Influence of endurance training on the age-related decline in hepatic glyconeogenesis.

Hepatic gluconeogenic and glyconeogenic capabilities were investigated in Fischer 344 rat livers (ages 7, 15 and 25 months; n = 66) to determine if endurance training affected age related decrements in these hepatic functions. Animals were trained 1 h/day, 5 days/week for 10 weeks at treadmill speeds of 75% of age-specific maximal capacity. After training, rats were injected (300 mg/kg) with a known gluconeogenic inhibitor, 3-mercaptopicolinic acid (MPA). Two endurance tests were performed to help assess the contribution of gluconeogenesis to exercise performance, an initial test 4 days prior to injection and a final test immediately post-injection. MPA significantly (P < 0.05) reduced running times in all trained groups compared to their control test: 89%, 81%, and 51% in the young, middle-aged, and old, respectively. MPA reduced running times in the untrained animals 19%, 11%, and 8%, respectively. Three days after the last exercise bout, the animals were anesthetized and liver sections were sliced and incubated in [14C]lactic acid or [14C]fructose. An age-related decline was found in [14C]lactate incorporation (middle-aged decreases 66%, old decreases 54%) and in [14C]fructose incorporation (middle-aged decreases 51%, old decreases 48%) into glycogen. Differences existed in lactate incorporation in trained compared to untrained animals for the young, middle-aged, and old groups: 150.1 +/- 11.3 vs. 102.1 +/- 10.0; 75.3 +/- 6.2 vs. 34.9 +/- 6.4; and 69.3 +/- 14.9 vs. 47.0 +/- 4.7 nmol/g/h, respectively. No differences were found with training in any of the age groups for fructose. Phosphoenolpyruvate carboxykinase (PEPCK) activity and messenger RNA (mRNA) were significantly reduced in the old compared to the young rats (decreases 64% and decreases 58%, respectively). No training effects were found for either PEPCK activity or mRNA for any age group. These results suggest that hepatic gluconeogenic and glyconeogenic capabilities declined with age. Training had an effect in attenuating the glyconeogenic decline but had a minimal effect in offsetting the age-related decline in PEPCK.

Postexercise thermoregulatory behavior and recovery from exercise in desert iguanas.

Desert iguanas (Dipsosaurus dorsalis) undergo respiratory recovery more rapidly and incur lower energetic costs when they recover from 40 degrees C burst activity at 20 degrees C than when they recover at 40 degrees C. However, a body temperature of 20 degrees C falls well outside the preferred activity temperature range of this species, and imposes several physiological and behavioral liabilities. To determine if exhausted animals would favor a thermal regimen that allows for rapid and inexpensive respiratory recovery, we exercised lizards to exhaustion and allowed them to recover in a laboratory thermal gradient for 180 min. Recovering animals allowed their body temperatures to cool significantly to a mean temperature of 33.5 degrees C during the first 60 min of recovery, and subsequently rewarmed themselves to an average temperature of 38 degrees C for the remainder of their recovery period. Control animals maintained a constant body temperature of 37.7 degrees C throughout the 180-min recovery period. We then exercised animals to exhaustion at 40 degrees C and allowed them to recover for 180 min under a thermal regimen that mimicked that selected by exhausted animals in the previous experiment. Animals recovering under this thermal regimen returned to rates of O2 consumption, removed exercise-generated blood lactate, and incurred energetic costs that were more similar to data previously collected for animals recovering from exercise at a constant 40 degrees C than to data from animals recovering at 20 degrees C. These results suggested that the energetic benefits associated with recovery at 20 degrees C are not of sufficient biological importance to cause a major shift in thermoregulatory behavior.

Maximal sprint speeds and muscle fiber composition of wild and laboratory house mice.

We compared males from four groups of house mice (Mus domesticus), all bred and raised under common conditions in the laboratory: randombred Hsd:ICR; a wild population from Wisconsin; hybrids from lab dams; hybrids from wild dams. Wild mice were much faster sprinters (maximal forced sprint speeds over 1.0 m ranged from 2.38 to 3.34 m/s) than were lab mice (range = 0.89-1.68 m/s). Hybrids exhibited intermediate speeds (range = 1.54-2.70 m/s) and body masses, indicating largely additive inheritance. Type-specific mean muscle fiber cross-sectional areas of the gastrocnemius muscle did not differ significantly among groups. Percentage cross-sectional areas occupied by each of the three identified fiber types also did not differ significantly among groups, nor did they covary with body mass. For their body mass, however, lab mice had smaller gastrocnemius muscles than did wild and hybrid mice, which had muscles of similar size. Although we cannot rule out the possibility that smaller gastrocnemius muscles or slight differences in fiber composition account for the lower sprint speeds of the lab mice, we suggest that differences in unmeasured physiological, behavioral or motivational factors are probably the primary cause. This interpretation is supported by a lack of correlation between individual differences in sprint speed and either relative gastrocnemius muscle mass or muscle fiber type composition.

Comparative analysis of fiber-type composition in the iliofibularis muscle of phrynosomatid lizards (Squamata).

The lizard family Phrynosomatidae comprises three subclades: the closely related sand and horned lizards, and their relatives the Sceloporus group. This family exhibits great variation in ecology, behavior, and general body plan. Previous studies also show that this family exhibits great diversity in locomotor performance abilities; as measured on a high-speed treadmill, sand lizards are exceptionally fast sprinters, members of the Sceloporus group are intermediate, and horned lizards are slowest. These differences are paralleled by differences in relative hindlimb span. To determine if muscle fiber-type composition also varies among the three subclades, we examined the iliofibularis (IF), a hindlimb muscle used in lizard locomotion, in 11 species of phrynosomatid lizards. Using histochemical assays for myosin ATPase, an indicator of fast-twitch capacity, and succinic dehydrogenase, denoting oxidative capacity, we classified fiber types into three categories based on existing nomenclature: fast-twitch glycolytic (FG), fast-twitch oxidative-glycolytic (FOG), and slow-twitch oxidative (SO). Sand lizards have a high proportion of FG fibers (64-70%) and a low proportion of FOG fibers (25-33%), horned lizards are the converse (FG fibers 25-31%, FOG fibers 56-66%), and members of the Sceloporus group are intermediate for both FG (41-48%) and FOG (42-45%) content. Hence, across all 11 species %FOG and %FG are strongly negatively correlated. Analysis with phylogenetically independent contrasts indicate that this negative relationship is entirely attributable to the divergence between sand and horned lizards. The %SO also varies among the three subclades. Results from conventional nested ANCOVA (with log body mass as a covariate) indicate that the log mean cross-sectional area of individual muscle fibers differs among species and is positively correlated with body mass across species, but does not differ significantly among subclades. The log cross-sectional area of the IF varies among species, but does not vary among subclades. Conversely, the total thigh muscle cross-sectional area does not vary among species, but does vary among subclades; horned lizards have slimmer thighs. Muscle fiber-type composition appears to form part of a coadapted suite of traits, along with relative limb and muscle sizes, that affect the locomotor abilities of phrynosomatid lizards.

Skeletal muscle substrate utilization is altered by acute and acclimatory temperature in the American bullfrog (Lithobates catesbeiana).

We investigated the effect of acute and acclimatory temperature on the relative contribution of g9lucose and lactate to metabolism in resting sartorius muscle of the American bullfrog (Lithobates catesbeiana). We examined the fate of these metabolites in vitro by supplying radiolabeled [(14)C]glucose, [(14)C]lactate and [(14)C]palmitate to isolated muscle bundles from frogs (1) acutely exposed to incubation conditions of 5, 15 or 25 degrees C, (2) acclimated for 2-6 weeks to 5 or 25 degrees C or (3) acclimated for 2-6 weeks to 5 or 25 degrees C and the muscles incubated at 15 degrees C. Under all three temperature conditions tested, net rate of lactate metabolism exceeded that of glucose. Acute exposure to 5 degrees C reduced net rate of glucose metabolism by 15x and net lactate metabolism by 10x as compared with 25 degrees C-exposed tissues. Acclimation to 5 degrees C favored glucose storage as glycogen and increased the proportion of lactate oxidized (versus stored or converted to glucose) when compared with 25 degrees C-acclimated tissues. Net rates of storage of lactate as glycogen (glyconeogenesis) were significantly higher in muscles from 5 degrees C-acclimated frogs during incubation at a common temperature of 15 degrees C. These data suggest that lactate is the predominant fuel for resting skeletal muscle over this temperature range, and particularly so under cold conditions. Ready use of lactate as a substrate, and enhancement of glyconeogenic pathways in response to cold acclimation, could play a role in the tolerance of this species to seasonal temperature changes by promoting sequestration and storage of available substrate under cold conditions.

Scaling the duration of activity relative to body mass results in similar locomotor performance and metabolic costs in lizards.

This study examines the physiological response to locomotion in lizards following bouts of activity scaled to body mass. We evaluate this method as a way to compare locomotor energetics among animals of varying body mass. Because most of the costs of brief activity in reptiles are repaid during recovery we focus on the magnitude and duration of the excess post-exercise oxygen consumption (EPOC). Lizards ranging from 3 g to 2400 g were run on a treadmill for durations determined by scaling the run time of each animal to the 1/4 power of body mass and allowing each animal to run at its maximum speed for that duration. This protocol resulted in each species traveling the same number of body lengths and incurring similar factorial increases in V(O(2)). Following activity, EPOC volume (ml O(2)) and the cost of activity per body length traveled (ml O(2) per body length) scaled linearly with body mass. This study shows that the mass-specific costs of activity over an equivalent number of body lengths are similar across a broad range of body mass and does not show the typical patterns of allometric scaling seen when cost of locomotion are expressed on a per meter basis. Under field conditions larger animals are likely to travel greater absolute distances in a given bout of activity than smaller animals but may travel a similar number of body lengths. This study suggests that if locomotor costs are measured on a relative scale (ml O(2) per body length traveled), which reflects these differences in daily movement distances, that locomotor efficiency is similar across a wide range of body mass.

Characterization of circannual patterns of metabolic recovery from activity in Rana catesbeiana at 15 degrees C.

We characterized carbohydrate metabolism following activity in the American bullfrog, Rana catesbeiana, and compared whole body metabolic profiles between two seasons. Forty-eight adult male Rana catesbeiana were chronically cannulated and injected with [U-(14)C]L-lactic acid sodium salt in either summer (June) or winter (January) after acclimation for 2 weeks at 15 degrees C with a 12 h:12 h L:D photoperiod. Following injection with [(14)C]lactate, frogs were either allowed to rest for 240 min (REST), hopped for 2 min on a treadmill and immediately sacrificed (PE), or hopped for 2 min on a treadmill and allowed to recover for 240 min (REC 4). Exercise caused a significant increase in blood lactate level from 2.7+/-0.1 mmol l(-1) at rest to 17.0+/-2.1 mmol l(-1) immediately following exercise. This increase persisted throughout the recovery period, with average blood lactate level only reduced to 13.7+/-1.1 mmol l(-1) after 240 min of recovery, despite complete recovery of intramuscular lactate levels. Lactate levels were not significantly different between seasons in any treatment (REST, PE, REC4), in either gastrocnemius muscle or blood. The vast majority of [(14)C]lactate was recovered in the muscle, in both winter (86.3%) and summer (87.5%). Season had no effect on total amount of (14)C label recovered. [(14)C]Lactate was measured in the forms of lactate, glucose and glycogen, in the liver and the muscle sampled. The most robust difference found in seasonal metabolism was that both the liver and the gastrocnemius contained significantly higher levels of intracellular free glucose under all treatments in winter. These data suggest that, overall, bullfrogs accumulate and slowly clear lactate in a manner quite similar to findings in fish, other amphibians and lizards. Additionally, our findings indicate that lactate metabolism is not highly influenced by season alone, but that intracellular glucose levels may be sensitive to annual patterns.

Post-exercise lactate metabolism: a comparative review of sites, pathways, and regulation.

Most vertebrates utilize supplemental lactate production to support the energetic demands of vigorous, brief exercise. Despite similar patterns of accumulation, there appears to be a trichotomy with regards to lactate processing post-exercise. Most fish retain most of their lactate intramuscularly, using it for in situ glycogen replenishment. Recent evaluation of fish muscle concludes that pyruvate kinase reversal is a probable gluconeogenic pathway. Amphibians and reptiles also utilize lactate as a muscle glyconeogenic substrate, but lactate is not sequestered post-exercise. None of these groups utilize hepatic gluconeogenesis to any significant extent post-exercise, and muscle glucose uptake is limited. Lactate oxidation plays a major role post-exercise in mammals, with hepatic and muscular gluco- and glyconeogenesis contributing to a lesser extent. Glucocorticoids may regulate lactate release from fish muscle, although catecholamines may influence glyconeogenesis in reptile muscle. Insulin affects lactate metabolism indirectly through its effects on muscle glucose metabolism.

A histochemical and enzymatic study of the muscle fiber types in the water monitor, Varanus salvator.

Histochemical analysis of five muscles from the water monitor, Varanus salvator, identified three major classes of fibers based on histochemical activities of the enzymes myosin ATPase (mATPase), succinic dehydrogenase (SDH), and alpha-glycerophosphate dehydrogenase (alpha GPDH). Fast-twitch, glycolytic (FG) fibers were the most abundant fiber type and exhibited the following reaction product intensities: mATPase, dark; SDH, light; alpha GPDH, moderate to dark. Fast-twitch, oxidative, glycolytic (FOG) fibers were characteristically mATPase, dark; SDH, light; alpha GPDH, moderate to dark. The third class of fibers had the following histochemical characteristics: mATPase, light; SDH, moderate to dark; alpha GPDH, light. These fibers were considered to be either slow twitch, or tonic, and oxidative (S/O). Pyruvate kinase (PK), alpha GPDH, and citrate synthase (CS) activities were measured in homogenates of the same muscles studied histochemically. There was a positive relationship between both PK and alpha GPDH activities and the percentage of glycolytic fiber types within a muscle. Likewise, CS activities were greater in muscles high in FOG and S/O content. Based on CS activities, Varanus S/O fibers were eight-fold more oxidative than FG fibers within the same muscle. PK/CS ratios suggested that FG fibers possess high anaerobic capacity, similar to the iguanid lizard Dipsosaurus. The fiber type composition of the gastrocnemius muscle, relative to that of other lizard species, suggests that varanid lizards may possess a greater proportion of FOG and S/O fibers than other lizards.

Glycogen synthesis from lactate in skeletal muscle of the lizard Dipsosaurus dorsalis.

The capacity of skeletal muscle to synthesize glycogen from lactate was tested in the iliofibularis muscle of the desert iguana, Dipsosaurus dorsalis. Like other reptiles, Dipsosaurus accumulates significant lactic acid concentrations following vigorous exercise. After 5 min of progressively faster treadmill running at 35 degrees C (final speed = 2.2 km/h), blood lactate concentration increased over 14 mM, which decreased 11 mM after 2 h of recovery. Blood glucose concentration remained unchanged throughout at 8.6 +/- 0.46 mM. The role that muscle gluconeogenesis might play in the removal of post-exercise lactate was evaluated. Animals were run to exhaustion at 1.5 km/h on a treadmill thermostatted at 35 degrees C. Animals (n = 43) ran 6.9 +/- 0.75 min prior to exhaustion. Animals were sacrificed and iliofibularis muscles of both hindlimbs removed and stimulated at 2 Hz for 5 min, reducing twitch tension to 6% of prestimulus tension. Fatigued muscles were then split into red and white fiber bundles and incubated 2 h or 5 h at 35 degrees C in Ringer solution or in Ringer plus 20 mM lactate. In muscles tested in August, red fiber bundles incubated in lactate demonstrated a rate of glycogen synthesis of approximately 1 mg/(g muscle . h). In muscles tested in December, red fiber bundles synthesized glycogen at a reduced rate that was not statistically different than in fiber bundles incubated in Ringer solution without lactate. Glycogen synthesis from lactate was not evident in white fiber bundles in either August or December. The period of peak gluconeogenic capacity coincides with the field active season of Dipsosaurus.(ABSTRACT TRUNCATED AT 250 WORDS)

Metabolic recovery from exhaustive activity by a large lizard.

Gas exchange (VO2 and VCO2) and blood lactate concentration were measured in the lizard Amblyrhynchus cristatus at 25 and 35 degrees C during resting, running, and recovery after exhaustion (less than or equal to 180 min) to analyze the temperature dependency of metabolic recovery in this lizard. Amblyrhynchus exhausted twice as fast (4.2 vs. 8.8 min) at 25 degrees C than when running at the same speed at 35 degrees C. At both temperatures, VO2 and VCO2 increased rapidly during activity and declined toward resting levels during recovery in a manner similar to other vertebrates. Respiratory quotients (R, where R = VCO2/VO2) exceeded 2.0 after exhaustion at both temperatures. Extensive lactate production occurred during activity; blood lactate concentrations ranged from 1.0 to 1.7 mg lactate/ml blood after activity. Net lactate removal exhibited a temperature dependence. Blood lactate concentrations remained elevated hours after VO2 returned to normal. Endurance was reduced in lizards that had recovered aerobically but still possessed high lactate concentrations. The temporal separation of the excess oxygen consumption and lactate removal suggests that the concept of the lactacid oxygen debt is not applicable to this animal. The temperature dependence of total metabolic recovery suggests a benefit for Amblyrhynchus that select warm basking temperatures following strenuous activity.

Activity before exercise influences recovery metabolism in the lizard Dipsosaurus dorsalis.

During recovery from even a brief period of exercise, metabolic rate remains elevated above resting levels for extended periods. The intensity and duration of exercise as well as body temperature and hormone levels can influence this excess post-exercise oxygen consumption (EPOC). We examined the influence of activity before exercise (ABE), commonly termed warm-up in endotherms, on EPOC in the desert iguana Dipsosaurus dorsalis. The rate of oxygen consumption and blood lactate levels were measured in 11 female D. dorsalis (mass 41.1 +/- 3.0 g; mean +/- s.e.m.) during rest, after two types of ABE and after 5 min of exhaustive exercise followed by 60 min of recovery. ABE was either single (15 s of maximal activity followed by a 27 min pause) or intermittent (twelve 15 s periods of exercise separated by 2 min pauses). Our results indicate that both single and intermittent ABE reduced recovery metabolic rate. EPOC volumes decreased from 0.261 to 0.156 ml of oxygen consumed during 60 min of recovery when lizards were subjected to intermittent ABE. The average cost of activity (net V(O2) during exercise and 60 min of recovery per distance traveled) was almost 40 % greater in lizards that exercised without any prior activity than in lizards that underwent ABE. Blood lactate levels and removal rates were greatest in animals that underwent ABE. These findings may be of particular importance for terrestrial ectotherms that typically use burst locomotion and have a small aerobic scope and a long recovery period.

Evidence for facilitated lactate uptake in lizard skeletal muscle.

To understand more fully lactate metabolism in reptilian muscle, lactate uptake in lizard skeletal muscle was measured and its similarities to the monocarboxylate transport system found in mammals were examined. At 2 min, uptake rates of 15 mmol l(-1) lactate into red iliofibularis (rIF) were 2.4- and 2.2-fold greater than white iliofibularis (wIF) and mouse soleus, respectively. alpha-Cyano-4-hydroxycinnamate (15 mmol l(-1)) caused little inhibition of uptake in wIF but caused a 42-54 % reduction in the uptake rate of lactate into rIF, suggesting that much of the lactate uptake by rIF is via protein-mediated transport. N-ethymaleimide (ETH) (10 mmol l(-1)) also caused a reduction in the rate of uptake, but measurements of adenylate and phosphocreatine concentrations show that ETH had serious effects on rIF and wIF and may not be appropriate for transport inhibition studies in reptiles. The higher net uptake rate by rIF than by wIF agrees with the fact that rIF shows much higher rates of lactate utilization and incorporation into glycogen than wIF. This study also suggests that lactate uptake by reptilian muscle is similar to that by mammalian muscle and that, evolutionarily, this transport system may be relatively conserved even in animals with very different patterns of lactate metabolism.

The effects of intensity on the energetics of brief locomotor activity.

The energetic costs associated with locomotion are often estimated only from the energy expended during activity and do not include the costs incurred during recovery. For some types of locomotion, this method overlooks important aspects of the metabolic costs incurred as a result of the activity. These estimates for energetic cost have also been predicted from long-duration, low-intensity activities that do not necessarily reflect all the behavior patterns utilized by animals in nature. We have investigated the effects of different activity intensities on the metabolic expenditure (per unit distance traveled) associated with brief exercise, and offer a more inclusive analysis of how the energetics of short-duration activities might be analyzed to estimate the costs to the animal. Mice ran on a treadmill for 15 or 60 s at 25 %, 50 % or 100 % of maximum aerobic speed (MAS) while enclosed in an open-flow respirometry system. Following the run, each mouse was allowed to recover while remaining enclosed in the respirometry chamber. Excess exercise oxygen consumption (EEOC), the excess volume of oxygen consumed during the exercise period, increased with the duration and increased linearly with the intensity of exercise. In contrast, the volume of oxygen consumed during the recovery period, or excess post-exercise oxygen consumption (EPOC), was independent of exercise intensity and duration and accounted for more than 90 % of the total metabolic cost. The net cost of activity (C(act)), calculated by summing EEOC and EPOC and then dividing by the distance run, increased as both activity duration and intensity decreased. The values for C(act) ranged from 553 ml O(2 )g(-)(1 )km(-)(1) for a 15 s run at 25 % MAS to 43 ml O(2 )g(-)(1 )km(-)(1) for a 60 s run at 100 % MAS. Combining these data with data from a companion paper, we conclude (1) that EPOC is independent of both the duration and intensity of activity when exercise duration is brief in mice, (2) that EPOC accounts for a majority of the oxygen consumed as a result of the activity when exercise durations are short, and (3) that animals can minimize their energy expenditure per unit distance by running faster for a longer period.

Muscle composition and its relation to sprint running in the lizard Dipsosaurus dorsalis.

Iguanid lizards exhibit considerable intraspecific variation in several aspects of their muscle composition. To determine the relationship of this variation to the variation in locomotor performance, running speeds of 20 male desert iguanas (Dipsosaurus dorsalis) of similar mass were measured from video recordings of animals as they sprinted down a 4.9-m runway maintained at 40 degrees C, the preferred body temperature of Dipsosaurus. Mean sprint speed ranged from 2.2 to 4.2 m/s. Selected muscles from these animals were then analyzed histochemically for fiber type size and composition, and the activities of citrate synthase, pyruvate kinase, and creatine kinase were measured. Muscle fiber cross-sectional areas were highly correlated within individuals, in three leg muscles and across all three fiber types, so that individuals could be characterized as possessing large or small fibers relative to the sample mean. Activities of all three enzymes also covaried within individuals so that individual lizards could be characterized as possessing high or low leg muscle catabolic capacity. There existed a significant and inverse relationship between fiber cross-sectional areas and muscle enzyme activities so that individuals with small muscle fibers tended to have higher catabolic capacities. Approximately 25-30% of the variation in mean sprint running speed could be predicted by variation in muscle fiber areas alone. The use of muscle fiber areas and snout vent length as independent variables in a multiple-regression equation explained approximately 50% of the sprint-running variation.(ABSTRACT TRUNCATED AT 250 WORDS)

Can energetic expenditure be minimized by performing activity intermittently?

Previous research has shown that the energetic expense per unit distance traveled for one bout of short-duration activity is much greater than the energetic expense associated with long-duration activity. However, animals are often seen moving intermittently, with these behaviors characterized by brief bouts of activity interspersed with brief pauses. We hypothesized that, when multiple bouts of brief activity are performed intermittently, the energetic cost per unit distance is less than when only one short bout is performed. Mice were run 1, 2, 3, 5, 9 or 13 times for 15 s at their maximal speed within a 375 s period while enclosed in an open-flow respirometry system on a treadmill. The mice were also run continuously for 375 s. Following the last sprint and the continuous run, the mice remained in the respirometry chamber until their vO2 reached resting levels. Excess exercise oxygen consumption (EEOC), the excess volume of oxygen consumed during the exercise period, increased from 0.03+/-0.01 to 0.40+/-0.02 ml O2g(-)(1) (mean +/- s.e.m., N=9) with activity frequency. However, the excess post-exercise oxygen consumption (EPOC), or volume of oxygen consumed during the recovery period, was independent of activity frequency (range 0.91-1.16 ml O2g(-)(1)) and accounted for more than 80 % of the total metabolic cost when activity was performed intermittently. Lactate concentration was measured at rest, immediately after running and immediately after recovering from running 1, 5 and 13 times within the 375 s period. After running, [lactate] was significantly higher than resting values, but following recovery, [lactate] had reached resting values. The net cost of activity, C(act), calculated by summing EEOC and EPOC and then dividing by the distance run, decreased significantly from 132+/-38 to 6+/-1 ml O2g(-)(1 )km(-)(1) as activity frequency increased. When these values for C(act) were compared with the cost of running continuously for the same amount of time, the values were identical. Therefore, we conclude that animals can minimize energetic expenditure by performing brief behaviors more frequently, just as they can minimize these costs if they increase the duration of continuous behaviors.

Acid-base imbalance in lizards during activity and recovery.

1. The effects of treadmill exercise on oxygen consumption (V02), carbon dioxide production (VCO2), arterial blood lactate concentration ([L-]a), arterial blood pH and arterial gas tensions (PaO2 and PaCO2) were measured in 3 species of lizards (Varanus salvator, V. exanthematicus, Iguana iguana) 2. Varanus salvator was exercised 45 min at an intensity which required 85% of its VO2 max. V. salvator utilized supplementary anaerobic metabolism during the first 10 min of this sustainable exercise, as evidenced by a 16 mmol/l increase in [L-]a. Respiratory exchange ratios (R, where R = VCO2/VO2) exceeded 1.2 when [L-]a and [H+]a were maximal. One half of the accumulated lactate was removed from the blood during the remainder of the 45 min exercise period, while blood pH returned to resting levels. 3. In a second set of experiments, high intensity exercise led to exhaustion after 5 to 10 min in all three species, resulting in large lactate (+ delta[L-]a = 14-20 mmol/l) and hydrogen ion (+ delta[H+]a = 23-57 nmol/l) accumulations. R values ranged from 1.2-1.8 at exhaustion. 4. Recovery from both sustainable and non-sustainable exercise was characterized as a period of rapid lactate removal. Respiratory exchange ratios were low (0.3-0.5) as metabolic CO2 was retained, replacing depleted bicarbonate stores. 5. We conclude that all three lizard species make ventilatory adjustments during and after exercise that minimize disturbances to resting hydrogen ion concentrations and acid-base balance. Varanus salvator demonstrate the ability to re-establish resting acid-base status during sustained exercise requiring 85% of their VO2,max. Changes in R appear to be a useful noninvasive indicator of net blood lactate accumulation.

Regulation of skeletal muscle metabolism in the lizard Dipsosaurus dorsalis by fructose-2,6-bisphosphate.

Changes in liver and skeletal muscle fructose-2,6-bisphosphate (Fru-2,6-P2) concentrations were compared during fasting, exercise, and recovery in the lizard Dipsosaurus dorsalis and in outbred mice (Mus musculus). We present the first correlative evidence that suggests that a decrease in the content of Fru-2,6-P2 may mediate elevated gluconeogenesis in lizard skeletal muscle. Contents of Fru-2,6-P2 in lizard gastrocnemius and red and white iliofibularis (IF) were significantly lower (as much as 55% in white IF) during recovery from exhaustive exercise than at rest. Recovery from exhaustive exercise had no significant effect on Fru-2,6-P2 concentrations in any mouse muscle examined. Fasting significantly depressed lizard and mouse liver Fru-2,6-P2 contents and decreased lizard red IF by over 84% from the fed condition. Lizard red and white muscle fiber bundles incubated in 20 mM lactate had significantly lower Fru-2,6-P2 (94 and 61% depression, respectively) than those incubated in 8.5 mM glucose. These results are consistent with the hypothesis that Fru-2,6-P2 acts as a signal for controlling gluconeogenesis in lizard skeletal muscle.

Cardiovascular response to treadmill exercise in untrained rats.

Oxygen consumption (Vo2), cardiac output (Q), heart rate (HR), stroke volume (SV), and oxygen extraction from blood (Cao2-Cvo2) were measured in untrained rats, both at rest and during treadmill running at various speeds (10-41 m/min). Vo2 increased linearly as a function of running speed, and maximal values (83 ml O2.kg-1 min-1) represented a five-fold increase over resting values. Q, HR, SV, and Cao2-Cvo2 increased linearly as functions of Vo2. Mixed venous oxygen content (Cao2) decreased with increasing Vo2; whereas arterial oxygen content (Cao2) remained independent of Vo2; whereas arterial oxygen content (Cao2) remained independent of Vo2, averaging 19 vol%. Maximal values of these variables and their relationship to Vo2 were as follows: Q = 4.3 Vo2 + 184; Qmax = 543 ml.kg-1.min-1, HR = 3.02 Vo2 + 340; HRmax 595 beats.min-1; SV = 0.004 Vo2 + 0.603; SVmax = 0.92 ml.kg-1.beat-1; Cao2-Cvo2 = 0.13 Vo2 + 5.62; Cao2-Cvo2max = 15.5 vol%.; Cvo2 = -0.12 Vo2 + 12.94; Cvo2min - 3.4 vol%. These data suggest that HR, SV, and Cao2-Cvo2 make significant contributions to the augmentation of Vo2 in the exercising rat.

Lactate and glucose metabolism in mouse (Mus musculus) and reptile (Anolis carolinensis) skeletal muscle.

The reliance on anaerobic metabolism during exercise in lizards has been the subject of a growing body of literature in activity metabolism. Prior studies have demonstrated that lizards rely more on postexercise lactate to regenerate depleted glycogen stores than do many mammals. These studies prompted an in vitro comparison between the metabolic mechanisms for the handling of lactate and glucose in the muscles of a small mammal and lizard. Hindlimb muscles of Mus and Anolis were stimulated to fatigue and then incubated in the presence of 15 mM lactate and either 5.5 (mice) or 8.5 (anoles) mM glucose. Oxidation rates of lactate and glucose were seven to eight times higher in mice. Both species oxidized more lactate than glucose (8 to 9 times). However, anole muscle showed a preference for lactate as a substrate for glycogenesis, incorporating 1.5 times as much lactate (expressed in glucose equivalents) as glucose. In contradistinction, mice incorporated 2.8 times as much glucose into glycogen as lactate. The quantitative differences in metabolic scope of mammals and reptiles are accompanied by fundamental differences in the capacity and patterns of skeletal muscle metabolism of lactate and glucose.