parent, are matters which have yet to be determined. We do not yet know whether such absolute blending is possible or not, or whether all apparent blending is only a more or less minutely subdivided “mosaic” of non-combinable characters of the parents, in fact whether the combinations due to heredity in reproduction are ever analogous to chemical compounds or are always comparable to particulate mixtures. The attempt to connect Mendel’s observation with the structure of the sperm-cells and egg-cells of plants and animals has already been made. The suggestion is obvious that the halving of the number of nuclear threads in the reproductive cells as compared with the number of those present in the ordinary cells of the tissues—a phenomenon which has now been demonstrated as universal—may be directly connected with the facts of segregation of hybrid characters observed by Mendel. The suggestion requires further experimental testing, for which the case of the parthenogenetic production of a portion of the offspring, in such insects as the bee, offers a valuable opportunity for research.
Another important development of Darwin’s conclusions deserves special notice here, as it is the most distinct advance in the department of bionomics since Darwin’s own writings, and at the same time Variation.touches questions of fundamental interest. The matter strictly relates to the consideration of the “causes of variation,” and is as follows. The fact of variation is a familiar one. No two animals, even of the same brood, are alike: whilst exhibiting a close similarity to their parents, they yet present differences, sometimes very marked differences, from their parents and from one another. Lamarck had put forward the hypothesis that structural alterations acquired by (that is to say, superimposed upon) a parent in the course of its life are transmitted to the offspring, and that, as these structural alterations are acquired by an animal or plant in consequence of the direct action of the environment, the offspring inheriting them would as a consequence not unfrequently start with a greater fitness for those conditions than its parents started with. In its turn, being operated upon by the conditions of life, it would acquire a greater development of the same modification, which it would in turn transmit to its offspring. In the course of several generations, Lamarck argued, a structural alteration amounting to such difference as we call “specific” might be thus acquired. The familiar illustration of Lamarck’s hypothesis is that of the giraffe, whose long neck might, he suggested, have been acquired by the efforts of a primitively short -necked race of herbivores who stretched their necks to reach the foliage of trees in a land where grass was deficient, the effort producing a distinct elongation in the neck of each generation, which was then transmitted to the next. This process is known as “direct adaptation”; and there is no doubt that such structural adaptations are acquired by an animal in the course of its life, though such changes are strictly limited in degree and rare rather than frequent and obvious.
Whether such acquired characters can be transmitted to the next generation is a separate question. It was not proved by Lamarck that they can be, and, indeed, never has been proved by actual observation. Nevertheless it has been assumed, and also indirectly argued, that such acquired characters must be transmitted. Darwin’s great merit was that he excluded from his theory of development any necessary assumption of the transmission of acquired characters. He pointed to the admitted fact of congenital variation, and he showed that congenital variations are arbitrary and, so to speak, non-significant. Causes of congenital variation.Their causes are extremely difficult to trace in detail, but it appears that they are largely due to a “shaking up” of the living matter which constitutes the fertilized germ or embryo-cell, by the process of mixture in it of the substance of two cells—the germ-cell and the sperm-cell—derived from two different individuals. Other mechanical disturbances may assist in this production of congenital variation. Whatever its causes, Darwin showed that it is all-important. In some cases a pair of animals produce ten million offspring, and in such a number a large range of congenital variation is possible. Since on the average only two of the young survive in the struggle for existence to take the place of their two parents, there is a selection out of the ten million young, none of which are exactly alike, and the selection is determined hi nature by the survival of the congenital variety which is fittest to the conditions 'of life. Hence there is no necessity for an assumption of the perpetuation of direct adaptations. The selection of the fortuitously (fortuitously,Transmis-
sion of acquired and inherited characters. that is to say, so far as the conditions of survival are concerned) produced varieties is sufficient, since it is ascertained that they will tend to transmit those characters with which they themselves were born, although it is not ascertained that they could transmit characters acquired on the way through life. A simple illustration of the difference is this: a man born with four fingers only on his right hand is ascertained to be likely to transmit this peculiarity to some at least of his offspring; on the other hand, there is not the slightest ground for supposing that a man who has had one finger chopped off, or has even lost his arm at any period of his life, will produce offspring who are defective in the slightest degree in regard to fingers, hand or arm. Darwin himself, influenced by the consideration of certain classes of facts which seem to favour the Lamarckian hypothesis, was of the opinion that acquired characters are in some cases transmitted. It should be observed, however, that Darwin did not attribute an essential part to this Lamarckian hypothesis of the transmission of acquired characters, but expressly assigned to it an entirely subordinate importance.
The new attitude which has been taken since Darwin’s writings on this question is to ask for evidence of the asserted transmission of acquired characters. It is held[1] that the Darwinian doctrine of selection of fortuitous congenital variations is sufficient to account for all cases, that the Lamarckian hypothesis of transmission of acquired characters is not supported by experimental evidence, and that the latter should therefore be dismissed. Weismann has also ingeniously argued from the structure of the egg-cell and sperm-cell, and from the way in which, and the period at which, they are derived in the course of the growth of the embryo from the egg—from the fertilized egg-cell—that it is impossible (it would be better to say highly improbable) that an alteration in parental structure could produce any exactly representative change in the substance of the germ or sperm-cells.
The one fact which the Lamarckians can produce in their favour is the account of experiments by Brown-Séquard, in which he produced epilepsy in guinea-pigs by section of the large nerves or spinal cord, and in the course of which he was led to believe that in a few rare instances the artificially produced epilepsy and mutilation of the nerves was transmitted. This instance does not stand the test of criticism. The record of Brown-Séquard’s original experiment is not satisfactory, and the subsequent attempts to obtain similar results have not been attended with success. On the other hand, the vast number of experiments in the cropping of the tails and ears of domestic animals, as well as of similar operations on man, are attended with negative results. No case of the transmission of the results of an injury can be produced. Stories of tailless kittens, puppies and calves, born from parents one of whom had been thus injured, are abundant, but they have hitherto entirely failed to stand before examination.
Whilst simple evidence of the fact of the transmission of an acquired character is wanting, the a priori arguments in its favour break down one after another when discussed. The very cases which are advanced as only to be explained on the Lamarckian assumption are found on examination and experiment to be better explained, or only to be explained, by the Darwinian principle. Thus the occurrence of blind animals in caves and in the deep sea was a fact which Darwin himself regarded as best explained by the atrophy of the organ of vision in successive generations through the absence of light and
- ↑ Weismann, Vererbung, &c. (1886).
