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- The Power of Movement in Plants - 70/98 -


lost; and the whole energy of the plant is expended in rendering the circumnutating movement rectilinear and rapid in one direction alone, namely, towards the light.

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. It must further be assumed that it is only a lateral light which has this extraordinary power of stopping circumnutation, for we know that the several plants above experimented on, and all the others which were observed by us whilst growing, continue to circumnutate, however bright the light may be, if it comes from above. Nor should it be forgotten that in the life of each plant, circumnutation precedes heliotropism, for hypocotyls, epicotyls, and petioles circumnutate before they have broken through the ground and have ever felt the influence of light.

We are therefore fully justified, as it seems to us, in believing that whenever light enters laterally, it is the [page 438] movement of circumnutation which gives rise to, or is converted into, heliotropism and apheliotropism. On this view we need not assume against all analogy that a lateral light entirely stops circumnutation; it merely excites the plant to modify its movement for a time in a beneficial manner. The existence of every possible gradation, between a straight course towards a lateral light and a course consisting of a series of loops or ellipses, becomes perfectly intelligible. Finally, the conversion of circumnutation into heliotropism or apheliotropism, is closely analogous to what takes place with sleeping plants, which during the daytime describe one or more ellipses, often moving in zigzag lines and making little loops; for when they begin in the evening to go to sleep, they likewise expend all their energy in rendering their course rectilinear and rapid. In the case of sleep-movements, the exciting or regulating cause is a difference in the intensity of the light, coming from above, at different periods of the twenty-four hours; whilst with heliotropic and apheliotropic movements, it is a difference in the intensity of the light on the two sides of the plant.

Transversal-heliotropismus (of Frank*) or Diaheliotropism.--The cause of leaves placing themselves more or less transversely to the light, with their upper surfaces directed towards it, has been of late the subject of much controversy. We do not here refer to the object of the movement, which no doubt is that their upper surfaces may be fully illuminated, but the means by which this position is gained. Hardly a better or more simple instance can be given

* 'Die natürliche Wagerechte Richtung von Pflanzentheilen,' 1870. See also some interesting articles by the same author, "Zur Frage über Transversal-Geo-und Heliotropismus," 'Bot. Zeitung,' 1873, p. 17 et seq. [page 439]

of diaheliotropism than that offered by many seedlings, the cotyledons of which are extended horizontally. When they first burst from their seed-coats they are in contact and stand in various positions, often vertically upwards; they soon diverge, and this is effected by epinasty, which, as we have seen, is a modified form of circumnutation. After they have diverged to their full extent, they retain nearly the same position, though brightly illuminated all day long from above, with their lower surfaces close to the ground and thus much shaded. There is therefore a great contrast in the degree of illumination of their upper and lower surfaces, and if they were heliotropic they would bend quickly upwards. It must not, however, be supposed that such cotyledons are immovably fixed in a horizontal position. When seedlings are exposed before a window, their hypocotyls, which are highly heliotropic, bend quickly towards it, and the upper surfaces of their cotyledons still remain exposed at right angles to the light; but if the hypocotyl is secured so that it cannot bend, the cotyledons themselves change their position. If the two are placed in the line of the entering light, the one furthest from it rises up and that nearest to it often sinks down; if placed transversely to the light, they twist a little laterally; so that in every case they endeavour to place their upper surfaces at right angles to the light. So it notoriously is with the leaves on plants nailed against a wall, or grown in front of a window. A moderate amount of light suffices to induce such movements; all that is necessary is that the light should steadily strike the plants in an oblique direction. With respect to the above twisting movement of cotyledons, Frank has given many and much more striking instances in the case of the leaves on [page 440] branches which had been fastened in various positions or turned upside down.

In our observations on the cotyledons of seedling plants, we often felt surprise at their persistent horizontal position during the day, and were convinced before we had read Frank's essay, that some special explanation was necessary. De Vries has shown* that the more or less horizontal position of leaves is in most cases influenced by epinasty, by their own weight, and by apogeotropism. A young cotyledon or leaf after bursting free is brought down into its proper position, as already remarked, by epinasty, which, according to De Vries, long continues to act on the midribs and petioles. Weight can hardly be influential in the case of cotyledons, except in a few cases presently to be mentioned, but must be so with large and thick leaves. With respect to apogeotropism, De Vries maintains that it generally comes into play, and of this fact we shall presently advance some indirect evidence. But over these and other constant forces we believe that there is in many cases, but we do not say in all, a preponderant tendency in leaves and cotyledons to place themselves more or less transversely with respect to the light.

In the cases above alluded to of seedlings exposed to a lateral light with their hypocotyls secured, it is impossible that epinasty, weight and apogeotropism, either in opposition or combined, can be the cause of the rising of one cotyledon, and of the sinking of the other, since the forces in question act equally on both; and since epinasty, weight and apogeotropism all act in a vertical plane, they cannot cause the twisting of the petioles, which occurs in seedlings under the

* 'Arbeiten des Bot. Instituts in Würzburg,' Heft. ii. 1872, pp. 223-277. [page 441]

above conditions of illumination. All these movements evidently depend in some manner on the obliquity of the light, but cannot be called heliotropic, as this implies bending towards the light; whereas the cotyledon nearest to the light bends in an opposed direction or downwards, and both place themselves as nearly as possible at right angles to the light. The movement, therefore, deserves a distinct name. As cotyledons and leaves are continually oscillating up and down, and yet retain all day long their proper position with their upper surfaces directed transversely to the light, and if displaced reassume this position, diaheliotropism must be considered as a modified form of circumnutation. This was often evident when the movements of cotyledons standing in front of a window were traced. We see something analogous in the case of sleeping leaves or cotyledons, which after oscillating up and down during the whole day, rise into a vertical position late in the evening, and on the following morning sink down again into their horizontal or diaheliotropic position, in direct opposition to heliotropism. This return into their diurnal position, which often requires an angular movement of 90o, is analogous to the movement of leaves on displaced branches, which recover their former positions. It deserves notice that any force such as apogeotropism, will act with different degrees of power* in the different positions of those leaves or cotyledons which oscillate largely up and down during the day; and yet they recover their horizontal or diaheliotropic position.

We may therefore conclude that diaheliotropic movements cannot be fully explained by the direct action of light, gravitation, weight, etc., any more

* See former note, in reference to Sachs' remarks on this subject. [page 442]

than can the nyctitropic movements of cotyledons and leaves. In the latter case they place themselves so that their upper surfaces may radiate at night as little as possible into open space, with the upper surfaces of the opposite leaflets often in contact. These movements, which are sometimes extremely complex, are regulated, though not directly caused, by the alternations of light and darkness. In the case of diaheliotropism, cotyledons and leaves place themselves so that their upper surfaces may be exposed to the light, and this movement is regulated, though not directly caused, by the direction whence the light proceeds. In both cases the movement consists of circumnutation modified by innate or constitutional causes, in the same manner as with climbing plants, the circumnutation of which is increased in amplitude and rendered more circular, or again with very young cotyledons and leaves which are thus brought down into a horizontal position by epinasty.

We have hitherto referred only to those leaves and cotyledons which occupy a permanently horizontal position; but many stand more or less obliquely, and some few upright. the cause of these differences of position is not known; but in accordance with Wiesner's views, hereafter to be given, it is probable that some leaves and cotyledons would suffer, if they were fully illuminated by standing at right angles to the light.

We have seen in the second and fourth chapters that those cotyledons and leaves which do not alter their positions at night sufficiently to be said to sleep, commonly rise a little in the evening and fall again on the next morning, so that they stand during the night at a rather higher inclination than during the middle of the day. It is incredible that a rising movement of 2o or 3o, or even of 10o or 20o, can be of [page 443] any service to the plant, so as to have been specially acquired. It must be the result of some periodical change in the conditions to which they are subjected, and there can hardly be a doubt that this is the daily alternations of light and darkness. De Vries states in the paper before referred to, that most petioles and midribs are apogeotropic;* and apogeotropism would account for the above rising movement, which is common to so many widely distinct species, if we suppose it to be conquered by diaheliotropism during the middle of the day, as long as it is of importance to the plant that its cotyledons and leaves should be fully exposed to the light. The exact hour in the afternoon at which they begin to bend slightly upwards, and the extent of the movement, will depend on their degree of sensitiveness to gravitation and on their power of resisting its action during the middle of the day, as well as on the amplitude of their ordinary circumnutating movements; and as these qualities differ much in different species, we might expect that the hour in the afternoon at which they begin to rise would differ much in different species, as is the case. Some other agency, however, besides apogeotropism, must come into play, either directly or indirectly, in this upward movement. Thus a young bean (Vicia faba), growing in a small pot, was placed in front of a window in a klinostat; and at night the leaves rose a little, although

* According to Frank ('Die nat. Wagerechte Richtung von Pflanzentheilen,' 1870, p. 46) the root-leaves of many plants, kept in darkness, rise up and even become vertical; and so it is in some cases with shoots. (See Rauwenhoff, 'Archives Néerlandaises,' tom. xii. p. 32.) These movements indicate apogeotropism; but when organs have been long kept in the dark, the amount of water and of mineral matter which they contain is so much


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