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‘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 super Summary 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 these 172 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 t 176 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 dhe as 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 corres 184 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 by 188 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

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