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Essential Pharmacology
Essential
Pharmacology
How Drugs Alter Signaling
Pathways in the Body
Carl E. Creutz, PhD
Harrison Professor of Medical Teaching in Pharmacology, Emeritus
University of Virginia, Charlottesville, VA
USA
Copyright © 2026 by John Wiley & Sons, Inc. All rights reserved, including rights for text and data mining
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Library of Congress Cataloging- in- Publication Data
Names: Creutz, Carl E. (Professor of Medical Teaching in Pharmacology)
author
Title: Essential pharmacology : how drugs alter signaling pathways in the
body / Carl E. Creutz, PhD, Harrison Professor of Medical Teaching in
Pharmacology, Emeritus, University of Virginia.
Description: First edition. | Hoboken, NJ, USA : Wiley, [2026] | Includes
index.
Identiers: LCCN 2026003498 (print) | LCCN 2026003499 (ebook) | ISBN
9781394347858 Paperback | ISBN 9781394347872 adobe pdf | ISBN
9781394347865 epub
Subjects: LCSH: Pharmacology | LCGFT: Handbooks and manuals
Classication: LCC RM301.12 .C74 2026 (print) | LCC RM301.12 (ebook)
LC record available at https://lccn.loc.gov/2026003498
LC ebook record available at https://lccn.loc.gov/2026003499
Cover Design: Wiley
Cover Images: Courtesy of Carl Eugene Creutz
Set in 11.5/14pt STIXTwoText by Straive, Pondicherry, India
v
Contents
About the Author viii
Chapter1: Introduction 1
Chapter2:
Chapter3:
Chapter4:
Chapter5:
vi CONTENTS
Chapter6:
Chapter7:
Chapter8:
Chapter9:
Contents vii
Chapter10:
Chapter11:
Chapter12:
Index
viii
About the Author
CARL E. CREUTZ After receiving a B.S. in Physics at Stanford
University (1969), an M.S. in Physics from the University of Wis-
consin (1970), and a Ph.D. in Biophysics from Johns Hopkins Uni-
versity (1976), Dr. Creutz conducted basic research in molecular
and cell biology at the National Institutes of Health in Bethesda,
MD (1976–1981). In 1981, he was recruited to the Department of
Pharmacology at the University of Virginia as an Assistant Profes-
sor. In 1987, he was promoted to Associate Professor, and in 1994
to Full Professor. In 2003 Dr. Creutz was elected as the Harrison
Professor of Medical Teaching in Pharmacology.
From 1985 to 1990, Dr. Creutz’s research was supported
by an Established Investigator Fellowship from the American
Heart Association. During a sabbatical leave from 1995 to 1996,
Dr.Creutz served as an NIH- funded Fogarty Senior International
Fellow at the Centre National de la Recherche Scientifique (CNRS)
in Gif- sur- Yvette, France. During a sabbatical in 2008, Dr. Creutz
served as the Thomas Jefferson Visiting Fellow at Downing Col-
lege, University of Cambridge, England.
About the Author ix
Dr. Creutz’s research has focused on the roles of calcium-
dependent, membrane- binding proteins in exocytosis and other
membrane- based phenomena in stimulated cells and has led to sev-
eral novel and expanding areas of research, including the follow-
ing. Annexins. Dr. Creutz was one of the discoverers of the annexin
family of calcium-dependent proteins and explored their functional
roles in exocytosis, endocytosis, and membrane repair in health and
disease. Synaptotagmins. Dr. Creutz’s group conducted some of the
original characterization of the biochemical and membrane-bind-
ing activities of the calcium-binding domains of synaptotagmin, a
protein that regulates membrane fusion in exocytosis in neurons
and other cells. Copines. During his sabbatical in France, Dr. Creutz
discovered a novel group of calcium- dependent proteins he named
copines in the model organism Paramecium tetraaurelia. Subse-
quently, his group characterized copines from a number of different
organisms and provided a detailed biochemical characterization of
the major human copine, copine I. The copines are calcium signal-
ing proteins that organize other peripheral proteins on membranes
in response to calcium mobilization.
Dr. Creutz has made major contributions to the teaching and
training programs of the University of Virginia. He has trained
13postdoctoral fellows in his laboratory, and 11 graduate students
who received Ph.D.’s in pharmacology, microbiology, biophysics, or
neuroscience. He also provided research training in his laboratory
to 28 undergraduate students. From 1990 to 2024, he taught the
General Principles of Pharmacology to over 2,400 first- and second-
year medical students.
1
Chapter 1
Introduction
M
ost drugs used as medicines in the body act by modifying
signaling systems. As examples, neurotransmitters and
hormones act on their receptors to control physiological
processes. This book describes the principles by which these signaling
systems control processes and how they can be influenced by drugs.
Pharmacology is one of the oldest sciences. It is also one of the
newest. Attempts to treat disease through the use of therapeutic sub-
stances can be traced to the earliest recorded human history. However,
the formal discipline of pharmacology was not recognized in medi-
cal schools until the beginning of the twentieth century. Today it is
a science that bridges and incorporates many traditional disciplines,
including physiology, biochemistry, cell biology, and microbiology. Its
most practical goal is to understand and develop therapeutic agents to
combat disease and promote human or animal welfare. Initial phar-
macological discoveries and advances, dating to the earliest times,
were largely empirical in nature. Certain substances were found to
be useful medicines, although their true molecular and physiological