What is an Adaptive Trait? Theodosius Dobzhansky The American Naturalist, Vol. 90, No. 855 (Nov. - Dec., 1956), 337-347. Stable URL: fip:flinks jstor-org/sici sici-0003-0147%28 195611 %2F 12%2990%3A855%3C337%3AWIAAT#9B2.0,CO%SB2-H The American Naturaist is currently published by The University of Chicupo Press Your use of the ISTOR archive indicates your acceptance of ISTOR's Terms and Conditions of Use, available at flip: feworwjtor org/aboutterms.htmal. ISTOR's Terms and Conditions of Use provides, in par, that unless you fave obtained pcior permission, you may not dowaload an cnt isus of @ journal or multiple copies of articles, and you may use content inthe ISTOR archive only for your personal, non-commercial uss. Please contact the publisher cegarding any further use of this work. Publisher contact information may be obtained at bhupsferww.jstoc.org/joumals‘ucpresshtel. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transtnission. ISTOR is an independent not-for-profit organization dedicated to creating and preserving a digital archive of scholarly journals. For more information regarding ISTOR, please contact support @jstor.org- hup:thrww stor. orgy ‘Tue fun 21 11:13:16 2005THE AMERICAN NATURALIST Vol. XC, November-December, 1956 No. 855 WRAT IS AN ADAPTIVE TRACT?* ‘THEODOSIUS NOBZHANSKY Deparcaent of Zoology, Columbia Universiey, New York ey MORPHOLOGICAL CONSERVATISH IN SIBLING SPECIES ‘The idea of the present article, which avempts te deal with the general problem of the nature af adaptedness of traits which dilferentiaee caces, species, and other groups of animals and planes, has been suggested by considetation of a tapic apparently nat closely zelaeed. Tae genus Drosae phila contains 2 camer of paics, or of groups af several, sibling species. This pheaomenon is patticularly common smang Drosophila of Brazil and of the American cropics in general. Sibling species are morphologically <0 inilar, or neatly identical, that distingutshing them by inspection is dite ficule of impessibie, Yer, they ace demonserably good biolagical species, since the reproductive isolation between them is camplete of neatly 50, Uhy is the genetic differentiation in sibliag species gor accompanied by morphological divergence? The quadruple of sibling species, Drosophila uallistom, 0. aulistorum D. tropicalis, and D. equinoxtalis, may be considered here as an example. Semial isolation heeneen ther even when no choice of mates is availatte; when interspecific copulation does occur no viable progeny is produced. The (our species are sympactic over extensive certitaries in South 2nd Cenceal Americas the populations of each species cagtain arrays of chromosomal vatizats of the inversion «ype, bur these is ao evidence of cither regular or occasignal cansfer of these ‘80 strong that crosmiosemination is rare variants from species to species, as would be expected if effective bys bridization occurred, however rarely, in anturat habitats, slarphological differences berween these species exist, especially ja the male geitali2, hue they are so slight chat i is not feasible to discinguish the species by inspection of living individuats under a binocular microscope, ac least not in the female sex (Burla er al. 1949, Spassky uapublished). Analysis of the salivary gland chromosomes shows that the gene arcangements in the The aubstane Acadenin Brasilel of this atticle was presented in aa address read before the fe Ciencias, at Rta de Jancieo, 0M May 30, 1956. 337238 ‘THE AMERICAN NATURALIST four species have diverged, especially in ane of the chromosomes, the hicd, which in chree of the four species is buile 99 differently chat homal- ‘ogous regions are no longer recogaizable, There is also gaod evidence of physiological and ecological divergences, Every species has a dist bation area differeat from the others, as well as hahirat preferences of ies own. Where the siblings are symparric they may vary in relarive abundance at different seasons. In laberatery cultures, the larval development of P. vwillistont ie faster thaa thae of D. paulistarum: D. tropicalis gives beau! fully clear polyteae chromosames in the sativary glands, D. equinoxialis gives poor chromosomes, while the other ewo siblings are intermediate. The reasons far the scanty morphological divergeace despice cansiderai genetic ant physiological diversification are a matter of speculation. have surmised thac the external morphology of some species gtoups of Deasophila has reached co high « level af adaptive specialization thar mast changes in the visible body srucrures are discouraged by natural selection; nevertheless, room is left for ecological diveryence based oa physiological differentiation (Dodzhanskey 1941} Artempts to apply this canjectuse to conctete eases lead however to difficulties, Indeed, most of the morphological traits which differentiate related species of Drosophila are of 2 kind che significance of which for the jeat, A partial lige of these craits will make this point clear: Number af branches in the arista, shape of the facial carina, width of the cheek celative to that of the eye, presence af one ys, tho praminent bristles in exe oral com, thorax more or less polisted or Zal!, presence or absence and kind of the calor patteza on the thorax, naner of cows of acrostichal microchactae on a certain portion of the thorax, anterior scucellar bristles divergent, parallel, ot conversent, re- lative lengths of certain bristles on the pleurse, proportions of certain sections af wing veins, presence or absence and intensity of color carke ings on the wings, pasicion aad kiod of bristles on some leg segments, Zecails of the color pattem on the abdomen, shapes of various parts of male genitalia, aumber of coils in the testes, shape of the speenathecae, number and kind of coils ia che ventesl seminal recepracle, ete. welfare of the acimats is nor at all ADAPTIVE AND NEUTRAL TRAITS IN CLASSEPICATION Mey should chese and other sinilar tains vary in some species groups of Drosophila, and yer be fixed in species of other groups abing the lawsershling species? ‘This problem ia only a epecial case of a tore reneral oa, that ofthe aaeive significance ofthe keleMdascopic diversity of morphological traies in different organisms, Darvin was quite aware thet che morphological differences which systematists use to disetiminate tsermeen species, subgenees, genera bes, families, and other groups often concen surprisingly tsivial. traits, of 09 appatent consequence co the crganians involved. Jk vas probably this, noce than any other dittteuley, wich Ted daring the first quarter of the cunent ceonuy te a cemportey ellnse of the eheory of natural selection, If most differences which weWHAT 6S AN ADAPTIVE TRAIT? 339 observe becween orgaaisins were adaptively aeutral, thea ic would seem thae factors other than selection must he iavoked to account for the origin and maincenance of these differences, le has bees tepestedly suggested in hielagical and anchropalogical Vrerature, even in recent years, that classification of organisms should be based on adaptively neural «waits. Allegedly, che waits which accu: rately ceflect homology aed common desceat are neucral, while positively adaptive teaits are too easily molified by the environment and are ape ¢0 show analogy instead of homology, ecological correspondence rather thaa common descent. But if so, chen the evalution of teaits on which the classification is based ig a nystery. To use Drosophila again az an illusteacion: che more than GOQ knows species of chi genus all have three erbital bristles on either side of theie heads, and the anterior of these bristles is always preclinate (bent forward), while the other two ace £e- clinate (Bene backward), Now, why should this character be retained $0 tenaciously in 30 many species? Is it really important far the ‘lies af this ‘genus to have one proclieate acd cwo cecliaare orbical beistles? ‘The basic postulate of the modern biological theary of evalurion is chat adaptation to the environment is the guiding force of evolutionary change. We need aot follow the extcemises who insist that all evolutionary changes are adapcive; however, we have to suppose chat most organs and functions ‘of most organisms ace, or at lease were 2¢ the time when they were formed, in some way useful co their possessars, Nothing less than this is acceptable if the modern cheery of evolurion is sustained, ‘Tne argumvenr is, then, thac our failure to See the adaptive singificance of most teaits is simply 2 measure of our igcorance. Some authors have even wged thac assuming seuteality of eaits is methodologically inad- missibte, since such an assumption allegedly prevents making observations and experiments needed co diccover their adaptive significance, The work of several investigators (e.g. Lamatte 1951, 1952) has at che very [east shown that supposing some traits to be clase to adaptive aeuerality may be 8 fruittul working hypothesis. The usefulness of a uaie muse be demons suated, ie cannot juse be eakea for granted. AL! that can be said is that ‘one's failure to See the adaptive significance of a teait is no proof that ie has none, The matter must be eettled on the basis of evidence, aot by theocetical arguments alone. Ie is a fact thar che variation in some traits which ar one time seemed neutral was Inter discovered wo be adaptive, The chromosomal inversions ia Drosophila populations are an example (Dobzhansky 1947); che colors and color patcerns of Cepaca snails ace another (Cain and Sheppard 1954). Suc ic is sil) as teue today as ie was fn Darwin's day chac the adaptive significance of most cuits which vary heween and within species is ob= scure, Suffice it € mention thar pegetically nothing $8 known about che significance of mosc traits which vary between races of wan as well as amoag individuals within 4 race, Although Coon (1955) bas made some340 THE AMERICAN NATURALIST inceteaciny sugzestions concerning the passibte soles of suck apparently nevteal human traits as che form af the mqse, lips, hair, and similar variacions, che macter awaics experimental verification. When one considers taits ia which species of laseces and ocaet organisms often differ, such as the differences berween Drosophila species meatianed sbave, che suspo- sition thar all of even most of them are directly usefal t0 chait possessars strecches eo much ove's eredulicy. CORRELATION OF TRACTS Ac this pater we should inevieably inquire whether the impasse which we seem to have ceached in out argument may be due to a faulty statement of che problem. A visible "trait"? is an eutcome of the occurrence of cettaia developmental, physiological, and ulvimacely physico-chemical, processes in the organism. Teaits, auch as the presence of one proclinate and cwo reelinate otbital bristles in Drosophila, need noc by themselves be of aay particular imporance to the animal, In fact, some Drosophila autaaes have fone or more of the orbital beistles missing, and rhe mueant flies seem to suffer no inconvenience ea this aeceuec. But the processes which cesuile in the formation of certain bristles may give rise also co the other traits, norpher logical and physiological, in the same organism. ‘The praclinace er ce= ctinate position of 2 bristle, though quite unimportant in itself, may be an ourmard visible sign af the aceurrence in the organism of quite importance developmental processes. The latcer axe not necessarily disturbed when other factors cause some particular bristle co be missing, a mutant may survive wighout it, However, if a bristle is formed at a certain poine of che hypoderm it is determined either as proclioate or recliaate (Stern 3954). ‘Tye fixation in the phylogeny of Drasophila of this and of ather apparently equatty unimporcant eaits as generic or specific characteristics is, then, fa consequence of the adapciveness of the processes of which ehese traits are the concomitants. lc cannot be stressed 109 often char garural selection does nt aperate wich separare “weairs."” Selection favors genotypes che cattiees of whick transmit their genes co suceceding generations more efficienely chan do the ‘cattiers of other genorypes. Now, the reproductive success of a genotype is determined by che totslity of che eraics and qualities which it produces in 2 given cavirooment. A disadvantage in some respects may be compen sated by advamages in other respects, Thus, in an experiment of che writer with laboratory populations of Drosopbila, a genorype which was weaker than another genotype in larval competition under crowding had nevertheless higher adaptive value, presumably because of a higher fecundity of adults (see also Sitch 1995). When a crait is a part of @ complex, 2 system, ot @ syadcome of developmentally cotrelated charneters, ic is obviously the whole system which is favored ar discriminated against by nacural selection. This is aot contradicted by the fact thar some one character may be of paramoucr significance in deciding che success otHAT IS AN ADAPTIVE TRAIT? ail failure of 2 genotype in a given envivonment and at a given time, Frest resistance may be decisive during hard winters, and drought cesiscance during dry suamers. PLEIOTROPLSH When genotyres which differ in s single gene yield phenotypes which are Unlike each ether in ewe or more not ebvigusly related eaits, this gene is said te have pleiateopie, oe manifold, effects. Examples af pleictrepiam are common, The classfeal mucancs of Drosophila were decected and identified almost always by some externally visible abnormalities in the steuccures Gf the adult fly. And yet, these mutants manifest very alten alsa some variations in che viahility, fectility, longevity, and/or other physiological characceristics. When such mutanes are submicted co the action of natural selection, their face is likely co he determined nee by their cbvious ex- ternally visible characteristics bue by che more subtle physiclogical anes. Reed and Reed (1950) and Mecrel] (1953) found thar the white-eyed mutant of Drosophila melanogaster is discriminaced agsinst by nacural selection in certain experimental enviconments; however, this occurs not because their pigmentless eyes make the white flies blind, but chiefly because the fe- males, bath red-eyed and white-eyed ones, accept the advances of normal redceyed males mote often than they de those of the white-eyed males. Quiee trivial, and by themselves adaptively acutral traits, may then, be established or eliminated by natural selection because they are componeats of genetic syndromes which contain also some selectively effective feances. [cis unimporant in this connection whether the different parts of a syndrome are due to a multiple ™ of a single "primary" action in che development. This distinction Grine+ beets 1958) has operatioaally lite meaning ae che present level af aur knowledge. Whether of aot genes produce difference “primary” products in the same or in different tissues at different stages of the development is an inwactable problem which has not heen solved. Regardless of wihac may be ies eventval solution, the characteristics of animals o¢ plants which appear striking co ow eyes ave often less important co heir possessers in the struggle for life than che less obvious genetic and developmental correlates of these characteristics. In looking for 29 explanation of the avolutionaey hehavior of a trait, the adaptive significance af correlared craits cannot he ignored The evalutionary importance of pleiatrapie:n is not lessened by che fact chat mast mutations of single genes are raflected on the phenotypic level by aleariag some one trait apparently mace strongly chan ochers. For example, keowe mutations at the white locus in Drosophila all affect the eye color, no matter what other characters they may influence at tha same time. Uhite is supposed to be aa “eye-color gene.” In microorganisms, mucatioas which are characterized biochemically usally affect ane and only one step in some ceaction chain. Sut ie should noe be fargatten az ic often is, primacy” action of a gene or co branciting382 ‘THE AMBRICAN NATURALIST that by comparing phenotypic effects of gene alleles A, and A, we learn at mast which developmental pracesses are influenced by the aifference between these alleles. We still do nat know the toral extent of the field ‘of action of either Aj and Ay To find this out, it ie necessary to compare the effects of the absence (deficiency) of the gene A with those of its presence. But even when this is possible, one eannor be sure that genes homologous to A are noc catried in the same genotype at other loci.. Recent work aa aseadoatieles has indeed show chat zene duplication occurs in the evolutionary process mare often chan previously kagwa, One cannot exclude the pessibility chat che presence in the generype of at least one init of every kind of gene whieh a species possesses may be essential for the life of the cattiers of this genotype. GENES AND DEVELOPMENTAL SYSTEMS Pleioerapiam ie cleatly a part of the story of the establishment io evolution of appaeently useless eraits, ut ic can haedly be the whole story. The problem is really insoluble as long as we think of every gene as controlling one or more ttaits o7 pracesses ia isolation from the develape nent of the organism as a whole. Phat, indeed, are the particular adaptively inporant tcaits which ace conelated with the presence of one peoclinate ‘and two seclioate oral bristles in Drosophita? Peshaps auch waits will some day be discovered, hur at present it may be useful co explore the possibilities of other aad moce genetal solutions. The development of an individual spans the inuerval of tiae which begios a fertilization of the ovum and concludes at death. The sequence of events which occur during this dime is described most easily in tems of the appearance, persistence, and disappearance of various “eaies" and “processes."" The same is true of phylogenetic development, except thae a¢ least the beginning of the sequence of rhe events is here always lose in the dimaess of the past. For example, we do know, or reasonably infee, thar in the human ancestry the trait “erect postute"” appeared, and the etait “facial progaathisa"” disazpeared, at a certaia time. Jo 20 describing our observations on the ontogeny or the phylogeny ove should, however, keep in mind thae she individual and phylogenetic Aevelopments do nee caasict of a series of diserere evens bue eee really continua in tine. The isolation of “ales” is, thea, only a semantic Aeviee valid aad useful in recording our observations. However, the use~ falness and validiey of this device lapse as soon as the “traits” segin co be thought of as having independent existence. But this is precisely what all oa alten happens in the ehinking of waay biolagisgs. Bath the ontogeny and the phylogeny ate visualized as building a kind of mosaic picture or as pucting together a jigsaw puzzle. The scones of the mosaic, or the pieces of che jigsaw puzzie, ate the waits, each derernined by ane o¢ by several genes, which are gralvally assembled by accretioa until she pietute (organism) is complete, of wax it zeaches the stare in which the observet anudies it, Now, this is a shockingly unrealistic way of representing eitherWHAT IS AN ADAPTIVE TRAIT? 343 the gatogeneti¢ of the phylogenetic development, Ir becomes 2 source of pseuipoblens, ane af which may be che appateat prevalence of traits which seem ca have no adaptive function to perform in the otganisms which possess them. Here we come upon a paradox ao biologist should avoid facing. Among tthe major achievemenes of modern biology is the discovery that the heredity which is handed down fram parents to offspring is a constellation af self= reproducing corpuscles of molecular dimensions, genes. Yer, the develope meat of individuals, as well as the evolucion of Mendelian populations, are highly incegrated processes. Even if the genes had each 2 single primary function, such as che production of a single enzyme (which may well be questioned), it would remain true chat their effects ate woven together ‘on the developmental Level into single system which undergoes orderly changes in time. Genes de cot determine each a sepacate traic; they deter~ tine, joincly and severally, the developmencal processes which resule in the appearance of afl the uaice which an organism possesses ac all stages of ies life eycle. ‘The face that the changes known at the locus “‘whice'* ia Drosophila alter the eye colar of the fly dacs nor exclude che possibilicy thar ehis locus hhas other functions as well. As stared above, che white mutane indeed shows biotically imporeane chatacteriscics which ate nor obviously relaced to the color of its eyes. It is even more misleading co say, as is frequently done, chat che eye color is formed by the gene white. Alrbough in order for the pigment to be formed and deposited in the eyes the presence of certain alleles of this gene is recessaty, many athet genes are known which also uence the formation of the sigment and the development of the eyes. Microcephalic idiocy in man may be due ta a mutation in a certain, gene, but ie does aot fallow chac the human head is formed by that gene. Formation of the head ie an outcome of embryonic and poscembryonic developmenc which is influenced by probably all the genes which the ocganism has, and the presence of most and perhaps of atl these genes is indispensable for the development 10 proceed normally. The genes are to some extent independent of each other in she processes of the transmission of hetedicy from parents to offepting, even thovgh studies on position effects and on partial allatism suggese char this independence is noc as complece as classical geneticists liked to think. By contrast, the processes of develop: ment cannot be divided in autonomous components governed by single genes, The verbaliems borrowed from studies on the transmission of heredity ate often misleading shea applied ta the gheaomena of develop~ ment. Ie is unforruasce chat this illegicimate ceansfer of words, and of habits of chought which go with them, is nevertheless # common practice with geneticists. ‘On the population level the genes are carely autonomous. Genes as suct, are subject to natutal selection pethaps only in simplest viruses. Else where ic is not a separate gene but the genarype of which ic is pare thatBad ‘THE AMERICAN NATURALIST is favored o¢ discriminated against oy selection. Moreover, the adaptive value of the zenotype Is manifested in the fitness of the phenotypes which it engenders in ics carciers when che Latter develop ia certain environments. is nor the genotype but « living individual who exhibits a certain succes sion of phenotypes at different stages of ies life, aad who eurvives ot dies and succeeds or fails to reproduce. And, 2s pointed out above, the success for failure ace decided by the competence of the phenotype not mecely ia che adule bur st all stages of the life eycle, No matter how healthy may be an infant or a larva or a seedling, the Daewinian fitness of the organise, ig zero if the adult is inviable or sterile. Conversely, high infaar mortality is a drawback even if che survivors ae vigorous as adults, ADAPTEDNESS AND THE DEVELOPMENT PATTERN ‘The phenotype of an individual may be said to follow a certain trajectory tira; this ecajectory is Aeceemined by the interactions of the genotype with the succession of che environments encounteced in the process of living. At oy givea polar on the trajectory che phenotyne is decided by ies preceding state and by the existiag environment, For example, the pheaotype of a person at s given moment is the outcome of transformations of his past phenorypes and the base of the future ones. The suecession of the changes in a person's phenotype is devetmined, of course, by the developmenc pattern induced by his genotype sad by the successive condi jane of the enviconment. Lec us now consider nat the phenotype as a whote but a wait, such as the weight oF the state of health of the disposition af a persoa, Obviously, my weight, health, and disposition ate at this moment what they were a omen ot several moments ago, except a¢ changed by the ageiag process, by diet, weather, aceupacions, concacts with ather persans, ete. Ihave aoe inbericed aay particular “weight,” since my weight bas vacied greadly during the embryonic and past-enbeyonic life. What [ have inhericed is a genotype which determines, joinely with che succession of the environments, the eajectory whic ny weight has followed thus far, and will follow dusing the renainder of ay life. Sheldon (1954) has ateenpted to trace the trajectories of weights st different agen and for different body heights for all. his somatorypes. Theae eeajcctories apply, as acknowledged by Shetdon himself, co develog- ineoe pactems only in environments which are repatded a5 normal or standard, In other eavironments the weighes aay be either higher o¢ lower, ad ic is aratuieaus to cegatd any weight as inherent of intrinsic for a given somacor type oc genotype. All wcights which a person may have in any environment, notmal of abnormal, salubtions oF morbid, ate equally inherear and fixed by the norm of teaction of a given genatype to 4 given succession of envicon- ents, This in 90 way ceduces the impercance af knowing just what eights individuals of a given somacorype will develap under different circumstances, sisce diffecent people show different development patternsWHAT IS AN ADAPTIVE TRAIT? 30 in environments which are very neatly similar. The same holds far intel= figence or for any other trait. There is oa such thing as “intrinsic intelligence," unless this is arbiteatily defined as that evolved in a ceteain cavionment; neither is there aay known upper or lower limits of intelligence, unteas certain eavicoanents ate considered to be the only possible ones. All this does nat mean that intelligence is aot conditioned genetically; i€ is, for each ser of eavizonmears, Ie would Se invaluable «0 kaow how persons af different genotypes wauld develop in differene envicon meats, and knawing this should be the sim of expecimencs with different educational systems. Although no phenotypic teaie ia any more "ncringic’ chen any other, certain phenotypes exhibit a fitness superior to that of athers of the environments in which the species normally lives, The superior Aevelopmentsl patcerns and superior phenotypic trajectories ate pramated by natural selection, while the inferiar ones fail ta be perperaated. There fore, ceetaia developmental pattems become estat iedividuals of every species oc population as its adaptive nore. Geno~ cynically conditioned deviations from this norm are heredicary diseases and naliornations. Some of them may be "eure?,"* that is, forced back te the normal development trajectory by special environments which are aot re- quired by the genotypes which consticate che adapeive norm. Others are incurable, inasmuch as no eaviconments capable of retecning ehem £9 normal phenotypic trajectories are knows, Fae fact of crucial impezcance is thar che develapmeet patterns which result in adapeively cucperent phenotypes ace homeastaric, They are prot tected by being bulte..i against the disrupeive effects of eovironmeatat changes which ace commonly met with in the habitas ia which che species normally accurs (Lerner 1954, Dabshansky and Wallace 1934, Dobzhansiey and Levene 1956, Lewontin 1954, and others), Rare or abnorial genotypes, such as homozygotes ia 2 normally ouired species, ate often deficient ia homeostatic buflering, Nar is nameaseasis necessarily shows when notmal genntypes are exposed to caviconments which are untsual for the species. ‘As poloced out especially by Schmalhsusea (1949), unasual environments ‘afcen provoke harmful morphoses instead of adaptive madifications, A nan= hereditary disease is, thea, a deviaar phenotype resulting from the exposure fof 2 cormal genotype to an abnotinal enviranienc, of a phenotype brought about by senescence which cesults in 4 gradual loss of hameastatie adjusce hed in a majority of A qualification which must 2¢ made at this poiee is that the adaptive norm need not be menclithic; on the contazy, maay species 2nd populations aze polymorphic, with cwe or more phenarypic norms which may possess superior fienesses in differeat ecatogical niches. Such polymorphic papu> lations thow, thea, twa or more homeostatically buffered developmencal parterns; che development is turaed t0 one of the alketnative teajsctories hy “owivch genes" with clearcut phenatypic effects, or by environmental346 ‘THE AMERICAN NATURALIST stimuli Gather 1953 and others). Sur whether monomorphic ot polymorphic, the developmencal patterns which confer high Darwinian fitness upon theit possessors ace hedged against environmental disturbances which are met ia the usual habiears of the species. ConcLusioxs ‘The cotorious difficulty of discovering the adaptive significance of waits in which races and species of animals ant plants commonly differ is due in parc co an incorrect seazement of the problem. “Ttaits" have no adaptive significance in isolation from cae whole development pactere of the ozgaaism which exhibits chem ae certain stages of ics life cycle. The phenotype of che individual changes celeatlessly from conception to maturity and to death; the path which these changes follow ic set by the genotype interacting with the succession of the eaviconments 0 which an individual is exposed. A “trait” may, chen, he defined as an aspect of all or of a certain portion of the path or the trajectory of the development. However, ic is che eajectory as 2 whole which confers upon the genotype ot the individual its fieness co survive and to reproduce. When the phenotypic manifestations of twa or wore genotypes are com pared, it may appear char they differ in adaptive values because of some few aspects, or aaits, of their developmene pacterns. Yer we muse beware of thinkiog of a developmen: pattern as chough it were a sum of independe ently variable waits, Then we wonder why a proclinace oral bristle has not become reclinate, or vice versa, ia some species of Drosophila, we pliciely assurve thac the position af a bristle is decided by a genetic Factor oF a developmental event which is insereed among other factors and events, bux which is not a par of any organized patcemn of intertelaced processes. Such aq assumption is moce likely to be weong than right. The same assumption is implicit when it is asked why some vestigial organs of the human body have nat disappeared completely, as some of chem have done io other species, of vice verss. We atill possess eraces of che maseles that in other mammals move the ears, still harbor the coccys, an utterly useless selic of the ancestral tail, and some of us are croubled by the veemiform appendix of the coscum, an ergan which, for example, che acs have dispased of encitely. Mammalian embryos are full of organs which are formed seemingly only co disappear without ever functioning: the gill bars and che notachord seem to be used only as weasured examples of the bingenetic law and as “proofs” of the occurrence of evolution. However, a complete disappearance of chese useless organs, both in the adules and in the embryos, would necessitate radical changes in che basic embryocic processes of organ formation snd diffecenciacion, The coccyx is a part of the axial skeleton, and the gill hars atise in the process of formation of the head and the neck. The problem should, therefore, be curned azound, and ie may be asked wher advancages would accrue to the arganism from such radical alterations of ics development. The develapment is moce thaaWHAT IS AN ADAPTIVE TRAIT? 347 summation of “unit chatacters" or unit processes. As emphasized ese pecially by Schmalhausen (1949), development is a highly integrated process, and progressive evalucion from the lower to higher organisms has on che whole cemded to make the incegration more and more thorough, and 60 make the basic developmental processes more and more auronamous from dizect stimuli emanating from the envicoamenc, suNMARY, A wale is a aspect of the whole of of ¢ certain portion of che develap= mental pattern of the organism. An adaptive teait is, then, an aspect of che developmental pattem which facilitates the survival and/or repeaduetion of es carrier in a certain succession of environments. LITERATURE QTED isch, Le C., 1955, Selection in Drosophila pseudoobscunt in relation to enwdiog. Evolution 9: 289309. Borla, Hay da Chcks, A, Bay Cordelro, 4, Ra, Dobahansky, The, Malegolewkio, C., ‘nd. 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