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When such considerations are fully appreciated it will be realised that medleys of most dissimilar occurrences are all confused together under the term Variation。 One of the first objects of genetic analysis is to disentangle this mass of confusion。
To those who have made no study of heredity it sometimes appears that the question of the effect of conditions in causing variation is one which we should immediately investigate； but a little thought will show that before any critical inquiry into such possibilities can be attempted； a knowledge of the working of heredity under conditions as far as possible uniform must be obtained。 At the time when Darwin was writing； if a plant brought into cultivation gave off an albino variety； such an event was without hesitation ascribed to the change of life。 Now we see that albino GAMETES； germs； that is to say； which are destitute of the pigment…forming factor； may have been originally produced by individuals standing an indefinite number of generations back in the ancestry of the actual albino； and it is indeed almost certain that the variation to which the appearance of the albino is due cannot have taken place in a generation later than that of the grandparents。 It is true that when a new DOMINANT appears we should feel greater confidence that we were witnessing the original variation； but such events are of extreme rarity； and no such case has come under the notice of an experimenter in modern times； as far as I am aware。 That they must have appeared is clear enough。 Nothing corresponding to the Brown… breasted Game fowl is known wild； yet that colour is a most definite dominant； and at some moment since Gallus bankiva was domesticated； the element on which that special colour depends must have at least once been formed in the germ…cell of a fowl； but we need harder evidence than any which has yet been produced before we can declare that this novelty came through over…feeding； or change of climate； or any other disturbance consequent on domestication。 When we reflect on the intricacies of genetic problems as we must now conceive them there come moments when we feel almost thankful that the Mendelian principles were unknown to Darwin。 The time called for a bold pronouncement； and he made it； to our lasting profit and delight。 With fuller knowledge we pass once more into a period of cautious expectation and reserve。
In every arduous enterprise it is pleasanter to look back at difficulties overcome than forward to those which still seem insurmountable； but in the next stage there is nothing to be gained by disguising the fact that the attributes of living things are not what we used to suppose。 If they are more complex in the sense that the properties they display are throughout so regular （I have in view； for example； the marvellous and specific phenomena of regeneration； and those discovered by the students of 〃Entwicklungsmechanik〃。 The circumstances of its occurrence here preclude any suggestion that this regularity has been brought about by the workings of Selection。 The attempts thus to represent the phenomena have resulted in mere parodies of scientific reasoning。） that the Selection of minute random variations is an unacceptable account of the origin of their diversity； yet by virtue of that very regularity the problem is limited in scope and thus simplified。
To begin with； we must relegate Selection to its proper place。 Selection permits the viable to continue and decides that the non…viable shall perish； just as the temperature of our atmosphere decides that no liquid carbon shall be found on the face of the earth： but we do not suppose that the form of the diamond has been gradually achieved by a process of Selection。 So again； as the course of descent branches in the successive generations； Selection determines along which branch Evolution shall proceed； but it does not decide what novelties that branch shall bring forth。 〃La Nature contient le fonds de toutes ces varietes； mais le hazard ou l'art les mettent en oeuvre；〃 as Maupertuis most truly said。
Not till knowledge of the genetic properties of organisms has attained to far greater completeness can evolutionary speculations have more than a suggestive value。 By genetic experiment； cytology and physiological chemistry aiding； we may hope to acquire such knowledge。 In 1872 Nathusius wrote （〃Vortrage uber Viehzucht und Rassenerkenntniss〃； page 120； Berlin； 1872。）： 〃Das Gesetz der Vererbung ist noch nicht erkannt； der Apfel ist noch nicht vom Baum der Erkenntniss gefallen； welcher； der Sage nach； Newton auf den rechten Weg zur Ergrundung der Gravitationsgesetze fuhrte。〃 We cannot pretend that the words are not still true； but in Mendelian analysis the seeds of that apple…tree at last are sown。
If we were asked what discovery would do most to forward our inquiry； what one bit of knowledge would more than any other illuminate the problem； I think we may give the answer without hesitation。 The greatest advance that we can foresee will be made when it is found possible to connect the geometrical phenomena of development with the chemical。 The geometrical symmetry of living things is the key to a knowledge of their regularity； and the forces which cause it。 In the symmetry of the dividing cell the basis of that resemblance we call Heredity is contained。 To imitate the morphological phenomena of life we have to devise a system which can divide。 It must be able to divide； and to segment asgrosslya vibrating plate or rod does； or as an icicle can do as it becomes ribbed in a continuous stream of water； but with this distinction； that the distribution of chemical differences and properties must simultaneously be decided and disposed in orderly relation to the pattern of the segmentation。 Even if a model which would do this could be constructed it might prove to be a useful beginning。
This may be looking too far ahead。 If we had to choose some one piece of more proximate knowledge which we would more especially like to acquire； I suppose we should ask for the secret of interracial sterility。 Nothing has yet been discovered to remove the grave difficulty； by which Huxley in particular was so much oppressed； that among the many varieties produced under domesticationwhich we all regard as analogous to the species seen in natureno clear case of interracial sterility has been demonstrated。 The phenomenon is probably the only one to which the domesticated products seem to afford no parallel。 No solution of the difficulty can be offered which has positive value； but it is perhaps worth considering the facts in the light of modern ideas。 It should be observed that we are not discussing incompatibility of two species to produce offspring （a totally distinct phenomenon）； but the sterility of the offspring which many of them do produce。
When two species； both perfectly fertile severally； produce on crossing a sterile progeny； there is a presumption that the sterility is due to the development in the hybrid of some substance which can only be formed by the meeting of two complementary factors。 That some such account is correct in essence may be inferred from the well…known observation that if the hybrid is not totally sterile but only partially so； and thus is able to form some good germ…cells which develop into new individuals； the sterility of these daughter…individuals is sensibly reduced or may be entirely absent。 The fertility once re…established； the sterility does not return in the later progeny； a fact strongly suggestive of segregation。 Now if the sterility of the cross…bred be really the consequence of the meeting of two complementary factors； we see that the phenomenon could only be produced among the divergent offspring of one species by the acquisition of at least TWO new factors； for if the acquisition of a single factor caused sterility the line would then end。 Moreover each factor must be separately acquired by distinct individuals； for if both were present together； the possessors would by hypothesis be sterile。 And in order to imitate the case of species each of these factors must be acquired by distinct breeds。 The factors need not； and probably would not； produce any other perceptible effects； they might； like the colour…factors present in white flowers； make no difference in the form or other characters。 Not till the cross was actually made between the two complementary individuals would either factor come into play； and the effects even then might be unobserved until an attempt was made to breed from the cross…bred。
Next； if the factors responsible for sterility were acquired； they would in all probability be peculiar to certain individuals and would not readily be distributed to the whole breed。 Any member of the breed also into which BOTH the factors were introduced would drop out of the pedigree by virtue of its sterility。 Hence the evidence that the various domesticated breeds say of dogs or fowls can when mated together produce fertile offspring； is beside the mark。 The real question is； Do they ever produce sterile offspring？ I think the evidence is clearly that sometimes they do； oftener perhaps than is commonly supposed。 These suggestions are quite amenable to e