Table Of ContentNucleus o f the
Solitary Tract
Edited by
I. Robin A. Barraco, Ph.D.
Department of Physiology
Wayne State University
School of Medicine
Detroit, Michigan
Boca Raton London New York
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PREFACE
The nucleus of the solitary tract (NTS) is a major integrative viscerosensory nucleus in the dorsal medulla;
it is characterized by a rich pool of neurochemically diverse neurons containing an extraordinarily vast
array of transmitters and modulators. The NTS serves as the first relay station for sensory signals
originating from viscera, including taste and other nutrient-related sensation and a spectrum of general
visceral sensations. The NTS may also respond to humoral factors in the blood and cerebrospinal fluid
via its reciprocal circuitry and subjacent location with the area postrema, a circumventricular organ
located in the fourth ventricle. This incoming visceral and humoral information is processed and
integrated in the nucleus with signals arriving from other regulatory regions of the brain involved in
cardiorespiratory control, nutrient homeostasis, and metabolic regulation. Efferent signals generated by
the intrinsic neuronal circuitry of the NTS are in turn either transmitted via its connections with autonomic
centers of the brainstem to appropriate viscera as integrated cardiorespiratory reflexes and homeostatic
visceral response patterns or transmitted rostrally for further integration at higher levels of the neuraxis
and the consequent production of complex visceroendocrine responses and nutrient-related behavioral
activities.
This volume provides a comprehensive, up-to-date summary of current information on the anatomy
and physiology of the NTS prepared by a distinguished group of internationally recognized leaders in the
field. The subject matter has been organized to cover the various areas in which this growing
multidisciplinary field has evolved. The broad topic areas include the functional morphology and
chemical neuroanatomy of the NTS; its central projections and functional neural connections; mecha-
nisms of synaptic processing and sensory integration by intrinsic neural circuits in the NTS; the role of
neurotransmitters and receptors in NTS-mediated mechanisms of cardiorespiratory control; and the
integrative role of the NTS in coordination of autonomic, visceroendocrine and viscerobehavioral
response patterns involved in nutrient homeostasis, ingestive behavior and metabolic regulation. Con-
tributors have provided chapters which are complete entities of themselves, providing the reader with a
concise and focused account of the relevant material including novel research trends and state-of-the-art
techniques used in brain research and potential directions for future research in the cellular, molecular and
behavioral neurosciences. Many chapters also provide useful illustrations and convenient, up-to-date
diagrams describing central neural pathways involved in autonomic control and the regulation of visceral
function.
This book is a detailed, informative volume which should serve as a standard reference work that will
be of value not only to established research scientists, but also beginning investigators, clinicians, and
students interested in this crucial area of the brain.
THE EDITOR
I. Robin A. Barraco, Ph.D., is Professor of Physiology in the School of Medicine at Wayne State
University, Detroit, Michigan. Dr. Barraco received his B.A. degree in 1966 from Georgetown Univer-
sity, Washington, D.C. and obtained his Ph.D. degree in Physiology from Wayne State University School
of Medicine in 1971. During his graduate training, Dr. Barraco received support as a Research Fellow
from the National Science Foundation and the DeVlieg Foundation. Dr. Barraco has been a member of
the faculty in the Department of Physiology at Wayne State University School of Medicine since 1972
and he has held the rank of Professor of Physiology at Wayne State University since 1987.
Dr. Barraco is a member of the American Physiological Society, Society for Neuroscience, American
Society for Neurochemistry, International Society for Neurochemistry, and the International Behavioral
Neuroscience Society.
Dr. Barraco is the recipient of research grants from the National Institutes of Health and the National
Science Foundation in addition to research training grants from the National Institute of Mental Health.
He also participates in a variety of scientific peer review activities including editorial service as journal
referee and by serving on Study Sections at the National Institutes of Health and Special Review Panels
at the National Science Foundation.
Dr. Barraco has presented over 40 invited lectures at national and international meetings and he has
published more than 100 research papers, review articles, and book chapters. His current research
interests are mainly concerned with the neurochemical and neuromolecular bases of both normal and
abnormal brain function with special focus on the physiologic integration of brainstem mechanisms
mediating both autonomic reflex response patterns and more complex homeostatic behaviors.
CONTRIBUTORS
Akira Adachi Ruben D. Bunag
Department of Physiology Department of Pharmacology
Okayama University Dental School University of Kansas Medical Center
Okayama, Japan Kansas City, Kansas
Luigi F. Agnati B. Bunnemann
Department of Human Physiology Department of Histology and Neurobiology
University of Modena Karolinska Institute
Modena, Italy Stockholm, Sweden
Michael C. Andresen Jean Champagnat
Department of Physiology Department de Biologie Fonctionelle du Neurone
School of Medicine Institut Alfred Fessard
Oregon Health Sciences University CNRS
Portland, Oregon Gif-sur-Yvette, France
M. Anwar
Julie Y.H. Chan
Department of Neurology and Neuroscience
Department of Medical Research
Division of Neurobiology
Veterans General Hospital
Cornell University Medical Center
Taipei, Taiwan, Republic of China
New York, New York
Samuel H.H. Chan
Karen L. Barnes
Institute of Pharmacology
Department of Neurosciences
National Yang-Ming Medical College
The Research Institute of the Cleveland
Taipei, Taiwan, Republic of China
Clinic Foundation
Cleveland, Ohio
John Ciriello
Department of Physiology
/. Robin A. Barraco
University of Western Ontario
Department of Physiology
Health Sciences Centre
Wayne State University
London, Ontario, Canada
School of Medicine
Detroit, Michigan
R. Covenas
Armand L. Bianchi Department of Histology and Neurobiology
Départment de Physiologie et Neurophysiologie Karolinska Institute
Laboratoire de Neurobiologie et Stockholm, Sweden
Neurophysiologie Fanctionelle
Faculté des Sciences et Techniques Saint-Jérôme Roger A.L. Dampney
CNRS Department of Physiology
Marseille, France University of Sydney
Sydney, Australia
Borje Bjelke
Department of Histology and Neurobiology John A. Drewe
Karolinska Institute Department of Molecular Physiology
Stockholm, Sweden and Biophysics
Baylor College of Medicine
Robert M. Bradley Houston, Texas
Department of Biologic and Materials Sciences
School of Dentistry Joseph C. Dunbar
Department of Physiology Department of Physiology
School of Medicine Wayne State University
The University of Michigan School of Medicine
Ann Arbor, Michigan Detroit, Michigan
Erhan Ergene Michael D. Hirsch
Department of Physiology National Institute of Mental Health
Wayne State University Neuroscience and Biobehavioral Science
School of Medicine Research Review Branch
Detroit, Michigan Rockville, Maryland
Robert B. Felder Susan L. Höchstenbach
Department of Internal Medicine Department of Physiology
College of Medicine University of Western Ontario
University of Iowa Health Sciences Centre
Iowa City, Iowa London, Ontario, Canada
Peter D. Feldman
Hiroshi Itoh
Department of Pharmacology and Experimental
Department of Pharmacology
Therapeutics
University of Kansas Medical Center
Louisiana State University Medical Center
Kansas City, Kansas
New Orleans, Louisiana
André Jean
Alastair V. Ferguson
Départment de Physiologie et Neurophysiologie
Department of Physiology
Laboratoire de Neurobiologie et
Queen s University
Neurophysiologie Fonctionnelle
Kingston, Ontario, Canada
Faculté des Sciences et Techniques Saint-Jérôme
CNRS
Edson Carlos Fraga da Silva
Marseille, France
Laboratories LIM
School of Medicine
Jean-Pierre Kessler
University of Sdo Paulo
Départment de Physiologie et Neurophysiologie
Säo Paulo, Brazil
Laboratoire de Neurobiologie et
Neurophysiologie Fonctionnelle
Kjell Fuxe
Faculté des Sciences et Techniques Saint-Jérôme
Department of Histology and Neurobiology
CNRS
Karolinska Institute
Marseille, France
Stockholm, Sweden
Barbara K. Giza Diana L. Kunze
Department of Molecular Physiology
Department of Psychology
University of Delaware and Biophysics
Newark, Delaware Baylor College of Medicine
Houston, Texas
Frank J. Gordon
Department of Pharmacology Raul Laguzzi
Emory University School of Medicine Neurobiologie Cellulaire et Fonctionnelle
Atlanta, Georgia INSERM
Faculté de Medicine Pitié-Salpètriêre
Paris, France
Laurent Grélot
Départment de Physiologie et Neurophysiologie
Faculté des Sciences et Techniques Saint-Jérôme James N. Livingston
CNRS Institute for Metabolic Diseases
Marseille, France Miles Research Center
West Haven, Connecticut
Naomi Shinomiya Hell
Laboratories LIM Vicki L. Lowes
Institute of Biomedical Sciences Department of Physiology
University of Säo Paulo Queen s University
Säo Paulo, Brazil Kingston, Ontario, Canada
Bruce E. Maley Alan Randich
Department of Anatomy Department of Psychology
University of Kentucky Medical Center University of Alabama
Lexington, Kentucky Birmingham, Alabama
Stephen T. Meller
Stefanie Roder
Department of Pharmacology
Department of Physiology
University of Iowa
University of Western Ontario
Iowa City, Iowa
Health Sciences Centre
London, Ontario, Canada
Steven W. Mifflin
Department of Pharmacology
The University of Texas Health Sciences Center David A. Ruggiero
at San Antonio Department of Neurology and Neuroscience
San Antonio, Texas Division of Neurobiology
Cornell University Medical Center
William R. Millington New York, New York
Division of Molecular Biology and
Biochemistry
Hreday N. Sapru
University of Missouri
Section of Neurological Surgery and Department
Kansas City, Missouri
of Pharmacology
TA, Milner New Jersey Medical School
Department of Neurobiology and Neuroscience Newark, New Jersey
Division of Neurobiology
Cornell University Medical Center
Thomas R, Scott
New York, New York
Department of Psychology
University of Delaware
E.P. Mtui
Newark, Delaware
Department of Neurology and Neuroscience
Division of Neurobiology
Cornell University Medical Center K. Michael Spyer
New York, New York Department of Physiology
Royal Free Hospital School of Medicine
JA, Narvaez London, England
Department of Physiology
Faculty of Medicine
Meera Srinivasan
University of Malaga
The Nobel Institute for Neurophysiology
Malaga, Spain
Karolinska Institute
Stockholm, Sweden
K. Otake
Department of Neurology and Neuroscience
Division of Neurobiology C. Streefland
Cornell University Medical Center Department of Animal Physiology
New York, New York University of Groningen
Groningen, The Netherlands
Doug H. Park
Department of Neurology and Neuroscience
Alan F, Sved
Division of Neurobiology
Department of Behavioral Neuroscience
Cornell University Medical Center
University of Pittsburgh
New York, New York
Pittsburgh, Pennsylvania
Virginia M. Pickel
Department of Neurology and Neuroscience William Talman
Division of Neurobiology Department of Neurology
Cornell University Medical Center University of Iowa and VAMC
New York, New York Iowa City, Iowa
Fabien Tell Mark C, Whitehead
Départment de Physiologie et Neurophysiologie Departments of Anatomy and Surgery
Faculté des Sciences et Techniques Saint-Jérôme School of Medicine
CNRS University of California at San Diego
Marseille, France La Jolla, California
Yuji Yamamoto
The Nobel Institute for Neurophysiology
Karolinska Institute
Gert J. Ter Horst
Stockholm, Sweden
Department of Biological Psychiatry
University of Groningen
Mingyong Yang
Groningen, The Netherlands
Department of Physiology
School of Medicine
Oregon Health Sciences University
Portland, Oregon
Cesar Timo-Iaria
Laboratories LIM Juergen W. Unger
School of Medicine Department of Anatomy
University of Säo Paulo University of Munich
Säo Paulo, Brazil Munich, Germany
ACKNOWLEDGMENTS
The Editor would like to express his deepest appreciation to the numerous authors who contributed to this
volume, many of whom have achieved more senior standing in this field. I am grateful for their prompt
and timely submission of manuscripts and for their kind cooperation and thoughtful professionalism
throughout the preparation of this volume. I would like to especially acknowledge my admiration of those
international contributors who have the praiseworthy ability of using a second language to write a
manuscript. I am grateful for their generous participation as contributors.
The Editor would also like to express his genuine thanks to the editorial staff at CRC Press, Inc. for
their cooperation, and outstanding professionalism in helping to prepare this volume. A special note of
thanks should go to Marsha Baker, Executive Editor, Cathy Walker, now Journals Editor, Helen Linna,
Project Editor, and especially to Monique Power, Managing Editor, Life Sciences.
Finally, I would like to acknowledge the love and support of my wife, Leslie, and my daughter, Nina,
to whom I would like to gratefully dedicate this volume.
