FOOTNOTES:

From Darwinism: An Exposition of the Theory of Natural Selection by Alfred Russel Wallace.

[Footnote 198: See the Duke of Argyll's letter in _Nature_, vol. xxxiv. p. 336.]

[Footnote 199: _Journal of the Anthropological Institute,_ vol. xv. pp. 246-260.]

[Footnote 200: The idea of the non-heredity of acquired variations was suggested by the summary of Professor Weismann's views, in _Nature_, referred to later on. But since this chapter was written I have, through the kindness of Mr. E.B. Poulton, seen some of the proofs of the forthcoming translation of Weismann's Essays on Heredity, in which he sets forth an explanation very similar to that here given. On the difficult question of the almost entire disappearance of organs, as in the limbs of snakes and of some lizards, he adduces "a certain form of correlation, which Roux calls 'the struggle of the parts in the organism,'" as playing an important part. Atrophy following disuse is nearly always attended by the corresponding increase of other organs: blind animals possess more developed organs of touch, hearing, and smell; the loss of power in the wings is accompanied by increased strength of the legs, etc. Now as these latter characters, being useful, will be selected, it is easy to understand that a congenital increase of these will be accompanied by a corresponding congenital diminution of the unused organ; and in cases where the means of nutrition are deficient, every diminution of these useless parts will be a gain to the whole organism, and thus their complete disappearance will, in some cases, be brought about directly by natural selection. This corresponds with what we know of these rudimentary organs.

It must, however, be pointed out that the non-heredity of acquired characters was maintained by Mr. Francis Galton more than twelve years ago, on theoretical considerations almost identical with those urged by Professor Weismann; while the insufficiency of the evidence for their hereditary transmission was shown, by similar arguments to those used above and in the work of Professor Weismann already referred to (see "A Theory of Heredity," in _Journ. Anthrop. Instit._, vol. v. pp. 343-345).]

[Footnote 201: This explanation is derived from Weismann's Theory of the Continuity of the Germ-Plasm as summarised in _Nature_.]

[Footnote 202: See a collection of his essays under the title, _The Origin of the Fittest: Essays on Evolution_, D. Appleton and Co. New York. 1887.]

[Footnote 203: _Origin of the Fittest_, p. 174.]

[Footnote 204: _Ibid._ p. 29. It may be here noted that Darwin found these theories unintelligible. In a letter to Professor E.T. Morse in 1877, he writes: "There is one point which I regret you did not make clear in your Address, namely, what is the meaning and importance of Professors Cope and Hyatt's views on acceleration and retardation? I have endeavoured, and given up in despair, the attempt to grasp their meaning" (_Life and Letters_, vol. iii. p. 233).]

[Footnote 205: _Origin of the Fittest_, p. 374.]

[Footnote 206: _Origin of the Fittest_, p. 40.]

[Footnote 207: _The Natural Conditions of Existence as they Affect Animal Life._ London, 1883.]

[Footnote 208: In Dr. Weismann's essay on "Heredity," already referred to, he considers it not improbable that changes in organisms produced by climatic influences may be inherited, because, as these changes do not affect the external parts of an organism only, but often, as in the case of warmth or moisture permeate the whole structure, they may possibly modify the germ-plasm itself, and thus induce variations in the next generation. In this way, he thinks, may possibly be explained the climatic varieties of certain butterflies, and some other changes which seem to be effected by change of climate in a few generations.]

[Footnote 209: This brief indication of Professor Geddes's views is taken from the article "Variation and Selection" in the _Encyclopedia Britannica_, and a paper "On the Nature and Causes of Variation in Plants" in _Trans. and Proc. of the Edinburgh Botanical Society_, 1886; and is, for the most part, expressed in his own words.]

[Footnote 210: Placostylis bovinus, 3½ inches long; Paryphanta Busbyi, 3 in. diam.; P. Hochstetteri, 2¾ in. diam.]

[Footnote 211: The general arguments and objections here set forth will apply with equal force to Professor G. Henslow's theory of the origin of the various forms and structures of flowers as due to "the responsive actions of the protoplasm in consequence of the irritations set up by the weights, pressures, thrusts, tensions, etc., of the insect visitors" (_The Origin of Floral Structures through Insect and other Agencies_, p. 340). On the assumption that acquired characters are inherited, such irritations may have had something to do with the initiation of variations and with the production of certain details of structure, but they are clearly incompetent to have brought about the more important structural and functional modifications of flowers. Such are, the various adjustments of length and position of the stamens to bring the pollen to the insect and from the insect to the stigma; the various motions of stamens and styles at the right time and the right direction; the physiological adjustments bringing about fertility or sterility in heterostyled plants; the traps, springs, and complex movements of various parts of orchids; and innumerable other remarkable phenomena.

For the explanation of these we have no resource but variation and selection, to the effects of which, acting alternately with regression or degradation as above explained (p. 328) must be imputed the development of the countless floral structures we now behold. Even the primitive flowers, whose initiation may, perhaps, have been caused, or rendered possible, by the irritation set up by insects' visits, must, from their very origin, have been modified, in accordance with the supreme law of utility, by means of variation and survival of the fittest.]

[Footnote 212: In an essay on "The Duration of Life," forming part of the translation of Dr. Weismann's papers already referred to, the author still further extends the sphere of natural selection by showing that the average duration of life in each species has been determined by it. A certain length of life is essential in order that the species may produce offspring sufficient to ensure its continuance under the most unfavourable conditions; and it is shown that the remarkable inequalities of longevity in different species and groups may be thus accounted for. Yet more, the occurrence of death in the higher organisms, in place of the continued survival of the unicellular organisms however much they may increase by subdivision, may be traced to the same great law of utility for the race and survival of the fittest. The whole essay is of exceeding interest, and will repay a careful perusal. A similar idea occurred to the present writer about twenty years back, and was briefly noted down at the time, but subsequently forgotten.]

[Footnote 213: The outline here given is derived from two articles in _Nature_, vol. xxxiii. p. 154, and vol. xxxiv. p. 629, in which Weismann's papers are summarised and partly translated.]

[Footnote 214: There are many indications that this explanation of the cause of variation is the true one. Mr. E.B. Poulton suggests one, in the fact that parthenogenetic reproduction only occurs in isolated species, not in groups of related species; as this shows that parthenogenesis cannot lead to the evolution of new forms. Again, in parthenogenetic females the complete apparatus for fertilisation remains unreduced; but if these varied as do sexually produced animals, the organs referred to, being unused, would become rudimentary.

Even more important is the significance of the "polar bodies," as explained by Weismann in one of his _Essays_; since, if his interpretation of them be correct, variability is a necessary consequence of sexual generation.]

[Footnote 215: Darwin's _Animals and Plants_, vol. ii. pp. 23, 24.]

[Footnote 216: In his essay on "Heredity," Dr. Weismann discusses many other cases of supposed inheritance of acquired characters, and shows that they can all be explained in other ways. Shortsightedness among civilised nations, for example, is due partly to the absence of selection and consequent regression towards a mean, and partly to its individual production by constant reading.]

[Footnote 217: Weismann explains instinct on similar lines, and gives many interesting illustrations (see _Essays on Heredity_). He holds "that all instinct is entirely due to the operation of natural selection, and has its foundation, not upon inherited experiences, but upon variations of the germ." Many interesting and difficult cases of instinct are discussed by Darwin in Chapter VIII of the _Origin of Species_, which should be read in connection with the above remarks.

Since this chapter was written my attention has been directed to Mr. Francis Galton's _Theory of Heredity_ (already referred to at p. 417) which was published thirteen years ago as an alternative for Darwin's theory of pangenesis.

Mr. Galton's theory, although it attracted little attention, appears to me to be substantially the same as that of Professor Weismann. Galton's "stirp" is Weismann's "germ-plasm." Galton supposes the sexual elements in the offspring to be directly formed from the residue of the _stirp_ not used up in the development of the body of the parent--Weismann's "continuity of the germ-plasm." Galton also draws many of the same conclusions from his theory. He maintains that characters acquired by the individual as the result of external influences cannot be inherited, unless such influences act directly on the reproductive elements--instancing the possible heredity of alcoholism, because the alcohol permeates the tissues and may reach the sexual elements. He discusses the supposed heredity of effects produced by use or disuse, and explains them much in the same manner as does Weismann. Galton is an anthropologist, and applies the theory, mainly, to explain the peculiarities of hereditary transmission in man, many of which peculiarities he discusses and elucidates. Weismann is a biologist, and is mostly concerned with the application of the theory to explain variation and instinct, and to the further development of the theory of evolution. He has worked it out more thoroughly, and has adduced embryological evidence in its support; but the views of both writers are substantially the same, and their theories were arrived at quite independently. The names of Galton and Weismann should therefore be associated as discoverers of what may be considered (if finally established) the most important contribution to the evolution theory since the appearance of the _Origin of Species_.]

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