Charles Darwin

“ Any one who will observe a flower-head burying itself, will be convinced that the rocking movement, due to the continued circumnutation of the peduncle, plays an important part in the act. Considering that the flower-heads are very light, that the peduncles are long, thin, and flexible, and that they arise from flexible branches, it is incredible that an object as blunt as one of these flower-heads could penetrate the ground by means of the growing force of the peduncle, unless it were aided by the rocking movement. ”

“ The direction which the apex takes at each successive period of the growth of a root, ultimately determines its whole course; it is therefore highly important that the apex should pursue from the first the most advantageous direction; and we can thus understand why sensitiveness to geotropism, to contact and to moisture, all reside in the tip, and why the tip determines the upper growing part to bend either from or to the exciting cause. ”

“ Circumnutation is of paramount importance in the life of every plant; for it is through its modification that many highly beneficial or necessary movements have been acquired. When light strikes one side of a plant, or light changes into darkness, or when gravitation acts on a displaced part, the plant is enabled in some unknown manner to increase the always varying turgescence of the cells on one side; so that the ordinary circumnutating movement is modified, and the part bends either to or from the exciting cause; or it may occupy a new position, as in the so-called sleep of leaves. ”

“ We do not pretend to say that if a part of a plant, whilst still growing, did not circumnutate—though such a supposition is most improbable—it could not bend towards the light; but, as a matter of fact, heliotropism seems always to consist of modified circumnutation. ”

“ Relation between Circumnutation and Heliotropism.—Any one who will look at the foregoing diagrams, showing the movements of the stems of various plants towards a lateral and more or less dimmed light, will be forced to admit that ordinary circumnutation and heliotropism graduate into one another. When a plant is exposed to a dim lateral light and continues during the whole day bending towards it, receding late in the evening, the movement unquestionably is one of heliotropism. ”

“ Many flowers close at night, and these are likewise said to sleep; but we are not here concerned with their movements, for although effected by the same mechanism as in the case of young leaves, namely, unequal growth on the opposite sides (as first proved by Pfeffer) , yet they differ essentially in being excited chiefly by changes of temperature instead of light; and in being effected, as far as we can judge, for a different purpose. Hardly any one supposes that there is any real analogy between the sleep of animals and that of plants, [2] whether of leaves or flowers. ”

“ Different parts or organs on the same plant, and the same part in different species, are thus excited to act in a widely different manner. We can see no reason why the attraction of gravity should directly modify the state of turgescence and subsequent growth of one part on the upper side and of another part on the lower side. We are therefore led to infer that both geotropic, apogeotropic, and diageotropic movements, the purpose of which we can generally understand, have been acquired for the advantage of the plant by the modification of the ever-present movement of circumnutation. ”

“ Light exerts a powerful influence on most vegetable tissues, and there can be no doubt that it generally tends to check their growth. But when the two sides of a plant are illuminated in a slightly different degree, it does not necessarily follow that the bending towards the illuminated side is caused by changes in the tissues of the same nature as those which lead to increased growth in darkness. We know at least that a part may bend from the light, and yet its growth may not be favoured by light. This is the case with the radicles of Sinapis alba, which are plainly apheliotropic ”

“ Owing to this movement—to the leaves being folded—and to the petioles rising, the whole plant is as much more compact at night than during the day, as a fastigiate Lombardy poplar is compared with any other species of poplar. It is remarkable that when our plants had grown a little older, viz., to a height of 2 or 3 feet, the petioles did not rise at night, and the midribs of the folded leaves were no longer bent back along one side of the petiole. We have noticed in some other genera that the petioles of very young plants rise much more at night than do those of older plants. ”

“ Our seedling now throws up a stem bearing leaves, and often branches, all of which whilst young are continually circumnutating. If we look, for instance, at a great acacia tree, we may feel assured that every one of the innumerable growing shoots is constantly describing small ellipses; as is each petiole, sub-petiole, and leaflet. The latter, as well as ordinary leaves, generally move up and down in nearly the same vertical plane, so that they describe very narrow ellipses. The flower-peduncles are likewise continually circumnutating. ”

“ The whole hypocotyl of a cabbage or the whole leaf of a Dionaea could not jerk forwards unless a very large number of cells on one side were simultaneously affected. Are we to suppose that these cells steadily become more and more turgescent on one side, until the part suddenly yields and bends, inducing what may be called a microscopically minute earthquake in the plant; or do the cells on one side suddenly become turgescent in an intermittent manner; each forward movement thus caused being opposed by the elasticity of the tissues? ”

“ The chief object of the present work is to describe and connect together several large classes of movement, common to almost all plants. The most widely prevalent movement is essentially of the same nature as that of the stem of a climbing plant, which bends successively to all points of the compass, so that the tip revolves. This movement has been called by Sachs “revolving nutation;” but we have found it much more convenient to use the terms circumnutation and circumnutate. ”

“ 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 Dionaea and the joints of the Gramineæ, the circumnutating movement when viewed under the microscope is seen to consist of innumerable small oscillations. ”

“ As the tip of the radicle has been found to be the part which is sensitive to geotropism in the members of such distinct families as the Leguminosae, Malvaceae, Cucurbitaceæ and Gramineæ, we may infer that this character is common to the roots of most seedling plants. Whilst a root is penetrating the ground, the tip must travel first; and we can see the advantage of its being sensitive to geotropism, as it has to determine the course of the whole root. ”

“ As soon as the faintest ray of light reaches a seedling, heliotropism will guide it through any crack in the soil, or through an entangled mass of overlying vegetation; for apogeotropism by itself can direct the seedling only blindly upwards. Hence probably it is that sensitiveness to light resides in the tip of the cotyledons of the Gramineæ, and in the upper part of the hypocotyls of at least some plants. ”

“ Strasburger has shown in his interesting work (‘Wirkung des Lichtes...auf Schwärmsporen,’ 1878) , that the movement of the swarm-spores of various lowly organised plants to a lateral light is influenced by their stage of development, by the temperature to which they are subjected, by the degree of illumination under which they have been raised, and by other unknown causes; so that the swarm-spores of the same species may move across the field of the microscope either to or from the light. Some individuals, moreover, appear to be indifferent to the light ”

“ If they had not possessed and exercised this curious power, they would infallibly have struck against the summit of the shoot and been arrested in their course. As soon as one of these tendrils with its three branches begins to stiffen itself and rise up vertically, the revolving motion becomes more rapid; and as soon as it has passed over the point of difficulty, its motion coinciding with that from its own weight, causes it to fall into its previously inclined position so quickly, that the apex can be seen travelling like the hand of a gigantic clock. ”

“ 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. ”

“ As a pulvinus is formed by the arrestment of the growth of its cells, movements dependent on their action may be long-continued without any increase in length of the part thus provided; and such long-continued movements seem to be one chief end gained by the development of a pulvinus. Long-continued movement would be impossible in any part, without an inordinate increase in its length, if the turgescence of the cells was always followed by growth. ”

“ Most seedling plants are strongly heliotropic, and it is no doubt a great advantage to them in their struggle for life to expose their cotyledons to the light as quickly and as fully as possible, for the sake of obtaining carbon. It has been shown in the first chapter that the greater number of seedlings circumnutate largely and rapidly; and as heliotropism consists of modified circumnutation, we are tempted to look at the high development of these two powers in seedlings as intimately connected. ”

“ When the tip is excited by light (though in the case of radicles this was ascertained in only a single instance) the adjoining part bends from the light; but when excited by gravitation the same part bends towards the centre of gravity. In almost every case we can clearly perceive the final purpose or advantage of the several movements. Two, or perhaps more, of the exciting causes often act simultaneously on the tip, and one conquers the other, no doubt in accordance with its importance for the life of the plant. ”

“ If they had not circumnutated, their tips would have been liable to have been doubled up, as often as they met with obstacles in their path; but as it is, they easily avoid them. This must be a considerable advantage to the plant in spreading from its parent-stock; but we are far from supposing that the power has been gained by the stolons for this purpose, for circumnutation seems to be of universal occurrence with all growing parts; but it is not improbable that the amplitude of the movement may have been specially increased for this purpose. ”

“ Similarly, we are led to believe in adaptation when we see the hypocotyl of a seedling, which contains chlorophyll, bending to the light; for although it thus receives less light, being now shaded by its own cotyledons, it places them—the more important organs—in the best position to be fully illuminated. ”

“ It is not that they cease to circumnutate in darkness, for in all the many cases observed by us they continued to do so; but the normal order of their movements in relation to the alternations of day and night is much disturbed or quite annulled. This holds good with species the cotyledons of which rise or sink so much at night that they may be said to sleep, as well as with others which rise only a little. But different species are affected in very different degrees by changes in the light. ”

“ Having made these few preliminary remarks, we will in imagination take a germinating seed, and consider the part which the various movements play in the life-history of the plant. The first change is the protrusion of the radicle, which begins at once to circumnutate. This movement is immediately modified by the attraction of gravity and rendered geotropic. The radicle, therefore, supposing the seed to be lying on the surface, quickly bends downwards, following a more or less spiral course, as was seen on the smoked glass-plates. Sensitiveness to gravitation resides in the tip ”

“ The common view seems to be that heliotropism is a quite distinct kind of movement from circumnutation; and it may be urged that in the foregoing diagrams we see heliotropism merely combined with, or superimposed on, circumnutation. But if so, it must be assumed that a bright lateral light completely stops circumnutation, for a plant thus exposed moves in a straight line towards it, without describing any ellipses or circles. If the light be somewhat obscured, though amply sufficient to cause the plant to bend towards it, we have more or less plain evidence of still-continued circumnutation. ”

“ The fact of the tip alone being sensitive is so remarkable a fact, that we will here give a brief summary of the foregoing experiments. The tips were cut off 29 horizontally extended radicles of Vicia faba, and with a few exceptions they did not become geotropic in 22 or 23 h., whilst unmutilated radicles were always bowed downwards in 8 or 9 h. It should be borne in mind that the mere act of cutting off the tip of a horizontally extended radicle does not prevent the adjoining parts from bending, if the tip has been previously exposed for an hour or two to the influence of geotropism. ”

“ After a radicle has been deflected by some obstacle, geotropism directs the tip again to grow perpendicularly downwards; but geotropism is a feeble power, and here, as Sachs has shown, another interesting adaptive movement comes into play; for radicles at a distance of a few millimeters from the tip are sensitive to prolonged contact in such a manner that they bend towards the touching object, instead of from it as occurs when an object touches one side of the tip. ”