THE METABOLIC COST OF BREATHING FOR EXERCISE VENTILATIONS: EFFECTS OF AGE AND SEX.

Given there are both sex-based structural differences in the respiratory system and age-associated declines in pulmonary function, the purpose of this study was to assess the effects of age and sex on the metabolic cost of breathing (VO2RM) for exercise ventilations in healthy younger and older, males and females. METHODS: Forty healthy participants (10 young males 23±3yrs; 10 young females 23±3yrs; 10 older males 63±3yrs, 10 older females 63±6yrs) mimicked their exercise breathing patterns in the absence of exercise across a range of exercise intensities. RESULTS: At peak exercise, VO2RM represented a significantly greater fraction of peak oxygen consumption (VO2peak) in young females, 12.8±3.9%, compared to young males, 10.7±3.0% (P=0.027), while VO2RM represented 13.5±2.3% of VO2peak in older females and 13.2±3.3% in older males. At relative ventilations, there was a main effect of age, with older males consuming a significantly greater fraction of VO2RM (6.6%±1.9)than younger males (4.4%±1.3;P=0.012), and older females consuming a significantly greater fraction of VO2RM (6.9%±2.5)than younger females (5.1%±1.4;P=0.004) at 65% max. Furthermore, both younger and older males had significantly better respiratory muscle efficiency than their female counterparts at peak exercise (P=0.011;P=0.015). Similarly younger participants were significantly more efficient than older participants (6.5%±1.5% vs. 5.5±2.0%;P=0.001). CONCLUSION: Age-related changes in respiratory function, and sex-based differences in airway anatomy, influence the cost to breathe during exercise. It is possible the higher fraction of VO2RM during peak exercise predispose young females and older individuals to divert more blood flow to respiratory muscles at the expense of other muscles.

Sports Bra Restriction on Respiratory Mechanics during Exercise.

PURPOSE: We set out to understand how underband tightness or pressure of a sports bra relates to respiratory function and the mechanical work of breathing ( during exercise. Our secondary purpose was to quantify the effects of underband pressure on O 2 during submaximal running. METHODS: Nine highly trained female runners with normal pulmonary function completed maximal and submaximal running in three levels of underband restriction: loose, self-selected, and tight. RESULTS: During maximal exercise, we observed a significantly greater during the tight condition (350 ± 78 J·min -1 ) compared with the loose condition (301 ± 78 J·min -1 ; P < 0.05), and a 5% increase in minute ventilation ( ) during the tight condition compared with the loose condition ( P < 0.05). The pattern of breathing also differed between the two conditions; the greater maximal during the tight condition was achieved by a higher breathing frequency (57 ± 6 vs. 52 ± 7 breaths·min -1 ; P < 0.05), despite tidal volume being significantly lower in the tight condition compared with the loose condition (1.97 ± 0.20 vs. 2.05 ± 0.23 L; P < 0.05). During steady-state submaximal running, O 2 increased 1.3 ± 1.1% (range: -0.3 to 3.2%, P < 0.05) in the tight condition compared with the loose condition. CONCLUSIONS: Respiratory function may become compromised by the pressure exerted by the underband of a sports bra when women self-select their bra size. In the current study, loosening the underband pressure resulted in a decreased work of breathing, changed the ventilatory breathing pattern to deeper, less frequent breaths, and decreased submaximal oxygen uptake (improved running economy). Our findings suggest sports bra underbands can impair breathing mechanics during exercise and influence whole-body metabolic rate.

Exercise-induced arterial hypoxemia in female masters athletes.

The purpose of the study was to characterize exercise induced arterial hypoxemia (EIAH) in female masters athletes (FMA). We hypothesized that FMA would experience EIAH during treadmill running. Eight FMA (48-57 years) completed pulmonary function testing and an incremental exercise test until exhaustion (V̇O2max⁡ = 45.7 ± 6.5, range:35-54 ml/kg/min). On a separate day, the participants were instrumented with a radial arterial catheter and an esophageal temperature probe. Participants performed three to four constant load exercise tests at 60-70 %, 75 %, 90 %, 95 %, and 100 % of maximal oxygen uptake while sampling arterial blood and recording esophageal temperature. We found that FMA decrease their partial pressure of oxygen (86.0 ± 7.6, range:73-108 mmHg), arterial saturation (96.2 ± 1.2, range:93-98 %), and widen their alveolar to arterial oxygen difference (23.2 ± 8.8, range:5-42 mmHg) during all exercise intensities however, with variability in terms of severity and pattern. Our findings suggest that FMA experience EIAH however aerobic fitness appears unrelated to occurrence or severity (r = 0.13, p = 0.756).

It is time to abandon single-value oxygen uptake energy equivalents.

Physiologists commonly use single-value energy equivalents (e.g., 20.1 kJ/LO2 and 20.9 kJ/LO2) to convert oxygen uptake (V̇o2) to energy, but doing so ignores how the substrate oxidation ratio (carbohydrate:fat) changes across aerobic intensities. Using either 20.1 kJ/LO2 or 20.9 kJ/LO2 can incur systematic errors of up to 7%. In most circumstances, the best approach for estimating energy expenditure is to measure both V̇o2 and V̇co2 and use accurate, species-appropriate stoichiometry. However, there are circumstances when V̇co2 measurements may be unreliable. In those circumstances, we recommend that the research report or compare only V̇o2.NEW & NOTEWORTHY We quantify that the common practice of using single-value oxygen uptake energy equivalents for exercising subjects can incur systematic errors of up to 7%. We argue that such errors can be greatly reduced if researchers measure both V̇o2 and V̇co2 and adopt appropriate stoichiometry equations.

Effects of sex and ageing on the human respiratory muscle metaboreflex.

Intense inspiratory muscle work evokes a sympathetically mediated pressor reflex, termed the respiratory muscle metaboreflex, in which young females demonstrate an attenuated response relative to males. However, the effects of ageing and female sex hormones on the respiratory muscle metaboreflex are unclear. We tested the hypothesis that the pressor response to inspiratory work would be similar between older males and females, and higher relative to their younger counterparts. Healthy, normotensive young (26 ± 3 years) males (YM; n = 10) and females (YF; n = 10), as well as older (64 ± 5 years) males (OM; n = 10) and females (OF; n = 10), performed inspiratory pressure threshold loading (PTL) to task failure. Older adults had a greater mean arterial pressure (MAP) response to PTL than young (P < 0.001). YF had a lower MAP compared to YM (+10 ± 6 vs. +19 ± 15 mmHg, P = 0.026); however, there was no difference observed between OF and OM (+26 ± 11 vs. +27 ± 11 mmHg, P = 0.162). Older adults had a lower heart rate response to PTL than young (P = 0.002). There was no effect of sex between young females and males (+19 ± 9 and +27 ± 11 bpm, P = 0.186) or older females and males (+17 ± 7 and +20 ± 7 bpm, P = 0.753). We conclude the respiratory muscle metaboreflex response is heightened in older adults, and the sex effect between older males and post-menopause females is absent, suggesting an effect of circulating sex hormones. KEY POINTS: The arterial blood pressure response to the respiratory muscle metaboreflex is greater in older males and females. Compared to sex-matched young individuals, there is no sex differences in the blood pressure response between older males and post-menopause females. Our results suggest the differences between males and females in the cardiovascular response to high levels of inspiratory muscle work is abolished with reduced circulating female sex hormones.

The effect of proportional assist ventilation on the electrical activity of the human diaphragm during exercise.

NEW FINDINGS: What is the central question of this study? What is the effect of lowering the normally occurring work of breathing on the electrical activity and pressure generated by the diaphragm during submaximal exercise in healthy humans? What is the main finding and its importance? Ventilatory assist during exercise elicits a proportional lowering of both the work performed by the diaphragm and diaphragm electrical activity. These findings have implications for exercise training studies using proportional assist ventilation to reduce diaphragm work in patients with cardiopulmonary disease. ABSTRACT: We hypothesized that when a proportional assist ventilator (PAV) is applied in order to reduce the pressure generated by the diaphragm, there would be a corresponding reduction in electrical activity of the diaphragm. Healthy participants (five male and four female) completed an incremental cycle exercise test to exhaustion in order to calculate workloads for subsequent trials. On the experimental day, participants performed submaximal cycling, and three levels of assisted ventilation were applied (low, medium and high). Ventilatory parameters, pulmonary pressures and EMG of the diaphragm (EMGdi ) were obtained. To compare the PAV conditions with spontaneous breathing intervals, ANOVA procedures were used, and significant effects were evaluated with a Tukey-Kramer test. Significance was set at P < 0.05. The work of breathing was not different between the lowest level of unloading and spontaneous breathing (P = 0.151) but was significantly lower during medium (25%, P = 0.02) and high (36%, P < 0.001) levels of PAV. The pressure-time product of the diaphragm (PTPdi ) was lower across PAV unloading conditions (P < 0.05). The EMGdi was significantly lower in medium and high PAV conditions (P = 0.035 and P < 0.001, respectively). The mean reductions of EMGdi with PAV unloading were 14, 22 and 39%, respectively. The change in EMGdi for a given lowering of PTPdi with the PAV was significantly correlated (r = 0.61, P = 0.01). Ventilatory assist during exercise elicits a reduction in the electrical activity of the diaphragm, and there is a proportional lowering of the work of breathing. Our findings have implications for exercise training studies using assisted ventilation to reduce diaphragm work in patients with cardiopulmonary disease.

Perceived exertion and dyspnea while cycling during a hypoxic and hyperoxic placebo.

Rating of perceived exertion (RPE) is used to subjectively quantify the perception of physical activity, breathlessness or dyspnea, and leg discomfort (RPElegs) during exercise. However, it is unknown how dyspnea or RPElegs can be influenced by expectations. Thirty healthy, active participants (19 males, 11 females) completed five, 5-minute submaximal cycling trials at 60% peak work rate. We deceived participants by telling them they were inspiring different hypoxic and hyperoxic gases, when in fact they breathed room air. Cardiorespiratory variables were similar between the trials, however, dyspnea and RPElegs evaluated with a Borg scale changed in a dose-response manner. When participants believed they were breathing 15% O2, they significantly increased dyspnea +0.70 ± 0.2 units (p = 0.03) compared to room air, whereas RPElegs was unchanged +0.35 ± 0.1 units (p = 0.70). When participants believed they were breathing 15% O2, they significantly increased dyspnea +1.05 ± 0.4 units (p = 0.003) compared to 23% hyperoxic condition, whereas RPElegs was unchanged +0.35 ± 0.1 units (p = 0.70). We found that dyspnea during exercise is susceptible to expectancy, without any accompanying physiological changes. Given coaches and clinicians use perceived exertion to prescribe exercise intensity and evaluate treatments, our findings show that the effect of expectations must be considered when interpreting sensations of breathlessness.

Predictors of Expiratory Flow Limitation during Exercise in Healthy Males and Females.

RATIONALE: It is unclear whether the frequency and mechanisms of expiratory flow limitation (EFL) during exercise differ between males and females. PURPOSE: This study aimed to determine which factors predispose individuals to EFL during exercise and whether these factors differ based on sex. We hypothesized that i) EFL frequency would be similar in males and females and ii) in females, EFL would be associated with indices of low ventilatory capacity, whereas in males, EFL would be associated with indices of high ventilatory demand. METHODS: Data from n = 126 healthy adults (20-45 y, n = 60 males, n = 66 females) with a wide range of cardiorespiratory fitness (81%-182% predicted maximal oxygen uptake) were included in the study. Participants performed spirometry and an incremental cycle exercise test to exhaustion. Standard cardiorespiratory variables were assessed throughout exercise. The tidal flow-volume overlap method was used to assess EFL based on a minimum threshold of 5% overlap between the tidal and the maximum expiratory flow-volume curves. Predictors of EFL during exercise were determined via multiple logistical regression using anthropometric, pulmonary function, and peak exercise data. RESULTS: During exercise, EFL occurred in 49% of participants and was similar between the sexes (females = 45%, males = 53%; P = 0.48). In males, low forced expired flow between 25% and 75% of forced vital capacity and high slope ratio as well as low end-expiratory lung volume, high breathing frequency, and high relative tidal volume at peak exercise were associated with EFL ( P < 0.001; Nagelkerke R2 = 0.73). In females, high slope ratio, high breathing frequency, and tidal volume at peak exercise were associated with EFL ( P < 0.001; Nagelkerke R2 = 0.61). CONCLUSIONS: Despite sex differences in respiratory system morphology, the frequency and the predictors of EFL during exercise do not substantially differ between the sexes.

Can We Quantify the Benefits of "Super Spikes" in Track Running?

The recent and rapid developments in track spike innovation have been followed by a wave of record-breaking times and top performances. This has led many to question what role "super spikes" play in improving running performance. To date, the specific contributions of new innovations in footwear, including lightweight, resilient, and compliant midsole foam, altered geometry, and increased longitudinal bending stiffness, to track running performance are unknown. Based on current literature, we speculate about what advantages these features provide. Importantly, the effects of super spikes will vary based on several factors including the event (e.g., 100 m vs. 10,000 m) and the characteristics of the athlete wearing them. Further confounding our understanding of super spikes is the difficulty of testing them. Unlike marathon shoes, testing track spikes comes with a unique challenge of quantifying the metabolic energy demands of middle-distance running events, which are partly anaerobic. Quantifying the exact benefits from super spikes is difficult and we may need to rely on comparison of track performances pre- and post- the introduction of super spikes.

Respiratory modulation of sympathetic vasomotor outflow during graded leg cycling.

Respiratory modulation of sympathetic vasomotor outflow to skeletal muscles (muscle sympathetic nerve activity; MSNA) occurs in resting humans. Specifically, MSNA is highest at end-expiration and lowest at end-inspiration during quiet, resting breathing. We tested the hypothesis that within-breath modulation of MSNA would be amplified during graded leg cycling. Thirteen (n = 3 females) healthy young (age: 25.2 ± 4.7 yr) individuals completed all testing. MSNA (right median nerve) was measured at rest (baseline) and during semirecumbent cycle exercise at 40%, 60%, and 80% of maximal workload (Wmax). MSNA burst frequency (BF) was 20.0 ± 4.0 bursts/min at baseline and was not different during exercise at 40%Wmax (21.3 ± 3.7 bursts/min; P = 0.292). Thereafter, MSNA BF increased significantly compared with baseline (60%Wmax: 31.6 ± 5.8 bursts/min; P < 0.001, 80%Wmax: 44.7 ± 5.3 bursts/min; P < 0.001). At baseline and all exercise intensities, MSNA BF was lowest at end-inspiration and greatest at mid-to-end expiration. The within-breath change in MSNA BF (ΔMSNA BF; end-expiration minus end-inspiration) gradually increased from baseline to 60%Wmax leg cycling, but no further increase appeared at 80%Wmax exercise. Our results indicate that within-breath modulation of MSNA is amplified from baseline to moderate intensity during dynamic exercise in young healthy individuals, and that no further potentiation occurs at higher exercise intensities. Our findings provide an important extension of our understanding of respiratory influences on sympathetic vasomotor control.NEW & NOTEWORTHY Within-breath modulation of sympathetic vasomotor outflow to skeletal muscle (muscle sympathetic nerve activity; MSNA) occurs in spontaneously breathing humans at rest. It is unknown if respiratory modulation persists during dynamic whole body exercise. We found that MSNA burst frequency was lowest at end-inspiration and highest at mid-to-end expiration during rest and graded leg cycling. Respiratory modulation of sympathetic vasomotor outflow remains intact and is amplified during dynamic whole body exercise.

An integrative approach to the pulmonary physiology of exercise: when does biological sex matter?

Historically, many studies investigating the pulmonary physiology of exercise (and biomedical research in general) were performed exclusively or predominantly with male research participants. This has led to an incomplete understanding of the pulmonary response to exercise. More recently, important sex-based differences with respect to the human respiratory system have been identified. The purpose of this review is to summarize current findings related to sex-based differences in the pulmonary physiology of exercise. To that end, we will discuss how morphological sex-based differences of the respiratory system affect the respiratory response to exercise. Moreover, we will discuss sex-based differences of the physiological integrative responses to exercise, and how all these differences can influence the regulation of breathing. We end with a brief discussion of pregnancy and menopause and the accompanying ventilatory changes observed during exercise.

Partitioning the work of breathing during running and cycling using optoelectronic plethysmography.

Work of breathing ([Formula: see text]) derived from a single lung volume and pleural pressure is limited and does not fully characterize the mechanical work done by the respiratory musculature. It has long been known that abdominal activation increases with increasing exercise intensity, yet the mechanical work done by these muscles is not reflected in [Formula: see text]. Using optoelectronic plethysmography (OEP), we sought to show first that the volumes obtained from OEP (VCW) were comparable to volumes obtained from flow integration (Vt) during cycling and running, and second, to show that partitioned volume from OEP could be utilized to quantify the mechanical work done by the rib cage ([Formula: see text]RC) and abdomen ([Formula: see text]AB) during exercise. We fit 11 subjects (6 males/5 females) with reflective markers and balloon catheters. Subjects completed an incremental ramp cycling test to exhaustion and a series of submaximal running trials. We found good agreement between VCW versus Vt during cycling (bias = 0.002; P > 0.05) and running (bias = 0.016; P > 0.05). From rest to maximal exercise,[Formula: see text]AB increased by 84% (range: 30%-99%; [Formula: see text]AB: 1 ± 1 J/min to 61 ± 52 J/min). The relative contribution of the abdomen increased from 17 ± 9% at rest to 26 ± 16% during maximal exercise. Our study highlights and provides a quantitative measure of the role of the abdominal muscles during exercise. Incorporating the work done by the abdomen allows for a greater understanding of the mechanical tasks required by the respiratory muscles and could provide further insight into how the respiratory system functions during disease and injury.NEW & NOTEWORTHY We demonstrated that optoelectronic plethysmography (OEP) is a reliable tool to determine ventilatory volume changes during cycling and running, without restricting natural upper arm movements. Second, using OEP volumes coupled with pressure-derived measures, we calculated the work done by the rib cage and abdomen, respectively, during exercise. Collectively, our findings indicate that pulmonary mechanics can be accurately quantified using OEP, and abdominal work performed during ventilation contributes substantially to the overall work of the respiratory musculature.

Extrapolating Metabolic Savings in Running: Implications for Performance Predictions.

Training, footwear, nutrition, and racing strategies (i.e., drafting) have all been shown to reduce the metabolic cost of distance running (i.e., improve running economy). However, how these improvements in running economy (RE) quantitatively translate into faster running performance is less established. Here, we quantify how metabolic savings translate into faster running performance, considering both the inherent rate of oxygen uptake-velocity relation and the additional cost of overcoming air resistance when running overground. We collate and compare five existing equations for oxygen uptake-velocity relations across wide velocity ranges. Because the oxygen uptake vs. velocity relation is non-linear, for velocities slower than ∼3 m/s, the predicted percent improvement in velocity is slightly greater than the percent improvement in RE. For velocities faster than ∼3 m/s, the predicted percent improvement in velocity is less than the percent improvements in RE. At 5.5 m/s, i.e., world-class marathon pace, the predicted percent improvement in velocity is ∼2/3rds of the percent improvement in RE. For example, at 2:04 marathon pace, a 3% improvement in RE translates to a 1.97% faster velocity or 2:01:36, almost exactly equal to the recently set world record.

The Biomechanics of Competitive Male Runners in Three Marathon Racing Shoes: A Randomized Crossover Study.

BACKGROUND: We have shown that a prototype marathon racing shoe reduced the metabolic cost of running for all 18 participants in our sample by an average of 4%, compared to two well-established racing shoes. Gross measures of biomechanics showed minor differences and could not explain the metabolic savings. OBJECTIVE: To explain the metabolic savings by comparing the mechanics of the shoes, leg, and foot joints during the stance phase of running. METHODS: Ten male competitive runners, who habitually rearfoot strike ran three 5-min trials in prototype shoes (NP) and two established marathon shoes, the Nike Zoom Streak 6 (NS) and the adidas adizero Adios BOOST 2 (AB), at 16 km/h. We measured ground reaction forces and 3D kinematics of the lower limbs. RESULTS: Hip and knee joint mechanics were similar between the shoes, but peak ankle extensor moment was smaller in NP versus AB shoes. Negative and positive work rates at the ankle were lower in NP shoes versus the other shoes. Dorsiflexion and negative work at the metatarsophalangeal (MTP) joint were reduced in the NP shoes versus the other shoes. Substantial mechanical energy was stored/returned in compressing the NP midsole foam, but not in bending the carbon-fiber plate. CONCLUSION: The metabolic savings of the NP shoes appear to be due to: (1) superior energy storage in the midsole foam, (2) the clever lever effects of the carbon-fiber plate on the ankle joint mechanics, and (3) the stiffening effects of the plate on the MTP joint.

Does Metabolic Rate Increase Linearly with Running Speed in all Distance Runners?

Running economy (oxygen uptake or metabolic rate for running at a submaximal speed) is one of the key determinants of distance running performance. Previous studies reported linear relationships between oxygen uptake or metabolic rate and speed, and an invariant cost of transport across speed. We quantified oxygen uptake, metabolic rate, and cost of transport in 10 average and 10 sub-elite runners. We increased treadmill speed by 0.45 m · s -1 from 1.78 m · s -1 (day 1) and 2.01 m · s -1 (day 2) during each subsequent 4-min stage until reaching a speed that elicited a rating of perceived exertion of 15. Average runners' oxygen uptake and metabolic rate vs. speed relationships were best described by linear fits. In contrast, the sub-elite runners' relationships were best described by increasing curvilinear fits. For the sub-elites, oxygen cost of transport and energy cost of transport increased by 12.8% and 9.6%, respectively, from 3.58 to 5.14 m · s -1 . Our results indicate that it is not possible to accurately predict metabolic rates at race pace for sub-elite competitive runners from data collected at moderate submaximal running speeds (2.68-3.58 m · s -1 ). To do so, metabolic rate should be measured at speeds that approach competitive race pace and curvilinear fits should be used for extrapolation to race pace.

What determines the metabolic cost of human running across a wide range of velocities?

The 'cost of generating force' hypothesis proposes that the metabolic rate during running is determined by the rate of muscle force development (1/tc, where tc=contact time) and the volume of active leg muscle. A previous study assumed a constant recruited muscle volume and reported that the rate of force development alone explained ∼70% of the increase in metabolic rate for human runners across a moderate velocity range (2-4 m s-1). We hypothesized that over a wider range of velocities, the effective mechanical advantage (EMA) of the lower limb joints would overall decrease, necessitating a greater volume of active muscle recruitment. Ten high-caliber male human runners ran on a force-measuring treadmill at 8, 10, 12, 14, 16 and 18 km h-1 while we analyzed their expired air to determine metabolic rates. We measured ground reaction forces and joint kinematics to calculate contact time and estimate active muscle volume. From 8 to 18 km h-1, metabolic rate increased 131% from 9.28 to 21.44 W kg-1tc decreased from 0.280 s to 0.190 s, and thus the rate of force development (1/tc) increased by 48%. Ankle EMA decreased by 19.7±11%, knee EMA increased by 11.1±26.9% and hip EMA decreased by 60.8±11.8%. Estimated active muscle volume per leg increased 52.8% from 1663±152 cm3 to 2550±169 cm3 Overall, 98% of the increase in metabolic rate across the velocity range was explained by just two factors: the rate of generating force and the volume of active leg muscle.

Comparison of running and cycling economy in runners, cyclists, and triathletes.

PURPOSE: Exercise economy is one of the main physiological factors determining performance in endurance sports. Running economy (RE) can be improved with running-specific training, while the improvement of cycling economy (CE) with cycling-specific training is controversial. We investigated whether exercise economy reflects sport-specific skills/adaptations or is determined by overall physiological factors. METHODS: We compared RE and CE in 10 runners, 9 cyclists and 9 triathletes for running at 12 km/h and cycling at 200 W. Gross rates of oxygen consumption and carbon dioxide production were collected and used to calculate gross metabolic rate in watts for both running and cycling. RESULTS: Runners had better RE than cyclists (917 ± 107 W vs. 1111 ± 159 W) (p < 0.01). Triathletes had intermediate RE values (1004 ± 98 W) not different from runners or cyclists. CE was not different (p = 0.20) between the three groups (runners: 945 ± 60 W; cyclists: 982 ± 44 W; triathletes: 979 ± 54 W). CONCLUSION: RE can be enhanced with running-specific training, but CE is independent of cycling-specific training.

Calculating metabolic energy expenditure across a wide range of exercise intensities: the equation matters.

We compared 10 published equations for calculating energy expenditure from oxygen consumption and carbon dioxide production using data for 10 high-caliber male distance runners over a wide range of running velocities. We found up to a 5.2% difference in calculated metabolic rate between 2 widely used equations. We urge our fellow researchers abandon out-of-date equations with published acknowledgments of errors or inappropriate biochemical/physical assumptions.