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 Lung Biology in Health Disease Volume 160 Gravity and the
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INTRODUCTION

Weightlessness certainly is not a new phenomenon. Indeed, it was about 400

million years ago that some vertebrates transitioned from an aquatic to a terrestrial
environment. An aquatic environment is a de facto weightless environment, since
the surrounding water provides support to its inhabitants. In contrast, the ter-
restrial vertebrate must support itself against the forces of gravity. Adaptation
to gravitational forces, in turn, leads to specific “structural” and functional
changes.
However, all this occurred over millions of years and, of course, no one
knew about it until the seventeenth century, when Galileo Galilei recognized
gravity as a physical entity. That century has been termed the “Age of the Scien-
tific Revolution,” and during that time speculation was replaced by experimenta-
tion. Among all the giants of the times, Galileo ranks as possibly the greatest;
he developed the mathematical basis of the laws of motion in the terrestrial envi-
ronment.
Today we live in another age of scientific revolution. Although we can
safely pinpoint its beginning—the middle of the twentieth century—there is no
end in sight, nor can we define the limits of this revolution. Just think, from space
to molecules and genes!
The International Space Station is being built and is becoming operational,
and space may also become the “recreational” area of the future (at least for
some). But we have to recognize that the full potential of the Space Station rests
on how physiological systems built to operate under forces of gravity can function
in the absence of these forces.
This is the subject of this monograph. The editors of Gravity and the Lung:
Lessons from Microgravity, Drs. G. Kim Prisk, Manuel Paiva, and John B. West,
and the contributors to this volume are all experts in their own fields and bring
with them years of experience and research.
Over the years, and by way of its many contributors, the Lung Biology in
Health and Disease series has focused on lung physiology and pathology in
the most common, not to say usual, environment. The series has also explored
known responses to one extreme environment, that is, the undersea world (Vol.

iii
iv Introduction

132). The current volume presents some aspects of human response to another
extreme!
As the Executive Editor of this series of monographs, I am as proud to
introduce this volume as I am grateful to the editors and the authors for having
given me the opportunity to do do.

Claude Lenfant, M.D.

Bethesda, Maryland
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PREFACE

Although humans have been traveling in space for 40 years (since the first flight
of Yuri Gagarin in 1961), for the first 20 or so years only a few scientific studies
were performed in space, and those studies were limited. Given the difficulties
of placing and sustaining humans in that distant and hostile environment, this is
hardly surprising. However, the first flight of the European-built Spacelab module
(a laboratory placed in the cargo bay of the space shuttle) in 1983 provided the
opportunity for well-designed and properly controlled experiments. The pressur-
ized Spacelab module flew 14 times during the period from 1983 to 1998. Of
those 14 flights, 11 carried a significant proportion of physiological experiments,
and 3 of those were dedicated to life sciences research. Many of those flights
carried payload specialists—professional scientists who were recruited because
of their highly specialized knowledge and skills. Spacelab was decommissioned
in 1998, following the Neurolab mission, which carried the most complex series
of life sciences experiments performed in space. In the future, the International
Space Station will provide the opportunity for more elaborate studies in micro-
gravity.
Knowledge of environmental physiology comes from understanding how
the organism reacts to changes in the environment. For example, thermoregula-
tory processes might be studied by raising or lowering ambient temperature.
However, for subjects here on earth, gravity is a constant force that cannot be
eliminated. We know that gravity has a large effect on the behavior between the
top and bottom of the lung in an upright subject. There are substantial vertical
gradients in alveolar size and ventilation, in perfusion, and in gas exchange.
Therefore, we would expect to see large differences in the behavior of the lung
in microgravity. In addition, because the lung presents such a large surface area
to the environment (⬃50–100 m2), it is vulnerable to polluted atmospheres such
as might occur in the closed environment of a spacecraft. Yet none of these
factors are unique to the microgravity environment. People routinely change the
gravitational influence on their lungs every time they lie down, and many terres-
trial dwellers are exposed to both indoor and outdoor atmospheric pollution.
Thus, this volume focuses on using measurements made in the absence of gravity

v
vi Preface

(or, in some cases, at high gravity levels) to understand how the lung is affected
by gravity here on earth. The title Gravity and the Lung: Lessons from Micrograv-
ity reflects this approach.
We had the good fortune to be closely involved with several experiments
on Spacelab, and its demise provides an ideal time to summarize those findings
in the field of respiratory physiology before the transition of the Space Station.
We are grateful to Claude Lenfant, Executive Editor, who agreed to place this
volume in the Lung Biology in Health and Disease series.
It is always challenging to perform first-class experiments in a difficult
environment. Performing studies in space presents numerous difficulties, but the
result is exhilarating. It has been a privilege to participate.

G. Kim Prisk
Manuel Paiva
John B. West
CONTRIBUTORS

Jay C. Buckey, Jr., M.D. Research Associate Professor, Department of Medi-

cine, Dartmouth Medical School, Hanover, New Hampshire

Bruce D. Butler, Ph.D. Professor and Vice Chairman of Research, Department

of Anesthesiology, The University of Texas–Houston Medical School, Houston,
Texas

Chantal Darquenne, Ph.D. Assistant Research Physiologist, Department of

Medicine, University of California, San Diego, La Jolla, California

Marc Estenne, M.D., Ph.D. Professor of Medicine, Chest Service, Erasme

University Hospital, Brussels, Belgium

David H. Glaister, O.St.J., Ph.D., M.B., B.S., F.F.O.M. Group Captain (re-
tired), Royal Air Force, Talywern Consultancy Ltd., Powys, United Kingdom

Robb W. Glenny, M.D. Associate Professor, Department of Medicine and

Department of Physiology and Biophysics, University of Washington, Seattle,
Washington

Michael P. Hlastala, Ph.D. Professor, Department of Physiology and Bio-

physics and Department of Medicine, University of Washington, Seattle, Wash-
ington

Dag Linnarsson, M.D., Ph.D. Professor, Department of Physiology and Phar-

macology, Karolinska Institutet, Stockholm, Sweden

Giuseppe Miserocchi, M.D. Professor of Physiology and Chairman, Depart-

ment of Experimental and Environmental Medicine, Università Milano-Bicocca,
Monza, Italy

vii
viii Contributors

Daniela Negrini Associate Professor, Department of Medicine, Surgery and

Dentistry, Università Degli Studi di Milano, Milan, Italy

William T. Norfleet, M.D. Principal Coordinating Scientist for Space Medi-

cine, National Aeronautics and Space Administration, Houston, Texas

Manuel Paiva, Ph.D. Professor, Biomedical Physics Laboratory, Université

Libre de Bruxelles, Brussels, Belgium

G. Kim Prisk, Ph.D. Professor, Department of Medicine, University of Cali-

fornia, San Diego, La Jolla, California

H. Thomas Robertson, M.D. Professor, Pulmonary and Critical Care Medi-

cine, Department of Medicine and Department of Physiology and Biophysics,
University of Washington, Seattle, Washington

John B. West, M.D., Ph.D. Professor, Department of Medicine, University of

California, San Diego, La Jolla, California
CONTENTS

Introduction Claude Lenfant iii

Preface v
Contributors vii

1. Historical Introduction 1
John B. West
I. Normal Gravity 1
II. Increased Acceleration 17
III. Microgravity 21
IV. Epilogue 34
References 34

2. Effects of Acceleration on the Lung 39

David H. Glaister
I. Introduction 39
II. Instrumentation 41
III. Pulmonary Ventilation 42
IV. Pulmonary Perfusion 50
V. Consequences of Acceleration-Induced Ventilation
and Perfusion Inequalities 58
VI. Effect of ⫾Gy Acceleration on Ventilation 65
VII. Modeling 69
VIII. Zero-G Predictions 71
IX. Summary and Conclusions 71
References 72

3. Lung Volumes and Chest Wall Mechanics 75

Marc Estenne
I. Introduction 75

ix
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