Comprehensive Physiology Wiley Online Library
For Librarians For Authors Editors About

Pulmonary Circulation at Exercise

Robert Naeije, N. Chesler

10.1002/cphy.c100091

Source: Volume 2, Issue 1, January 2012

Published online: January 2012

Full Article on Wiley Online Library



Abstract

The pulmonary circulation is a high‐flow and low‐pressure circuit, with an average resistance of 1 mmHg/min/L in young adults, increasing to 2.5 mmHg/min/L over four to six decades of life. Pulmonary vascular mechanics at exercise are best described by distensible models. Exercise does not appear to affect the time constant of the pulmonary circulation or the longitudinal distribution of resistances. Very high flows are associated with high capillary pressures, up to a 20 to 25 mmHg threshold associated with interstitial lung edema and altered ventilation/perfusion relationships. Pulmonary artery pressures of 40 to 50 mmHg, which can be achieved at maximal exercise, may correspond to the extreme of tolerable right ventricular afterload. Distension of capillaries that decrease resistance may be of adaptative value during exercise, but this is limited by hypoxemia from altered diffusion/perfusion relationships. Exercise in hypoxia is associated with higher pulmonary vascular pressures and lower maximal cardiac output, with increased likelihood of right ventricular function limitation and altered gas exchange by interstitial lung edema. Pharmacological interventions aimed at the reduction of pulmonary vascular tone have little effect on pulmonary vascular pressure‐flow relationships in normoxia, but may decrease resistance in hypoxia, unloading the right ventricle and thereby improving exercise capacity. Exercise in patients with pulmonary hypertension is associated with sharp increases in pulmonary artery pressure and a right ventricular limitation of aerobic capacity. Exercise stress testing to determine multipoint pulmonary vascular pressures‐flow relationships may uncover early stage pulmonary vascular disease. © 2012 American Physiological Society. Compr Physiol 2:711‐741, 2012.

Comprehensive Physiology offers downloadable PowerPoint presentations of figures for non-profit, educational use, provided the content is not modified and full credit is given to the author and publication.

Download a PowerPoint presentation of all images


Figure 1. Figure 1.

Starling resistor model to explain the concept of closing pressure within a circulatory system. Flow (Q) is determined by the gradient between an inflow pressure, or mean pulmonary artery pressure (Ppa), and an outflow pressure which is either closing pressure (Pc) or left atrial pressure (Pla). When Pla > Pc, the (Ppa–Pla)/Q relationship crosses the origin (A curve) and PVR is constant. When Pc > Pla, the (Ppa–Pla)/Q relationship has a positive pressure intercept (B and C curves), and PVR decreases curvilinearly with increasing Q. The B and C curves are curvilinear a low flow representing recruitment. Also shown are possible misleading PVR calculations: PVR, the slope of (Ppa–Pla)/Q may remain unchanged in the presence of a vasoconstriction (from 1 to 2) or decrease (from 1 to 3) with no change in the functional state of the pulmonary circulation (unchanged pressure/flow line). Adapted from reference 155. Permission pending.
Figure 2. Figure 2.

Mean pulmonary artery pressure (Ppa) as a function of cardiac output (Q) at constant left atrial pressure (Pla), left panel, and Ppa as a function of Pla at constant Q in an anesthetized dog before (stippled line) and after (full line) injection of oleic acid (OA) to produce an acute lung injury. Lung injury was associated with a shift of linear Ppa‐Q relationships to higher pressures, with increased extrapolated pressure intercept (small stipple line). Pla was transmitted to Ppa in a close to 1:1 relationship before oleic acid, but only at a pressure equal to the extrapolated pressure intercept of Ppa‐Q after oleic acid, which is compatible with an increased closing pressure becoming the effective downstream pressure of the pulmonary circulation. Adapted, with permission, from reference 121.
Figure 3. Figure 3.

Pressure‐flow relationships of an isolated perfused mouse lung before (empty sqares) and after (full squares) embolization. The shape of the multipoint pressure‐flow relationship is increasingly curvilinear at decreasing flow. Extrapolated pressure intercept from best adjustments on “physiological” values for pressure and flow leads to spurious estimations of increased closing pressures. Adapted, with permission, from reference 219.
Figure 4. Figure 4.

Pulmonary vascular impedance spectra in a dog, at rest and running. Running was associated with a decrease in 0 Hz impedance, but an increase in the ratio of pressure and flow moduli (in dyne/s/cm5) at all frequencies, and a decrease in low‐frequency phase angle, suggestive of decreased proximal compliance of the pulmonary arterial tree. Drawn, with permission, after reference 48.
Figure 5. Figure 5.

Effects of exercise (shaded columns) on pulmonary vascular resistance (PVR), characteristic impedance (Zc), and arterial compliance (Ca) in healthy human subjects. Exercise decreased PVR and increased Ca, while there was no significant change in Zc. Drawn, with permission, from data in reference 193.
Figure 6. Figure 6.

Effects of increasing levels of exercise in the supine and in the sitting position on mean ± SD (vertical bars) pulmonary vascular resistance (PVR, Wood units or mmHg/L) in healthy volunteers. Initial PVR was higher in the sitting position, but otherwise PVR decreased slightly with increasing levels of workload, and this was similar in the supine and in the sitting positions. Redrawn, with permission, after reference 179.
Figure 7. Figure 7.

Rapid recovery of mean pulmonary artery pressure (mPpa) and cardiac output (Q) after maximal exercise. Values are reported as mean ± SD (vertical bars). *P< 0.05 compared to resting baseline. After 20 min, Q is still higher than resting baseline. Drawn from reference 4, permission pending.
Figure 8. Figure 8.

Linear relationship between mean pulmonary artery pressure (Ppam) and cardiac output (Q) in 25 healthy subjects during progressively severe exercise until maximum tolerated. The slope of Ppam‐Q was 1.37 mmHg/L/min. Adapted from reference 4. Permission pending.
Figure 9. Figure 9.

Mean ± SD (vertical bars) values of mean pulmonary artery wedge pressure (Ppw) and right atrial pressure (Pra) as a function of cardiac output (Q) during progressive exercise in normal volunteers, left panel, and Ppw versus Pra of the same subjects, right panel. Both Ppw and Pra increase with Q, but the gradient between the pressures tended to increase. The right panel shows that Ppw is correlated to Pra, but is higher than Pra by 5 mmHg at rest, and this increases with a slope of 1.49 mmHg increase in Ppw for every mmHg increase of Pra at exercise. Adapted, with permission, from references 180 and 181.
Figure 10. Figure 10.

Mean ± SD (vertical bars) values of mean pulmonary artery pressure (Ppa) and wedged Ppa (Ppw) as a function of cardiac output (Q) during progressive exercise in normal volunteers, left panel, and Ppa versus Prw of the same subjects, right panel. Both Ppa and Ppw increase with Q, but the gradient between the pressures tends to remain unchanged. The left panel shows that Ppa is correlated to Prw, but is higher than Ppw by 9 mmHg at rest, and this increases with a slope of 1.1 mmHg increase in Ppa for every mmHg increase of Ppw at exercise. Adapted, with permission, from references 180 and 181.
Figure 11. Figure 11.

Individual mean ± SE values of pulmonary artery wedge pressure (Ppw) as a function of cardiac output (Q) in subjects with a low (untrained) versus a high exercise capacity (trained). The increase in Ppw was delayed until higher Q in the fittest subjects, suggesting improved ventricular compliance. Adapted, with permission, from reference 201.
Figure 12. Figure 12.

Predicted minus measured mean pulmonary artery pressure (Ppa) as a function of mean Ppa in healthy exercising volunteers. The stippled line shows 2 SD, which is equal to 1.7 mmHg. This presentation is suggestive of a good agreement. The prediction of Ppa was obtained using the distensibility model of Linehan. Adapted, with permission, from reference 182.
Figure 13. Figure 13.

Mean values for pulmonary artery pressure (Ppa) and pulmonary artery wedge pressure (Ppw) as a function of cardiac output (Q) in exercising horses. Exercise in horses is associated with marked increases in pulmonary vascular pressures. Adapted, with permission, from reference 183.
Figure 14. Figure 14.

Relation of pulmonary carbon monoxide diffusion capacity (DLCO) to pulmonary blood flow (Q) at increasing levels of exercise. DLCO increases in proportion of Q, even in Olympic cyclists with very high Q. Adapted, with permission, from references 104 and 183.
Figure 15. Figure 15.

Echocardiographic apical 4 chamber views at rest (left panel) and at exercise (middle and right panels) after the injection of agitated contrast, showing appearance in the left heart chambers after 4 to 5 beats at moderate exercise, and more so at intense exercise. Courtesy of A La Gerche.
Figure 16. Figure 16.

Curvilinear relationships between mean pulmonary artery pressure (PPA) and arterial oxygen saturation (SaO2) in healthy highlanders (empty symbols) and highlanders with chronic mountain sickness (full symbols), left palel, and between mean pulmonary artery pressure (PPA) and altitude of locations (right panel). Extremes are observed in Lhasa and in Leadville. Hypoxia is a major determinant of Ppa, but there is ethnic variability. Adapted from reference 172. Permission pending.
Figure 17. Figure 17.

Mean pulmonary artery pressure (Ppa) as a function of cardiac output (Q) at exercise in healthy subjects at sea level, barometric pressure (Pb) 760 mmHg, and after acclimatization at two levels of simulated altitudes. Baseline Ppa and the slope of Ppa‐Q increase with altitude. Adapted, with permission, from reference 73.
Figure 18. Figure 18.

Averaged mean pulmonary artery pressures (Ppa) versus indexed cardiac output (Q) plots measured invasively (cardiac cath, full lines) or noninvasively (echo‐Doppler, stippled lines) in highlanders exercising at high altitude and in lowlanders exercising at sea leve. From references 1,4,6,11,13,73,173,204,223, with indications of names of first authors in the figure. With the exception of Tibetans in lhasa, highlanders present with higher resting Ppa and increased slopes of Ppa‐Q. Slopes of Ppa‐Q are particularly steep on patients with chronic mountain sickness (CMS). Courtesy of Dante Penaloza.
Figure 19. Figure 19.

Correlation between hypoxia‐induced increase in mean pulmonary artery pressure (Ppam) and decrease maximal oxygen uptake (o2 max) in healthy volunteers in either acute normobaric or more chronic hypobaric hypoxic conditions. Adapted from reference 158. Permission pending.
Figure 20. Figure 20.

Two‐point mean pulmonary artery pressure (Ppa) cardiac output (Q) plots in normal subjects (N), or patients with mitral stenosis (MS), acute respiratory distress syndrome (ARDS), chronic obstructive pulmonary disease (COPD), left ventricular failure (LVF), and pulmonary arterial hypertension (PAH). All Ppa‐Q plots are shifted upward to higher pressure with an increased extrapolated pressure intercept (stippled lines). Left atrial pressure was equal to the extrapolated pressure intercept in LVF. Constructed, with permission, with mean data reported in references 50,83,103,106,160,241.
Figure 21. Figure 21.

Plots of mean pulmonary artery pressure (Ppa) as a function cardiac output (Q) in patients with idiopathic pulmonary arterial pressure, with Q increased at exercise or by an infusion of low‐dose dobutamine. The slope of Ppa‐Q was higher with exercise. Adapted from reference 106. Permission pending.


Figure 1.

Starling resistor model to explain the concept of closing pressure within a circulatory system. Flow (Q) is determined by the gradient between an inflow pressure, or mean pulmonary artery pressure (Ppa), and an outflow pressure which is either closing pressure (Pc) or left atrial pressure (Pla). When Pla > Pc, the (Ppa–Pla)/Q relationship crosses the origin (A curve) and PVR is constant. When Pc > Pla, the (Ppa–Pla)/Q relationship has a positive pressure intercept (B and C curves), and PVR decreases curvilinearly with increasing Q. The B and C curves are curvilinear a low flow representing recruitment. Also shown are possible misleading PVR calculations: PVR, the slope of (Ppa–Pla)/Q may remain unchanged in the presence of a vasoconstriction (from 1 to 2) or decrease (from 1 to 3) with no change in the functional state of the pulmonary circulation (unchanged pressure/flow line). Adapted from reference 155. Permission pending.



Figure 2.

Mean pulmonary artery pressure (Ppa) as a function of cardiac output (Q) at constant left atrial pressure (Pla), left panel, and Ppa as a function of Pla at constant Q in an anesthetized dog before (stippled line) and after (full line) injection of oleic acid (OA) to produce an acute lung injury. Lung injury was associated with a shift of linear Ppa‐Q relationships to higher pressures, with increased extrapolated pressure intercept (small stipple line). Pla was transmitted to Ppa in a close to 1:1 relationship before oleic acid, but only at a pressure equal to the extrapolated pressure intercept of Ppa‐Q after oleic acid, which is compatible with an increased closing pressure becoming the effective downstream pressure of the pulmonary circulation. Adapted, with permission, from reference 121.



Figure 3.

Pressure‐flow relationships of an isolated perfused mouse lung before (empty sqares) and after (full squares) embolization. The shape of the multipoint pressure‐flow relationship is increasingly curvilinear at decreasing flow. Extrapolated pressure intercept from best adjustments on “physiological” values for pressure and flow leads to spurious estimations of increased closing pressures. Adapted, with permission, from reference 219.



Figure 4.

Pulmonary vascular impedance spectra in a dog, at rest and running. Running was associated with a decrease in 0 Hz impedance, but an increase in the ratio of pressure and flow moduli (in dyne/s/cm5) at all frequencies, and a decrease in low‐frequency phase angle, suggestive of decreased proximal compliance of the pulmonary arterial tree. Drawn, with permission, after reference 48.



Figure 5.

Effects of exercise (shaded columns) on pulmonary vascular resistance (PVR), characteristic impedance (Zc), and arterial compliance (Ca) in healthy human subjects. Exercise decreased PVR and increased Ca, while there was no significant change in Zc. Drawn, with permission, from data in reference 193.



Figure 6.

Effects of increasing levels of exercise in the supine and in the sitting position on mean ± SD (vertical bars) pulmonary vascular resistance (PVR, Wood units or mmHg/L) in healthy volunteers. Initial PVR was higher in the sitting position, but otherwise PVR decreased slightly with increasing levels of workload, and this was similar in the supine and in the sitting positions. Redrawn, with permission, after reference 179.



Figure 7.

Rapid recovery of mean pulmonary artery pressure (mPpa) and cardiac output (Q) after maximal exercise. Values are reported as mean ± SD (vertical bars). *P< 0.05 compared to resting baseline. After 20 min, Q is still higher than resting baseline. Drawn from reference 4, permission pending.



Figure 8.

Linear relationship between mean pulmonary artery pressure (Ppam) and cardiac output (Q) in 25 healthy subjects during progressively severe exercise until maximum tolerated. The slope of Ppam‐Q was 1.37 mmHg/L/min. Adapted from reference 4. Permission pending.



Figure 9.

Mean ± SD (vertical bars) values of mean pulmonary artery wedge pressure (Ppw) and right atrial pressure (Pra) as a function of cardiac output (Q) during progressive exercise in normal volunteers, left panel, and Ppw versus Pra of the same subjects, right panel. Both Ppw and Pra increase with Q, but the gradient between the pressures tended to increase. The right panel shows that Ppw is correlated to Pra, but is higher than Pra by 5 mmHg at rest, and this increases with a slope of 1.49 mmHg increase in Ppw for every mmHg increase of Pra at exercise. Adapted, with permission, from references 180 and 181.



Figure 10.

Mean ± SD (vertical bars) values of mean pulmonary artery pressure (Ppa) and wedged Ppa (Ppw) as a function of cardiac output (Q) during progressive exercise in normal volunteers, left panel, and Ppa versus Prw of the same subjects, right panel. Both Ppa and Ppw increase with Q, but the gradient between the pressures tends to remain unchanged. The left panel shows that Ppa is correlated to Prw, but is higher than Ppw by 9 mmHg at rest, and this increases with a slope of 1.1 mmHg increase in Ppa for every mmHg increase of Ppw at exercise. Adapted, with permission, from references 180 and 181.



Figure 11.

Individual mean ± SE values of pulmonary artery wedge pressure (Ppw) as a function of cardiac output (Q) in subjects with a low (untrained) versus a high exercise capacity (trained). The increase in Ppw was delayed until higher Q in the fittest subjects, suggesting improved ventricular compliance. Adapted, with permission, from reference 201.



Figure 12.

Predicted minus measured mean pulmonary artery pressure (Ppa) as a function of mean Ppa in healthy exercising volunteers. The stippled line shows 2 SD, which is equal to 1.7 mmHg. This presentation is suggestive of a good agreement. The prediction of Ppa was obtained using the distensibility model of Linehan. Adapted, with permission, from reference 182.



Figure 13.

Mean values for pulmonary artery pressure (Ppa) and pulmonary artery wedge pressure (Ppw) as a function of cardiac output (Q) in exercising horses. Exercise in horses is associated with marked increases in pulmonary vascular pressures. Adapted, with permission, from reference 183.



Figure 14.

Relation of pulmonary carbon monoxide diffusion capacity (DLCO) to pulmonary blood flow (Q) at increasing levels of exercise. DLCO increases in proportion of Q, even in Olympic cyclists with very high Q. Adapted, with permission, from references 104 and 183.



Figure 15.

Echocardiographic apical 4 chamber views at rest (left panel) and at exercise (middle and right panels) after the injection of agitated contrast, showing appearance in the left heart chambers after 4 to 5 beats at moderate exercise, and more so at intense exercise. Courtesy of A La Gerche.



Figure 16.

Curvilinear relationships between mean pulmonary artery pressure (PPA) and arterial oxygen saturation (SaO2) in healthy highlanders (empty symbols) and highlanders with chronic mountain sickness (full symbols), left palel, and between mean pulmonary artery pressure (PPA) and altitude of locations (right panel). Extremes are observed in Lhasa and in Leadville. Hypoxia is a major determinant of Ppa, but there is ethnic variability. Adapted from reference 172. Permission pending.



Figure 17.

Mean pulmonary artery pressure (Ppa) as a function of cardiac output (Q) at exercise in healthy subjects at sea level, barometric pressure (Pb) 760 mmHg, and after acclimatization at two levels of simulated altitudes. Baseline Ppa and the slope of Ppa‐Q increase with altitude. Adapted, with permission, from reference 73.



Figure 18.

Averaged mean pulmonary artery pressures (Ppa) versus indexed cardiac output (Q) plots measured invasively (cardiac cath, full lines) or noninvasively (echo‐Doppler, stippled lines) in highlanders exercising at high altitude and in lowlanders exercising at sea leve. From references 1,4,6,11,13,73,173,204,223, with indications of names of first authors in the figure. With the exception of Tibetans in lhasa, highlanders present with higher resting Ppa and increased slopes of Ppa‐Q. Slopes of Ppa‐Q are particularly steep on patients with chronic mountain sickness (CMS). Courtesy of Dante Penaloza.



Figure 19.

Correlation between hypoxia‐induced increase in mean pulmonary artery pressure (Ppam) and decrease maximal oxygen uptake (o2 max) in healthy volunteers in either acute normobaric or more chronic hypobaric hypoxic conditions. Adapted from reference 158. Permission pending.



Figure 20.

Two‐point mean pulmonary artery pressure (Ppa) cardiac output (Q) plots in normal subjects (N), or patients with mitral stenosis (MS), acute respiratory distress syndrome (ARDS), chronic obstructive pulmonary disease (COPD), left ventricular failure (LVF), and pulmonary arterial hypertension (PAH). All Ppa‐Q plots are shifted upward to higher pressure with an increased extrapolated pressure intercept (stippled lines). Left atrial pressure was equal to the extrapolated pressure intercept in LVF. Constructed, with permission, with mean data reported in references 50,83,103,106,160,241.



Figure 21.

Plots of mean pulmonary artery pressure (Ppa) as a function cardiac output (Q) in patients with idiopathic pulmonary arterial pressure, with Q increased at exercise or by an infusion of low‐dose dobutamine. The slope of Ppa‐Q was higher with exercise. Adapted from reference 106. Permission pending.

References
 1. Aaron CP, ETandri H, Barr RG, Johnson WC, Bagiella E, Chahal H, Jain A, Kizer JR, Bertoni AG, Lima JA, Bluemke DA, Kawut SM. Physical activity and right ventricular structure and function. The MESA‐right ventricle study. Am J Respir Crit Care Med 183: 396‐404, 2010.
 2. Allemann Y, Rotter M, Hutter D, Lipp E, Sartori C, Scherrer U, Seiler C. Impact of acute hypoxic pulmonary hypertension on LV diastolic function in healthy mountaineers at high altitude. Am J Physiol Heart Circ Physiol 286: H856‐H862, 2004.
 3. Anand I, Malhotra R, Chandrashekhar Y, Bali HK, Chauhan SS, Jindal SK, Bhandari RK, Wahi PL. Adult subacute mountain sickness‐a syndrome of congestive heart failure in man at very high altitude. Lancet 335: 561‐565, 1990.
 4. Argiento P, Chesler N, Mulè M, D'Alto M, Bossone E, Unger P, Naeije R. Exercise stress echocardiography for the study of the pulmonary circulation. Eur Respir J 35: 1273‐1278, 2010.
 5. Astrand PO, Cuddy TE, Saltin B, Tenberg J. Cardiac output during submaximal and maximal work. J Appl Physiol 19: 268‐274, 1964.
 6. Banchero N, Sime F, Penaloza D, Cruz J, Gamboa R, Marticorena E. Pulmonary pressure, cardiac output, and arterial oxygen saturation during exercise at high altitude and at sea level. Circulation 33: 249‐262, 1966.
 7. Barer GR, Howard P, Shaw JW. Stimulus‐response curves for the pulmonary vascular bed to hypoxia and hypercapnia. J Physiol 21: 139‐155, 1970.
 8. Bärtsch P, Mairbäurl H, Maggiorini M, Swenson E. Physiologic aspects of high‐altitude pulmonary edema. J Appl Physiol 98: 1101‐1110, 2005.
 9. Bernard SL, Glenny RW, Erickson HH, Fedde MR, Polissar N, Basaraba RJ, Hlastala MP. Minimal redistribution of blood flow with exercise in race horses. J Appl Physiol 81: 1062‐1070, 1996.
 10. Bevegaard S, Holmgren A, Jonsson B. The effect of body position on the circulation at rest and during exercise, with special reference to the influence of stroke volume. Acta Physiol Scand 49: 279‐298, 1960.
 11. Bevegaard S, Holmgren A, Jonsson B. Circulatory studies in well trained athletes at rest and during heavy exercise, with special reference to stroke volume and the influence of body position. Acta Physiol Scand 57: 26‐50, 1963.
 12. Bonderman D, Martischnig AM, Vonbank K, Nikfardjam M, Meyer B, Heinz G, Naeije R, Lang IM. Reduced exercise capacity after successful pulmonary endarterectomy is due to decreased pulmonary arterial compliance. Chest 139: 122‐127, 2011
 13. Bossone E, Rubenfire M, Bach DS, Ricciardi M, Armstrong WF. Range of tricuspid regurgitation velocity at rest and during exercise in normal adult males: Implications for the diagnosis of pulmonary hypertension. J Am Coll Cardiol 33: 1662‐1666, 1999.
 14. Boussuges A, Molenat F, Burnet H, Cauchy E, Gardette B, Sainty JM, Jammes Y, Richalet JP. Operation Everest III (Comex ‘97): Modifications of cardiac function secondary to altitude‐induced hypoxia. Am J Respir Crit Care Med 161: 264‐270, 2000.
 15. Brimioulle S, Lejeune P, Vachiéry JL, Delcroix M, Hallemans R, Leeman M, Naeije R. The stimulus‐response curve of hypoxic pulmonary vasoconstriction in intact dogs: Effects of ASA. J Appl Physiol 77: 476‐480, 1994.
 16. Brimioulle S, Maggiorini M, Stephanazzi J, Vermeulen F, Lejeune P, Naeije R. Effects of low flow on pulmonary vascular pressure‐flow curves and pulmonary vascular impedance. Cardiovasc Res 42: 183‐192, 1999.
 17. Brimioulle S, Wauthy P, Ewalenko P, Rondelet B, Vermeulen F, Kerbaul F, Naeije R. Single‐beat estimation of right ventricular end‐systolic pressure‐volume relationship. Am J Physiol Heart Circ Physiol 284: H1625‐H1630, 2003.
 18. Brofman BL, Charms BL, Kohn PM, Elder J, Newman B, Rizika M. Unilateral pulmonary occlusion in man. Control studies. J Thoracic Surg 34: 206‐227, 1957.
 19. Bryan AC, Bentibaglio LG, Beerel F, MacLeish H, Zidulka A, Bates DV. Factors affecting regional distribution of ventilation and perfusion in the lung. J Appl Physiol 19: 395‐402, 1964.
 20. Bshouty Z, Younes M. Arterial occlusion versus isofiltration pulmonary capillary pressures during very high flow. J Appl Physiol 62: 1174‐1178, 1987.
 21. Bshouty Z, Younes M. Distensibility and pressure‐flow relationship of the pulmonary circulation. II. Multibranched model. J Appl Physiol 68: 1514‐1527, 1990.
 22. Burnham KJ, Arai TJ, Dubowitz DJ, Henderson AC, Holverda S, Buxton RB, Prisk GK, Hopkins SR. Pulmonary perfusion heterogeneity in increased in sustained heavy exercise in humans. J Appl Physiol 107: 1559‐1568, 2009.
 23. Calbet JA, Boushel R, Radegran G, Sondergaard H, Wagner PD, Saltin B. Why is VO2 max after altitude acclimatization still reduced despite normalization of arterial O2 content? Am J Physiol Regul Integr Comp Physiol 284: R304‐R316, 2003.
 24. Castelain V, Chemla D, Humbert M, Sitbon O, Simonneau G, Lecarpentier Y, Hervé P. Pulmonary artery pressure‐flow relations after prostacyclin in primary pulmonary hypertension. Am J Respir Crit Care Med 165: 338‐340, 2002.
 25. Chammas JH, Rickaby DA, Guarin M, Linehan JH, Hanger CC, Dawson CA. Flow‐induced vasodilation in the ferret lung. J Appl Physiol 83: 495‐502, 1997
 26. Champion HC, Michelakis ED, Hassoun PM. Comprehensive invasive and noninvasive approach to the right ventricle‐pulmonary circulation unit: State of the art and clinical and research implications. Circulation 120: 992‐1007, 2009.
 27. Chemla D, Castelain V, Humbert M, Hébert JL, Simonneau G, Lecarpentier Y, Hervé P. New formula for predicting mean pulmonary artery pressure using systolic pulmonary artery pressure. Chest 126: 1313‐1317, 2004.
 28. Christie J, Sheldahl LM, Tristani FE, Sagar KB, Ptacin MJ, Wann S. Determination of stroke volume and cardiac output during exercise: Comparison of two‐dimensional and Doppler echocardiography, Fick oximetry, and thermodilution. Circulation 76: 539‐547, 1987.
 29. Cope DK, Grimbert F, Downey JM, Taylor AE. Pulmonary capillary pressure: A review. Crit Care Med 20: 1043‐1056, 1992.
 30. Cournand A, Riley RL, Himmelstein A, Austrial R. Pulmonary circulation and alveolar ventilation‐perfusion relationships after pneumonectomy. J Thoracic Surg 19: 80‐116, 1950.
 31. D'Alto M, Ghio S, D'Andrea A, Pazzano AS, Argiento P, Camporotondo R, Allocca F, Scelsi L, Cuomo G, Caporali R, Cavagna L, Valentini G, Calabrò R. Inappropriate exercise‐induced increase inpulmonary artery pressure in patients with systemic sclerosis. Heart 97: 112‐117, 2011
 32. de Beer VJ, de Graaff HJD, Hoekstra M, Duncker DJ, Merkus D. Integrated control of pulmonary vascular tone by endothelin and angiotensin II in exercising swine depends on gender. Am J Physiol Heart Circ Physiol 298: H1976‐H1985, 2010.
 33. Delcroix M, Mélot C, Vachiery JL, Lejeune P, Leeman M, Vanderhoeft P, Naeije R. Effects of embolic size on hemodynamics and gas exchange in canine embolic pulmonary hypertension. J Appl Physiol 69: 2254‐2261, 1990.
 34. Dempsey JA. Is the lung build for exercise? Med Sci Sports Exerc 18: 143‐155, 1986.
 35. Dempsey JA, Hanson P, Henderson K. Exercise‐induced alveolar hypoxemia in healthy human subjects at sea level. J Physiol (Lond) 355: 161‐175, 1984.
 36. Dempsey JA, Wagner PD. Exercise‐induced arterial hypoxemiua. J Appl Physiol 87: 1997‐2006, 1999.
 37. Dexter L, Whittenberger JL, Haynes FW, Goodale WT, Gorlin R, Sawyer CG. Effects of exercise on circulatory dynamics of normal individuals. J Appl Physiol 3: 439‐453, 1951.
 38. Dorrington KL, Clar C, Young JD, Jonas M, Tansley JG, Robbins PA. Time course of the human pulmonary vascular response to 8 hours of isocapnic hypoxia. Am J Physiol 273: H1126‐1134, 1997.
 39. Douglas PS, O'Toole ML, Hiller WD, Reichek N. Different effects of prolonged exercise on the right and the left ventricles. J Am Coll Cardiol 15: 64‐69, 1990.
 40. Dripps RD, Comroe JH Jr. The effect of the inhalation of high and low oxygen concentrations on respiration, pulse rate, ballistocardiogram and arterial oxygen saturation (oximeter) of normal individuals. Am J Physiol 149: 277‐283, 1947.
 41. Dujardin JP, Stone DN, Forcino CD, Paul LT, Pieper HP. Effects of blood volume changes on characteristic impedance of the pulmonary artery. Am J Physiol 242: H197‐H202, 1982.
 42. Duncker DJ, Stubenitsky R, Tonino PA, Verdouw PD. Nitric oxide contributes to the regulation of vasomotor tone but does not modulate O2 consumption in exercising swine. Cardiovasc Res 47: 738‐748, 2000.
 43. Ekelund LG, Holmgren A. Circulatory and respiratory adaptation during long‐term, non‐steady state exercise in the sitting position. Acta Physiol Scand 62: 240‐255, 1964.
 44. Ekelund LG, Holmgren A, Ovenfors CO. Heart volume during prolonged exercise in the supine and sitting position. Acta Physiol Scand 70: 88‐98, 1967.
 45. Eldridge MW, Braun RK, Yoneda KY, Walby WF. Effects of altitude and exercise on pulmonary capillary integrity: Evidence for subclinical high‐altitude pulmonary edema. J Appl Physiol 100: 972‐980, 2006.
 46. Eldridge MW, Dempsey JA, Haverkamp HC, Lovering AT, Hokanson JS. Exercise‐induced intrapulmonary arteriovenous shunting in healthy humans. J Appl Physiol 97: 797‐805, 2004.
 47. Eldridge MW, Podolsky A, Richardson RS, Johnson DH, Knight DR, Johnson EC, Hopkins SR, Michimata H, Grassi B, Feiner J, Kurdak SS, Bickler PE, Wagner PD, Severinghaus JW. Pulmonary hemodynamic response to exercise in subjects with prior high‐altitude pulmonary edema. J Appl Physiol 81: 911‐921, 1996.
 48. Elkins RC, Milnor WR. Pulmonary vascular response to exercise in the dog. Circ Res 29: 591‐599, 1971.
 49. Elzinga G, Westerhof N. The effect of an increase in inotropic state and end‐diastolic volume on the pumping ability of the feline left heart. Circ Res 42: 620‐628, 1978.
 50. Even P, Duroux P, Ruff F, Caubarrere I, de Vernejoul P, Brouet G. The pressure/flow relationship of the pulmonary circulation in normal man and in chronic obstructive pulmonary diseases. Effects of muscular exercise. Scand J Respir Dis 77: 72‐76, 1971.
 51. Faoro V, Boldingh S, Moreels M, Martinez S, Lamotte M, Unger P, Brimioulle S, Huez S, Naeije R. Bosentan decreases pulmonary vascular resistance and improves exercise capacity in acute hypoxia. Chest 135: 1215‐1222, 2009.
 52. Faoro V, Huez S, Giltaire S, Pavelescu A, van Osta A, Moraine JJ, Guenard H, Martinot JB, Naeije R. Effects of acetazolamide on aerobic exercise capacity and pulmonary hemodynamics at high altitudes. J Appl Physiol 103: 1161‐1165, 2007.
 53. Faoro V, Lamotte M, Deboeck G, Pavelescu A, Huez S, Guenard H, Martinot JB, Naeije R. Effects of sildenafil on exercise capacity in hypoxic normal subjects. High Alt Med Biol 8: 155‐163, 2007.
 54. Farber HW, Loscalzo J. Pulmonary arterial hypertension. N Engl J Med 351: 1655‐1665, 2004.
 55. Fesler P, Pagnamenta A, Rondelet B, Kerbaul F, Naeije R. Effects of sildenafil on hypoxic pulmonary vascular function in dogs. J Appl Physiol 101: 1085‐1090, 2006.
 56. Fischler M, Maggiorini M, Dorschner L, Debrunner J, Bernheim A, Kiencke S, Mairbäurl H, Bloch KE, Naeije R, Brunner‐La Rocca HP. Dexamethasone but not tadalafil improves exercise capacity in adults prone to high altitude pulmonary edema. Am J Respir Crit Care Med 180: 346‐352, 2009.
 57. Fishman AP, Mc Clement J, Himmelstein A, Cournand A. Effects of acute anoxia on the circulation and respiration in patients with chronic obstructive pulmonary disease studied during the “steady state”. J Clin Invest 31: 770‐781, 1952.
 58. Fowler NO. The normal pulmonary arterial pressure‐flow relationships during exercise. Am J Med 47: 1‐6, 1969.
 59. Fowler NO, Holmes JC. Pulmonary arterial pressure at high pulmonary flow. J Clin Invest 44: 2040‐2050, 1965.
 60. Fulco CS, Rock PB, Cymerman A. Maximal and submaximal exercise performance at altitude. Aviat Space Environ Med 69: 793‐801, 1998.
 61. Fung YC, Sobin SS. Pulmonary alveolar blood flow. Circ Res 30: 470‐490, 1972.
 62. Gaar KA Jr, Taylor AE, Owens LJ, Guyton AC. Pulmonary capillary pressure and filtration coefficient in the isolated perfused lung. Am J Physiol 213: 910‐914, 1967.
 63. Gale GE, Torre‐Bueno JR, Moon RE, Saltzman HA, Wagner PD. Ventilation‐perfusion inequality in normal humans during exercise at sea level and simulated altitude. J Appl Physiol 58: 978‐988, 1985.
 64. Geary GG, Buchholz JN. Effects of aging on cerebrovascular tone and [Ca2+]i. J Appl Physiol 95: 1746‐1754, 2003
 65. Ghofrani HA, Reichenberger F, Kohstall MG, Mrosek EH, Seeger T, Olschewski H, Seeger W, Grimminger F. Sildenafil increased exercise capacity during hypoxia at low altitudes and at mount Everest base camp: A randomized, double‐blind, placebo‐controlled crossover trial. Ann Intern Med 141: 169‐177, 2004.
 66. Gledhill N, Cox D, Jamnik R. Endurance athletes stroke volume does not plateau: Major advantage is diastolic function. Med Sci Sports Exerc 26: 1116‐1124, 1994.
 67. Gledhill N, Froese AB, Buick FJ, Bryan AC. VA/Q inhomogeneity and AaDO2 in man during exercise: Effect of SF6 breathing. J Appl Physiol 45: 512‐515, 1978.
 68. Granath A, Jonsson B, Strandell T. Circulation in healthy old men, studied by right heart catheterization at rest and during exercise in supine and sitting position. Acta Med Scand 176: 425‐446, 1964.
 69. Granath A, Strandell T. Relationships between cardiac output, stroke volume, and intracardiac pressures at rest and during exercise in supine position and some anthropometric data in healthy old men. Acta Med Scand 176: 447‐466, 1964.
 70. Grant BJ, Davies EE, Jones HA, Hughes JM. Local regulation of pulmonary blood flow and ventilation‐perfusion ratios in the coatimundi. J Appl Physiol 40: 216‐228, 1976.
 71. Grover RF, Wagner WW, McMurtry IF, Reeves JT. Pulmonary Circulation. In: Shepherd JT, Abboud FM, editors. Handbook of Physiology. The Cardiovascular System. Peripheral Circulation and Organ Blood Flow. Bethesda, MD: Am Physiol Soc sect 2, Vol. III, part 1, Chap 4, pp. 103‐36, 1983.
 72. Groves BM, Droma T, Sutton JR, McCullough RG, McCullough RE, Zhuang J, Rapmund G, Sun S, Janes C, Moore LG. Minimal hypoxic pulmonary hypertension in normal Tibetans at 3,658 m. J Appl Physiol 74: 312‐318, 1993.
 73. Groves BM, Reeves JT, Sutton JR, Wagner PD, Cymerman A, Malconian MK, Rock PB, Young PM, Houston CS. Operation Everest II: Elevated high altitude pulmonary resistance unresponsive to oxygen. J Appl Physiol 63: 521‐530, 1987.
 74. Grünig E, Mereles D, Hildebrandt W, Swenson ER, Kübler W, Kuecherer H, Bärtsch P. Stress Doppler echocardiography for identification of susceptibility to high altitude pulmonary edema. J Am Coll Cardiol 35: 980‐987, 2000.
 75. Grünig E, Weissmann S, Ehlken N, Fijalkowska A, Fischer C, Fourme T, Galié N, Ghofrani A, Harrison RE, Huez S, Humbert M, Janssen B, Kober J, Koehler R, Machado RD, Mereles D, Naeije R, Olschewski H, Provencher S, Reichenberger F, Retailleau K, Rocchi G, Simonneau G, Torbicki A, Trembath R, Seeger W. Stress‐Doppler‐echocardiography in relatives of patients with idiopathic and familial pulmonary arterial hypertension: Results of a multicenter European analysis of pulmonary artery pressure response to exercise and hypoxia. Circulation 119; 1747‐57, 2009.
 76. Guazzi M, Tumminello G, Di Marco F, Fiorentini C, Guazzi M. D. The Effects of Phosphodiesterase‐5 Inhibition with sildenafil on pulmonary hemodynamics and diffusion capacity, exercise ventilatory efficiency, and oxygen uptake kinetics in chronic heart failure. J Am Coll Cardiol 44: 2339‐48, 2004.
 77. Gurtner HP, Walser P, Fassler B. Normal values for pulmonary hemodynamics at rest and during exercise in man. Progr Resp Res 9: 295‐315, 1975.
 78. Guyton AC, Lindsay AE. Effects of elevated left atrial pressures and decreased plasma protein in the development of lung edema. Circ Res 7: 649‐655, 1959.
 79. Halpern SD, Taichman DB. Misclassification of pulmonary hypertension due to reliance on pulmonary capillary wedge pressure rather than left ventricular end‐diastolic pressure. Chest 136: 37‐43, 2009
 80. Hammond HK, White FC, Bhargava V, Shabetai R. Heart size and maximal cardiac output are limited by the pericardium. Am J Physiol Heart Circ Physiol 263: H1675‐H1681, 1992.
 81. Hammond MD, Gale GE, Kapitan KS, Reis A, Wagner PD. Pulmonary gas exchange in humans during exercise at sea level. J Appl Physiol 60: 1590‐1598, 1986.
 82. Harf A, Pratt T, Hughes JMB. Regional distribution of VA/Q in man at rest and with exercise measured with krypton‐81m. J Appl Physiol 44: 115‐123, 1978.
 83. Harris P, Segel N, Bishop JM. The relation between pressure and flow in the pulmonary circulation of normal subjects and in patients with chronic bronchitis and mitral stenosis. Cardiovasc Res 2: 73‐84, 1968.
 84. Hasinoff L, Ducas J, Schick U, Prewitt RM. Pulmonary vascular pressure‐flow characteristics in canine pulmonary embolism. J Appl Physiol 68: 462‐467, 1990.
 85. Hecht HH, Kuida H, Lange RL, Thorne JL, Brown AM. Brisket disease. II. Clinical features and hemodynamic observations in altitude‐dependent right heart failure of cattle. Am J Med 32: 171‐183, 1962.
 86. Heidbuchel H, Hoogsteen J, Fagard R, Vanhees L, Ector H, Willems R, Van Lierde J. High prevalence of right ventricular involvement in endurance athletes with ventricular arrythmias. Role of an electrophysiological study in risk stratification. Eur Heart J 24: 1473‐1480, 2003.
 87. Hickman JB, Cargill WH. Effect of exercise on cardiac output and pulmonary artery pressure in normal persons and in patients with cardiovascular disease and pulmonary emphysema. J Clin Invest 27: 10‐23, 1948.
 88. Holmgren A, Jonsson B, Sjostrand T. Circulatory data in normal subjects at rest and during exercise in the recumbent position, with special reference to the stroke volume at different working intensities. Acta Physiol Scand 49: 343‐363, 1960.
 89. Hopkins SR. Last word on PointCounterpoint: Pulmonary edema does/does not occur in human athletes performing sea level exercise. J Appl Physiol 109: 1281, 2010.
 90. Hopkins SR, Bayly WM, Slocombe RF, Wagner H, Wagner PD. Effect of prolonged heavy exercise on exercise gas exchange in horses. J Appl Physiol 84: 1723‐1730, 1998.
 91. Hopkins SR, Garg J, Bolar DS, Balouch J, Levin DL. Pulmonary blood flow heterogeneity during hypoxia and high altitude pulmonary edema. Am J Respir Crit Care Med 171: 83‐87, 2005.
 92. Hopkins SR, Kleinsasser A, Bernard S, Loeckinger A, Faber E, Neradilek B, Polissar NL, Hlastala MP. Hypoxia has a greater effect than exercise on the distribution of pulmonary blood flow in swine. J Appl Physiol 103: 2112‐1254, 2007.
 93. Hopkins SR, Levin DL, Emami K, Kadlecek S, Yu J, Ishii M, Rizi RR. Advances in magnetic resonance imaging on lung physiology. J Appl Physiol 102: 1244‐1254, 2007.
 94. Hopkins SR, Olfert IM, Wagner PD. Point: Exercise‐induced intrapulmonary shunting is imaginary. J Appl Physiol 107: 993‐994, 2009.
 95. Hopkins SR, Stary CM, Falor E, Wagner H, Wagner PD, McKirnan MD. Pulmonary gas exchange during exercise in pigs. J Appl Physiol 86: 93‐100, 1999.
 96. Hsu AR, Barnholt KE, Grundmann NK, Lin JH, McCullum SW, Friedlander AL. Sildenafil improves cardiac output and exercise performance during acute hypoxia, but not normoxia. J Appl Physiol 100: 2031‐2040, 2006.
 97. Huez S, Faoro V, Guénard H, Martinot JB, Naeije R. Echocardiographic and tissue Doppler imaging of cardiac adaptation to high altitude in native highlanders versus acclimatized lowlanders. Am J Cardiol 103: 1605‐1609, 2009.
 98. Huez S, Faoro V, Vachiery JL, Unger P, Martinot JB, Naeije R. Images in cardiovascular medicine. High‐altitude‐induced right‐heart failure. Circulation 115: e308‐e309, 2007.
 99. Huez S, Retailleau K, Unger P, Pavelescu A, Vachiery JL, Derumeaux G, Naeije R. Right and left ventricular adaptation to hypoxia: A tissue Doppler imaging study. Am J Physiol Heart Circ Physiol 289: H1391‐H1398, 2005.
 100. Huez S, Roufosse F, Vachièry JL, Pavelescu A, Derumeaux G, Wautrecht JC, Cogan E, Naeije R. Isolated right ventricular dysfunction in systemic sclerosis: Latent pulmonary hypertension? Eur Respir J 30: 928‐936, 2007.
 101. Hultgren HN, Grover RF, Hartly LH. Abnormal circulatory responses to high altitude in subjects with a previous history of high‐altitude pulmonary edema. Circulation 44: 759‐770
 102. Janicki JS. Influence of the pericardium and ventricular interdependence on left ventricular diastolic and systolic function in patients with heart failure. Circulation 81: III15‐III20, 1990.
 103. Janicki JS, Weber KT, Likoff MJ, Fishman AP. The pressure‐flow response of the pulmonary circulation in patients with heart failure and pulmonary vascular disease. Circulation 72: 1270‐1278, 1985.
 104. Johnson RL, Spicer WS, Bishop JM, Forster RE. Pulmonary capillary blood volume, flow and diffusing capacity during exercise. J Appl Physiol 15: 893‐902, 1960.
 105. Julien M, Hakim TS, Vahi R, Chang HK. Effect of hematocrit on vascular profile in dog lungs. J Appl Physiol 58: 743‐748, 1985.
 106. Kafi SA, Mélot C, Vachiéry JL, Brimioulle S, Naeije R. Partitioning of pulmonary vascular resistance in primary pulmonary hypertension. J Am Coll Cardiol 31: 1372‐1376, 1998.
 107. Kane DW, Tesauro T, Newman JH. Adrenergic modulation of the pulmonary circulation during strenuous exercise in the sheep. Am Rev Respir Dis 147: 1233‐1238, 1993.
 108. Kawashima A, Kubo K, Kobayashi T, Sekiguchi M. Hemodynamic response to acute hypoxia, hypobaria, and exercise in subjects susceptible to high‐altitude pulmonary edema. J Appl Physiol 67: 1982‐1989, 1989.
 109. Kobs RW, Muvarak NE, Eickhoff JC, Chesler NC. Linked mechanical and biological aspects of remodelling in mouse pulmonary arteries with hypoxia‐induced pulmonary hypertension. Am J Physiol Heart Circ Physiol 288; H1209‐H2017, 2005.
 110. Koizumi T, Gupta R, Banerjee M, Newman JH. Changes in pulmonary vascular tone during exercise. Effects of nitric oxide (NO) synthase inhibition, L‐arginine infusion and NO inhalation. J Clin Invest 94: 2285‐2282, 1994.
 111. Kovacs G, Berghold A, Scheid S, Olschewski H. Pulmonary artery pressure during rest and exercise in healthy subjects: A systematic review. Eur Respir J 34: 888‐894, 2009.
 112. Kovacs G, Maier R, Aberer E, Brodmann M, Scheidl S, Tröster N, Hesse C, Salmhofer W, Graninger W, Gruenig E, Rubin LJ, Olschewski H. Borderline pulmonary arterial pressure is associated with decreased exercise capacity in scleroderma. Am J Respir Crit Care Med 180: 881‐886, 2009.
 113. Krenz GS, Dawson CA. Flow and pressure distributions in vascular networks consisting of distensible vessels. Am J Physiol Heart Circ Physiol 284: H2192‐H2203, 2003.
 114. Kuehne T, Yilmaz S, Steendijk P, Moore P, Groenink M, Saaed M, Weber O, Higgins CB, Ewert P, Fleck E, Nagel E, Schulze‐Neick I, Lange P. Magnetic resonance imaging analysis of right ventricular pressure‐volume loops: In vivo validation and clinical application in patients with pulmonary hypertension. Circulation. 110: 2010‐2016, 2004.
 115. La Gerche A, Heidbuchel H, Burns AT, Mooney DJ, Taylor AJ, Pfluger HB, Inder WJ, Macisaac AI, Prior DL. Disproportionate exercise load and remodeling of the athlete's right ventricle. Med Sci Sports Exerc 43: 974‐981, 2010.
 116. La Gerche A, MacIsaac AL, Burns AT, Mooney DJ, Inder WJ, Voigt JU, Heidbüchel H, Prior DL. Pulmonary transit of agitated contrast is associated with enhanced pulmonary vascular reserve and right ventricular function at exercise. J Appl Physiol 109: 1307‐1317, 2010.
 117. La Gerche A, Robberecht C, Kuiperi C, Nuyens D, Willems R, de Ravel T, Matthijs G, Heidbüchel H. Lower than expected desmosomal gene mutation prevalence in endurance athletes with complex ventricular arrhythmias of right ventricular origin. Heart 96: 1268‐1274, 2010.
 118. Lankhaar JW, Westerhof N, Faes TJ, Gan CT, Marques KM, Boonstra A, van den Berg FG, Postmus PE, Vonk‐Noordegraaf A. Pulmonary vascular resistance and compliance stay inversely related during treatment of pulmonary hypertension. Eur Heart J 29: 1688‐1695, 2008.
 119. Lankhaar JW, Westerhof N, Faes TJ, Marques KM, Marcus JT, Postmus PE, Vonk‐Noordegraaf A. Quantification of right ventricular afterload in patients with and without pulmonary hypertension. Am J Physiol Heart Circ Physiol 291: H1731‐H1737, 2006.
 120. Lategola MT. Pressure‐flow relationships in the dog lung during acute subtotal vascular occlusion. Am J Physiol 192: 613‐619, 1958.
 121. Leeman M, Lejeune P, Closset J, Vachiéry JL, Mélot C, Naeije R. Nature of pulmonary hypertension in canine oleic acid pulmonary edema. J Appl Physiol 69: 293‐298, 1990.
 122. Lejeune P, Vachiéry JL, Leeman M, Brimioulle S, Hallemans R, Mélot C, Naeije R. Absence of parasympathetic control of pulmonary vascular pressure/flow plots in hyperoxic and hypoxic dogs. Respir Physiol 78: 123‐133, 1989.
 123. Lenfant C, Sullivan KN. Adaptation to high altitude. N Engl J Med 284: 1298‐309, 1971.
 124. León‐Velarde F, Maggiorini M, Reeves JT, Aldashev A, Asmus I, Bernardi L, Ge RL, Hackett P, Kobayashi T, Moore LG, Penaloza D, Richalet JP, Roach R, Wu T, Vargas E, Zubieta‐Castillo G, Zubieta‐Calleja G. Consensus statement on chronic and subacute high altitude diseases. High Alt Med Biol 6: 147‐157, 2005.
 125. Leung H, Wang JJ, Rochtchina E, Tan AG, Wong TY, Klein R, Hubbard LD, Mitchell P. Relationships between age, blood pressure, and retinal vessel diameters in an older population. Invest Ophthalmol Vis Sci 44: 2900‐2904, 2003
 126. Levine BD, Lane LD, Buckey JC, Friedman BD, Blomqvist CG. Left ventricular pressure‐volume and Frank‐Starling relation in endurance athletes. Implications for orthostatic tolerance and exercise performance. Circulation 84: 1016‐1023, 1991.
 127. Lindenfeld J, Reeves JT, Horwitz LD. Low exercise pulmonary resistance is not dependent on vasodilator prostaglandins. J Appl Physiol 55: 558‐561, 1983.
 128. Linehan JH, Haworth ST, Nelin LD, Krenz GS, Dawson CA. A simple distensible model for interpreting pulmonary vascular pressure‐flow curves. J Appl Physiol 73: 987‐994, 1992.
 129. Lovering AT, Eldridge MW, Stickland MK. Counterpoint: Exercise‐induced intrapulmonary shunting is real. J Appl Physiol 107: 994‐998, 2009.
 130. Lovering AT, Stickland MK. Not hearing is believing: Novel insight into cardiopulmonary function using agitated contrast and ultrasound. J Appl Physiol 109: 1290‐1291, 2010.
 131. MacDougall JD, McKelvie RS, Moroz DE, Sale DG, McCartney N, Buick F. Factors affecting blood pressure during heavy weight lifting and static contractions. J Appl Physiol 73: 1590‐1597, 1992.
 132. Maggiorini M, Brimioulle S, De Canniere D, Delcroix M, Wauthy P, Naeije R. Pulmonary vascular impedance responses to hypoxia in dogs and in minipigs : Effects of inhaled nitric oxide. J Appl Physiol 79: 1156‐1162, 1995.
 133. Maggiorini M, Mélot C, Gilbert E, Vermeulen F, Naeije R. Nature of pulmonary vascular resistance in dogs and minipigs. Respir Physiol 111: 213‐222, 1998.
 134. Maggiorini M, Mélot C, Pierre S, Pfeiffer F, Greve I, Sartori C, Lepori M, Hauser M, Scherrer U, Naeije R. High altitude pulmonary edema is initially caused by an increased capillary pressure. Circulation 103: 2078‐2083, 2001.
 135. Maignan M, Privat C, Leon‐Velarde F, Richalet JP, Pham I. Pulmonary pressure and cardiac function in chronic mountain sickness patients. Chest 135: 499‐504, 2008.
 136. Manohar M, Goetz TE. L‐NAME does not affect exercise‐induced pulmonary hypertension in thoroughbred horses. J Appl Physiol 84: 1902‐1908, 1998.
 137. Manohar M, Hutchens E, Corney E. Frusemide attenuates the exercise‐induced rise in pulmonary capillary blood pressure in horses. Equine Vet J 26: 51‐54, 1994.
 138. Marcus JT, Gan CT, Zwanenburg JJ, Boonstra A, Allaart CP, Götte MJ, Vonk‐Noordegraaf A. Interventricular mechanical asynchrony in pulmonary arterial hypertension: Left‐to‐right delay in peak shortening is related to right ventricular overload and left ventricular underfilling. J Am Coll Cardiol 51: 750‐757, 2008.
 139. Marshall BE, Marshall C. Site and sensitivity for stimulation of hypoxic pulmonary vasoconstriction. J Appl Physiol 55: 711‐716, 1983.
 140. Marshall BE, Marshall C. A model for hypoxic constriction of the pulmonary circulation. J Appl Physiol 64: 68‐77, 1988.
 141. Marshall BE, Marshall C, Benumof J, Saidman LJ. Hypoxic pulmonary vasoconstriction in dogs: Effects of lung segment size and oxygen tension. J Appl Physiol 51: 1543‐1551, 1981.
 142. McGregor M, Sniderman A. On pulmonary vascular resistance: The need for more precise definition. Am J Cardiol 55: 217‐221, 1985.
 143. McLaughlin VV, Archer SL, Badesch DB, Barst RJ, Farber HW, Lindner JR, Mathier MA, McGoon MD, Park MH, Rosenson RS, Rubin LJ, Tapson VF, Varga J, Harrington RA, Anderson JL, Bates ER, Bridges CR, Eisenberg MJ, Ferrari VA, Grines CL, Hlatky MA, Jacobs AK, Kaul S, Lichtenberg RC, Lindner JR, Moliterno DJ, Mukherjee D, Pohost GM, Rosenson RS, Schofield RS, Shubrooks SJ, Stein JH, Tracy CM, Weitz HH, Wesley DJ. ACCF/AHA. ACCF/AHA 2009 Expert Consensus Document on Pulmonary Hypertension. A Report of the American College of Cardiology Foundation Task Force on Expert Consensus Documents and the American Heart Association. Circulation 119: 2250‐2294, 2009.
 144. Mélot C, Delcroix M, Lejeune P, Leeman M, Naeije R. Starling resistor versus viscoelastic models for embolic pulmonary hypertension. Am J Physiol 267: H817‐H827, 1995.
 145. Mélot C, Naeije R. Pulmonary vascular diseases. Compr Physiol 1: 593‐619, 2011.
 146. Mélot C, Naeije R, Hallemans R, Lejeune P, Mols P. Hypoxic pulmonary vasoconstriction and pulmonary gas exchange in normal man. Respir Physiol 68: 11‐27, 1987.
 147. Merkus D, de Beer VJ, Houweling B, Duncker DJ. Control of pulmonary vascular tone during exercise in health and pulmonary hypertension. Pharmacol Ther 119: 242‐263, 2008.
 148. Merkus D, Houweling B, Mirza A, Boomsma F, van den Meiracker AH, Duncker DJ. Contribution of endotheline and its receptors to the regulation of vascular tone during exercise is different in the systemic, coronary and pulmonary circulation. Cardiovasc Res 59: 745‐754, 2003.
 149. Merkus D, Houweling B, Zarbanoui A, Duncker DJ. Interaction between prostanoids and nitric oxide in regulation of systemic, pulmonary and coronary vascular tone in exercising swine. Am J Physiol 286: H1114‐H1123, 2004.
 150. Milnor WR, Bergel DH, Bargainer JD. Hydraulic power associated with pulmonary blood flow and its relation to heart rate. Circ Res 19: 467‐480, 1966.
 151. Mitzner W, HK Chang. Hemodynamics of the pulmonary circulation. In: Chang HK, Paiva M, editors. Respiratory Physiology: An Analytical Approach. New York, Dekker, p. 561‐631, 1989.
 152. Motley HL, Cournand A, Werkö L, Himmelstein A, Dresdale D. The influence of short periods of induced acute anoxia upon pulmonary artery pressures in man. Am J Physiol 150: 315‐320, 1947.
 153. Moussavi N, Czarnecki A, Kumar K, Fallah‐Rad N, Lytwyn M, Han SY, Francis A, Walker JR, Kirkpatrick ID, Neilan TG, Sharma S, Jassal DS. Relation of biomarkers and cardiac magnetic resonance imaging after marathon running. Am J Cardiol 103: 1467‐1472, 2009.
 154. Murgo JP, Westerhof N. Input impedance of the pulmonary arterial system in normal man. Effects of respiration and comparison to systemic impedance. Circ Res 54: 666‐673, 1984.
 155. Naeije R. Pulmonary vascular resistance: A meaningless variable? Intens Care Med 29: 526‐529, 2003.
 156. Naeije R. In defence of exercise stress tests for the diagnosis of pulmonary hypertension. Heart 97: 94‐95, 2011
 157. Naeije R, Faoro V. Pathophysiological insight into bubbles. J Appl Physiol 107: 999‐1000, 2010.
 158. Naeije R, Huez S, Lamotte M, Retailleau K, Neupane S, Abramowicz D, Faoro V. Pulmonary artery pressure limits exercise capacity at high altitudes. Eur Respir J 36: 1049‐1055, 2010.
 159. Naeije R, Lejeune P, Leeman M, Melot C, Closset J. Pulmonary vascular responses to surgical chemodenervation and chemical sympathectomy in dogs. J Appl Physiol 66: 42‐50, 1989.
 160. Naeije R, Lipski A, Abramowicz M, Lejeune P, Mélot C, Antoine M, De Smet JM, Leclerc JL, Primo G. Nature of pulmonary hypertension in congestive heart failure. Effects of cardiac transplantation. Am J Respir Crit Care Med 147: 881‐887, 1994.
 161. Naeije R, Mélot C, Mols P, Hallemans R. Effects of vasodilators on hypoxic pulmonary vasoconstriction in normal man. Chest 82: 404‐410, 1982.
 162. Naeije R, Mélot C, Niset G, Delcroix M, Wagner PD. Improved arterial oxygenation by a pharmacological increase in chemosensitivity during hypoxic exercise in normal subjects. J Appl Physiol 74: 1666‐1671, 1993.
 163. Nagueh S, Middleton K, Kopelen H, Zoghbi W, Quinones M. Doppler tissue imaging: a noninvasive technique for evaluation of left ventricular relaxation and estimation of filling pressures. J Am Coll Cardiol 30: 1527‐1533, 1997.
 164. Nandiwada PA, Hyman AL, Kadowitz PJ. Pulmonary vasodilator responses to vagal stimulation and acetylcholine in the cat. Circ Res 53: 86‐95, 1983.
 165. Neilan TG, Januzzi JL, Lee‐Lewandowski E, Ton‐Nu TT, Yoerger DM, Jassal DS, Lewandrowski KB, Siegel AJ, Marshall JE, Douglas PS, Lawlor D, Picard MH, Wood MJ. Myocardial injury and ventricular dysfunction related to training levels among nonelite participants to the Boston marathon. Circulation 114: 2325‐2333, 2006.
 166. Nelin LD, Krenz GS, Rickaby DA, Linehan JH, Dawson CA. A distensible vessel model applied to hypoxic pulmonary vasoconstriction in the neonatal pig. J Appl Physiol 73: 987‐994, 1992.
 167. Newman JH, Butka BJ, Brigham KL. Thromboxane A2 and prostacyclin do not modulate pulmonary hemodynamics during exercise in sheep. J Appl Physiol 61: 1706‐1711, 1986.
 168. Newman JH, Cochran CP, Roselli RJ, Parker RE, King LS. Pressure and flow changes in the pulmonary circulation of exercising sheep: Evidence for elevated microvascular pressure. Am Rev Respir Dis 147: 921‐926, 1993.
 169. Overbeek MJ, Lankhaar JW, Westerhof N, Voskuyl AE, Boonstra A, Bronzwaer JG, Marques KM, Smit EF, Dijkmans BA, Vonk‐Noordegraaf A. Right ventricular contractility in systemic sclerosis‐associated and idiopathic pulmonary arterial hypertension. Eur Respir J 31: 1160‐1166, 2008.
 170. Pace JB. Sympathetic control of pulmonary vascular impedance in anesthetized dogs. Circ Res 29: 555‐567, 1971.
 171. Patterson SW, Piper H, Starling EH. The regulation of the heart beat. J Physiol (London) 48: 465‐513, 1914.
 172. Penazola D, Arias‐Stella J. The heart and pulmonary circulation at high altitudes. Healthy highlanders and chronic mountain sickness. Circulation 115: 1132‐1146, 2007.
 173. Penaloza D, Sime F. Chronic cor pulmonale due to loss of acclimatization (chronic mountain sickness). Am J Med 50: 728‐743, 1971.
 174. Permutt S, Bromberger‐Barnea B, Bane HN. Alveolar pressure, pulmonary venous pressure and the vascular waterfall. Med Thorac 19: 239‐260, 1962.
 175. Podolsky A, Eldridge MW, Richardson RS, Knight DR, Johnson EC, Hopkins SR, Johnson DH, Michimata H, Grassi B, Feiner J, Kurdak SS, Bickler PE, Severinghaus JW, Wagner PD. Exercise‐induced VA/Q inequality in subjects with prior high‐altitude pulmonary edema. J Appl Physiol 81: 922‐932, 1996.
 176. Poon CS. Analysis of linear and mildly nonlinear relationships using pooled subject data. J Appl Physiol 64: 854‐858, 1988.
 177. Quaife RA, Chen MY, Lynch D, Badesch DB, Groves RM, Wolfel E, Robertson AD, Bristow MR, Voelkel NF. Importance of right ventricular end‐systolic regional wall stress in idiopathic pulmonary hypertension: A new method for estimation of right ventricular wall stress. Eur J Med Res 11: 214‐220, 2006.
 178. Reeves JT, Dempsey JA, Grover RF. Pulmonary circulation during exercise. In: Weir EK, Reeves JT, editors. Pulmonary Vascular Physiology and Physiopathology. New York: Marcel Dekker, chap. 4, p. 107‐133, 1989.
 179. Reeves JT, Groves BM. Approach to the patient with pulmonary hypertension. In: Weir EK, Reeves JT, editors. Pulmonary Hypertension. Mt Kisco, NY: Futura, p. 1‐44, 1984.
 180. Reeves JT, Groves BM, Cymerman A, Sutton JR, Wagner PD, Turkevich D, Houston CS. Operation Everest II: Cardiac filling pressures during cycle exercise at sea level. Respir Physiol 80: 147‐154, 1990.
 181. Reeves JT, Groves BM, Sutton JR, Wagner PD, Cymerman A, Malconian MK, Rock PB, Young PM, Houston CS. Operation Everest II: Preservation of cardiac function at extreme altitude. J Appl Physiol 63: 531‐539, 1987.
 182. Reeves JT, Linehan JH, Stenmark KR. Distensibility of the normal human lung circulation during exercise. Am J Physiol Lung Cell Mol Physiol 288: L419‐L425, 2005.
 183. Reeves JT, Taylor AE. Pulmonary hemodynamics and fluid exchange in the lungs at exercise. In: Rowell LB, Shepherd JT, editors. Handbook of Physiology. American Physiological Society, New York, New York: Oxford University Press, sect. 12, p. 585‐613, 1996.
 184. Riley RL, Shepard RH, Cohn JE, Carroll DG, Armstrong BW. Maximal diffusing capacity of the lungs. J Appl Physiol 10: 573‐587, 1954.
 185. Riley RL, Himmelstein A, Motley HL, Weiner H, Cournand A. Studies of the pulmonary circulation at rest and during exercise in normal individuals and in patients with chronic pulmonary disease. Am J Physiol 152: 372‐382, 1948.
 186. Rowland T. Echocardiography and circulatory response to progressive endurance exercise. Sports Med 38: 541‐551, 2008.
 187. Sagawa K, Maughan L, Suga H, Sunagawa K. Cardiac Contraction and the Pressure‐Volume Relationship. New York: Oxford University Press, 1988.
 188. Sancetta SM, Rakita L. Response of pulmonary artery pressure and total pulmonary resistance of untrained, convalescent man to prolonged mild steady state exercise. J Clin Invest 36: 1138‐1149, 1957.
 189. Saouti N, Westerhof N, Helderman F, Marcus JT, Stergiopulos N, Westerhof BE, Boonstra A, Postmus PE, Vonk‐Noordegraaf A. RC time constant of single lung equals that of both lungs together: A study in chronic thromboembolic pulmonary hypertension. Am J Physiol Heart Circ Physiol. 297: H2154‐H2160, 2009.
 190. Saouti N, Westerhof N, Helderman F, Marcius JT, Boonstra A, Postmus PE, Vonk Noordegraaf A. Right ventricular oscillatory power is a constant fraction of total power irrespective of pulmonary artery pressure. Am J Respir Crit Care Med 182: 1315‐1320, 2010.
 191. Schaffartzik WD, Polle DC, Derion T, Tsukimoto K, Mathieu CO, Wagner PD. Pulmonary interstitial edema in the pig after heavy exercise. J Appl Physiol 72: 1657‐1667, 1992.
 192. Scharhag J, Schneider G, Urhausen A, Rochette V, Kramann B, Kindermann W. Athlete's heart: Right and left ventricular mass and function in male endurance athletes and untrained individuals determined by magnetic resonance imaging. J Am Coll Cardiol 40: 1856‐1863, 2002.
 193. Slife DM, Latham RD, Sipkema P, Westerhof N. Pulmonary arterial compliance at rest and at exercise in normal humans. Am J Physiol 258: H1823‐H1828, 1990.
 194. Slonim NB, Ravin A, Balchum OJ, Dressler RH. The effect of mild exercise in the supine position on the pulmonary arterial pressure of five normal human subjects. J Clin Invest 33: 1022‐1030, 1954.
 195. Sobin SS, Fung YC, Tremer HM, Rosenquist TH. Elasticity of the pulmonary alveolar microvascular sheet in the cat. Circ Res 30: 440‐450, 1972.
 196. Sobin SS, Lindal RG, Fung YC, Tremer HM. Elasticity of the smallest noncapillary pulmonary blood vessels in the cat. Microvasc Res 15: 57‐68, 1978.
 197. Stamler JS, Loh E, Reddy MA, Creager MA. Nitric oxide regulates basal systemic and pulmonary vascular resistance in healthy humans. Circulation 89: 2035‐40, 1994.
 198. Stickland MK, Lovering AT. Exercise‐induced intrapulmonary arteriovenous shunting and pulmonary gas exchange. Exerc Sport Sci Rev 34: 99‐106, 2006.
 199. Stickland MK, Welsh RC, Haykowsky MJ, Petersen SR, Anderson WD, Taylor DA, Bouffard M, Jones RL. Intrapulmonary shunt and pulmonary gas exchange during exercise in humans. J Physiol 561: 321‐329, 2004.
 200. Stickland MK, Welsh RC, Haykowsky MJ, Peterson SR, Anderson WD, Taylor DA, Bouffart M, Jones RL. Effect of acute increases in pulmonary vascular pressures on exercise pulmonary gas exchange. J Appl Physiol 100: 1910‐1917, 2006.
 201. Stickland MK, Welsh RC, Petersen SR, Tyberg JV, Anderson WD, Jones RL, Taylor DA, Bouffard M, Haykowsky MJl. Does fitness level modulate the cardiovascular hemodynamic response to exercise? J Appl Physiol 100: 1895‐901, 2006.
 202. Stray‐Gunderson J, Musch TI, Haidot GC, Swain DP, Ordway GA, Mitchell JH. The effects of pericardiectomy on maximal oxygen consumption and maximal cardiac output in untrained dogs. Circ Res 58: 523‐530, 1986.
 203. Stubenetsky R, Verdouw PD, Duncker DJ. Autonomic control of cardiovascular performance and whole body O2 delivery and utilization in swine during treadmill exercise. Cardiovasc Res 39: 459‐474, 1999.
 204. Stuber T, Sartori C, Schwab M, Jayet PY, Rimoldi SF, Garcin S, Thalmann S, Spielvogel H, Salmòn CS, Villena M, Scherrer U, Allemann Y. Exaggerated pulmonary hypertension during mild exercise in chronic mountain sickness. Chest 137; 288‐292, 2010.
 205. Suarez J, Alexander JK, Houston CS. Enhanced left ventricular systolic performance at high altitude during Operation Everest II. Am J Cardiol 60: 137‐142, 1987.
 206. Suga H, Sagawa K, Shoukas AA. Load independence of the instantaneous pressure‐volume ratio of the canine left ventricle and the effect of epinephrine and heart rate on the ratio. Circ Res 32: 314‐322, 1973.
 207. Sui G, Liu Y, Cheng X, Anand IS, Harris E, Harris P, Heath D. Subacute infantile mountain sickness. J Pathol 155: 161‐170, 1988.
 208. Sullivan MJ, Cobb FR, Higginbotham MB. Stroke volume increases by similar mechanisms during upright exercise in men and women. Am J Cardiol 67: 1405‐1412, 1991.
 209. Sunagawa K, Maughan WL, Sagawa K. Optimal arterial resistance for the maximal stroke work studied in the isolated canine left ventricle. Circ Res 56: 586‐595, 1985.
 210. Sunagawa K, Yamada A, Senda Y, Kikuchi Y, Nakamura M, Shibahara T. Estimation of the hydromotive source pressure from ejecting beats of the left ventricle. IEEE Trans Biomed Eng 57: 299‐305, 1980.
 211. Sylvester JT, Harabin AL, Peake MD, Frank RS. Vasodilator and constrictor responses to hypoxia in isolated pig lungs. J Appl Physiol 49: 820‐825, 1980.
 212. Syyed R, Reeves JT, Welsh D, Raeside D, Johnson MK, Peacock AJ. The relationship between the components of pulmonary artery pressure remains constant under all conditions in both health and disease. Chest 133: 633‐639, 2008.
 213. Tabima, DM, Chesler NC. The effects of vasoactivity and hypoxic pulmonary hypertension on extralobar pulmonary artery biomechanics. J Biomech 43: 1864‐1869, 2010
 214. Talreja DR, Nishimura RA, Oh JK. Estimation of left ventricular filling pressure with exercise by Doppler echocardiography in patients with normal systolic function: A simultaneous echocardiographic‐cardiac catheterization study. J Am Soc Echocardiogr 20: 477‐79, 2007.
 215. Thadani U, Parker JO. Hemodynamics at rest and during supine and sitting exercise in normal subjects. Am J Cardiol 41: 52‐59, 1978.
 216. Tolle JJ, Waxman AB, Van Horn TL, Pappagianopoulos PP, Systrom DM. Exercise‐induced pulmonary arterial hypertension. Circulation 118: 2183‐2189, 2008
 217. Tonelli AR, Mubarak KK, Li N, Carrie R, Alnuaimat H. Effect of balloon inflation volume on pulmonary artery occlusion pressure in patients with and without pulmonary hypertension. Chest 139: 115‐121, 2011
 218. Trivax JE, Franklin BA, Goldstein JA, Chinnaiyan KM, Gallagher MJ, de Jong AT, Colar JM, Haines DE, McCullough PA. Acute effects of marathon running. J Appl Physiol 108: 1148‐1153, 2010.
 219. Tuchscherer HA, Webster EB, Chesler NC. Pulmonary vascular resistance and impedance in isolated mouse lungs: Effects of pulmonary emboli. Ann Biomed Eng 34: 660‐668, 2006.
 220. Vanderpool RR, Naeije R, Chesler NC. Impedance in isolated mouse lungs for the determination of site of action of vasoactive agents and disease. Ann Biomed Eng 38: 1854‐1861, 2010.
 221. Varnauskas E, Korsgren M, Hoak P, Prerovsky I. Studies in hypertensive cardiovascular disease. Scand J Clin Lab Invest suppl 86: 100‐117, 1965.
 222. Viswanathan R, Jain SK, Subramanian S, Subramanian TAV, Dua GL, Giri J. Pulmonary edema of high altitude. II. Clinical, aerohemodynamic and biochemical studies in a group with history of pulmonary edema of high altitude. Am Rev Respir Dis 100: 334‐349, 1969.
 223. Vogel JHK, Weaver WF, Rose RL, Blount SG, Grover RF. Pulmonary hypertension on exertion in normal man living at 10150 feet (Leadville, Colorado). Med Thorac 19: 269‐285, 1962.
 224. Wagner PD. Reduced maximal cardiac output at altitude. Mechanisms and significance. Respir Physiol 120: 1‐11, 2000.
 225. Wagner PD, Araoz M, Boushel R, Calbet JA, Jessen B, Rådegran G, Spielvogel H, Søndegaard H, Wagner H, Saltin B. Pulmonary gas exchange and acid‐base state at 5,260 m in high‐altitude Bolivians and acclimatized lowlanders. J Appl Physiol 92: 1393‐1400, 2002.
 226. Wagner PD, Gale GE, Moon RE, Torre‐Bueno J, Stolpe BW, Saltzman HA. Pulmonary gas exchange in humans at sea level and simulated altitude. J Appl Physiol 61: 260‐270, 1986.
 227. Wagner PD, Gillespie JR, Landgren GL, Fedde MR, Jones BW, de Bowes RM, Peischel RL, Erickson HH. Mechanisms of exercise‐induced hypoxemia in horses. J Appl Physiol 66: 1227‐1233, 1989.
 228. Wagner PD, Sutton JR, Reeves JT, Cymerman A, Groves BM, Malconian MK. Operation Everest II: Pulmonary gas echange during a simulated ascent of Mt Everest. J Appl Physiol 63: 2348‐2359, 1987.
 229. Wauthy P, Pagnamenta A, Vassali F, Brimioulle S, Naeije R. Right ventricular adaptation to pulmonary hypertension. An interspecies comparison. Am J Physiol Heart Circ Physiol 286: H1441‐H1447, 2004.
 230. Weir EK, Archer SL. The mechanism of acute hypoxic pulmonary vasoconstriction: the tale of two channels. FASEB J 9: 183‐189, 1995.
 231. West JB. The physiological basis of high altitude diseases. Ann Intern Med 141: 789‐800, 2004.
 232. West JB, Dollery CT, Naimark A. Distribution of blood flow in isolated lung: Relation to vascular and alveolar pressures. J Appl Physiol 19: 713‐724, 1964.
 233. West JB, Mathieu‐Costello O, Jones JH, Birks EK, Logemann RB, Pascoe JR, Tyler WS. Stress failure of pulmonary capillaries in race horses with exerecise‐induced pulmonary hemorrhage. J Appl Physiol 75: 1097‐1109, 1993.
 234. West JB, Tsukimoto K, Mathieu‐Costello O, Prediletto R. Stress failure in pulmonary capillaries. J Appl Physiol 70: 1731‐1742, 1991.
 235. Westerhof N, Stergiopulos N, Noble MIM. Snapshots of Hemodynamics an Aid for Clinical Research and Graduate Education.Second Edition. New York, NY: Springer, 2010.
 236. Williams MH Jr. Relationships between pulmonary artery pressure and blood flow in the dog lung. Am J Physiol 179: 243‐245, 1954.
 237. Yagi H, Yamada H, Kobayashi T, Sekiguchi M. Doppler assessment of pulmonary hypertension induced by hypoxic breathing in subjects susceptible to high altitude pulmonary edema. Am Rev Respir Dis 142: 796‐801, 1990.
 238. Yen RT, Fung YC. Model experiments on apparent blood viscosity and hematocrit in pulmonary alveoli. J Appl Physiol 35: 510‐517, 1973.
 239. Yen RT, Fung YC, Bingham N. Elasticity of small pulmonary arteries in the cat. J Biomech Eng 102: 170‐177, 1980.
 240. Yock P, Popp R. Noninvasive estimation of right ventricular systolic pressure by Doppler ultrasound in patients with tricuspid regurgitation. Circulation 70: 657‐662, 1984.
 241. Zapol WM, Snider MT. Pulmonary hypertension in severe acute respiratory failure. N Engl J Med 296: 476‐480, 1977.
 242. Zhuang FY, Fung YC, Yen RT. Analysis of blood flow in cat's lung with detailed anatomical and elasticity data. J Appl Physiol 55: 1341‐1348, 1983.

Related Articles:

Pulmonary Vascular Disease

Contact Editor

Submit a note to the editor about this article by filling in the form below.

* Required Field

Article Title: Pulmonary Circulation at Exercise
 

How to Cite

Robert Naeije, N. Chesler. Pulmonary Circulation at Exercise. Compr Physiol 2012, 2: 711-741. doi: 10.1002/cphy.c100091