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sessors, and therefore cannot have been influenced through natural selection. Bronn adduces the length of the ears and tails in the different species of hares and mice,—the complex folds of enamel in the teeth of many animals, and a multitude of analogous cases. With respect to plants, this subject has been discussed by Nägeli in an admirable essay. He admits that natural selection has effected much, but he insists that the families of plants differ chiefly from each other in morphological characters, which appear to be quite unimportant for the welfare of the species. He consequently believes in an innate tendency towards progressive and more perfect development. He specifies the arrangement of the cells in the tissues, and of the leaves on the axis, as cases in which natural selection could not have acted. To these may be added the numerical divisions in the parts of the flower, the position of the ovules, the shape of the seed, when not of any use for dissemination, &c.
There is much force in the above objection. Nevertheless, we ought, in the first place, to be extremely cautious in pretending to decide what structures now are, or have formerly been, of use to each species. In the second place, it should always be borne in mind that when one part is modified, so will be other parts, through certain dimly seen causes, such as an increased or diminished flow of nutriment to a part, mutual pressure, an early developed part affecting one subsequently developed, and so forth,—as well as through other causes which lead to the many mysterious cases of correlation, which we do not in the least understand. These agencies may be all grouped together, for the sake of brevity, under the expression of the laws of growth. In the
third place, we have to allow for the direct and definite action of changed conditions of life, and for so-called spontaneous variations, in which the nature of the conditions apparently plays a quite subordinate part. Budvariations, such as the appearance of a moss-rose on a common rose, or of a nectarine on a peach-tree, offer good instances of spontaneous variations; but even in these cases, if we bear in mind the power of a minute drop of poison in producing complex galls, we ought not to feel too sure that the above variations are not the effect of some local change in the nature of the sap, due to some change in the conditions. There must be some efficient cause for each slight individual difference, as well as for more strongly marked variations which occasionally arise; and if the unknown cause were to act persistently, it is almost certain that all the individuals of the species would be similarly modified.
In the earlier editions of this work I under-rated, 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 than 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.
It may be worth while to illustrate some of the foregoing remarks. With respect to the assumed inutility of various parts and organs, it is hardly necessary to observe that even in the higher and best-known animals many structures exist, which are so highly developed that no one doubts that they are of importance, yet their
use has not been, or has only recently been, ascertained. As Bronn gives the length of the ears and tail in the several species of mice as instances, though trifling ones, of differences in structure which can be of no special use, I may mention that, according to Dr. Schöbl, the external ears of the common mouse are supplied in an extraordinary manner with nerves, so that they no doubt serve as tactile organs; hence the length of the ears can hardly be quite unimportant. We shall, also, presently see that the tail is a highly useful prehensile organ to some of the species; and its use would be much influenced by its length.
With respect to plants, to which on account of Nägeli's essay I shall confine myself in the following remarks, it will be admitted that the flowers of orchids present a multitude of curious structures, which a few years ago would have been considered as mere morphological differences without any special function; but they are now known to be of the highest importance for the fertilisation of the species through the aid of insects, and have probably been gained through natural selection. No one until lately would have imagined that in dimorphic and trimorphic plants the different lengths of the stamens and pistils, and their arrangement, could have been of any service, but now we know this to be the case.
In certain whole groups of plants the ovules stand erect, and in others they are suspended; and within the same ovarium of some few plants, one ovule holds the former and a second ovule the latter position. These positions seem at first purely morphological, or of no physiological signification; but Dr. Hooker informs me that within the same ovarium, the upper ovules alone in
some cases, and in other cases the lower ones alone are fertilised; and he suggests that this probably depends on the direction in which the pollen-tubes enter the ovarium. If so, the position of the ovules, even when one is erect and the other suspended within the same ovarium, would follow from the selection of any slight deviations in position which favoured their fertilisation, and the production of seed.
Several plants belonging to distinct orders habitually produce flowers of two kinds,—the one open of the ordinary structure, the other closed and imperfect. These two kinds of flowers sometimes differ wonderfully in structure, yet may be seen to graduate into each other on the same plant. The ordinary and open flowers can be intercrossed; and the benefits which certainly are derived from this process are thus secured. The closed and imperfect flowers are, however, manifestly of high importance, as they yield with the utmost safety a large stock of seed, with the expenditure of wonderfully little pollen. The two kinds of flowers often differ much, as just stated, in structure. The petals in the imperfect flowers almost always consist of mere rudiments, and the pollen-grains are reduced in diameter. In Ononis columnæ five of the alternate stamens are rudimentary; and in some species of Viola three stamens are in this state, two retaining their proper function, but being of very small size. In six out of thirty of the closed flowers in an Indian violet (name unknown, for the plants have never produced with me perfect flowers), the sepals are reduced from the normal number of five to three. In one section of the Malpighiaceæ the closed flowers, according to A. de Jussieu, are still further modified, for the five stamens
which stand opposite to the sepals are all aborted, a sixth stamen standing opposite to a petal being alone developed, and this stamen is not present in the ordinary flowers of these species; the style is aborted; and the ovaria are reduced from three to two. Now although natural selection may well have had the power to prevent some of the flowers from expanding, and to reduce the amount of pollen, when rendered by the closure of the flowers superfluous, yet hardly any of the above special modifications can have been thus determined, but must have followed from the laws of growth, including the functional inactivity of parts, during the progress of the reduction of the pollen and the closure of the flowers.
It is so necessary to appreciate the important effects of the laws of growth, that I will give some additional cases of another kind, namely of differences in the same part or organ, due to differences in relative position on the same plant. In the Spanish chestnut, and in certain fir-trees, the angles of divergence of the leaves differ, according to Schacht, in the nearly horizontal and in the upright branches. In the common rue and some other plants, one flower, usually the central or terminal one, opens first, and has five sepals and petals, and five divisions to the ovarium; whilst all the other flowers on the plant are tetramerous. In the British Adoxa the uppermost flower generally has two calyx-lobes with the other organs tetramerous, whilst the surrounding flowers generally have three calyx-lobes with the other organs pentamerous. In many Compositæ and Umbelliferæ (and in some other plants) the circumferential flowers have their corollas much more developed than those of the centre; and this seems often connected with the