Rat soleus muscle ultrastructure after hindlimb suspension.

The aim of the present investigation was to determine, by quantitative electron microscopy, the effects of a 5-wk tail-suspension period on rat soleus muscle ultrastructure. A marked decline (-60%) in muscle mass occurred. The mean fiber cross-sectional area decreased to a greater extent (-75%) than the capillary-to-fiber ratio (-37%), leading to a higher capillary density (+148%) after hypokinesia. The total mitochondrial volume density remained unchanged, whereas the volume density of myofibrils was slightly but significantly reduced (-6%). A shift from subsarcolemmal to interfibrillar mitochondria occurred. Interfibrillar mitochondrial volume density was highest near the fiber border and decreased toward the fiber center. An increase in volume density of satellite cells suggested muscle regenerative events. Soleus atrophy with tail suspension greatly decreases the muscular volume but leaves the ultrastructural composition of muscle fibers relatively unaffected.

Effect of spontaneous recovery or retraining after hindlimb suspension on aerobic capacity.

The purpose of this study was to compare the effects of spontaneous recovery or recovery by treadmill training (180 min/day, 5 days/wk, 30 m/min for 8 wk) on maximal O2 uptake (VO2max), histochemical and biochemical muscular properties (soleus), of rats subsequent to 5 wk of hindlimb suspension. Spontaneous recovery reversed the 15% reduction in VO2max, whereas training posthypokinesia induced a 20% increase over control values. In the spontaneous recovery group, both citrate synthase and 3-hydroxyacyl-CoA dehydrogenase activities, decreased by hypokinesia (-40%), increased but remained 20% below the control level. In the training posthypokinesia group, an increase of these activities over control occurred (+50 and +20%, respectively). Recovery or training led to a 100% type I distribution in soleus muscle and to a recovery of all fibers' cross-sectional areas. In the spontaneous recovery group, capillaries per fiber, decreased by 46%, returned to the normal range. In the training posthypokinesia group, training induced an increase in capillaries per fiber above their control values (+23%). These results point to the plasticity of the muscle and indicate the necessity of a posthypokinesia training program for recovery of the total oxidative enzyme capacity.

Free dopamine in dog plasma: lack of relationship with sympathoadrenal activity.

To investigate the relationship between dopamine (DA) released into the bloodstream and sympathoadrenal activity, levels of free DA, norepinephrine (NE), and epinephrine (E) in plasma were recorded in four dogs subjected to three tests: treadmill exercise at two work levels [55 and 75% maximal O2 uptake; 15 min], normobaric hypoxia (12% O2; 1 h), combined exercise and hypoxia. Normoxic exercise induced slight nonsignificant decreases in the arterial partial pressure of O2 (PaO2), increases in NE [median values and ranges during submaximal work vs. rest: 1086 (457-1,637) vs. 360 (221-646) pg/ml; P less than 0.01] and E [277 (151-461) vs. 166 (95-257) pg/ml; P less than 0.05], but it failed to alter the DA level. Hypoxia elicited large decreases in PaO2 [hypoxia vs. normoxia: 42.8 (40.3-50.0) vs. 97.6 (83.2-117.6) Torr; P less than 0.01], increases in DA [230 (105-352) vs. 150 (85-229) pg/ml; P less than 0.01] and NE [383 (219-1,165) vs. 358 (210-784) pg/ml; P less than 0.05], but it failed to alter the E level. Combined exercise and hypoxia further increased NE levels but did not alter the DA response to hypoxia alone. The data indicate that free DA in plasma may vary independently of the sympathoadrenal activity.

Structural and functional responses to prolonged hindlimb suspension in rat muscle.

The purpose of this study was to investigate alterations in structural and functional properties in the soleus (SOL) and extensor digitorum longus (EDL) muscles of rats after 1, 2, and 5 wk of tail suspension. Maximal O2 uptake was 19% lower after 5 wk suspension. Loss of muscle mass was greater in SOL (63%) than in EDL (22%) muscle. A reduction of type I distribution was accompanied by an increase of intermediate fiber subgroups (int I in SOL, int II in EDL). The cross-sectional area of all three fiber types was reduced by hypokinesia. The decrease in capillaries per fiber in SOL was greater than the decrease in citrate synthase and 3-hydroxyacyl-CoA dehydrogenase activities after 5 wk. No alteration in lactate dehydrogenase activity was noted. In EDL, no changes in fiber area, capillarization, and enzymatic activities occurred. Energy charge remained unchanged (0.91) whatever the muscle. These results suggest that type I fibers showed an earlier and greater susceptibility than type II fibers to suspension which is also accompanied by a decreased aerobic capacity.

Effects of gluconeogenic precursor flux alterations on glycogen resynthesis after prolonged exercise.

The importance of gluconeogenic substrates (i.e., lactate, glycerol, and alanine) in the glycogen resynthesis observed in fasting rats after exhausting submaximal exercise [R.D. Fell et al. Am. J. Physiol. 238 (Regulatory Integrative Comp. Physiol. 7): R328-R332, 1980] was examined in muscles and liver in response to pharmacological alterations of gluconeogenic precursor flux. The minor role of lactate for glycogen resynthesis after prolonged submaximal exercise was confirmed by the insignificant accumulation of lactate neither in muscles nor in plasma. When the rate of lipolysis is reduced either by beta-blockade or by nicotinic acid injection, the replenishment of muscle glycogen persisted, suggesting that glycerol released by triglycerides hydrolysis did not play an important role in glycogen resynthesis. On the other hand, when pyruvate oxidation is enhanced by dichloroacetate (DCA), thus reducing plasma levels of lactate and alanine, glycogen resynthesis was completely blocked in liver and partly in some but not all muscles. This failure in total inhibition of glycogen resynthesis associated with the significant reduction of the plasma alanine level could be attributed to the possible stimulation of gluconeogenesis from alanine by DCA (R.A. Harris and D.W. Crabb. Arch. Biochem. Biophys. 189: 364-371, 1978). The results could point out alanine as the major gluconeogenic substrate during recovery from exhaustive exercise in fasting conditions.

Skeletal muscle adaptation in rats flown on Cosmos 1667.

Seven male Wistar rats were subjected to 7 days of weightlessness on the Soviet biosatellite Cosmos 1667. Muscle histomorphometry and biochemical analyses were performed on the soleus (SOL) and extensor digitorum longus (EDL) of flight rats (group F) and compared with data from three groups of terrestrial controls: one subjected to conditions similar to group F in space except for the state of weightlessness (group S) and the others living free in a vivarium (V1, V2). Relative to group V2 (its age and weight-matched control group), group F showed a greater decrease of muscle mass in SOL (23%) than in EDL (11%). In SOL a decrease in the percentage of type I fibers was counterbalanced by a simultaneous increase in type IIa fibers. The cross-sectional area of type I fiber was reduced by 24%. No statistically significant difference in capillarization and enzymatic activities was observed between the groups. In EDL a reduction in type I fiber distribution and 3-hydroxyacyl-CoA-dehydrogenase activity (27%) occurred after the flight. The small histochemical and biochemical changes reported suggest the interest in studying muscular adaptation during a flight of longer duration.

[Recent data on respiratory control during muscular exercise in man]. / Données récentes sur le contrôle ventilatoire à l'exercice musculaire chez l'homme.

During the last two decades, numerous investigations have been conducted to determine the degree of involvement and mechanism of action of various factors responsible for ventilatory adaptation to exercise in human. The neuromechanical ventilatory system plays a role, exercise ventilation representing a compromise between the necessity of maintaining an adequate level of chemical arterial stimulus and of avoiding mechanical overload and, consequently, respiratory fatigue. The role of arterial chemoreceptors is essential and that of muscular chemoreceptors likely. The limb mechanoreceptors also play a role essentially via small nerve fibers on the other hand. To date, there is no experimental evidence of ventilatory reflexes elicited by pulmonary chemoreceptors. Further, the action of cardiopulmonary baroreceptors was proved only in extraphysiological conditions in animals. It is also possible that a feed-forward mechanism originating in the central nervous system plays an important role. In conclusion, the time course of pulmonary ventilation during exercise could be explained by the action of humoral and neurogenic stimuli.

[Control of pulmonary ventilation during muscular exercise (author's transl)]. / Le contrôle de la ventilation pulmonaire au cours de l'exercice musculaire.

The ventilatory changes as function of time or power depend on the type of exercise. However, in any case, two kinds of factors are involved in these changes : fast, neurogenic, factors and other factors, slower, which are thought to be humoral. Neurogenic stimuli arise from the periphery and from the central nervous system. Peripheral stimuli are induced by the repetitive changes of muscle length and by the motion of articulations. Ventilatory output is also increased by stimuli coming from motor centers. These two kinds of factors induce a ventilatory increase at the start of exercise ("accrochage ventilatoire"). Both chemical and physical stimuli act as humoral factors. The arterial PO2 is scarcely reduced even during maximal exercise, but the increased blood level of norepinephrine may account for the enhanced sensitivity of the centers to the oxygen drive. The sensitivity of the respiratory centers to PaCO2 is not increased. The pH is lowered and can reach 7.15 during very intense exercise. tthis acidity may account for the increased ventillatoy output under these circumstances. The increased body temperature may partly explain the progressive increase of ventilatory output during long lasting exercise, but it does not change the alveolar ventilation. It thus appears that the exercise ventilatory behaviour meets the metabolic, acid-base balance and thermolytic requirements.

[Role of aerobic metabolism in prolonged intensive exercise]. / Le rôle du métabolisme aérobie dans l'exercice intense de longue durée.

The purpose of this review is to analyze the different factors which intervene with respect to the ability to perform intense exercises of long duration. This ability implies : (a) a high level of maximal oxygen uptake (Vo 2 max) and (b) a great endurance, i.e. the capacity of maintaining for a long time a high percentage of this Vo2 max. First of all, the values Vo2 max usually recorded on normal subjects are given, as well as those of subjects specially trained for endurance events ; then the factors which condition a high level of Vo2 max are described. These factors consist on great capacity in all of the links of the oxygen transport system. These capacities appear to be essentially dependant on the genetic endowment, but their maximal size can only by attained through intensive training. Endurance can be defined as the maximal period of time during which a given percentage of the Vo2 max can be sustained. The values recorded on normal subjects and on those specially trained for endurance events are given, and then the factors conditioning a high level of endurance are studied. Endurance appears to depend on the feeding of muscular fibers with energentic substrates. Training permits the muscles to store large quantities of glycogen. It also seems to enable the organism to increase the percentage of energetic substrates made up by lipids. This latter ability in associated with an alteration of the endocrine reactions induced by exercise. Finally, endurance is accompanied by an ability of the body to store and to eliminate the heat produced during exercise.

[Metabolic and respiratory parameters during muscular exercise in man (author's transl)]. / Evolution des paramètres métaboliques et respiratoires au cours de l'exercice musculaire chez l'homme.

Muscle can use ATP exclusively as the direct source of energy for contraction. The muscle ATP stores cannot provide more than 1 or 2 kcal of muscular work. tthe energy for resynthetizing ATP is supplied by three processes : the breakdown of creatine phosphate, anaerobic glycolysis and aerobic processes. These three mechanisms are characterized by different inertia, maximal output and capacity. Taking into account the part of aerobic processes in energy production, the physical fitness of an individual is usually expressed by its maximal oxygen uptake (VO2 max.). When the energy requirements cannot be met by aerobic reactions, the subject contracts an oxygen deficit which is compensated for during the recovery period by an oxygen uptake exceeding the rest requirements. During exercise tidal volume and ventilatory frequency are increased. The increase in ventilatory output is directly related to the workload until 75% of the maximal aerobic power is reached. For higher relative workloads the increase in ventilatory output is steeper. This increased ventilation allows the organism to limit the decreases in PaO2 and pH during exercises of high intensity.

Thermoregulation at rest and during exercise in prepubertal boys.

Thermal balance was studied in 11 boys, aged 10-12 years, with various values for maximal oxygen uptake (VO2max), during two standardized sweating tests performed in a climatic chamber in randomized order. One of the tests consisted in a 90-min passive heat exposure [dry bulb temperature (Tdb) 45 degrees C] at rest. The second test was represented by a 60-min ergocycle exercise at 60% of individual VO2max (Tdb 20 degrees C). At rest, rectal temperature increased during heat exposure similar to observations made in adults, but the combined heat transfer coefficient reached higher values, reflecting greater radiative and convective heat gains in the children. Children also exhibited a greater increase in mean skin temperature, and a greater heat dissipation through sweating. Conversely, during the exercise sweating-test, although the increase in rectal temperature did not differ from that of adults for similar levels of exercise, evaporative heat loss was much lower in children, suggesting a greater radiative and convective heat loss due to the relatively greater body surface area. Thermophysiological reactions were not related to VO2max in children, in contrast to adults.

Increase in sweating sensitivity by endurance conditioning in man.

Sweating sensitivity has been evaluated at rest in 10 competitive athletes (cross-country skiers and swimmers). Three sedentary men underwent a 3-mo period of endurance training in a temperate climate, (dry bulb temperature (Tdb): 18 degrees C) and had their sweating sensitivity measured before and after the training period. Mean maximum oxygen uptake (Vo2max, ml.min(-1).kg(-1)) was: skiers: 66.5; swimmers 65.8; sedentary men, pretraining 40.9; posttraining: 48.3 (+18%). Sweat output of athletes under a given stress (passive heating) was markedly higher than that of sedentary men. Skiers exhibited a high level of heat tolerance and were better acclimatized than swimmers, although they had never experienced exposure to heat. The increase in Vo2max of sedentary men was accompanied by 1) an increase in sweating sensitivity with a decrease of body heat storage at steady state (pretraining: 5.4 kJ.kg(-1); posttraining: 3.5 kJ.kg(-1); P less than 0.05); 2) significant shift down the temperature scale with reduced rectal temperature (Tre) for sweat onset; 3) an increase of gain constants of sweating (W.m-2 degrees C(-1) (pretraining: 168; posttraining: 269; gain constant of swimmers: 222). It was suggested that endurance training in cold or temperate conditions with significant increase of Vo2max could act on the thermoregulatory function in a way similar to body heating procedures, such as work in heat, and could contribute to heat acclimatization.

Effects of chronic lung denervation on breathing pattern and respiratory gas exchanges during hypoxia, hypercapnia and exercise.

The influence of vagal fibres from the lung on ventilatory responses to hypercapnia, hypoxia and exercise was studied in two intact dogs (C) and two chronically lung denervated dogs (C.L.D.). In intact dogs, inspiration duration did not change as tidal volume increased in response to increased chemical drives. Chronic lung denervation did not affect the hypercapnia- or hypoxia-induced elevations in V, despite significant changes in breathing pattern. During exercise, oxygen consumption was similar for C and C.L.D. animals. V for a given oxygen uptake was the same in C and C.L.D. dogs, but VT was higher in C.L.D. animals at all levels of exercise. It is concluded that vagal fibres from the lung play a role in determining the breathing pattern, but are not required for a normal ventilatory response to hypercapnia, hypoxia and exercise. Interaction between vagal sensory input and specific structures sensitive to chemical or physical stimuli is discussed.

[Adrenergic response to intense muscular activity in sedentary subjects as a function of emotivity and training]. / Réponse adrénergique à l'exercice musculaire intense chez le sujet sédentaire en fonction de l'emotivité et de l'entrainement.

Seven male sedentary human subjects were studied during intense muscular work (80% of maximal oxygen uptake) performed either for 15 min or until exhaustion (mean duration: 47 +/- 2 min). Plasma catecholamines were estimated before and after the experiment by means of an original fluorimetric assay. Epinephrine or norepinephrine were individually isolated from plasma and assayed in single extracts by a highly sensitive fluorimetric method. Epinephrine and norepinephrine levels as low as 15 ng per liter were detectable by this procedure in human plasma. The adrenergic pattern was found to be greatly different from one subject to another and related to emotivity: the effect of this factor was revealed by the predominance of epinephrine in plasma at rest or under exercise (ratio NA/A less than 1). In nonemotive subjects (ratio NA/A greater than 1 at rest) plasma epinephrine and norepinephrine increased progressively during exercise. Increments after exercise were higher for norepinephrine changes; however, the fact that epinephrine concentrations correlated significantly with norepinephrine suggests a simulataneous and coordinated stimulation of adrenal glands and orthosympathetic nervous system. In emotive subjects (ratio NA/A less than 1 at rest) the apprehension of muscular work promoted a difference in catecholamine responses: norepinephrine release was not affected by subject's anxiety, while epinephrine secretion, already elevated before the test, reached a high degree of magnitude in the first minutes of muscular work, remaining nearly constant until exhaustion. Physical training of nonemotive subjects, during 2 months with two intense exercises by a week, reduced strongly norepinephrine release after exhaustive muscular work. In the same conditions, the adrenal-medullary response was not significantly modified when compared with untrained subjects. Our results suggest that the adrenergic behaviour during exercise is a function of effort intensity to be supplied; catecholamines seem to be important factors in regulating body homeostasy during muscular work in man. In addition, emotive subjects exhibit amplified adrenal-medullary response, which may be related to psychological stimuli.

[Chemoreflexive drive of ventilation and noradrenaline stimulus in man]. / Commande chémoréflexe de la ventilation et stimulus noradrénaline chez l'homme.

O2 chemoreflex drive of ventilation was studied before and after an intravenous infusion of L-norepinephrine (9 microgram/min), inducing a plasmatic hormone concentration similar to that obtained during submaximal exercise. The ventilation increasing rapidly at the beginning of the infusion was stabilized after 30 min : the ventilation was two times the reference value. The O2 chemoreflex drive of ventilation increased during norepinephrine infusion. When man was transiently switched from hypoxia to pure O2 (O2 test) the maximal fall of ventilation was two times the reference response. This increase in the chemoreflex drive, although the physico-chemical blood state was unchanged, may be explained by a norepinephrine chemoreceptor sensitization. Such a mechanism could partly explain the increase of O2 chemoreflex drive observed during muscular exercise.

Structural and metabolic properties of rat muscle exposed to weightlessness aboard Cosmos 1887.

Male Wistar rats were subjected to 12.5 days of weightlessness aboard Cosmos 1887. Histomorphometric and biochemical analyses were investigated in soleus (SOL), plantaris (PL) and extensor digitorum longus (EDL) muscles of flight rats (group F) and compared with data from two groups of terrestrial controls: one group living free in a vivarium (group V) and another subjected to a flight simulation except for the state of weightlessness (group S). Relative to groups V and S, no alteration in the percentage distribution of fibres had occurred in SOL, PL or EDL, after the flight. In SOL muscles from group F animals, cross-sectional areas of all fibre types were reduced to a greater extent (-40%) than capillary to fibre ratio (-24%) leading to a higher capillary density (+33%) than in V and S groups. In PL, type I, IIA and IIB fibre cross-sectional areas were less decreased (-25%). In EDL, only fast-twitch fibre cross-sectional areas showed an average decrease of 30%. Capillary per fibre ratio was reduced by 15% and 28% respectively in PT and EDL muscles from group F rats compared to control groups V and S. Citrate synthase and 3-hydroxyacyl-coenzyme A dehydrogenase activities remained unchanged in SOL, PL and EDL following spaceflight. These findings indicate greater atrophy and functional alterations (capillarity) compared to those observed after 7 days of microgravity on Cosmos 1667.

Effects of hypoxia on catecholamine and cardiorespiratory responses in exercising dogs.

The sympathoadrenal contribution to cardiorespiratory response elicited by hypoxia and/or exercise was assessed in the dog. The increased plasma norepinephrine (NE) and dopamine (DA) levels which follow hypoxia (fraction of inspired O2 equals 0.12) while epinephrine (E) remained unchanged ruled out the possibility of a primacy of the adrenal medulla in the response to hypoxia. In contrast to the lack of effect of hypoxic exposure, the adrenal medulla was substantially stimulated during exercise. The exercise-induced sympathoadrenal response remained unchanged during hypoxia as compared to normoxia when expressed as function of relative work intensity. Nevertheless at a given oxygen uptake, all plasma catecholamines were increased by hypoxia. These modifications in hormonal milieu failed, however, to alter the cardiac responses to exercise but were associated with a change in breathing pattern.

Aerobic performance capacity in paraplegic subjects.

To determine adaptation to prolonged exercise in paraplegics, maximal O2 uptake (VO2max) and lactate threshold (LT) were evaluated during an arm cranking exercise in nine patients (P) and nine able-bodied (AB) subjects. Mean VO2max averaged 25.1 and 31.6 ml X min-1 X kg-1 in P and AB groups respectively. VO2max in P was found to be directly related to the level of spinal injury: the higher the lesion the lower the uptake. Lactate threshold expressed as a percentage of VO2max was higher in P (59%) than in AB (43%), and close to that observed in arm-trained athletes. Since training has less effect on VO2max in paraplegics than in able-bodied subjects, attributable to a deficiency in the circulatory adaptation of paraplegics to exercise, the observed differences between AB and P in lactate threshold and submaximal exercise indicate that the possible effect of training in paraplegics is located at the level of intracellular chemistry, with a diminution in glycogenolysis (higher LT) and a higher rate of lipid utilization (lower RQ).