Hypoxia and Exercise PDF
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HYPOXIA AND EXERCISE ADVANCES IN EXPERIMENTAL MEDICINE AND BIOLOGY Editorial Board: NATHAN BACK,State University ofNew York at Buffalo IRUN R.COHEN,The Weizmann Institute ofScience DAVID KRITCHEVSKY,Wistar Institute ABEL LAJTHA,N.S.KlineInstitute for Psychiatric Research RODOLFO PAOLETTI,University ofMilan Recent Volumes in this Series Volume 580 THE ARTERIAL CHEMORECEPTORS Edited by Yoshiaki Hayashida,Constancio Gonzalez,and Hisatake Condo Volume 581 THE NIDOVIRUSES:THE CONTROL OF SARS AND OTHER NIDOVIRUS DISEASES Edited by Stanley Perlman and Kathryn Holmes Volume 582 HOT TOPICS IN INFECTION AND IMMUNITY IN CHILDREN III Edited by Andrew J.Pollard and Adam Finn Volume 583 TAURINE 6:UPDATE 2005 Edited by Simo S.Oja and Pirjo Saransaari Volume 584 LYMPHOCYTE SIGNAL TRANSDUCTION Edited by Constantine Tsoukas Volume 585 TISSUE ENGINEERING Edited by John P.Fisher Volume 586 CURRENT TOPICS IN COMPLEMENT Edited by John D.Lambris Volume 587 NEW TRENDS IN CANCER FOR THE 21ST CENTURY 2006 Edited by López-Guerrero,J.A.,Llombart-Bosch,A.,Felipo,V.(Eds.), Volume 588 HYPOXIA AND EXERCISE 2007 Roach,R.,Wagner,P.D.,Hackett,P.(Eds.), A Continuation Order Plan is available for this series.A continuation order will bring delivery of each new volume immediately upon publication. Volumes are billed only upon actual shipment.For further information please contact the publisher. HYPOXIA AND EXERCISE Edited by ROBERT C. ROACH Colorado Center for Altitude Medicine and Physiology University ofColorado at Denver Health Sciences Center Denver,Colorado,USA PETER D. WAGNER Department ofMedicine University ofCalifornia San Diego La Jolla,California,USA PETER H. HACKETT Telluride Medical Clinic Ridgway,Colorado,USA Editors: Robert C.Roach,Ph.D. Peter D.Wagner Peter H.Hackett Research Director UCSD Dept Medicine Telluride Medical Center Altitude Research Center 0623 500 W.Pacific Mail Stop F524, 9500 Gilman Avenue Telluride,Colorado 81435 PO Box 6508 La Jolla,CA 92093-0623 [email protected] University of Colorado [email protected] Health Sciences Center Aurora,Colorado 80045-0508 [email protected] Library ofCongress Control Number:2006927789 Printed on acid-free paper. ISBN-10:0-387-34816-6 ISBN-13:978-0387-34816-2 e-ISBN:0-387-34817-4 Proceedings of the 14th International Hypoxia Symposium,held in Chateau Lake Louise, Lake Louis,Alberta,Canada,February 22-27,2005. © 2006 Springer Science+Business Media,LLC All rights reserved.This work may not be translated or copied in whole or in part without the written permission ofthe publisher (Springer Science+Business Media,LLC,233 Spring Street, New York,NY 10013,USA),except for brief excerpts in connection with reviews or scholarly analysis. Use in connection with any form of information storage and retrieval, electronic adaptation,computer software,or by similar or dissimilar methodology now known or hereafter developed is forbidden. The use in this publication oftrade names,trademarks,service marks and similar terms,even if they are not identified as such,is not to be taken as an expression of opinion as to whether or not they are subject to proprietary rights. Printed in the United States ofAmerica. 9 8 7 6 5 4 3 2 1 springer.com The 14th International Hypoxia Symposium was dedicated to the memory of Dr. John T. “Jack” Reeves. As most of you know, Dr. John T. “Jack” Reeves passed away in September, 2004 fol- lowing a motor vehicle-bicycle accident. Jack was a former Professor of Medicine, Pedi- atrics and Surgery and Professor Emeritus at the University of Colorado Health Sciences Center, having joined the faculty in 1972. He made exceptional contributions in teach- ing/mentoring, research, administration and leadership. He was a scientist of international stature. He made major advances at the molecular, cellular, animal and human level with regard to the pulmonary circulation and adaptation to high altitude. For many years Jack was a senior member of the Cardiovascular Pulmonary Laboratory of the School of Medi- (cid:70)(cid:76)(cid:81)(cid:72)(cid:3)(cid:90)(cid:76)(cid:87)(cid:75)(cid:76)(cid:81)(cid:3)(cid:87)(cid:75)(cid:72)(cid:3)(cid:39)(cid:72)(cid:83)(cid:68)(cid:85)(cid:87)(cid:80)(cid:72)(cid:81)(cid:87)(cid:3)(cid:82)(cid:73)(cid:3)(cid:48)(cid:72)(cid:71)(cid:76)(cid:70)(cid:76)(cid:81)(cid:72)(cid:3)(cid:68)(cid:81)(cid:71)(cid:3)(cid:80)(cid:82)(cid:86)(cid:87)(cid:3)(cid:85)(cid:72)(cid:70)(cid:72)(cid:81)(cid:87)(cid:79)(cid:92)(cid:3)(cid:83)(cid:79)(cid:68)(cid:92)(cid:72)(cid:71)(cid:3)(cid:68)(cid:3)(cid:86)(cid:76)(cid:74)(cid:81)(cid:76)(cid:191)(cid:70)(cid:68)(cid:81)(cid:87)(cid:3)(cid:85)(cid:82)(cid:79)(cid:72)(cid:3)(cid:76)(cid:81)(cid:3)(cid:87)(cid:75)(cid:72)(cid:3) establishment of the Colorado Center for Altitude Medicine and Physiology (CCAMP). He was a key, active and memorably vocal member of the Hypoxia family. No one who ever met Jack at Hypoxia will forget him; his spirit lives on within the Hypoxia community. In honor of Jack, we established The Reeves Prize for Presentation Excellence. This prize will be awarded at the biennial International Hypoxia Symposium to the speaker (cid:77)(cid:88)(cid:71)(cid:74)(cid:72)(cid:71)(cid:3)(cid:87)(cid:82)(cid:3)(cid:83)(cid:85)(cid:72)(cid:86)(cid:72)(cid:81)(cid:87)(cid:3)(cid:87)(cid:75)(cid:72)(cid:3)(cid:80)(cid:82)(cid:86)(cid:87)(cid:3)(cid:82)(cid:88)(cid:87)(cid:86)(cid:87)(cid:68)(cid:81)(cid:71)(cid:76)(cid:81)(cid:74)(cid:3)(cid:86)(cid:70)(cid:76)(cid:72)(cid:81)(cid:87)(cid:76)(cid:191)(cid:70)(cid:3)(cid:87)(cid:68)(cid:79)(cid:78)(cid:15)(cid:3)(cid:90)(cid:76)(cid:87)(cid:75)(cid:3)(cid:86)(cid:83)(cid:72)(cid:70)(cid:76)(cid:68)(cid:79)(cid:3)(cid:72)(cid:80)(cid:83)(cid:75)(cid:68)(cid:86)(cid:76)(cid:86)(cid:3)(cid:82)(cid:81)(cid:3)(cid:70)(cid:79)(cid:68)(cid:85)(cid:76)(cid:87)(cid:92)(cid:3)(cid:82)(cid:73)(cid:3) presentation skills that were cherished, taught and practiced by Jack. The recipient will be named at the closing banquet by the organizers of the International Hypoxia Symposia with funds coming from the John Sutton Fund, McMaster University, the Colorado Cen- ter for Altitude Medicine and Physiology, University of Colorado at Denver and Health Sciences Center, and the International Hypoxia Symposia. The winner for 2005 was Dr. Randy Sprague. See his written contribution starting on page 207. v PREFACEANDACKNOWLEDGEMENTS The International Hypoxia Symposia convenes every other year to bring together in- (cid:87)(cid:72)(cid:85)(cid:81)(cid:68)(cid:87)(cid:76)(cid:82)(cid:81)(cid:68)(cid:79)(cid:3)(cid:72)(cid:91)(cid:83)(cid:72)(cid:85)(cid:87)(cid:86)(cid:3)(cid:73)(cid:85)(cid:82)(cid:80)(cid:3)(cid:80)(cid:68)(cid:81)(cid:92)(cid:3)(cid:191)(cid:72)(cid:79)(cid:71)(cid:86)(cid:3)(cid:87)(cid:82)(cid:3)(cid:72)(cid:91)(cid:83)(cid:79)(cid:82)(cid:85)(cid:72)(cid:3)(cid:87)(cid:75)(cid:72)(cid:3)(cid:86)(cid:87)(cid:68)(cid:87)(cid:72)(cid:3)(cid:82)(cid:73)(cid:3)(cid:87)(cid:75)(cid:72)(cid:3)(cid:68)(cid:85)(cid:87)(cid:3)(cid:76)(cid:81)(cid:3)(cid:81)(cid:82)(cid:85)(cid:80)(cid:68)(cid:79)(cid:3)(cid:68)(cid:81)(cid:71)(cid:3)(cid:83)(cid:68)(cid:87)(cid:75)(cid:82)- physiological responses to hypoxia. Representatives from 18 countries joined together in (cid:41)(cid:72)(cid:69)(cid:85)(cid:88)(cid:68)(cid:85)(cid:92)(cid:3)(cid:21)(cid:19)(cid:19)(cid:24)(cid:3)(cid:73)(cid:82)(cid:85)(cid:3)(cid:73)(cid:82)(cid:88)(cid:85)(cid:3)(cid:71)(cid:68)(cid:92)(cid:86)(cid:3)(cid:82)(cid:73)(cid:3)(cid:76)(cid:81)(cid:87)(cid:72)(cid:81)(cid:86)(cid:72)(cid:3)(cid:86)(cid:70)(cid:76)(cid:72)(cid:81)(cid:87)(cid:76)(cid:191)(cid:70)(cid:3)(cid:71)(cid:76)(cid:86)(cid:70)(cid:82)(cid:88)(cid:85)(cid:86)(cid:72)(cid:3)(cid:76)(cid:81)(cid:3)(cid:87)(cid:75)(cid:72)(cid:3)(cid:71)(cid:85)(cid:68)(cid:80)(cid:68)(cid:87)(cid:76)(cid:70)(cid:3)(cid:80)(cid:82)(cid:88)(cid:81)(cid:87)(cid:68)(cid:76)(cid:81)(cid:3)(cid:86)(cid:72)(cid:87)(cid:87)(cid:76)(cid:81)(cid:74)(cid:3) of Lake Louise, Canada. The 14th International Hypoxia Symposium was a rewarding experience due to the outstanding faculty and the lively participation of our largest-ever group of participants. At this, our fourth meeting as the organizers, we were especially pleased that the experience known as the Hypoxia Meetings can continue to prosper. We remain always thankful for the kind and wise guidance of Charlie Houston, the originator of the Hypoxia meetings. As editors of the Proceedings of the International Hypoxia Symposia, we strive to main- tain a 28 year tradition of presenting a stimulating blend of clinical and basic science papers focused on hypoxia. Topics covered in 2005 included a history of high altitude physiology, arterial hypoxemia in exercise, sleep and hypoxia, genetic components of adaptation to hypoxia, erythropoietin, cell stress pathways and hypoxia, the role of red blood cells, ATP release and hemoglobin in the control of vasoreactivity, exercise and altitude training, the eye at high altitude and pulmonary hypertension in high altitude residents. An update on methods to measure the hypoxic ventilatory response was also featured. The abstracts from the 2005 meeting were published in High Altitude Medicine & Biol- ogy Dec 2004, Vol. 5, No. 4: 471-509. Late abstracts are presented staring on page 311, this volume. We hope that this collection of papers especially prepared for this volume allows us to share with a broader audience some of the intellectual excitement that embodies the spirit of the Hypoxia meetings. In 2005 we had the generous support of a number of organizations and individuals, including the U.S. Army Research and Development Command, The White Mountain Re- search Station, our International Advisory Committee. At the meeting we were greatly helped by Barbara Lommen, Paige Sheen, Mollie Pritcher, Gene and Rosann McCullough and Andy Subudhi who each made a tremendous effort to make every delegate feel at home, and to make the meeting go very smoothly. Please join us by the light of the full moon in February 2007 at the Chateau Lake Louise, Lake Louise, Alberta, Canada for the 15th International Hypoxia Symposium. Robert C. Roach, Peter D. Wagner, Peter H. Hackett, Editors, Fall 2005 (www.hypoxia.net) vii CONTENTS HYPOXIA HONOREE 1. A Tribute To John Burnard West 1 James S Milledge 2. Adventures In High-Altitude Physiology 7 John B. West EXERCISE-INDUCED HYPOXEMIA (EIH): CAUSES AND CONSEQUENCES 3. Exercise Induced Arterial Hypoxemia: The Role Of Ventilation-Perfusion 17 Inequality And Pulmonary Diffusion Limitation Susan R. Hopkins 4. Intrapulmonary Shunt During Normoxic And Hypoxic Exercise In Healthy 31 Humans Andrew T. Lovering, Michael K. Stickland, Marlowe W. Eldridge 5. Exercise-Induced Arterial Hypoxemia: Consequences For Locomotor Muscle 47 Fatigue Lee M. Romer, Jerome A. Dempsey, Andrew Lovering, Marlowe Eldridge CAUSES AND EFFECTS OF SLEEP-DISORDERED BREATHING 6. Mechanisms Of Sleep Apnea At Altitude 57 William Whitelaw 7. Control Of Cerebral Blood Flow During Sleep And The Effects Of Hypoxia 65 Douglas R. Corfi eld and Guy E. Meadows xi x CONTENTS 8. Neural Consequences Of Sleep Disordered Breathing: The Role Of 75 Intermittent Hypoxia Mary J Morrell and Gillian Twigg GENETICS, GENOMICSANDADAPTATION TO HIGHALTITUDE 9. Finding The Genes Underlying Adaptation To Hypoxia Using Genomic Scans 89 For Genetic Adaptation And Admixture Mapping Mark D. Shriver, Rui Mei, Abigail Bigham, Xianyun Mao,Tom D. Brutsaert, Esteban J. Parra, Lorna G. Moore 10. An Evolutionary Model For Identifying Genetic Adaptation To High Altitude 101 Lorna G. Moore, Mark Shriver, Lynne Bemis, and EnriqueVargas 11. Hypoxic Preconditioning And Erythropoietin Protect Retinal Neurons From 119 Degeneration Christian Grimm, A. Wenzel, N. Acar, S. Keller, M. Seeliger, Max Gassmann BLOCKINGSTRESSPATHWAYS WITHCELL-PERMEABLE MEMBRANES 12. Blocking Stress Signaling Pathways With Cell Permeable Peptides 133 Christophe Bonny 13. Jnk Pathway As Therapeutic Target To Prevent Degeneration In The 145 Central Nervous System Mariaelena Repici and Tiziana Borsello 14. Salvage Of Ischemic Myocardium: A Focus On Jnk 157 Hervé Duplain THEIMPACT OF IMMOBILIZATIONAND HYPOXIAONSKELETAL MUSCLE (cid:20)(cid:24)(cid:17)(cid:3)(cid:48)(cid:76)(cid:87)(cid:82)(cid:70)(cid:75)(cid:82)(cid:81)(cid:71)(cid:85)(cid:76)(cid:68)(cid:79)(cid:3)(cid:53)(cid:72)(cid:68)(cid:70)(cid:87)(cid:76)(cid:89)(cid:72)(cid:3)(cid:50)(cid:91)(cid:92)(cid:74)(cid:72)(cid:81)(cid:3)(cid:54)(cid:83)(cid:72)(cid:70)(cid:76)(cid:72)(cid:86)(cid:3)(cid:36)(cid:85)(cid:72)(cid:3)(cid:53)(cid:72)(cid:84)(cid:88)(cid:76)(cid:85)(cid:72)(cid:71)(cid:3)(cid:41)(cid:82)(cid:85)(cid:3)(cid:43)(cid:92)(cid:83)(cid:82)(cid:91)(cid:76)(cid:70)(cid:3)(cid:43)(cid:76)(cid:73)(cid:302) 165 Stabilization M. Celeste Simon 16. Hypoxia-Induced Gene Activity In Disused Oxidative Muscle 171 Christoph Däpp, Max Gassmann, Hans Hoppeler, Martin Flück CONTENTS xi HYPOXIC VASOREGULATION:INTERACTIONSOF REDCELLS, ENDOTHELIUMANDSMOOTH MUSCLE 17. Role Of The Red Blood Cell In Nitric Oxide Homeostasis And Hypoxic 189 Vasodilation Mark T. Gladwin 18. Expression Of The Heterotrimeric G Protein Gi And Atp Release Are 207 Impaired In Erythrocytes Of Humans With Diabetes Mellitus Randy Sprague, Alan Stephenson, Elizabeth Bowles, Madelyn Stumpf, Gregory Ricketts and Andrew Lonigro 19. Red Blood Cells And Hemoglobin In Hypoxic Pulmonary Vasoconstriction 217 Steven Deem HYPOXIA:STATEOF THE ART 20. Dose-Response Of Altitude Training: How Much Altitude Is Enough? 233 Benjamin D. Levine, James Stray-Gundersen 21. The Eye At Altitude 249 Daniel S Morris, John Somner, Michael J Donald, Ian JC McCormick, Rupert RA Bourne, Suber S Huang, Peter Aspinall, Baljean Dhillon 22. Lake Louise Consensus Methods For Measuring The Hypoxic Ventilatory 271 Response Frank L. Powell FUTUREDIRECTIONSIN HYPOXIARESEARCH 23. Pulmonary Hypertension In High-Altitude Dwellers: Novel Mechanisms, 277 Unsuspected Predisposing Factors Urs Scherrer, Pierre Turini, Sébastien Thalmann, Damian Hutter, Carlos Salinas Salmon, Thomas Stuber, Sidney Shaw, Pierre-Yves Jayet, Céline Sartori-Cucchia, Mercedes Villena, Yves Allemann and Claudio Sartori 24. Gene Hunting In Hypoxia And Exercise 293 Kenneth B. Storey LATEABSTRACTS 311 AUTHORINDEX 347 SUBJECT INDEX 351 AUTHORS FOR CORRESPONDENCE Christophe Bonny Martin Flück Unit of Molecular Genetics, CHUV Department of Anatomy- University of Bern CH1011 Lausanne, Switzerland Baltzerstrasse 2 Email:[email protected] CH3000 Bern Switzerland (Chapter 12) (cid:40)(cid:80)(cid:68)(cid:76)(cid:79)(cid:29)(cid:3)(cid:192)(cid:88)(cid:72)(cid:70)(cid:78)(cid:35)(cid:68)(cid:81)(cid:68)(cid:17)(cid:88)(cid:81)(cid:76)(cid:69)(cid:72)(cid:17)(cid:70)(cid:75)(cid:3)(cid:3) (Chapter 16) Tiziana Borsello BDCM, rue du Bugnon 9 Max Gassmann CH1005 Switzerland University of Zurich Email: [email protected] Institute of Veterinary Physiology (Chapter 13) Winterthurerstrasse 260 CH-8057 Zurich Switzerland (cid:39)(cid:82)(cid:88)(cid:74)(cid:79)(cid:68)(cid:86)(cid:3)(cid:38)(cid:82)(cid:85)(cid:191)(cid:72)(cid:79)(cid:71) E-mail: [email protected] Keele University, School of Life Sciences (Chapter 11) Keele, Staffordshire, United Kingdom, ST5 5BG (cid:48)(cid:68)(cid:85)(cid:78)(cid:3)(cid:55)(cid:17)(cid:3)(cid:42)(cid:79)(cid:68)(cid:71)(cid:90)(cid:76)(cid:81) (cid:40)(cid:80)(cid:68)(cid:76)(cid:79)(cid:29)(cid:3)(cid:71)(cid:17)(cid:70)(cid:82)(cid:85)(cid:191)(cid:72)(cid:79)(cid:71)(cid:35)(cid:78)(cid:72)(cid:72)(cid:79)(cid:72)(cid:17)(cid:68)(cid:70)(cid:17)(cid:88)(cid:78)(cid:3) National Institutes of Health (Chapter 7) NHLBI-CVB 10 Center Drive Bldg 10-CRC StevenDeem Room 5-5142 University of Washington Baltimore MD United States 20892-1454 Dept. of Anesthesiology, Box 359724 Email: [email protected] Harborview Medical Center (Chapter 17) Seattle WA United States 98104 Email: [email protected] SusanR. Hopkins (Chapter 19) University of California, San Diego Division of Physiology Jerome A.Dempsey 9500 Gilman Dr Univ of Wisconsin, Med Sci Bldg La Jolla CA United States 92093-0623 1300 University Ave Rm4245 Email: [email protected] Madison WI United States 53706 (Chapter 3) Email: [email protected] (Chapter 2) BenjaminD. Levine Institute for Exercise and Environmental HérveDuplain Medicine CHUV, Département de Médecine Interne 7232 Greenville Ave, Suite 435 CH1011 Lausanne, Switzerland Dallas TX United States 75231 Email: [email protected] Email: [email protected] (Chapter 14) (Chapter 20) xiii
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