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PubMed1990

[Changes in body fluids of the frog Leptodactylus fuscus during estivation (Anura, Leptodactylidae)].

The frog, L. fuscus, becomes dormant during the dry season in southeastern Brazil. Plasma and urine were obtained and analyzed for K+, Na+, and osmotic concentrations in active and estivating frogs. Soil water potential from the estivation sites was compared with the osmotic concentrations of the frog. Plasma and urine osmotic concentrations (286.2 +/- 13.8 and 242.3 +/- 17.2 mOsm1(-1), respectively) were higher in the estivating than in active frogs (240.3 +/- 12.8 and 112.7 +/- 15.6 mOsm1(-1); plasma and urine), and the same holds true for plasma K+ content. The Na+ concentration was the same for active and estivating frogs. Soil water potential corresponded to osmotic pressure of 110 mOsm1(-1), showing that L. fuscus may uptake water from the soil during the estivation.

Animals↗
PubMedDate not supplied

Xanthine oxidase and xanthine dehydrogenase from an estivating land snail.

During arousal from estivation in land snails, Otala lactea, active metabolic functions are restored within minutes and oxygen consumption increases dramatically. During the transition from the hypoxic conditions of estivation to normoxia it is possible that xanthine oxidase (XO) in hepatopancreas contributes to the observed lipid peroxidation. Using a fluorometric assay that is based on the oxidation of pterin, the activities and some properties of XO and XO+XDH (sum of XO and xanthine dehydrogenase activities) were measured in hepatopancreas extracts. Km values for pterin for XO and XO+XDH were 9 and 6 microM, respectively, and the Km of XDH for methylene blue was 5 microM. Both XO+XDH and XO activities were inhibited by allopurinol (I50 = 2 microM), pre-incubation at 40 degrees C, and by 5 min H2O2 pre-exposure. Inclusion of azide in the reaction promoted a rise of approximately 70-fold in the inactivation power of H2O2 due to inhibition of high endogenous catalase activity. The I50 for H2O2 of XO+XDH and XO activities in the presence of azide was 0.04 and 0.11 mM, respectively. Unlike the situation for mammalian XO, a previous reduction of O. lactea XO (by pterin) was not necessary to make the enzyme susceptible to H2O2 effects. Interestingly, methylene blue partially prevented both heat- and H2O2-induced inactivation of XO+XDH activity. These data indicate that the formation of an enzyme-methylene blue complex induces protection against heat and oxidative damage at the FAD-active site. Both XO and XO+XDH activites were significantly higher in snails after 35 days of estivation compared with active snails 24 h after arousal from dormancy. The ratio of XO/(XO+XDH) activities was also slightly increased in estivating O. lactea (from 0.07 to 0.09; P < 0.025). XO activity was 0.03 nmol.min-1.mg protein-1 in estivating snails. Compared with hepatopancreas catalase, XO activity is probably too low to contribute significantly to the net generation of oxyradicals, and hence to peroxidative damage. Rather, the low potential of XO to induce oxidative stress may constitute an adaptive advantage for O. lactea during arousal periods.

Acclimatization↗
PubMed2000

Intrinsic metabolic depression in cells isolated from the hepatopancreas of estivating snails.

Many animals across the phylogenetic scale are routinely capable of depressing their metabolic rate to 5-15% of that at rest, remaining in this state sometimes for years. However, despite its widespread occurrence, the biochemical processes associated with metabolic depression remain obscure. We demonstrate here the development of an isolated cell model for the study of metabolic depression. The isolated cells from the hepatopancreas (digestive gland) of the land snail (Helix aspersa) are oxygen conformers; i.e., their rate of respiration depends on pO(2). Cells isolated from estivating snails show a stable metabolic depression to 30% of control (despite the long and invasive process of cell isolation) when metabolic rate at the physiological pH and pO(2) of the hemolymph of estivating snails is compared with metabolic rate at the physiological pH and pO(2) of the hemolymph of control snails. When the extrinsic effects of pH and pO(2) are excluded, the intrinsic metabolic depression of the cells from estivating snails is still to below 50% of control snails. The in vitro effect of pO(2) on metabolic rate is independent of pH and state (awake or estivating), but the effects of pH and state significantly interact. This suggests that pH and state change affect metabolic depression by similar mechanisms but that the metabolic depression by hypoxia involves a separate mechanism.

Animals↗
PubMed2005

Lessons from an estivating frog: sparing muscle protein despite starvation and disuse.

Long (6- to 9-mo) bouts of estivation in green-striped burrowing frogs lead to 28% atrophy of cruralis oxidative fibers (P < 0.05) and some impairment of in vitro gastrocnemius endurance (P < 0.05) but no significant deficit in maximal twitch force production. These data suggest the preferential atrophy of oxidative fibers at a rate slower than, but comparable to, laboratory disuse models. We tested the hypothesis that the frog limits atrophy by modulating oxidative stress. We assayed various proteins at the transcript level and verified these results for antioxidant enzymes at the biochemical level. Transcript data for NADH ubiquinone oxidoreductase subunit 1 (71% downregulated, P < 0.05) and ATP synthase (67% downregulated, P < 0.05) are consistent with mitochondrial quiescence and reduced oxidant production. Meanwhile, uncoupling protein type 2 transcription (P = 0.31), which is thought to reduce mitochondrial leakage of reactive oxygen species, was maintained. Total antioxidant defense of water-soluble (22.3 +/- 1.7 and 23.8 +/- 1.5 microM/microg total protein in control and estivator, respectively, P = 0.53) and membrane-bound proteins (31.5 +/- 1.9 and 42.1 +/- 7.3 microM/microg total protein in control and estivator, respectively, P = 0.18) was maintained, equivalent to a bolstering of defense relative to oxygen insult. This probably decelerates muscle atrophy by preventing accumulation of oxidative damage in static protein reserves. Transcripts of the mitochondrially encoded antioxidant superoxide dismutase type 2 (67% downregulated, P < 0.05) paralleled mitochondrial activity, whereas nuclear-encoded catalase and glutathione peroxidase were maintained at control values (P = 0.42 and P = 0.231), suggesting a dissonance between mitochondrial and nuclear antioxidant expression. Pyruvate dehydrogenase kinase 4 transcription was fourfold lower in estivators (P = 0.11), implying that, in contrast to mammalian hibernators, this enzyme does not drive the combustion of lipids that helps spare hypometabolic muscle.

Animals↗
PubMed1977

Oxygen-binding properties of hemoglobins from estivating and active African lungfish.

The oxygen-binding characteristics and the multiplicity of the stripped hemoglobiin from active lungfish Protopterus amphibius, are the same as in specimens that have been estivating for about 30 months, showing that alteration in the hemoglobin molecules is not involved in the earlier reported increase in oxygen affinity of whole blood during estivation (Johansen et al., '76). At pH 7.0 and 26 degrees C the hemolysates show a high oxygen affinity (P50 = 3.1 Torr), a Bohr factor (delta log P50/delta pH) of - 0.33, and a cooperativity coefficient (n) of 1.7. Between 15 and 26 degrees C, the apparent heat of oxygenation (delta H) is - 8.6 Kcal-mole-1 at pH 7.0, corresponding with data for other fish. A low sensitivity of oxygen affinity to urea appears to be adaptive to the high urea concentrations in estivating lungfish. The salt sensitivity is, however, similar to human hemoglobin. The hemoglobin consists of two major (electrophoretically anodal) components, which differ slightly in oxygen affinity but are both sensitive to pH and nucleoside triphosphates (NTP). Guanosine triphosphate (GTP), the major erythrocytic organic phosphate, however, depresses the oxygen affinity of the composite and separated hemoglobins more effectively than ATP suggesting that GTP is the primary modulator of oxygen affinity. Comparative measurements reveal only one major hemoglobin component in P. annectens which has a markedly lower oxygen affinity and phosphate sensitivity than P. amphibius hemoglobins and thus seems less pliable to phosphate-mediated variation in oxygen affinity. The data are discussed in relation to the hemoglobin systems of other fish.

Adaptation, Physiological↗
PubMedDate not supplied

Physiological and behavioral variation in estivation among mud turtles (Kinosternon spp.).

Kinosternid mud turtles, a primarily aquatic group, exhibit variable degrees of terrestrial activity in the Sonoran and Chihuahuan Deserts. We compared behavioral and physiological responses to dry conditions in four populations representing three species, Kinosternon sonoriense, Kinosternon flavescens, and Kinosternon hirtipes. All four groups were subjected to simulated dry season conditions in the laboratory, during which activity was monitored and physiological responses (blood chemistry and rates of resting metabolism and evaporative water loss) were measured. Kinosternon flavescens and K. hirtipes represented extremes in apparent ability to estivate, based on activity and rate of increase of plasma osmolality. Two populations of K. sonoriense exhibited intraspecific differences in behavioral and physiological measures that were related to extant environmental conditions. Large numbers of K. sonoriense from Arizona and K. hirtipes, the poorest estivators, had to be rehydrated after only 30 d out of water. Kinosteron flavescens had the lowest metabolic rates, but no evidence of metabolic depression during dehydration was found for any of the four populations. We conclude that the differences in capacity for estivation among populations are primarily linked to variable behavioral responses to dry conditions, though high rates of evaporative water loss in K. hirtipes represent a probable physiological constraint.

Adaptation, Physiological↗
PubMed1996

Effectors of metabolic depression in an estivating pulmonate snail (Helix aspersa): whole animal and in vitro tissue studies.

We have examined metabolic depression in the land snail (Helix aspersa) during estivation, and have developed a tissue model of metabolic depression using an in vitro mantle preparation. The metabolic rate of H. aspersa is depressed by 84% in vivo within 4 weeks of onset of estivation, and this metabolic depression is accompanied by a decrease in haemolymph PO2 and pH, and an increase in haemolymph PCO2. The in vitro mantle preparation has a stable O2 consumption and energy charge, and an energy charge similar to that of mantle in vivo. The in vitro mantle is an O2-conforming tissue, with VO2 varying curvilinearly with PO2. Consequently, we have developed a mathematical method of calculating tissue VO2 at any PO2. These calculations show that under appropriate incubation conditions of pH and PO2, the mantle from estivating animals shows a stable in vitro metabolic depression of 48% compared to mantle from control snails. The extrinsic effects of PO2 and pH account for 70% of the total in vitro metabolic depression of mantle tissue; intrinsic effectors contribute a further 30%.

Adenosine Diphosphate↗
PubMed2000

Comparison of hibernation, estivation and daily torpor in the edible dormouse, Glis glis.

Three major forms of dormancy in mammals have been classified: hibernation in endotherms is characterised by reduced metabolic rate (MR) and body temperature (Tb) near ambient temperature (Ta) over prolonged times in the winter. Estivation is a similar form of dormancy in a dry and hot environment during summertime. Daily torpor is defined as reduced MR and Tb lower than 32 degrees C, limited to a duration of less than 24 h. The edible dormouse (Glis glis) is capable for all three distinct forms of dormancy. During periods of food restriction and/or low Ta, daily torpor is displayed throughout the year, alternating with hibernation and estivation in winter and summer respectively. We recorded Tb, O2-consumption and CO2-production in unrestrained dormice at different Ta's for periods of up to several months. Cooling rate and rate of metabolic depression during entrance into the torpid state was identical in all three forms of dormancy. The same was true for thermal conductance, maximum heat production, duration of arousal and cost of an arousal. The only difference between hibernation and daily torpor was found in the bout duration. A daily torpor bout lasted 3-21 h, a hibernation bout 39-768 h. As a consequence of prolonged duration, MR, Tb and also the Tb - Ta gradient decreased to lower values during hibernation bouts when compared to daily torpor bouts. Our findings suggest that all three forms of dormancy are based on the same physiological mechanism of thermal and metabolic regulation.

Animals↗
PubMed1990

Metabolic rate depression and biochemical adaptation in anaerobiosis, hibernation and estivation.

For many animals, the best defense against harsh environmental conditions is an escape to a hypometabolic or dormant state. Facultative metabolic rate depression is the common adaptive strategy of anaerobiosis, hibernation, and estivation, as well as a number of other arrested states. By reducing metabolic rate by a factor ranging from 5 to 100 fold or more, animals gain a comparable extension of survival time that can support months or even years of dormancy. The present review focuses on the molecular control mechanisms that regulate and coordinate cellular metabolism for the transition into dormancy. These include reversible control over the activity state of enzymes via protein phosphorylation or dephosphorylation reactions, pathway regulation via the association or dissociation of particle-bound enzyme complexes, and fructose-2,6-bisphosphate regulation of the use of carbohydrate reserves for biosynthetic purposes. These mechanisms, their interactions, and the regulatory signals (e.g., second messenger molecules, pH) that coordinate them form a common molecular basis for metabolic depression in anoxia-tolerant vertebrates (goldfish, turtles) and invertebrates (marine molluscs), hibernation in small mammals, and estivation in land snails and terrestrial toads.

Adaptation, Physiological↗
PubMed2002

In vivo downregulation of protein synthesis in the snail Helix apersa during estivation.

Protein synthesis is downregulated during metabolic depression in a number of systems where the metabolic depression is effected by obvious extrinsic cues. The metabolic depression of the estivating land snail Helix apersa occurs in the absence of any obvious physiological stress and has an intrinsic component independent of temperature, pH, O(2) status, or osmolality. We show that this metabolic depression is accompanied by a downregulation of protein synthesis in vivo. The rate of protein synthesis decreases in two major tissues during estivation: to 23% and 53% of the awake rate in hepatopancreas and foot muscle, respectively. We show from calculations of the theoretical contribution of protein synthesis to total O(2) consumption that the depression of protein synthesis must be a significant, obligate, in vivo component of metabolic depression in H. aspersa.

Animals↗
PubMedDate not supplied

Estivation in South American amphibians and reptiles.

A number of amphibians and reptiles have cyclic behavior, becoming inactive with the coming of the dry season. In South America this pattern of activity is common, particularly in savannah-like vegetation. During the dry season amphibians burrow into the mud or soil, and either form a cocoon or increase the osmotic concentration of body fluids to reduce evaporative water loss. Some phyllomedusid tree frogs coat their body surface with skin secretion and excrete uric acid to minimize water loss. Reptiles also retreat into shelter deep enough to avoid temperature fluctuation during estivation or reduce metabolic response to temperature. Reduction of temperature sensitivity of the metabolism seems to be a strategy common to estivating amphibians and reptiles. Despite seasonal change of the environment, some species of reptiles are active all year round.

Amphibians↗
PubMed1997

Glycolytic controls in estivation and anoxia: a comparison of metabolic arrest in land and marine molluscs.

Facultative metabolic rate depression is the common adaptive strategy underlying various animal mechanisms for surviving harsh environmental conditions. This strategy is common among molluscs, enabling animals to survive over days or even months in the absence of oxygen or under extremely dry conditions. The large reductions in metabolic rate during estivation and anoxia can translate into considerable energy savings when dormant animals are compared to active animals. A complex metabolic coordination is required during the transition into the dormant state to maintain cellular homeostasis and involves both energy-consuming and energy-producing pathways. With regard to energy-producing pathways, several different mechanisms have been identified that participate in controlling flux. One such mechanism, enzyme phosphorylation, can have a wide-ranging effect. For example, phosphorylated enzymes exhibit altered substrate, activator, and inhibitor affinities. This effect may be magnified by changes in the concentrations of allosteric effectors, such as fructose 2,6-bisphosphate, that occur during hypometabolic states. Changes in fructose 2,6-bisphosphate are related to changes in enzyme phosphorylation through changes in the relative activity of phosphofructokinase-2. Alterations in glycolytic enzyme binding can also be brought about through changes in enzyme phosphorylation. The present review focuses on identifying hypometabolism-related changes in enzyme phosphorylation as well as characterizing the mechanisms involved in mediating these phosphorylation events.

Animals↗
PubMed2006

The potential role of CO2 in initiation and maintenance of estivation in the land snail Helix lucorum.

Elevated CO(2) levels are hypothesized to play a role in the initiation and maintenance of estivation in snails through disturbances of acid-base status. The aim of our study was to identify the ambient CO(2) threshold that induces disturbances in acid-base status in the air-breathing land snail Helix lucorum. Acid-base parameters were determined in the hemolymph of snails acclimated to 0.5%, 1%, 2%, 4%, and 8% CO(2) in air for 20 d. In addition, we evaluated the effects of long-term acclimation on metabolic rate and on levels of D-lactate dehydrogenase activity (D-LDH) and of D-lactate in snails after 20 d of exposure to increased CO(2) levels. Helix lucorum proved to be unable to compensate for a decrease in extracellular pH (pH(e)) when acclimated to levels higher than 1% CO(2) in air. The rate of oxygen consumption started to decrease when snails were acclimated to 0.5% CO(2) in air. However, there was no correlation between the drops in pH(e) and in metabolic rate. Long-term acclimation to elevated CO(2) levels induced an increase in the activity of D-LDH with a concomitant accumulation of D-lactate in tissues. This indicates that long-term acclimation to elevated ambient CO(2) levels could reduce the aerobic capacity of land snails and trigger expression of anaerobic pathways of ATP turnover. The threshold levels of ambient CO(2) that induce changes in acid-base status and elicit metabolic depression in adult land snails H. lucorum are higher than the future atmospheric levels that are expected to result from human use of fossil energy resources.

Acclimatization↗
PubMed2002

Urea and KCl have differential effects on enzyme activities in liver and muscle of estivating versus nonestivating species.

The effects of 300 mM urea or 300 mM KCl on the maximal activities of 25 enzymes of intermediary metabolism were assessed in extracts of liver and muscle from spadefoot toads (Scaphiopus couchii), leopard frogs (Rana pipiens), and rats to assess their sensitivity to these osmolytes. During estivation, toads can lose -50% of total body water, and urea, which is known for its action as a protein denaturant, accumulates to 200-300 mM. The data show that the maximal activities of toad liver enzymes were not affected when assayed in the presence of 300 mM urea in vitro whereas urea inhibited the activities of seven enzymes in frog and 11 enzymes in rat liver. High KCl affected 12 or 13 enzymes in liver of each species causing inhibition in eight or nine cases each, and for frog and rat enzymes, inhibition was frequently more pronounced than for urea. Both urea and KCl affected enzyme activities in muscle extracts of all three species, but whereas their effects were largely negative for frog and rat enzymes, the enzymes affected by urea or KCl in toad muscle were primarily activated by these osmolytes (six out of nine enzymes affected by urea and eight of 15 enzymes affected by KCl). Urea, KCl, and polyethylene glycol (a protein crowding agent) also had species-specific effects on the dissociation constant (Ka) for cAMP of protein kinase A. The data suggest that the accumulation of urea by water-stressed anurans not only contributes to minimizing cell volume reduction, but by doing so also limits the increase in intracellular ionic strength that occurs and thereby helps to minimize the potential inhibitory effects of high salts on metabolic enzymes.

Animals↗
PubMed2005

ESTIV questionnaire on the acquisition and use of primary human cells and tissue in toxicology.

The ability to use human cells and tissues in toxicology research and testing has the benefit that it obviates the need to undertake species extrapolation when assessing human hazard. However, obtaining and using human cells and tissues is logistically difficult, ethically complex and is a potential source of infections to those coming into contact with human cell material. The issue is also controversial, with the recent EU legislation draft on tissue engineering, and also due to some instances of human material being obtained and used without informed consent. There are also varying regulations and attitudes relating to the use of human cells and tissues throughout Member States of the EU, and there is a need for harmonisation. The European Society of Toxicology in Vitro (ESTIV) Executive Board and the European Network of Human Research Tissue Banks (ENRTB) have conducted a survey to ascertain the extent to which human cells and tissues are used by its members, how these are obtained, what local regulations are in force, how the material is used, and the advantages and disadvantages experienced by members in using such material, as opposed to cell lines. The results obtained have been compared with the results from a previous survey conducted in 2000. It is hoped that this information will help to facilitate the process of acquiring and using human cells and tissues in a safe and effective way to promote the use of non-animal approaches for investigating the mechanisms of toxicity, and for predicting the toxic hazard of substances.

Animal Testing Alternatives↗
PubMed1979

Sleep and estivation (shallow torpor): continuous processes of energy conservation.

Estivation (shallow torpor) in the round-tailed ground squirrel (Citellus tereticaudus) is entered through electrophysiologically defined states of sleep. Rapid-eye-movement sleep diminishes as body temperature falls in such a way that, at a body temperature of 26 degrees to 28 degrees C, torpor is characterized by almost continuous slow-wave sleep isomorphic with that observed at euthermic body temperatures.

Animals↗