scholarly journals Adjustments of pulmonary O2 uptake and muscle deoxygenation during ramp incremental exercise and constant-load moderate-intensity exercise in young and older adults

2012 ◽  
Vol 113 (9) ◽  
pp. 1466-1475 ◽  
Author(s):  
Braden M. R. Gravelle ◽  
Juan M. Murias ◽  
Matthew D. Spencer ◽  
Donald H. Paterson ◽  
John M. Kowalchuk
Keyword(s):  
Older Adults ◽  
Lactate Threshold ◽  
Near Infrared ◽  
Vastus Lateralis ◽  
Constant Load ◽  
Moderate Intensity ◽  
Sigmoid Function ◽  
Moderate Intensity Exercise ◽  
Muscle Deoxygenation ◽  
Age Related

The matching of muscle O2 delivery to O2 utilization can be inferred from the adjustments in muscle deoxygenation (Δ[HHb]) and pulmonary O2 uptake (V̇o2p). This study examined the adjustments of V̇o2p and Δ[HHb] during ramp incremental (RI) and constant-load (CL) exercise in adult males. Ten young adults (YA; age: 25 ± 5 yr) and nine older adults (OA; age: 70 ± 3 yr) completed two RI tests and six CL step transitions to a work rate (WR) corresponding to 1) 80% of the estimated lactate threshold (same relative WR) and 2) 50 W (same absolute WR). V̇o2p was measured breath by breath, and Δ[HHb] of the vastus lateralis was measured using near-infrared spectroscopy. Δ[HHb]-WR profiles were normalized from baseline (0%) to peak Δ[HHb] (100%) and fit using a sigmoid function. The sigmoid slope ( d) was greater ( P < 0.05) in OA (0.027 ± 0.01%/W) compared with YA (0.017 ± 0.01%/W), and the c/ d value (a value corresponding to 50% of the amplitude) was smaller ( P < 0.05) for OA (133 ± 40 W) than for YA (195 ± 51 W). No age-related differences in the sigmoid parameters were reported when WR was expressed as a percentage of peak WR. V̇o2p kinetics compared with Δ[HHb] kinetics for the 50-W transition were similar between YA and OA; however, Δ[HHb] kinetics during the transition to 80% of the lactate threshold were faster than V̇o2p kinetics in both groups. The greater reliance on O2 extraction displayed in OA during RI exercise suggests a lower O2 delivery-to-O2 utilization relationship at a given absolute WR compared with YA.

Effects of ageing on muscle O2 utilization and muscle oxygenation during the transition to moderate-intensity exercise

2007 ◽  
Vol 32 (6) ◽  
pp. 1251-1262 ◽  
Author(s):  
Darren S. DeLorey ◽  
Donald H. Paterson ◽  
John M. Kowalchuk
Keyword(s):  
Older Adults ◽  
Near Infrared ◽  
Oxidative Capacity ◽  
Delivery Systems ◽  
Enzyme Activation ◽  
Uptake Kinetics ◽  
Moderate Intensity ◽  
Moderate Intensity Exercise ◽  
Age Related ◽  
Integrated Response

At the onset of exercise, an increase in muscle and pulmonary O2 consumption is met by increases in muscle O2 delivery and muscle O2 extraction. Thus, the study of pulmonary O2 uptake kinetics reflects the integrated response between the convective and diffusive O2 delivery systems and the muscle metabolic machinery (i.e., mitochondrial enzyme activation and provision of acetyl groups to the tricarboxcylic acid cycle) to increase muscle O2 consumption. Pulmonary O2 uptake kinetics are slowed in older adults compared with young adults and previous studies suggest that the slower O2 uptake kinetics may be the result of an age-associated decline in the ability of older adults to increase O2 delivery to active muscles. However, an inherent limitation to understanding the control of and limitations to pulmonary O2 uptake kinetics is that it is methodologically difficult to examine the adaptation of muscle perfusion and O2 delivery and muscle O2 utilization in the muscle microcirculation of active muscles in the dynamically exercising human. In this review, we provide an overview of the effect of ageing on pulmonary O2 uptake kinetics (reflecting the activation of muscle O2 consumption) during the transition to moderate-intensity exercise. Age-related changes in O2 delivery systems and muscle oxidative capacity are examined as potential limitations to pulmonary O2 uptake kinetics. We then review recent studies from our laboratory that have investigated the control of pulmonary O2 uptake kinetics at the level of the muscle microcirculation by examining the adaptation of muscle O2 delivery and muscle O2 utilization using near-infrared spectroscopy during the transition to exercise in healthy young and older adults.

Relationship between pulmonary O2 uptake kinetics and muscle deoxygenation during moderate-intensity exercise

2003 ◽  
Vol 95 (1) ◽  
pp. 113-120 ◽  
Author(s):  
Darren S. DeLorey ◽  
John M. Kowalchuk ◽  
Donald H. Paterson
Keyword(s):  
Near Infrared ◽  
Vastus Lateralis ◽  
Metabolic Activation ◽  
Moderate Intensity ◽  
Vastus Lateralis Muscle ◽  
Phase 2 ◽  
Moderate Intensity Exercise ◽  
Muscle Deoxygenation ◽  
Monoexponential Model ◽  
Kinetics Of

The temporal relationship between the kinetics of phase 2 pulmonary O2 uptake (V̇o2p) and deoxygenation of the vastus lateralis muscle was examined during moderate-intensity leg-cycling exercise. Young adults (5 men, 6 women; 23 ± 3 yr; mean ± SD) performed repeated transitions on 3 separate days from 20 W to a constant work rate corresponding to 80% of lactate threshold. Breath-by-breath V̇o2p was measured by mass spectrometer and volume turbine. Deoxyhemoglobin (HHb), oxyhemoglobin, and total hemoglobin and myoglobin were sampled each second by near-infrared spectroscopy (Hamamatsu NIRO-300). V̇o2p data were filtered, interpolated to 1 s, and averaged to 5-s bins; HHb data were averaged to 5-s bins. Phase 2 V̇o2p data were fit with a monoexponential model. For HHb, a time delay (TDHHb) from exercise onset to an increase in HHb was determined, and thereafter data were fit with a monoexponential model. The time constant for V̇o2p (30 ± 8 s) was slower ( P < 0.01) than that for HHb (10 ± 3 s). The TDHHb before an increase in HHb was 13 ± 2 s. The possible mechanisms of the TDHHb are discussed with reference to metabolic activation and matching of local muscle O2 delivery and O2 utilization. After this initial TDHHb, the kinetics of local muscle deoxygenation were faster than those of phase 2 V̇o2p (and presumably muscle O2 consumption), reflecting increased O2 extraction and a mismatch between local muscle O2 consumption and perfusion.

Effects of prior heavy-intensity exercise on pulmonary O2 uptake and muscle deoxygenation kinetics in young and older adult humans

2004 ◽  
Vol 97 (3) ◽  
pp. 998-1005 ◽  
Author(s):  
Darren S. DeLorey ◽  
John M. Kowalchuk ◽  
Donald H. Paterson
Keyword(s):  
Older Adults ◽  
Near Infrared ◽  
Vastus Lateralis ◽  
Muscle Blood Flow ◽  
Moderate Intensity ◽  
Vastus Lateralis Muscle ◽  
Warm Up ◽  
Muscle Deoxygenation ◽  
Older Subjects ◽  
Young Subjects

Pulmonary O2 uptake (V̇o2p) and muscle deoxygenation kinetics were examined during moderate-intensity cycling (80% lactate threshold) without warm-up and after heavy-intensity warm-up exercise in young ( n = 6; 25 ± 3 yr) and older ( n = 5; 68 ± 3 yr) adults. We hypothesized that heavy warm-up would speed V̇o2p kinetics in older adults consequent to an improved intramuscular oxygenation. Subjects performed step transitions ( n = 4; 6 min) from 20 W to moderate-intensity exercise preceded by either no warm-up or heavy-intensity warm-up (6 min). V̇o2p was measured breath by breath. Oxy-, deoxy-(HHb), and total hemoglobin and myoglobin (Hbtot) of the vastus lateralis muscle were measured continuously by near-infrared spectroscopy (NIRS). V̇o2p (phase 2; τ) and HHb data were fit with a monoexponential model. After heavy-intensity warm-up, oxyhemoglobin (older subjects: 13 ± 9 μM; young subjects: 9 ± 8 μM) and Hbtot (older subjects: 12 ± 8 μM; young subjects: 14 ± 10 μM) were elevated ( P < 0.05) relative to the no warm-up pretransition baseline. In older adults, τV̇o2p adapted at a faster rate ( P < 0.05) after heavy warm-up (30 ± 7 s) than no warm-up (38 ± 5 s), whereas in young subjects, τV̇o2p was similar in no warm-up (26 ± 7 s) and heavy warm-up (25 ± 5 s). HHb adapted at a similar rate in older and young adults after no warm-up; however, in older adults after heavy warm-up, the adaptation of HHb was slower ( P < 0.01) compared with young and no warm-up. These data suggest that, in older adults, V̇o2p kinetics may be limited by a slow adaptation of muscle blood flow and O2 delivery.

Prior heavy exercise elevates pyruvate dehydrogenase activity and muscle oxygenation and speeds O2 uptake kinetics during moderate exercise in older adults

2009 ◽  
Vol 297 (3) ◽  
pp. R877-R884 ◽  
Author(s):  
Brendon J. Gurd ◽  
Sandra J. Peters ◽  
George J. F. Heigenhauser ◽  
Paul J. LeBlanc ◽  
Timothy J. Doherty ◽  
...  
Keyword(s):  
Older Adults ◽  
Pyruvate Dehydrogenase ◽  
Near Infrared ◽  
Muscle Oxygenation ◽  
Moderate Intensity ◽  
Moderate Intensity Exercise ◽  
Younger Adults ◽  
Warm Up ◽  
Muscle Deoxygenation ◽  
Exercise Activity

The adaptation of pulmonary oxygen uptake (V̇o2p) kinetics during the transition to moderate-intensity exercise is slowed in older compared with younger adults; however, this response is faster following a prior bout of heavy-intensity exercise. We have examined V̇o2p kinetics, pyruvate dehydrogenase (PDH) activation, muscle metabolite contents, and muscle deoxygenation in older adults [ n = 6; 70 ± 5 (67–74) yr] during moderate-intensity exercise (Mod1) and during moderate-intensity exercise preceded by heavy-intensity warm-up exercise (Mod2). The phase 2 V̇o2p time constant (τV̇o2p) was reduced ( P < 0.05) in Mod2 (29 ± 5 s) compared with Mod1 (39 ± 14 s). PDH activity was elevated ( P < 0.05) at baseline prior to Mod2 (2.1 ± 0.6 vs. 1.2 ± 0.3 mmol acetyl-CoA·min−1·kg wet wt−1), and the delay in attaining end-exercise activity was abolished. Phosphocreatine breakdown during exercise was reduced ( P < 0.05) at both 30 s and 6 min in Mod2 compared with Mod1. Near-infrared spectroscopy-derived indices of muscle oxygenation were elevated both prior to and throughout Mod2, while muscle deoxygenation kinetics were not different between exercise bouts consistent with elevated perfusion and O2 availability. These results suggest that in older adults, faster V̇o2p kinetics following prior heavy-intensity exercise are likely a result of prior activation of mitochondrial enzyme activity in combination with elevated muscle perfusion and O2 availability.

Time-course of V̇O2 kinetics responses during moderate-intensity exercise subsequent to HIIT vs moderate-intensity continuous training in type 2 diabetes.

2021 ◽  
Author(s):  
Norita Gildea ◽  
Adam McDermott ◽  
Joel Rocha ◽  
Donal O'Shea ◽  
Simon Green ◽  
...  
Keyword(s):  
Type 2 Diabetes ◽  
Time Course ◽  
Near Infrared ◽  
Vastus Lateralis ◽  
Moderate Intensity ◽  
Vastus Lateralis Muscle ◽  
Maximal Heart Rate ◽  
Continuous Training ◽  
Muscle Deoxygenation

We assessed the time course of changes in oxygen uptake (V̇O2) and muscle deoxygenation (i.e., deoxygenated haemoglobin and myoglobin, [HHb+Mb]) kinetics during transitions to moderate-intensity cycling following 12-weeks of low-volume high-intensity interval training (HIIT) vs. moderate-intensity continuous training (MICT) in adults with type 2 diabetes (T2D). Participants were randomly assigned to MICT (n=10, 50 min of moderate-intensity cycling), HIIT (n=9, 10x1 min at ~90% maximal heart rate) or non-exercising control (n=9) groups. Exercising groups trained 3 times per week and measurements were taken every 3 weeks. [HHb+Mb] kinetics were measured by near-infrared spectroscopy at the vastus lateralis muscle. The local matching of O2 delivery to O2 utilization was assessed by the Δ[HHb+Mb]/ΔV̇O2ratio. The pretraining time constant of the primary phase of V̇O2 (τV̇O2p ) decreased (P<0.05) at wk 3 of training in both MICT (from 44±12 to 32±5 s) and HIIT (from 42±8 to 32 ± 4 s) with no further changes thereafter; while no changes were reported in controls. The pretraining overall dynamic response of muscle deoxygenation (τ'[HHb+Mb]) was faster than τV̇O2p in all groups, resulting in Δ[HHb+Mb]/V̇O2p showing a transient "overshoot" relative to the subsequent steady-state level. After 3 wks, the Δ[HHb+Mb]/V̇O2p overshoot was eliminated only in the training groups, so that τ'[HHb+Mb] was not different to τV̇O2p in MICT and HIIT. The enhanced V̇O2 kinetics response consequent to both MICT and HIIT in T2D was likely attributed to a training-induced improvement in matching of O2 delivery to utilization.

Prior heavy-intensity exercise speeds V̇o2 kinetics during moderate-intensity exercise in young adults

2005 ◽  
Vol 98 (4) ◽  
pp. 1371-1378 ◽  
Author(s):  
Brendon J. Gurd ◽  
Barry W. Scheuermann ◽  
Donald H. Paterson ◽  
John M. Kowalchuk
Keyword(s):  
Young Adults ◽  
Lactate Threshold ◽  
Near Infrared ◽  
Linear Regression Analysis ◽  
Cycle Ergometer ◽  
Moderate Intensity ◽  
Moderate Intensity Exercise ◽  
Warm Up ◽  
Total Hemoglobin ◽  
Concentration Changes

The effect of prior heavy-intensity warm-up exercise on subsequent moderate-intensity phase 2 pulmonary O2 uptake kinetics (τV̇o2) was examined in young adults exhibiting relatively fast (FK; τV̇o2 < 30 s; n = 6) and slow (SK; τV̇o2 > 30 s; n = 6) V̇o2 kinetics in moderate-intensity exercise without prior warm up. Subjects performed four repetitions of a moderate (Mod1)-heavy-moderate (Mod2) protocol on a cycle ergometer with work rates corresponding to 80% estimated lactate threshold (moderate intensity) and 50% difference between lactate threshold and peak V̇o2 (heavy intensity); each transition lasted 6 min, and each was preceded by 6 min of cycling at 20 W. V̇o2 and heart rate (HR) were measured breath-by-breath and beat-by-beat, respectively; concentration changes of muscle deoxyhemoglobin (HHb), oxyhemoglobin, and total hemoglobin were measured by near-infrared spectroscopy (Hamamatsu NIRO 300). τV̇o2 was lower ( P < 0.05) in Mod2 than in Mod1 in both FK (20 ± 5 s vs. 26 ± 5 s, respectively) and SK (30 ± 8 s vs. 45 ± 11 s, respectively); linear regression analysis showed a greater “speeding” of V̇o2 kinetics in subjects exhibiting a greater Mod1 τV̇o2. HR, oxyhemoglobin, and total hemoglobin were elevated ( P < 0.05) in Mod2 compared with Mod1. The delay before the increase in HHb was reduced ( P < 0.05) in Mod2, whereas the HHb mean response time was reduced ( P < 0.05) in FK (Mod2, 22 ± 3 s; Mod1, 32 ± 11 s) but not different in SK (Mod2, 36 ± 13 s; Mod1, 34 ± 15 s). We conclude that improved muscle perfusion in Mod2 may have contributed to the faster adaptation of V̇o2, especially in SK; however, a possible role for metabolic inertia in some subjects cannot be overlooked.

scholarly journals Limitations of skeletal muscle oxygen delivery and utilization during moderate-intensity exercise in moderately impaired patients with chronic heart failure

2016 ◽  
Vol 311 (6) ◽  
pp. H1530-H1539 ◽  
Author(s):  
Victor M. Niemeijer ◽  
Ruud F. Spee ◽  
Thijs Schoots ◽  
Pieter F. F. Wijn ◽  
Hareld M. C. Kemps
Keyword(s):  
Heart Failure ◽  
Skeletal Muscle ◽  
Chronic Heart Failure ◽  
Near Infrared ◽  
Vastus Lateralis ◽  
Oxygen Uptake Kinetics ◽  
Moderate Intensity ◽  
Moderate Intensity Exercise ◽  
Exercise Tests ◽  
Mean Response Time

The extent and speed of transient skeletal muscle deoxygenation during exercise onset in patients with chronic heart failure (CHF) are related to impairments of local O2 delivery and utilization. This study examined the physiological background of submaximal exercise performance in 19 moderately impaired patients with CHF (Weber class A, B, and C) compared with 19 matched healthy control (HC) subjects by measuring skeletal muscle oxygenation (SmO2) changes during cycling exercise. All subjects performed two subsequent moderate-intensity 6-min exercise tests (bouts 1 and 2) with measurements of pulmonary oxygen uptake kinetics and SmO2 using near-infrared spatially resolved spectroscopy at the vastus lateralis for determination of absolute oxygenation values, amplitudes, kinetics (mean response time for onset), and deoxygenation overshoot characteristics. In CHF, deoxygenation kinetics were slower compared with HC (21.3 ± 5.3 s vs. 16.7 ± 4.4 s, P < 0.05, respectively). After priming exercise (i.e., during bout 2), deoxygenation kinetics were accelerated in CHF to values no longer different from HC (16.9 ± 4.6 s vs. 15.4 ± 4.2 s, P = 0.35). However, priming did not speed deoxygenation kinetics in CHF subjects with a deoxygenation overshoot, whereas it did reduce the incidence of the overshoot in this specific group ( P < 0.05). These results provide evidence for heterogeneity with respect to limitations of O2 delivery and utilization during moderate-intensity exercise in patients with CHF, with slowed deoxygenation kinetics indicating a predominant O2 utilization impairment and the presence of a deoxygenation overshoot, with a reduction after priming in a subgroup, indicating an initial O2 delivery to utilization mismatch.

High-intensity interval training speeds the adjustment of pulmonary O2 uptake, but not muscle deoxygenation, during moderate-intensity exercise transitions initiated from low and elevated baseline metabolic rates

2013 ◽  
Vol 114 (11) ◽  
pp. 1550-1562 ◽  
Author(s):  
Alexandra M. Williams ◽  
Donald H. Paterson ◽  
John M. Kowalchuk
Keyword(s):  
High Intensity ◽  
Time Course ◽  
Interval Training ◽  
Constant Load ◽  
Moderate Intensity ◽  
Metabolic Rates ◽  
High Intensity Interval Training ◽  
Moderate Intensity Exercise ◽  
Muscle Deoxygenation ◽  
High Intensity Interval

During step transitions in work rate (WR) within the moderate-intensity (MOD) exercise domain, pulmonary O2 uptake (V̇o2p) kinetics are slowed, and V̇o2p gain (ΔV̇o2p/ΔWR) is greater when exercise is initiated from an elevated metabolic rate. High-intensity interval training (HIT) has been shown to speed V̇o2p kinetics when step transitions to MOD exercise are initiated from light-intensity baseline metabolic rates. The effects of HIT on step transitions initiated from elevated metabolic rates have not been established. Therefore, this study investigated the effects of HIT on V̇o2p kinetics during transitions from low and elevated metabolic rates, within the MOD domain. Eight young, untrained men completed 12 sessions of HIT (spanning 4 wk). HIT consisted of 8–12 1-min intervals, cycling at a WR corresponding to 110% of pretraining maximal WR (WRmax). Pre-, mid- and posttraining, subjects completed a ramp-incremental test to determine maximum O2 uptake, WRmax, and estimated lactate threshold (θ̂L). Participants additionally completed double-step constant-load tests, consisting of step transitions from 20 W → Δ45% θ̂L [lower step (LS)] and Δ45 → 90% θ̂L [upper step (US)]. HIT led to increases in maximum O2 uptake ( P < 0.05) and WRmax ( P < 0.01), and τV̇o2p of both lower and upper MOD step transitions were reduced by ∼40% (LS: 24 s → 15 s; US: 45 s → 25 s) ( P < 0.01). However, the time course of adjustment of local muscle deoxygenation was unchanged in the LS and US. These results suggest that speeding of V̇o2p kinetics in both the LS and US may be due, in part, to an improved matching of muscle O2 utilization to microvascular O2 delivery within the working muscle following 12 sessions of HIT, although muscle metabolic adaptations cannot be discounted.

Effect of pedal cadence on the heterogeneity of muscle deoxygenation during moderate exercise

2013 ◽  
Vol 38 (12) ◽  
pp. 1206-1210 ◽  
Author(s):  
Houssem Zorgati ◽  
Katia Collomp ◽  
Virgile Amiot ◽  
Fabrice Prieur
Keyword(s):  
Near Infrared ◽  
Vastus Lateralis ◽  
Muscle Blood Flow ◽  
Relative Dispersion ◽  
Moderate Intensity ◽  
Delivery Ratio ◽  
Gas Exchanges ◽  
Muscle Deoxygenation ◽  
Exercise Muscle ◽  
Gas Exchange Threshold

This study examined the effect of pedal cadence on the heterogeneity of muscle deoxygenation during exercise of moderate intensity. Twelve healthy subjects performed 6 min of cycling at 40 and 100 r·min–1 at 80% of the workload corresponding to the gas exchange threshold. Gas exchanges were measured breath by breath during each exercise. Muscle deoxygenation (HHb, i.e., O2 extraction) was monitored continuously by near-infrared spectroscopy at eight sites on the vastus lateralis. The heterogeneity of HHb was assessed using the relative dispersion of the signal measured at the eight sites (i.e., 100 × standard deviation / mean). HHb was not altered by the pedal cadence, whereas pulmonary V̇O2 was higher at 100 r·min–1 than at 40 r·min–1 (p < 0.001). The relative dispersion of HHb was significantly higher at 100 r·min–1 than at 40 r·min–1 (p < 0.001). These results indicate that pedal cadence has no effect on O2 extraction but that an elevated cadence would increase muscle V̇O2, suggesting an increase in muscle blood flow. Elevated cadence also induced greater heterogeneity of the muscle’s V̇O2/Q̇O2 delivery ratio, suggesting a change in the adequacy between O2 demand and O2 delivery in some regions of active muscle.

O2 uptake kinetics, pyruvate dehydrogenase activity, and muscle deoxygenation in young and older adults during the transition to moderate-intensity exercise

2008 ◽  
Vol 294 (2) ◽  
pp. R577-R584 ◽  
Author(s):  
B. J. Gurd ◽  
S. J. Peters ◽  
G. J. F. Heigenhauser ◽  
P. J. LeBlanc ◽  
T. J. Doherty ◽  
...  
Keyword(s):  
Older Adults ◽  
Young Adults ◽  
Pyruvate Dehydrogenase ◽  
Lactate Threshold ◽  
Time Course ◽  
Moderate Intensity ◽  
Moderate Intensity Exercise ◽  
Similar Time ◽  
O2 Uptake Kinetics ◽  
The Relationship

The adaptation of pulmonary O2 uptake (V̇o2p) kinetics is slowed in older compared with young adults during the transition to moderate-intensity exercise. In this study, we examined the relationship between V̇o2p kinetics and mitochondrial pyruvate dehydrogenase (PDH) activity in young ( n = 7) and older ( n = 6) adults. Subjects performed cycle exercise to a work rate corresponding to ∼90% of estimated lactate threshold. Phase 2 V̇o2p kinetics were slower ( P < 0.05) in older (τ = 40 ± 17 s) compared with young (τ = 21 ± 6 s) adults. Relative phosphocreatine (PCr) breakdown was greater ( P < 0.05) at 30 s in older compared with young adults. Absolute PCr breakdown at 6 min was greater ( P < 0.05) in older compared with young adults. In young adults, PDH activity increased ( P < 0.05) from baseline to 30 s, with no further change observed at 6 min. In older adults, PDH activity during baseline exercise was similar to that seen in young adults. During the exercise transition, PDH activity did not increase ( P > 0.05) at 30 s of exercise but was elevated ( P < 0.05) after 6 min. The change in deoxyhemoglobin (HHb) was greater for a given V̇o2p in older adults, and there was a similar time course of HHb accompanying the slower V̇o2p kinetics in the older adults, suggesting a slower adaptation of bulk O2 delivery in older adults. In conclusion, the slower adaptation of V̇o2p in older adults is likely a result of both an increased metabolic inertia and lower O2 availability.

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