Advances in Comparative and Environmental Physiology 16 Editor-in-Chief R. Gilles, Liege Editorial Board P. J. Butler, Birmingham R. Greger, Freiburg Ch. P. Mangum, Williamsburg G. N. Somero, Corvallis K. Thkahashi, Tokyo R. E. Weber, Aarhus Volumes already published Vol. 4: Animal Adaptation to Cold Edited by L. C. H. Wang (1989) Vol. 5: From the Contents: Osmoregulation - Chemoreception - Temperature - Spectroscopy - Metalloproteins Prostaglandins (Eicosanoids) - Maternal-Fetal Relationships (1989) Vol. 6: Vertebrate Gas Exchange: From Environment to Cell Edited by R. G. Boutilier (1990) Vol. 7: From the Contents: Protein Turnover - Xenobiotic Metabolism Lipoproteins (1991) Vol. 8: From the Contents: Dormancy in Aquatic Invertebrates - Root Effect - Red Blood Cell Functions in Fish - Diving Mammals (1991) Vol. 9: Volume and Osmolality Control in Animal Cells Edited by R. Gilles, E.K. Hoffmann, and L. Bolis (1991) Vol. 10: Comparative Aspects of Mechanoreceptor Systems Edited by F. Ito (1992) Vol. 11: Mechanics of Animal Locomotion Edited by R. MeN. Alexander (1992) Vol. 12: Muscle Contraction and Cell Motility: Molecular and Cellular Aspects Edited by H. Sugi (1992) Vol. 13: Blood and Tissue Oxygen Carriers Edited by Ch. P. Mangum (1992) Vol. 14: Interaction of Cell Volume and Cell Function Edited by F. Lang and D. Hiiussinger (1993) Vol. 15: From the Contents: Salivary Glands - Nematocyst Discharge - Crustacean Physiology - Steroid-Sensitive Areas (1993) Vol. 16: Ion Transport in Vertebrate Colon Edited by W. Clauss (1993) Volumes in preparation Vol. 17: Effects of High Pressure on Biological System Edited by A. G. Macdonald (1993) Vol. 18: Biomechanics of Feeding in Vertebrates Edited by V. L. Bels, M. Chardon and P. Vandewalle (1993) Advances in Comparative and Environmental Physiology 16 Ion Transport in Vertebrate Colon Guest Editor: W. Clauss With Contributions by Martine Avella · Kim E. Barrett · H. J. Binder · W. Clauss D. C. Dawson · W. von Engelhardt · R. Greger · J. P. Hayslett R. Krattenmacher · K. Kunzelmann · B. Lahlou · M. A. Post V. M. Rajendran · G. Rechkemmer · G. 1. Sandle E. Skadhauge · M. Tilman With 73 Figures Springer-V erlag Berlin Heidelberg GmbH Guest Editor: Professor Dr. WOLFGANG CLAUSS Institut fiir Tierphysiologie Justus-Liebig-Universitat Wartweg 95 W-6300 Giessen, Germany ISSN 0938-2673 ISBN 978-3-642-77120-0 ISBN 978-3-642-77118-7 (eBook) DOI 10.1007/978-3-642-77118-7 This work is subject to copyright. AII rights are reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broad casting, reproduction on microfilm or in any other way, and storage in data banks. Duplication of this publication or parts thereof is permitted only under the provisions of the German Copyright Law of September 9, 1965, in its current version, and permission for use must always be obtained from Springer-Verlag. Violations are liable for prosecution under the German Copyright Law. © Springer-Verlag Berlin Heidelberg 1993 Softcover reprint of the hardcover 1st edition 1993 The use of general descriptive names, registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant pro tective laws and regulations and therefore free for general use. Product liability: The publishers cannot guarantee the accuracy of any information about dosage and application contained in this book. In every individual case the user must check such information by consulting the relevant literature. lYPesetting: Best-set 'JYpesetter Ltd., Hong Kong 31/3145-5 4 3 2 1 O - Printed on acid-free paper Foreword to the Series The aim of the series is to provide comprehensive, integrated reviews giving sound, critical, and provocative summaries of our present knowledge in environmental and comparative physiology, from the mo lecular to the organismic level. Living organisms have evolved a widespread range of basic solu tions to cope with the different problems, both organismal and envi ronmental, with which they are faced. A clear understanding of these solutions is of course of fundamental interest for all biologists, zoolog ically or medically oriented. They can be best comprehended in the framework of the environmental and/or comparative approaches. These approaches demand either wide surveys of animal forms or a knowledge of the specific adaptive features of the species considered. This diversity of requirements, both at the conceptual and techno log icallevel, together with the fact that physiology and biochemistry have long been mainly devoted to the service of medicine, can account for the fact that these approaches emerged only slowly amongst the other new, more rapidly growing disciplines of the biological sciences. The field has now gained the international status it deserves and the organization of a series devoted to it appeared timely to me in view of its actual rapid development and of the interest it arouses for a growing number of biologists, physiologists, and biochemists, independently of their basic, major orientation. Liege, Belgium, Spring 1988 Raymond Gilles List of Editors Editor-in-Chief R. Gilles Laboratory of Animal Physiology University of Liege 22, Quai Van Beneden 4020 Liege, Belgium Editorial Board P. J. Butler G.N. Somera School of Biological Sciences Department of Zoology University of Birmingham Oregon State University P.O. Box 363 Corvallis, OR 97331-2914, USA Birmingham B 15 2TT, England K. Takahashi R. Greger Zoological Institute Physiologisches Institut Faculty of Sciences der UniversiHit Freiburg University of Tokyo Lehrstuhl II Hongo, Tokyo 113, Japan Hermann-Herder-Strasse 7 7800 Freiburg, FRG R.E. Weber Ch. P. Mangum Zoophysiology Laboratory Department of Biology University of Aarhus The College of William & Mary Building 131 Williamsburg, VA 23185, USA 8000 Aarhus C, Denmark Those interested in contributing to this series should contact Professor R. Gilles or one of the Series Editors. Review proposals should include a short outline - and a Table of Contents - briefly defining the aims and scope of the review. Preface The absorptive and secretory capacities of the large intestine are of special importance in the refinement and termination of the digestive processes. For a long time colonic properties were poorly investigated, as the main interest focussed on the major exchange site of ions and nutrients, the small intestine. In textbooks colonic properties were usually dealt with in a small section, noting that the colon is the site of water and electrolyte conservation. Whereas the overall absorptive and secretory features of the large intestine had been studied in vivo with perfusions or instillation tech niques, details about the mechanisms of ion exchange have only been worked out since Us sing laid the theoretical background with his flux ratio equation and double membrane model, and pioneered in vitro studies under controlled conditions with his famous chamber and short-circuit technique. Over the following years many of the new tech niques in cellular physiology, such as intracellular microelectrodes, noise analysis, patch clamping, membrane vesicles, and reconstitution techniques, have also been applied to colonic epithelia and isolated enterocytes of various vertebrates and have yielded interesting results. At first, researchers viewed the colon as a tube, and their interest focussed mainly on mammalian colonic epithelia (e.g. rat and rabbit). Later, they recognized segmental and diurnal differences, and func tional properties similar to the organization of ion transport in kidney tubules. Comparative physiologists noted that the colon does not serve exclusively digestive functions, but is in some vertebrates involved in osmoregulation, and even in urine concentration. The regulation of colonic ion transport was investigated, and was found regulated not only be enteric gut hormones, but by the classical target hormones of the kidney such as adrenal steroids and antidiuretic hormones. Many of these fine studies have led to models and conclusions of general interest. Therefore, the time seems ripe to collect some of this information, and to present an integrated view of ion transport in the vertebrate large intestine. Research on the colon not only serves medical interest or clinical purposes, but is a highly interesting subject of comparative physiology, from which we can learn much about the basic principles in physiology. The present book presents examples of research on col- VIII Preface onic ion transport of each vertebrate group, not only the classical models of epithelia, rat and rabbit, but also on the special and unique features in fish, amphibian, reptiles, birds and selected mammals such as the guinea pig. Physiology and pathophysiology of ion transport in the human large intestine are as well covered as the use of colonic cell cultures as model systems for crypt cell properties. I am especially grateful to the authors, each a specialist and leader in their field, who have contributed to this comparative approach to colonic ion transport, and hope that these interesting contributions will stimulate further research in this area. Giessen, February 1993 WOLFGANG CLAUSS Contents Chapter 1 Electrolyte Transport Mechanisms in Fish Intestine B. Lahlou and Martine Avella 1 Introduction ......................................... . 1 2 Morphology ......................................... . 2 3 Solute-Linked Water Transport ......................... . 5 4 Ion Transport ........................................ . 7 5 Control of Ion Transport by Second Messenger Systems ... . 15 6 Sodium-Linked Transport of Organic Solutes ............. . 18 7 Conclusion .......................................... . 20 References 20 Chapter 2 Mechanisms of Electrogenic Sodium Transport in the Amphibian Colon R. Krattenmacher and W. Clauss 1 Introduction ......................................... . 27 2 The Zoological System of Amphibians .................. . 28 3 The Function of the Amphibian Colon .................. . 29 4 Electrogenic Sodium Absorption ........................ . 30 5 Conclusions .......................................... . 43 References 43 Chapter 3 Ion Transport by Turtle Colon: A Role for Volume-Sensing Transporters in the Basolateral Membrane D. C. Dawson and M. A. Post 1 "Throughput": The Other Problem for Epithelial Cells 49 2 Functional Signatures and the Importance of Polarity ...... 50 3 Basolateral K + Channels Activated by Cell Swelling ........ 53 4 Basolateral NaiR Exchangers Activated by Cell Shrinking .. 56 5 Models for NaiR Exchange. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 61 6 Carrier to Channel and Back Again ..................... 63