The electric organs, present in several fish, offer a case of special difficulty to the selection theory. When well developed, as in the Torpedo and in Gymnotus, it is conceivable that it may serve as an organ of defence, but in other forms the shock is so weak that it is not to be supposed that it can have any such function. Romanes, who in many ways was one of the stanchest followers of Darwin, admits that, so far as he can see, the evolution of the electric organs cannot be explained by the selection theory. Darwin offers no explanation, but bases his defence on the grounds that we do not know of what use this organ can be to the animal.

Darwin also refers to the phosphorescent, or luminous, organs as a supposed case of difficulty for his theory.

“The luminous organs which occur in a few insects, belonging to widely different families, and which are situated in different parts of the body, offer, under our present state of ignorance, a difficulty almost exactly parallel with that of the electric organs.”

In this case also, as in that of the electric organs, the structures appear in entirely different parts of the body of the insect in different species, so that their occurrence in this group cannot be accounted for on a common descent. In whatever way they have arisen, they must have evolved independently in different species. Darwin advances no explanation of the origin of the luminous organs, but states that they “offer under our present state of ignorance a difficulty almost exactly parallel with that of the electric organs.” It will be noticed that the difficulty referred to rests on the assumption that since the organs are well developed they must have some important use!

We may next consider “organs of little apparent importance as affected by natural selection.” Darwin says:—

“As natural selection acts by life and death,—by the survival of the fittest, and by the destruction of the less well-fitted individuals,—I have sometimes felt great difficulty in understanding the origin or formation of parts of little importance; almost as great, though of a very different kind, as in the case of the most perfect and complex organs.”

His answers to this difficulty are: (1) we are too ignorant “in regard to the whole economy of any one organic being to say what slight modifications would be of importance or not,”—thus such apparently trifling characters as the down on fruit, or the colors of the skin and hair of quadrupeds, which from being correlated with constitutional differences or from determining the attacks of insects might be acted on by natural selection; (2) organs now of trifling importance have in some cases been of high importance to an early progenitor; (3) the changed conditions of life may account for some of the useless organs; (4) reversion accounts for others; (5) the complex laws of growth account for still others, such as correlation, compensation of the pressure of one part on another, etc.; (6) the action of sexual selection is responsible for many characters not to be explained by natural selection. Admitting that there may be cases that can be accounted for on one or the other of these six possibilities, yet there can be no doubt that there are still a considerable number of specific characters that cannot be explained in any of these ways. I do not think that Darwin has by any means met this objection, even if all these six possibilities be admitted as generally valid.

Amongst the “miscellaneous objections” to his theory that Darwin considers we may select the most important cases. The following paragraph has been sometimes quoted by later writers to show that Darwin saw, to a certain extent, the insufficiency of fluctuating variations as a basis for selection. What he calls here “spontaneous variability” refers to sudden and extensive variations, or what we may call discontinuous variations. “In the earlier editions of this work I underrated, as it now seems probable, the frequency and importance of modifications due to spontaneous variability. But it is impossible to attribute to this cause the innumerable structures which are so well adapted to the habits of life of each species. I can no more believe in this, that the well-adapted form of a race-horse or greyhound, which before the principle of selection by man was well understood, excited so much surprise in the minds of the older naturalists, can thus be explained.”

Darwin appears to mean by the latter part of this statement, that he cannot believe that such sudden and great variations as have caused a peach tree to produce nectarines can account for the wonderful adaptations of organisms; but it is not really necessary to suppose that this would often occur, for the same result could be reached by several stages, even if the discontinuous variations had been small, and had appeared in many individuals simultaneously. After showing that in a number of flowers, especially of the Compositæ and Umbelliferæ, the individual flowers in the closely crowded heads are sometimes formed on a different type, Darwin concludes: “In these several cases, with the exception of that of the well-developed ray-florets, which are of service in making the flowers conspicuous to insects, natural selection cannot, as far as we can judge, have come into play, or only in a quite subordinate manner. All these modifications follow from the relative position and interaction of the parts; and it can hardly be doubted that if all the flowers and leaves on the same plant had been subjected to the same external and internal condition, as are the flowers and leaves in certain positions, all would have been modified in the same manner.”

Further on we meet with the following remarkable statement: “But when, from the nature of the organism and of the conditions, modifications have been induced which are unimportant for the welfare of the species, they may be, and apparently often have been, transmitted in nearly the same state to numerous, otherwise modified, descendants. It cannot have been of much importance to the greater number of mammals, birds, or reptiles, whether they were clothed with hair, feathers, or scales; yet hair has been transmitted to almost all mammals, feathers to all birds, and scales to all true reptiles. A structure, whatever it may be, which is common to many allied forms, is ranked by us as of high systematic importance, and consequently is often assumed to be of high vital importance to the species. Thus, as I am inclined to believe, morphological differences, which we consider as important,—such as the arrangement of the leaves, the divisions of the flower or of the ovarium, the position of the ovules, etc.,—first appeared in many cases as fluctuating variations, which sooner or later became constant through the nature of the organism and of the surrounding conditions, as well as through the intercrossing of distinct individuals, but not through natural selection; for as these morphological characters do not affect the welfare of the species, any slight deviations in them could not have been governed or accumulated through this latter agency. It is a strange result which we thus arrive at, namely, that characters of slight vital importance to the species are the most important to the systematist; but, as we shall hereafter see when we treat of the genetic principle of classification, this is by no means so paradoxical as it may at first appear.”