Measurement of forearm blood flow by venous occlusion plethysmography: Influence of hand blood flow during sustained and intermittent isometric exercise

Summary The requirement for using an arterial occlusion cuff at the wrist when measuring forearm blood flows by plethysmography was tested on a total of 8 subjects at rest and during and after sustained and intermittent isometric exercise. The contribution of the venous effluent from the hand to the forearm flow during exercise was challenged by immersing the arm in water at 20, 34, and 40 C. Occlusion of the circulation to the hand reduced the blood flow through the resting forearm at all water temperatures. There was an inverse relationship between the temperature of the water and the proportion in the reduction of forearm blood flow upon inflation of the wrist-cuff, ranging from 45 to 19% at 20 to 40 C, respectively. However, during sustained isometric exercise at 10% of the subjects maximum voluntary contraction (MVC) there was no reduction in the measured forearm flow when an arterial occlusion cuff was inflated around the wrist. Similarly, there was no alteration in the blood flow measured 2 s after each of a series of intermittent isometric contractions exerted at 20% or 60% MVC for 2 s whether or not circulation to the hand was occluded nor of the post-exercise hyperemia following 1 min of sustained contraction at 40% MVC. These results indicate that a wrist-cuff is not required for accurate measurement of forearm blood flows during or after isometric exercise.This work was supported by N.I.H. training grant HLO 7050-03, H.E.W. contract 210-77-0044 and Air Force grant AFOSR-76-3084 B     

Physiological response in the forearm during and after isometric intermittent handgrip

Summary The aim of the present paper was to study the development of fatigue during isometric intermittent handgrip exercise. Using a handgrip dynamometer, four combinations of contraction-relaxation periods were studied (10+10, 10+5, 10+2s and continuous contraction) at three contraction intensities (10, 25 and 40% maximum voluntary contraction, MVC). Local blood flow (BF) in the forearm (venous occlusion plethysmography) was followed before, during and after the exercise period. Electromyography (EMG) (frequency analysis) and the perceived effort and pain were recorded during the exercise period. Forearm BF is insufficient even at isometric contractions of low intensity (10% MVC). The results indicate that vasodilating metabolites play an active role for BF in low-intensity isometric contractions. It is shown that maximal BF in the forearm during relaxation periods (25–30 ml min^–1 · 100 ml^–1) is already reached at 25% MVC. Only intermittent exercise at 10% MVC and (10+5s) and (10+10s) at 25% MVC was considered acceptable with regard to local fatigue, which was defined as a switch of local BF to the post-exercise period, a decrease in the number of zero-crossings (EMG) and marked increases in subjective ratings.     

Tissue oxygenation by near-infrared spectroscopy and muscle blood flow during isometric contractions of the forearm.

The relationship between tissue oxygenation measured by near-infrared spectroscopy (NIRS) and forearm muscle blood flow (FBF) measured by Doppler ultrasound was tested during isometric contractions at 10 and 30% maximal voluntary contraction (MVC) under conditions of normoxia and hypoxia (14% inspired O2). Six subjects maintained contractions at 10% MVC for 5 min and 30% for 2 min in both gas conditions. FBF was elevated during exercise at 10% MVC in hypoxia compared to normoxia, but there was no further increase in flow at 30% MVC. Median power frequency calculations from electromyographic recordings suggested progressive development of fatigue throughout both 10 and 30% MVC contractions. NIRS indicated no change in muscle oxygenation at 10% MVC, but deep venous blood O2 saturation was reduced in normoxia and more so in hypoxia. At 30% MVC, both NIRS and venous O2 saturation were reduced, with no effect of hypoxia on the NIRS signal. While NIRS might provide an indication of muscle oxygenation during isometric exercise, the conflicting findings for NIRS and direct venous blood sampling at 10 vs 30% MVC suggest caution in the application of this noninvasive technique.     

Plasma potassium concentration and Doppler blood flow during and following submaximal handgrip contractions

The aim of the present study was to investigate the time-course of blood velocity in the forearm during and folllowing isometric handgrip contractions and to reveal a possible temporal relationship between the circulatory response and venous effluent potassium concentration ([K]) not only during contractions but also during the post-exercise recovery period. Contractions of 15% maximal voluntary contraction (MVC) and 30% MVC with and without 3 min of artirial occlusion following the contractions were studied. All contractions induced a significant increase in venous plasma [K] from an average resting level of 4.0 to 5.0 mM during 15% MVC and 5.8 mM during 30% MVC. Blood velocity increased from a resting level of 0.07 to 0.22 m s^-1 diromg 15% and 30% MVC, respectively. MCC of 30% always elicited a larger blood velocity and [K] response than 15% MVC. Following the contractions hyperaemia was elicited. Recovery of the local blood velocity was markedly slower than the K recovery, since [K] remained significantly above resting level for only 25 s following 15% MVC and 45 s following 15 and 30% MVC, respectively. Further, a larger hyperaemia following the occlusion was elicited as compared to the contraction without occlusion, in spite of [K] being lower immediately after the occlusion period than immediately after the contraction. Finally, [K] decreased below resting level in the recovery period while the blood velocity remained elebvated. Therefore, the present study showed that the venous plasma [K] is not causally related to the prolonged post-exercise hyperaemia. The skin temperature remained unchanged during the contractions, while during the recovery period the skin temperature increased for several minutes. The major part of the temperature increase was likely to be due to conductance of heat from muscles to skin surface as a consequence of muscle hyperaemia.     

Pain and changes in peripheral resistance at high vascular transmural pressure in the human forearm

A hydrostatic rise in forearm vascular transmural pressure may be associated with an increase in forearm blood flow (FBF) that causes pain. To test this hypothesis, forearm vascular transmural pressure was elevated in eight male volunteers by a series of 1-min hypobaric exposures of the left arm to incrementing differential pressures of 40, 80, 120, 140, 160 and 200 mmHg. The series was repeated after a 30-min interval. Forearm venous pressure (FVP) was measured in the median antecubital vein, and FBF was determined by ultrasound Doppler in the axillary artery. Pain level was recorded by numerical rating scale. In all subjects, an increase in FBF and forearm vascular conductance (FVC) occurred (P < 0.05) at high FVP (mean ± SE, 184 ± 8 mmHg). Pain was linearly related to the increase in FVC. In the second series of exposures, increases in FBF, FVC and pain occurred at a lower transmural pressure (FVP 152 ± 15 mmHg, P < 0.01). It is concluded that intense forearm pain is associated with a failure of autoregulation in the peripheral vascular bed and is worsened on repeated exposure to high transmural pressure. This may explain the overt forearm pain experienced by the crews of high performance military aircraft during manoeuvring.     

The exercise pressor response to sustained handgrip does not augment blood flow in the contracting forearm skeletal muscle

Previous studies have advanced the concept that during sustained handgrip (SHG) reflex increases in blood pressure are able to partially offset increases in tissue pressure and thus effectively maintain increases in muscle blood flow during mild to moderate levels of sustained handgrip. However, this concept is based upon measurements of blood flow to the entire forearm. The aim of this study was to evaluate this concept by simultaneously measuring time-dependent changes in systemic arterial pressure and blood flow in an active muscle during the actual period of exercise. To accomplish this aim, we measured ¹³³Xenon washout from the extensor carpi radialis longus muscle over 3 min of SHG at 15, 30 and 45% of maximal voluntary contraction (MVC). During sustained handgrip at 15% MVC, muscle blood flow increased more than 20 fold from rest to exercise (P < 0.05), even though mean arterial pressure increased by only 12 ± 4 mmHg. This large exercise-induced hyperaemia was abolished during SHG at both 30 and 45% MVC, despite large and progressive increases in mean arterial pressure of 29 ± 3 and 54 ± 5 mmHg, respectively. We conclude that at levels of handgrip above 15% MVC blood pressure ceases to be an important determinant of blood flow in the active skeletal muscle. Importantly, the increases in forearm blood flow that have been reported previously with such levels of static handgrip do not appear to be directed to the most active muscle.     

Muscle blood flow during isometric activity and its relation to muscle fatigue

Summary The effect of isometric exercise on blood flow, blood pressure, intramuscular pressure as well as lactate and potassium efflux from exercising muscle was examined. The contractions performed were continuous or intermittent (5 s on, 5 s off) and varied between 5% and 50% maximal voluntary contraction (MVC). A knee-extensor and a hand-grip protocol were used. Evidence is presented that blood flow through the muscle is sufficient during low-level sustained contractions (<10% MVC). Despite this muscle fatigue occurs during prolonged contractions. One mechanism for this fatigue may be the disturbance of the potassium homeostasis. Such changes may also play a role in the development of fatigue during intermittent isometric contractions and even more so in the recovery from such exercise. In addition the role of impaired transport of substances within the muscle, due to longlasting daily oedema formation, is discussed in relation to fatigue in highly repetitive, monotonous jobs.     

Muscle weakness following sustained and rhythmic isometric contractions in man

Summary The effects of sustained and rhythmically performed isometric contractions on electrically evoked twitch and tetanic force generation of the triceps surae have been investigated in 4 healthy male subjects. The isometric contractions were performed separately and on different occasions at 30%, 60% and 100% of the force of maximal voluntary contraction (MVC). The area under the maximal voluntary contraction (MVC) force/ time curve during the rhythmic and sustained contractions was the same for each experiment. The results showed that following rhythmic isometric exercise there was a small decrease in low (10 and 20 Hz) and high (40 Hz) frequency tetanic tension which was associated with % MVC. However, there was no change in the 20/40 ratio of tetanic forces, MVC or the contraction times and force of the maximal twitch. In contrast, following sustained isometric exercise tetanic forces were markedly reduced, particularly at low frequencies of stimulation. The 20/40 ratio decreased and the induced muscle weakness was greater at 30% than 60% or 100% MVC. The performance of sustained isometric contractions also effected a decrease in contraction time of the twitch and MVC. The results are in accord with previous findings for dynamic work (Davies and White 1982), and show that if isometric exercise is performed rhythmically the effect on tetanic tensions is small and there is no evidence of a preferential loss of electrically evoked force at either high or low frequencies of stimulation following the contractions. For sustained contractions, however, the opposite is true, the ratio of 20/40 Hz forces is markedly reduced and following 30% sustained MVC there is a significant (p<0.05) change in the time to peak tension (TPT) of the maximal twitch.     

Relationship between cortical alpha and skeletal muscle blood flow in a feedback task

During operant control of forearm blood flow (FBF) cortical alpha was recorded. A significant inverse relationship was found between FBF and alpha activity. Decreased FBF was associated with increased alpha. One possible reason that several investigators found spurious relationships between alpha and 2 other cardiovascular parameters, heart rate and blood pressure, is that the range of response for HR and BF is relatively restricted compared to that of alpha.     

The influence of temperature on the amplitude and frequency components of the EMG during brief and sustained isometric contractions

Summary The influence of temperature on the amplitude and frequency components of the EMG power spectra of the surface EMG recorded over the forearm muscles was examined in five male and five female subjects during brief and fatiguing isometric contractions of their handgrip muscles. Brief (3 s) isometric contractions were exerted at tensions ranging between 10 and 100% of each subject's maximum strength while fatiguing contractions were exerted at tensions of 25, 40, and 70% of their maximum strength. The temperature of the muscles during those contractions was varied by placing the forearms of the subjects in a controlled temperature water bath at temperatures of 10, 20, 30, and 40 C. The results of these experiments showed that the center frequency of the power spectra of the surface EMG was directly related to the temperature of the exercising muscles during brief isometric contractions. During fatiguing isometric contractions, the amplitude of the EMG increased while the center frequency of the EMG power spectra decreased for all tensions examined.     

Blood flow in the triceps brachii muscle in humans during sustained submaximal isometric contractions

The main purpose of this study was to determine the extent to which blood flow through the profunda artery within the triceps brachii muscle may be compromised during maintained low-force isometric fatiguing contractions. Doppler ultrasound techniques were used to record mean blood velocity and arterial diameter of the profunda brachii artery during sustained isometric contractions of 20% maximal voluntary contraction. The arterial diameter did not change throughout the contraction. Thus, blood velocity was considered to be an indicator of blood flow. The mean blood velocity increased initially and then remained constant during the contraction period. When compared to rest [0.06 (SD 0.03) m s^–1] mean blood velocity was significantly larger at the start of the contraction [0.13 (SD 0.07) m s^–1] and larger yet during recovery following the contraction [0.30 (SD 0.14) m s^–1]. Although blood flow through the conduit artery did not drop during the contraction, the post-contraction hyperaemia suggested that circulatory compromise might have occurred at the level of the capillary beds. Electronic Publication     

Effect of hypoxia on arterial and venous blood levels of oxygen,carbon dioxide,hydrogen ions and lactate during incremental forearm exercise

Summary The purpose of the present study was to investigate whether, in humans, hypoxia results in an elevated lactate production from exercising skeletal muscle. Under conditions of both hypoxia [inspired oxygen fraction (F_IO₂): 11.10%] and normoxia (F_IO₂: 20.94%), incremental exercise of a forearm was performed. The exercise intensity was increased every minute by 1.6 kg·m·min^–1 until exhaustion. During the incremental exercise the partial pressure of oxygen (PO₂) and carbon dioxide (PCO₂), oxygen saturation (SO₂), pH and lactate concentration [HLa] of five subjects, were measured repeatedly in blood from the brachial artery and deep veins from muscles in the forearm of both the active and inactive sides. The hypoxia (arterial SO₂ approximately 70%) resulted in (1) the difference in [HLa] in venous blood from active muscle (values during exercise — resting value) often being more than twice that for normoxia, (2) a significantly greater difference in venous-arterial (v-a) [HLa] for the exercising muscle compared to normoxia, and (3) a difference in v-a [HLa] for non-exercising muscle that was slightly negative during normoxia and more so with hypoxia. These studies suggest that lower O₂ availability to the exercising muscle results in increased lactate production.     

Cardiovascular responses to heat stress and blood volume displacements during exercise in man

Summary Subjects exercised in the upright position at approximately 50% of maximal oxygen consumption in four situations: in 25 C air, in 45 C air [mean skin temperature ( _sk) 35 C], in 35 C water immersed to the level of the xiphoid process, and finally wearing a suit perfused with 35 C water. The water immersion prevented gravitational shifts of blood volume to the legs. In this situation the forearm blood flow (FBF) rose continually with increasing core temperature (T_es) in contrast to the attenuation in rise above 38 C T_es in 45 C air. The differences were significant above 38.6 C T_es in experiments in eight subjects. The effects of immersion on cardiac output (CO), stroke volume (SV), and heart rate (HR) were studied in five of the subjects in relation to T_es, since the rate of rise of T_es was different in the four situations. CO and SV tended to be higher during both rest and exercise in the water than in the other three conditions, while HR rose in the same manner with increasing core temperature, except that it was lower in 25 C air, where T_es was lower. Thus, the prevention of hydrostatic shifts of peripheral venous volume permitted the maintenance of a higher SV and peripheral blood flow, and enhanced the ability of the circulation to deal with the combined exercise and heat stress.Supported by the Danish Space Board (1112-32/81), the Danish Medical Research Council (512–15983), and the Danish Sports Research Council. Dr. Rowell was supported in part by NHLBI Grant HL 16910     

Effects of dynamic leg exercise on subcutaneous blood flow rate in the lower limb of man

Subcutaneous blood flow (SBF) was studied simultaneously in the upper arm at heart level and in the lower limb during positional changes and during leg exercise in seven healthy males. SBF was estimated by local clearance of ‘“Xenon registered by portable cadmium telluride detectors. Venous pressure was recorded directly on dorsum on the foot. Changinr the position from supine to head-up tilt, SBF decreased by 43 % (P < 0.01) at the arm level, 40% at the thigh (P < 0.01), 47% at the calf (P < 0.01) and decreased by 51 % at the ankle level (P < 0.01). Performing 20 heel-raisings per min in nearly erect posture, SBF increased by 96% at the thigh (P < 0.01), 25% at the calf (P > 0.1) and increased by 18% at the ankle level (P > 0.1). At 40 heel-raisings per min SBF increased by 99% at the thigh (P < 0.0 1), 121 % at the calf (P < 0.0 1), but only 44% at the ankle level (P > 0.1). During leg exercise subcutaneous vascular resistance was significantly increased at arm and ankle levels. In contrast, a vasodilatory response was noticed at the thigh and calf levels and seemed associated with a decrease in local venous pressure to below the trigger level of the sympathetic veno-arteriolar reflex mechanism. In conclusion, SBF in the lower limb of man was increased during exercise. The increase in SBF could only partly be ascribed to the concomitant increase in perfusion pressure. The local blood flow response seemed modified by changes in sympathetic nervous activity and metabolic rate.     

Effects of dynamic leg exercise on subcutaneous blood flow rate in the lower limb of man

Subcutaneous blood flow (SBF) was studied simultaneously in the upper arm at heart level and in the lower limb during positional changes and during leg exercise in seven healthy males. SBF was estimated by local clearance of ¹³³Xenon registered by portable cadmium telluride detectors. Venous pressure was recorded directly on dorsum on the foot. Changing the position from supine to head-up tilt, SBF decreased by 43% (P < 0.01) at the arm level, 40% at the thigh (P < 0.01), 47% at the calf (P < 0.01) and decreased by 51 % at the ankle level (P < 0.01). Performing 20 heel-raisings per min in nearly erect posture, SBF increased by 96% at the thigh (P < 0.01), 25% at the calf (P > 0.1) and increased by 18% at the ankle level (P > 0.1). At 40 heel-raisings per min SBF increased by 99% at the thigh (P < 0.01), 121% at the calf (P < 0.01), but only 44% at the ankle level (P > 0.1). During leg exercise subcutaneous vascular resistance was significantly increased at arm and ankle levels. In contrast, a vasodilatory response was noticed at the thigh and calf levels and seemed associated with a decrease in local venous pressure to below the trigger level of the sympathetic veno-arteriolar reflex mechanism. In conclusion, SBF in the lower limb of man was increased during exercise. The increase in SBF could only partly be ascribed to the concomitant increase in perfusion pressure. The local blood flow response seemed modified by changes in sympathetic nervous activity and metabolic rate.     

Leg blood flow during static exercise

Summary Leg blood flow was studied with the constant infusion dye technique during static exercise of the thigh muscles (quadriceps) and during hand-grips at 15 and 25–30% of MVC.Blood flow and oxygen uptake in the leg increased in quadriceps exercise and reached their highest values (around 1.2 l/min and 165 ml/min respectively) at 25–30% of MVC, whereas leg vascular resistance decreased. Regional circulatory adaptations and the oxygen uptake — leg blood flow relationship were in close agreement with the responses found in dynamic leg exercise. In view of the marked rise in intramuscular pressure previously observed during quadriceps contractions, a restriction of blood flow and an increased vascular resistance had been expected. Involuntary activation of leg muscles other than the quadriceps may explain the finding.Contractions of the contralateral quadriceps induced a slight increase in leg blood flow, whereas hand-grips had no influence on blood flow or vascular resistance in the leg. The distribution of the cardiac output during static contractions is discussed, and it is concluded that during hand-grips the increase in blood flow is predominantly distributed to the upper part of the body.     

Autonomic nervous control of the heart rate during isometric exercise in normal man

The relative contribution of the efferent components of the autonomic nervous system to the regulation of tachycardia induced by isometric exercise was assessed in 23 normal males. The isometric exercise (handgrip) was performed at the maximum intensity tolerated by the individual over a period of 10 s (maximal voluntary contraction — MVC) and at levels equivalent to 75, 50 and 25% of MVC for 20, 40 and 10 s, respectively. The study was performed both under control conditions and after pharmacological blockade with atropine (12 individuals) or propranolol (11 individuals). Under control conditions, the heart rate (HR) responses to isometric effort were dependent on the intensity and duration of the exercise, showing a tendency towards progressive elevation with the maintenance of muscular contraction at the levels studied. The tachycardia evoked by this effort was of considerable magnitude and of rapid onset, especially at the more intense levels of activity. Parasympathetic blockade markedly decreased tachycardia, which manifested itself during the first 10 s of exercise at all levels of intensity, whereas sympathetic blockade markedly modified the HR response after 10 s of effort at the 75 and 50% MVC levels. A slight depression of the tachycardiac response could be observed already after 10 s of maximum effort after propranolol. The present results suggest that the autonomic regulation of these responses is based on a biphasic mechanism, with the initial phase depending on the rapid withdrawal of the parasympathetic influence, followed by a marked sympathetic contribution to the induction of tachycardia after 10 s of isometric contraction or even a little before at maximum exertion.     

Dynamic exercise in man as influenced by experimental restriction of blood flow in the working muscles

The effects of reduced muscle perfusion pressure on dynamic exercise performance and cardiovascular and respiratory functions were investigated. Eight subjects were studied during supine cycle ergometry at stepwise increasing workloads until exhaustion with and without the legs exposed to a supra-atmospheric pressure of 50 mmHg (Leg Positive Pressure, LPP), a novel and convenient means of reducing the perfusion pressure in the working muscles. In the LPP condition exercise performance was reduced by 40% which, judging from assessments of perceived exertion, was due to premature muscle fatigue, indicating local or overall underperfusion of the working muscles. At any given work load, the arterial pressure response was considerably stronger during LPP than in the control condition. LPP also caused greater increases in blood lactate concentration and pulmonary ventilation, the differences from control increasing with the work load. Furthermore, the ventilatory equivalent for O₂ at a given work load was markedly higher in the LPP than in the control condition, while exercise-induced decreases in end-tidal P_CO2 were considerably exaggerated by LPP. The augmented pressor response during flow-restricted exercise, together with the strong ventilatory response which was out of proportion to overall O₂ uptake, suggests increased activation of muscle chemoreflexes by accumulation of metabolic end products, the increased pressor response tending to reduce the local flow error in the working muscles.     

Right atrial pressure and forearm blood flow during prolonged exercise in a hot environment

Right atrial pressure (RAP) at rest is known to be reduced by an increase in skin blood flow (SkBF) in a hot environment. However, there is no clear evidence that this is so during exercise. To clarify the effect of the increase in SkBF on RAP during exercise, we measured forearm blood flow (FBF) (as an index of SkBF) and RAP continuously using a Swan-Ganz catheter in five male volunteers exercising on a cycle ergometer at 60% of peak aerobic power for 50 min in a hot environment (30°C, relative humidity 20%). Cardiac output increased from 5.5±0.21/min at rest to 17.9±1.21/min (mean±SE, P<0.01) in the first 10 min of exercise and then remained steady until the end of exercise. FBF did not change significantly during the first 5 min, but then increased from 2.7±0.5 ml/100 ml per min at rest to 10.8±1.7 ml/100 ml per min (P<0.001) by 25 min as pulmonary arterial blood temperature (T _b) rose from 37.0±0.1°C to 38.1±0.1°C (P<0.001). FBF then reached a plateau, despite a continuing increase in T _b. RAP increased significantly from 4.3±0.8 to 7.6±1.2 mm Hg (P<0.001) during the first 5 min of exercise and then gradually declined to 6.1±1.0 mm Hg by 25 min (P<0.001 vs. 5 min) and further to 5.7±1.0 mm Hg by 50 min, a value not significantly higher than at rest. This reduction in RAP during exercise was significantly correlated with the increase in FBF (r=–0.97, P<0.001) with a regression equation of RAP=–0.25×FBF+8.8. These results suggest that the decrease in RAP after 5 min exercise was caused by an increase in SkBF during exercise in a hot environment.Part of this work has been published in abstract form [FASEB J 5A1400 (1991)]