Just Enough Physiology

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Library of Congress Cataloging-in-Publication Data

Munis, James R.
Just enough physiology / James R. Munis.
p. ; cm.
ISBN 978–0–19–979779–0 (pbk.)
I. Title.
[DNLM: 1. Physiological Phenomena. QT 4]
LC classification not assigned
612.1—dc23 2011030222

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987654321
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For Lisa
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 PREFACE

THIS BOOK EMERGED FROM a long tradition of one-on-one

teaching, and I hope it reads that way. What began as notes on
chalkboards and scraps of paper eventually became a teaching mono-
graph. These essays, aided and abetted by various illustrations and
observations, were used as a core resource for a second-year physi-
ology course at Mayo Medical School, starting in 2002. It was also
used since that time as a handout for rotating and visiting residents
in anesthesiology and critical care. Following a kind and enthusi-
astic recognition by the Anesthesia Foundation in 2008, Oxford
University Press and Mayo Clinic Scientific Press have shaped Just
Enough Physiology into a real textbook. My own illustrations have
been replaced with the far more professional work of James Tidwell,
and each essay has been edited and rewritten, but with an emphasis
on preserving a consistent voice. I say “essay” because that is really
what each chapter is—a story. People remember stories longer than
they do arguments, equations, or concepts. So, by embedding argu-
ments, equations, and concepts—the necessary stuff of physiology—
within stories, I hope that you will learn more and remember more.
For more than 2 decades, I have taught what I hope have been
a few surprising but useful concepts about cardiopulmonary physi-
ology to college, medical, and other professional students in the
classroom, as well as to residents and fellows in the operating room
and hospital. Like many other teachers, I have discovered that our
brains can hold only so much at one time. That is why Just Enough
Physiology is divided into small, easily digested topics. I have kept
each one short in the hope that you will be able to dine on an entire
topic at one sitting and still have room for dessert. Left to our own
viii / PREFACE

devices as teachers and students, I believe that is how we best learn,

remember, and apply.
In one sense, Just Enough Physiology takes an inverted approach
to teaching. Many textbooks are written explicitly for board prepa-
ration and hope to sneak in some real science along the way. This
book will do just the opposite: it will take you by the hand through
some of the theory, history, and quirky places where the science of
physiology makes its home (Mt Everest, Mars, or beneath the sea,
for example); preparing you to answer questions is an imperceptible
side benefit. By way of reassurance (but without making any extraor-
dinary claim to comprehensiveness), I think you are unlikely to field
questions about the physiology of the heart, lungs, or circulation that
Just Enough Physiology doesn’t equip you to answer. Most of what
you need to know for the boards or on rounds is in here somewhere.
One of my mentors during my postgraduate training was Dr
Richard Teplick at the Massachusetts General Hospital. He taught a
year-long physiology and pathophysiology course for critical care fel-
lows that began with Newton’s Laws of Motion. Dr Lawrence Wood
did the same at the University of Chicago. The benefit of beginning
at the beginning—that is, with basic physical principles—cannot be
overstated. Physiology is the science that is applied at the boundary
between life and death; this is why it’s so important to those of us
who tread that same boundary every day in the practice of anesthe-
siology and critical care. The functional difference between a patient
who has just died and one who is still alive is physiology.
What the heart, lungs, and circulation do in life is best under-
stood through some simple, unifying principles of mechanics and
chemistry. But that is not enough. There is also a way of thinking
about these concepts that helps pull them together. Interestingly,
that form of logic is almost identical to the way our brains work
when we are solving logic puzzles. To that effect, brain teasers are
included at the end of each chapter. I hope that your brain is both
enriched and teased as you work your way through these stories.

James Munis, MD, PhD

Rochester, Minnesota
 ACKNOWLEDGMENTS

I HAVE BEEN BLESSED with a few good mentors. Their finger-

prints are present in every insight and absent in every oversight of this
book. Their curiosity drove mine, and I hope to convey its full force
to my own students and readers. Lawrence D. H. Wood, MD, PhD,
of the University of Chicago; John B. West, MD, PhD, DSc, James W.
Covell, MD, Richard S. Kornbluth, MD, PhD, and Douglas D.
Richman of the University of California, San Diego; and Warren M.
Zapol, MD, of Massachusetts General Hospital and Solomon Snyder,
MD, DSc, of The Johns Hopkins University taught me to think like a
physiologist. In return, I taught them to be patient and persevering.
I am indebted to Sarah Flannery, whose charming book In Code
reminded me of some long-forgotten brain teasers and also taught
me some new ones. I only hope that these little logic puzzles stir
my own readers to, like me, stare absently past companions while
trying to solve riddles in the middle of the day. These puzzles
are a delightful addiction, and they teach reasoning well suited to
physiology.
Along with the Cleveland Clinic, I hold a patent on an infusion
pump that is intended to measure peripheral venous pressure (PVP).
I have a great deal to say about PVP in this book for other reasons,
and to teach other lessons, but I do not mention or advocate the use
of this specific device.
Finally, I would like to acknowledge the excellent professionals
in Mayo Clinic Scientific Publications and Media Support Services
who brought this book to life. They include my editor, June Oshiro,
PhD, my illustrator, James Tidwell, and the project coordinators who
x / ACKNOWLEDGMENTS

made it all happen: Kenna Atherton, Roberta Schwartz, and LeAnn

Stee. Barb Golenzer, our editorial assistant, and John Hedlund, our
proofreader, kept us all on the same page. Special thanks go to Joseph
G. Murphy, MD, of Mayo Clinic Scientific Press and Doris K. Cope,
MD, of the University of Pittsburgh for thinking outside of the box to
champion the cause of Just Enough Physiology.
 CONTENTS

1. Pressure and Its Measurement 1

2. Atmospheric and Alveolar Pressures 12
3. Hydrostatic Pressure 19
4. Doctor Dolittle Visits a Sitting Case 27
5. In the Loop—Left Ventricular Pressures 42
6. What Goes Around Comes Around—Venous Return 48
7. Pushmi-Pullyu and the Right Atrium 55
8. Pressure and Flow—Chickens and Eggs 63
9. Down But Not Out—Circulatory Arrest Pressures 70
10. Starling’s Riddle of the Broken Heart 77
11. Oxygen and the Gradients of Life 88
12. The Two Doctors Fick 94
13. A Breath of Fresh Air—Ventilation 101
14. Pulmonary Function Tests 108
15. Where Breath Meets Blood—Lung Perfusion 122
16. Bird Brains and Bird Breath 129
17. Diffusion Limitation—Montana Style 135
18. Man, Machine, and Homeostasis 141
19. Putting It All Together—Manned Space Flight 149

INDEX 157
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 ∙1∙
PRESSURE AND ITS MEASUREMENT

Definition of Pressure

IN PHYSIOLOGIC TERMS, WE are exposed to 3 main sources

of pressure: 1) the weight of the atmosphere; 2) hydrostatic forces
exerted by the weight of body fluids; and 3) mechanical pressure
generated by the heart or other muscles that contract around those
fluids. Because cardiopulmonary physiology deals so much with
pressure measurements, let’s start by defining what pressure really
is. Simply put, pressure is force divided by area.
It’s also important to understand what pressure is not. For
example, pressure is not energy. Only when pressure is coupled to a
volume change (ie, movement or pressure-volume work) is it a com-
ponent of energy. This is more than just a semantic point. Although
we’re fond of saying that fluids move from high to low pressure,
that isn’t always true. The reason why highlights a fundamental dif-
ference between pressure and energy. For example, in Figure 1.1,
the pipe contains a fluid that is flowing from left to right through a
constricted region. The vertical tube in each segment is a pressure
manometer.
The eighteenth-century Swiss mathematician Daniel Bernoulli
taught us that a fluid in motion contains a total quantity of energy,
termed total fluid energy. Total fluid energy is simply the sum of 3
elements: 1) pressure energy, a form of potential energy available to
push fluid out from the tube if a hole is placed in its wall; 2) hydrostatic
2 / JUST ENOUGH PHYSIOLOGY

Fluid moves from high to low total fluid energy,

not necessarily from high to low pressure
Pressure ≠ Energy

P1 P2 P3

Figure 1.1. Water Flows Through a Constricted Region.

energy, the potential energy due to elevation of the fluid above a refer-
ence point; and 3) kinetic energy of motion.

Total Fluid Energy (per unit of volume) = P + ρgz + 1/2 ρv2

where P indicates pressure energy; ρ, density of the fluid; g, accel-

eration due to gravity; z, height above a reference point; and v, fluid
velocity.
As the fluid moves through the constricted area, it gains speed
(just like a rapid in a narrowing river). More speed means more
kinetic energy. By the law of conservation of energy, the increase
in kinetic energy must be counterbalanced by a decrease in pres-
sure energy; thus, in our example, the pressure manometer in the
narrower segment shows a lower pressure. The law also explains
why the fluid can move from an area of lower pressure (P2) to
an area of higher pressure (P3)—that is, because total fluid energy
remains constant (neglecting energy lost through frictional heat).
This application of the law of conservation of energy, widely
 1. Pressure and Its Measurement / 3

Q: Since P1 > P2,

why doesn’t water
flow from P1 to P2 ?
P2 A: Because the
total fluid energy
of P1 = total fluid
energy of P2

P1

Figure 1.2. Pressure in a Glass of Water.

known as the Bernoulli principle, also pertains to airflow over

a wing.
An even simpler example illustrating that fluids do not always
move from high to low pressure comes from the physiologist A.C.
Burton in his book Physiology and Biophysics of the Circulation. In
Figure 1.2, the water at the bottom of the glass has a higher pres-
sure than the water at the top, but we know that water doesn’t flow
spontaneously along the pressure gradient from bottom to top.
The reason is the same as before: the total fluid energy, which
includes pressure energy and the energy of position, is the same at
the bottom of the glass as it is at the top. P1 has greater pressure
energy than P2, but P2 has more energy of position than P1—and
the 2 offset each other exactly.
Let’s take another case that’s a bit closer to home to see whether
you understand the distinction between pressure and energy. Is it
accurate to say that the heart generates a pressure gradient that
drives blood flow through a resistance? The answer is no. When
the heart contracts and generates a pressure gradient, the gradient is
abolished as soon as blood flows from one place in the circulation to
another. The heart needs to contract again to maintain the difference
4 / JUST ENOUGH PHYSIOLOGY

in total fluid energy between the arteries and veins. The business of
the heart is not to maintain pressure gradients—it is to impart fluid
energy, which includes a kinetic energy component, to the blood,
thereby moving blood from veins to arteries. Pressure gradients are
a consequence, not the cause, of that movement of blood. You’ll
learn more about this in Chapter 8 (Pressure and Flow—Chickens
and Eggs).
For the time being, though, consider what happens when we
temporarily replace the heart with a mechanical cardiopulmonary
bypass system. (It’s always instructive to learn from the experience
of replacing the function of a normal organ with a mechanical ana-
logue. When we do that, we learn more about the physiology of
the natural organ.) Which do we adjust on the machine, pressure or
flow? If you didn’t know anything else about how cardiopulmonary
bypass works, would you say that the machine generates flow and
that the pressure gradients observed are the result? Or would you say
that the machine generates pressure and flow results? In Chapter 8
(Pressure and Flow—Chickens and Eggs), you’ll see the rationale
for adopting the former rather than the latter statement.

Measuring Pressure

Pressure is surprisingly difficult to measure. Often, when we

think we are measuring pressure, we are actually measuring stretch
or movement. A pressure transducer contains a small strain gauge
that stretches (or otherwise deforms) in proportion to the pressure
difference between the 2 sides. This stretch is coupled to an electri-
cal signal, which is converted to a pressure reading that is refer-
enced (“zeroed”) against atmospheric pressure.
The carotid sinus and aortic arch baroreceptors operate with a
similar mechanism. Although “baro” means pressure (see the dis-
cussion of Mr Torricelli below), these receptors don’t measure pres-
sure directly—instead, they measure stretch. In a classic experiment,
the carotid sinuses of an anesthetized dog were encased in plaster
of Paris while the activity of the carotid sinus nerve ending was
recorded. The firing rate did not change with an increase in carotid