Table Of ContentCOLON
Structure and Function
TOPICS IN GASTROENTEROLOGY
Series Editor: Howard M. Spiro, M.D.
Yale University School ofM edicine
PANCREATITIS
Peter A. Banks, M.D.
MEDICAL ASPECTS OF DIETARY FIBER
Edited by Gene A. Spiller, Ph.D., and Ruth McPherson Kay, Ph.D.
NUTRITION AND DIET THERAPY
IN GASTROINTESTINAL DISEASE
Martin H. Floch, M.S., M.D., F.A.C.P.
COLON
Structure and Function
Edited by Luis Bustos-Fernandez, M.D.
A Continuation Order Plan is available for this series. A continuation order will bring delivery
of each new volume immediately upon publication. Volumes are billed only upon actual ship
ment. For further information please contact the publisher.
COLON
Structure and Function
Edited by
Luis Bustos-Fernandez, M. D.
Buenos Aires University School of Medicine
Buenos Aires, Argentina
PLENUM MEDICAL BOOK COMPANY
NEW YORK AND LONDON
Library of Congress Cataloging in Publication Data
Main entry under title:
Colon, structure and function.
(Topics in gastroenterology)
Includes bibliographical references and index.
I. Colon (Anatomy) I. Bustos-Fernandez, Luis. II. Series.
QPI56.C64 1982 612'.36 82-20413
ISBN 978-1-4757-0309-2 ISBN 978-1-4757-0307-8 (eBook)
DOl 10.1007/978-1-4757-0307-8
(c) 1983 Plenum Publishing Corporation
Softcover reprint of the hardcover 1s t edition 1983
233 Spring Street, New York, N.Y. 10013
Plenum Medical Book Company is an imprint of Plenum Publishing Corporation
All rights reserved
No part of this book may be reproduced, stored in a retrieval system, or transmitted
in any form or by any means, electronic, mechanical, photocopying, microfilming,
recording, or otherwise, without written permission from the Publisher
Contributors
Adrian Allen, Department of Physiological Sciences, The Medical School, The Uni
versity of Newcastle upon Tyne, Newcastle upon Tyne, England
C. Bernard, Instituto de Gastroenterolgia, "Dr. Jorge Perez Companc," Universidad
Cat6lica Argentina, Hospital Italiano, Buenos Aires, Argentina
Anne E. Bishop, Department of Histochemistry and Medicine, Royal Postgraduate
Medical School, Hammersmith Hospital, London, England
S. R. Bloom, Department of Histochemistry and Medicine, Royal Postgraduate Med
ical School, Hammersmith Hospital, London, England
Luis Bustos-Fernandez, Instituto de Gastroenterologia, "Dr. Jorge Perez Com
panc," Universidad Cat6lica Argentina, Hospital Italiano, Buenos Aires, Argen
tina
Ghislain Devroede, Departements de Chirurgie Generale et de Physiologie, Unite de
Recherche Gastrointestinale, Faculte de Medecine, Universite de Sherbrooke,
Sherbrooke, Quebec, Canada
Gregory L. Eastwood, Gastroenterology Division, Department of Medicine, Univer
sity of Massachusetts Medical School, Worcester, Massachusetts
M. A. Eastwood, Wolfson Gastrointestinal Laboratory, Gastrointestinal Unit, Depart
ment of Medicine, Western General Hospital, Edinburgh, Scotland
K. Ewe, I. Medizinische Klinik und Poliklinik, Universitat Mainz, Mainz, Federal
Republic of Germany
E. Gonzalez, Instituto de Gastroenterologia, "Dr. Jorge Perez Companc,"
Universidad Cat6lica Argentina, Hospital Italiano, Buenos Aires, Argentina
Sherwood L. Gorbach, Division of Infectious Diseases, The George Washington Uni
versity Medical Center, Washington, D.C., and Tufts-New England Medical
Center, Boston, Massachusetts
S. Hamamura, Instituto de Gastroenterologia, "Dr. Jorge Perez Companc,"
Universidad Cat6lica Argentina, Hospital Italiano, Buenos Aires, Argentina
M. J. Hill, Central Public Health Laboratory, London, England
D. P. Jewell, The John Radcliffe Hospital, Headington, Oxford, England
Mats Jodal, Department of Physiology, University of G6teborg, G6teborg, Sweden
Paul Kerlin, Mayo Foundation and Mayo Medical School, Rochester, Minnesota
v
vi CONTRIBUTORS
J. A. Kofoed, Instituto de Gastroenterologfa, "Dr. Jorge Perez Companc,"
Universidad Cat6lica Argentina, Hospital Italiano, Buenos Aires, Argentina
I. Ledesma-Paolo, Instituto de Gastroenterologfa, "Dr. Jorge Perez Companc,"
Universidad Cat6lica Argentina, Hospital Italiano, Buenos Aires, Argentina
Ove Lundgren, Department of Physiology, University of G6teberg, G6teborg,
Sweden
H. Menge, Abteilung flir Innere Medizin mit Schwerpunkt Gastroenterologie,
Klinikum Steglitz der Freien Universitat, Berlin, Federal Republic of Germany
K. Ogawa-Furuya, Instituto de Gastroenterologfa, "Dr. Jorge Perez Companc,"
Universidad Cat6lica Argentina, Hospital Italiano, Buenos Aires, Argentina
Sidney Phillips, Mayo Foundation and Mayo Medical School, Gastroenterology Unit,
Mayo Clinic, Rochester, Minnesota
Julia M. Polak, Department of Histochemistry and Medicine, Royal Postgraduate
Medical School, Hammersmith Hospital, London, England
J. A. Robertson, Wolfson Gastrointestinal Laboratory, Gastrointestinal Unit, Depart
ment of Medicine, Western General Hospital, Edinburgh, Scotland
J. W. L. Robinson, Departement de Chirurgie Experimentale, Centre Hospitalier,
Universitaire Vaudois, Lausanne, Switzerland
W. S. Selby, The John Radcliffe Hospital, Headington, Oxford, England
Gary L. Simon, Division of Infectious Diseases, The George Washington University
Medical Center, Washington, D.C., and Tufts-New England Medical Center,
Boston, Massachusetts
R. Wanitschke, I. Medizinische Klinik and Poliklinik, Universitat Mainz, Mainz,
Federal Republic of Germany
Preface
The functional and organic alterations of the colon constitute one of the leading reasons
why patients consult gastroenterologists. The irritable colon is one of the most com
mon causes of discomfort in human beings. The organic pathology of the large bowel
(malignancy and chronic inflammatory disease) contributes, particularly among Occi
dental peoples, to discouragingly high levels of morbidity and mortality.
One realizes the importance of having a thorough physiologic knowledge of the
colon in order to scientifically plan the functional treatment of organic colonic dis
eases.
If we consider the large amount of material published on the physiology of the
esophagus, stomach, small bowel, pancreas, and liver, we realize that the colon has
been relatively neglected. The chapters in this book have been written by people who
have done their utmost to alter this imbalance.
I want to thank all the contributors for their generous collaboration that allows me
to present in one volume virtually all the information known about the structure and
function of the colon, and to record my deep graditude to Dr. Howard Spiro for his
willingness to include this volume in his series. I would also like to express my sincere
appreciation to Plenum Publishing Corporation for making this book possible. A spe
cial thanks goes to Dr. Isable Ledesma de Paolo for her cooperation in selecting the
material to be published, to my daughter Marfa Angelica Bustos-Fernandez de
Aragone for her diversified help in the preparation of the work, and also to my secre
tary Marcela Claudia Alvarez for her correspondence with the authors, as well as her
diligent processing and retyping of some of the revised chapters.
My wife Susana was once more an important supporter of my work.
Luis Bustos-Fernandez
vii
Contents
1. Colon Structure
Gregory L. Eastwood
I. Development of the Colon. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
II. Macroanatomy............................................. 3
III. Microanatomy............................................. 5
A. The Four Layers of the Colon . . . . . . . . . . . . . . . . . . . . . . . . . . .. 5
B. Epithelial Renewal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 6
C. Epithelial Cell Types. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
IV. Conditions Which Affect Colonic Structure. . . . . . . . . . . . . . . . . . . .. 10
A. Food .................................................. 10
B. Bowel Resection. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13
C. Colonic Disease. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13
D. Irradiation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13
References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 14
2. Absorption and Secretion of Electrolytes by the Human Colon
Paul Kerlin and Sidney Phillips
I. Introduction............................................... 17
A. Aims and Overview ..................................... 17
B. Comparison of Man and Other Species. . . . . . . . . . . . . . . . . . . .. 18
C. Experimental Approaches to the Study of Colonic Absorption.. 18
II. Concept of Colonic Epithelium as a "Tight" Mucosa. . . . . . . . . . .. 20
A. Estimates of Pore Size. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 20
B. Permeability of the Colon to Water and Hydrophilic Solutes ... 20
C. Transepithelial Potential Difference ........................ 20
D. Regional Differences. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 21
ix
x CONTENTS
III. Absorption in Vitro ......................................... 21
A. Sodium Transport. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 21
B. Chloride............................................... 23
C. Bicarbonate............................................ 23
D. Potassium .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 24
E. Short-Chain Fatty Acids ................................. 24
IV. Secretion in Vitro . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 25
V. Absorption by the Colon in Vivo . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 26
A. Estimates of Ileal Input into the Colon . . . . . . . . . . . . . . . . . . . .. 26
B. Colonic Absorptive Capacity. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 27
C. Specifics of Electrolyte and Water Transport. . . . . . . . . . . . . . .. 28
VI. Intraluminal Metabolism and Absorption. . . . . . . . . . . . . . . . . . . . . .. 29
A. Dietary Fiber and Carbohydrates .......................... 29
B. Ammonia Generation and Absorption . . . . . . . . . . . . . . . . . . . . .. 31
VII. Factors Modifying Colonic Absorption: Secretion in Vivo. . . . . . . .. 31
A. Hormones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 31
B. Laxatives.............................................. 32
C. Bile Acids and Fatty Acids. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 33
VIII. Divalent Cations: Calcium, Magnesium, and Zinc . . . . . . . . . . . . . .. 34
IX. Colonic Absorption in Disease. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 34
A. Colonic Overload . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 34
B. Diarrhea after Ileal Resection . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 34
C. Congenital Chloride Diarrhea. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 34
D. Mucosal Disease. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 35
E. Villous Adenoma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 35
F. Constipation ........................................... 36
G. Enteric Hyperoxaluria ................................... 36
H. The Role of Fecal Electrolyte Measurements in
Clinical Medicine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 36
X. Summary and Future Directions .............................. 36
References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 37
3. The Structure of Colonic Mucus
Adrian Allen
I. Introduction............................................... 45
II. Isolation and Purification of Mucus Glycoproteins . . . . . . . . . . . . . .. 46
m. Carbohydrate Chain Structure in Mucus Glycoproteins ........... 50
IV. Protein Core and Polymeric Structure in Mucus Glycoproteins ..... 55
V. The Relationship between Glycoprotein Structure and the Gel
Forming Properties of Mucus Secretions . . . . . . . . . . . . . . . . . . . . . .. 57
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 60
