Ordinary circumnutation is converted into a nyctitropic movement, firstly by an increase in its amplitude, but not to so great a degree as in the case of climbing plants, and secondly by its being rendered periodic in relation to the alternations of day and night. But there is frequently a distinct trace of periodicity in the circumnutating movements of non-sleeping leaves and cotyledons. The fact that nyctitropic movements occur in species distributed in many families throughout the whole vascular series, is intelligible, if they result from the modification of the universally present movement of circumnutation; otherwise the fact is inexplicable.
In the seventh chapter we have given the case of a Porlieria, the leaflets of which remained closed all day, as if asleep, when the plant was kept dry, apparently for the sake of checking evaporation. Something of the same kind occurs with certain Gramineae. At the close of this same chapter, a few observations were appended on what may be called the embryology of leaves. The leaves produced by young shoots on cut-down plants of Melilotus Taurica slept like those of a Trifolium, whilst the leaves on the older branches on the same plants slept in a very different manner, proper to the genus; and from the reasons assigned we are tempted to look at this case as one of reversion to a former nyctitropic habit. So again with Desmodium gyrans, the absence of small lateral leaflets on very young plants, makes us suspect that the immediate progenitor of this species did not possess lateral leaflets, and that their appearance in an almost rudimentary condition at a somewhat more advanced age is the result of reversion to a trifoliate predecessor. However this may be, the rapid circumnutating or [page 563] gyrating movements of the little lateral leaflets, seem to be due proximately to the pulvinus, or organ of movement, not having been reduced nearly so much as the blade, during the successive modifications through which the species has passed.
We now come to the highly important class of movements due to the action of a lateral light. When stems, leaves, or other organs are placed, so that one side is illuminated more brightly than the other, they bend towards the light. This heliotropic movement manifestly results from the modification of ordinary circumnutation; and every gradation between the two movements could be followed. When the light was dim, and only a very little brighter on one side than on the other, the movement consisted of a succession of ellipses, directed towards the light, each of which approached nearer to its source than the previous one. When the difference in the light on the two sides was somewhat greater, the ellipses were drawn out into a strongly-marked zigzag line, and when much greater the course became rectilinear. We have reason to believe that changes in the turgescence of the cells is the proximate cause of the movement of circumnutation; and it appears that when a plant is unequally illuminated on the two sides, the always changing turgescence is augmented along one side, and is weakened or quite arrested along the other sides. Increased turgescence is commonly followed by increased growth, so that a plant which has bent itself towards the light during the day would be fixed in this position were it not for apogeotropism acting during the night. But parts provided with pulvini bend, as Pfeffer has shown, towards the light; and here growth does not come into play any more than in the ordinary circumnutating movements of pulvini. [page 564]
Heliotropism prevails widely throughout the vegetable kingdom, but whenever, from the changed habits of life of any plant, such movements become injurious or useless, the tendency is easily eliminated, as we see with climbing and insectivorous plants.
Apheliotropic movements are comparatively rare in a well-marked degree, excepting with sub-a๋rial roots. In the two cases investigated by us, the movement certainly consisted of modified circumnutation.
The position which leaves and cotyledons occupy during the day, namely, more or less transversely to the direction of the light, is due, according to Frank, to what we call diaheliotropism.