement distances and/or higher average movement speeds. The maximal rate of oxygen consumption (VO2max) generally sets an upper limit to the intensity of work (e.g., speed of locomotion) that an animal can sustain without fatigue. Therefore, home range area and VO2max are predicted to evolve in concert (coadapt). We gathered literature data on home range and VO2max for 55 species of mammals. We computed residuals from log-log (allometric) regressions on body mass with two different regression models: ordinary least squares (OLS) and phylogenetic generalized least squares (PGLS). Residuals were weakly positively related for both the OLS (r = 0.278, one-tailed P < 0.05) and PGLS (r = 0.210, P > 0.05) regressions. For VO2max, the PGLS regression model had a slightly higher likelihood than the OLS model, but the situation was reversed for home range area. In addition, for both home range area and VO2max, models that fit better than either OLS or PGLS were obtained by modeling residual variation with the Ornstein-Uhlenbeck process to mimic stabilizing selection (RegOU), indicating that phylogenetic signal is present in both size-adjusted traits, consistent with findings of previous studies. (However, residuals from the RegOU models cannot be tested for correlation due to mathematical complexities.) We conclude that the best estimate of the residual correlation is probably somewhere between these two values reported above. Possible reasons for the low correlation between residual home range area and VO2max are discussed.