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As far back as 1864 (Principles of Biology, Sec. 166) I named in illustration an animal carrying heavy horns–the extinct Irish elk; and indicated the many changes in bones, muscles, blood-vessels, nerves, composing the fore-part of the body, which would be required to make an increment of size in such horns advantageous. Here let me take another instance–that of the giraffe: an instance which I take partly because, in the sixth edition of the Origin of Species, issued in 1872, Mr. Darwin has referred to this animal when effectually disposing of certain arguments urged against his hypothesis. He there says:–

“In order that an animal should acquire some structure specially and largely developed, it is almost indispensable that several other parts should be modified and co-adapted. Although every part of the body varies slightly, it does not follow that the necessary parts should always vary in the right direction and to the right degree” (p. 179).

And in the summary of the chapter, he remarks concerning the adjustments in the same quadruped, that “the prolonged use of all the parts together with inheritance will have aided in an important manner in their co-ordination” (p. 199): a remark probably having reference chiefly to the increased massiveness of the lower part of the neck; the increased size and strength of the thorax required to bear the additional burden; and the increased strength of the fore-legs required to carry the greater weight of both. But now I think that further consideration suggests the belief that the entailed modifications are much more numerous and remote than at first appears; and that the greater part of these are such as cannot be ascribed in any degree to the selection of favourable variations, but must be ascribed exclusively to the inherited effects of changed functions. Whoever has seen a giraffe gallop will long remember the sight as a ludicrous one. The reason for the strangeness of the motions is obvious. Though the fore limbs and the hind limbs differ so much in length, yet in galloping they have to keep pace–must take equal strides. The result is that at each stride, the angle which the hind limbs describe round their centre of motion is much larger than the angle described by the fore limbs. And beyond this, as an aid in equalizing the strides, the hind part of the back is at each stride bent very much downwards and forwards. Hence the hind-quarters appear to be doing nearly all the work. Now a moment’s observation shows that the bones and muscles composing the hind-quarters of the giraffe, perform actions differing in one or other way and degree, from the actions performed by the homologous bones and muscles in a mammal of ordinary proportions, and from those in the ancestral mammal which gave origin to the giraffe. Each further stage of that growth which produced the large fore-quarters and neck, entailed some adapted change in sundry of the numerous parts composing the hind-quarters; since any failure in the adjustment of their respective strengths would entail some defect in speed and consequent loss of life when chased. It needs but to remember how, when continuing to walk with a blistered foot, the taking of steps in such a modified way as to diminish pressure on the sore point, soon produces aching of muscles which are called into unusual action, to see that over-straining of any one of the muscles of the giraffe’s hind-quarters might quickly incapacitate the animal when putting out all its powers to escape; and to be a few yards behind others would cause death. Hence if we are debarred from assuming that co-operative parts vary together even when adjacent and closely united–if we are still more debarred from assuming that with increased length of fore-legs or of neck, there will go an appropriate change in any one muscle or bone in the hind-quarters; how entirely out of the question it is to assume that there will simultaneously take place the appropriate changes in all those many components of the hind-quarters which severally require re-adjustment. It is useless to reply that an increment of length in the fore-legs or neck might be retained and transmitted to posterity, waiting an appropriate variation in a particular bone or muscle in the hind-quarters, which, being made, would allow of a further increment. For besides the fact that until this secondary variation occurred the primary variation would be a disadvantage often fatal; and besides the fact that before such an appropriate secondary variation might be expected in the course of generations to occur, the primary variation would have died out; there is the fact that the appropriate variation of one bone or muscle in the hind-quarters would be useless without appropriate variations of all the rest–some in this way and some in that–a number of appropriate variations which it is impossible to suppose.