Mechanisms of Systemic Regulation: Acid—Base Regulation, Ion-Transfer and Metabolism

275 Pages·1995·4.73 MB·English

by  D. E. Atkinson

Advances in Comparative and Environmental Physiology 22
Editor-in-Chief
R Gilles, Liege
Editorial Board
P.J. Butler, Birmingham
R Greger, Freiburg
Ch. P. Mangum, Williamsburg
G.N. Somero, Corvallis
K. Takahashi, Tokyo
RE. Weber, Aarhus
Volumes already published
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 (1993)
Vol. 14: Interaction of Cell Volume and Cell Function
Edited by F. Lang and D. Hiiussinger (1993)
Vol. 15: From the Contents: Salivary Gland Secretion
Nematocyst Discharge-
Adaptations in Decapodan Crustaceans -
Steroid-Sensitive Areas Mediating Reproductive
Behaviors (1993)
Vol. 16: Ion Transport in Vertebrate Colon
Edited by W. Clauss (1993)
Vol. 17: Effects of High Pressure on Biological Systems
Edited by A.G. Macdonald (1993)
Vol. 18: Biomechanics of Feeding in Vertebrates
Edited by V.L. Bels, M. Chardon and P. Vandewalle (1994)
Vol. 19: Electrogenic Cl- Transporters in Biological Membranes
Edited by G.A. Gerencser (1994)
Vol. 20: From the Contents: Motile Activities of Fish
Chromatophores - Epithelial Transport of Heavy Metals
Heavy Metal Cytotoxicity in Marine Organisms
Comparative Pulmonary Morphology and Morphometry
Molecular Adaptations in Resistance to Penicillins (1994)
Vol. 21: Mechanisms of Systemic Regulation:
Respiration and Circulation
Edited by N. Heisler (1995)
Vol. 22: Mechanisms of Systemic Regulation:
Acid-Base Regulation, Ion Transfer and Metabolism
Edited by N. Heisler (1995)
Volumes in preparation
Comparative Immunology: Invertebrate Immune Responses.
Vol. 23: Cells and Molecular Products.
Vol. 24: Cell Activities and the Environment.
Edited by E.L. Cooper
Advances in
Comparative
and Environmental
Physiology 22
Mechanisms of Systemic Regulation:
Acid-Base Regulation, Ion-Transfer
and Metabolism
Guest Editor: N. Heisler
With Contributions by
D.E. Atkinson . E. Bourke . W.H. Dantzler . D.H. Evans
M.K. Grieshaber· B.J. Harvey. N. Heisler· P. Laurent
S.F. Perry. H.O. Portner· T.J. Shuttleworth· V. Urbach
s.c.
S. Volkel . Wood
With 67 Figures
Springer
Guest Editor:
Prof. Dr. NORBERT HEISLER
Lehrstuhl fur Tierphysiologie
Math. Naturwiss. Fak. I
Humboldt UniversiHit zu Berlin
Abderhaldenhaus
Philippstr. 13
10 115 Berlin
Germany
ISSN 0938-2673
ISBN 978-3-642-52365-6 ISBN 978-3-642-52363-2 (eBook)
DOI 10.1007/978-3-642-52363-2
CIP data applied for
Die Deutsche Bibliothek - CIP-Einheitsaufnahme
Mechanisms of systemic regulation/guest ed.: Heisler.
Berlin; Heidelberg; New York: Springer
NE: Heisler. Norbert [Hrsg.J
Acid base regulation, ion transfer and metabolism/with
contributions by D. E. Atkinson ... -1995
(Advances in comparative and environmental physiology; 22)
NE: Atkinson, D. E.; GT
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Foreword to the Series
The aim of the series is to provide comprehensive, integrated reviews
giving sound, critical and provocative summaries of our present knowl
edge in environmental and comparative physiology, from the molecular
to the organismic level.
Living organisms have evolved a widespread range of basic solu
tions to cope with the different problems, both organismal and environ
mental, with which they are faced. A clear understanding of these
solutions is of course of fundamental interest for all biologists, zoo
logically or medically oriented. They can be best comprehended in the
framework ofthe 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 technological level, 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 slow 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.
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. Somero
School of Biological Sciences
Department of Zoology
University of Birmingham
Oregon State University
P.O. Box 363
Corvallis, OR 97331-2914, USA
Birmingham B15 2TT, England
R. Greger K. Takahashi
Physiologisches Institut Zoological Institute
der UniversiHit Freiburg Faculty of Sciences
Lehrstuhl II University of Tokyo
Hermann-Herder-Strasse 7 Hongo, Tokyo 113, Japan
79104 Freiburg, Germany
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
Preface
Various endogenous and environmental challenges of homoiostasis
have resulted in the evolution of apparently quite different mechanisms
for the same or similar functions in individual representatives of the
animal kingdom. One of the prominent achievements of comparative
physiology over the last few decades has been the description of regula
tory features common to many studied species beyond the extreme
diversity of their morphological forms. Delineation offunctional princi
ples universally applicable to the physiology and biochemistry of living
systems became often possible through technical advances in the devel
opment of numerous new techniques, in many cases modified and
adopted from other fields of science, but also by approaching certain
problems using multifactorial analysis. The advance in technology has
facilitated studies of minute functional details of mechanisms, which
finally lead to better understanding of generally similar functions,
covered by the multiple developments of Nature as a response to an
extreme variety of different conditions. Improved understanding of
specific mechanisms, however, has presented new problems at the level of
system integration. The importance of the integrative aspect became
particularly apparent during an international symposium on 'Mecha
nisms of Systemic Regulation in Lower Vertebrates: Respiration, Circu
lation, Ion Transfer and Metabolism' (organized in 1990 by Norbert
Heisler and Johannes Piiper at the Max-Planck-Institut fUr experimen
telle Medizin at Gottingen/Germany). Interaction, discussion and scien
tific cooperation among participants of the symposium initiated further
progress in the understanding of the role of specific mechanisms in the
light of organismic function, but at the same time indicated still largely
fragmentary knowledge and existing deficits. The resulting progress in
understanding homoiostatic control prompted the idea of publishing
a number of concise reviews on selected aspects of this scientific field,
which are presented in volumes 21 and 22 of this book series.
These reviews do not attempt to present encyclopedic completeness,
but are intended to selectively delineate recent progress in some fields of
comparative animal physiology, qualified by new approaches, interest
ing new data, novel interpretation or particular importance in terms of
integrative merits. The large weight of the last point is reflected in the
VIII Preface
title of the two volumes. The general aim is to provide a synthesis of
established ideas, models and the state of knowledge with new (some
times controversial) ideas, approaches, results and interpretations.
The various chapters present aspects of cellular mechanisms and
generalized functions related to import and export of substance, intra
corporeal transfer and maintenance of the overall status of animals.
Ammonia is the energetically most efficient nitrogenous metabolic end
product, but there are problems of toxicity and effects on acid-base
status. Metabolic elimination of ammonia by the synthesis of urea is one
way to deal with these problems. The interpretation of the regulatory
logic of the involved metabolic processes and of the interaction with the
acid-base status is still controversial (Chaps. 1 and 3). Elimination of
urate as another less toxic and acid-base relevant nitrogenous waste and
of other organic acids in the kidney in general is another mechanism to
reduce metabolic effects on homoiostatic control (Chap. 2). Direct elim
ination of ammonia to the environment is limited to aquatic species
because of its extreme toxicity. The form in which ammonia is transfer
red through membranes and epithelia has been the subject of intense
study and discussion over the last few years, with recent data clearly
suggesting transfer by nonionic diffusion (Chap. 4). Acid-base and ionic
regulation in fish takes place mainly at the branchial epithelium, the
morphological structure of which may be modified considerably in
response to various external and internal homoiostatic conditions
(Chap. 5). Natriuretic peptide hormones, well known from mammals,
are also found in heart muscle, brain and plasma of various fish groups.
Changes in activity of natriuretic peptide hormones initiate osmoregula
tory and haemodynamic effects in fish, which may be closely related to
systemic regulation (Chap. 6). Changes in intracellular pH regulate Na +
and K + channel activity, indirectly affect the Na + /K + pump in the
principal cells of high resistance epithelia and will also affect the activity
of the proton pump of mitochondria-rich cells in order to reestablish
equilibrium after acid loading (Chap. 7). Extensive studies on nasal
gland cells, particularly involved in the osmoregulation of marine birds,
have provided new information with respect to the character of intracel
lular signals controlling ionic and acid-base regulation (Chap. 8). The
amount of oxygen available for metabolic energy production is limited
in hypoxia. Recent studies indicate that in both ectotherms and en
dotherms an alternative and energetically advantageous approach to
increasing oxygen supply under such conditions is the reduction of
metabolic demand by behaviorally decreasing body temperature
(Chap. 9). In spite of the extreme cellular toxicity of hydrogen sulfide
numerous invertebrate species as well as a few vertebrates have develop
ed various types of effective internal and external defense mechanisms in
order to cope with hydrogen sulfide as an environmental challenge
(Chap. 10).
Preface IX
All of the chapters have been prepared by authorities in the respective
fields and as such represent the state of the art of our present knowledge,
sometimes spiced with controversial aspects delineated from various
sides. With this high standard of the individual contributions, it is still
hoped that this collection of reviews represents more than the sum of the
parts. The volume is designed to attract readers from a variety of
disciplines in biology and medicine, in addition to specialized fields in
physiology, biochemistry and biophysics, providing solid information
for established scientists as well as for students and postgraduates.
The editor would like to express his gratitude to the authors, who have
provided an excellent selection of chapters and have made a great effort
to make this volume comprehensible and wide ranging in a limited
amount of print space. Numerous other people involved in the prepara
tion of the manuscript and in the printing process, in particular my
assistant Dr. Sabine Heinz and the editorial staff of Springer, in particu
lar Ursula Gramm and Theodora Krammer, facilitated the publication
process.
April 7, 1995 Norbert Heisler
Contents
Acid-Base Regulation and Ammonia Elimination
Chapter 1
pH Homeostasis in Terrestrial Vertebrates;
Ammonium Ion as a Proton Source
D.E. Atkinson and E. Bourke
1 Introduction ....................................... 3
2 Renal Glutamine Metabolism ......................... 4
3 Catabolism of Protein Generates Bicarbonate ............ 8
4 Disposal of Metabolic Bicarbonate . . . . . . . . . . . . . . . . . . . . . 10
5 Summary. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
References .......................................... 23
Chapter 2
Renal Transport of Organic Acids and Bases
in Non-Mammalian Vertebrates
WHo Dantzler
1 Introduction ....................................... 27
2 Organic Acids (or Anions) ........ . . . . . . . . . . . . . . . . . . . . 28
3 Organic Bases (or Cations) ........................... 41
References .......................................... 46
Chapter 3
pH Homeostasis in Terrestrial Vertebrates:
A Comparison of Traditional and New Concepts
H. 0. Portner
1 Introduction 51
2 Nutrients and Acid-Base Status ...................... . 52