For Darwin there were two modes of racial preparation, (1) natural selection, and (2) the establishment of individually acquired habit. He showed that instincts are subject to hereditary variation; he saw that instincts are also subject to modification through acquisition in the course of individual life. He believed that not only the variations but also, to some extent, the modifications are inherited. He therefore held that some instincts (the greater number) are due to natural selection but that others (less numerous) are due, or partly due, to the inheritance of acquired habits. The latter involve Lamarckian inheritance, which of late years has been the centre of so much controversy. It is noteworthy however that Darwin laid especial emphasis on the fact that many of the most typical and also the most complex instincts--those of neuter insects--do not admit of such an interpretation. "I am surprised," he says ("Origin of Species" (6th edition), page 233.), "that no one has hitherto advanced this demonstrative case of neuter insects, against the well-known doctrine of inherited habit, as advanced by Lamarck." None the less Darwin admitted this doctrine as supplementary to that which was more distinctively his own--for example in the case of the instincts of domesticated animals.
Still, even in such cases, "it may be doubted," he says (Ibid. pages 210, 211.), "whether any one would have thought of training a dog to point, had not some one dog naturally shown a tendency in this line...so that habit and some degree of selection have probably concurred in civilising by inheritance our dogs." But in the interpretation of the instincts of domesticated animals, a more recently suggested hypothesis, that of organic selection (Independently suggested, on somewhat different lines, by Profs.
J. Mark Baldwin, Henry F. Osborn and the writer.), may be helpful.
According to this hypothesis any intelligent modification of behaviour which is subject to selection is probably coincident in direction with an inherited tendency to behave in this fashion. Hence in such behaviour there are two factors: (1) an incipient variation in the line of such behaviour, and (2) an acquired modification by which the behaviour is carried further along the same line. Under natural selection those organisms in which the two factors cooperate are likely to survive. Under artificial selection they are deliberately chosen out from among the rest.
Organic selection has been termed a compromise between the more strictly Darwinian and the Lamarckian principles of interpretation. But it is not in any sense a compromise. The principle of interpretation of that which is instinctive and hereditary is wholly Darwinian. It is true that some of the facts of observation relied upon by Lamarckians are introduced. For Lamarckians however the modifications which are admittedly factors in survival, are regarded as the parents of inherited variations; for believers in organic selection they are only the foster parents or nurses.
It is because organic selection is the direct outcome of and a natural extension of Darwin's cardinal thesis that some reference to it here is justifiable. The matter may be put with the utmost brevity as follows.
(1) Variations (V) occur, some of which are in the direction of increased adaptation (+), others in the direction of decreased adaptation (-). (2)Acquired modifications (M) also occur. Some of these are in the direction of increased accommodation to circumstances (+), while others are in the direction of diminished accommodation (-). Four major combinations are (a) + V with + M, (b) + V with - M, (c) - V with + M, (d) - V with - M.
Of these (d) must inevitably be eliminated while (a) are selected. The predominant survival of (a) entails the survival of the adaptive variations which are inherited. The contributory acquisitions (+M) are not inherited;but they are none the less factors in determining the survival of the coincident variations. It is surely abundantly clear that this is Darwinism and has no tincture of Lamarck's essential principle, the inheritance of acquired characters.
Whether Darwin himself would have accepted this interpretation of some at least of the evidence put forward by Lamarckians is unfortunately a matter of conjecture. The fact remains that in his interpretation of instinct and in allied questions he accepted the inheritance of individually acquired modifications of behaviour and structure.