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Respiratory properties of blood in awake and estivating lungfish, Protopterus amphibius.

Blood respiratory properties have been studied in awake and estivating African lungfish, Protopterus amphibius. Fish had been estivating 28-30 months when blood was sampled. Hematocrit, O2 capacity and blood hemoglobin concentration increased by about 50% during estivation. Red cell Hb concentration (MCHC) was unaltered. Total blood concentration of nucleoside triphosphates (NTP) was reduced 50% in estivation. Thin-layer chromatography showed that the change in NTP resulted from a GTP/Hb (guanosine triphosphate) reduction to 20% of the value in awake fish. ATP (adenosine triphosphate) concentration remained largely unaltered. GTP/Hb changes were accompanied by marked changes in O2 affinity. The P50 value in blood from awake fish was 33 mm Hg at pH 7.5 compared to 9 mm Hg for the estivating fish. The n-value changed insignificantly. An increased Bohr effect was present in estivating fish, but its importance will depend on circulating pH in the two states. Standard bicarbonate ranged from a low of 8.6 mMol-L-1 (P plasma) at pH 7.5 in an awake fish to 49.6 mMol-L-1 (P) in an estivating fish. CO2 dissociation curves showed a variable Haldane effect having its highest value in the in vivo range of PCO2. The striking increase in O2-Hb affinity during estivation is regarded as an adaptation to a reduced alveolar O2 availability associated with estivation. Altered mechanics of pulmonary ventilation and reduced ambient O2 availability caused by the subterranean habitat may have been selection pressures for the development of the high O2 availability caused by the subterranean habitat may have been selection pressures for the development of the high O2 affinity. The mechanism behind it rests with the recorded change in erythrocytic concentration of GTP.

Adaptation, Physiological

Reduction of thyrotropin-releasing hormone concentrations in central nervous system of African lungfish during estivation.

Thyrotropin-releasing hormone (TRH) has been implicated as an important modulator of arousal state in mammals. Changes in the content of TRH in several brain regions accompany hibernation in the ground squirrel. In the present study, the involvement of TRH in the regulation of arousal was further investigated in the African lungfish, Protopterus annectens, which contain high concentrations of TRH throughout its central nervous system and enter a hibernation-like state, estivation. Lungfish were divided into three groups. Group 1 was fed normally, group 2 was starved while aquatic, and group 3 was allowed to enter into a state of estivation. After 3 months, the lungfish were sacrificed and the concentrations of TRH, norepinephrine, dopamine, and serotonin were determined in the telencephalon, diencephalon, medulla, and spinal cord. In estivation, there was a significant decline in the concentration of TRH in the diencephalon, with no alteration in other regions. Starvation had no effect on regional TRH concentrations. The concentration of norepinephrine, dopamine, and serotonin did not change in estivation; however, a significant elevation of norepinephrine in the diencephalon and dopamine in the telencephalon was observed in starvation. Starvation and estivation were associated with significant declines in the protein content of the diencephalon and medulla. The estivation-linked decline in TRH in the diencephalon of the lungfish is similar to the decrease in TRH content in the hypothalamus in hibernating ground squirrels. These findings lend further support to the importance of TRH in the regulation of arousal state.

Adaptation, Physiological

[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

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

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

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

[Estivation of Biomphalaria tenagophila (Pulmonata, Planorbidae)].

This study reports the finding of Biomphalaria tenagophila naturally aestived in two municipalities of the State of S. Paulo (Brazil): Ubatuba and Conchas. This ethological characteristic was discovered in 15 specimens collected in Ubatuba, and in 6 specimens collected in Conchas. The snails aestived showed vitality after being placed in water.

Adaptation, Physiological

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

[Idiopathic light eruption].

Idiopathic polymorphous light eruptions (PLE) are photodermatites due to an as yet unidentified photosensitizing agent. Benign estival PLE is the most frequent form: the face is spared, and as soon as the patient has acquired a slight tan the eruption will no longer occur. Juvenile spring eruption affects the free border of the helix and may be regarded as a localized and mild form of estival PLE. Polymorphous light eruptions appear as lesions of the face which may resemble lupus erythematosus; photobiological exploration reproduces the lesions experimentally in 70% of the cases; immunohistochemistry studies are in favour of an immunological mechanism. Solar urticaria is a rare physical urticaria started by UVA, UVB and/or visible light: tolerance to sunlight can be induced by repeated exposures, but it does not last. Hydroa vacciniform is a rare PLE in children with leaves smallpox-like scars; photobiological exploration reproduces the lesions with high doses of UVA applied for 3 consecutive days. The remanent photosensitivity syndrome is characterized by an extreme photosensitivity which may be very disabling, preventing the patient to go outdoors; photo-allergological exploration discloses numerous positive patch-tests or photopatch-tests to various allergens, but the role played by these allergens in the occurrence of photosensitivity remains unclear. The syndrome probably results from an exogenous photosensitivity accident that has not regressed after exclusion of the presumably responsible photoallergens. Many treatments tested for these PLE seem to be effective, but they have not yet been controlled by double-blind trials.

Humans

Bioenergetic functions of sleep and activity rhythms and their possible relevance to aging.

The hypothesis is proposed that sleep constitutes a period of dormancy in which energy is conserved to partially offset the increased energy demands of homeothermy. Phylogenetic data indicate that the complete psysiological and behavioral manifestations of sleep are unique to homeotherms; furthermore "ontogeny recapitulates phylogeny" in the parallel development of slow wave sleep and thermoregulation as exemplified in the opossum. Thus, sleep constitutes a state of reduced metabolism that may represent a variation on the theme of dormancy, functionally lying on a continuum of energy conservation processes, ranging from inactivity and estivation to torpor and hibernation. The high amounts of sleep in infancy may involve conservation of energy and its consequent availability for growth. Decreased amounts of stage 4 and total sleep with aging in humans may represent reduced energy demands reflected by parallel declines in basal metabolic rate and physical activity. Disruptions of circadian rhythms of sleep and wakefulness in humans produce impairments in mood and performance independent of total amounts of sleep obtained, and reduce the amplitude of physiological rhythms. It is suggested that aging processes might also be affected by such disruptions in activity rhythms.

Aging

Vitamin D (soltriol), light, and reproduction.

Evidence from autoradiographic studies with 1,25(OH)2-vitamin D3 (vitamin D, soltriol) labeled with tritium and from the literature indicates that the steroid hormone soltriol regulates and modulates reproductive processes in the female, as it does in the male. Nuclear receptors for soltriol have been discovered in the uterus, oviduct, ovary, mammary gland, placenta, and fetal membranes, as well as in the pituitary and hypothalamus. Soltriol is recognized as a transducer and hormonal messenger of sunlight, acting as a somatotropic activator and modulator of vital processes for the seasonal and estival adaptation of growth, development, and procreation. Its influence on calcium equilibrium is just one of its many functions to serve this goal. This article reviews experimental, clinical, and epidemiologic evidence that suggests the involvement of soltriol in the control of reproductive processes, noting its importance for the onset of puberty, fertility, pregnancy, lactation, and probably sexual behavior. Cooperative actions between soltriol and other steroid hormones, especially estradiol, are pointed out.

Animals

Seasonal fluctuations in hormones and behavior of free-living male California ground squirrels (Spermophilus beecheyi).

Plasma testosterone (T), prolactin (PRL), and behavior were monitored concurrently in free-living male California ground squirrels (Spermophilus beecheyi). Male plasma T peaked in January, 2-3 months after males emerged from their annual period of estivation. PRL peaked 6 to 10 weeks after T. Temporal patterns of space use by males were directly correlated with plasma T levels, but seasonal maxima in rates of most monitored male behaviors occurred 4 weeks after the T peak. No monitored behavior rates were correlated with plasma PRL values. Yearling and adult males did not differ with respect to circulating levels of either T or PRL. Both groups of males mated in our study population, and did not differ significantly with respect to rates of most monitored behaviors. However, yearlings pursued fewer females, mated less frequently, and required more mounts to complete each copulation than did adults. We suggest that T serves an important priming function for various seasonally fluctuating behaviors, but also that mating experience might influence the efficiency of copulatory behavior in this species.

Animals

Systemic administration of kainic acid produces elevations in TRH in rat central nervous system.

Several studies have suggested that the concentration of thyrotropin releasing hormone (TRH) in the central nervous system (CNS) is influenced by the level of CNS activation. Hibernation in the ground squirrel and estivation in the lungfish result in region-specific decreases in TRH concentrations. Repeated electroconvulsive shock (ECS) and amygdaloid kindling have been shown to result in elevations of TRH in limbic brain regions. In the present study, limbic seizures induced by systemic administration of kainic acid resulted in substantial increases in the TRH content of posterior cortex and of dorsal and ventral hippocampus, and in moderate elevations in anterior cortex, amygdala/piriform cortex and corpus striatum. Maximal elevations in TRH were observed 2-4 days after kainic acid administration, and by 14 days TRH levels were similar to control values, with the exception of the dorsal hippocampus, which exhibited more prolonged elevations in TRH levels. Prior exposure to limbic seizure activity attenuated the magnitude of TRH elevation in response to a second administration of kainic acid in the posterior cortex but in no other region. These results indicate that seizure-related processes or events influence TRH systems in the CNS. Neuronal populations involved in limbic seizure induced damage may be involved in the modulation of posterior cortical TRH levels.

Animals