A steady-state model of maximal oxygen and carbon dioxide transport in anuran amphibians.

A steady-state model, incorporating pulmonary ventilation, pulmonary diffusion capacity, cardiovascular transport capacity, and tissue diffusion capacity, was developed to describe the maximal O2 and CO2 transport capacity for an anuran amphibian (Bufo). Solution of the model by iterative calculation closely predicted 1) the empirical maximal O2 consumption (VO2max) for Bufo, 2) variation in empirical VO2max for three other genera (Rana, Xenopus, Scaphiopus), and the empirically observed effects on VO2max of 3) hypobaric hypoxia, 4) artificially induced anemia, and 5) beta-blockade of heart rate increment with activity. The model indicates that cardiovascular transport is the rate-limiting step to VO2max in amphibians and that an increase in circulatory O2 transport is a major physiological adaptation for increasing total aerobic capacity. CO2 transport and body fluid PCO2 values were primarily determined by pulmonary ventilatory capacity, and to a lesser extent by cardiovascular transport. The model should be generally applicable to other terrestrial vertebrates.

Water loss, desiccation tolerance, and survival under desiccating conditions in 11 species of Caribbean Anolis : Evolutionary and ecological implications.

1. Rates of water loss and tolerance to desiccation were examined in 11 species of Caribbean Anolis and the Sonoran desert iguanid lizard, Urosaurus ornatus. 2. Rates of water loss ranged from 0.07% body wt/h (A. bonairensis) to 0.43% body wt/h (A. distichus). 3. There were significant correlations between habitat rainfall and both the rates of water loss (P<0.005) and the maximum time of survival (LTmax) (P<0.01) for the 11 species of Anolis. Species from areas of low rainfall generally had lower rates of water loss and survived longer than species from areas of high rainfall. 4. There was no correlation between habitat rainfall and the ability to withstand desiccation; therefore differences in LTmax are probably the result of differences in rates of water loss. 5. Rates of water loss did not significantly correlate with either structural niche or ecomorphic category. 6. Since rates of water loss varied to a large degree within closely related species groups, there is apparently little phylogenetic inertia for this physiologic parameter.

The role of pulmocutaneous baroreceptors in the control of lymphatic heart rate in the toad Bufo marinus.

The influence of increased pulmocutaneous arterial systolic pressure on amphibian lymph heart activity was determined in Bufo marinus. Arginine vasotocin-induced increases in systolic arterial pressure of greater than 0.5 kPa significantly (P<0.0002) decreased lymph heart rate. Denervation of the recurrent laryngeal nerve led to an increase in arterial pressure of 3.88 kPa in test animals compared to control animals. Denervation abolished lymph heart response to increased arterial pressure for an average of 38 min. Direct stimulation to the recurrent laryngeal nerve stopped lymph heart activity without an increase in arterial pressure. These data indicate that baroreceptors in the pulmocutaneous artery can decrease lymph heart rate during periods of either direct stimulation or elevated arterial pressure. The data also support lymph hearts as effectors for aortic and carotid baroreceptors.

Whole-body systemic transcapillary filtration rates, coefficients, and isogravimetric capillary pressures in Bufo marinus and Rana catesbeiana.

Whole-body and organ-level transcapillary filtration rates and coefficients are virtually unexamined in ectothermal vertebrates. These filtration rates appear to be greater than in mammals when plasma volume shifts and lymphatic function are analyzed. Gravimetric techniques monitoring whole-body mass changes were used to estimate net systemic filtration in Bufo marinus and Rana catesbeiana while perfusing with low-protein Ringer's and manipulating venous pressure. Capillary pressures were estimated from arterial and venous pressures after measuring the venous to arterial resistance ratio of 0.23. The capillary filtration coefficient (CFC) for the two species was 25.2+/-1.47 mL min-1 kg-1 kPa-1. Isogravimetric capillary pressure (Pci), the pressure at which net fluid is neither filtered nor reabsorbed, was 1.12+/-0.054 kPa and was confirmed by an independent method. None of these variables showed a significant interspecific difference. The anuran CFC and Pci are significantly higher than those found using the same method on rats (7.6+/-2.04 mL min-1 kg-1 kPa-1 and 0.3+/-0.37 kPa, respectively) and those commonly reported in mammals. Despite the high CFC, the high Pci predicts that little net filtration will occur at resting in vivo capillary pressures.

Effects of temperature and physical activity on blood flow shunts and intracardiac mixing in the toad Bufo marinus.

Blood flow in systemic (.Qsys) and pulmocutaneous (.Qpul) arteries was measured as a function of body temperature (10 degrees, 20 degrees, and 30 degrees C) at rest and following enforced physical activity in conscious, adult cane toads (Bufo marinus). Arterial and mixed venous hemoglobin concentration (CHb) and total oxygen content (Co2, tot) were measured in a separate group under identical conditions. Heart rate (fH) and total flow (.Qtot) increased significantly (P<0.001) with elevated temperature and with activity, whereas stroke volume (VS) increased (P<0.001) only with activity. .Qtot ranged about 10-fold, from 10 degrees C (rest) to 30 degrees C (activity); increases in both fH and VS contributed to the increase in .Qtot. The overall distribution of blood to the pulmocutaneous circuit (net L-R shunt) increased with both temperature and activity and was significantly correlated with .Qtot. These data indicate that blood flow distribution in toads is a direct function of cardiac output, and this is linked to relative changes in resistance in the major outflow vessels. Arterial O2 saturation (Sa) was high (mean=93%) in all conditions except activity at 30 degrees C, when it decreased to 74% and contributed to a decrease in the arteriovenous O2 difference. Venous O2 saturation (Sv) was high at rest (76%) and dropped significantly during activity to about 30% at all temperatures. Intracardiac arterial-venous mixing (systemic mixing index) showed the strongest correlation with variation in fH with minimal mixing (17%) occurring at about 50 beats min-1. The most mixing occurred at the lowest fH (13 beats min-1) and at the highest fH (103 beats min-1). The results indicate that the heart of a 0.25-kg toad becomes more efficient from an oxygen transport perspective from low fH to 50 beats min-1 and then less efficient at higher fH, contributing to an uncoupling of blood flow and metabolic rates at these high rates.

Nursing council ... coordination within decentralization.

A nursing council modeled after the traditional five nursing roles--manager, educator, practitioner, evaluator and researcher--improves patient outcomes and enhances nurse work satisfaction. The council provides a participative forum to address nursing and patient-care issues within a decentralized matrix organization.

The effects of in vivo and in vitro hyperosmolality on skeletal muscle performance in the amphibians Rana pipiens and Scaphiopus couchii.

1. The effect of in vivo and in vitro hyperosmolality on skeletal muscle function was investigated in two species of anuarans Scaphiopus couchii and Rana pipiens. 2. Muscle contractile performance, measured as peak tetanic tension declined to greater degree when tissue dehydration occurred in vitro rather than in vivo, even though tissue water contents were greater in vivo. 3. The muscles from S. couchii, a more dehydration tolerant species than R. pipiens, maintained tension at lower tissue water contents than R. pipiens. 4. Data for the effects of in vivo dehydration on plasma sodium, urea and osmotic concentration, as well as tissue water contents, are also presented for both species.

The hemodynamic consequences of hemorrhage and hypernatremia in two amphibians.

1. Graded hypovolemia was induced by hemorrhagic blood loss and graded hypernatremia by salt load in the toad, Bufo marinus, and the bullfrog, Rana catesbeiana. Maximal blood flow rates in the systemic arches and arterial and venous pressures were measured during activity after each stress. 2. Maximal blood flow rates in the B. marinus did not decline until blood loss exceeded 5% of initial body mass. In R. catesbeiana, losses of 2% initial body mass caused a decline (Fig. 1). 3. Maximal heart rates did not change with hemorrhage (Fig. 2). The decline in blood flow rates with hemorrhage was due to declining pulse volumes in both species (Fig. 3). 4. Arteriovenous pressure difference declined with hemorrhage in both species (Fig. 4). Peripheral resistance increased with hemorrhage in parallel with compromised blood flow rates (Fig. 5). 5. Plasma sodium concentration slightly increased with hemorrhage, while plasma protein concentration and hematocrit declined. Lymphatic compensation for hemorrhagic loss is indicated in both species (Fig. 6). 6. Induced hypernatremia compromised blood flow rates in both species at plasma sodium concentrations above 175 mM. The decline in flow rates was principally a result of a decrease in pulse volume, though maximal heart rates also declined (Figs. 2, 3, 7). 7. Induced hypernatremia had no effect on the arteriovenous pressure difference in B. marinus but caused it to decline in R. catesbeiana. Peripheral resistance increased in only B. marinus but not R. catesbeiana (Figs. 4, 5). Hematocrit did not change with salt load, indicative of a constant vascular volume.

Adrenergic control of the anuran cutaneous hydroosmotic response.

Adrenergic effectors were shown to play a major role in increasing cutaneous water uptake (the hydroosmotic response) in dehydrated anurans. A beta-adrenergic antagonist, propranolol, blocked 61% of the cutaneous response to dehydration and 67% of the response elicited by salt loading in the toad Bufo cognatus. A beta-adrenergic agonist, isoproterenol, elicited a response in normally hydrated animals equal to the propranolol-sensitive portion of the cutaneous hydroosmotic response of dehydrated animals. The isoproterenol-induced cutaneous water balance response in hydrated animals greatly exceeded the response induced by arginine vasotocin, the endogenous antidiuretic hormone.

An analysis of pH tolerance and substrate preference of isolated skeletal muscle mitochondria from Bufo marinus and Rana catesbeiana.

1. The effects of varying pH and substrate on isolated skeletal muscle mitochondria from Bufo marinus and Rana catesbeiana were investigated. 2. For both species, VO2 max significantly decreased at all pH < 7.3 (P < 0.05), while maximum values were observed at a pH range of 7.3-7.6 with B. marinus maintaining a greater VO2 max than R. catesbeiana. 3. Respiratory control values (RCR) decreased significantly at all pH < 6.9 for both species (P < 0.05). 4. Isolated mitochondria from both species were maintained at pH = 7.2 and O2 consumption measured under five separate substrate conditions. 5. A rank preference was established based upon state 3 and RCR values. 6. Substrate preference was identical for both species and interspecific comparisons revealed differences in state 3 respiration and coupling.

Effects of activity, hemorrhage, and dehydration on plasma catecholamine levels in the marine toad (Bufo marinus).

Resting plasma epinephrine and norepinephrine levels were 13.1 and 2.1 nmol liter-1 for the marine toad (Bufo marinus). Plasma catecholamine levels increased during enforced activity by five- to sixfold. Marine toads are remarkably tolerant of graded hemorrhagic loss of blood (over 10% mass loss). Plasma catecholamine levels did not increase at moderate blood loss, but increased substantially when cardiovascular variables (blood pressure, blood flow) were compromised and peripheral resistance was increased. Plasma catecholamine levels did not increase with dehydrational mass loss until a 15-20% loss of mass. The increase in plasma catecholamine concentration was correlated with an increase in vivo vascular resistance. Vascular resistance measured in vitro was unaltered at physiological catecholamine concentrations, although systemic resistance increased at pharmacological concentrations. The lack of effects of adrenalectomy on plasma catecholamine levels suggests that nerve terminal release, rather than adrenal secretion, may be the primary source of circulating catecholamines. We therefore suggest that circulating catecholamine levels are not an important endocrinological mechanism for defense of activity blood pressure, at least until it is compromised to the resting value.

Individual variation in maximum aerobic capacity: cardiovascular and enzymatic correlates in Rana catesbeiana.

We quantified natural variation in maximum aerobic capacity (V02max) exhibited by a free-living population of bullfrogs (Rana catesbeiana) and examined the degree to which such variation is associated with key parameters of the systemic oxygen transport apparatus and oxidative enzyme (citrate synthase) activity at the tissue level. Regression analysis of these data revealed that only ventricle mass and hemoglobin concentration accounted for significant fractions of the variation in V02max. Neither variation in maximum heart rate nor in citrate synthase activity were significantly correlated with individual variation in maximum aerobic capacity. These results support the contention that, in at least some taxa, maximum aerobic capacity is limited by the ability of the cardiovascular system to deliver oxygen to the tissues.

Water loss and nitrogen excretion in sharp-nosed reed frogs (Hyperolius nasutus: anura, Hyperoliidae).

Sharp-nosed African reed frogs, Hyperolius nasutus Gunther, are small (0.4 g) hyperoliids which have minimal rates of evaporative water loss (4.5 mg g-1 h-1; 0.3 mg cm-2 h-1) that are only 1/10 to 1/20 that of a typical frog, Hylaregilla, of comparable size (171 mg g-1 h-1, 4.8 mg cm-2 h-1). The surface-area-specific resistance to water flux of H. nasutus dorsal skin (96-257 sec cm-1) is similar to that of other 'waterproof' frogs (300-400), of cocooned frogs (40-500), and of desert reptiles (200-1400). However, H. nasutus can greatly increase the rate of evaporative water loss during radiative heat stress by mucous gland discharge, and by exposing the ventral skin. Urea is the principal nitrogenous waste product of H. nasutus and uric acid comprises less than 1% of the total nitrogen excretion for both H. nasutus and H. regilla. Other 'waterproof' frogs, in contrast, are uricotelic. Lethal dehydration requires less than two weeks in H. nasutus, despite its low surface-area-specific rate of water loss, because of its small size and concomitantly high surface-to-volume ratio. The rate of urea accumulation during dehydration was 23 mM g-1 day-1, which is sufficiently low that urea accumulation would not be lethal before the frog had succumbed to dehydrational death. Consequently, there appears to be little or no selective advantage for uricotely in small 'waterproof' frogs, such as H. nasutus.

Optimal hematocrit theory during activity in the bullfrog (Rana catesbeiana).

1. There is an exponential relationship between blood viscosity (cP) and hematocrit (%) for the bullfrog; eta = 1.81 e0.033Hct. The in vitro optimal hematocrit calculated for blood flow through tubes, from this relationship for bullfrog blood, is 30%. 2. Amphibian blood is a non-Newtonian fluid with viscosity dependent on shear rate. It has a finite yield shear stress of about 1.5 dynes cm-2. 3. Hematocrit of bullfrogs was increased from 27% (control) to 57% by isovolemic erythrocythemia (constant volume blood-doping). There was a slight increase in systolic, diastolic and venous blood pressure with elevated hematocrit. 4. Systemic arch blood flow rate was inversely related to blood viscosity for erythrocythemic bullfrogs. The decrease in systemic arch blood flow at high hematocrits was due primarily to reduced pulse volume rather than reduced heart rate. 5. Systemic arch blood flow, when standardised between individuals, was inversely related to blood viscosity; Qbl = 0.185 + 3.73 eta -1. This relationship was significantly different from that predicted by the Poiseuille-Hagen flow formula. The in vivo optimal hematocrit calculated from this relationship was 41%. 6. Optimal hematocrit theory appears to be generally applicable for Rana catesbeiana in vitro and in vivo. Most individuals had an in vivo optimal hematocrit, but the absence of a clear optimal hematocrit for some individuals could reflect methodological variability, or in vivo physiological compensation for the increased blood viscosity at high hematocrit.

The effects of erythrocythemia on blood viscosity, maximal systemic oxygen transport capacity and maximal rates of oxygen consumption in an amphibian.

Graded erythrocythemia was induced by isovolemic loading of packed red blood cells in the toad, Bufo marinus. Blood viscosity, hematocrit, hemoglobin concentration, maximal aortic blood flow rate and maximal rates of oxygen consumption were determined after each load. Blood viscosity was related to hematocrit in the expected exponential manner; ln eta = 0.43 + 0.035 Hct. Maximal blood flow rates in the dorsal aorta were inversely proportional to blood viscosity and fit predictions of the Poiseuille-Hagen flow formula. The effect of increased blood viscosity was to reduce aortic pulse volume, but not maximal heart rate. Maximal systemic oxygen transport capacity (aortic blood flow rate X hemoglobin concentration X O2 binding capacity of hemoglobin) was linearly correlated with the maximal rate of oxygen consumption. These date indicate that optimal hematocrit theory is applicable for maximal blood flow rates in vivo, and that systemic oxygen transport is the primary limitation to aerial VO2 max in amphibians.