Whether use and disuse are to be regarded as sources of origin of variations is, again, a matter in which there is wide difference of opinion. But if we admit that any variations can take their origin in the body (as distinguished from the germ), then there is no à priori reason for rejecting use and disuse as factors. As such, we are, I think, justified, in the present state of our knowledge, in reckoning them, at all events, provisionally.

It is clear, however, that they are a proximate, not an ultimate, source of origin. I mean that the structures must be there before they can be either strengthened or weakened by use or disuse. They are at most a source of positive or negative variations of existing structures. They cannot be a direct source of origin of superficial variations. Gain or loss of colour; form-variations not correlated with organic variations;—these cannot be directly due to use or disuse. It is in the nervous and muscular systems and the glandular organs that use and disuse are mainly operative. When, however, organs are brought into relation, or fail to be brought into relation, to their appropriate stimuli, we speak of this, too, as use and disuse. We say, for example, that persistent disuse may impair the essential tissues of the recipient end-organs of the special senses, implying that these tissues require to be brought into continued relation to the appropriate stimuli in order that their efficiency be maintained. So, too, we say that the epidermis is thickened by use, meaning that it is brought into relation with certain mechanical stresses. Through correlation, too, the effects of use and disuse may be widespread. Thus increase in the size of a group of muscles may be correlated with increase in the size of the bones to which they are in relation. In fact, so knit together and co-ordinated is the organism into a unity, it is probable that hardly any variation could take place through use or disuse without modifying to some extent the whole organic being.

Once more, let it be clearly remembered that a large and important school of zoologists reject altogether use or disuse as a factor in variation. They believe that those germs are selected through natural selection in which there is an increased tendency to use or disuse of certain organs. In this, however, we are all agreed. The real question is what is the source of origin of this tendency. On the view of germinal origin, we are forced back on unknown physical or chemical influences in no wise related in origin (though, of course, related in result) with the use or disuse to which they give rise.

So far the main distinction between the two biological schools seems to be that the one, placing the origin of variation in the body-tissues, regards the variations as evoked in direct reaction to physical or chemical influences; while the other, placing the origin of variation in the germ, regards the variations as of fortuitous origin.

I do not use the phrase, "of fortuitous origin," as in any sense discrediting the theory. I am not attempting the cheap artifice of damning a view that does not happen to be my own with a phrase or a nickname. And I therefore hasten to point out what variations I do believe to have had a fortuitous origin. The phrase is often misunderstood, and they will serve to explain its meaning.

If the reader will kindly refer to the tables of variations in the bats' wings (Figs. 14-17), he will see that there are a great number of bones which vary in length and vary independently. And if he will also refer to [Fig. 18], in which seven species of bats are compared, he will see that the differences arise from the increased length of one set of bones in one species and another set of bones in another species. Now, let us suppose that the long, swallow-like wing of the noctule, a high flyer with rapid wing-strokes, that catches insects in full flight, and the broad wings of the horse-shoe, a low flyer, flapping slowly, and, at any rate, sometimes catching insects on the ground, and covering them with its wings as with a net; let us suppose, I say, that to each species its special form of wing is an advantage. Among thousands of independent variations in the lengths of the bones there would be occasional combinations of variations, giving either increased length or increased breadth to the wing. In the noctule, the former would tend to be selected; in the horse-shoe, the latter. Thus the wing of the noctule would be lengthened, and that of the horse-shoe broadened, through the selection of fortuitous combinations of variations which chanced to be favourable. Now, each individual bone-variation is, we believe, due to some special cause; but the fortunate combination is fortuitous, due to what we term "mere chance."

Darwin believed that chance, in this sense, played a very important part in the origin of those favourable variations for which, as he said, natural selection is constantly and unceasingly on the watch. And there can be little question that Darwin was right.

We must now consider very briefly some of the proximate causes of variations. In most of these cases we cannot hope to unravel the nexus of causation. When a plexus of environing circumstances acts upon a highly organized living animal, the most we can do in the present state of knowledge is to note—we cannot hope to explain—the effects produced.

All readers of Darwin's works know well how insistent he was that the nature of the organism is more important than the nature of the environing conditions. "The organization or constitution of the being which is acted on," he says,[EF] "is generally a much more important element than the nature of the changed conditions in determining the nature of the variation." And, again,[EG] "We are thus driven to conclude that in most cases the conditions of life play a subordinate part in causing any particular modification; like that which a spark plays when a mass of combustible matter bursts into flame—the nature of the flame depending on the combustible matter, and not on the spark."

Recent investigations have certainly not lessened the force of Darwin's contention. From which there follows the corollary that the vital condition of the organism is a fact of importance. Darwin was led to believe that among domesticated animals and plants good nutritive conditions were favourable to variation. "Of all the causes which induce variability," he says,[EH] "excess of food, whether or not changed in nature, is probably the most powerful." Darwin also held that the male is more variable than the female—a view that has been especially emphasized by Professor W. K. Brooks. Mr. Wallace, as we have already seen, regards the secondary sexual characters of male birds as the direct outcome of superabundant health and vigour. "There is," he says,[EI] "in the adult male a surplus of strength, vitality, and growth-power which is able to expend itself in this way without injury." And Messrs. Geddes and Thomson contend[EJ] that "brilliancy of colour, exuberance of hair and feathers, activity of scent-glands, and even the development of weapons, are in origin and development outcrops of a male as opposed to a female constitution."