But apart from the invention of this reasonable hypothesis, which may well, as Huxley estimated, "be the guide of biological and psychological speculation for the next three or four generations," Darwin made a more significant and imperishable contribution. Not for a few generations, but through all ages he should be remembered as the first who showed clearly that the problems of Heredity and Variation are soluble by observation, and laid down the course by which we must proceed to their solution.[61] The moment of inspiration did not come with the reading of Malthus, but with the opening of the "first note-book on Transmutation of Species."[62] Evolution is a process of Variation and Heredity. The older writers, though they had some vague idea that it must be so, did not study Variation and Heredity. Darwin did, and so begat not a theory, but a science.
The extent to which this is true, the scientific world is only beginning to realise. So little was the fact appreciated in Darwin's own time that the success of his writings was followed by an almost total cessation of work in that special field. Of the causes which led to these remarkable consequences I have spoken elsewhere. They proceeded from circumstances peculiar to the time; but whatever the causes there is no doubt that this statement of the result is historically exact, and those who make it their business to collect facts elucidating the physiology of Heredity and Variation are well aware that they will find little to reward their quest in the leading scientific Journals of the Darwinian epoch.
In those thirty years the original stock of evidence current and in circulation even underwent a process of attrition. As in the story of the Eastern sage who first wrote the collected learning of the universe for his sons in a thousand volumes and by successive compression and burning reduced them to one and from this by further burning distilled the single ejaculation of the Faith "There is no god but God and Mohammed is the Prophet of God," which was all his maturer wisdom deemed essential:—so in the books of that period do we find the corpus of genetic knowledge dwindle to a few prerogative instances and these at last to the brief formula of an unquestioned creed.
And yet in all else that concerns biological science this period was, in very truth, our Golden Age, when the natural history of the earth was explored as never before; morphology and embryology were exhaustively ransacked; the physiology of plants and animals began to rival chemistry and physics in precision of method and in the rapidity of its advances; and the foundations of pathology were laid.
In contrast with this immense activity elsewhere the neglect which befel the special physiology of Descent, or Genetics as we now call it, is astonishing. This may of course be interpreted as meaning that the favoured studies seemed to promise a quicker return for effort, but it would be more true to say that those who chose these other pursuits did so without making any such comparison; for the idea that the physiology of Heredity and Variation was a coherent science, offering possibilities of extraordinary discovery, was not present to their minds at all. In a word, the existence of such a science was well nigh forgotten. It is true that in ancillary periodicals, as for example those that treat of entomology or horticulture, or in the writings of the already isolated systematists,[63] observations with this special bearing were from time to time related, but the class of fact on which Darwin built his conceptions of Heredity and Variation was not seen in the highways of biology. It formed no part of the official curriculum of biological students, and found no place among the subjects which their teachers were investigating.
During this period nevertheless one distinct advance was made, that with which Weismann's name is prominently connected. In Darwin's genetic scheme the hereditary transmission of parental experience and its consequences played a considerable role. Exactly how great that role was supposed to be, he with his habitual caution refrained from specifying, for the sufficient reason that he did not know. Nevertheless much of the process of Evolution, especially that by which organs have become degenerate and rudimentary, was certainly attributed by Darwin to such inheritance, though since belief in the inheritance of acquired characters fell into dispute, the fact has been a good deal overlooked. The Origin without "use and disuse" would be a materially different book. A certain vacillation is discernible in Darwin's utterances on this question, and the fact gave to the astute Butler an opportunity for his most telling attack. The discussion which best illustrates the genetic views of the period arose in regard to the production of the rudimentary condition of the wings of many beetles in the Madeira group of islands, and by comparing passages from the Origin[64] Butler convicts Darwin of saying first that this condition was in the main the result of Selection, with disuse aiding, and in another place that the main cause of degeneration was disuse, but that Selection had aided. To Darwin however I think the point would have seemed one of dialetics merely. To him the one paramount purpose was to show that somehow an Evolution by means of Variation and Heredity might have brought about the facts observed, and whether they had come to pass in the one way or the other was a matter of subordinate concern.
To us moderns the question at issue has a diminished significance. For over all such debates a change has been brought by Weismann's challenge for evidence that use and disuse have any transmitted effects at all. Hitherto the transmission of many acquired characteristics had seemed to most naturalists so obvious as not to call for demonstration.[65] Weismann's demand for facts in support of the main proposition revealed at once that none having real cogency could be produced. The time-honoured examples were easily shown to be capable of different explanations. A few certainly remain which cannot be so summarily dismissed, but—though it is manifestly impossible here to do justice to such a subject—I think no one will dispute that these residual and doubtful phenomena, whatever be their true nature, are not of a kind to help us much in the interpretation of any of those complex cases of adaptation which on the hypothesis of unguided Natural Selection are especially difficult to understand. Use and disuse were invoked expressly to help us over these hard places; but whatever changes can be induced in offspring by direct treatment of the parents, they are not of a kind to encourage hope of real assistance from that quarter. It is not to be denied that through the collapse of this second line of argument the Selection hypothesis has had to take an increased and perilous burden. Various ways of meeting the difficulty have been proposed, but these mostly resolve themselves into improbable attempts to expand or magnify the powers of Natural Selection.
Weismann's interpellation, though negative in purpose, has had a lasting and beneficial effect, for through his thorough demolition of the old loose and distracting notions of inherited experience, the ground has been cleared for the construction of a true knowledge of heredity based on experimental fact.
In another way he made a contribution of a more positive character, for his elaborate speculations as to the genetic meaning of cytological appearances have led to a minute investigation of the visible phenomena occurring in those cell-divisions by which germ-cells arise. Though the particular views he advocated have very largely proved incompatible with the observed facts of heredity, yet we must acknowledge that it was chiefly through the stimulus of Weismann's ideas that those advances in cytology were made; and though the doctrine of the continuity of germ-plasm cannot be maintained in the form originally propounded, it is in the main true and illuminating.[66] Nevertheless in the present state of knowledge we are still as a rule quite unable to connect cytological appearances with any genetic consequences and save in one respect (obviously of extreme importance—to be spoken of later) the two sets of phenomena might, for all we can see, be entirely distinct.