As twining is the simplest means of ascending a support, and forms the basis of our series, it may naturally be asked how did plants acquire this power in an incipient degree, afterwards to be improved and increased through natural selection. The power of twining depends, firstly, on the stems while young being extremely flexible (but this is a character common to many plants which are not climbers); and, secondly, on their continually bending to all points of the compass, one after the other in succession, in the same order. By this movement the stems are inclined to all sides, and are made to move round and round. As soon as the lower part of a stem strikes against any object and is stopped, the upper part still goes on bending and revolving, and thus necessarily twines round and up the support. The revolving movement ceases after the early growth of each shoot. As in many widely separated families of plants, single species and single genera possess the power of revolving, and have thus become twiners, they must have independently acquired it, and cannot have inherited it from a common progenitor. Hence, I was led to predict that some slight tendency to a movement of this kind would be found to be far from uncommon with plants which did not climb; and that this had afforded the basis for natural selection to work on and improve. When I made this prediction, I knew of only one imperfect case, namely, of the young flower-peduncles of a Maurandia which revolved slightly and irregularly, like the stems of twining plants, but without making any use of this habit. Soon afterwards Fritz Muller discovered that the young stems of an Alisma and of a Linum-- plants which do not climb and are widely separated in the natural system-- revolved plainly, though irregularly, and he states that he has reason to suspect that this occurs with some other plants. These slight movements appear to be of no service to the plants in question; anyhow, they are not of the least use in the way of climbing, which is the point that concerns us. Nevertheless we can see that if the stems of these plants had been flexible, and if under the conditions to which they are exposed it had profited them to ascend to a height, then the habit of slightly and irregularly revolving might have been increased and utilised through natural selection, until they had become converted into well-developed twining species.

With respect to the sensitiveness of the foot-stalks of the leaves and flowers, and of tendrils, nearly the same remarks are applicable as in the case of the revolving movements of twining plants. As a vast number of species, belonging to widely distinct groups, are endowed with this kind of sensitiveness, it ought to be found in a nascent condition in many plants which have not become climbers. This is the case: I observed that the young flower-peduncles of the above Maurandia curved themselves a little towards the side which was touched. Morren found in several species of Oxalis that the leaves and their foot-stalks moved, especially after exposure to a hot sun, when they were gently and repeatedly touched, or when the plant was shaken. I repeated these observations on some other species of Oxalis with the same result; in some of them the movement was distinct, but was best seen in the young leaves; in others it was extremely slight. It is a more important fact that according to the high authority of Hofmeister, the young shoots and leaves of all plants move after being shaken; and with climbing plants it is, as we know, only during the early stages of growth that the foot-stalks and tendrils are sensitive.

It is scarcely possible that the above slight movements, due to a touch or shake, in the young and growing organs of plants, can be of any functional importance to them. But plants possess, in obedience to various stimuli, powers of movement, which are of manifest importance to them; for instance, towards and more rarely from the light--in opposition to, and more rarely in the direction of, the attraction of gravity. When the nerves and muscles of an animal are excited by galvanism or by the absorption of strychnine, the consequent movements may be called an incidental result, for the nerves and muscles have not been rendered specially sensitive to these stimuli. So with plants it appears that, from having the power of movement in obedience to certain stimuli, they are excited in an incidental manner by a touch, or by being shaken. Hence there is no great difficulty in admitting that in the case of leaf-climbers and tendril-bearers, it is this tendency which has been taken advantage of and increased through natural selection. It is, however, probable, from reasons which I have assigned in my memoir, that this will have occurred only with plants which had already acquired the power of revolving, and had thus become twiners.

I have already endeavoured to explain how plants became twiners, namely, by the increase of a tendency to slight and irregular revolving movements, which were at first of no use to them; this movement, as well as that due to a touch or shake, being the incidental result of the power of moving, gained for other and beneficial purposes. Whether, during the gradual development of climbing plants, natural selection has been aided by the inherited effects of use, I will not pretend to decide; but we know that certain periodical movements, for instance the so-called sleep of plants, are governed by habit.

I have now considered enough, perhaps more than enough, of the cases, selected with care by a skilful naturalist, to prove that natural selection is incompetent to account for the incipient stages of useful structures; and I have shown, as I hope, that there is no great difficulty on this head. A good opportunity has thus been afforded for enlarging a little on gradations of structure, often associated with strange functions--an important subject, which was not treated at sufficient length in the former editions of this work. I will now briefly recapitulate the foregoing cases.

With the giraffe, the continued preservation of the individuals of some extinct high-reaching ruminant, which had the longest necks, legs, etc., and could browse a little above the average height, and the continued destruction of those which could not browse so high, would have sufficed for the production of this remarkable quadruped; but the prolonged use of all the parts, together with inheritance, will have aided in an important manner in their co-ordination. With the many insects which imitate various objects, there is no improbability in the belief that an accidental resemblance to some common object was in each case the foundation for the work of natural selection, since perfected through the occasional preservation of slight variations which made the resemblance at all closer; and this will have been carried on as long as the insect continued to vary, and as long as a more and more perfect resemblance led to its escape from sharp-sighted enemies. In certain species of whales there is a tendency to the formation of irregular little points of horn on the palate; and it seems to be quite within the scope of natural selection to preserve all favourable variations, until the points were converted, first into lamellated knobs or teeth, like those on the beak of a goose--then into short lamellae, like those of the domestic ducks--and then into lamellae, as perfect as those of the shoveller-duck--and finally into the gigantic plates of baleen, as in the mouth of the Greenland whale. In the family of the ducks, the lamellae are first used as teeth, then partly as teeth and partly as a sifting apparatus, and at last almost exclusively for this latter purpose.

With such structures as the above lamellae of horn or whalebone, habit or use can have done little or nothing, as far as we can judge, towards their development. On the other hand, the transportal of the lower eye of a flat-fish to the upper side of the head, and the formation of a prehensile tail, may be attributed almost wholly to continued use, together with inheritance. With respect to the mammae of the higher animals, the most probable conjecture is that primordially the cutaneous glands over the whole surface of a marsupial sack secreted a nutritious fluid; and that these glands were improved in function through natural selection, and concentrated into a confined area, in which case they would have formed a mamma. There is no more difficulty in understanding how the branched spines of some ancient Echinoderm, which served as a defence, became developed through natural selection into tridactyle pedicellariae, than in understanding the development of the pincers of crustaceans, through slight, serviceable modifications in the ultimate and penultimate segments of a limb, which was at first used solely for locomotion. In the avicularia and vibracula of the Polyzoa we have organs widely different in appearance developed from the same source; and with the vibracula we can understand how the successive gradations might have been of service. With the pollinia of orchids, the threads which originally served to tie together the pollen-grains, can be traced cohering into caudicles; and the steps can likewise be followed by which viscid matter, such as that secreted by the stigmas of ordinary flowers, and still subserving nearly but not quite the same purpose, became attached to the free ends of the caudicles--all these gradations being of manifest benefit to the plants in question. With respect to climbing plants, I need not repeat what has been so lately said.

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