[Histophysiological study of central monoamines in the garden dormouse (Eliomys quercinus) in estival and prehibernating phases].
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Animals↗
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[Palliative treatment of estival rhinitis in children. A double blind combined study of an antiamine, pimethixen, and an antihistamine, Paradryl].
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Adolescent↗
[Decontamination and estivation of ponds in the control of khawiasis in carp].
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Animals↗
[Effect of rest on functional state of the cardiovascular system and on physical endurance in estival conditions].
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Adult↗
Adaptations of the reed frog Hyperolius viridiflavus (Amphibia, Anura, Hyperoliidae) to its arid environment. VII. The heat budget of Hyperolius viridiflavus nitidulus and the evolution of an optimized body shape.
Estivating reed frogs of the superspecies Hyperolius viridiflavus are extraordinarily resistant to the highly adverse climatic conditions prevailing in their African savanna habitats during dry season (air temperature up to 45 degrees C, solar radiation load up to 1000 W.m-2, no water replenishment possible for up to 3 months). They are able to withstand such climatic stress at their exposed estivation sites on dry plants without evaporative cooling. We developed a heat budget model to understand the mechanisms of how an anuran can achieve this unique tolerance, and which allows us to predict the anuran's core and surface temperature for a given set of environmental parameters, to within 4% of the measured values. The model makes it possible to quantify some of the adaptive mechanisms for survival in semiarid habitats by comparing H. viridiflavus with anurans (H. tuberilinguis and Rana pipiens) of less stressful habitats. To minimize heat gain and maximize heat loss from the frog, the following points were important with regard to avoiding lethal heat stress during estivation: 1) solar heat load is reduced by an extraordinarily high skin reflectivity for solar radiation of up to 0.65 under laboratory and even higher in the field under dry season conditions. 2) The half-cylindrical body shape of H. viridiflavus seems to be optimized for estivation compared to the hemispheroidal shape usually found for anurans in moist habitats. A half-cylinder can be positioned relative to the sun so that large surface areas for conductive and convective heat loss are shielded by a small area exposed to direct solar radiation. 3) Another important contribution of body shape is a high body surface area to body mass ratio, as found in the estivating subadult H. viridiflavus (snout-vent lengths of 14-20 mm and body weights of 350-750 mg) compared to adult frogs (24-30 mm, 1000-2500 mg) which have never been observed to survive a dry season. 4) These mechanisms strongly couple core temperature to air temperature. The time constant of the core temperature is 29 +/- 10 s. Since air temperature can be 43-45 degrees C, H. viridiflavus must have a very unusual tolerance to transient core temperatures of 43-45 degrees C. 5) If air temperature rises above this lethal limit, the estivating frog would die despite all its optimizations, but moving from an unsuited to a more favorable site during estivation can be extremely costly in terms of unavoidably high evaporative water loss. Therefore, H. viridiflavus must have developed behavioral strategies for reliably choosing estivation sites with air temperature staying on average within the vital range during the whole dry season.
Acclimatization↗
Hypometabolism, antioxidant defenses and free radical metabolism in the pulmonate land snail Helix aspersa.
The aim of this work was to evaluate the effect of a cycle of estivation and awakening on free radical metabolism in selected organs of the land snail Helix aspersa. Estivation for 20 days induced a 4.9- and 1.8-fold increase in selenium-dependent glutathione peroxidase activity (Se-GPX) and in total glutathione levels (GSH-eq), respectively, in hepatopancreas when compared to activity in active animals 24 h after awakening. Foot muscle Se-GPX activity was also increased 3.9-fold during estivation, whereas GSH-eq did not vary. The activities of other antioxidant enzymes (catalase, superoxide dismutase, glutathione reductase and glutathione S-transferase) and glucose 6-phosphate dehydrogenase were unchanged in both organs. After 15 min of awakening, the glutathione disulphide (GSSG)/GSH-eq ratio increased significantly by 55% in hepatopancreas, slowly returning to the levels observed during estivation. The higher GSSG/GSH-eq ratio may be caused by increased formation of reactive oxygen species (ROS) during awakening. The levels of thiobarbituric acid reactive substances (TBARS) decreased from 49 to 30.7 nmol g(-1) wet mass in hepatopancreas after 5 min arousal and, after 30 min, TBARS rose significantly to 39.6 nmol g(-1) wet mass, gradually declining thereafter. The levels of lipid hydroperoxides in hepatopancreas and of carbonyl protein in foot muscle both decreased during awakening. The higher levels of products of free radical damage during estivation may have resulted from low levels of ROS formation associated with decreased rates of lipid hydroperoxide detoxification and oxidized protein turnover caused by metabolic depression. The regulation of the antioxidant system during hypometabolism may constitute a mechanism to minimize oxidative stress during cycles of estivation and awakening.
Animals↗
Glucose-6-phosphate dehydrogenase regulation during hypometabolism.
Glucose-6-phosphate dehydrogenase (G6PDH) from hepatopancreas of the land snail, Otala lactea, shows distinct changes in properties between active and estivating (dormant) states, providing the first evidence of pentose phosphate cycle regulation during hypometabolism. Compared with active snails, G6PDH Vmax increased by 50%, Km for glucose-6-phosphate decreased by 50%, Ka Mg x citrate decreased by 35%, and activation energy (from Arrhenius plots) decreased by 35% during estivation. DEAE-Sephadex chromatography separated two peaks of activity and in vitro incubations stimulating protein kinases or phosphatases showed that peak I (low phosphate) G6PDH was higher in active snails (57% of activity) whereas peak II (high phosphate) G6PDH dominated during estivation (71% of total). Kinetic properties of peaks I and II forms mirrored the enzyme from active and estivated states, respectively. Peak II G6PDH also showed reduced sensitivity to urea inhibition of activity and greater stability to thermolysin protease treatment. The interconversion of G6PDH between active and estivating forms was linked to protein kinase G and protein phosphatase 1. Estivation-induced phosphorylation of G6PDH may enhance relative carbon flow through the pentose phosphate cycle, compared with glycolysis, to help maintain NADPH production for use in antioxidant defense.
Animals↗
Antioxidant defenses and metabolic depression. The hypothesis of preparation for oxidative stress in land snails.
The roles of enzymatic antioxidant defenses in the natural tolerance of environmental stresses that impose changes in oxygen availability and oxygen consumption on animals is discussed with a particular focus on the biochemistry of estivation and metabolic depression in pulmonate land snails. Despite reduced oxygen consumption and PO2 during estivation, which should also mean reduced production of oxyradicals, the activities of antioxidant enzymes, such as superoxide dismutase and catalase, increased in 30 day-estivating snails. This appears to be an adaptation that allows the snails to deal with oxidative stress that takes place during arousal when PO2 and oxygen consumption rise rapidly. Indeed, oxidative stress was indicated by increased levels of lipid peroxidation damage products accumulating in hepatopancreas within minutes after arousal was initiated. The various metabolic sites responsible for free radical generation during arousal are still unknown but it seems unlikely that the enzyme xanthine oxidase plays any substantial role in this despite being implicated in oxidative stress in mammalian models of ischemia/reperfusion. We propose that the activation of antioxidant defenses in the organs of Otala lactea during estivation is a preparative mechanism against oxidative stress during arousal. Increased activities of antioxidant enzymes have also observed under other stress situations in which the actual production of oxyradicals should decrease. For example, antioxidant defenses are enhanced during anoxia exposure in garter snakes Thamnophis sirtalis parietalis (10 h at 5 degrees C) and leopard frogs Rana pipiens (30 h at 5 degrees C) and during freezing exposure (an ischemic condition due to plasma freezing) in T. sirtalis parietalis and wood frogs Rana sylvatica. It seems that enhancement of antioxidant enzymes during either anoxia or freezing is used as a preparatory mechanism to deal with a physiological oxidative stress that occurs rapidly within the early minutes of recovery during reoxygenation or thawing. Thus, a wide range of stress tolerant animals display coordinated changes in antioxidant defenses that allow them to deal with oxidative stress that occurs as part of natural cycles of stress/recovery that alter oxygen levels in tissues. The molecular mechanisms that trigger and regulate changes in antioxidant enzyme activities in these species are still unknown but could prove to have key relevance for the development of new intervention strategies in the treatment of cardiovascular ischemia/reperfusion injuries in humans.
Animals↗
Primary causes of decreased mitochondrial oxygen consumption during metabolic depression in snail cells.
Cells isolated from the hepatopancreas of estivating snails (Helix aspersa) have strongly depressed mitochondrial respiration compared with controls. Mitochondrial respiration was divided into substrate oxidation (which produces the mitochondrial membrane potential) and ATP turnover and proton leak (which consume it). The activity of substrate oxidation (and probably ATP turnover) decreased, whereas the activity of proton leak remained constant in estivation. These primary changes resulted in a lower mitochondrial membrane potential in hepatopancreas cells from estivating compared with active snails, leading to secondary decreases in respiration to drive ATP turnover and proton leak. The respiration to drive ATP turnover and proton leak decreased in proportion to the overall decrease in mitochondrial respiration, so that the amount of ATP turned over per O2 consumed remained relatively constant and aerobic efficiency was maintained in this hypometabolic state. At least 75% of the total response of mitochondrial respiration to estivation was caused by primary changes in the kinetics of substrate oxidation, with only 25% or less of the response occurring through primary effects on ATP turnover.
Adenosine Triphosphate↗
Antioxidant defenses and metabolic depression in a pulmonate land snail.
During arousal from estivation oxygen consumption by land snails (Otala lactea) increases severalfold. To determine whether snails prepared for an accompanying rise in the rates of oxyradical generation by altering their antioxidant defense mechanisms, changes in the activities of antioxidant enzymes and lipid peroxidation products were quantified in foot and hepatopancreas of control, 30-day estivating, and aroused snails. Compared with controls, estivating O. lactea showed significant increases in the activities of foot muscle superoxide dismutase (SOD) (increasing by 56-67%), catalase (51-72%), and glutathione S-transferase (79-108%), whereas, in hepatopancreas, SOD (57-78%) and glutathione peroxidase (93-144%) increased. Within 40 min after arousal began, hepatopancreas glutathione peroxidase activity had returned to control values, but SOD showed a further 70% increase in activity but then returned to control levels by 80 min. Estivation had no effect on total glutathione (GSH + 2 GSSG) concentrations in tissues, but GSSG content had increased about twofold in both organs of 30-day dormant snails. Lipid peoxidation (quantified as thiobarbituric acid reactive substances) was significantly enhanced at the onset of arousal from dormancy, indicating that oxidative stress and tissue damage occurred at this time. The data suggest that antioxidant defenses in snail organs are increased while snails are in the hypometabolic state as a preparation for oxidative stress during arousal.
Animals↗
Sprouting by intact Helisoma neurons: role for glutamate.
We examined the possible relationship between sprouting by intact neurons and the elevation of blood acidic amino acids that is reported to occur in Helisoma in response to osmotic stress. In the present study, buccal ganglion neurons 5 were examined following exposure of animals to conditions that induce estivation, a behavioral state exhibited by these freshwater snails in nature. Surprisingly, sprouting activity occurred in some neurons (31%) in normal animals, this being transiently elevated (to 71%) after 4 days of estivation. Blood acidic amino acids (i.e., glutamate and aspartate) were also elevated under this condition, their levels reaching a plateau on day 4. These observations give rise to two hypotheses: 1) the acidic amino acids can induce sprouting, or 2) these agents can release trophic factors, which, in turn, induce sprouting. The first hypothesis was tested by exposing semi-intact preparations in vitro to a range of concentrations of the acidic amino acids; no promotion of sprouting was observed by either glutamate or aspartate. The second hypothesis was supported, however, by the following observations. First, blood from estivated animals was tested for trophic activity. Such blood (but not that from control animals) could promote sprouting by intact neurons in vitro, a response that depended on factor(s) whose filtration behavior on Amicon membranes suggests an approximate molecular weight of 30-100 kD. Second, since medium conditioned by Helisoma ganglia is known to contain a protein(s) that is necessary for sprouting by isolated neurons, we tested the ability of such medium to promote sprouting by intact neurons. Conditioned medium promoted sprouting by intact neurons in vitro in a manner quantitatively and qualitatively similar to that observed in vivo during estivation. Additionally, sprouting by intact neurons in conditioned medium was also attributable to factor(s) that acted as 30-100 kD molecules on Amicon membranes. To test the ability of the acidic amino acids to release or activate these factors, medium was conditioned for short periods in the presence of glutamate or aspartate; glutamate, but not aspartate, was found to enhance conditioned medium. We conclude that glutamate can promote sprouting by intact neurons, but that its action is indirect and is mediated by macromolecular trophic factors.
Animals↗
Microanatomy and ultrastructure of the kidney of the African lungfish Protopterus dolloi.
The Dipnoi (lungfishes) have developed true lungs, having the ability to take oxygen from both the gills and the lungs. During the tropical dry season, many lungfish estivate on land, breathing only air. The estivation period is accompanied by profound functional modifications, including the suppression of urine. Thus, the lungfish kidney must be designed to cope with these dramatic cyclic changes in renal function. We study here the microanatomy and the structure of the kidney of the African lungfish Protopterus dolloi, maintained under controlled freshwater conditions. Chemical microdissection, light microscopy, and scanning and transmission electron microscopy have been used. The nephrons of P. dolloi are composed of a renal corpuscle (RC) and of a renal tubule that appears divided into five morphologically distinct segments: neck segment (NS), proximal tubule (PT), intermediate segment (IS), distal tubule (DT), and collecting tubule (CT). Paired CTs abut into a collecting duct, the latter emptying into an archinephric duct. The RCs lie in the mid-zone of the kidney, between the PTs and the convoluted DTs. The spatial distribution of these elements allows recognition of a kidney zonation. The RCs group into clusters (3-4 RCs per cluster) that are supplied by a single arteriole surrounded by pericytes. Each cluster appears to represent a functional unit with a common hemodynamic regulatory mechanism. The major processes of the podocytes form flattened networks that appear to constitute an integrated system due to the presence of gap junctions. The existence of mesangial cells with large cell processes, and of mesangial cells with a dendritic appearance, suggests a complex functional role (contractile and phagocytic) for the mesangium. The NS and the IS are the narrowest nephron segments, formed only by multiciliated cells. The PT and the DT can be subdivided, based on the tubular morphology and on cell composition, into portions I and II: PTI is formed only by brush border (BB) cells, while PTII contains BB and multiciliated cells. The DTI is formed by segment-specific cells, while the DTII contains segment-specific and a small number of flask cells. The CT contains principal and flask cells in a 5:1 ratio. The flask cells adopt two different configurations (with a narrow canaliculus or with a large cavity). The main goal of this study was to disclose specific kidney features that could be related to function, phylogeny, and habitat. In addition, the present results constitute the basis for a study of the morphologic changes that should occur in the kidney of P. dolloi during estivation.
Animals↗
The muscle fatty acid binding protein of spadefoot toad (Scaphiopus couchii).
Fatty acid binding protein was purified from skeletal muscle of the spadefoot toad (Scaphiopus couchii), an estivating species. While estivating, this animal relies on the fatty acid oxidation for energy. Hence we were interested in the behaviour of fatty acid binding protein under conditions of elevated urea (up to 200 mM) and potassium chloride such as exist during estivation. Also we examined whether there were interactions between glycolytic intermediates and the binding ability of the protein. The amount of bound fatty acid (a fluorescence assay using cis-parinarate) was not affected (P < 0.05) by glucose, fructose 6-phosphate or phosphoenolpyruvate at physiological concentrations. By contrast, glucose 6-phosphate increased the amount of bound cis-parinarate but the apparent dissociation constant was not different from the control. Fructose 1,6-bisphosphate but not fructose 2,6-phosphate decreased cis-parinarate binding by 40%, commensurate with doubling the apparent dissociation constant (1.15-2.62 microM). Urea, guanidinium and trimethylamine N-oxide at 200 mM increased cis-parinarate binding 60% over controls. Urea (1 M) and KCl (200 mM) did not affect cis-parinarate binding compared to controls. The interaction of this fatty acid transporter with fructose 1,6-bisphosphate is discussed in terms of reciprocal interaction with phosphofructokinase since fatty acid is also an inhibitor of phosphofructokinase.
Animals↗
Induction and inhibition of diapause by the same photoperiod: experimental evidence for a "double circadian oscillator clock".
On the southern Iberian Peninsula, the seasonal life history of the large white butterfly, Pieris brassicae, comprises 2 different photoperiodically induced developmental arrests: a hibernation diapause at photophases < 11 h and an estivation diapause at photophases > 14 h. At intermediate photophases (12 h to 13 h), the butterfly responds with a nondiapause. Combined with the experimental setup to determine photosensitivity in insects, the different photoperiodic responses at long-, intermediate-, and short-night conditions were examined to gain more insight into the time measurement mechanism in P. brassicae. The study reveals evidence for a "double circadian oscillator clock" mechanism that is based on 2 submechanisms, a "short-night determining system" and a separate "long-night determining system." This conclusion was drawn from the facts that an LD 9:15 long-night induces a hibernation diapause but inhibits an estivation diapause and, conversely, that an LD 16:8 short-night inhibits a hibernation diapause but induces an estivation diapause. This opposite effect of the same photoperiod supports the argument for the existence of 2 independent targets for light-dark cycles, interpreted as 2 antagonistic time measurement systems. The existence and independence of 2 systems was further shown by differences in long-night versus short-night responses regarding photosensitivity, temperature dependence, and heritable factors. The long-night measurement system is most effective in the 5th larval stage, is highly affected by temperature, and is easy to manipulate by selective inbreeding. The short-night measurement system is most effective in the 4th larval stage, is largely temperature compensated, and is not affected by experimental manipulation of the longnight measurement system.
Animals↗
Sublethal soil copper concentrations increase mortality in the earthworm Aporrectodea caliginosa during drought.
Simultaneous effects of drought and a sublethal concentration of copper on the earthworm Aporrectodea caliginosa were investigated in the laboratory. Experiments were performed with exposure to 150 mg Cu kg(-1) dry soil and different levels of drought during a 3-wk period. Soil water potentials from pF 1.5 (wet) to pF 5 (very dry) were obtained. The soil water potential resulting in 50% mortality was significantly reduced from pF 4.48 in worms not exposed to copper to pF 4.09 in copper-exposed worms, thus demonstrating synergy-like effects of drought and copper. Development of estivation cells was significantly depressed in copper-exposed worms compared to worms not exposed to copper. For all drought levels (except the highest, where 100% mortality occurred), copper-exposed earthworms with no estivation cells had more than twice as high mortality and, at severe drought exposure, also lower water content than those which had developed the estivation cells. With increasing drought level, the whole-body burden of copper increased from about 40 microg Cu g(-1) dry weight to about 90 microg Cu g(-1). When the worms were exposed to drought, the osmolality in their body fluids increased. This was due to a greater concentration of already existing solutes resulting from extensive loss of water, and not to mobilization or synthesis of additional solutes.
Animals↗
In vitro nitric oxide effects on basal and gonadotropin-releasing hormone-induced gonadotropin secretion by pituitary gland of male crested newt (Triturus carnifex) during the annual reproductive cycle.
The objective of this study was to test the possible nitric oxide (NO) involvement in pituitary gonadotropin secretion in the male crested newt, Triturus carnifex. Pituitaries were incubated in vitro with medium alone, GnRH, NO donor (NOd, sodium nitroprusside), NO synthase inhibitor (NOSi, Nomega-nitro-L-arginine methyl ester), cGMP analogue (cGMPa, 8-bromo-cGMP), soluble guanylate cyclase inhibitor (sGCi, cystamine), GnRH plus NOSi, GnRH plus sGCi, and NOd plus sGCi during the annual reproductive cycle: pre-reproduction, reproduction (noncourtship and courtship), and the refractory, recovery, and estivation periods. To determine pituitary gonadotropin secretion indirectly, newt testes were superfused in vitro with preincubated pituitaries, and androgen release was determined. NO synthase (NOS) activity and cGMP levels were assessed in the preincubated pituitaries. Medium alone- and GnRH-preincubated pituitary increased androgen secretion during pre-reproduction, noncourtship, courtship, and recovery; the GnRH-induced increase was higher than the medium alone-induced increase during pre-reproduction, noncourtship, and recovery. NOd and cGMPa increased androgens in all reproductive phases considered except courtship; the NOd- and cGMP-induced increase was higher than the medium alone-induced increase during pre-reproduction, noncourtship, and recovery. NOS activity was highest during courtship and lowest during the refractory and estivation periods. GnRH increased NOS activity during pre-reproduction, noncourtship, and recovery. Cyclic GMP levels were highest during courtship and lowest during the refractory period and estivation. GnRH increased cGMP levels during pre-reproduction, noncourtship, and recovery, while NOd did so during all reproductive phases considered. These results suggest that basal and GnRH-induced gonadotropin secretion are up-regulated by NO in the pituitary gland of the male Triturus carnifex.
Androgens↗
Biomarker response and biomass change of earthworms exposed to chlorpyrifos in microcosms.
Background levels of chlorpyrifos and earthworm abundance were determined in an orchard and adjacent areas on a farm in the Western Cape, South Africa before these areas were again sprayed with this organophosphate. The background concentrations ranged from 0.2 microg/kg dm in the spray drift area adjacent to the orchard to 10.18 microg/kg dm on the slope in the run off area. In the target area the chlorpyrifos concentrations varied from a mean of 15.25 +/- 10.0 microg/kg directly after spraying to a mean of 7.0 +/- 0.9 microg/kg 6 months later and in the nontarget area they varied from a mean of 55.0 +/- 35 microg/kg to 12.0 +/- 5 microg/kg after 6 months. Chlorpyrifos was therefore still present in the field soils, but at lower concentrations, up to 6 months after the last spraying event. Earthworm abundance and population densities were very low. Only Aporrectodea caliginosa was found and the densities were much lower in the orchards (22 per m(2)) than in the nontarget areas (98.3 per m(2)). Microcosm studies were undertaken to relate biomarker responses to chlorpyrifos with biomass changes. Microcosms were filled with soil from the same areas and earthworms of the species A. caliginosa were introduced. The microcosms were treated with a series of concentrations of chlorpyrifos in the laboratory under controlled conditions. These concentrations were chosen to fall within the background ranges found in the soils. The biomass of the worms was determined regularly for a period of 5 weeks and worms in a state of estivation were noted. Earthworms were removed from the microcosms for biomarker tests: for cholinesterase (ChE) inhibition assays every week and for a neutral red retention determination 2 weeks after the exposures started. The most prominent biomass loss was noted in earthworms exposed to the highest pesticide concentration of 8.0 microg/kg. Estivation was higher among earthworms exposed to higher exposure concentrations. Inhibition of ChE increased with higher exposure concentrations and with time but there was no clear dose-related response. A clear dose-related response with exposure concentration was established for the neutral red retention assay. A correlation between ChE inhibition and biomass change existed directly after the second application of chlorpyrifos.
Animals↗
Cardiac natriuretic peptide response to water restriction in the hormonal adaptation of two semidesert rodents from West Africa (Steatomys caurinus, Taterillus gracilis).
Two African rodents, Taterillus gracilis and Steatomys caurinus, native to regions of alternate dry and wet seasons, were studied under laboratory conditions. These species differ in estivation behavior, one undergoing pseudoestivation and the other strong estivation. One group of animals of each species was provided with unlimited access to seed and vegetables rich in water, mimicking the food availability of the wet season (control group). A second group of animals of each species was subjected to water restriction for 8 days, mimicking the natural drought that occurs during the dry-hot season. The effects of water restriction on osmoregulation and body water content were assessed from hematocrit, and plasma and urinary osmolalities (PO, UO). Whether the natriuretic peptide system was modified by the osmoregulator adaptation to aridity of these semidesert rodents was examined from measurements of atrial natriuretic peptide (ANP) levels in plasma, atria, and ventricles, in parallel with morphological studies. In both species, UO was increased by water restriction. In water-deprived T. gracilis, ANP levels were about twice (right atria: 1.08 +/- 0.16 microg/mg protein vs control: 0.40 +/- 0.06 microg/mg protein) and plasma concentrations half (0.28 +/- 0.06 ng/ml vs control: 0.64 +/- 0.07 ng/ml) those in control animals. In S. caurinus these variables were not affected by water availability (right atria water restricted: 2. 20 +/- 0.15 microg/mg protein vs control: 2.86 +/- 0.37 microg/mg protein; plasma ANP water restricted: 0.80 +/- 0.12 ng/ml vs control: 0.90 +/- 0.16 ng/ml). Consistent with these quantitative results, immunohistochemical and ultrastructural observations showed an increase in immunostaining for both the N- and the C-terminal ANP and a larger number of granules in the atria of T. gracilis following water restriction, whereas there was no visible change in S. caurinus. Thus, water restriction induced a decrease in ANP secretion in T. gracilis, increasing cardiac storage alongside a reduced urine production. In contrast, in S. caurinus, the natriuretic system was not affected by an 8-day period of water restriction.
Adaptation, Physiological↗