Lowered cutaneous sensitivity to nonpainful electrical stimulation during isometric exercise in humans

Summary The effect of isometric exercise on cutaneous sensitivity to nonpainful electric stimulation was studied in human subjects. The exercises consisted of brief (duration: 1–10 s) palmar flexions of the hand or foot against varying loads (10–30% of the maximal force). A visual go cue was used to indicate the start and end of the exercise. Isometric hand exercise produced a load-dependent increase of electrotactile thresholds of the fingers. The threshold elevation was rapidly attenuated with prolonged duration of the exercise. The hand exercise-induced threshold elevation was of equal magnitude in the glabrous and hairy skin of the fingers. Thresholds were not changed for the hand contralateral to the exercising hand. Passive static pressure of the hand did not produce threshold changes, whereas activation of afferent inhibition by a vibrotactile stimulus (100 Hz, 0.1 mm) did produce a significant threshold elevation. Exerciseinduced threshold elevation was also significant immediately prior to the EMG response of the arm but not at the time of the visual go signal, or before it. The threshold increase found during the EMG response was not significantly stronger than that found prior to the EMG response. These results suggest that isometric exercise load-dependently produces a phasic, rapidly attenuating increase in cutaneous tactile thresholds in the exercising limb but not multisegmentally. Corollary efferent barrage from motor to sensory structures of the brain could be underlying the threshold changes produced by isometric exercise, whereas afferent inhibitory mechanisms seem to have only a minor role.     

Central command increases cardiac output during static exercise in humans

Neural control of the circulation during static two-leg exercise was evaluated in 10 subjects. External compression of the legs was employed to assess muscle mechano-receptor influence by achieving the same intramuscular pressure (80 mmHg) as developed during exercise. The muscle metabo-reflex contribution was assessed by post-exercise muscle ischaemia, and the influence from higher centres in the central nervous system (‘central command’) was taken as the part of the response that could not be accounted for by the two reflex contributions. During static exercise, mean arterial pressure was higher (26±3 mmHg; P<0.01) as compared with leg compression (10±2 mmHg) and with post-exercise muscle ischaemia (11±2 mmHg). Heart rate (25±4 b.p.m.) and cardiac output (0.8±0.3 L min^-1) were increased only during static exercise (P<0.05). Increase in total peripheral resistance were similar during static exercise, post-exercise muscle ischaemia and leg compression. The pressor response to static exercise with a large muscle group was equally attributable to mechanical and metabolic stimulation of afferent nerves; and the two influences were redundant in their effect on total peripheral resistance. In contrast, the influence from central command was directed to the heart with elevation of its rate and minute volume.     

Cardiopulmonary responses to combined rhythmic and isometric exercise in humans

Summary A rhythmic (R) and an isometric (I) exercise were performed separately and in combination to assess their additive effects on arterial systolic (P _as) and diastolic (P _ad) blood pressures, heart rate (f _c), and minute ventilation (V _I). The isometric effort consisted of a 40% maximal voluntary handgrip contraction (MVC) performed for a duration of 80% of a previously determined 40% MVC fatiguing effort. The R effort consisted of a 13-min cycle effort at 75% maximum oxygen consumption (VO_2maX). For the combined efforts, I was performed starting simultaneously with or ending simultaneously with R. Data on nine subjects yield statistically significant evidence (P<0.05) that the effects of I and R are not additive for the following three cases: (1)P _as when I and R are ended simultaneously (I alone=4.9, SEM 0.5 kPa increase; R alone=no significant change from steady state; I+R=1.2, SEM 0.4 kPa increase), (2)P _ad when I and R are started simultaneously (I alone=4.1, SEM 0.4 kPa increase; R alone=0.7, SEM 0.3 kPa decrease; I+R=1.9, SEM 0.4 kPa increase), and (3)P _ad when I and R are ended simultaneously (I alone=4.1, SEM 0.4 kPa increase; R alone=0.3, SEM 0.5 kPa decrease; I+R=0.8, SEM 0.3 kPa increase). For all other variables and cases, there is not sufficient evidence to conclude that the effects of I and R are not additive. We conclude that R and I exercises do not invariably produce strictly additive cardiopulmonary responses. When R and I exercises are started simultaneously, however,P _as,f _c, andV _I are additive. Finally, cardiopulmonary responses to I efforts superimposed on steady-state 75%VO_2max R efforts suggest that cardiopulmonary controls already established during R efforts attenuate responses to the I effort.     

Sex differences in the sympatho-adrenal response to isometric exercise

Summary The effects on heart rate, blood pressure, and plasma catecholamines (epinephrine, norepinephrine, dopamine) of sustained isometric contraction (SIC) were studied in six women and nine men. Each subject held a tension equivalent to 30% of maximal handgrip strength until exhaustion. There were no significant differences between women and men in the duration of handgrip. Rise of heart rate and blood pressure were similar for women and men. Considering the absolute plasma levels of each catecholamine, no sex differences was observed at rest and at any time during SIC, except for epinephrine whose concentration was higher in men at first min of SIC. On the other hand, women and men exhibited different adrenergic patterns in response to SIC: in the first min of exercise the plasma level of the three catecholamines increased in men whereas for women plasma catecholamines levels were essentially unaffected. Thus, epinephrine seems to play a minor role in the regulation of heart rate and blood pressure during SIC for women. Another interesting result of our study is that SIC is able to induce an increase in dopamine plasma level for women as well as for men.     

Muscle afferent contributions to the cardiovascular response to isometric exercise

The cardiovascular response to isometric exercise is governed by both central and peripheral mechanisms. Both metabolic and mechanical stresses on the exercising skeletal muscle produce cardiovascular change, yet it is often overlooked that the afferent signal arising from the muscle can be modified by factors other than exercise intensity. This review discusses research revealing that muscle fibre type, muscle mass and training status are important factors in modifying this peripheral feedback from the active muscles. Studies in both animals and humans have shown that the pressor response resulting from exercise of muscle with a faster contractile character and isomyosin content is greater than that from a muscle of slower contractile character. Athletic groups participating in training programmes that place a high anaerobic load on skeletal muscle groups show attenuated muscle afferent feedback. Similarly, longitudinal studies have shown that specific local muscle training also blunts the pressor response to isometric exercise. Thus it appears that training may decrease the metabolic stimulation of muscle afferents and in some instances chronic exposure to the products of anaerobic metabolism may blunt the sensitivity of the muscle metaboreflex. There may be surprising parallels between the local muscle conditions induced in athletes training for longer sprint events (e.g. 400 m) and by the low-flow conditions in, for example, the muscles of chronic heart failure patients. Whether their similar attenuations in muscle afferent feedback during exercise are due to decreased metabolite accumulation or to a desensitization of the muscle afferents is not yet known.     

Influence of central command and ergoreceptors on the splanchnic circulation during isometric exercise

The splanchnic circulation can make a major contribution to blood flow changes. However, the role of the splanchnic circulation in the reflex adjustments to the blood pressure increase during isometric exercise is not well documented. The central command and the muscle chemoreflex are the two major mechanisms involved in the blood pressure response to isometric exercise. This study aimed to examine the behaviour of the superior mesenteric artery during isometric handgrip (IHG) at 30% maximal voluntary contraction (MVC). The pulsatility index (PI) of the blood velocity waveform of the superior mesenteric artery was taken as the study parameter. A total of ten healthy subjects [mean age, 21.1 (SEM 0.3) years] performed an IHG at 30% MVC for 90 s. At 5 s prior to the end of the exercise, muscle circulation was arrested for 90 s to study the effect of the muscle chemoreflex (post exercise arterial occlusion, PEAO). The IHG at 30% MVC caused a decrease in superior mesenteric artery PI, from 4.84 (SEM 1.57) at control level to 3.90 (SEM 1.07) (P = 0.015). The PI further decreased to 3.17 (SEM 0.70) (P = 0.01) during PEAO. Our results indicated that ergoreceptors may be involved in the superior mesenteric artery vasodilatation during isometric exercise.     

Effect of activated sweat glands on the intensity-dependent sweating response to sustained static exercise in mildly heated humans

Changes in the number of activated sweat glands (ASGs) and sweat output per gland (SGO) with increased exercise intensity during sustained static exercise were investigated. Fourteen male subjects performed 20, 35, and 50% maximal voluntary contraction (MVC) for 60 s with the right hand (exercised arm) at an ambient temperature of 35 degrees C and 50% relative humidity. Although sublingual, local skin, and mean skin temperatures remained essentially constant throughout the exercise at each intensity, the sweating rate (SR) of nonglabrous skin on the nonexercised left forearm increased significantly with a rise in exercise intensity (p<0.05). Changes in the number of ASGs with rising exercise intensity paralleled changes in the SR, but the SGO did not change markedly with altered exercise intensity. These results suggest that in mildly heated humans, at less than 50% MVC, the increase in the SR from nonglabrous skin with rising exercise intensity during sustained static exercise is dependent on changes in the number of ASGs and not on SGO.     

Effects of varying central command and muscle mass on the cardiovascular responses to isometric exercise.

To determine if the central command signal associated with isometric exercise is mass-dependent, 20 subjects (nine male, 11 female; 23 +/- 1 years) performed four 5-min bouts of supine isometric exercise with a large (quadriceps; LEG) and small (forearm; ARM) muscle mass. For each extremity, one bout entailed maintaining a constant force (CF; 20% maximal voluntary contraction) and the other constant electromyographic activity (CE; approximately 20% MVC initially). Central command was assumed to increase with CF and remain unchanged with CE. Heart rate increased more with LEG than ARM (P<0.001) and, in LEG, was higher in CF than CE at min 5 (P<0.001). Mean arterial pressure was higher in LEG (P<0.001) by min 2 and 10 +/- 3 mmHg higher in LEG CF than LEG CE by min 5 (P<0.001). Ratings of perceived exertion were highest in LEG CF (P<0.001); LEG CE did not differ from ARM CE (P<0.001) by min 4. The ARM responses did not differ between CF and CE in any variable. These data suggest that muscle mass influences the central command signal during isometric exercise and central command modulates this response in larger muscle masses.     

FPA-based infrared thermography as applied to the study of cutaneous perspiration and stimulated vascular response in humans

This review gives an overview of focal plane array (FPA)-based infrared (IR) thermography as a powerful research method in the field of physiology and medicine. Comparison of the gained results with the data previously obtained by other authors with other research tools is given. Outer thermoregulatory manifestations displayed by the human organism subjected to whole-body heating (sauna bath) and physical loads (exercise bicycling) are quantitatively analysed. Some details of human body emotional sweating (psycho-physiological effect) are reported. Particular attention is paid to studying active sweat glands as individual objects. All experimental data were obtained with the help of a high-sensitivity (0.03 degrees C) fast 128 x 128 InAs IR detector-based thermal imaging system operating in the short-wave spectral region (2.5 to 3 microm) and perfectly suiting medical purposes. It is shown that IR thermography makes it possible to overcome limitations inherent to contact measuring means that were traditionally used before in thermal studies. It is also shown that heterogeneous thermograms displayed by organisms with disturbed inner equilibrium can be quantitatively analysed in terms of statistical parameters of related surface-temperature histograms, such as the mean temperature and the standard deviation of temperature (SDT). The increase and the decrease in SDT turned out to be typical of prolonged physical load and subsequent relaxation, and of external whole-body heating, respectively. Explanation of this result based on a hypothesis advanced within the context of the doctrine of human-organism evolution is given. Skin-temperature distribution function accompanying the relaxed organism in normality was found to closely resemble normal-distribution function. Symmetry break down and variation of the shape of this characteristic may serve as an indicator of homeostasis shift and can be used as a quantitative criterion for the latter. A new phenomenon, stable punctate hidrosis, is discovered and described. The term sweatology is introduced to refer to the discussed specific research area in biomedical science.     

Cardiovascular and respiratory responses to changes in central command during isometric exercise at constant muscle tension

1. Experiments were designed to show whether elements of the command descending from higher centres to exercising muscles provide an input for cardiovascular and respiratory control. Vibration, known to be a powerful stimulus to the primary afferents from muscle spindles, was applied to the biceps tendon of human subjects performing sustained isometric contractions with the biceps or the triceps muscle. When the biceps was contracting this activation of muscle spindle primary afferents in it provided an element of reflex excitation, so that less central command was required to achieve a given tension. When triceps was contracting, the activation of muscle spindle primary afferents in its antagonist, biceps, contributed an element of reflex inhibition, so that more central command than normally was required to achieve a given tension. The cardiovascular and respiratory responses to an isometric effort could thus be investigated at any tension when the central command was normal, decreased, or increased.2. Blood pressure, heart rate, and pulmonary ventilation all increase in an isometric effort. The increase in each is less when the central command is reduced. The increase in each is greater when the central command is increased.3. It is concluded that there is irradiation of cardiovascular and respiratory control centres by the descending central command during voluntary muscular contractions in man.     

Physiological unloading of cardiopulmonary mechanoreceptors by posture changes does not influence the pressor response to isometric exercise in healthy humans

Summary In recent studies in humans the role of cardiopulmonary baroreflexes in modulating the cardiovascular responses to isometric exercise (somatic pressor reflex) has been investigated by performing static hand-grip exercise during deactivation of cardiopulmonary receptors produced by low levels of lower body negative pressure; however, findings from these studies have not been consistent. The purpose of this study was to investigate whether a more physiological unloading stimulus of cardiopulmonary baroreceptors, obtained by sequentially changing posture, could influence the pressor response to somatic afferent stimulation induced by isometric, exercise. To accomplish this, ten healthy subjects performed a 2-min isometric handgrip (IHG) at 30% maximal voluntary contraction after 10 min of supine rest and, in rapid sequence, after 10 min of sitting and 10 min of standing, at the time when, owing to their transitory nature, the cardiovascular effects, due to arterial baroreceptor intervention should have been minimal. During IHG arterial pressure (BP_a) was continuously and noninvasively measured to quantify accurately the blood pressure response to IHG both in magnitude and time course. Results showed that the pressor response to IHG was not significantly influenced by change in posture, either in magnitude or in time course. The mean arterial pressure increased by 17.4 (SEM 2.5), 18.6 (SEM 1.2) and 17.0 (SEM 1.3) mmHg in supine, sitting and standing [2.3 (SEM 0.3), 2.5 (SEM 0.2) and 2.3 (SEM 0.2) kPa] positions, respectively. Also the heart rate response to IHG was unaffected by change in posture. Most important, the sum of the separate BP_a responses induced by supine IHG and by posture change from supine to sitting (summation of reflexes) was not significantly different from the pressor response observed during sitting IHG (interaction of reflexes). Likewise, the sum of the separate BP_a. responses induced by sitting IHG and by changing postures from sitting to standing was not significantly different from the pressor response to standing IHG. These data indicate that, under physiological conditions, cardiopulmonary baroreflexes do not exert a significant role in modulating the reflex pressor drive from muscles during isometric exercise in healthy humans.     

Decreased muscle sympathetic nerve activity does not explain increased vascular conductance during contralateral isometric exercise in humans

At the onset of both electrically evoked (STIM) and voluntary (VOL) isometric calf exercise there is an increase in vascular conductance of the contralateral lower limb, suggesting withdrawal of muscle sympathetic nerve activity (MSNA). Seven subjects performed STIM or VOL ischaemic calf exercise at 30% maximum voluntary contraction in a seated position. Blood pressure, heart rate and peroneal MSNA in the resting contralateral lower limb were recorded. During both STIM and VOL exercise blood pressure increased (P < 0.05). Blood flow increased by 40 +/- 3 and 35 +/- 3% and conductance increased by 37 +/- 3 and 31 +/- 4% (P < 0.05) after 10 s of STIM and VOL, respectively, and thereafter declined. The time course and direction of these changes persisted with subjects in a semisupine position, confirming that the transient conductance changes were not an artefact of the dependent leg position. Thigh cuff inflation for 1 min without exercise caused a 47 +/- 7.5% (P < 0.05) reduction in MSNA, which recovered when the circulation was restored. However, when cuff inflation was followed by STIM or VOL exercise, MSNA did not fall further. These data suggest that the transient increase in vascular conductance at the onset of contralateral electrically evoked or voluntary lower limb exercise is unrelated to MSNA.     

Identifying cardiorespiratory neurocircuitry involved in central command during exercise in humans

For almost one hundred years, the exact role of human brain structures controlling the cardiorespiratory response to exercise ('central command') has been sought. Animal experiments and functional imaging studies have provided clues, but the underlying electrophysiological activity of proposed relevant neural sites in humans has never been measured. In this study, local field potentials were directly recorded in a number of 'deep' brain nuclei during an exercise task designed to dissociate the exercise from peripheral feedback mechanisms. Several patient groups had electrodes implanted sterotaxically for the treatment of movement disorder or chronic pain. Fast Fourier transform analysis was applied to the neurograms to identify the power of fundamental spectral frequencies. Anticipation of exercise resulted in increases in heart rate, blood pressure and ventilation. The greatest neural changes were found in the periaqueductal grey area (PAG) where anticipation of exercise was accompanied by an increase of 43% in the power of the 12–25 Hz frequency band ( P = 0.007). Exercise increased the activity by 87% compared to rest ( P = 0.006). Changes were also seen in the 60–90 Hz band when anticipation or exercise increased power by 32% ( P = 0.006) and 109% ( P < 0.001), respectively. In the subthalamic nucleus there was a reduction in the power of the beta frequency during both anticipation (7.6 ± 0.68% P = 0.001) and exercise (17.3 ± 0.96% P < 0.001), whereas an increase was seen with exercise only at higher frequencies (93 ± 1.8% P = 0.007). No significant changes were seen in the globus pallidus during anticipation of exercise. We provide direct electrophysiological evidence highlighting the PAG as an important subcortical area in the neural circuitry of the cardiorespiratory response to exercise, since stimulation of this structure is known to alter blood pressure in awake humans.     

The interaction of central command and the exercise pressor reflex in mediating baroreflex resetting during exercise in humans

Central command and the exercise pressor reflex can independently reset the carotid baroreflex (CBR) during exercise. The present investigation assessed the interactive relationship between these two neural mechanisms in mediating baroreflex resetting during exercise. Six men performed static leg exercise at 20% maximal voluntary contraction under four conditions: control, no perturbation; neuromuscular blockade (NMB) induced by administration of the neuromuscular blocking agent Norcuron (central command activation); MAST, application of medical antishock trousers inflated to 100 mmHg (exercise pressor reflex activation); and Combo, NMB plus MAST (concomitant central command and exercise pressor reflex activation). With regard to CBR control of heart rate (HR), both NMB and Combo conditions resulted in a further resetting of the carotid-cardiac stimulus-response curve compared to control conditions, suggesting that CBR-HR resetting is predominately mediated by central command. In contrast, it appears that CBR control of blood pressure can be mediated by signals from either central command or the exercise pressor reflex, since both NMB and MAST conditions equally augmented the resetting of the carotid-vasomotor stimulus-response curve. With regard to the regulation of both HR and blood pressure, the extent of CBR resetting was greater during the Combo condition than during overactivation of either central command or the exercise pressor reflex alone. Therefore, we suggest that central command and the exercise pressor reflex interact such that signals from one input facilitate signals from the other, resulting in an enhanced resetting of the baroreflex during exercise.     

The influence of exercise/rest schedule on the physiological and psychophysical response to isometric shoulder-neck exercise

Six female subjects, aged 24-34 years, performed shoulder-neck exercise for 1 h or until they were exhausted by holding out their arms horizontally at 60° to the sagittal plane. One continuous and six intermittent protocols were applied, all with a mean load corresponding to the torque of the arms, i.e. about 15% maximal voluntary contraction (MVC). The intermittent protocols varied according to cycle time (10 s, 60 s, 360 s) and duty cycle (0.33, 0.50, 0.67, 0.83). Electromyogram (EMG), mean arterial blood pressure ( _a), heart rate (f _c) and perceived fatigue were monitored at regular intervals during exercise. Blood concentrations of potassium, lactate and ammonia were determined in pre- and postexercise samples of venous blood. Before and up to 4 h after exercise, measurements were made of MVC, pressure pain threshold, proprioceptive performance, and of EMG, _a and f _c during 1-min arm-holding at 25% MVC. Endurance times ranged from about 10 min to more than 1 h, significantly relating to both cycle time and duty cycle. The _a, f _c EMG amplitude and perceived fatigue increased early during all protocols and continued to increase throughout the exercise period. Duty cycle influenced all of these variables, while only _a and fatigue perception were related to cycle time. Cardiovascular and neuromuscular recovery was incomplete for hours after several of the protocols, as indicated for example by a sensitizised response to the 1-min armholding. The protocols differed substantially as regards the relationship between different responses. Thus, ranking of the protocols in terms of physiological strain was different, depending on the criterion variable. The result stresses the relevance of applying a comprehensive selection of variables when evaluating the responses to intermittent shoulder-neck exercise.     

Role of central command in carotid baroreflex resetting in humans during static exercise

The purpose of the experiments was to examine the role of central command in the exercise-induced resetting of the carotid baroreflex. Eight subjects performed 30 % maximal voluntary contraction (MVC) static knee extension and flexion with manipulation of central command (CC) by patellar tendon vibration (PTV). The same subjects also performed static knee extension and flexion exercise without PTV at a force development that elicited the same ratings of perceived exertion (RPE) as those observed during exercise with PTV in order to assess involvement of the exercise pressor reflex. Carotid baroreflex (CBR) function curves were modelled from the heart rate (HR) and mean arterial pressure (MAP) responses to rapid changes in neck pressure and suction during steady state static exercise. Knee extension exercise with PTV (decreased CC activation) reset the CBR-HR and CBR-MAP to a lower operating pressure ( P < 0.05) and knee flexion exercise with PTV (increased CC activation) reset the CBR-HR and CBR-MAP to a higher operating pressure ( P < 0.05). Comparison between knee extension and flexion exercise at the same RPE with and without PTV found no difference in the resetting of the CBR-HR function curves ( P > 0.05) suggesting the response was determined primarily by CC activation. However, the CBR-MAP function curves were reset to operating pressures determined by both exercise pressor reflex (EPR) and central command activation. Thus the physiological response to exercise requires CC activation to reset the carotid-cardiac reflex but requires either CC or EPR to reset the carotid-vasomotor reflex.