It has now been shown that the following important classes of movement all arise from modified circumnutation, which is omnipresent whilst growth lasts, and after growth has ceased, whenever pulvini are present. These classes of movement consist of those due to epinasty and hyponasty,--those proper to climbing plants, commonly called revolving nutation,--the nyctitropic or sleep movements of leaves and cotyledons,--and the two immense classes of movement excited by light and gravitation. When we speak of modified circumnutation we mean that light, or the alternations of light and darkness, gravitation, slight pressure or other irritants, and certain innate or constitutional states of the plant, do not directly cause the movement; they merely lead to a temporary increase or diminution of those spontaneous changes in the turgescence of the cells which are already in progress. In what manner, light, gravitation, etc., act on the cells is not known; and we will here only remark that, if any stimulus affected the cells in such a manner as to cause some slight tendency in the affected part to bend in a beneficial manner, this tendency might easily be increased through the preservation of the more sensitive individuals. But if such bending were injurious, the tendency would be eliminated unless it was overpoweringly strong; for we know how commonly all characters in all organisms vary. Nor can we see any reason to doubt, that after the complete elimination of a tendency to bend in some one direction under a certain stimulus, the power to bend in a directly [page 570] opposite direction might gradually be acquired through natural selection.*

Although so many movements have arisen through modified circumnutation, there are others which appear to have had a quite independent origin; but they do not form such large and important classes. When a leaf of a Mimosa is touched it suddenly assumes the same position as when asleep, but Brucke has shown that this movement results from a different state of turgescence in the cells from that which occurs during sleep; and as sleep-movements are certainly due to modified circumnutation, those from a touch can hardly be thus due. The back of a leaf of Drosera rotundifolia was cemented to the summit of a stick driven into the ground, so that it could not move in the least, and a tentacle was observed during many hours under the microscope; but it exhibited no circumnutating movement, yet after being momentarily touched with a bit of raw meat, its basal part began to curve in 23 seconds. This curving movement therefore could not have resulted from modified circumnutation. But when a small object, such as a fragment of a bristle, was placed on one side of the tip of a radicle, which we know is continually circumnutating, the induced curvature was so similar to the movement caused by geotropism, that we can hardly doubt that it is due to modified circumnutation. A flower of a Mahonia was cemented to a stick, and the stamens exhibited no signs of circumnutation under the microscope, yet when they were lightly touched they suddenly moved towards the pistil. Lastly, the curling of the extremity of a tendril when

* See the remarks in Frank's 'Die wagerechte Richtung von Pflanzentheilen' (1870, pp. 90, 91, etc.), on natural selection in connection with geotropism, heliotropism, etc. [page 571]

touched seems to be independent of its revolving or circumnutating movement. This is best shown by the part which is the most sensitive to contact, circumnutating much less than the lower parts, or apparently not at all.*

Although in these cases we have no reason to believe that the movement depends on modified circumnutation, as with the several classes of movement described in this volume, yet the difference between the two sets of cases may not be so great as it at first appears. In the one set, an irritant causes an increase or diminution in the turgescence of the cells, which are already in a state of change; whilst in the other set, the irritant first starts a similar change in their state of turgescence.

Charles Darwin

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