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- The Power of Movement in Plants - 3/98 -


the bending of an organ to or from the light; but it is much more convenient to confine the word heliotropism to bending towards the light, and to designate as apheliotropism bending from the light. There is another reason for this change, for writers, as we have observed, occasionally drop the adjectives positive and negative, and thus introduce confusion into their discussions. Diaheliotropism may express a position more or less transverse to the light and induced by it. In like manner positive geotropism, or bending towards the centre of the earth, will be called by us geotropism; apogeotropism will mean bending in opposition to gravity or from the centre of the earth; and diageotropism, a position more or less transverse to the radius of the earth. The words heliotropism and geotropism properly mean the act of moving in relation to the light or the earth; but in the same manner as gravitation, though defined as "the act of tending to the centre," is often used to express the cause of a body falling, so it will be found convenient occasionally to employ heliotropism and geotropism, etc., as the cause of the movements in question.

The term epinasty is now often used in Germany, and implies that the upper surface of an organ grows more quickly than the

* The highly useful terms of Heliotropism and Geotropism were first used by Dr. A. B. Frank: see his remarkable 'Beiträge zur Pflanzenphysiologie,' 1868. [page 6] lower surface, and thus causes it to bend downwards. Hyponasty is the reverse, and implies increased growth along the lower surface, causing the part to bend upwards.*

Methods of Observation.--The movements, sometimes very small and sometimes considerable in extent, of the various organs observed by us, were traced in the manner which after many trials we found to be best, and which must be described. Plants growing in pots were protected wholly from the light, or had light admitted from above, or on one side as the case might require, and were covered above by a large horizontal sheet of glass, and with another vertical sheet on one side. A glass filament, not thicker than a horsehair, and from a quarter to three-quarters of an inch in length, was affixed to the part to be observed by means of shellac dissolved in alcohol. The solution was allowed to evaporate, until it became so thick that it set hard in two or three seconds, and it never injured the tissues, even the tips of tender radicles, to which it was applied. To the end of the glass filament an excessively minute bead of black sealing-wax was cemented, below or behind which a bit of card with a black dot was fixed to a stick driven into the ground. The weight of the filament was so slight that even small leaves were not perceptibly pressed down. another method of observation, when much magnification of the movement was not required, will presently be described. The bead and the dot on the card were viewed through the horizontal or vertical glass-plate (according to the position of the object), and when one exactly covered the other, a dot was made on the glass-plate with a sharply pointed stick dipped in thick Indian-ink. Other dots were made at short intervals of time and these were afterwards joined by straight lines. The figures thus traced were therefore angular; but if dots had been made every 1 or 2 minutes, the lines would have been more curvilinear, as occurred when radicles were allowed to trace their own courses on smoked glass-plates. To make the dots accurately was the sole difficulty, and required some practice. Nor could this be done quite accurately, when the movement was much magnified, such as 30 times and upwards; yet even in this case the general course may be trusted. To test the accuracy of the above method of observation, a filament was fixed to an

* These terms are used in the sense given them by De Vries, 'Würzburg Arbeiten,' Heft ii 1872, p. 252.

[page 7] inanimate object which was made to slide along a straight edge and dots were repeatedly made on a glass-plate; when these were joined, the result ought to have been a perfectly straight line, and the line was very nearly straight. It may be added that when the dot on the card was placed half-an-inch below or behind the bead of sealing-wax, and when the glass-plate (supposing it to have been properly curved) stood at a distance of 7 inches in front (a common distance), then the tracing represented the movement of the bead magnified 15 times.

Whenever a great increase of the movement was not required, another, and in some respects better, method of observation was followed. This consisted in fixing two minute triangles of thin paper, about 1/20 inch in height, to the two ends of the attached glass filament; and when their tips were brought into a line so that they covered one another, dots were made as before on the glass-plate. If we suppose the glass-plate to stand at a distance of seven inches from the end of the shoot bearing the filament, the dots when joined, will give nearly the same figure as if a filament seven inches long, dipped in ink, had been fixed to the moving shoot, and had inscribed its own course on the plate. The movement is thus considerably magnified; for instance, if a shoot one inch in length were bending, and the glass-plate stood at the distance of seven inches, the movement would be magnified eight times. It would, however, have been very difficult to have ascertained in each case how great a length of the shoot was bending; and this is indispensable for ascertaining the degree to which the movement is magnified.

After dots had been made on the glass-plates by either of the above methods, they were copied on tracing paper and joined by ruled lines, with arrows showing the direction of the movement. The nocturnal courses are represented by straight broken lines. the first dot is always made larger than the others, so as to catch the eye, as may be seen in the diagrams. The figures on the glass-plates were often drawn on too large a scale to be reproduced on the pages of this volume, and the proportion in which they have been reduced is always given.* Whenever it could be approximately told how much the movement had been magnified, this is stated. We have perhaps

* We are much indebted to Mr. Cooper for the care with which he has reduced and engraved our diagrams.

[page 8] introduced a superfluous number of diagrams; but they take up less space than a full description of the movements. Almost all the sketches of plants asleep, etc., were carefully drawn for us by Mr. George Darwin.

As shoots, leaves, etc., in circumnutating bend more and more, first in one direction and then in another, they were necessarily viewed at different times more or less obliquely; and as the dots were made on a flat surface, the apparent amount of movement is exaggerated according to the degree of obliquity of the point of view. It would, therefore, have been a much better plan to have used hemispherical glasses, if we had possessed them of all sizes, and if the bending part of the shoot had been distinctly hinged and could have been placed so as to have formed one of the radii of the sphere. But even in this case it would have been necessary afterwards to have projected the figures on paper; so that complete accuracy could not have been attained. From the distortion of our figures, owing to the above causes, they are of no use to any one who wishes to know the exact amount of movement, or the exact course pursued; but they serve excellently for ascertaining whether or not the part moved at all, as well as the general character of the movement.]

In the following chapters, the movements of a considerable number of plants are described; and the species have been arranged according to the system adopted by Hooker in Le Maout and Decaisne's 'Descriptive Botany.' No one who is not investigating the present subject need read all the details, which, however, we have thought it advisable to give. To save the reader trouble, the conclusions and most of the more important parts have been printed in larger type than the other parts. He may, if he thinks fit, read the last chapter first, as it includes a summary of the whole volume; and he will thus see what points interest him, and on which he requires the full evidence.

Finally, we must have the pleasure of returning our [page 9] sincere thanks to Sir Joseph Hooker and to Mr. W. Thiselton Dyer for their great kindness, in not only sending us plants from Kew, but in procuring others from several sources when they were required for our observations; also, for naming many species, and giving us information on various points. [page 10]

CHAPTER I.

THE CIRCUMNUTATING MOVEMENTS OF SEEDLING PLANTS.

Brassica oleracea, circumnutation of the radicle, of the arched hypocotyl whilst still buried beneath the ground, whilst rising above the ground and straightening itself, and when erect--Circumnutation of the cotyledons-- Rate of movement--Analogous observations on various organs in species of Githago, Gossypium, Oxalis, Tropaeolum, Citrus, Aesculus, of several Leguminous and Cucurbitaceous genera, Opuntia, Helianthus, Primula, Cyclamen, Stapelia, Cerinthe, Nolana, Solanum, Beta, Ricinus, Quercus, Corylus, Pinus, Cycas, Canna, Allium, Asparagus, Phalaris, Zea, Avena, Nephrodium, and Selaginella.

THE following chapter is devoted to the circumnutating movements of the radicles, hypocotyls, and cotyledons of seedling plants; and, when the cotyledons do not rise above the ground, to the movements of the epicotyl. But in a future chapter we shall have to recur to the movements of certain cotyledons which sleep at night.

[Brassica oleracea (Cruciferae)'.--Fuller details will be given with respect to the movements in this case than in any other, as space and time will thus ultimately be saved.

Radicle.--A seed with the radicle projecting .05 inch was fastened with shellac to a little plate of zinc, so that the radicle stood up vertically; and a fine glass filament was then fixed near its base, that is, close to the seed-coats. The seed was surrounded by little bits of wet sponge, and the movement of the bead at the end of the filament was traced (Fig. 1) during sixty hours. In this time the radicle increased in length from .05 to .11 inch. Had the filament been attached at first close to the apex of the radicle, and if it could have remained there all the time, the movement exhibited would have [page 11] been much greater, for at the close of our observations the tip, instead of standing vertically upwards, had become bowed downwards through geotropism, so as almost to touch the zinc plate. As far as we could roughly ascertain by measurements made with compasses on other seeds, the tip alone, for a length of only 2/100 to 3/100 of an inch, is acted on by geotropism. But the tracing shows that the basal part of the radicle continued to circumnutate irregularly during the whole time. The actual extreme amount of movement of the bead at the end of the filament was nearly .05 inch, but to what extent the movement of the radicle was magnified by the filament, which was nearly 3/4 inch in length, it was impossible to estimate.

Fig. 1. Brassica oleracea: circumnutation of radicle, traced on horizontal glass, from 9 A.M. Jan. 31st to 9 P.M. Feb. 2nd. Movement of bead at end of filament magnified about 40 times.

Another seed was treated and observed in the same manner, but the radicle in this case protruded .1 inch, and was not Fig. 2. Brassica oleracea: circumnutating and geotropic movement of radicle, traced on horizontal glass during 46 hours.

fastened so as to project quite vertically upwards. The filament was affixed close to its base. The tracing (Fig. 2, reduced by half) shows the


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