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It may be useful to the reader if we briefly sum up the chief conclusions, which, as far as we can judge, have been fairly well established by the observations given in this volume. All the parts or organs in every plant whilst they continue to grow, and some parts which are provided with pulvini after they have ceased to grow, are continually circumnutating. This movement commences even before the young seedling has broken through the ground. The nature of the movement and its causes, as far as ascertained, have been briefly described in the Introduction. Why every part of a plant whilst it is growing, and in some cases after growth has ceased, should have its cells rendered more turgescent and its cell-walls more extensile first on one side and then on another, thus inducing circumnutation, is not known. It would appear as if the changes in the cells required periods of rest.
In some cases, as with the hypocotyls of Brassica, the leaves of Dionæa and the joints of the Gramineæ, the circumnutating movement when viewed under the microscope is seen to consist of innumerable small oscillations. The part under observation suddenly jerks forwards for a length of ·002 to ·001 of an inch, and then slowly retreats for a part of this distance; after a few seconds it again jerks forwards, but with many intermissions. The retreating movement apparently is due to the elasticity of the resisting tissues.
No one can look at the plants growing on a bank or on the borders of a thick wood, and doubt that the young stems and leaves place themselves so that the leaves may be well illuminated. They are thus enabled to decompose carbonic acid. But the sheath-like cotyledons of some Gramineæ, for instance, those of Phalaris, are not green and contain very little starch; from which fact we may infer that they decompose little or no carbonic acid. Nevertheless, they are extremely heliotropic; and this probably serves them in another way, namely, as a guide from the buried seeds through fissures in the ground or through overlying masses of vegetation, into the light and air. This view is strengthened by the fact that with Phalaris and Avena the first true leaf, which is bright green and no doubt decomposes carbonic acid, exhibits hardly a trace of heliotropism. The heliotropic movements of many other seedlings probably aid them in like manner in emerging from the ground; for apogeotropism by itself would blindly guide them upwards, against any overlying obstacle.
Heliotropism prevails so extensively among the higher plants, that there are extremely few, of which some part, either the stem, flower-peduncle, petiole, or leaf, does not bend towards a lateral light. Drosera rotundifolia is one of the few plants the leaves of which exhibit no trace of heliotropism. Nor could we see any in Dionæa, though the plants were not so carefully observed.
The radicles, hypocotyls and epicotyls of seedling plants, even before they emerge from the ground, and afterwards the cotyledons, are all continually circumnutating. So it is with the stems, stolons, flowerpeduncles, and leaves of older plants. We may, therefore, infer with a considerable degree of safety that all the growing parts of all plants circumnutate. Although this movement, in its ordinary or unmodified state, appears in some cases to be of service to plants, either directly or indirectly–for instance, the circumnutation of the radicle in penetrating the ground, or that of the arched hypocotyl and epicotyl in breaking through the surface–yet circumnutation is so general, or rather so universal a phenomenon, that we cannot suppose it to have been gained for any special purpose. We must believe that it follows in some unknown way from the manner in which vegetable tissues grow.
We shall now consider the many cases in which circumnutation has been modified for various special purposes; that is, a movement already in progress is temporarily increased in some one direction, and temporarily diminished or quite arrested in other directions. These cases may be divided in two sub-classes; in one of which the modification depends on innate or constitutional causes, and is independent of external conditions, excepting in so far that the proper ones for growth must be present. In the second sub-class the modification depends to a large extent on external agencies, such as the daily alternations of light and darkness, or light alone, temperature, or the attraction of gravity.