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.     

Autonomic Interactions in the Control of Heart Rate in the Monkey

Each of 5 monkeys (Macaca mulatta) was operantly conditioned to raise and to lower heart rate consistently and reliably. Following such training the animals were tested using autonomic blocking agents (methyl-atropine bromide and 1-propranolol) to characterize the autonomic mechanisms mediating such control. The results were: 1) In the undrugged animal the extent to which it decreases its heart rate over a 2048-sec period is a linear function of the baseline heart rate; 2) A linear relationship between baseline heart rate and heart rate decrease also is present within the first 128 sec; 3) There is a less consistent relationship between baseline heart rate and change in heart rate when animals must increase heart rate; 4) Vagal blockade significantly attenuates the ability of most animals to increase heart rate, primarily by reducing their ability to produce large, relatively rapid increases; 5) Sympathetic blockade significantly attenuates the ability of most animals to increase heart rate both in terms of overall changes and in terms of large, relatively rapid responses; 6) Vagal blockade very significantly attenuates the ability of all animals to slow heart rate; 7) Sympathetic blockade facilitates the ability of most animals to slow heart rate. These findings show that both branches of the autonomic nervous system participate in the operant control of heart rate. The relative role of one branch or the other in a given experiment will depend upon the baseline conditions at the time of testing, and upon the requirements—i.e., raising or lowering of heart rate.     

Autonomic nervous control of heart rate at altitude (5050 m)

To investigate possible changes in autonomic regulation of heart rate as a result of acclimatization to high altitude, indexes of autonomic nervous activity were obtained non invasively by spectrum analysis of heart rate variability on five healthy male subjects [age, 31 (SEM 2) years] during a postural change from supine to seated, both at sea level and after 1 month of exposure to an altitude of 5050 m. Heart rate fluctuations at the respiratory frequency (high frequency, HF) are mediated by the parasympathetic system whereas fluctuations at about 0.1 Hz (low frequency, LF) are due to both sympathetic and parasympathetic nervous systems. Maximal heart rate, as measured during an incremental exercise test, decreased from 184 (SEM 5) beats · min^–1 at sea level to 152 (SEM 2) beats · min^–1 at 5050 m. At sea level, the change in posture from supine to seated induced an increase in LF amplitude accompanied by an increase or a decrease in HF amplitude, whereas after 1 month at altitude the HF amplitude decreased in all subjects, with little or no change in LF amplitude. These results indicate a changed strategy of heart rate regulation after acclimatization to high altitude. At sea level, the postural change induced an increase in sympathetic activity in all subjects with different individual vagal responses, whereas at altitude the postural change induced a net decrease in vagal tone in all subjects, with little or no change in sympathetic activity. These results corroborate the reported reduced sensitivity of the heart to adrenergic drive in chronic hypoxia, which may, at least in part, explain the decreased maximal heart rate in altitude-acclimatized human subjects.     

Autonomic nervous control of heart rate during blood-flow restricted exercise in man

Summary Power spectra of instantaneous heart rate (f _c) allows the estimation of the contribution of sympathetic and parasympathetic control of f _c during steady-state conditions. The present study was designed to examine autonomic control of f _c as influenced by normal dynamic leg exercise and by ischemic leg exercise. Eight subjects performed supine cycle ergometry at 30% of their control peak work rate, with and without blood-flow restriction. Blood-flow restriction was induced by exposing the exercising legs to a supra-atmospheric pressure of 6.7 kPa (leg positive pressure; LPP). The exercise responses of arterial pressure and f _c increased (P<0.05) by LPP exposure. The exaggerated pressor response may be attributed to a chemoreflex drive originating in the ischemic muscles. Exposure to LPP during exercise also produced a significant decrease in parasympathetically mediated high frequency (HF; 0.15-1.00 Hz) fluctuation of f _c, as indicated by a decrease (P<0.05) in percent HF power compared to the control exercise level. During LPP exercise, the sympathetically mediated very low frequency (VLF; 0–0.05 Hz) fluctuation of f _c increased, as indicated by an increase (P<0.05) in percent VLF power above control exercise levels. Both LPP and control exercise conditions decreased (P<0.05) power in all frequency ranges of interest compared to their respective resting conditions. The results suggest that the increase in f _c associated with normal dynamic exercise was mediated predominantly by parasympathetic withdrawal, whereas the exaggerated f _c response during ischemic exercise resulted from a combination of cardiac sympathetic drive and parasympathetic withdrawal. The increase in sympathetic activity is attributable to a muscle chemoreflex drive, which also may have attenuated parasympathetic activity by reciprocal inhibition. Alternatively, augmented central command mediated parasympathetic withdrawal during ischemic exercise.     

Autonomic nervous control of the heart in exercising man

Effects of pharmacological denervation with propranolol (0.2 mg· kg^–1) and/or atropine (0.04 mg· kg^–1) on the control system of heart rate during exercise were studied using a frequency domain analysis, to evaluate the cardiac control function of autonomic nervous systems in man. Propranolol decreased the gain of the system in low frequency range and increased the gain slightly in middle and high frequency ranges. The phase angle of the system was advanced over the whole frequency range observed. Atropine decreased and delayed the gain and phase angle, respectively, especially in high frequency range. The combined administration of blockades significantly decreased the gain and delayed the phase angle in the whole and high frequency ranges, respectively. These findings suggest that sympathetic and parasympathetic nervous systems act nearly independently in controlling heart rate during exercise; that they act in different manners; and that the characteristics of the systems can feasibly be used to estimate clinically the cardiac control function of autonomic nerves in man.     

Effect of low-dose endurance training on heart rate variability at rest and during an incremental maximal exercise test

We evaluated the effects of low-dose endurance training on autonomic HR control. We assessed the heart rate variability (HRV) of 11 untrained male subjects (36.8 +/- 7.2 years) at rest and during an incremental maximal aerobic exercise test prior to a 7-week preparatory period and prior to and following a 14-week endurance training period, including a low to high intensity exercise session twice a week. Total (0.04-1.2 Hz), low (0.04-0.15 Hz) and high (0.15-1.2 Hz) frequency power of HRV were computed by short-time Fourier transform. The preparatory period induced no change in aerobic power or HRV. The endurance training period increased peak aerobic power by 12% (P < 0.001), decreased the HR (P < 0.01) and increased all HRV indices (P < 0.05-0.01) at absolute submaximal exercise intensities, but not at rest. In conclusion, low-dose endurance training enhanced vagal control during exercise, but did not alter resting vagal HR control.     

Effects on blood pressure and autonomic nervous system function of a 12-week exercise or exercise plus DASH-diet intervention in individuals with elevated blood pressure

Aim: Hypertension is related to abnormalities in autonomic nervous system (ANS) function, with increased sympathetic output and decreased parasympathetic tone. Lifestyle interventions are the first line of treatment in hypertension, and decreased blood pressure (BP) effects may be related to changes in ANS function. Using heart rate recovery (HRR) from exercise as an index of parasympathetic tone and plasma noradrenaline as an index of sympathetic tone, we investigated the effects of lifestyle interventions on ANS function in patients with elevated BP. Methods: Sedentary participants with elevated BP were randomly assigned to either an exercise only (N = 25), exercise plus dietary approaches to stop hypertension (DASH) diet (N = 12), or waitlist control (N = 15) 12‐week intervention. Plasma noradrenaline was measured at rest and participants performed a peak exercise test before and after the intervention. HRR was calculated as peak heart rate (HR) minus HR at 1 min post‐exercise. Results: Heart rate recovery showed a significant group by time interaction; both intervention groups showed increases in HRR from pre‐ to post‐intervention, while waitlist showed no change. Similarly, both exercise plus diet and exercise groups, but not waitlist, showed significant reductions in BP from pre‐ to post‐intervention. Linear regression revealed that BP post‐intervention was significantly predicted by change in HRR when controlling for pre‐BP, age, gender and BMI. Conclusions: Lifestyle interventions induced training‐reduced BP and altered autonomic tone, indexed by HRR. This study indicates the importance of behavioural modification in hypertension and that increased parasympathetic function is associated with success in reduction of BP.     

Computer simulation of the baroregulation in response to moderate dynamic exercise

A baroregulation model, based on a previous pulsatile non-linear multielement cardiovascular model, is extended and used to study short-term regulation mechanisms. Using this model, the responses of several cardiovascular variables to different exercise levels are simulated and compared with the experimental data reported in the literature. The impact of physiological or pathological changes on the short-term regulation of arterial pressure under the stimulus of moderate dynamic exercise is then studied. The simulation results indicate that baroreflex feedback plays a critical role in the short-term regulation of arterial pressure. When the baroreflex gain decreases to one-third of the normal value, the response of the mean arterial pressure to moderate dynamic exercise and post-exercise recovery time increases by factors of 1.7 and 2.3, respectively. Clinical data from 36 subjects (two groups: normal and hypertensive) are collected to validate the model. Computer simulations for the hypertensive group show that the elastic modulus of the arterial vessel wall is increased by 1.5 times, and peripheral resistance is increased by 1.3 times the normal value, and the baroreflex gain decreases from 0.55 (for the normal group) to 0.40. The simulation results for normal and hypertensive groups agree well with the clinical data.     

Heart rate variability during dynamic exercise in elderly males and females

It has been proposed that cardiac control is altered in the elderly. Power spectral analysis of heart rate variability (HRV) was performed on 12 male and 11 female elderly subjects (mean age 74 years) while at rest in supine and sitting positions, and at steady states during 5 min of exercise (35–95% peak oxygen consumption, V˙O_2peak). There were no differences in power, measured as a percentage of the total of the high frequency peak (HF, centred at about 0.25 Hz; 13% in males vs 12% in females), low frequency peak (LF, centred at 0.09 Hz; 25% in males and 22% in females), and very low frequency component (VLF, at 0.03 Hz; 66% in males and 69% in females) between body positions at rest. There was no difference in spectral power between male and female subjects. Total power decreased as a function of oxygen consumption during exercise, LF% did not change up to about 14 ml · kg⁻¹ · min⁻¹ (40% and 80% V˙O_2peak in males and females, respectively), then decreased towards minimal values in both genders. HF% power and central frequency increased linearly with metabolic demand, reaching higher values in male subjects than in female subjects at V˙O_2peak, while VLF% remained unchanged. Thus, the power spectra components of HRV did not reflect the changes in autonomic activity that occur at increasing exercise intensities, confirming previous findings in young subjects, and indicated similar responses in both genders. Accepted: 30 November 1999     

Identification of low and high frequency ranges for heart rate variability and blood pressure variability analyses using pharmacological autonomic blockade with atropine and propranolol in swine

Understanding autonomic nervous system functioning, which mediates behavioral and physiological responses to stress, offers great potential for assessing farm animal stress and welfare. Evaluation of heart rate variability (HRV) and blood pressure variability (BPV), using time and frequency domain analyses may provide a sensitive and reliable measure of affective states and stress-mediated changes in sympathetic and parasympathetic tones. The aim of this research was to define low (LF) and high frequency (HF) power spectral ranges using pharmacological autonomic blockade, and to examine HRV and BPV parameter changes in response to atropine and propranolol in swine. Ten, 13-week old, barrows (n = 6) and gilts (n = 4) underwent surgery to place an intra-cardiac electrode and a blood pressure catheter attached to a biotelemetric transmitter; pigs had a 3-week recovery period prior to data collection. Each pig was subjected to administration of 4 intravenous (i.v.) drug treatments: a control treatment, 3 mL of saline, and 3 blockade treatments; 0.1 mg/kg of atropine, 1.0 mg/kg of propranolol, and .1 mg/kg of atropine together with 1.0 mg/kg of propranolol. All treatments were delivered by injection in the jugular vein with a minimum of 48 h between individual treatments. Behavior, ECG and blood pressure data were recorded continuously for a total of 1 h, from 30 min pre-injection to 30 min post-injection. For data analyses, two 512-beat intervals were selected for each treatment while the pig was lying and inactive. The first interval was selected from the pre-injection period (baseline), and the second was selected between 10 and 30 min post-injection. Time and frequency domain (power spectral density) analyses were performed on each data interval. Subsequent, LF and HF bands from the power spectral densities were defined based on general linear and regression analyses. The HRV and BPV were computed with a covariate (baseline) factorial analysis of treatment by sex interaction, and day of injection, with mixed models and Tukey's post-hoc tests. The best-fit range for LF was 0.0-0.09 Hz, and HF was 0.09-2.0 Hz (r²: 0.41 and 0.43, respectively). Propranolol and saline injections led to a greater overall total power and overall higher inter-beat interval, HF and LF power. Atropine led to a dominant sympathovagal balance of the cardiac activity in pigs. In addition, atropine led to an increase in LF power of both systolic and diastolic blood pressures in gilts suggesting vagal tone mediation of BPV. The understanding of autonomic regulation of HRV and BPV in domestic swine facilitates our ability to detect and quantify stress responses, and broadens its application in assessing farm animal welfare.     

The influence of exercise intensity on the power spectrum of heart rate variability

Summary The power spectral analysis of R-R interval variability (RRV) has been estimated by means of an autoregressive method in seven sedentary males at rest, during steady-state cycle exercise at 21 percent maximal oxygen uptake. (% V _{O 2max}), SEM 2%, 49% V_{O 2max}, SEM 2% and 70% V_{O 2max}, SEM 2% and during recovery. The RRV, i.e. the absolute power of the spectrum, decreased 10, 100 and 500 times in the three exercise intensities, returning to resting value during recovery. In the RRV power spectrum three components have been identified: (1) high frequency peak (HF), central frequency about 0.24 Hz at rest and recovery, and 0.28 Hz, SEM 0.02, 0.37 Hz, SEM 0.03 and 0.48 Hz, SEM 0.06 during the three exercise intensities, respectively; (2) low frequency peak (LF), central frequency about 0.1 Hz independent of the metabolic state; (3) very low frequency component (VLF), <0.05 Hz, no peak observed. The HF peak power, as a percentage of the total power (HF%), averaged 16%, SEM 5% at rest and did not change during exercise, whereas during recovery it decreased to 5%–10%. The LF% and VLF% were about 50% and 35% at rest and during low exercise intensity, respectively. At higher intensities, LF% decreased to 16% and VLF% increased to 70%. During recovery a return to resting values occurred. The HF component may reflect the increased respiratory rate and the LF peak changes the resetting of the baroreceptor reflex with exercise. The hypothesis is made that VLF fluctuations in heart rate might be partially mediated by the sympathetic system.     

Spectral evaluation of aging effects on blood pressure and heart rate variations in healthy subjects

The background to heart rate variability (HRV) and blood pressure variability (BPV), and their determinants and physiological correlates, remain obscure. The impact of age must be taken into account if HRV and BPV are used for predictive purposes in clinical settings. Healthy subjects show wide inter-individual variation in their heart rate behaviour and the factors affecting heart rate dynamics are not well known. This paper has undertaken to evaluate heart rate variability (HRV) and baroreflex sensitivity (BRS) in a random sample of subjects without evidence of heart disease, and to estimate the relation of HRV and BPV behaviour to age. The aim of this study was to analyse the effects of ageing on HRV and BPV for simultaneous recordings of electrocardiograph (ECG) and blood pressure (BP) signals at rest in healthy subjects. We studied eight young (21 – 34 years old) and eight elderly (68 – 85 years old) rigorously screened subjects from the Fantasia Database to make the reproducibility and comparability of the results more extensive. Time- and frequency-domain analysis of HRV and BPV was performed on 5-minute ectopic-free recordings. BRS on the heart was estimated by frequency-domain analysis of spontaneous variability of systolic blood pressure (SBP) and RR interval. It has been observed that compared to young the elderly subjects have (i) diminished HRV; (ii) a shift in the power spectral density and median frequency to low frequency side for HRV and to higher frequency side for BPV; and (iii) increased low-frequency alpha index and decreased high-frequency alpha index of BRS with overall alpha index augmented. The results convey that normal ageing in the absence of disease is associated with lesser parasympathetic regulation of heart rate. Thus it is concluded that the age is an important factor to be considered for prognosis and diagnosis by HRV and BPV. For reliable clinical applications, more research needs to be done on a broad spectrum of subjects. In addition, these observations will prove to be useful for dynamic modelling of cardiovascular regulation for testing the authentication of new techniques for analysis purposes.     

Autonomic nervous system activity in emotion: A review

Autonomic nervous system (ANS) activity is viewed as a major component of the emotion response in many recent theories of emotion. Positions on the degree of specificity of ANS activation in emotion, however, greatly diverge, ranging from undifferentiated arousal, over acknowledgment of strong response idiosyncrasies, to highly specific predictions of autonomic response patterns for certain emotions. A review of 134 publications that report experimental investigations of emotional effects on peripheral physiological responding in healthy individuals suggests considerable ANS response specificity in emotion when considering subtypes of distinct emotions. The importance of sound terminology of investigated affective states as well as of choice of physiological measures in assessing ANS reactivity is discussed.     

Compression on trigger points in the leg muscle increases parasympathetic nervous activity based on heart rate variability

Massotherapy, the therapeutic use of massage, is used to treat various chronic pain syndromes. One type of massotherapy, pressure stimulus applied over trigger points (TPs), is reported to have excellent therapeutic effects. Its effect is possibly mediated through changes in the autonomic nervous system although little research has been conducted to assess autonomic activity during TP compression. We have investigated how compression applied over TPs affects the autonomic nervous system. Six healthy young adult females whose daily working routine was carried out predominantly in a standing position were enrolled in the study cohort. After a day's work, the subjects were asked to rest supine, and electrocardiograms (ECGs), instantaneous lung volume (ILV) and systolic and diastolic blood pressures (SBP, DBP) were measured before and after pressure application over the TPs in those lower limb muscles where the subjects felt muscle fatigue or discomfort. The subjects were also asked to coordinate breathing with the beeping sounds. The therapeutic effects of TP compression were assessed by a subjective fatigue scale. Parasympathetic nervous activity was also assessed by spectral analysis of heart rate (HR) variability. The transfer function from ILV to HR was evaluated using linear analysis. The results indicated that TP compression (1) decreased HR, SBP and DBP, (2) increased parasympathetic activity, (3) increased the gain from ILV to HR, and (4) improved the fatigue scores. These findings suggest that an increase in parasympathetic nervous activity after the TP compression induced a reduction of fatigue. The therapeutic mechanisms of TP compression to enhance parasympathetic nervous system are discussed.     

Mitigating the effect of non-stationarity in spectral analysis—An application to neonate heart rate analysis

In order to mitigate the effect of non-stationarity in frequency domain analysis of data, we propose a modification to the power spectral estimation, a widely used technique to characterize physiological signals. Spectral analysis requires partitioning data into smaller epochs determined by the desired frequency resolution. The modified approach proposed here involves dividing the data within each epoch by the standard deviation of the data for that epoch. We applied this modified approach to cardiac beat-to-beat interval data recorded from a newborn infant undergoing hypothermia treatment for birth asphyxia. The critically ill infant had episodes of tachyarrhythmia, distributed sporadically throughout the study, which affected the stationarity of the heart rate. Over the period of continuous heart rate recording, the infant's clinical course deteriorated progressively culminating in death. Coinciding with this clinical deterioration, the heart rate signal showed striking changes in both low-frequency and high-frequency power indicating significant impairment of the autonomic nervous system. The standard spectral approach failed to capture these phenomena because of the non-stationarity of the signal. Conversely, the modified approach proposed here captured the deteriorating physiology of the infant clearly.     

心室复极化时程与自主神经系统关系的研究

目前,随着心血管疾病日益增多,国外有少量研究者开始分析心室极化时程变异（Ｒｅｐｏｌａｒｉｚｔｉｏｎ ｄｕｒａｔｉｏｎ ｖａｒｉａｂｉｌｉｔｙ,ＲＤＶ）某些心脏疾病会在心室复极化时程上会有所表现。我们用乌拉坦麻醉大白鼠,研究用阿托品阻断迷走神经前后,ＲＤＶ的变化规律。通过正常人体位变化来改变人体自主神经系统的平衡状态,以研究心室复极化时程与自主神经系统的关系,发现大白鼠的ＨＲＶ和ＲＤＶ的谱分析的峰值     

Practical problems of determining the dimensions of heart rate data

The practical problems are explored of determining the dimension of the phase space set generated from real, experimental and simulated data of the times between consecutive hearbeats in normal and diseased rabbits. It is determined how different measures of dimension have depended on the procedures used to construct the phase space set and on such properties of the data as the amount of data, noise, long-term trends and stationarity. Reproducible estimates of the dimensions of different physiological states are found to require considerable amounts of data recorded under stationary conditions.     

Effect of training on the resting heart rate of rats

Summary Adult male rats were progressively trained 5 days/week on a motor-driven treadmill. The training period lasted 12 weeks and consisted of 60 min/day of wind-sprints and endurance work. No significant difference in resting heart rates was observed between the control and exercise groups during week 1 (394±7 vs. 388±5). However, at week 12 the exercise group had a lower resting heart rate (359±6 vs. 331±4). Heart rates observed following saline, propranolol, atropine, and propranolol plus atropine injections were lower in the exercise group in all cases. The difference in heart rates between the control and exercise groups was 19 beats/min following propranolol plus atropine which was less than the 28 beats/min difference observed under control conditions. With atropine and then with propranolol the differences were 33 and 27 beats/min. These heart rate differences were observed without the presence of cardiac hypertrophy as assessed from ventricle weights.Our data indicate that the bradycardia resulting from exercise training is due primarily to changes other than neural influences on the heart.Supported by a USPHS Career Development Award HL 00052-01     

Moderate alcohol intake is related to increased heart rate variability in young adults: Implications for health and well‐being

Epidemiological literature indicates that the relationship between alcohol consumption and health outcomes reflects a J‐shaped curve such that moderate alcohol consumption confers a protective effect in comparison to abstinence, while heavy consumption is associated with poorer health. While heart rate variability (HRV) may underpin the relationship between drinking and poor health in heavy drinkers, it is unclear whether HRV is increased in moderate, habitual drinkers relative to nonhabitual drinkers. HRV and drinking habits were assessed in 47 volunteers. Results supported hypotheses suggesting that moderate, habitual drinking increases HRV. Although not supported by a significant interaction between drinking group and sex, planned follow‐up analysis also revealed that these findings may be specific to males. Regardless, results highlight HRV as a candidate mechanism for the findings reported in the epidemiological literature.     

Cardiovascular response to exercise in humans following acclimatization to extreme altitude

The purpose of this study was to assess the effects of acclimatization to extreme altitude on the cardiovascular system, using vagal and adrenergic blockade and acute restoration of normoxia during exercise to maximum with one and two legs. Fourteen climbers on an expedition to the Himalayas were studied at a lower base camp (5250 m) following 56–81 days at altitudes between 5250 and 8700 m. After acclimatization, peak heart rate (HR_peak), oxygen uptake (o_2k) and noradrenaline (NA) were similar during maximal one- and two-legged cycling, whereas peak plasma lactate was higher during the one-legged protocol. HR_peak (range 113–168 beats min“¹) was lowest when subjects returned from the higher camps. The degree of partial restoration of HR_peak to more normal values within seconds of 60% 0₂ inhalation (range 5–35 beats min?^l HR_peak increase) was greatest in subjects with low HR_peak. HR responses to /?-l blockade increased as a function of HR_peak and the HR responses to atropine were the least in subjects with high HRpeak- These findings suggest that (a) the reduction in HR_peak is linked to the duration and severity of the hypoxaemia, (b) the degree of restoration of HR_peak with acute normoxia is dependent on the level of attenuation or down-regulation of cardiac sympathetic activation (SNA), (c) cardiac vagal drive is masked to a lesser extent in chronic hypoxia because of attenuated SNA and lower HR_peak values, and (d) the lower blood lactate levels at altitude is a function of muscle mass involvement rather than adrenergic activation, as normal peak values were reached during exercise with a small muscle mass.