Table Of ContentORDER IN LIVING ORGANISMS
Order in
Living Organisms
A Systems Analysis of Evolution
by
RUPERT RIEDL
Chairman of the Zoological Institute
University of Vienna, Austria
translated by
R. P. S. JEFFERIES
Department of Palaeontology
British Museum (Natural History)
A Wiley-Interscience Publication
JOHN WILEY & SONS
Chichester New York Brisbane Toronto
The original edition was published by Verlag Paul Parey, Hamburg and
Berlin, under the title Die Ordnung des Lebendigen © 1975 Verlag Paul
Parey, Hamburg and Berlin, who retain copyright of all illustrations in
this edition.
Copyright © 1978 by John Wiley & Sons Ltd.
Library of Congress Cataloging in Publication Data:
Riedl, Rupert.
Order in living organisms.
Translation of Die Ordnung des Lebendigen. Includes
bibliographical references and indexes. 1. Evolution.
2. Life (Biology). I. Title.
QH366.2.R5313 575 77-28245
ISBN 0 471 99635 1
Printed in Great Britain by The Pitman Press, Bath.
fsfW'VERSITY
OF BRISTOL
ZCC1.06Y
CONTENTS
PREFACE TO THE GERMAN EDITION.......................................................................................................... xi
PREFACE TO THE ENGLISH EDITION......................................................................................................... xv
INTRODUCTION......................................................................................................................................................... xvii
a. Accident and necessity.................................................................................................................... xvii
b. Decision and event........................................................................................................................... xviii
c. Mutation and selection.................................................................................................................... xviii
d. The system of phenomena............................................................................................................. xix
e. Cause and effect.................................................................................................................................. xix
f. Material and methods....................................................................................................................... xx
CHAPTER I WHAT IS ORDER?.................................................................................................................... 1
A Three Approaches to the Problems of Order..................................................................................... I
1. For and against order.............................................................................................................................. 1
a. Order as presupposition................................................................................................................. 1
b. Order and reality............................................................................................................................... 3
2. Entropy, n^entropy, order, and chaos............................................................................................. 3
3. Accident and necessity, certainty and uncertainty..................................................................... 4
B. Order as Probability..................................................................................................................................... 5
1. Indeterminacy and determinacy......................................................................................................... 5
a. Information content........................................................................................................................ 5
b. Predictability..................................................................................................................................... 6
c. Measuring the improbability of accident.................................................................................. 7
d. The probability of accident versus the probability of determinacy.............................. 7
e. The probability of determinacy ................................................................................................ 8
f. Specifying the determinacy content......................................................................................... 9
g. The limits of systems and methods............................................................................................. 10
2. Redundancy content and law content............................................................................................ 11
a. Redundancy content........................................................................................................................ 12
b. Useful and empty redundancy....................................................................................................... 12
c. Redundancy content and law content...................................................................................... 13
d. Visible and hidden redundancy................................................................................................... 14
e. The origin and fate of redundancy............................................................................................ 16
3. Order, determinacy, and negentropy ............................................................................................ 17
a. Entropy and negentropy................................................................................................................. 18
b. Certainty and uncertainty.............................................................................................................. 18
c. Information as entropy and negentropy.................................................................................. 20
4. Order as law times the number of instances................................................................................. 22
a. Solution of the information paradoxes...................................................................................... 23
b. Instance, decision, and event....................................................................................................... 24
VI
CHAPTER II THE DIMENSIONS AND FORMS OF LIVING ORDER........................................ 27
A. The Parameters of Biological Order......................................................................................................... 27
1. Order as energy...................................................................................................................................... 27
2. Order as improbability of state........................................................................................................ 28
3. Order as the extent of possible predictions................................................................................ 29
B. The Forms of Biological Order................................................................................................................ 30
1. The qualitative aspects of order........................................................................................................ 31
2. The building blocks : Identicality of individualities................................................................... 32
a. The seven forms of similarity..................................................................................................... 32
b. Homology and identicality............................................................................................................ 39
c. Individuality and law content..................................................................................................... 43
d. Homology and order...................................................................................................................... 45
e. The problem of degrees of similarity....................................................................................... 47
f. Freedom and necessity................................................................................................................... 47
3. The patterns of open questions : The identicality of regularities.......................... 48
a. The standard part............................................................................................................................. 49
b. Hierarchy........................................................................................................................................... 53
c. Interdependence................................................................................................................................ 55
d. Traditive inheritance...................................................................................................................... 56
4. The interconnections of the patterns ............................................................................................ 58
C. Biological Order as a Problem.................................................................................................................... 59
1. The controversy of complexity......................................................................................................... 60
2. The controversy of ‘internal causes’ ............................................................................................... 60
3. The controversy of essential structures........................................................................................... 62
4. The controversy about thought patterns........................................................................................ 63
CHAPTER III THE MOLECULAR CAUSE OF THE PATTERNS OF ORDER............................ 66
A. On Cause in General...................................................................................................................................... 66
a. The cause of the cause................................................................................................................... 66
b. The results of this cause................................................................................................................ 67
B. Determinative Decisions in the Organism........................................................................................... 68
1. The importance of the single decisions............................................................................................. 68
a. Adaptability - the designer plays dice ...................................................................................... 68
b. The holes in the punched tape - accidental programming................................................ 70
2. The advantage of dismantling redundant decisions..................................................................... 70
a. The principle of economy............................................................................................................. 70
b. Adaptability and redundancy...................................................................................................... 71
c. The necessity for systemization.................................................................................................. 72
C. The Systemization of Decisions.................................................................................................................. 73
1. The model and its molecular realization, Part I ............................................................................ 73
a. The repeat switch —‘repeat on demand’................................................................................. 73
b. The nucleic-acid systems................................................................................................................ 75
c. The selection switch - ‘applies until further notice’ ......................................................... 76
d. The operon system........................................................................................................................... 78
2. The first consequences of systemization.......................................................................................... 80
a. Burden and canalization................................................................................................................ 80
b. Freedom, determinacy, and superdeterminacy.................................................................... 82
c. The building-up and dismantling of decisions....................................................................... 83
3. The model and its molecular realization. Part II............................................................................ 84
a. Synchronous switching - ‘if N, then M’ ................................................................................. 84
b. The regulator-repressor system................................................................................................... 85
c. Sequential switching - ‘N only after A’ .................................................................................. 86
d. The order-on-order system.............................................................................................................. 88
Vll
D. Patterns of Systemization and Patterns of Features....................................................................... 88
1. Patterns of features are systemization patterns........................................................................... 89
a. Repeat switching, nucleic acids and the standard part..................................................... 89
b. Selector switching, operon and hierarchy............................................................................. 89
c. Synchronous switching, the regulator and interdependence........................................... 90
d. Sequential switching, order-on-order, traditive inheritance........................................... 91
2. Systemization patterns are patterns of features.......................................................................... 92
a. Copying of functional patterns by the epigenetic system.............................................. 92
b. Conservation of the original pattern....................................................................................... 93
CHAPTER IV THE STANDARD-PART PATTERN OF ORDER...................................................... 95
A. Introduction and Definition...................................................................................................................... 95
a. A fantasy world without standard-part order....................................................................... 95
b. Masses and classes.......................................................................................................................... 96
B. The Morphology of standard parts........................................................................................................ 97
1. Complexity, quantities and transformations.................................................................................. 97
a. The limits of identicality............................................................................................................... 97
b. Complexity and quantity............................................................................................................ 98
c. Origin and fate................................................................................................................................ 99
2. The placement of standard parts into systems............................................................................... 101
a. Positional standard parts and symmetrical standard parts................................................ 101
b. The substrate for single homologues........................................................................................ 101
c. Diversification of placement and function.............................................................................. 103
3. Burden, change and constancy.......................................................................................................... 104
a. The forms of burden...................................................................................................................... 104
b. The forms of freedom and of change........................................................................................ 105
c. The degrees of constancy and fixation.................................................................................... 106
C. Standard-part Selection............................................................................................................................. 107
1. The advantages of standardization.................................................................................................. 107
a. The prospects of success for blind accident.......................................................................... 107
b. The prospects of success for established facts....................................................................... 108
c. Economy and the increase of order........................................................................................... 109
d. Quick breakthroughs to new forms of organization............................................................. 110
2. Canalization and fixation.................................................................................................................... Ill
a. The prospects of successful alteration..................................................................................... Ill
b. Burden and selection....................................................................................................................... 112
c. Standard-part superselection......................................................................................................... 114
3. Norms and standards of civilization................................................................................................... 114
a. Success and mass-production...................................................................................................... 115
b. Tolerance and the collective......................................................................................................... 115
CHAPTER V THE HIERARCHICAL PATTERN OF ORDER.......................................................... 117
A. Introduction and Definition........................................................................................................................ 117
a. A fantasy world without hierarchy............................................................................................ 119
b. Preconditions and forms of hierarchy..................................................................................... 119
B. The Morphology of the Hierarchical Pattern......................................................................................... 125
1. Identification of general properties................................................................................................... 125
a. The rule for separating homologues............................................................................................ 125
b. The arrangement of systems.......................................................................................................... 125
c. Degree of complexity....................................................................................................................... 126
d. The degree of integration............................................................................................................. 126
e. Position within a system (and burden)..................................................................................... 127
f. Similarity............................................................................................................................................ 129
g. Age.......................................................................................................................................................... 130
VIU
h. Constancy (and fixation).......................................................................................................... 131
2. Extreme degrees of freedom and fixation.................................................................................... 132
a. Systems of maximal freedom.................................................................................................... 132
b. Systems of maximal fixation.................................................................................................... 135
3. The correlation of burden with fixation........................................................................................ 140
a. Quantitative characteristics....................................................................................................... 140
b. The qualitative characteristics associated with high and low burden.......................... 144
c. The pattern of distribution within the natural classification............................................ 145
d. Living fossils...................................................................................................................................... 145
e. Correlation of burden and constancy with the hierarchy of
representation within a phyletic group.................................................................................... 146
f. Correlation of burden and constancy with the hierarchy of position.......................... 147
4. The path towards burden and fixation........................................................................................ 151
a. The zero instant for a feature.................................................................................................... 152
b. An example of a fixation path.................................................................................................... 153
c. The principle of fixation............................................................................................................... 158
5. The rhythm of free and fixated phases........................................................................................ 158
a. A hyperbolic course for cladogenesis........................................................................................ 158
b. New freedom by means of new features................................................................................. 159
c. The hierarchy of the fixation patterns.................................................................................... 162
6. A summary and an anticipation......................................................................................................... 163
a. The reality of the hierarchical condition................................................................................ 163
b. The origin of hierarchy.................................................................................................................. 163
c. The causes.......................................................................................................................................... 163
C. The Selection of Ranks.............................................................................................................................. 164
1. Organization of events and decisions............................................................................................... 164
a. The degree of organization of determination complexes.................................................... 165
b. The gap in organization................................................................................................................ 165
c. The prospects of success................................................................................................................ 166
d. Three hypothetical solutions...................................................................................................... 167
e. The consequence of the organizational gap........................................................................... 168
2. The advantages of ranked decisions................................................................................................... 169
a. The origin of redundancy............................................................................................................. 169
b. The possibility of dismantling hidden redundancy............................................................. 169
0. The necessity of ranking................................................................................................................ 170
d. The imitative hierarchy of decisions........................................................................................ 170
3. Canalization and fixation.................................................................................................................... 171
a. The turning point.............................................................................................................................. 171
b. The effectiveness of superselection............................................................................................ 172
c. Counterselection and superselection........................................................................................ 173
d. Accident and necessity. ................................................................................................................. 173
4. The hierarchical pattern of civilization............................................................................................. 175
a. The success of ranking.................................................................................................................... 175
b. Qasses and tolerance........................................................................................................................ 176
CHAPTER VI THE INTERDEPENDENT PATTERN OF ORDER.................................................... 179
A. Introduction and Definition........................................................................................................................ 179
a. A fantasy world without interdependence.............................................................................. 180
b. The preconditions and forms of interdependence................................................................ 181
B. The Morphology of Interdependence....................................................................................................... 182
1. Interdependence in the phene system............................................................................................. 182
a. The recognition of interdependence........................................................................................ 182
b. The average degree of interdependence..................................................................................... 183
c. Synorganization or coadaptation................................................................................................ 183
d. The ubiquity of the interdependence phenomenon........................................................... 183
2. The organization of determination complexes............................................................................... 185
a. Pleiotropy and polygeny................................................................................................................ 185
b. Ho moeotic mutations.................................................................................................................... 187
c. All successful change is organized............................................................................................... 189
IX
3. The organization of the stream of determination...................................................................... 190
a. Phenocopies...................................................................................................................................... 190
b. Heteromorphosis............................................................................................................................ 190
c. Regeneration and propagation.................................................................................................... 192
C. Selection of Interdependence................................................................................................................... 194
1. The advantages of imitative interdependence............................................................................. 194
a. Narrowing the play of accident................................................................................................. 194
b. The imitation of phene patterns by gene patterns............................................................ 195
c. An imitative epigenotype........................................................................................................... 196
2. The canalization of interdependent patterns................................................................................. 197
a. Disruption and alteration........................................................................................................... 197
b. Burden and fixation..................................................................................................................... 197
c. Superselection and canalization................................................................................................. 198
3. Interdependence in civilization......................................................................................................... 199
a. Success and long-term success..................................................................................................... 200
b. Dependence and tolerance........................................................................................................... 200
CHAPTER VII THE TRADITIVE PATTERN OF ORDER.................................................................... 202
A. Introduction and Definition....................................................................................................................... 202
a. A fantasy world without traditive inheritance. ..................................................................... 202
b. Preconditions and forms................................................................................................................. 203
B. The Morphology of Traditive Patterns.................................................................................................. 204
1. The conservation of ancient patterns................................................................................................ 205
a. Spontaneous atavism....................................................................................................................... 205
b. The cryptotype — relict homoeostasis..................................................................................... 206
2. The general characteristics of ancient patterns............................................................................. 207
a. The topography of the determination complexes................................................................ 208
b. The functions of interphenes...................................................................................................... 209
c. The recapitulation of the determination patterns................................................................ 211
d. Decisions and phyletic relationships........................................................................................ 212
3. Freedom and fixation of interphenes................................................................................................ 214
a. The change from metaphene functions to interphene functions..................................... 214
b. Burden and fixation....................................................................................................................... 216
c. The freedom to simplify ontogeny........................................................................................... 218
4. Traditively inherited metaphenes...................................................................................................... 221
a. Atavism and the slowness of vestigialization........................................................................... 221
b. Adaptation and toleration............................................................................................................. 223
C. Traditive Selection........................................................................................................................................ 224
1. The advantages of traditively inherited data.................................................................................. 224
a. The necessity of traditive inheritance........................................................................................ 225
b. Traditive inheritance of past metaphene laws....................................................................... 225
c. Copies of old genotype patterns.................................................................................................. 226
d. The archigenotype.......................................................................................................................... 226
2. Canalization by traditively inherited organization.................................................................... 227
a. Functions, burden, and fixation................................................................................................... 227
b. The dimensions and direction of superselection.................................................................... 227
c. Counterselection and canalization............................................................................................ 228
3. Tradition in civilization........................................................................................................................ 230
a. The necessity of adopting old practice..................................................................................... 230
b. Tradition and tolerance................................................................................................................ 232
CHAPTER VIII THE THEORY AND ITS CONSEQUENCES................................................................. 235
A. A Theory of Systemic Conditions. . . . ; ............................................................................................ 235
1. The survivsJ prospects of molecular decisions.............................................................................. 236
a. Accident and necessity of decisions........................................................................................... 236
