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THE _——— DIALECTICAL BIOLOGIST Richard Levins and Richard Lewontin Harvard University Press ‘Cambridge, Massachusetts and Cpt © 1985 te Prien an aos of avr Clee Pind he Und Sa of than ae los FaaonyAdtions, sr tu 2." nelog seca sptee—acre aa Gusti peso aeaie Ravecrtat tw) (sav osteo x ape) To Frederick Engels, ‘who gor it wrong a lot of the time but who got it right where it counted 08 EVOLUTION te Cartesian mechanical for the greatest number. At the same time ei ‘Su by atone ad een of ‘Sie rom the carer worl. aes he ogy of eulivio on rami ably arcs smote canny ter) 2 much 8 doe Bout poi ny gtr wiih by Wh ony Te rn kn pes, wth sens 1 demons hat a erga nc expectel optima, kel oe conte eee soup come nto area x ent oh Sen pening inven. Icons Drv, od ies a th see of ima HTACS, Pt Sa lng bb the consi er ment Thre aig sly wen hs Streit ame anode so hs adam psi, te sought my 8 oF ra ery nein mei see acai La cl, a hat hole acl chan aut ran they need oKerp pwns dane 3 The Organism as the Subject and Object of Evolution SS ee ae Se aecarecereenttuensoene a eee Ba coosuncnincnest tas eer pointes tore oan a eerie eee se es ep ses eter ee ee eee pancneteter iemtay route fects terais ornare a eee Sictasmuncksia viene sane iene ood ee eecsam re sce secretes SS ee Be ovens eerie re ee oar es coer es Se a a tapi wa plied i Son 18 90: 6-2 Inconirast, Darwin proposed a vritional principe that individval enters of theensembe diffe from each ater in some properties and that the system evolves by changes inthe proportion ofthe diferent types. Theres asrting-out process in which some variant types persist While others disappear, so the nature of the ensemble as a whole ‘changes without any successive changes inthe individual members, Ths variation among objessin space transformed qualitatively into temporal variation. A dynamic process in time aries as the conse (quence ofa sai variation in space. There isn historical process oth- than the elation of living organisms tha has this variational form, atleast as far as we know. In iransformational theories the individual elements are the subject of the evolutionary process; change inthe elements themselves pro- tlces the evolution, These subjects change because of forces that tentiely internal co them; the change isa kind of unfolding of stages that are immanent in them. The clements “develop,” and indeed the ‘nord “development” originally meant an unfolding or unrling of Predetermined pattern, a meaning ic til retains in photography and {eometry. The role of the external woed in such developmental theo fesisrestricted toa inital iggering to sete proces ia motion. Even ‘Lamatet's theory of organic evolution dd not make the environment the crestor of change bit only the impetus forthe orgasm to change its through wil and striving. Two characterises Now from such a ‘tansformational view. Fist, the tages through which each individual pases are themelves the preconiton forthe next stage. There are no horteuts posible, no reordering ofthe transformation, and only one posible end tthe procst Indeed, the tension and contradictions of ‘ne stage are actually the motive forces ofthe change tothe next tae. Marx's theory of history ie precisely sucha theory of well-ordered hi torical tages, each of which gives ret the next as a consequence of forces interna to each tp, Theories of psychi development, such as those of Freud and Page, are derived from theories of embryologica development of the netenth century. Each stags, whether of the body or ofthe poyehe, sw necessary preconciton ofthe next stage and leads to it because of forces that ate purely internal st each moment The role ofthe oer worlds coset the process in mation and to alow ‘he sccesfl completion of eshstep. “Tis roe of the environment provide the second charac transformational theories, the possibility of arrested development. IF nie of ‘external forces block the unvollng,thesystem may become permanent: ly fixed at an erly sige, and itis this premature fixation that explains any observed variation from individual to individual, In Freudian the ‘o1y the personality may become ied at an anal or ral rote stage or 1 the stage of Oedipal resolution and a0 givers tothe manifest var ations amons neurotic spon. Ta the theory of neoten, evolutionary theory retains notions of ‘ar arrays of stages and arrested development. Accodina to is view ‘organisms tht appear later n evotion hive the form of erier devel ‘opmental stages oF thelr ancestral species. Gorilla and human embryos resemble cach other much more th the adults do, and ado. Ramaas fe morphologically like the gorilla fetus. Humans are thus grills ‘born to soon and fixed ata gorilla fetal sage It follows from such a theory that ifthe development ofa human being could somehow be ‘unblocked, it would develop the long ams, receding jaw, and sagittal ‘est ofthe adlt gorilla ha present bu hidden. Iescem obvious that ‘ neotenie view of evolution is severely ited in is cope since adult ‘humans cannot be sid to resemble the carly embryonic stages of sh Indeed, evolution cannot be any kind of simple unfolding, for such a homunculus theory implies the absurdity that mammals are already edn the carl singl-eled organisms. ional theory's theory of the organism asthe obec, not the subject, of evoluionary forces. Variation among organisins arises as a consequence of internal forces that are autonomous and Alienate from the organism as a whole. The organi the abject of these internal Fores, which operat independently of its funcional needs or of is relations to the outer world, Tht is what is meant by imitations being random.” Its not that mutations are uncaused or outside of e deterministic word (except as quantum uncertainty may enter into the actual process of moleular chang), but that the Forces overning the nature of ne variations operate without influence from the organism or is mies. Once varation as cared, some variants survive and reproduce wile oters are los to the species, according 10 the rari between the varlan (pes andthe envionment in which they live. Once again the oranism isthe object, this time of external forces, which are again autonomous and alenated from the organise, 4s whole. Te environment changes as a consequence of cexmolon ‘al, geological, and meteorological eens that have thelr own laws dependent of the ife and death of the species. Even when the environ: ment of given species includes ocher species, the hiories of those species are autonomous and independent ofthe species being consis- ed The roles ofthe external and the internal are not symmetrical in Dat- ‘winism. Pre-Darwinian variational theories placed the internal forces (of development in the dominant position and understood history a8 & consequence of development. Neoteny belongs to this Paton, pre- Darwinian tradition fori portrays the evolution of oasis as noth- ing but various sages of arrested developmen; ontogeny dominates history. In Darwinian chery the reverses tue. Historia frees are dominant, and development docs nothing but provide the raw material forthe forces of natural selestion. The enteral chooses which of many possible internal states shall survive. Thus the developmental pathways that we se are the consequence of history, not is cause. Ernst Haeck c's theory of recapitulation i, in his sens, erly Darwinian, fr it Ilds chat the embryonic tages through which an organism passes are ‘the rae ofits evolutionary pas, not the mage of its evolutionary fu ture. Human embryos have gl sits because thei fish and amphibian ancestors ha them, but n human beings theglelts disappear because ‘human beings have evolved further. Through evolution, new sages of development have been added, stages that were not immanent in De- ‘vonian fh So istry in Darwinism dominates ontogeny. “Thos classical Darwinism place the organism atthe menus of intr nal and external frees, each of which has itsown laws, independent of ‘each other and of the organism tha is ther creation In cutis way the organism, the objet of thee forces, becomes irlevat forthe evo lutions, because the evolution of rzanisms i only a tansformation ‘of the evolution ofthe environment. The organism is metly the medi tum by which the exeral forces ofthe environment contont the ite. nal fores that produce variation Ir snot surprising, then hat some ‘vulgar Darwiiss make the gene the only real uit of seleton and see evolution as a process of eiferential survival of genes in response 10 the external worl. tn The Sesh Gene Richard Dawns (1976) speaks ‘of organisms as "robots controlled body and mind” by the genes, asmothing but a gene's way of making another gene. Ihe species isin: Seed the passive nexus of gene and elctive environment, ithe genes propose and the environment dispose, then na deep sens organisms really aeirrelevat, andthe tudy af evolution x nothing but acombi- nation of motecula Biology and geology. ORGANISM AS SUBIECT AND OBJECT © Bur such view givesa false picture of organic evolution and cannot -successflly cope withthe problems posed by evolutionary biology, for itignores two fundamental properties of living organisms that arin di- feet contradiction to a superficial Darwinism. Fis, its not ue that the development ofan individual organism san unfolding or uroling ‘of an internal program. A‘ a symposium in 1982 commemorating the ‘hundredth aniversary of Darwin's death, a leading molecular bilo- ‘st exposed the bei that ifthe compat sequence of an organisn’s| DNA wore known and a large enough computer were available, it ‘would be posible, in principle, to compute the organism. But that is szely false, because an oxganism does not compute itself from is DNA. The organism sth consequence of ahistorical process that goes con from the moment of conception ust the momeat of death; atevery ‘moment gene, environment, chance, and the organism asa whole are allpartiipaing. Second, it isnot rue that the fe and death and repro- Secon of an ovanism aie consequence ofthe way in which theliving being is acted upon by an autonomous environment. Natural selection is nota consequence of how well the organism solves a set of ined problems posed bythe environment; onthe contrary, the envionment fd the organam actively codetermine eachother. Theinternal andthe external factors, genes and environment, act upon esc eter through themed ofthe organism, Juste the orgeism isthe nexus of inter ‘al and external factor, tia the locus of thei intcaction. The oF ‘nism cannot be regarded as simply the passive abject of autonomous {ternal and external forces; itis lo the subjex fis own evolution. (GENE AND DEVELOPMENT nis common, even in textbooks of genetics, to speak of genes deter- ait, a knowing the gene means the tat ofthe organism i. ier. This notion drives from several historical soutes. Fis, since the nineteenth century, embryoogits have taken their poblematl 0 be explaining how a fertilized egg of a frog always Becomes afro, while that ofa ehicken always develop into another chicken, Even ‘when the environment in which development staking place is severely Aisturbed, a process of regulation often assures that the final outcome [isthe same. I the developing ib bud ofan amphibian embry i cut ‘out, the cel dsagaregated, then put back together again, andthe ump of cells simplanted in the embryo, a normal eg will develop. And a0 ‘environmental disturbance has ever caused an amphibian embry (2

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