The strategy of the genes : a discussion of some aspects of theoretical biology

275 Pages·2014·11.64 MB·English

by  C H Waddington

ROUTLEDGE LIBRARY EDITIONS:
20TH CENTURY SCIENCE
Volume 20
THE STRATEGY OF THE GENES
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THE STRATEGY OF THE GENES
A Discussion of Some Aspects of
Theoretical Biology
C. H. WADDINGTON
Firstpublishedin1957
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THE STRATEGY OF
THE GENES
A Discussi011 of Some Aspects of Theoretical Biology
BY
c.
H. WADDINGTON
SC.D., F.R.S.
WITH AN APPENDIX
BY
H. KAeSER
PH.D.
Ruskin House
GEORGE ALLEN & UNWIN LTD
MUSEUM STREET LONDON
fIRST PUBLISHED IN 1957
This book is copyright under the Berne Convention. Apart
from any fair dealing for the purposes of private study, research,
criticism or review, as permitted under the Copyright Act 1911,
no portion may be reproduced by any process withol4t written
permission. Enquiry should be made to the puhlisher
© George Allen & Unwin Ltd., 1957
PRINTED IN GREAT BRITAIN
in 12 point Bembo type
BY J. W. ARROWSMITH LTD.
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PREFACE
THE FIVE essays which form the body of this book, and the sixth
one which my friend Dr. Henrik Kacser has kindly allowed me
to include, are concerned with two of the major classical problems
of biology-firstly, the nature of biological organisation and the
developmental processes which bring it into being, and secondly,
the theory of evolution. These matters have been debated for
centuries. At the present time they tend, perhaps, to be somewhat
thrown into the shadow by the bright light which beats on the
growing points of our understanding of analytical biology. But
the search for the secret of gene action, for a comprehension of
how enzymes operate, or of the significance of nucleic acids and
proteins in determining specificity, although exciting and of
profound importance, nevertheless do not comprise the whole
aim of biology. One can adequately appreciate their relevance
only by reference to a framework of ideas sufficiently wide to
embrace all aspec'rs ofl iving systems. It is the purpose of Theoret
ical Biology-a young, and it must be confessed not as yet very
substantial branch of science-to provide such a conceptual
scheme. These essays are offered as a contribution towards the
scaffolding in one wing of the whole edifice:
Part of the fIrst essay is taken from a paper printed in the
Proceedings oft he Tenth International Congress ofP hilosophy ; the
second essay incorporates some paragraphs from an article which
appeared in Recent Developments in Cell Physiology (Butterworths,
1954); part of the third is from a paper published in Evolution;
of
Seventh Symposium of the Society Experimental Biology (Cam
bridge University Press 1953), and a few pages of the fifth are
quoted from an article published in Endeavour (Vol. 12, July 1953).
A few of the figures have been taken from published sources, to
which reference is made in the captions. I should like to thank
the editors and publishers concerned for permission to use this
material.
C.H.W.
v
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CONTENTS
Page
PREFACB V
I. FORM, END AND TIME I
Physical thing and biological organism. Latter charac
terised by Mind, Form and End, of which the first will
not be discussed. Organisation as the relation of parts to
wholes; but one must know according to what rule the
wholes are to be dissected into parts. A major character-
istic of life is that three time scales are relevant to it;
those ofe volution, development and physiological func
tioning. Form (wholeness, integration) and End (direc
tiveness, goal-seeking) appear extraordinary only be
cause we usually view them on the time scale of physi
ology, whereas they are appropriate to development and
evolution respectively, and are the major problems of
these two sciences.
2. THE CYBERNETICS OF DEVELOPMENT II
The three basic processes of development: regionalisa-
tion, histogenesis and morphogenesis. Competing
reaction systems with feed-backs. Canalised pathways,
which represent systems of simultaneous differential
equations, and can only be fully expressed in multi
dimensional phase space. The inherent trajectory of a
system, or 'creode'. The creodic profile; periodic
phenomena in development i recapitulation. 'Homeo
rhesis'; physiological and genetical homeostasis; feed
back, developmental flexibility. The canalisation cross
section and its genetical control. Developmental noise.
The tactical organisation of a developmental channel;
structured effectors; focal and peripheral processes;
sequential and concurrent syntheses.
3. SELECTION OF, FOR AND BY 59
Is the problem of evolution solved i Modes of selection.
Individual selection, of first and higher orders; progeny
selection; inter-population selection; metaselection.
vii