‘The question of the connection between law and chance in the
evolution of life is one of the cenwal philosophical problems in
In this book, fundamental aspects of this question have
n treated from the viewpoint of modern evolutionary biology.
The investigation starts from the assumption that the objectof its
analysis counts asa wellfounded part of natural science and does
not need to be specially justified. It presupposes that evolution is,
an established fact, and that the synthetic evolution theory built up
on the basis of Darwin's ideas provides an appropriate framework
for the understanding and explanation of evolutionary phenom-
ena (chapter 1)
However, the issue of the theoretical character of synthetic
evolution theory isa problem in the philosophy of science that
possesses a special significance with regard t the more gener:
question of the connection beoveen law and chance in evolution.
Access o this problems afforded in particular by the analysis of the
so-called molecular theory of evolution, which links the principles
of synthetic evolution theory with the fundamental laws of physics
and chemistry. At the same time, many aspects of this connection
throw new light upon the general problems of the formation of
concepts and theories in the border region of physics, chemistry,
and bioloj
‘The miolécular theory of evohution has been a result of rapid
progress in biology and physics in the last two decades. I is based.
on the one hand upon the discovery in molecular biology that all
basic phenomena of life such as metabolism and heredity can be
traced back to regular interactions between biological macr
and thereby to the laws of physies and chemistry, and on the
other hand upon the discovery in physics of open systems that, far
Summary — 169
from equilibrium, can spontaneously assemble states of material
order (so-called dissipative structures) that are also characteristic of
living systems,
The motecular theory of evolution describes the origin and the
early evolution of life as a process of material self organization, in
the process of which the two most important classes of biological
macromolecule (nucleic acids and proteins) were able to organize
themselves spontancously into a system, governed by information,
that possessed the three basic properties of living matter: metabo
lism, selfreproduction and mutability
The modern theory of the origin of life is a physicochemical
theory. As such, it attempts to base the historical process of
biological selforganization upon its fundamental, unchanging
principles and mechanisms. The theory does not claim to be able
to reconstruct the process in its historical details.
In the overall process of biological self organization, three phases,
each corresponding to the degree of optimization attained at that
time, can be distinguished (chapter 2):
1. the phase of noninstructed prebiotic synthesis of
macromolecules,
logical
2, the phase of selCorganization of biological macromolecules to 2
selfiinstructed biosynthetic cycle, and
3. the phase of evolutionary optimization of the biosynthetic cycle.
‘The actual transition from inert to living matter took place during
the second phase, and was essentially a transition from noninstra-
cted to instructed synthesis of biological macromolecules. T
phase of biological self organization ended with the nucleation of
a self'instructed biosynthetic cycle, which (after the formation of
compartments) can be regarded as a primitive precursor of the
ving cell
Every kind of instruction requires information, The information
used by the selfinstructed biosythetic cycle is encoded in the
detailed sequence of its nucleic acid components. These instruct
the construction of a protein machinery that in turn catalyzes the
reproduction of the entire cycle,
Ithas emerged that the information for the construction of a
iving organism is not simply encoded in the linear sequence of the
monomers of its DNA molecules, but that this linear arrange-
ment—as in human language—possesses a hicrarchical superSummary
1. the phase of the origin of syntactic information,
2, the phase of the origin of semantic information, and
nary optimization of semantic information.
8. the phase of evolu
The syuhevie aspect of the origin of biological information
Unproblemadc fom a philosopliel sandposnt, since it's eon.
cemed solely with the formation of sructires—for example, of
shacromolecules such as protein or nucleic acids Incontras, the
semantlcaspectraises many problems sinesinrlerstothe content,
that sine sense and mcening of the information encoded in
nactomolecalar struct Tine raises the queston of whether
thu iso, towhatextent Sense" and "meaning can be objecified
tind studied ciently
‘As the caample of human language show, semantics cannot
extn an absolute sense, but only relaive toa semanti frame of
feference (chapuer 4). Geneve information, oo, poseeses No
Stvolutestmnantc value, But only » elave one, relerred to the
Summary 171
specific environmental conditions to which the organism in ques:
Yon has become adapted. The environment thus represents
externally localized information, on the basis of which the seman.
des of genetic information are selectively evaluated,
According to Darwinian evolution theory, the purpose-directed,
or information-controlled, element of biological structures pos:
sesses.a certain function (and thus sense and meaning) with regard
to the preservation of the dynamic order that is characteristic of
living systems, Fhe semantics of genetic information can thus be
interpreted directly in terms of evolution; an organism is the better
adapted to its environment (that is, the semantic information it
contains has the greater value), the more accurately and rapidly
it—for a given lifespan—reproduces its information content.
‘This principle provides a link with Gvo philosophical theses of
‘arl Friedrich von Weizsicker, according to which the semantic
aspect of information can only be objectitied via the pragmatic
aspect of information (“Information is only that which is under
stood”; “Information is only that which produces information”)
This thought runs through the entire discussion like a leitmotiv. Tc
mplies that even the “protosemantics" of biological information
can be objectified by a dynamic criterion of the generation of
information (see below)
The question of whether and, if so, to what extent the origin of
biological information can be explained in terms of regularity
within the framework of modem science is taken up in part Hl
‘Three suggested solutions are presented: (a) the chance hypathe-
sis, in chapter 6, (b) the teleological approach, in chapter 7, and (c)
the molecular-Darwinistic approach, in chapter 8.
The chance hypothesis interprets the origin of biological infor-
ation as a singular event, whose description by the laws of physics
and chemistry can only take place on the syntactic level, that is, on
the level of chemical evolution. The semantic aspect of the origin
of biological information, in contrast, is understood in the chance:
hypothesis as an epiphenomenon, which by its very nature is
determined purely by chance and possesses none of the regular.
ties of natural law. The chance hypothesis is justified by the
assertion, at first seemingly contradictory, that within equilibeiurn
statistics the probability of the chance synthesis of an information.
carrying macromolecule is extremely low, because the informa-
Hon-carrying macromolecules represent only a minute fraction of
their physically equivalent alternative sequences, and all these172 Summary
alternasives have under equilibrivm condi
prior probabilies,Consequenthy, in the context of che chance
Pypothesis the origin of lifes interpreted posterior axa singular
reeebnn events which Iecause of is exstemely low prior probability
Inunique and unrepeatable inthe whole universe
Taancicological approach proceeds similarly Seom the explans-
ton deficit of equilibrum statistics, jst described, In contrast eo
Re chance hypothesis, which does Rot offer a causal explanation
Tes ie ongin otbiotogical informacion, the teleological approach
Rhotularerthe exisence of a goabdirecced and at the same tine
Tradable, lavike bebasion,ahich narvows down the immense
Hiologicaly relevant. In the teleological model of explanation,
rear icacanning biomolecules must be regarded as the mate:
tha expresion of a finale principe at work in Nature.’ The
Teleclegical approach and the chance hypothesis agree dhat the
seeSinSarrangement of the monomers of te DNA molecules na
seen cngading regularity and purposiveness, are fundamen
aby inexplicable t the lovels of physics and chemistry.
Ththe dibubon of roles beoseen lay andl chance the origin
of biological information, the molecularDerwinstic appro
Shope aa intermediate position. Ia the molecular-Darwinistic
tol, the origin of blogs formation is derived fom a
Terplay benveen undirected, random process (rmstation) an
reigabar betavior of matter (nasal selection), ‘The mathematical
funmutation of this approach i the molecular cheory of evolution
MQothed our atthe beginning. However, and here the moteculae-
Dasialise approach differs fandameptally from its rivals dis
Cred above, ihe mespay peorcenshuzon and sleton
Sutal in as easental way upon the principles of nonequilibrium
Thoinedyaamice, Further, the molecular-Darwinistic approach
se upon the working hypothesis that natural selection in the
Doan seze appears alveady in the realm of inanimate matter
“Phe veunsion in part Hl makes ie clear Ghat a philosophical
analjis of eustent hypotheses concerning the origin of biological
see Gon presapposes « formulation Of the concept of bv £0
reer that ven sten diverse approaches asthe chance hypothesis
Beth ihe tseglogical approach ean be selated to one another, s0°0
speak cna higher level of the formation of concepts and theories
Noun chasacteriatic feature of a satural law is that it contains
information about natural events in a compressed form. The
Summary 173
logical next step is to look for ageneral formulation of the concept
of law within the framework of information theory.
The information for the construction of a living organism is
encoded in the detailed sequence of the monomers in its DNA
molecules, This sequence ean be found out, usually by experimen-
tal (chemical) sequence determination. Like any other empirical
data in science, it can be represented by a finite row of ones and
zeros (chapter 10, figure 19).
The representation of observational data as binary sequences
leads directly to an information-theoretical interpretation of the
concept of natural law, Clearly, a lawlike relationship is always
contained in a series of observational data when there is a compact
algorithm with the help of which the observed data can be repre-
sented in compressed form. The central feature of such an
algorithm is the property that it can be represented (on the
syntactic level} by using less information than the informati
needed for the representation of the data sequences that it gener-
ates. Conversely, a sequence of observational data must be re-
garded as random until an appropriate compact algorithm has
been found. The term “random sequence” in this way acquires
a precise algorithmic definition (chapter 9). However, the irre-
ucibility, that is, the randomness, of a binary sequence is inher-
enily incapable of proof, since the nonexistence of a compact
algorithm capable of generating the sequence can never be proven
(che randomness theorem).
From the randomness theorem, the roots of which go back to
Godel’s incompleteness theorem, ovo fundamental limits of objec
tive knowledge in science can be deduced (chapter 10). One such
limit is due to dhe fundamental problems of biological discovery
mentioned in part II the chance hypothesis is fundamentally
incapable of proof, while the teleological approach is fundamen-
tally irrefutable. The other limitis related to the ability of natural
law to describe reality asa whole: itcan never be proved that agiven
algorithm is the smallest capable of describing a class of natural
events in a lawlike way, since the smallest algorithm (in binary
representation) is by definition a random sequence, whose irre-
ducibility—because of the randomness theorem—cannotbe proved.
‘This is in particular true of the case when an algorithm is given for
all classes of natural event. In other words, the completeness of a
scientific theory, as would be expressed information-theoretically
in the statement of a unique and irreducible algorithm, is funda-74 Susmery
mentally incapable of proof, It is only efutable by pragmatic
eee a eae memento a nev and more compact aigovh
Fae ine sane rreatine conceptofmatural law intresuced on
eee ePestauon ean wal Rave lmplieadoasforastumber
aera ei enms in the philosophy of scence. The supposition
Cae ree acca in chapter 10 that chere may oe an
a ere ection bemveen the algorithm detinion ofa water
serie Cte iden developed by CF won Welasicherof so-called
se itomatives, as outined in ote 168.
ae aernattiedon to the chanes hypothesis and to the cle
cgi sporoach, the molecular-Darwhiste approach remains
Chukely within the confines of taditional scienuie explanatory
See However the molecular Darwinioucapproach ison valid
Fee reer selecaon really does tae ploce inthe Fea
ean matter at's comequencr of pure physicochemical
Seguin (ee abowe)
eT Scalar Darwinistic approach raises paradigmatically the
quetion of the physial foundation of biciogy tie scaled
ween of reduction). Consequently te reduction ise a also
aeoetee ef pein inthe daruston of the posites and limits:
see Othe olccular arvinisic explanatory model, vis consid
wee RSer 1 Gestaon philosophical problem. The election
cine eel how oibg phystallyjustfable (chapter 12). In
aa ae ana recial question of ie explanatory uructure of
th nnolechlae Darwinisie spprosch is kes up. “
Feenecean ot reauction as long been the object ofa vigorous
acon ecercen ceductioniatcand ofganiomie biclogy. Thee
SaaS Enos unlvingeysteraproceedsfrom the asumption that
He proceden nun fn’a stacy camabgenette way and an be
See eee ele bats of the matcsl properties of hele
molecular carriers (the so-called genetic determinism). If limits
eee ce ghyaieal desniption Sf iving systems, den theve are
2S Risdametil in nance, Rutare due soley to the complet of
Rot fondamnen at Paagarlemlc theory of lving sstemh om te
ae eee atders dhe phenomens of ie rom heli ovat
cee ane ouanates'a councctlon beeween the level of integra
Soe eat af eauling properties, This\eabore all
Seen aa tne senae datas kucgrated genetic ecm at
orci of oopanbanton can poms preperdcs that cannot Bt
Bey Gnea by sephyaical and ‘chemica!anayss of fs subesterme
sees perdction onganismie biology denies the possibilty of
Summary 175
physical foundation of the principle of selection, as the natural
selection is considered in organismic biology to be an irreducible
property of systems that are already living.
Organismic biology is based upon two central theses: (1) the
whole is more than the sum of its parts (this leads to the principle
of emergence); (2) the whole determines the behavior of i
(this leads to the principle of macrodetermination)
The universal validity of these two theses, and in particular their
relevance to the phenomena of life, are not questioned here. What
we do criticize is the claim that reductionistic biology is fundamen-
tally incapable of explaining holistic phenomena, such as the
emergence of macrodctermination, within the framework of its
analytical method. However, the analysis of concrete examples
from physics and chemistry shows that in the branches of science
to which the reductionist program attempts to reduce the phenom-
ena of life, the concept of system has all along been a genuine
component of explanations. Therefore, the concept of system
cannot be used as a criterion for distinguishing between between
animate and inanimate matter. On the contrary, the special
position allotted to the “system” within organismic biology is
beginning to allow a methodological mysticism to creep back into
biology, at a time when this had only just begun to be eliminated by
the development of molecular biology. Tt can be shown that a
number of barriers to research in biology have been erected merely
by the unreflecting adoption of holistic thinking. A principal
victim of this is organismic biology itself, which is now confronted
with several self-sustained contradictions (chapter 12).
After showing in chapter 11 that the methodological position of
‘organismic biology is already anchored in the reductionistic pro-
gram, so that it does not present a methodology that goes beyond
that of reductionism, itwas demonstrated in chapter 12 that one of
the central assertions of organismic biology is incorrect. natural
selection is not an irreducible property of living systems, but is
demonstrably a physically justifiable extremum principle that al-
ready appears in inanimate material systems as long as certain
material and physical prerequisites are met.
The physical justification of the selection principle provides
profound insight into the mechanism of evolutionary processes. It
is seen that the Darwinian concept of adaption is related primarily
to the functional aspects of the adaption and less to the structural—
in agreement with the thesis with which we started, according t176 Summary
which the semantic aspect of biological information is objectifiable
only via a dynamic criterion of generation of information,
‘When relaced to the phenomena of life, the concept of con-
straints has an expanded meaning—in comparison with its tradi-
tional application in physics. In this case it embraces both biologi-
cal constraints, as laid down in the primary structares of biological
macromolecules, and also the physical environmental conditions
Under which the biological macromolecules operate as carriers of
information or of function, respectively. Itis primarily the biologi-
cal constraints that are incorporated into the explanandum in
evolutionary explanatory models, while the physical environment
is given as an antecedent condition.
‘The molecular-Darwinistic model possesses oe important fear
ure in which itdiffers from traditional models of physical explana-
tion: while in traditional explanatory models the constraints
present contingent quantities, which are imposed upon the ex
planans as irreducible antecedence conditions, the molecular-
Darwinistic model considers the constraints to be a part of the
explanandum (chapter 13)
“The issue becomes more complicated in consequence of the fact,
that on the evolutionary level the biological constraints and the
system dynamics on which they operate are interdependent. The
feedback between the constraints of a system and the system
dynamics induced by the constraints is so characteristic for all self
organization processes that this phenomenon can be used to
define the term “selforganization” (chapter 13)
In the historical origin of life, the development of selforganizing
systems began from unspecific constraints, as they must have been
mt the end of the phase of chemical evolution, when macromole-
Gules did notyet embody any genetic information in their primary
Structure. During the period of self organization, the unspecific
constraints were then transformed successively into biological
ones, that is, ones encoding genetic information, The biological
Constraints are thus noncontingent quantities insofar as they
represent a limited and, from an evolutionary standpoint, reg-
ular selection from an immense number of physically equivalent
alternatives.
"The characteristic features of the evolutionary optimization
process can be illustrated with the help of a simple model (chapter
Be figure 16). [fall combinatorially possible sequences of a biologi.
eal macromolecule are used as coordinates in a "sequence space,”
Summary 177
then the proces ofthe origi of biologie information resembles
halk through a multecimensional landscape, che profile of which
W'Rtcrminad bythe sclonton valves associated with a vale
oontinte ale prof). Tne ete taken subject only othe
rule thatitalways leads from a low peak toa higher one, thatis, fron
a low (local) maximum to a higher (local) maximum. :
“Tras only the sign ot the graien ef the optimization ute
determined by nial law wo tedeaeapath there ae bane
‘¥0 reasons for the indeterminacy of the detailed path, One is that
the “direction” taken by the opsimicaton proves depen om
genetic variation, and the in sen ise rest a fundamen
thc value prot depends upon ie naviduals taking pati the
Crotusonary proces” Each iferental shift in contention
Ginstboson deforms the value profi iwc so tari comes ca
Infuence the gradient and the “airecton” of te optinizacon
route Gonl ad gon diectedens in a procen or blogic! sale
Srenistanareinamnioyimereonnecs na
otgiesl structures exe but oot wharbilogea scares see,
Te traciores ha are found elect the hie! uniguenes a
thing nystome, and the delale of their exigin tc in pence
Imactemlbie ir deverption in terms ef natal awh nese
the ongin of bloga information cay indeed be explained to
general phonomeno, but the conee content Of tog
Ean on be sd oe he of ps ed
erdependent.Notes
1. Larnarek (1809),
2, Darwin (1859)
3. Independendly of Charles Darwin, Alfred
sel Wallace had arrived at similar
larch 9, 1858, Wallace
Tendency of Varieties 10 De
ich Darwin recognised the
‘The manuscript was therefore read
1¢ meeting of the Linnean Society of
ly later were published by Feancis Darwin under the tie The
Foundations of the Origin of Species. Wallace himself never attempted to cast doubt
‘on the Darvan’s priority and expertise in repectof the development of dhe theory
of evolution
4. The term“
alongside RA.
le population genetics. An
ven by Earl Hanson (1981)
8, The phenomenon of purposiveness was dealt with by Kant in his Kritk der
co ). He was particularly occupied with the question of
{gpossible toexplain che obvious purposiveness
‘goal or cause,
9, Pitendrigh (1958).
10, Monod (1972) p. 15.
11. Monod (1972) pp. 31F
Nas 179
12, Excerptfrom the poster "Metabolic Pathways," revised and published each year
by the firm Boehringer Mannhicira,
13. Perutz (1972) p. 354,
14 Perue: (1972).
15. A gppical example of this is cytochrome c, with its 104 aminoacid residues.
Cytochrome ¢ hasan importantfunctionasan electsonstransferring proteinin the
respiratory chain.
16. A chain of m symbols of 2 different kinds can be arranged into N= a
combinatorially possible sequences. For nucleic acids, 1= 4; for proteins, 2~ 20,
17. Compare Lohrmana et al. (1980).
18, Fierset al, (1976). The sequence excerpt shown is from the replicase gene of
se.
24, Polanyi (1968) p. 59.
25, Weiesicker, CF. von (19
26. Weiasicker, CF. von (1972) p. 522
27. Seltfert (1968),
28, Shannon and Weaver (1949).
29. Weaver (3949) p. 12
80. Welzsicker, CF, von (I97L) p22. ~
31. Hartley (1928).
82. Shannon and Weaver (1949).
38. Wiener (1950).
34. Details can be found in the appropriate texthooks. See, for exan
sn),
35. Brillouin (198
36. Woinsicker, GF. von (1972)
87. Rényi (1970)
38, Ruch and Lesche (1978),
39. Chaitin
ly a tong asthe recipi
-VOLUTION THEORY a phrase:
of the appearance of any particular letes
the sequences of
Englshlanguage. The probability
differs feom that expected on dheas been generalized by Carl Fried
rafound way. Weivsicker takes u
tion asa measure of the
fy of heredity informa
This is discussed in chapter 12,
50. This appears to confirm the assumptions made by En
of the coding problem, in his book Wha
the goaldlireeted activity.” Ac
the semantics ofa messageare not measured by how the
t by how he could react in particular relev
(1968) pp. 65tf.182 Nous
63. Weizsacker, E.von (1974).
64, Weizsicker, E. von
65, Weirsicke!
666, Welasacker, E,
E. von (1974)
69. Jacob (
spsbe used as the basis For such ‘confiematory i
ical approach to the degree,
hen beimaginableasa gain in confirmation thr
hed by
Noes 189
3 (ye MSR, etc.) of the bac
Property: they Induce in their host
the phage RNA possesses special
RNA before the copying star
sgl RNA reproducing gem of the Q, phage ca bes
cee the corres184 Notes Nows 185
In sequence and are adapted to the reaction conditions under which they were
formed.
istence ofa teleological princi
own representatives of
Kuppers (19796, 198)
96. Popper (1989) p. 58,
97. Kppers (19790),
198. Concerning the problem of the synthesis de novsof RNA structures (see note
95), Popper writes: "We sere best
adapted RNA sequences there eres, which normally develop further
that correspond to an enzyn the embryo, give complete embryos when they are isolated. Tis seemed to
contradict the mechanistic picture of the structure and
process
Driesch (1908) had shown experimentally that every fragment of two
e cells of a seaurchin can dev ete larva, Even
is cut down the middle. These experiments tn
led the neovitalists tothe conclusion that there must
the biosphere that directs the processes of ife in a
(2968) p. 1308.
104. Potanyi (1967) p. 62.
106. Polanyi (1967) p. 64,
107. Polanyi (1967) pp. 64E
ler (1983) pp. 640M.
109. Ifthe schema of DN-explanations (see chapter 13) isused asa systematization
‘model, the “formal” teleology denotes the ease of "reflection with respect io ime”
terized aptly by Monod: "For vitaisin,
ifmot of actual paradoxes at leastofs few “mysteries.” Develo
Tor the time being suil ‘reserved,’ such speculation will prove as sterile asin all the
others where ithas been practised up to now" (Monod, 1972, p.37). In this way,
scienufic vitalism was pushed further and further back from areas that previously
hhad not been—and in some cates sill are not—completely accessible for physica”Nous 187
5 his own
We idea of random sequence is an age-old problem of proba
and hat given ss
1964). An elementary
these sources, has been given by188 Notes
2. Kolmogorov (1968)
8. See, for example, Ebeling and Fei
sh coule generate a Lmembered
wand 2"" algorithms of
thas given by
heats
hrof the algorithms with K