Gas Exchange in Exercise

Respiratory Physiology > Gas Exchange > Physiological Principles of Pulmonary Gas Exchange

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Paolo Cerretelli, Pietro E. Di Prampero

10.1002/cphy.cp030416

Source: Supplement 13: Handbook of Physiology, The Respiratory System, Gas Exchange

Originally published: 1987

Published online: January 2011

Full Article on Wiley Online Library

Abstract
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References

Abstract

The sections in this article are:

1 Steady State

1.1 External Gas Exchange

1.2 Alveolar Gas Exchange

1.3 Blood O2‐Carrying Capacity

1.4 Energy Cost of Blood Pumping

1.5 Cardiovascular Adjustments

1.6 Oxygen Utilization and its Limiting Factors

1.7 Comparative Aspects of Gas Exchange

2 Unsteady State

2.1 Pulmonary Gas‐Exchange Transients

2.2 Cardiovascular Transients

2.3 Control and Kinetics of Gas Exchange in Muscle

3 Conclusions

	Figure 1. Increase of overall O₂ uptake per unit increase of expired minute ventilation (, kJ/liter) in normoxia (N, •, –) and in chronic hypoxia (H, ○, –––; 5,800 m). Data from Pugh et al. 390. O₂ cost of ventilation (Δ /η Δ ) in normoxia (N, ‐, ▪, ▴) and in chronic hypoxia (H, ‐ ‐ ‐), as calculated on the basis of efficiency values (η) of 0.25 (lower curves) and 0.1 (upper curves). [Data from Cruz 114, Margaria et al. 336, and Thoden et al. 470.] Pulmonary ventilation expressed at body temperature, ambient pressure, saturated with water vapor in liters per minute. “Critical level” for exercise ventilation is ∼15 liters/min lower in hypoxia (A' and B') than in normoxia (A and B). This indicates that in hypoxia the fraction of absolute utilized by respiratory muscles is necessarily greater than in normoxia
	Figure 2. Mean arterial O₂ partial pressure (Pa_o2, •) and (○) values (±SD) as a function of O₂ consumption (). Thin lines, calculated alveolar values. Shaded areas, alveolar‐arterial difference in partial pressure of O₂ ( − ) and CO₂ ( − ) , P < 0.05. , P < 0.01. **, P < 0.001. Data from Table 1 (ref. 133a not included)
	Figure 3. Mean (±SD) diffusing capacity for O₂ (D_o2; 44 subj.) and D_co (318 subj.) as a function of by various methods. , P < 0.05. , P < 0.01. **, P < 0.001. (Data from refs. 13,35,48,55,75,119,149,169,199,265,269,326,337,359,363,387,392,402,415,461,478,511.)
	Figure 4. Myocardial O₂ consumption (M_o2) per 100 g left ventricular tissue as a function of overall O₂ consumption (), cardiac output (), and left ventricular blood flow (LV) at rest (•) and exercise (○). (a ‐ v)_o2, Arteriovenous O₂ difference in milliliters of O₂ per 100 ml of blood. (Data from refs. 52,241,280,346,347,397.)
	Figure 5. Cardiac output () as a function of O₂ uptake () for different groups of subjects: • boys (11‐14 yr); O, girls (11‐14 yr); ⋄, adult males; ⋄, adult females; +, male athletes; x, female athletes. Group Vmax_o2 values shown by ↑. Isopleths showing arterial‐mixed‐venous O₂ difference in milli‐liters of O₂ per 100 ml of blood, ()_ov are also drawn. (Data from refs. 26,36,86,155,161,201,217,227,430.)
	Figure 6. A: heart rate (HR) as a function of . B: relationship between stroke volume (q_st) and cardiac output (). Isopleth HR lines: 50, 100, 200/min. C: cardiac output as a function of . Isopleths show arterial‐mixed‐venous O₂ difference: 5, 10, 15 ml/100 ml. • Walking and running (P. Cerretelli, unpublished observations); ▪, during aquatic activity 400; x, in hot environments 419; ⋄, during arm exercise 51; ○, during arm and leg exercise 51; o, during isometric (I) exercise (148; P. Cerretelli, unpublished observations).
	Figure 7. Maximum O₂ consumption [max_o2 (ml·kg⁻¹·min⁻¹), lower panel] and ratio of max_o2 to resting O₂ consumption (rest_o2, upper panel) as a function of body mass [M (g, log scale)] for various mammals. U, untrained; T, trained; S, sedentary; A, athlete. Data for small rodents 312,358,370,413,414,438; data for average‐size mammals (springhares, pigs, baby lions: 100, 437, 438, 467); data for the dog 32,90,101,145,253,325,357,368,379,410,437,438,484,510; data for trained horses (G. Cortili, unpublished observations); data for untrained horses 471; data for humans 303
	Figure 8. Breath‐by‐breath O₂ uptake (V_o₂, Δ) and CO₂ output (V_co₂, ○) as a function of time during very strenuous exercise (treadmill running at 18 km/h, on a +15% incline, exhaustion time = 22 s) and in recovery period. Exhausting exercise was preceded by ∼10‐min walk at 4.5 km/h on a +15% incline. (Note 3 different time scales.)
	Figure 9. Output of CO₂ in excess of O₂ uptake (excess V_co₂) as a function of peak blood lactate concentration above resting (Δ_ab) in recovery after very strenuous exercise (treadmill running at 18 km/h; +10%, +15%, and +20% incline; exhaustion time 14‐50 s) in 4 subjects. Excess V_co₂ was calculated as V_co₂ · dt ‐ V_o₂ · dt, where t₁ is the time at which V_co₂ = V_o₂, usually ∼10 min (see Fig. 8).
	Figure 10. Excess V_co₂ as a function of corresponding PA_co₂ in 4 subjects (○, x, ○, ⋄). Excess V_co₂ is here the amount of CO₂ eliminated in excess of O₂ uptake, up to the moment at which PA_co₂ was as indicated. Thus, as recovery proceeds, the subjects' respiratory conditions move toward the left, along the curve; points to the extreme left correspond to ∼10 min in all subjects. Three subjects are characterized by the same function; 4th subject (⋄) follows a markedly different pattern. For a given excess V_co₂, drop in [] can be considered equal in all subjects (see Fig. 9). Therefore, this last subject (⋄), having a higher PA_co₂ for a given excess V_co₂, is presumably characterized by a lesser sensitivity to CO₂.
	Figure 11. Time course of energetic processes at onset of exercise in humans. Ordinate in arbitrary units: 1 = Vmax_o₂. Top: energy requirement of exercise (E) is 0.8 Vmax_o₂. Curve 1, time course of energy yield at muscle level, resulting from sum of V_o₂ through the mouth (curve 3, t_1/2 = 40 s), early lactate production, and O₂‐stores depletion. Amount of energy contributed by latter 2 factors is indicated by areas labeled VeLa_o₂ and ΔVst_o₂, respectively. Area labeled V_o₂, amount of O₂ taken in through the mouth during exercise. (For mild aerobic exercise eLa = 0, so that curve 3 overlaps curve 2.) Difference between energy requirement of exercise (constant from time 0) and curve 1 must be supplied by net PC breakdown. Alactic O₂ debt (VPC_o₂), amount of phosphocreatine (PC) split in O₂ equivalents. Curve 1, time course of VPC_o₂ contraction; curve 1′, time course of VPC_o₂ payment; t_1/2 of curves 1 and 1' = 15 s. No La production occurs in recovery; thus curves 2' and 3' coincide (t_1/2 of curves 2 and 2' = 25 s). Bottom: energy requirement of exercise is 2.33 Vmax_o₂; duration of exercise to exhaustion is 1.15 min. Curve 1, time course of energy yield at muscle level, resulting from sum of V_o₂ through the mouth (curve 2) and O₂‐stores depletion. Lactate production (——) begins when V_o₂ at muscle level has reached its maximum value (areas La, amount of energy supplied by lactic sources). (Time course of La production, as well as time at which glycolysis begins, is hypothetical.) After exercise, VPC_o₂ is paid by the oxidative processes that remain at maximum level for ∼25 s; t_1/2 of curves 1 and 1' = 15 s (asymptote of curve 1 = energy requirement); t_1/2 of curve 2 = 30 s (asymptote = energy requirement); t_1/2 of curve 2' = 25 s. Adapted from di Prampero 137
	Figure 12. Model for control of muscle via mitochondrial creatinephosphokinase (CPK). Subscripts: e, extramitochondrial; i, intramitochondrial; m, mitochondrial. Glyc., glycogen. Adapted from Mahler (329,528)

Figure 1.

Increase of overall O₂ uptake per unit increase of expired minute ventilation (, kJ/liter) in normoxia (N, •, –) and in chronic hypoxia (H, ○, –––; 5,800 m).

Data from Pugh et al. 390. O₂ cost of ventilation (Δ /η Δ ) in normoxia (N, ‐, ▪, ▴) and in chronic hypoxia (H, ‐ ‐ ‐), as calculated on the basis of efficiency values (η) of 0.25 (lower curves) and 0.1 (upper curves). [Data from Cruz 114, Margaria et al. 336, and Thoden et al. 470.] Pulmonary ventilation expressed at body temperature, ambient pressure, saturated with water vapor in liters per minute. “Critical level” for exercise ventilation is ∼15 liters/min lower in hypoxia (A' and B') than in normoxia (A and B). This indicates that in hypoxia the fraction of absolute utilized by respiratory muscles is necessarily greater than in normoxia

Figure 2.

Mean arterial O₂ partial pressure (Pa_o2, •) and (○) values (±SD) as a function of O₂ consumption (). Thin lines, calculated alveolar values. Shaded areas, alveolar‐arterial difference in partial pressure of O₂ ( − ) and CO₂ ( − ) *, P < 0.05. **, P < 0.01. ***, P < 0.001.

Data from Table 1 (ref. 133a not included)

Figure 3.

Mean (±SD) diffusing capacity for O₂ (D_o2; 44 subj.) and D_co (318 subj.) as a function of by various methods. *, P < 0.05. **, P < 0.01. ***, P < 0.001. (Data from refs. 13,35,48,55,75,119,149,169,199,265,269,326,337,359,363,387,392,402,415,461,478,511.)

Figure 4.

Myocardial O₂ consumption (M_o2) per 100 g left ventricular tissue as a function of overall O₂ consumption (), cardiac output (), and left ventricular blood flow (LV) at rest (•) and exercise (○). (a ‐ v)_o2, Arteriovenous O₂ difference in milliliters of O₂ per 100 ml of blood. (Data from refs. 52,241,280,346,347,397.)

Figure 5.

Cardiac output () as a function of O₂ uptake () for different groups of subjects: • boys (11‐14 yr); O, girls (11‐14 yr); ⋄, adult males; ⋄, adult females; +, male athletes; x, female athletes. Group Vmax_o2 values shown by ↑. Isopleths showing arterial‐mixed‐venous O₂ difference in milli‐liters of O₂ per 100 ml of blood, ()_ov are also drawn. (Data from refs. 26,36,86,155,161,201,217,227,430.)

Figure 6.

A: heart rate (HR) as a function of . B: relationship between stroke volume (q_st) and cardiac output (). Isopleth HR lines: 50, 100, 200/min. C: cardiac output as a function of . Isopleths show arterial‐mixed‐venous O₂ difference: 5, 10, 15 ml/100 ml. • Walking and running (P. Cerretelli, unpublished observations); ▪, during aquatic activity 400; x, in hot environments 419; ⋄, during arm exercise 51; ○, during arm and leg exercise 51; o, during isometric (I) exercise (148; P. Cerretelli, unpublished observations).

Figure 7.

Maximum O₂ consumption [max_o2 (ml·kg⁻¹·min⁻¹), lower panel] and ratio of max_o2 to resting O₂ consumption (rest_o2, upper panel) as a function of body mass [M (g, log scale)] for various mammals. U, untrained; T, trained; S, sedentary; A, athlete.

Data for small rodents 312,358,370,413,414,438; data for average‐size mammals (springhares, pigs, baby lions: 100, 437, 438, 467); data for the dog 32,90,101,145,253,325,357,368,379,410,437,438,484,510; data for trained horses (G. Cortili, unpublished observations); data for untrained horses 471; data for humans 303

Figure 8.

Breath‐by‐breath O₂ uptake (V_o₂, Δ) and CO₂ output (V_co₂, ○) as a function of time during very strenuous exercise (treadmill running at 18 km/h, on a +15% incline, exhaustion time = 22 s) and in recovery period. Exhausting exercise was preceded by ∼10‐min walk at 4.5 km/h on a +15% incline. (Note 3 different time scales.)

Figure 9.

Output of CO₂ in excess of O₂ uptake (excess V_co₂) as a function of peak blood lactate concentration above resting (Δ_ab) in recovery after very strenuous exercise (treadmill running at 18 km/h; +10%, +15%, and +20% incline; exhaustion time 14‐50 s) in 4 subjects. Excess V_co₂ was calculated as V_co₂ · dt ‐ V_o₂ · dt, where t₁ is the time at which V_co₂ = V_o₂, usually ∼10 min (see Fig. 8).

Figure 10.

Excess V_co₂ as a function of corresponding PA_co₂ in 4 subjects (○, x, ○, ⋄). Excess V_co₂ is here the amount of CO₂ eliminated in excess of O₂ uptake, up to the moment at which PA_co₂ was as indicated. Thus, as recovery proceeds, the subjects' respiratory conditions move toward the left, along the curve; points to the extreme left correspond to ∼10 min in all subjects. Three subjects are characterized by the same function; 4th subject (⋄) follows a markedly different pattern. For a given excess V_co₂, drop in [] can be considered equal in all subjects (see Fig. 9). Therefore, this last subject (⋄), having a higher PA_co₂ for a given excess V_co₂, is presumably characterized by a lesser sensitivity to CO₂.

Figure 11.

Time course of energetic processes at onset of exercise in humans. Ordinate in arbitrary units: 1 = Vmax_o₂. Top: energy requirement of exercise (E) is 0.8 Vmax_o₂. Curve 1, time course of energy yield at muscle level, resulting from sum of V_o₂ through the mouth (curve 3, t_1/2 = 40 s), early lactate production, and O₂‐stores depletion. Amount of energy contributed by latter 2 factors is indicated by areas labeled VeLa_o₂ and ΔVst_o₂, respectively. Area labeled V_o₂, amount of O₂ taken in through the mouth during exercise. (For mild aerobic exercise eLa = 0, so that curve 3 overlaps curve 2.) Difference between energy requirement of exercise (constant from time 0) and curve 1 must be supplied by net PC breakdown. Alactic O₂ debt (VPC_o₂), amount of phosphocreatine (PC) split in O₂ equivalents. Curve 1, time course of VPC_o₂ contraction; curve 1′, time course of VPC_o₂ payment; t_1/2 of curves 1 and 1' = 15 s. No La production occurs in recovery; thus curves 2' and 3' coincide (t_1/2 of curves 2 and 2' = 25 s). Bottom: energy requirement of exercise is 2.33 Vmax_o₂; duration of exercise to exhaustion is 1.15 min. Curve 1, time course of energy yield at muscle level, resulting from sum of V_o₂ through the mouth (curve 2) and O₂‐stores depletion. Lactate production (——) begins when V_o₂ at muscle level has reached its maximum value (areas La, amount of energy supplied by lactic sources). (Time course of La production, as well as time at which glycolysis begins, is hypothetical.) After exercise, VPC_o₂ is paid by the oxidative processes that remain at maximum level for ∼25 s; t_1/2 of curves 1 and 1' = 15 s (asymptote of curve 1 = energy requirement); t_1/2 of curve 2 = 30 s (asymptote = energy requirement); t_1/2 of curve 2' = 25 s.

Adapted from di Prampero 137

Figure 12.

Model for control of muscle via mitochondrial creatinephosphokinase (CPK). Subscripts: e, extramitochondrial; i, intramitochondrial; m, mitochondrial. Glyc., glycogen.

Adapted from Mahler (329,528)

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Paolo Cerretelli, Pietro E. Di Prampero. Gas Exchange in Exercise. Compr Physiol 2011, Supplement 13: Handbook of Physiology, The Respiratory System, Gas Exchange: 297-339. First published in print 1987. doi: 10.1002/cphy.cp030416

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