Table Of ContentThe development of the shapes of living organisms and their parts is a field of
science in which there are no generally accepted theoretical principles. What
form these principles are likely to take, when they emerge, is a subject in
which there is a wide gulf of disagreement between physical scientists and
experimental biologists.
This book contains both an extensive philosophical commentary on this
dichotomy in views and an exposition of the type of theory most favoured by
physical scientists. In this theory, living form is a manifestation of the dynam-
ics of chemical change and physical transport or other physics of spatial
communication. The reaction-diffusion theory, as initiated by Turing in 1952
and since elaborated by Prigogine and by Gierer and Meinhardt and others, is
discussed in detail at a level that requires a good knowledge of a first course in
calculus, but no more than that. In some respects this book takes up the theme
that "the things which we see in the cell are less important than the actions
which we recognize in the cell," which was a major theme of D'Arcy W.
Thompson's classic 1917 work On Growth and Form. The rapid growth of the
field of molecular biology has tended to overshadow the increase in our
understanding of the nature of these kinetic processes. This book seeks to
reawaken interest in dynamics in the hope that a better balance between the
importance of things and the importance of actions may gradually emerge in
the field of biology in the twenty-first century.
DEVELOPMENTAL AND CELL BIOLOGY SERIES 28
EDITORS
P. W. BARLOW D. BRAY P. B. GREEN J. M. W. SLACK
KINETIC THEORY OF LIVING PATTERN
Developmental and cell biology series
SERIES EDITORS
Dr P. W. Barlow, Long Ashton Research Station, Bristol
Dr D. Bray, King's College, London
Dr P. B. Green, Dept of Biology, Stanford University
Dr J. M. W. Slack, ICRF Developmental Biology Unit, Oxford
The aim of the series is to present relatively short critical accounts of areas of
developmental and cell biology where sufficient information has accumulated to
allow a considered distillation of the subject. The fine structure of the cells,
embryology, morphology, physiology, genetics, biochemistry and biophysics are
subjects within the scope of the series. The books are intended to interest and
instruct advanced undergraduates and graduate students and to make an important
contribution to teaching cell and developmental biology. At the same time, they
should be of value to biologists who, while not working directly in the area of a
particular volume's subject matter, wish to keep abreast of developments relative to
their particular interests.
B O O K S IN THE SERIES
R. Maksymowych Analysis of leaf development
L. Roberts Cytodijferentiation in plants: xylo gene sis as a model system
P. Sengel Morphogenesis of skin
A. McLaren Mammalian chimaeras
E. Roosen-Runge The process of spermatogenesis in animals
F. D'Amato Nuclear cytology in relation to development
P. Nieuwkoop & L. Sutasurya Primordial germ cells in the chordates
J. Vasiliev & I. Gelfand Neoplastic and normal cells in culture
R. Chaleff Genetics of higher plants
P. Nieuwkoop & L. Sutasurya Primordial germ cells in the invertebrates
K. Sauer The biology of Physarum
N. Le Douarin The neural crest
M. H. Kaufman Early mammalian development: parthenogenic studies
V. Y. Brodsky & I. V. Uryvaeva Genome multiplication in growth and development
P. Nieuwkoop, A. G. Johnen & B. Albers The epigenetic nature of early chor date
development
V. Raghavan Embryogenesis in angiosperms: a development and experimental study
C. J. Epstein The consequences of chromosome imbalance: principles, mechanisms, and
models
L. Saxen Organogenesis of the kidney
V. Raghaven Developmental biology of fern gametophytes
R. Maksymowych Analysis of growth and development in Xanthium
B. John Meiosis
J. Bard Morphogenesis: the cellular and molecular processes of developmental anatomy
R. Wall This side up: spatial determination in the early development of animals
T. Sachs Pattern formation in plant tissues
J. M. W. Slack From egg to embryo: regional specification in early develoment
A. I. Farbman Cell biology of olfaction
KINETIC THEORY
OF LIVING PATTERN
LIONEL G. HARRISON
University of British Columbia
CAMBRIDGE
UNIVERSITY PRESS
CAMBRIDGE UNIVERSITY PRESS
Cambridge, New York, Melbourne, Madrid, Cape Town, Singapore, Sao Paulo
Cambridge University Press
The Edinburgh Building, Cambridge CB2 2RU, UK
Published in the United States of America by Cambridge University Press, New York
www.cambridge.org
Information on this title: www.cambridge.org/9780521306911
© Cambridge University Press 1993
This publication is in copyright. Subject to statutory exception
and to the provisions of relevant collective licensing agreements,
no reproduction of any part may take place without
the written permission of Cambridge University Press.
First published 1993
This digitally printed first paperback version 2005
A catalogue record for this publication is available from the British Library
Library of Congress Cataloguing in Publication data
Harrison, Lionel G.
Kinetic theory of living pattern / Lionel G. Harrison.
p. cm. - (Developmental and cell biology series : 28)
Includes bibliographical references and index.
ISBN 0-521-30691-4
1. Biology—Philosophy. 2. Matter, Kinetic theory of.
3. Biology-Mathematical models. I. Title. II. Series.
QH331.H368 1992
574'.01-dc20 92-12578
CIP
ISBN-13 978-0-521-30691-1 hardback
ISBN-10 0-521-30691-4 hardback
ISBN-13 978-0-521-01991-0 paperback
ISBN-10 0-521-01991-5 paperback
To the memory of Joyce and Julian
Contents
Preface page xiii
Part I: Macroscopics without mathematics 1
1 Introduction 3
1.1 Philosophy: lumping, spliting, abstraction, and reality 3
1.1.1 Lumping and spliting 4
1.1.2 Abstraction is reality 6
1.1.3 Macroscopics and the structure of proceses 8
1.2 Strategies of research 10
1.2.1 On starting to build a bridge from both ends 10
1.2.2 Making the join: bridges versus brains 11
1.2.3 Visions of the invisible 16
1.2.4 Preconceptions and experimental programs 17
1.3 Mixing and unmixing: molecules versus large objects 21
1.3.1 Molecules and individual identity: classical
physical chemistry versus modern molecular
biology 21
1.3.2 Arrangements, aggregations, amplifiers, automata 24
1.3.3 The kinetic preconception 28
2 Morphogen: one word for at least two concepts 31
2.1 Type I morphogens: Wolpert positional signalers 33
2.2 Activators versus morphogens 37
2.3 Inhibitors, such as chalones, versus morphogens 46
2.4 Type I morphogens: Turing morphogen pairs 47
2.5 The two-morphogen interaction 49
2.6 Activation and inhibition versus activated and
inhibited regions 53
3 Pictorial reasoning in kinetic theory of patern and form 56
3.1 The fit of pattern to boundaries and the dynamics
of patern growth 58
3.1.1 The fit of patern to boundaries 59
3.1.2 The dynamics of patern growth 63
vii
viii Contents
3.1.3 The guests of Procrustes: responses to chopping
and stretching 65
3.2 Modes of cel division in plants 67
3.2.1 One dimension: filaments 67
3.2.2 Location of determinative events within the cell 72
3.2.3 Division sequence of the Azolla root meristem 74
3.3 Animal development: response to damage and grafting 80
3.3.1 Teratogenesis and nonlinearities in the
Gierer-Meinhardt model 80
3.3.2 "Firing" a persistent gradient: grafting
behaviour of Hydra 84
Structure, equilibrium, kinetics 89
4.1 Definitions of the categories 90
4.1.1 Structure 90
4.1.2 Equilibrium 91
4.1.3 Kinetics 92
4.1.4 Mechanochemistry 94
4.1.5 Semantics of the term "field theory,"
and electric fields 96
4.2 Developmental control of the shapes of crystals 101
4.2.1 Structural aspect: just a few symmetry elements 101
4.2.2 Equilibrium shapes: surface free energy
and WulfFs theorem 102
4.2.3 Kinetic aspect: the diverse shapes of snow crystals 104
4.2.4 Biological control: the echinoid spicule 106
4.3 Division of plant cells: Is control kinetic,
thermodynamic, or mechanical? 108
4.4 Animal morphogenesis: rearrangement, deformation, and
proliferation of cels 110
4.4.1 Some phenomena, and the cell-as-molecule
concept 10
4.4.2 Equilibrium aspects: differential adhesion 118
4.4.3 From equilibrium to kinetics: incompleteness of
adhesive-gradient theory 123
4.4.4 Kinetic aspects involving mechanical forces 129
4.5 A few problems (without solutions) 137
Part II: Pattern-forming processes 143
5 The making and breaking of symmetry 145
5.1 Symmetry is in the spline of the beholder 145
5.2 Open and closed traverses: the accuracy of
self-organization 149
