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Effects of thermal acclimation on nervous conduction and muscle contraction in the frog Rana temporaria.

The effects of season and acclimation temperature on the latency of the leg withdrawal reflex and three of its components have been studied: conduction velocity in the sciatic nerve, spinal conduction time, and contraction time of gastrocnemius muscle. The latency of the leg withdrawal reflex was markedly shortened by cold acclimation: the reaction times were at 6 degrees C 1.54 s in 4 degrees C acclimated and 3.97 s in 24 degrees C acclimated winter frogs. Also, the temperature dependence of the reflex latency was reduced by cold acclimation. Thus, frogs acclimated to cold responded to external stimuli in cold more rapidly than warm-acclimated ones. This cold adaptation of the reflex could not be explained by changes in its studied components. These made up only one-tenth of the reflex response time, and either did not show significant cold acclimation (muscle contraction and spinal conduction times in summer) or showed inverse acclimation, especially when measured at high temperatures (i.e. conduction velocities were reduced by acclimation to cold). Thus, the cold acclimation of the reflex response probably resides in the sensory component of the response. The inverse temperature adaptation response of conduction velocities may reflect a reduced ion permeability across cellular membranes in cold which decreases metabolic energy expenditure during inactive periods.

Adaptation, Physiological↗

Thermal acclimation, growth, and burst swimming of threespine stickleback: enzymatic correlates and influence of photoperiod.

Threespine sticklebacks (Gasterosteus aculeatus) that had been reared in the laboratory under natural photoperiods were acclimated to 23 degrees and 8 degrees C in late spring under increasing day lengths and again in late fall under decreasing day lengths. The parents of these fish were from the anadromous Isle Verte population. In the spring, cold- and warm-acclimated fish grew at the same rates and attained similar condition factors (mass L(-3)), although food intake was considerably higher at 23 degrees C. As both groups had similar increases in mass and condition, the higher axial muscle activities of citrate synthase and phosphofructokinase (measured at 20 degrees C) after cold acclimation were likely a direct response to temperature. Multiple regression analysis showed that axial muscle levels of cytochrome C oxidase and citrate synthase were correlated with the burst swimming speeds of the spring sticklebacks, while growth rates were positively correlated with lactate dehydrogenase levels in pectoral and axial muscles and creatine kinase levels in the axial muscle. In the fall, the fish in both acclimation groups grew little, although they fed at similar rates as in the spring experiment. Overall, the sticklebacks showed lower burst swimming speeds in the fall. In both spring and fall, the burst speeds of cold- and warm-acclimated sticklebacks only differed at warm temperatures. In the spring experiment, the cold-acclimated fish swam faster, whereas in the fall experiment the warm-acclimated fish swam faster despite their lower percentage of axial muscle. Swimming speeds were measured both at a fish's acclimation temperature and after 12 h at the other temperature. Cold-acclimated sticklebacks seem to have more facility in rapidly adjusting to warm temperatures when they have experienced increasing rather than decreasing day lengths, perhaps as a result of the requirements of the spring migration to the intertidal breeding grounds.

Acclimatization↗

Fast skeletal myosin isoforms in thermally acclimated carp.

Fast skeletal myosins were isolated from carp acclimated to 10 and 30 degrees C, and their structural and enzymatic properties were compared. Myosins in 0.5 M KCl were subjected to limited proteolysis by using various proteases including alpha-chymotrypsin, trypsin, and papain, and different SDS-PAGE patterns were seen for the 10- and 30 degrees C-acclimated myosins in all cases. Myosin subfragment-1 (S1) prepared from the 10 degrees C-acclimated myosin by alpha-chymotryptic digestion in 0.12 M NaCl showed higher acto-S1 Mg(2+)-ATPase activity and lower thermostability than S1 from the warm-acclimated myosin. The peptide maps and ATP-induced spectral changes of tryptophan fluorescence also showed an obvious difference between the two types of S1. Temperature acclimation further caused changes in the rod region of myosin, since the apparent sizes of light meromyosin were different from each other for the two types of myosin. Myosin from carp acclimated to 20 degrees C showed intermediate properties between those of the 10- and 30 degrees C-acclimated myosins. Myosin isoforms might be expressed in a temperature-dependent manner to compensate for the effect of seasonal environmental temperature variation on swimming ability.

Acclimatization↗

Impact of temperature on the relationship between respiration and nitrogen concentration in roots: an analysis of scaling relationships, Q10 values and thermal acclimation ratios.

* The impact of nitrogen (N) supply on the temperature response of root respiratory O(2) uptake (R) was assessed in several herbaceous species grown in solution culture. Warm-grown (25 : 20 degrees C, day:night) plants differing in root N concentration were shifted to 13 : 8 degrees C for 7 d to cold-acclimate. * Log-log plots of root R vs root N concentration both showed that R increased with increasing tissue N concentration, irrespective of the growth temperature. Although the regression slopes of the log-log plots did not differ between the warm-grown and cold-acclimated plants, cold-acclimated plants did exhibit a higher y-axis intercept than their warm-grown counterparts. This suggests that cold acclimation of root R is not entirely dependent on cold-induced increases in tissue N concentration and that scaling relationships (i.e. regression equations fitted to the log-log plots) between root R and N concentration are not fixed. * No systematic differences were found in the short-term Q(10) (proportional change in R per 10 degrees C change in temperature), or degree of cold acclimation (as measured by the proportional difference between warm- and cold-acclimated roots) among roots differing in root N concentration. The temperature response of root R is therefore insensitive to tissue N concentration. * The insensitivity of Q(10) values and acclimation to tissue N concentration raises the possibility that root R and its temperature sensitivity can be predicted for a range of N supply scenarios.

Acclimatization↗

Effects of diet and thermal acclimation on oxygen consumption in rats.

The relationship between food intake and oxygen consumption was studied in a group of 60 rats acclimated at environmental temperatures of either 30 or 10 degrees C. Three separate experiments were performed. In the first, 28 rats were divided into two groups: control, which received 20 and 32 g of food/day at 30 and 10 degrees C, respectively, from 0800 to 1700 and experimental, which received 10 and 25 g of food/day at 30, and 10 degrees C, respectively. The experimental period lasted 6 weeks. Oxygen consumption was measured weekly at environmental temperatures of 5, 15, 25, 30, and 35 degrees C. In the second experiment, 16 rats were subjected to the same food intake as the animals in the first experiment. After 1 week, their oxygen consumption was measured at 25 degrees C over a period of 24 h. The third experiment was carried out with 16 other rats in which the control groups received the same amount of food as in the first experiment, and the experimental groups were fed 6 and 11 g/day at 30 and 10 degrees C, respectively, during 1 week. In the first experiment, no changes in oxygen consumption (per kilogram 0.67) were apparent in the experimental rats during 6 weeks. However, after 1 week on severe food restriction a significant decrease in oxygen consumption (per kilogram 0.67) was observed. A long-lasting thermic effect of food was observed in control rats from the second experiment and a rapid effect was apparent in restricted rats.(ABSTRACT TRUNCATED AT 250 WORDS)

Acclimatization↗

Influence of thermal acclimation on membrane lipid composition of rainbow trout liver.

Rainbow trout (Salmo gairdneri) acclimated to 5 degrees C possessed larger livers and less neutral lipid per gram of liver than 20 degrees C-acclimated animals; quantities of liver glycolipid, phospholipid, and cholesterol did not vary significantly with acclimation temperature. The relative proportions of phosphatidylethanolamine increased significantly following cold exposure, whereas the quantities of sphingomyelin and cardiolipin declined. For all phosphatides examined (phosphatidylethanolamine, phosphatidylcholine, phosphatidylserine, phosphatidylinositol, lysolecithin, cardiolipin, sphingomyelin) cold acclimation resulted in 1) an increase in the quantity of polyunsaturated fatty acids, 2) a reduction in the level of saturated fatty acids, and 3) little change in the total content of monoenes and dienes. The increased content of polyunsaturated fatty acids in choline and ethanolamine phosphatides following cold acclimation was confined to the 2-position and occurred at the expense of monoenes and dienes. The relative proportions of n - 3 fatty acids, and less frequently n - 6 fatty acids, increased in phosphatides of cold-acclimated trout, whereas the relative proportions of n - 9 fatty acids declined. These data suggest a preferential incorporation of fatty acids belonging to the linolenic acid family at reduced temperatures. Temperature-induced changes in the chemical composition of trout liver phospholipids counteracted the effects of acute temperature change on nonelectrolyte permeability of isolated liposomes.

Adaptation, Physiological↗

Influence of thermal acclimation on glucose production and ketogenesis in isolated eel hepatocytes.

Hepatocytes were isolated by collagenase perfusion of the liver from adult eels (Anguilla anguilla L.) acclimated to different temperatures. Whereas the relative weight of the liver increased in cold-acclimated fish, hepatocytes from 10- and 20 degrees C-acclimated animals did not differ in cellular weight, dry weight, or protein content. Endogenous rates of oxygen consumption and respiratory control ratios were independent of acclimation temperature. There was no effect of temperature on triacylglycerol content, but glycogen concentration was significantly higher in hepatocytes of cold-acclimated fish. Liver cells from cold-acclimated eels exhibited higher rates of glucose release and ketogenesis than those from warm-acclimated animals. It is concluded that the increase in acetoacetate production induced by cold acclimation results primarily from a higher rate of lipolysis. Cellular interactions between ketogenesis and gluconeogenesis are demonstrated and discussed.

Acetoacetates↗

Thermal acclimation induces adaptive changes in subcellular structure of fish skeletal muscle.

Stereological analyses of electron micrographs were used to quantify physiologically important ultrastructures of slow-twitch oxidative (red) and fast-twitch glycolytic (white) muscle fibers from striped bass (Morone saxatilis) acclimated to 25 and 5 degrees C. The fraction of cell volume occupied by the mitochondria [volume density, Vv (mit,f)] of red fibers increases from 0.286 +/- 0.018 to 0.448 +/- 0.024 between 25 and 5 degrees C; Vv (mit,f) of white fibers increased from 0.027 +/- 0.003 and 0.040 +/- 0.004 at 25 and 5 degrees C, respectively. Because of a concomitant increase in the mass of oxidative muscle, acclimation from 25 to 5 degrees C results in an increase in total mitochondrial volume per 100 g body wt from 2.58 to 6.73 cm3 in oxidative muscle and from 2.46 to 3.40 cm3 in fast glycolytic muscle. Mitochondria of red fibers are in more clustered arrays after cold acclimation. Size and cristae surface densities of individual mitochondria are not affected markedly by acclimation, suggesting true cold-induced proliferation rather than enlargement of organelles. Harmonic means of intermitochondrial spacing in red fibers decreases from 2.64 to 1.43 micron between 25 and 5 degrees C. This reduces diffusion path lengths between sarcoplasmic and mitochondrial compartments proportionately, compensating for decreases in diffusivity of aqueous solutes. Intracellular lipid droplets of red fibers markedly increase in volume density from 0.006 +/- 0.003 at 25 degrees C to 0.079 +/- 0.014 at 5 degrees C.(ABSTRACT TRUNCATED AT 250 WORDS)

Adaptation, Physiological↗

The temperature dependence of the time course of growth and decay of miniature end-plate currents in carp extraocular muscle following thermal acclimation.

1. The effect of temperature (5-35 degrees C) on the decay and growth phases of miniature end-plate currents (MEPCs) was investigated in extraocular muscle from freshwater carp acclimated to either high (28 degrees C) or low (8 degrees C) temperature. 2. The temperature dependence of the time constant of decay (TD) was found to follow an Arrhenius relationship; the relationship between logTD and reciprocal of absolute temperature (1/K) being linear in both groups. The TD of MEPCs recorded from cold-acclimated carp was not statistically significant from that of the warm group. 3. TD was moderately temperature-dependent. Regression gave a Q10 of 1.78 for the warm-acclimated carp, corresponding to an activation energy, Ea, of 41.15 +/- 2.17 kJ mol-1. For the cold-acclimated carp, the Q10 was 1.79, and Ea was 41.43 +/- 2.46 kJ mol-1. 4. Growth time (TG) was less susceptible than TD to temperature change. The relationship between growth time (taken as the time for MEPCs to rise from 20 to 80% of maximum) and temperature was linear for the cold-acclimated group, with a Q10 of 1.34 and Ea of 20.94 +/- 4.75 kJ mol-1. The data for the warm group, were, in contrast, best fitted by two linear regressions meeting at 15.1 degrees C. At temperatures below 15.1 degrees C Q10 was 3.16 and Ea was 82.20 +/- 15.47 kJ mol-1; above 15 degrees C, Q10 was 1.22 and Ea was 14.15 +/- 12.24 kJ mol-1. 5. The acetylcholinesterase inhibitor neostigmine increased TD by approximately twofold and raised TG to approximately 1.4 times control values. These effects were observed across the temperature range scanned for both groups. 6. The results are discussed with reference to the documented effects of temperature and temperature acclimation on membrane lipids and proteins.

Acclimatization↗

The influence of thermal acclimation on power production during swimming. I. In vivo stimulation and length change pattern of scup red muscle.

Ectothermal animals are able to locomote in a kinematically similar manner over a wide range of temperatures. It has long been recognized that there can be a significant reduction in the power output of muscle during swimming at low temperatures because of the reduced steady-state (i.e. constant activation and shortening velocity) power-generating capabilities of muscle. However, an additional reduction in power involves the interplay between the non-steady-state contractile properties of the muscles (i.e. the rates of activation and relaxation) and the in vivo stimulation and length change pattern the muscle undergoes during locomotion. In particular, it has been found that isolated scup (Stenotomus chrysops) red muscle working under in vivo stimulus and length change conditions (measured in warm-acclimated scup swimming at low temperatures) generates very little power for swimming. Even though the relaxation of the muscle has slowed greatly, warm-acclimated fish swim with the same tail-beat frequencies and the same stimulus duty cycles at cold temperatures, thereby not affording the slow-relaxing muscle any extra time to relax. We hypothesize that considerable improvement in the power output of the red muscle at low temperatures could be achieved if cold acclimation resulted in either a faster muscle relaxation rate or in the muscle being given more time to relax (e.g. by shortening the stimulus duration or reducing the tail-beat frequency). We test these hypotheses in this paper and the accompanying paper. Scup were acclimated to 10 degrees C (cold-acclimated) and 20 degrees C (warm-acclimated) for at least 6 weeks. Electromyograms (EMGs) and high-speed cine films were taken of fish swimming steadily at 10 degrees C and 20 degrees C. At 10 degrees C, we found that, although there were no differences in tail-beat frequency, muscle strain or stimulation phase between acclimation groups, cold-acclimated scup had EMG duty cycles approximately 20 % shorter than warm-acclimated scup. In contrast at 20 degrees C, there was no difference between acclimation groups in EMG duty cycle, nor in any other muscle length change or stimulation parameter. Thus, in response to cold acclimation, there appears to be a reduction in EMG duty cycle at low swimming temperatures that is probably due to an alteration in the operation of the pattern generator. This novel acclimation probably improves muscle power output at low temperatures compared with that of warm-acclimated fish, an expectation we test in the accompanying paper using the work-loop technique.

Adaptation, Physiological↗

The influence of thermal acclimation on power production during swimming. II. Mechanics of scup red muscle under in vivo conditions.

We have previously shown that the power output of red muscle from warm-acclimated scup is greatly reduced when the fish swim at low temperatures. This reduction occurs primarily because, despite the slowing of muscle relaxation rate at cold temperatures, warm-acclimated scup swim with the same tail-beat frequency and the same stimulation durations, thereby not affording the slower-relaxing muscle any extra time to relax. We hypothesize that power output during swimming could be increased if the stimulus duration were reduced or if the relaxation rate of the red muscle were increased during cold acclimation. Scup were acclimated to 10 degrees C (cold-acclimated) and 20 degrees C (warm-acclimated) for at least 6 weeks. Cold acclimation dramatically increased the ability of scup red muscle to produce power at 10 degrees C. Power output measured from cold-acclimated muscle bundles driven through in vivo conditions measured from cold-acclimated scup swimming at 10 degrees C (i.e. work loops) was generally much greater than that from warm-acclimated muscle driven through its respective in vivo conditions at 10 degrees C. The magnitude of the increase depended both on the anatomical location of the muscle and on swimming speed. Integrated over the length of the fish, the red musculature from cold-acclimated fish generated 2.7, 8.9 and 5.8 times more power than the red musculature from warm-acclimated fish while swimming at 30 cm s(-)(1), 40 cm s(-)(1) and 50 cm s(-)(1), respectively. Our analysis suggests that the cold-acclimated fish should be able to swim in excess of 40 cm s(-)(1) with just their red muscle whereas the warm-acclimated fish must recruit their pink muscle well below this speed. Because the red muscle is more aerobic than the pink muscle, cold acclimation may increase the sustained swimming speed at which scup perform their long seasonal migrations at cool temperatures. We then explored the underlying mechanisms for the increase in muscle power output in cold-acclimated fish. Contrary to our expectations, cold-acclimated muscle did not have a faster relaxation rate; instead, it had an approximately 50 % faster activation rate. Our work-loop studies showed that this faster activation rate, alone, can increase the mechanical power production during cyclical contractions to a surprising extent. By driving cold-acclimated muscle through warm- and cold-acclimated in vivo conditions, we were able to partition the improvement in power production associated with increased activation rate and the approximately 20 % reduction in the duration of electromyographic activity found in the accompanying study. Depending on the position and swimming speed, approximately 60 % of the increase in power output was due to the change in the red muscle's contractile properties (i.e. faster activation); the remainder was due to the shorter stimulus duty cycle of cold-acclimated scup. Thus, by both shortening the in vivo stimulation duration and speeding up the rate of muscle activation as part of cold-acclimation, scup achieve a very large increase in the power output of their red muscle during swimming at low temperature. This increase in power output probably results in an increase in muscle efficiency and, hence, a reduction in the energetic cost of swimming. This increase in power output also reduces reliance on the less aerobic and less fatigue-resistant pink muscle. Both these abilities may increase the swimming speed at which prolonged aerobic muscle activity can occur and thus reduce the travel time for the long seasonal migrations in which scup engage.

Adaptation, Physiological↗

Thermal acclimation in a crustacean neuromuscular system.

1. Effects of temperature on the muscle fibre membrane and synapses of stretcher muscle preparations made from autotomized limbs of the Pacific shore crab (Pachygrapsus crassipes) were investigated. 2. Acclimation of the crabs to different temperatures modified properties of both muscle fibre membrane and synapses. 3. Increased temperature produced an increase in membrane potential of the muscle fibres. A semi-log plot of these data revealed two linear phases of the membrane potential-temperature relationship, with a change in slope near the acclimation temperature. 4. Maximum values for excitatory junction potential (EJP) amplitude and time constant of EJP decay, and minimum values for facilitation were obtained at temperatures close to the acclimation temperature. It is suggested that the decline in EJP amplitude and time constant of decay produced by deviations in temperature from the acclimation temperature is compensated for by an increase in the amount of facilitation. In this way, maximum tension can be produced by the stretcher muscle in a range of at least 8 degrees C around the acclimation temperature.

Adaptation, Physiological↗