_ American Scientist 4 Wol-49,.No. 1 JANUARY 1952 | EVOLUTIGN-AS VIEWED BYON SJOURNALA ARTICLE EVOLUTION, AS VIEWED BY ONE GENETICIST = RICHARD B.GOLDSCHMIDI American Scientist, Vol. 40; No. 1 GANUARY 1952), pp. 84-98, 135 (16 pages) https://wwwjstor. org/stable/27826418 > About this issue > Referenceswine Warcur (letter, June ?, 1908) lution_of the organie-world, from the synthesis of the first molecules endowed with the faculty of reproducing their ___kind-te-the most advanced-type of life, must have taken place roughly “within-the-past two -billion-years on our planet. All the facts of biology, geology, paleontology,-biochemistry, and radiology not only agree with ——_this-statement but actually prove it. Evolution of the animal and plant — world -is-considered by all thase entitled to judgment to be a fact for —_____Whichne further pr f isneeded. But in spite of nearly a century of work and discussion there is still no unanimity in regard to the details of the =—=means-of evolution. This statement should not be misunderstood: all 0 have mastered the available facts are agreed upon the s, the Lig outlines of the explanation of evolution. Nobody doubts that the main theses of Darwin have settled this point ance and forall evolution results from (1) the presence of hereditary jatin ich furnishes-the material thut is able to fit into all the different niches-of- environment and therefore allows organisms thus en- “dowed-to-oeeupy those niches; (2) the action of selection, which puts at _isadvantage the types that, not properly fitting into their environment, -are-not-adapted. This allows the better-ndapted ones alone to multiply “more rapidly-and to survive, Hefeditary variation (mutation) and ini Le eel i) the “ | ih Hi wil i ti ee sis GT % i a of iin it P ty i a er tie Rte aa Nath bLd| gh ul ne ul Wee (iP che aN a ieee ye Pe it nee) on 1 es ! ie nn gave | Pr ent Hi i fl cy per etiqgan nip mea " RetaHN il hil h ul sti a St it : m m ipl h wh stn nM at i ih te =e ‘ ca : 7 il — aan ae ke Sree os a sii 1 ind te mt | | Genetics | the ah al eR | | th 4 le Hew tool] han! ben Wdded ie the i it | raat ty in evolution. This is nee of wa Arte y hity years of xis rate the central fields bf biologi a Tet ati all ‘thit genet “Darwin's i atid all mi ence of Sarton. "This Pe | I} nn | Pee MM re hasta upon thelt-xist= Pans thut within a population of an animal or slight atl upon whieb se caull Werk to. produces rather vague, ‘The early ie Hi ely ch Ta~The second important contribution of genetics to the ames of - variation is thus the firm establishment of the theory of mutation. This ‘theory states that all hereditary differences arise in single steps or mu- tations, that is, changes in the hereditary material, the chromosomes and their subsegments, the genes-Such a change occurring more or less rarely produces, af once,-the mutated gene-and the mutant organism. The mu- tated-gene now reproduces its own Kid; the mutant breeds true, and can be extracted-and isolated as anew constant form. Mutants, whether they-result in_smail visite changes or in large ones, are the material of —arduti = “Phe third basic fact concerning Variation is derived from Meéndel’s dis- very that individual mutants behave as units in heredity, which segre- ——_#ite and-recombine after ae necording to the laws of chance in volved in Mendelian heredity. The necessary corollary, to this is that is a renmaine for tion and recombination. To this must be added the ed, or reshuffled; by the process of erossing over. y defined hereditary variation as the result interbreeding of the mutants with the origi- if érassing. The consequence of the tants ces population will: therefore be the formation of = :tiptivs way becomes ole as lle I “e ee oe eye = a: i soi atest is hi i i fy ‘ ii uy tat i it i i cha ne cutIt of mutation, interbreed- i ‘bination also includes cross- mo over, and vicar upon all this is the phenomenon of domi- with ‘the proper elucidation of variation, selection, the second concept basic to the understanding of evolution, also received a definite meaning. Selection is not some more or less mystical agent which acts with purpose, ‘as the breeder does when he selects his stock. It now becomes a term which describes a simple fact, namely, that in a mixed population— “mixed” meaning the free interbreeding of a jure of mutants, with the consequences just described-—not all combinations are equally fit to compete and to survive. Mutants have a tendency to reduce vitality, Recombinations of different mutants may increase vitality, reduce it, or Teave it unchanged, Some recombinations may enhance fertility, others -may reduce it. Some recombinations may allow the form to survive _____usual sirain (eg, heat, frost); others may permit the bearer to live in thus far-inaccessible surroundings, for instance, to stand higher saline concentration-oF to live on a special soil, The consequence of all this is —= that the less vital, lees fertile variants as well as those not fitted for the ——grironment will lose in numbers and finully disappear; they are being selected against. On the other hand, the better-fitted ones will not only hold their own but will increase in number} they are being selected. ‘Thus, negative or positive selection in a mixed population is the descrip. is in such perfect equi q@nvironment that its Soni the Sta and therefore|sdlection will fhviir sacion, bsiedlly sat rs i f rn 1 retin, 6 hb an UC a i ile fit nl ii b Ha re Bet ia vil| i mn i | Ml ~ il en OTA A epee I Hl ty nn ‘ N a ns Wy nO Bonnar : nyt ws me rt | - al So ea Hf aa lad a ve Adapitati{ins thy spe eh taniy, Ih Sede the Oa of be adaptive fea | daptation, thet win did not obj " iants. Genetics Lutatir ct ‘hance and com- ) without any relation {¢ the environment. The at- ve value must therefore proceed in & way | that assumed by Darwin, This met honk has been termed * This means that among the chance mutants and their re| NT | nti i ‘ Hl Mac " na ! i Dan eat i! i “it i 4 i iy ad from Hs d called sub papecies “incipient spe- i = generally hell by taxt he is at hone a, Eee ision int) subspecies, Yange pf the species and can be distiliguished from 'a| number "of these subspecies are present in a typical geo- dal) arrangement, one ks of a rassenkreis. The de= fratures of a rassenkreds are: i - ‘the subspecies species aid Completel: therefore, where two subspecies meet xq they ae based upon mendelizing differences, rarely im-one matan yin multiple alleles, and most frequently of the ty factors, To this tnust = = a fact that in man) - character is af extemal exprvssior{ of a physiol ition. af adap tive value, duuptations in ¢ Hion are those of the most elementary, - Ff : a i sae gs t - ual mcs i t d Max ie ’ } | es ‘{Neo-Darwinians,”” are y sen which the genetirists a operating at the subspecific level, called microevolutio rt iss at work it maerodvolition. ‘This means that specien, aenert, ordléts, etndece|| andl phyla also produced by the oe secumilation of small ats, elthier by n. OF ii drift, and their final selection by the ~— “adaptive ridiation. are incipient species ad that all spegies have arisen from i rably at the end of a sab+ ies have became so different from those hat the two types, if brought together, ———=-no tonger interbrend. L ‘rather disappointing Uy|see that only i fet sel quoted over and over again, exist to demonstrate whist should it frequent occurrence. It is, of Couns, nut assumed that all trinsform into species, though potentially Neate adalat haa a ) tet | i‘isolation. : n originated by mutation on the spot, followed by further di- fe) PS transformed into species by chance development (through mutation) of in mechanisms and afterwards ence is due to existent -adaptation. It must therefore be assumed that to the area of the old one has to develop anew = _the proper existential adaptations whieh would make it again identical ~ With the former species, at least for adaptive traits. Neo-Darwinians are ~__fikely to gioss over such difficulties. —Tet_us_continue the picture in Neo-Darwinian terms. The newly formed species, just like the subspecies, may have accumulated mutants :combinations, or may begin accumulating them, permitting the oceu- of vacant new environmental niches. This time, however, the aime necessary existential aclaptations cannot be involved, but rather the special adaptation which constitutes the bulk of the differences in the higher categories. Consider as an example a bird which formed sub- species adapted to different climates—an instance of existential adapta- tion. Aiter isolation a subspecies becomes a species, which might again SSS ies of the existential type. But some “variants,” sub- it, must be filted to occupy a completely new niche. The SSS species- might have been a grain-eater, while the empty niche available is fora honeysucker. Adaptive radiation produces this type, is. But how does such a complicated genetic change, leading, by accumulation of small mutational steps, to the perfect mechanism for honeysucking in the structure of bill and ‘tongue, become available just when iti has a chance for successful selec tion? If one tries to work out this idea in detail, one soon comes to a point where it is evident: that something besides the Neo-Darwinian tenets is — heeded to explain such macroevolutionary: processes. The difficulties al- these and generic level seem to the present the level of families, orders, classes, and phyla. a umei Mili; ee | 3 ‘ i i ee ! bywith honeysucking nite ecological nicl it had reached a woi ty?) Neo-Daltwvini circumvent this difficult intr bo ee for accumulation and fixation of ne tants nM small populations. But this great deus ex mac hardly be ed to work up to bel jit the most integrated june eoiiplicated ddlapt just by baphasard || ceil einnuinssy i tt i i Macromutation mautants ce dev lepn to disturb the fabric of interwoven and cl might atest hb ‘step. eo or less divengent pattern of the tion might hin € logical changes at the level ocr ae i se ime HT Mh oo ' — rae Mavens on WN Wy) We nT Hf Hie ae a no flys yc i if forest is replaced by steppe, or the like, large attires ate affected, and this would make for a few big ‘Thus we should expect a convergent adaptation with ul he the available forms, Actually such changes » diversification. On the basis of the Neo- » should expect the production of a very few diversification. ¢tomutations, it may then be assumed that major de- nk of higher ¢ategories are attained initially by single iehi tippers producing at once the essentials of the new I} nption is violently opposed by the majority of geneti- cists, who claim that the facts found on the subspecifie level must apply also to the higher categories. Incessant repetition of this unproved claim. = glossing aw over the difficulties, and the assumption of an arrogant attitude towatd those who are not so easily swayed by fashions ir scienee- are considered to afford scientific proof of the doctrine. ft is trae that nobody thus far has Prluced species-or-genus, ete., by macro- mutation. Lt is equally true, ody eed even 2 species by— the sel i micrornuute st-known organisms, tike : we were able to combine: se of such-mitants in a single individual, this-still would have ib Liki nae tee: to any +t 4 Speci in nature. ‘Three questions now (2) Ar¢ macromutations Fidaieieee known to exist, and ean they be used as ‘model for happenings inna ture? (2) Are stich ant sufficient hi | Wl ie a dnb yi i iM ah io stamens ‘a i ce iatother rare mutant should not produce) at the level of any higher category. These macromutants, ih in themselves only freakish events within a. species, ara good models for possible evolutionary hap- penings. Macromutants may. Havel ridiidadk viability: ‘or normal viability, or may reach normal viability by the accumulation of modifiers. They may be completely interfertile with the original form, or the fertility may be im- paired. They may be dominant or recessive, Finally, if they are extreme and therefore less viable and fertile; they may vary in expressivity, within the same homozygote, from extreme hee poorly-viable and fertile to ‘The-next problem, the genetic isolation of a macromutant from the original form, spf course not: dierent fram the situation: in micro- IV etek ai eonchusioor-entablished population genetics apply also to oft eat to light rather recently, re~ ialtitis tool furnished by the giant sali- i sen i Me “ il sgt nally on uo yeep Pst yb mn oe ia | ne tipi St a dbnes ofleeoleast ehi ee TT i i ] Hl wien i : ints an faleual 6 osomal ae parallels ee can Neo:Durwinism fit these facts into its Pk ca ah see only one way, namely, by assuming that, ‘evolution by selection of micromutants affecting id physiological characters, small changes of the chromo- Appeat which for unknown reasons are of selective ji wumulated until the entire chromosome is changed. Tf one tries to give such an interpretation a real meaning, serious diffi- culties arise. In i bate tafe large inve protect certain sections of the chromosome from crossing over when But why dow to produces is nm ous parallel with the same gene divest I tiast be clear to anybodly lacing Uhese-tacts: athe} intimate architecture of the chromo- = some up to the assumed a¢etimiilation of Vita By taking into ‘account thes cytal about the necessity of itation for ai of higher categories and Lap tare modern ideas about: ; 8 the gene und it to sinall Kon cad del nn al Dr Latin Panetta NTL WALei Tui abel bit bal ay Cn picture, ithould be welcome. It is of course at such a concept is. Apart from Diptera, he chiymosome is inaccessible. If we have two | i ane 24 rather similar-looking chromo- ile to their intimate structure (apart ae —_ not to undergo sy around the centromeres de eulty in this point. “Thus the the fact of simple marromu been established a Neo-Dai tion which makes sense. | The science which ‘she ‘should be able to contribute important: information. facts is available, but there i 1 ity interpretation. ‘The —— io ah “ea h n feoin| LH fol ne | 4 wip te i cai wen eaaL i ce vi ne cha tariatls trie. Tn spite of tha nmense amount of the paleon- ‘material and the existence of long series of intact stratigraphic es with perfect records for the lower categories, transitions be- ween the higher categories are missing. ‘These and many other major facts of paleontology have found very different i in lations. If we disregard the obsolete ideas of Lamarek- ion and of mystic forces of orthogenesis, we may_aay that ists are agreed that the basic tenets of evolution, and adaptive radiation, apply also-to the ifference of opinion is encountered at the —— as in the case genetics. One group tries to account for the facts of palontology-on the basis of the Neo-Darw jan outlook.Ac————_ ie View, accumulation of micronutations can-also-explain: nit ide paleontological record. \ Where. difficulties arise; evolution oie paees pines ‘up into — a units. ___ 38 frequently -nppealed to without any explanation of how szch-a-setep—— continued unintem uceeptance of a point of view identical eee earlier in this pap rs ieee formation of the major categories by large ear the nat crac imaeromurt affecting decisive early em- thats a gre i Penny cs in Ips) mH pt rt |It might. be noted that this work has gone on independently of Kor- aybski, and perhaps many workers in those fields might even resent the implication of Korzybski’s intellectual paternity. But one must-not confine oneself to Korzybski’s intellectual progeny ———— influence of his ideas. One must also count his m in a is-why questions of priority -our day be taken-seriously. Relations between ____€ver, are important.What we have attempted here is to show, 1 not tro eared what from—whom,—but-rather_what-telation some leading ia ine age ee antec coe Emotes have toacho = oe gre Problene data Fete ten 4 ear and ‘evolution: ‘in plants. Calne University “Press, a Sidi