Article

The Contribution of "Resting" Body Muscles to the Slow Component of Pulmonary Oxygen Uptake During High-Intensity Cycling

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Abstract

Oxygen uptake (VO2) kinetics during moderate constant-workrate (WR) exercise (>lactate-threshold (θL)) are well described as exponential. AboveθL, these kinetics are more complex, consequent to the development of a delayed slow component (VO2sc), whose aetiology remains controversial. To assess the extent of the contribution to the VO2sc from arm muscles involved in postural stability during cycling, six healthy subjects completed an incremental cycle-ergometer test to the tolerable limit for estimation of θL and determination of peak VO2. They then completed two constant-WR tests at 90% of θL and two at 80% of ∆ (difference between θL and VO2peak). Gas exchange variables were derived breath-by-breath. Local oxygenation profiles of the vastus lateralis and biceps brachii muscles were assessed by near-infrared spectroscopy, with maximal voluntary contractions (MVC) of the relevant muscles being performed post-exercise to provide a frame of reference for normalising the exercise-related oxygenation responses across subjects. Above supra-θL, VO2 rose in an exponential-like fashion ("phase 2), with a delayed VO2sc subsequently developing. This was accompanied by an increase in [reduced haemoglobin] relative to baseline (∆[Hb]), which attained 79 ± 13 % (mean, SD) of MVC maximum in vastus lateralis at end-exercise and 52 ± 27 % in biceps brachii. Biceps brachii ∆[Hb] was significantly correlated with VO2 throughout the slow phase. In contrast, for sub- L exercise, VO2 rose exponentially to reach a steady state with a more modest increase in vastus lateralis ∆[Hb] (30 ± 11 %); biceps brachii ∆[Hb] was minimally affected (8 ± 2 %). That the intramuscular O2 desaturation profile in biceps brachii was proportional to that for VO2sc during supra-θL cycle ergometry is consistent with additional stabilizing arm work contributing to the VO2sc.

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... It is well established that other factors such as the work of ventilation may make small but significant contributions to the pulmonary VȮ 2 response during highintensity exercise. Additionally, it has been shown that the VȮ 2 slow component significantly correlated with increased oxygen utilization of the arm muscles during cycling, 6 presumably due to an alteration in cycling technique as fatigue developed. In this regard, Özyener et al. 6 raised some questions regarding whether the data of Poole et al. 2 were "distorted" by a single data point at the end of the trial, suggesting that a larger proportion of the slow component may reside outside of the working muscles than typically thought. ...
... Additionally, it has been shown that the VȮ 2 slow component significantly correlated with increased oxygen utilization of the arm muscles during cycling, 6 presumably due to an alteration in cycling technique as fatigue developed. In this regard, Özyener et al. 6 raised some questions regarding whether the data of Poole et al. 2 were "distorted" by a single data point at the end of the trial, suggesting that a larger proportion of the slow component may reside outside of the working muscles than typically thought. However, Özyener et al. 6 go on to state that "Even taking this possible 'distorting' influence into account, there is little question that the exercising legs make a dominant contribution to the VȮ 2 slow component during supra-threshold constant work rate cycle ergometry." ...
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... e., pulling on the handlebars during cycling) contribute to the SC [38]. A recent study by Ozyener et al. [39] confirmed this contribution by estimating the muscle O 2 saturation of the biceps brachii at high work rates. A significant correlation was observed between V O 2 and oxygen extraction (deoxy-haemoglobin) as measured by near-infrared spectroscopy and it was concluded that additional work of the arm muscle might contribute to the SC. ...
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Near infrared spectroscopy (NIRS) has been frequently used to assess intra-muscular oxygenation past few decades. In recent years, refinement of NIRS hardware and algorithms used to convert changes in optical absorption to changes in concentration improved the validity of oxygenated haemoglobin (HbO2), deoxyhaemoglobin (Hb), myoglobin (Mb) and the oxidised copper compound of cytochrome aa3 , (cyt aa3 ) measurements. Subsequently, the use of NIRS to study the muscle oxygenation profiles during various types of exercise and to monitor differences in oxygenation levels in patients under various pathological conditions is being increased. It would be thought-provoking to combine future muscle studies with MRS and/or electromyography techniques to improve the understanding of intramuscular oxygenation.
Article
The pulmonary oxygen uptake (VO2) response to incremental-ramp cycle ergometry typically demonstrates lagged-linear first-order kinetics with a slope of ~10-11 ml·min(-1)·W(-1), both above and below the lactate threshold (θL), i.e. there is no discernible VO2 slow component (or "excess" VO2) above θL. We were interested in determining whether a reverse ramp profile would yield the same response dynamics. Ten healthy males performed a maximum incremental -ramp (15-30 W·min(-1), depending on fitness). On another day, the work rate (WR) was increased abruptly to the incremental maximum and then decremented at the same rate of 15-30 W.min(-1) (step-decremental ramp). Five subjects also performed a sub-maximal ramp-decremental test from 90% of θL. VO2 was determined breath-by-breath from continuous monitoring of respired volumes (turbine) and gas concentrations (mass spectrometer). The incremental-ramp VO2-WR slope was 10.3 ± 0.7 ml·min(-1)·W(-1), whereas that of the descending limb of the decremental ramp was 14.2 ± 1.1 ml·min(-1)·W(-1) (p < 0.005). The sub-maximal decremental-ramp slope, however, was only 9. 8 ± 0.9 ml·min(-1)·W(-1): not significantly different from that of the incremental-ramp. This suggests that the VO2 response in the supra-θL domain of incremental-ramp exercise manifest not actual, but pseudo, first-order kinetics. Key pointsThe slope of the decremental-ramp response is appreciably greater than that of the incremental.The response dynamics in supra-θL domain of the incremental-ramp appear not to manifest actual first-order kinetics.The mechanisms underlying the different dynamic response behaviour for incremental and decremental ramps are presently unclear.
Article
Near infrared spectroscopy (NIRS) has been frequently used to assess intra-muscular oxygenation past few decades. In recent years, refinement of NIRS hardware and algorithms used to convert changes in optical absorption to changes in concentration improved the validity of oxygenated haemoglobin (HbO 2), deoxyhaemoglobin (Hb), myoglobin (Mb) and the oxidised copper compound of cytochrome aa 3 , (cyt aa 3) measurements. Subsequently, the use of NIRS to study the muscle oxygenation profiles during various types of exercise and to monitor differences in oxygenation levels in patients under various pathological conditions is being increased. It would be thought-provoking to combine future muscle studies with MRS and/or electromyography techniques to improve the understanding of intramuscular oxygenation. KEY WORDS: Near infrared spectroscopy, muscle oxygenation. İNSANLARDA EGZERSİZ SIRASINDA KAS İÇİ OKSİJENLENMENİN DEĞERLEN-DİRİLMESİ Özet Geçtiğimiz birkaç on-yıldan beri kas-içi oksijenlenmeyi değerlendirmede "near infrared spektroscopy" (NIRS) yöntemi sıklıkla kullanılmaktadır. Son yıllarda NIRS cihazlarının ve bunlarda kullanılan optik absorbsiyon değişikliklerini konsantrasyon farklarına çeviren algoritmaların gittikçe gelişmesi oksijenlenmiş hemoglobin (HbO 2), oksijensiz hemoglobin (Hb), miyoglobin (Mb) ve okside bakır bileşiği olan sitoktom aa 3 , (cyt aa 3) ölçümlerinin güvenilirliğini arttırmıştır. Böylece NIRS farklı egzersiz tiplerinde kas içi oksijenlenme profillerini ortaya koymanın yanı sıra değişik patolojik bozuklukları olan hastalarda oksijenlenme düzeylerini gözlemede kullanılması giderek artmaktadır. Gelecekteki kas çalışmalarında NIRS ile beraber MRS veya elektromiyografi tekniklerini birleştirmek kas içi oksijenlenmenin anlaşılmasını geliştirmede yeni çarpıcı düşüncelerin oluşmasına yol açabilir.
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The sections in this article are:
Article
The aim of the present study was to investigate whether the sigmoid pattern of deoxy[Hb + Mb] during incremental exercise is specific to non-steady-state conditions. Ten highly trained cyclists performed an incremental step (40 W x 3 min(-1)) and ramp (35 W x min(-1)) exercise. Deoxy[Hb + Mb] was measured at the distal and proximal sites of the musculus vastus lateralis throughout the exercises using near-infrared spectroscopy. Deoxy[Hb + Mb] was set out as a function of work rate (% peak power), and using curve-fitting techniques, the best-fitting model was determined. These procedures showed that the sigmoid pattern also provided the best fit for the pattern of deoxy[Hb + Mb] in the step exercise. Furthermore, it was observed that the sigmoid model was similar for the ramp (d = 6.9% +/- 1.1% and 6.9% +/- 1.4% x %(-1) peak power; c/d = 52.1% +/- 3.8% and 52.1% +/- 4.5% peak power, for the proximal and distal measurement sites, respectively) and the step exercise (d = 7.4% +/- 1.5% and 6.4% +/- 1.5% x %(-1) peak power; c/d = 52.3% +/- 6.0% and 52.5% +/- 4.2% peak power, for the proximal and distal measurement sites, respectively). The pattern of deoxy[Hb + Mb] was not influenced by measurement site. From the present study, it can be concluded that the sigmoid pattern of deoxy[Hb + Mb] during incremental exercise is not specific to non-steady-state conditions. It was hypothesized that this pattern is an expression of a nonlinear Q x m/V x O2m relationship, related to changes in muscle fiber-type recruitment.
Article
A simple muscle tissue spectrophotometer is adapted to measure the recovery time (TR) for hemoglobin/myoglobin (Hb/Mb) desaturation in the capillary bed of exercising muscle, termed a deoxygenation meter. The use of the instrument for measuring the extent of deoxygenation is presented, but the use of TR avoids difficulties of quantifying Hb/Mb saturation changes. The TR reflects the balance of oxygen delivery and oxygen demand in the localized muscles of the quadriceps following work near maximum voluntary contraction (MVC) in elite male and female rowers (a total of 22) on two occasions, 1 yr apart. TR ranged from 10 to 80 s and was interpreted as a measure of the time for repayment of oxygen and energy deficits accumulated during intense exercise by tissue respiration under ADP control. The Hb/Mb resaturation times provide a noninvasive localized indication of the degree of O2 delivery stress as evoked by rowing ergometry and may provide directions for localized muscle power output improvement for particular individuals in rowing competitions.
Article
1. At work rates which do not result in a sustained increase in blood lactate ([L-]), oxygen uptake (VO2) approaches the steady state with first-order kinetics. However, when [L-] is increased, at least two kinetic components are required to characterize the VO2 response dynamics. The purpose of the present investigation was to determine whether these more-complex kinetics are best represented as: (a) two components which operate throughout the exercise or (b) a delayed slow component which is consequent to the lactic acidaemia and which does not influence the early development of the O2 deficit. 2. Six healthy subjects underwent an incremental ramp test on a cycle ergometer, to the limit of tolerance, for determination of the maximum VO2 (micro VO2) and and estimation [symbol: see text] of the threshold for lactic acidaemia (theta L) non-invasively. Subjects then performed, on different days, two to four repetitions of square-wave exercise from a baseline of unloaded pedalling ('O' Watts (W)) to work rates (WR) less than theta L (90% theta L) and greater than theta L (half-way between theta L and micro VO2). Ventilatory and pulmonary gas exchange variables were determined breath-by-breath. For each subject, the VO2 transitions were averaged prior to fitting a least-squares algorithm to the on- and off-transient responses. 3. The less than theta L test resulted in a mono-exponential VO2 response, with a time constant of 31.3 and 31.5 s for the on- and off-transients, respectively. 4. The VO2 responses to the greater than theta L test were fitted to three competing models: (a) a single exponential for the entire period; (b) a double exponential for the entire period; and (c) an initial single exponential with a subsequent phase of delayed onset. Model (c) yielded a significantly lower residual mean-squares error than methods (a) and (b), with a time constant for the initial component of 40.2 s for the on-transient and 32.9 s for the off-transient and a subsequent phase of VO2 increase for the on-transient which averaged 230 ml min-1. The delta VO2/delta WR for the early kinetics of the greater than theta L test were not different from the less than theta L test (9.6 and 9.5 ml min-1 W-1, respectively). 5. These data suggest that the slow phase of the greater than theta L VO2 kinetics is a delayed-onset process. This being the case, the O2 deficit during heavy exercise, as conventionally estimated, would be overestimated.
Article
The present study was undertaken to determine whether near-infrared spectroscopy can be used to noninvasively assess skeletal muscle oxygenation in patients with heart failure. The difference between light absorption at 760 and 800 nm was used to assess hemoglobin-myoglobin oxygenation. Initial studies conducted in isolated canine gracilis muscle demonstrated that 760-800-nm absorption correlated closely (r = -0.97 +/- 0.01) with venous hemoglobin O2 saturation when the muscle was stimulated to contract at 0.25-5.0 Hz. In normal subjects (n = 6) and patients with heart failure (n = 8), 760-800-nm absorption changes from the vastus lateralis muscle were monitored at rest, during progressive maximal bicycle exercise, and during thigh cuff inflation to suprasystolic pressure, an intervention designed to assess minimal hemoglobin-myoglobin oxygenation. Absorption changes were expressed relative to the full physiologic range noted from rest to thigh cuff inflation. During exercise, normal subjects exhibited an initial increase in hemoglobin-myoglobin oxygenation followed by a progressive decrease in oxygenation to 27 +/- 13% of the physiologic range at the peak exercise workload of 140 +/- 9 W. In contrast, patients exhibited an initial decrease in hemoglobin-myoglobin oxygenation with the first workload, followed by a progressive further decrease to 26 +/- 13% of the physiologic range at a peak exercise workload of 60 +/- 8 W, less than half the peak workload noted in the normal subjects. At all exercise loads, hemoglobin-myoglobin oxygenation was significantly less in the patients than in the normal subjects. These data suggest that near-infrared spectroscopy can detect impaired skeletal muscle O2 delivery in patients with heart failure. This technique could provide a valuable method of assessing muscle O2 delivery in patients, particularly before and after therapeutic interventions.
Article
A portable software package (TIDAL--Tools for Intelligent Data Acquisition in the Laboratory) for acquiring and analyzing respiratory physiological data is described. TIDAL supports flow-, volume-, and concentration-measuring devices, and any other instruments that produce linear analog outputs. The system allows users to specify the names and types of channels to be sampled, and the calculations involved in reducing samples to breath-to-breath values. The specification of channels and calculations is given in EDL, an Experiment Description Language designed for respiratory physiology. EDL comprises a set of internal functions (primitives) which can be combined into arbitrarily complex expressions. To simplify EDL programming, TIDAL includes a macro processor and a standard macro library; the library contains definitions for a wide variety of respiratory variables. Examples are inspiratory and expiratory times and total volumes, mean inspired and expired gas volumes and concentrations, and end-tidal concentrations. Variables that are derived from these primary data, such as respiratory quotient, are also easily specified. TIDAL is written entirely in the C programming language, with special attention to portable coding practices. The code is organized in a modular structure that eases porting to multiple hardware/compiler/operating system environments.
Article
Breathing has inherent irregularities that produce breath-to-breath fluctuations ("noise") in pulmonary gas exchange. These impair the precision of characterizing nonsteady-state gas exchange kinetics during exercise. We quantified the effects of this noise on the confidence of estimating kinetic parameters of the underlying physiological responses and hence of model discrimination. Five subjects each performed eight transitions from 0 to 100 W on a cycle ergometer. Ventilation, CO2 output, and O2 uptake were computed breath by breath. The eight responses were interpolated uniformly, time aligned, and averaged for each subject; and the kinetic parameters of a first-order model (i.e., the time constant and time delay) were then estimated using three methods: linear least squares, nonlinear least squares, and maximum likelihood. The breath-by-breath noise approximated an uncorrelated Gaussian stochastic process, with a standard deviation that was largely independent of metabolic rate. An expression has therefore been derived for the number of square-wave repetitions required for a specified parameter confidence using methods b and c; method a being less appropriate for parameter estimation of noisy gas exchange kinetics.
Article
To determine the precise nonsteady-state characteristics of ventilation (VE), O2 uptake (VO2), and CO2 output (VCO2) during moderate-intensity exercise, six subjects each underwent eight repetitions of 100-W constant-load cycling. The tests were preceded either by rest or unloaded cycling ("0" W). An early component of VE, VO2, and VCO2 responses, which was obscured on any single test by the breath-to-breath fluctuations, became apparent when the several repetitions were averaged. These early responses were abrupt when the work was instituted from rest but were much slower and smaller from the 0-W base line and corresponded to the phase of cardiodynamic gas exchange. Some 20 s after the onset of the work a further monoexponential increase to steady state occurred in all three variables, the time constants of which did not differ between the two types of test. Consequently, the exponential behavior of VE, VO2, and VCO2 in response to moderate exercise is best described by a model that incorporates only the second phase of the response.
Article
We compared the slow rise in VO2 during heavy exercise (i.e., greater than lactic acidosis threshold (LAT)) with changes in muscle oxyhemoglobin+oxymyoglobin (O2Hb/O2Mb) saturation by reflectance near infrared spectroscopy. Ten subjects performed four 6-min cycle ergometer tests with two constant work rates less than and two greater than the LAT, equivalent to 20, 40, 65 and 75% peak VO2. During less than LAT exercise, O2Hb/O2Mb saturation decreased to a minimum by 2 min and then remained constant or rose slightly. For greater than LAT work rates, the initial fall in O2Hb/O2Mb saturation was greater the higher the work rate and continued to decrease with time after 3 min. Between minutes 3 and 6, the rate of decrease in O2Hb/O2Mb saturation correlated with the increase in VO2 (r = -0.69, P < 0.0001). These studies support the hypothesis that the slow rise in VO2 during heavy constant work rate exercise is associated with a progressive decline in O2Hb/O2Mb saturation in the contracting muscles themselves that may be facilitated by capillary oxyhemoglobin dissociation owing to tissue lactic acidosis (Bohr effect).
Article
Near-infrared (NIR) spectroscopy is a noninvasive technique that uses the differential absorption properties of hemoglobin to evaluate skeletal muscle oxygenation. Oxygenated and deoxygenated hemoglobin absorb light equally at 800 nm, whereas at 760 nm absorption is primarily from deoxygenated hemoglobin. Therefore, monitoring these two wavelengths provides an index of deoxygenation. To investigate whether venous oxygen saturation and absorption between 760 and 800 nm (760-800 nm absorption) are correlated, both were measured during forearm exercise. Significant correlations were observed in all subjects (r = 0.92 +/- 0.07; P < 0.05). The contribution of skin flow to the changes in 760-800 nm absorption was investigated by simultaneous measurement of skin flow by laser flow Doppler and NIR recordings during hot water immersion. Changes in skin flow but not 760-800 nm absorption were noted. Intra-arterial infusions of nitroprusside and norepinephrine were performed to study the effect of alteration of muscle perfusion on 760-800 nm absorption. Limb flow was measured with venous plethysmography. Percent oxygenation increased with nitroprusside and decreased with norepinephrine. Finally, the contribution of myoglobin to the 760-800 nm absorption was assessed by using 1H-magnetic resonance spectroscopy. At peak exercise, percent NIR deoxygenation during exercise was 80 +/- 7%, but only one subject exhibited a small deoxygenated myoglobin signal. In conclusion, 760-800 nm absorption is 1) closely correlated with venous oxygen saturation, 2) minimally affected by skin blood flow, 3) altered by changes in limb perfusion, and 4) primarily derived from deoxygenated hemoglobin and not myoglobin.
Article
We applied near-infrared spectroscopy (NIRS) for the simultaneous measurement of forearm blood flow (FBF) and oxygen consumption (VO2) in the human by inducing a 50-mmHg venous occlusion. Eleven healthy subjects were studied both at rest and after hand exercise during vascular occlusion. FBF was also measured by strain-gauge plethysmography. FBF measured by NIRS was 1.9 +/- 0.8 ml.100 ml-1.min-1 at rest and 8.2 +/- 2.9 ml.100 ml-1.min-1 after hand exercise. These values showed a correlation (r = 0.94) with those obtained by the plethysmography. VO2 values were 4.6 +/- 1.3 microM O2 x 100 ml-1.min-1 at rest and 24.9 +/- 11.2 microM O2 x 100 ml-1.min-1 after hand exercise. The scatter of the FBF and VO2 values showed a good correlation between the two variables (r = 0.93). The results demonstrate that NIRS provides the particular advantage of obtaining the contemporary evaluation of blood flow and VO2, allowing correlation of these two variables by a single maneuver without discomfort for the subject.
The purpose of this study was to compare the rates of muscle deoxygenation in the exercising muscles during incremental arm cranking and leg cycling exercise in healthy men and women. Fifteen men and 10 women completed arm cranking and leg cycling tests to exhaustion in separate sessions in a counterbalanced order. Cardiorespiratory measurements were monitored using an automated metabolic cart interfaced with an electrocardiogram. Tissue absorbency was recorded continuously at 760 nm and 850 nm during incremental exercise and 6 min of recovery, with a near infrared spectrometer interfaced with a computer. Muscle oxygenation was calculated from the tissue absorbency measurements at 30%, 45%, 60%, 75% and 90% of peak oxygen uptake (VO2) during each exercise mode and is expressed as a percentage of the maximal range observed during exercise and recovery (%Mox). Exponential regression analysis indicated significant inverse relationships (P < 0.01) between %Mox and absolute VO2 during arm cranking and leg cycling in men (multiple R = -0.96 and -0.99, respectively) and women (R = -0.94 and -0.99, respectively). No significant interaction was observed for the %Mox between the two exercise modes and between the two genders. The rate of muscle deoxygenation per litre of VO2 was 31.1% and 26.4% during arm cranking and leg cycling, respectively, in men, and 26.3% and 37.4% respectively, in women. It was concluded that the rate of decline in %Mox for a given increase in VO2 between 30% and 90% of the peak VO2 was independent of exercise mode and gender.
Article
Indices of mechanical power output were obtained from twelve subjects during high intensity leg cycle ergometry tests (20 second duration; 75 grams per kilogram total body mass) using two protocols: one with a standard handle-bar grip (with-grip), and one with supinated wrists (without-grip). Peak mechanical power, mean mechanical power, fatigue index and total mechanical work values were calculated for each subject during each test, and the sample mean differences associated with the two protocols were compared using paired Student t-tests. The with-grip protocol yielded significantly greater peak mechanical power output and greater fatigue index than the without-grip protocol (886 +/- 124W and 815 +/- 151W, respectively; and 35 +/- 10% and 25 +/- 8%, respectively; p<0.01). The electrical activity of the anterior forearm musculature was measured in the twelfth subject during the performance of each of the test protocols. While peak mechanical power output was greater during the with-grip protocol, than during the without-grip protocol, the electromyographs showed much greater forearm muscle activity during the with-grip protocol. Thus the protocol which allowed for the greatest measure of peak leg power output was also associated with considerable arm muscle activity. These findings should be considered when biochemical and physiological measurements are obtained from arm blood samples.
Article
Traditionally, leg cycle ergometry is used to assess the power output of the lower limbs. However, it is suspected that the upper body makes a significant, albeit as yet unknown, contribution to the measured power output, and as such, the lean mass of the whole body should be considered during ergometric assessment. To test this idea, indices of mechanical power output were obtained from 11 subjects during high intensity leg cycle ergometry tests (20 second duration; 75 grams per kilogram total body mass) using two protocols: one with a standard handle-bar grip (with grip) and one with supinated wrists (without-grip). Peak mechanical power, mean mechanical power, fatigue index and total mechanical work values were calculated for each subject during each test and the sample mean differences associated with the two protocols were compared using paired Student t-tests. The with-grip protocol yielded significantly greater peak mechanical power output than the without-grip protocol (886+/-124 W and 815+/-151 W, respectively), suggesting a significant upper body contribution to the maximum power output measured for the legs. As a first step towards quantifying the upper body involvement during leg cycle ergometry, surface electromyography of the forearm musculature was measured in a twelfth subject whilst performing each of the test protocols. During the with-grip ergometer tests, the intensity of electrical activity in the forearm musculature was similar, if not greater than, the intensity of electrical activity recorded for the forearm musculature during 100% maximum voluntary hand grip-dynamometer contractions, suggesting maximum isometric-type forearm muscle contraction during the with-grip leg ergometry tests. These findings suggest that the performance of traditional-style leg cycle ergometry requires a muscular contribution from the whole body. As such, researchers should be mindful of this, both in terms of the allocation of ergometer loads, and in the analysis of blood-borne metabolites.
Article
The aim of the present study was to examine whether ATP production increases and mechanical efficiency decreases during intense exercise and to evaluate how previous exercise affects ATP turnover during intense exercise. Six subjects performed two (EX1 and EX2) 3-min one-legged knee-extensor exercise bouts [66.2 +/- 3.9 and 66.1 +/- 3.9 (+/-SE) W] separated by a 6-min rest period. Anaerobic ATP production, estimated from net changes in and release of metabolites from the active muscle, was 3.5 +/- 1.2, 2.4 +/- 0.6, and 1.4 +/- 0.2 mmol ATP x kg dry wt(-1) x s(-1) during the first 5, next 10, and remaining 165 s of EX1, respectively. The corresponding aerobic ATP production, determined from muscle oxygen uptake, was 0.7 +/- 0.1, 1.4 +/- 0.2, and 4.7 +/- 0.4 mmol ATP x kg dry wt(-1) x s(-1), respectively. The mean rate of ATP production during the first 5 s and next 10 s was lower (P < 0.05) than during the rest of the exercise (4.2 +/- 1.2 and 3.8 +/- 0.7 vs. 6.1 +/- 0.3 mmol ATP x kg dry wt(-1) x s(-1)). Thus mechanical efficiency, expressed as work per ATP produced, was lowered (P < 0.05) in the last phase of exercise (39.6 +/- 6.1 and 40.7 +/- 5.8 vs. 25.0 +/- 1.3 J/mmol ATP). The anaerobic ATP production was lower (P < 0.05) in EX2 than in EX1, but the aerobic ATP turnover was higher (P < 0.05) in EX2 than in EX1, resulting in the same muscle ATP production in EX1 and EX2. The present data suggest that the rate of ATP turnover increases during intense exercise at a constant work rate. Thus mechanical efficiency declines as intense exercise is continued. Furthermore, when intense exercise is repeated, there is a shift toward greater aerobic energy contribution, but the total ATP turnover is not significantly altered.
Article
1. The maximal oxygen uptake (V(O(2),peak)) during dynamic muscular exercise is commonly taken as a crucial determinant of the ability to sustain high-intensity exercise. Considerably less attention, however, has been given to the rate at which V(O(2)) increases to attain this maximum (or to its submaximal requirement), and even less to the kinetic features of the response following exercise. 2. Six, healthy, male volunteers (aged 22 to 58 years), each performed 13 exercise tests: initial ramp-incremental cycle ergometry to the limit of tolerance and subsequently, on different days, three bouts of square-wave exercise each at moderate, heavy, very heavy and severe intensities. Pulmonary gas exchange variables were determined breath by breath throughout exercise and recovery from the continuous monitoring of respired volumes (turbine) and gas concentrations (mass spectrometer). 3. For moderate exercise, the V(O(2)) kinetics were well described by a simple mono-exponential function, following a short cardiodynamic phase, with the on- and off-transients having similar time constants (tau(1)); i.e. tau(1,on) averaged 33 +/- 16 s (+/- S.D.) and tau(1,off) 29 +/- 6 s. 4. The on-transient V(O(2)) kinetics were more complex for heavy exercise. The inclusion of a second slow and delayed exponential component provided an adequate description of the response; i.e. tau(1,on) = 32 +/- 17 s and tau(2,on) = 170 +/- 49 s. The off-transient V(O(2)) kinetics, however, remained mono-exponential (tau(1,off) = 42 +/- 11 s). 5. For very heavy exercise, the on-transient V(O(2)) kinetics were also well described by a double exponential function (tau(1,on) = 34 +/- 11 s and tau(2,on) = 163 +/- 46 s). However, a double exponential, with no delay, was required to characterise the off-transient kinetics (i.e. tau(1,off) = 33 +/- 5 s and tau(2,off) = 460 +/- 123 s). 6. At the highest intensity (severe), the on-transient V(O(2)) kinetics reverted to a mono-exponential profile (tau(1,on) = 34 +/- 7 s), while the off-transient kinetics retained a two-component form (tau(1,off) = 35 +/- 11 s and tau(2,off) = 539 +/- 379 s). 7. We therefore conclude that the kinetics of V(O(2)) during dynamic muscular exercise are strikingly influenced by the exercise intensity, both with respect to model order and to dynamic asymmetries between the on- and off-transient responses.
Article
The fundamental pulmonary O(2) uptake (.VO(2)) response to moderate, constant-load exercise can be characterized as (d.VO(2)/dt)(tau)+Delta.VO(2) (t)=Delta.VO(2SS) where Delta.VO(2SS) is the steady-state response, and tau is the time constant, with the .VO(2) kinetics reflecting intramuscular O(2) uptake (.QO(2)) kinetics, to within 10%. The role of phosphocreatine (PCr) turnover in .QO(2) control can be explored using (31)P-MR spectroscopy, simultaneously with .VO(2). Although tau.VO(2) and tauPCr vary widely among subjects (approx. 20-65 s), they are not significantly different from each other, either at the on- or off-transient. A caveat to interpreting the "well-fit" exponential is that numerous units of similar Delta.VO(2SS) but with a wide tau distribution can also yield a .VO(2) response with an apparent single tau. This tau is, significantly, inversely correlated with lactate threshold and .VO(2max)(but is poorly predictive; a frail stamen, therefore), consistent with tau not characterizing a compartment with uniform kinetics. At higher intensities, the fundamental kinetics become supplemented with a slowly-developing phase, setting .VO(2)on a trajectory towards maximum .VO(2). This slow component is also demonstrable in Delta[PCr]: the decreased efficiency thereby reflecting a predominantly high phosphate-cost of force production rather than a high O(2)-cost of phosphate production. We also propose that the O(2)-deficit for the slow-component is more likely to reflect shifting Delta.VO(2SS) rather than a single one with a single tau.
Article
The purpose of this study was to examine the upper-body contribution via handgrip to power profiles and blood lactate concentrations during high-intensity cycle ergometry. Nine trained male subjects each completed a 20-s high-intensity cycle ergometer test twice, in a random manner, using two protocols, with a handgrip (WG), and without handgrip (WOHG). Capillary (earlobe) blood samples were obtained pre- and post-exercise. Blood samples were corrected for plasma volume changes and analyzed to determine blood lactate concentrations. In the WG protocol, mean (+/-SEM) blood lactate concentrations sampled over the three conditions were 0.98 +/- 0.33 mmol.L-1, 5.68 +/- 0.46 mmol.L-1, and 9.14 +/- 0.38 mmol.L-1, respectively. During the WOHG protocol, blood lactate values recorded were 0.99 +/- 0.26 mmol.L-1, 5.58 +/- 0.58 mmol.L-1, and 7.62 +/- 0.65 mmol.L-1, respectively. Differences were found (P < 0.05) from rest to 4 min after exercise for both groups. Differences in concentrations were also observed between groups at the 4-min postexercise blood-sampling stage. Peak power output values recorded using the WG protocol were also greater (1461 +/- 94 W vs 1136 +/- 88 W; P < 0.05). No differences were recorded for mean power output (MPO), fatigue index (FI), or work done (WD). Results indicate significant differences in power output and blood lactate concentrations between protocols. These findings suggest that the performance of traditional style leg-cycle ergometry requires a muscular contribution from the whole body. As such, researchers should consider this, both in terms of the allocation of ergometer loads, and in the analysis of blood-borne metabolites.
Article
The on- and off-transient (i.e. phase II) responses of pulmonary oxygen uptake (V(O(2))) to moderate-intensity exercise (i.e. below the lactate threshold, theta;(L)) in humans has been shown to conform to both mono-exponentiality and 'on-off' symmetry, consistent with a system manifesting linear control dynamics. However above theta;(L) the V(O(2)) kinetics have been shown to be more complex: during high-intensity exercise neither mono-exponentiality nor 'on-off' symmetry have been shown to appropriately characterise the V(O(2)) response. Muscle [phosphocreatine] ([PCr]) responses to exercise, however, have been proposed to be dynamically linear with respect to work rate, and to demonstrate 'on-off' symmetry at all work intenisties. We were therefore interested in examining the kinetic characteristics of the V(O(2)) and [PCr] responses to moderate- and high-intensity knee-extensor exercise in order to improve our understanding of the factors involved in the putative phosphate-linked control of muscle oxygen consumption. We estimated the dynamics of intramuscular [PCr] simultaneously with those of V(O(2)) in nine healthy males who performed repeated bouts of both moderate- and high-intensity square-wave, knee-extension exercise for 6 min, inside a whole-body magnetic resonance spectroscopy (MRS) system. A transmit-receive surface coil placed under the right quadriceps muscle allowed estimation of intramuscular [PCr]; V(O(2)) was measured breath-by-breath using a custom-designed turbine and a mass spectrometer system. For moderate exercise, the kinetics were well described by a simple mono-exponential function (following a short cardiodynamic phase for V(O(2))), with time constants (tau) averaging: tauV(O(2))(,on) 35 +/- 14 s (+/- S.D.), tau[PCr](on) 33 +/- 12 s, tauV(O(2))(,off) 50 +/- 13 s and tau[PCr](off) 51 +/- 13 s. The kinetics for both V(O(2)) and [PCr] were more complex for high-intensity exercise. The fundamental phase expressing average tau values of tauV(O(2))(,on) 39 +/- 4 s, tau[PCr](on) 38 +/- 11 s, tauV(O(2))(,off) 51 +/- 6 s and tau[PCr](off) 47 +/- 11 s. An associated slow component was expressed in the on-transient only for both V(O(2)) and [PCr], and averaged 15.3 +/- 5.4 and 13.9 +/- 9.1 % of the fundamental amplitudes for V(O(2)) and [PCr], respectively. In conclusion, the tau values of the fundamental component of [PCr] and V(O(2)) dynamics cohere to within 10 %, during both the on- and off-transients to a constant-load work rate of both moderate- and high-intensity exercise. On average, approximately 90 % of the magnitude of the V(O(2)) slow component during high-intensity exercise is reflected within the exercising muscle by its [PCr] response.
Article
For moderate-intensity exercise (below lactate threshold, thetaL), muscle O(2) consumption (VO(2)) kinetics are expressed in a first-order phase 2 (or fundamental) pulmonary O(2) uptake (VO(2)) response: dVO(2)/dt . tau + DeltaVO(2)((t)) = DeltaVO(2)((ss)); where DeltaVO(2)(ss) is the steady-state VO(2) increment, and tau the VO(2) time constant (which is within approximately 10% of tauQVO(2)). A likely source of VO(2) control in this intensity domain is ADP-mediated, for which intramuscular phosphocreatine (PCr) may serve as a proxy variable. Whether, in reality, this behavior reflects the operation of a single homogeneous compartment is unclear, however; a multicompartment structure comprised of units having a similar DeltaVO(2)((ss)) but with widely varying tau can also yield a "well-fit" exponential response with an apparent single tau. In support of this is the inverse (although poorly predictive) correlation between tau and both theta(L) and VO(2max). Above theta(L), the fundamental VO(2) kinetics are supplemented with a delayed, slowly developing component that can set VO(2) on a trajectory towards VO(2max), and that has complex temporal- and intensity-related kinetics. This VO(2) slow component is also demonstrable in [PCr], suggesting that the decreased efficiency above theta(L) predominantly reflects a high phosphate cost of force production rather than a high O(2) cost of phosphate production. In addition, the oxygen deficit for the slow component is more likely to reflect a progressive shifting of DeltaVO(2)((ss)) rather than a single DeltaVO(2)((ss)) having a single tau.
Article
The effect of prior exercise on pulmonary O2 uptake (V̇O2p), leg blood flow (LBF), and muscle deoxygenation at the onset of heavy-intensity alternate-leg knee-extension (KE) exercise was examined. Seven subjects [27 (5) yr; mean (SD)] performed step transitions (n = 3; 8 min) from passive KE following no warm-up (HVY 1) and heavy-intensity (Δ50%, 8 min; HVY 2) KE exercise. V̇O2p was measured breath-by-breath; LBF was measured by Doppler ultrasound at the femoral artery; and oxy (O2Hb)-, deoxy (HHb)-, and total (Hbtot) hemoglobin/myoglobin of the vastus lateralis muscle were measured continuously by near-infrared spectroscopy (NIRS; Hamamatsu NIRO-300). Phase 2 V̇O 2p, LBF, and HHb data were fit with a monoexponential model. The time delay (TD) from exercise onset to an increase in HHb was also determined and an HHb effective time constant (HHb - MRT = TD + τ) was calculated. Prior heavy-intensity exercise resulted in a speeding (P < 0.05) of phase 2 V̇O2p kinetics [HVY 1: 42 s (6); HVY 2: 37 s (8)], with no change in the phase 2 amplitude [HVY 1: 1.43 l/min (0.21); HVY 2: 1.48 l/min (0.21)] or amplitude of the V̇O2p slow component [HVY 1: 0.18 l/min (0.08); HVY 2: 0.18 l/min (0.09)]. O2Hb and Hbtot were elevated throughout the on-transient following prior heavy-intensity exercise. The τLBF [HVY 1: 39 s (7); HVY 2: 47 s (21); P = 0.48] and HHb-MRT [HVY 1: 23 s (4); HVY 2: 21 s (7); P = 0.63] were unaffected by prior exercise. However, the increase in HHb [HVY 1: 21 μM (10); HVY 2: 25 μM (10); P < 0.001] and the HHb-to-V̇O2p ratio [(HHb/V̇O2p) HVY 1: 14 μM·l-1·min-1 (6); HVY 2: 17 μM·l-1·min-1 (5); P < 0.05] were greater following prior heavy-intensity exercise. These results suggest that the speeding of phase 2 τV̇O2p was the result of both elevated local O2 availability and greater O2 extraction evidenced by the greater HHb amplitude and HHb/V̇O2p ratio following prior heavy-intensity exercise.
Influence of muscle fibre type and motor recruitment on VO 2 kinetics
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