The Origin of Species by means of Natural Selection (6th ed), Charles Darwin [the beginning after the end read novel .TXT] 📗

one could have been converted into the other, but it by no means follows from this that such gradations have not existed.

As the chelae of Crustaceans resemble in some degree the avicularia of Polyzoa, both serving as pincers, it may be worth while to show that with the former a long series of serviceable gradations still exists. In the first and simplest stage, the terminal segment of a limb shuts down either on the square summit of the broad penultimate segment, or against one whole side, and is thus enabled to catch hold of an object, but the limb still serves as an organ of locomotion. We next find one corner of the broad penultimate segment slightly prominent, sometimes furnished with irregular teeth, and against these the terminal segment shuts down. By an increase in the size of this projection, with its shape, as well as that of the terminal segment, slightly modified and improved, the pincers are rendered more and more perfect, until we have at last an instrument as efficient as the chelae of a lobster. And all these gradations can be actually traced.

Besides the avicularia, the polyzoa possess curious organs called vibracula. These generally consist of long bristles, capable of movement and easily excited. In one species examined by me the vibracula were slightly curved and serrated along the outer margin, and all of them on the same polyzoary often moved simultaneously; so that, acting like long oars, they swept a branch rapidly across the object-glass of my microscope. When a branch was placed on its face, the vibracula became entangled, and they made violent efforts to free themselves. They are supposed to serve as a defence, and may be seen, as Mr. Busk remarks, “to sweep slowly and carefully over the surface of the polyzoary, removing what might be noxious to the delicate inhabitants of the cells when their tentacula are protruded.” The avicularia, like the vibracula, probably serve for defence, but they also catch and kill small living animals, which, it is believed, are afterwards swept by the currents within reach of the tentacula of the zooids. Some species are provided with avicularia and vibracula, some with avicularia alone and a few with vibracula alone.

It is not easy to imagine two objects more widely different in appearance than a bristle or vibraculum, and an avicularium like the head of a bird; yet they are almost certainly homologous and have been developed from the same common source, namely a zooid with its cell. Hence, we can understand how it is that these organs graduate in some cases, as I am informed by Mr.

Busk, into each other. Thus, with the avicularia of several species of Lepralia, the movable mandible is so much produced and is so like a bristle that the presence of the upper or fixed beak alone serves to determine its avicularian nature. The vibracula may have been directly developed from the lips of the cells, without having passed through the avicularian stage; but it seems more probable that they have passed through this stage, as during the early stages of the transformation, the other parts of the cell, with the included zooid, could hardly have disappeared at once. In many cases the vibracula have a grooved support at the base, which seems to represent the fixed beak; though this support in some species is quite absent. This view of the development of the vibracula, if trustworthy, is interesting; for supposing that all the species provided with avicularia had become extinct, no one with the most vivid imagination would ever have thought that the vibracula had originally existed as part of an organ, resembling a bird’s head, or an irregular box or hood. It is interesting to see two such widely different organs developed from a common origin; and as the movable lip of the cell serves as a protection to the zooid, there is no difficulty in believing that all the gradations, by which the lip became converted first into the lower mandible of an avicularium, and then into an elongated bristle, likewise served as a protection in different ways and under different circumstances.

In the vegetable kingdom Mr. Mivart only alludes to two cases, namely the structure of the flowers of orchids, and the movements of climbing plants.

With respect to the former, he says: “The explanation of their ORIGIN is deemed thoroughly unsatisfactory—utterly insufficient to explain the incipient, infinitesimal beginnings of structures which are of utility only when they are considerably developed.” As I have fully treated this subject in another work, I will here give only a few details on one alone of the most striking peculiarities of the flowers of orchids, namely, their pollinia. A pollinium, when highly developed, consists of a mass of pollen-grains, affixed to an elastic footstalk or caudicle, and this to a little mass of extremely viscid matter. The pollinia are by this means transported by insects from one flower to the stigma of another. In some orchids there is no caudicle to the pollen-masses, and the grains are merely tied together by fine threads; but as these are not confined to orchids, they need not here be considered; yet I may mention that at the base of the orchidaceous series, in Cypripedium, we can see how the threads were probably first developed. In other orchids the threads cohere at one end of the pollen-masses; and this forms the first or nascent trace of a caudicle. That this is the origin of the caudicle, even when of considerable length and highly developed, we have good evidence in the aborted pollen-grains which can sometimes be detected embedded within the central and solid parts.

With respect to the second chief peculiarity, namely, the little mass of viscid matter attached to the end of the caudicle, a long series of gradations can be specified, each of plain service to the plant. In most flowers belonging to other orders the stigma secretes a little viscid matter. Now, in certain orchids similar viscid matter is secreted, but in much larger quantities by one alone of the three stigmas; and this stigma, perhaps in consequence of the copious secretion, is rendered sterile. When an insect visits a flower of this kind, it rubs off some of the viscid matter, and thus at the same time drags away some of the pollen-grains.

>From this simple condition, which differs but little from that of a multitude of common flowers, there are endless gradations—to species in which the pollen-mass terminates in a very short, free caudicle—to others in which the caudicle becomes firmly attached to the viscid matter, with the sterile stigma itself much modified. In this latter case we have a pollinium in its most highly developed and perfect condition. He who will carefully examine the flowers of orchids for himself will not deny the existence of the above series of gradations—from a mass of pollen-grains merely tied together by threads, with the stigma differing but little from that of the ordinary flowers, to a highly complex pollinium, admirably adapted for transportal by insects; nor will he deny that all the gradations in the several species are admirably adapted in relation to the general structure of each flower for its fertilisation by different insects. In this, and in almost every other case, the enquiry may be pushed further backwards; and it may be asked how did the stigma of an ordinary flower become viscid, but as we do not know the full history of any one group of beings, it is as useless to ask, as it is hopeless to attempt answering, such questions.

We will now turn to climbing plants. These can be arranged in a long series, from those which simply twine round a support, to those which I have called leaf-climbers, and to those provided with tendrils. In these two latter classes the stems have generally, but not always, lost the power of twining, though they retain the power of revolving, which the tendrils likewise possess. The gradations from leaf-climbers to tendril bearers are wonderfully close, and certain plants may be differently placed in either class. But in ascending the series from simple twiners to leaf-climbers, an important quality is added, namely sensitiveness to a touch, by which means the footstalks of the leaves or flowers, or these modified and converted into tendrils, are excited to bend round and clasp the touching object. He who will read my memoir on these plants will, I think, admit that all the many gradations in function and structure between simple twiners and tendril-bearers are in each case beneficial in a high degree to the species. For instance, it is clearly a great advantage to a twining plant to become a leaf-climber; and it is probable that every twiner which possessed leaves with long footstalks would have been developed into a leaf-climber, if the footstalks had possessed in any slight degree the requisite sensitiveness to a touch.

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 footstalks 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