Table Of ContentTYPE-B
CYTOCHROMES:
Sensors and Switches
TYPE-B
CYTOCHROMES:
Sensors and Switches
Johnathan L. Kiel
United States Air Force
Air Force Materiel Command
Human Systems Center
Armstrong Laboratory
Radiofrequency Radiation Division
Biotechnology Branch
Brooks Air Force Base, Texas
CRC Press
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Library of Congress Cataloging-in-Publication Data
Kiel, Johnathan L.
Type B cytochromes : sensors and switches / Johnathan L. Kiel.
p.(cid:9) cm.
Includes bibliographical references and index.
ISBN 0-8493-4576-6
1. Cytochrome b. 2. Cellular signal transduction.(cid:9) 3. Cell
metabolism. I. Title.
QP671 .C78K54 1995
574.87’61—dc20(cid:9) 94-36841
A Library of Congress record exists under LC control number: 94036841
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ISBN 13: 978-1-315-89831-5 (hbk)
ISBN 13: 978-1-351-07741-5 (ebk)
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PREFACE
When I proposed this book to CRC Press on July 22, 1991, the connections
among feeder cell and thiol support of lymphocytes in vitro, green hemopro-
teins of red blood cells and the liver, NADPH oxidase of granulocytes and
macrophages, nitric oxide synthase of leukocytes and neurons, and nitrogen
oxide reductases of plants and bacteria were very tenuous. The feeder cell
phenomenon was considered totally replaceable by exogenous simple
nonphysiologic thiols. This requirement was seen as an in vitro artifact result-
ing from nonphysiologic levels of oxygen and its toxic products in cell culture
media. Yet experiments with exogenous oxidase and peroxidase preparations
in vivo showed that energetic redox processes that did not generate ATP were
not the haphazard, nonspecific injurious processes seen in vitro. These nonspe-
cific oxidative killing mechanisms used against cancer cells and infectious
agents by granulocytes and macrophages seemed to be inexplicably linked to
development of a specific immune response. The discovery of the redox-
mediated effects of tumor necrosis factor (which were very similar if not
identical to those of bacterial lipopolysaccharide and gamma-interferon) sug-
gested a close redox cooperation between macrophages and lymphocytes
regulated by cytokines. In addition, the discovery of nitric oxide synthase
activity as essential to the physiologic responses of vasculature, the brain, and
insulin-producing beta cells of the pancreas indicated that this specialized
NADPH oxidase played roles well beyond the one of wholesale killing by
phagocytes.
In my letter to CRC Press, I stated that it was time to draw together the
information on electron transport chains and the central role and repeating
theme played in them by type-b cytochromes. I further stated that nitric oxide
synthase was at the same place NADPH oxidase was 12 years ago. I proposed
to show evidence that predicted a type-b cytochrome or similar hemoprotein
was required for the second messenger or neurotransmitter function of nitric
oxide synthase. On July 28, 1992, White and Marietta reported in Biochemistry
(31, 6627, 1992) that nitric oxide synthase was the first catalytically self-
sufficient mammalian cytochrome P450 (type-b cytochrome), containing both
reductase and heme domains on the same polypeptide. This discovery con-
nected the most important redox second messenger, cell-to-cell communicat-
ing enzyme with type-b cytochrome biochemistry. It mechanistically con-
nected nonlethal specific cellular responses to nonspecific redox agent effects.
Overall, this book attempts to portray type-b cytochromes as primordial pro-
teins that have played and continue to play a central role in the evolution and
functions of metabolism, cell growth, and survival in bacteria, plants, animals,
and man.
THE AUTHOR
Johnathan L. Kiel, D.V.M., Ph.D., received his B.S. degree in 1973 and
his D.V.M. in 1974 (both summa cum laude) from Texas A & M University.
He received his interdisciplinary Ph.D. in biochemistry and microbiology from
Texas Tech University Health Sciences Center, School of Medicine, in 1981.
He is a diplomate of the American College of Veterinary Microbiologists
(since 1984) and served on the National Examination Committee for the
College from 1988-1991.
Dr. Kiel is the Chief of the Biotechnology Branch, Radiofrequency Radia-
tion Division, Occupational and Environmental Health Directorate of the United
States Air Force Armstrong Laboratory. He is author of over 50 scientific
publications and holder of ten patents with four more pending. He received the
Otis 0. Benson Award for the Greatest Scientific Contribution to the U.S.A.F.
School of Aerospace Medicine in 1984 and 1989, the Air Force Systems
Command Science and Technology Award in 1991, the Research and Devel-
opment 100 Award for one of the most technologically significant new prod-
ucts of the year 1991 (the Quantitative Luminescence Imaging System), the
Harry G. Armstrong Scientific Excellence Award for outstanding research in
occupational and environmental health, and the Air Force Association Scientist
of the Year Award (Texas Branch) for 1994. He is the United States Air Force's
1994 recipient of the Basic Research Award for his research in mechanisms of
interaction of biological systems with nonionizing electromagnetic radiation.
CONTENTS
Chapter 1
General Properties of a Primordial Cellular Sensor and Switch (cid:9)1
Chapter 2
Primitive Type-B Cytochromes: Green Hemoproteins (cid:9) 13
Chapter 3
NADPH Oxidase and its Terminal Cytochrome (cid:9) 27
Chapter 4
Nitric Oxide Synthase (cid:9) 67
Chapter 5
Extracellular Disulfide Reductase (cid:9) 103
Chapter 6
Nitrogen Oxide Reductases (cid:9) 135
Chapter 7
Summary Schematics (cid:9) 155
Chapter 8
Future Research and Applications (cid:9) 167
References (cid:9) 173
Index (cid:9) 209
