Cardiopulmonary Function in Iguanids
Includes abstracts on post-exertional recovery
Compiled by Melissa Kaplan
Circulatory impairment induced by exercise in the lizard Iguana iguana. Farmer CG; Hicks JW Department of Ecology and Evolutionary Biology, University of California at Irvine, Irvine, CA 92697, USA. J Exp Biol 2000 Sep;203 (Pt 17):2691-7
Mechanical integration of the cardiac, muscular and ventilatory pumps enables mammals to vary cardiac output over a wide range to match metabolic demands. We have found this integration lacking in a lizard (Iguana iguana) that differs from mammals because blood flow from the caudal body and ventilation are maximal after, rather than during, exercise. Because Iguana iguana are constrained from ventilation during intense locomotion, they appear to be unable to recruit the abdomen and thorax as a pump for venous return. This constraint on simultaneous running and costal breathing arises from their musculoskeletal design, which is similar to that of basal tetrapods, and so a constraint on venous return during exercise may be ancestral for tetrapods. We suggest that mechanical coupling of the pulmonary and cardiac pumps may have been important for the evolution of high-speed locomotor stamina in terrestrial vertebrates.
Ventilation and gas exchange in lizards during treadmill exercise. Wang T; Carrier DR; Hicks JW. Department of Ecology, University of California at Irvine, Irvine, CA 92717, USA. J Exp Biol 1997 Oct;200 (Pt 20):2629-39
The extent to which lizards ventilate their lungs during locomotion is controversial. Direct measurements of airflow across the nostrils suggest a progressive reduction in tidal volume and minute ventilation with increased running speed, while other studies have demonstrated that arterial PO2 remains constant during exercise. To resolve these conflicting findings, we measured minute ventilation and gas exchange rate in five specimens of Varanus exanthematicus and five specimens of Iguana iguana during treadmill locomotion at speeds between 0.14 and 1.11ms-1 at 35 degrees C. These speeds are much lower than maximal running speeds, but are greater than the maximal aerobic speed. In both species, the ventilatory pattern during locomotion was highly irregular, indicating an interference between locomotion and lung ventilation. In Varanus exanthematicus, treadmill locomotion elicited a six- to eightfold increase in minute ventilation from a pre-exercise level of 102mlkg-1min-1, whereas the rate of oxygen uptake increased approximately threefold (from 3.9 to 12.6mlkg-1min-1). After exercise, both minute ventilation and gas exchange rate decreased immediately. Because minute ventilation increased more than did oxygen consumption, an increase in lung PO2 during exercise is predicted and, thus, Varanus exanthematicus appears effectively to ventilate its lungs to match the increased metabolic rate during locomotion at moderate speed. In Iguana iguana, both minute ventilation and gas exchange rate increased above resting values during locomotion at 0.28ms-1, but both decreased with further increases in locomotor speed. Furthermore, following exercise, both minute ventilation and oxygen uptake rate increased significantly. Iguana iguana, therefore, appears to be unable to match the increased oxygen demand with adequate ventilation at moderate and higher speeds.
Characterization of the renal portal system of the common green iguana (Iguana iguana) by digital subtraction imaging. Benson KG; Forrest L. Department of Medical Sciences, School of Veterinary Medicine, University of Wisconsin, Madison 53706, USA. J Zoo Wildl Med 1999 Jun;30(2):235-41
Digital subtraction angiography was used to map the venous blood flow from the pelvic limb and the tail in the common green iguana (Iguana iguana). The majority of blood returning from the pelvic limb bypassed the kidney and entered the general circulation, whereas venous blood flow from the tail entered the renal portal circulation. No evidence was found of a renal portal valve.
Activity before exercise influences recovery metabolism in the lizard Dipsosaurus dorsalis. Scholnick DA; Gleeson TT. Department of Biology, Eckerd College, Marine Sciences, St Petersburg, FL 33711, USA. J Exp Biol 2000 Jun;203 Pt 12:1809-15
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.
Contribution of gular pumping to lung ventilation in monitor lizards. Owerkowicz T; Farmer CG; Hicks JW; Brainerd EL. Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138, USA. Science 1999 Jun 4;284(5420):1661-3
A controversial hypothesis has proposed that lizards are subject to a speed-dependent axial constraint that prevents effective lung ventilation during moderate- and high-speed locomotion. This hypothesis has been challenged by results demonstrating that monitor lizards (genus Varanus) experience no axial constraint. Evidence presented here shows that, during locomotion, varanids use a positive pressure gular pump to assist lung ventilation. Disabling the gular pump reveals that the axial constraint is present in varanids but it is masked by gular pumping under normal conditions. These findings support the prediction that the axial constraint may be found in other tetrapods that breathe by costal aspiration and locomote with a lateral undulatory gait.
The influence of corticosterone and glucagon on metabolic recovery from exhaustive exercise in the desert iguana Dipsosaurus dorsalis. Scholnick DA; Weinstein RB; Gleeson TT. University of Colorado, Department of Environmental Population and Organismic Biology, Boulder 80309-0334, USA. Gen Comp Endocrinol 1997 May;106(2):147-54
The skeletal muscles of ectothermic vertebrates possess an elevated glyconeogenic capacity that is responsible for a major portion of lactate removal and glycogen resynthesis following exercise. In lizards, changes in plasma hormone levels and the influence of differing hormone levels on muscle metabolism postexercise are poorly understood. We measured the effects of 5 min of exhaustive exercise on plasma levels of glucagon and corticosterone in the desert iguana Dipsosaurus dorsalis. We also determined the extent to which these hormones influence, or are influenced by, postexercise plasma lactate concentrations postexercise. Exercise resulted in the accumulation of 20 mM blood lactate, while plasma glucose levels remained stable throughout 90 min of recovery. Plasma glucagon was elevated sevenfold during 5 min of exercise and returned to resting levels within 45 min of recovery. Glucagon stimulated lactate incorporation into glycogen in isolated red muscle fiber bundles. Plasma corticosterone was also elevated to three times normal resting values, but only after 45 min of recovery. Blocking corticosterone elevation with metyrapone did not alter the kinetics of plasma lactate removal. In lizards, the dramatic rise in plasma glucagon occurs at the same time as previously reported elevated skeletal muscle glyconeogenesis and elevated glucagon stimulates lactate removal in vitro, strongly suggesting a role for glucagon in postexercise skeletal muscle metabolism.
Postexercise thermoregulatory behavior and recovery from exercise in desert iguanas. Wagner EL; Gleeson TT. Department of Environmental, Population and Organismic Biology, University of Colorado, Boulder 80309-0334, USA. Physiol Behav 1997 Feb;61(2):175-80
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.
Plasma catecholamine and corticosterone and their in vitro effects on lizard skeletal muscle lactate metabolism. Gleeson TT; Dalessio PM; Carr JA; Wickler SJ; Mazzeo RS; Department of Environmental, Population, and Organismic Biology, University of Colorado, Boulder 80309-0334. Am J Physiol 1993 Sep;265(3 Pt 2):R632-9
Lizard skeletal muscles utilize primarily lactate as a gluconeogenic substrate for glycogen replenishment following exercise. To understand the influence of selected hormones on this process, we measured changes in plasma catecholamines and corticosterone resulting from exercise in the lizard Dipsosaurus dorsalis and then investigated the physiological effects of those hormones on skeletal muscle lactate and glucose metabolism in vitro. Plasma epinephrine (Epi), norepinephrine, and corticosterone (Cort) increased 5.8, 10.2, and 2.2 times, respectively, after 5 min of exhaustive exercise. Epi and Cort levels remained elevated after 2 h of recovery. Skeletal muscle fiber bundles isolated from the red and white regions of the iliofibularis muscle were incubated 2 h at 40 degrees C in the presence of postexercise concentrations of [14C]lactate (15 mm) and glucose (8.5 mm) in the presence and absence of Epi or Cort. Red muscle oxidized both substrates at 2-3 times the rate of white muscle, and both red and white fibers oxidized lactate at 5-10 times the rate of glucose oxidation. Epi had a stimulatory effect on lactate oxidation by white muscle. Lactate incorporation into glycogen proceeded at 2-3 times the rate of glucose incorporation in both muscle types, with rates in red muscle again 2-3 times that for white muscle. Epi stimulated lactate carbon incorporation into glycogen by 50-140% in both red and white muscle but had no effect on glucose incorporation into glycogen in either tissue. We interpret these data as evidence that epinephrine stimulates lactate removal by skeletal muscle. Cort had no effect on lactate metabolism in either muscle type. [Abstract truncated at 250 words]
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