Popular Science Monthly/Volume 65/October 1904/On the Perception of the Force of Gravity by Plants

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By FRANCIS DARWIN, F.R.S., Fellow of Christ College.


WHEN I had the honor of addressing this association at Cardiff as president of the mother section from which ours has sprung by fission—I spoke of the mechanism of the curvatures commonly known as tropisms. To-day I propose to summarize the evidence—still far from complete—which may help us to form a conception of the mechanism of the stimulus which calls forth one of these movements—namely, geotropism. I have said that the evidence is incomplete, and perhaps I owe you an apology for devoting the time of this section to an unsolved problem. But the making of theories is the romance of research; and I may say, in the words of Diana of the Crossways, who indeed spoke of romance, 'The young who avoid that region escape the title of fool at the cost of a celestial crown.' I am prepared for the risk in the hope that in not avoiding the region of hypothesis I shall at least be able to interest my hearers.

The modern idea of the behavior of plants to their environment has been the growth of the last twenty-five years; though, as Pfeffer has shown, it was clearly stated in 1824 by Dutrochet, who conceived the movements of plants to be 'spontaneous'—i. e., to be executed at the suggestion of changes in the environment, not as the direct and necessary result of such changes. I have been in the habit of expressing the same thought in other words, using the idea of a guide or signal, by the interpretation of which plants are able to make their way successfully through the difficulties of their surroundings. In the existence of the force of gravity we have one of the most striking features of the environment, and in the sensitiveness to gravity which exists in plants we have one of the most widespread cases of a plant reading a signal and directing its growth in relation to its perception. I use the word perception not of course to imply consciousness, but as a convenient form of expression for a form of irritability. It is as though the plant discovered from its sensitiveness to gravity the line of the earth's radius, and then chose a line of growth bearing a certain relation to the vertical line so discovered, either parallel to it or across it at various angles. This, the reaction or reply to the stimulus, is, in my judgment, an adaptive act forced on the species by the struggle for life. This point of view, which, as I regret to think, is not very fashionable, need not trouble us. We are not concerned with why the plant grows up into the air or down into the ground; we are only concerned with the question of how the plant perceives the existence of gravitation. Or, in other words, taking the reaction for granted, what is the nature of the stimulus? If a plant is beaten down by wind or by other causes into a horizontal position, what stimulative change is wrought in the body of the plant by this new posture?

It is conceivable in the case of a stem supported by one end and projecting freely in the air that the unaccustomed state of strain might act as a signal. The tissues on one side (the upper) are stretched, and they are compressed below; this might guide the plant; it might, in fact, have evolved the habit of rapid growth in the compressed side. This is only given as an illustration, for we know that the stimulus does not arise in this way, since such a plant, supported throughout its length, and, therefore, suffering no strain, is geotropically stimulated. The illustration is so far valuable, as it postulates a stimulus produced by weight, and we know from Knight's centrifugal experiment that weight is the governing factor in the conditions. Since we can not believe that the stimulus arises from the strain as affecting the geotropic organ as a whole, we must seek for weight-effects in the individual cell of which the plant is built. We must, in fact, seek for weight-effects on the ectoplasm of those cells which are sensitive to the stimulus of gravity.

If we imagine a plant consisting of a single apogeotropic cell we shall see that the hydrostatic pressure of the cell-contents might serve as a signal.

As long as the cell is vertical the hydrostatic pressure of the cell-sap upon the ectoplasm at C (Fig. 1) is equal to that at D. But the pressure on the basal wall, B, differs from that at A (the apical wall)

PSM V65 D537 Hydrostatic pressure on a single apogeotropic cell.png

Fig. 1.

by the weight of the column AB. If the plant be forced into the horizontal, the pressure at A and B becomes the same, while the pressure at C no longer equals that at D, but differs by the weight of the column CD. Here undoubtedly is a possible means by which the plant could perceive that it was no longer vertical, and would have the means of distinguishing up from down. So that if it were an apogeotropic plant it would need to develop the instinct of relatively accelerated growth on the side D, on which the pressure is greatest.

What is here roughly sketched is the groundwork of the theory of graviperception suggested by Pfeffer and supported by Czapek, which I shall speak of as the radial pressure theory, and to which I shall return later.

It is obvious that there is another consideration to be taken into account, namely, that cells do not contain cell-sap only, but various bodies—nucleus, chloroplasts, crystals, etc.—and that these bodies, differing in specific gravity from the cell-sap, will exert pressure on the physically lower or physically higher cell-walls according as they are heavier or lighter than the cell-sap. Here we have the possibility of a sense-organ for verticality. As long as the stem is vertical and the apex upwards the heavy bodies rest on the basal wall, and the plant is not stimulated to curvature; but if placed horizontally, so that the heavy bodies rest on the lateral cell-walls, which are now horizontal, the plant is stimulated to curve. This is known as the statolith theory. It seems to me quite certain that the stimulus must originate either in the weight of solid particles or in the weight of the fluid in the cells, or by both these means together. And for this reason. Take the statolith theory first. There undoubtedly are heavy bodies in cells; for instance, certain loose, movable starch-grains. Now, either these starch-grains are specialized to serve the purpose of graviperception or they are not. If they are so specialized, cadit quæstio; if they are not, there still remains this interesting point of view; the starch-grains fall to the lower end of the cells in which they occur; therefore, shortly before every geotropic curvature which has taken place since movable starch-grains came into existence, there has been a striking change in the position of these heavy cell contents. Now, if we think of the evolution of geotropism as an adaptive manner of growth we must conceive plants growing vertically upwards and succeeding in life, others not so behaving, and consequently failing. There will be a severe struggle tending to pick out those plants which associated certain curvatures with certain preceding changes, and therefore it seems to me that, if movable starch-grains were originally in no way specialized as part of the machinery of graviperception, they would necessarily become an integral part of that machinery, since the act of geotropism would become adherent to or associated with the falling of the starchgrains.

This argument must in fairness be applied to any other physical conditions which constantly precede geotropic curvature; it is, therefore, not an argument in favor of the statolith theory alone, but equally for the pressure theory, and can not help us to decide between the two points of view.

Are there any general considerations which can help us to decide for or against the statolith theory? I think there are—namely, (1) analogy with the graviperceptive organs of animals; (2) the specialization and distribution of the falling bodies in plants.

Berthold (to whom the credit is due of having first suggested that Dehnecke's falling starch-grains might function as originators of geotropic reaction) is perhaps somewhat bold in saying that 'the primary effect of gravity' as regards stimulation must depend on the passive sinking of the heavier parts. Noll, too, says that Knight's experiment depends on weight, and not the weight of complete parts of the plantbody, but of weight within the irritable structure. I can not see that these downright statements are justified on direct evidence, and I accordingly lay some stress on the support of zoological evidence. It has been conclusively proved by Kreidl's beautiful experiment that in the Crustacean Palæmon the sense of verticality depends on the pressure of heavy bodies on the inside of cavities now known as statocysts, and formerly believed to be organs of hearing. The point of the experiment is that when the normal particles are replaced by fragments of iron the Palæmon reacts towards the attraction of a magnet precisely as it formerly reached towards gravity.

It is unfortunate that Noll's arguments in favor of the existence of a similar mechanism in plants were not at once followed by the demonstration of those easily visible falling bodies, which, in imitation more flattering than accurate, are called statoliths, after the bodies in the statocysts of animals. Personally I was convinced by Kreidl, as quoted by Noll, that here was the key to graviperception in plants. But it was not until the simultaneous appearance of Haberlandt's and Němec's papers that my belief became active, and this, I think, was the case with others. The whole incident is an instance of what my father says somewhere about the difficulty of analyzing the act of belief. I find it impossible to help believing in the statolith theory, though I own to not being able to give a good account of the faith that is in me. It is a fair question whether the analogy drawn from animals gives any support to the theory for plants. The study of sense-organs in plants dates, I think, in its modern development, at least, from my father's work on root-tips, and on the light-perceiving apices of certain seedlings. And the work on the subject is all part of the wave of investigation into adaptations which followed the publication of the 'Origin of Species.' It is very appropriate that one of the two authors to whom we owe the practical working out of the statolith theory should also be one of the greatest living authorities on adaptation in plants. Haberlandt's work on sense-organs, especially on the apparatus for the reception of contact stimuli, is applicable to our present case, since lie has shown that the organs for intensifying the effect of contact are similar in the two kingdoms. No one supposes that the whisker of a cat and the sensitive papilla of a plant are phylogenetically connected. It is a case of what Ray Lankester called homoplastic resemblance. Necessity is the mother of invention, but invention is not infinitely varied, and the same need has led to similar apparatus in beings which have little more in common than that both are living organisms.

But, whether we are or are not affected in our belief by the general argument from analogy, we can not neglect the important fact that Kreidl proves the possibility of gravisensitiveness depending on the possession of statoliths. We must add to this a very important consideration—namely, that we know from Němec's work that an alteration in the position of the statoliths does stimulate the statocyte. Such, at least, is, to my mind, the only conclusion to be drawn from the remarkable accumulation of protoplasm which occurs, for instance, on the basal wall of a normally vertical cell when that wall is cleared of statoliths by temporary horizontality. The fact that a visible disturbance in the plasmic contents of the statocyte follows the disturbance of the starch-grains seems to me a valuable contribution to the evidence.

There is one other set of facts of sufficiently general interest to find a place in this section. I mean Haberlandt's result, also independently arrived at by myself, that when a plant is placed horizontally and rapidly shaken up and down in a vertical plane the gravistimulus is increased. This is readily comprehensible on the statolith theory, since we can imagine the starch-grains would give a greater stimulus if made to vibrate on one of the lateral walls, or if forced into the protoplasm, as Haberlandt supposes. I do not see that the difference in the pressure of the cell-sap on the upper and lower walls (i. e., the lateral walls morphologically considered) would be increased. It would, I imagine, be rendered uneven; but the average difference would remain the same. But in the case of the starch-grains an obvious new feature is introduced by exchanging a stationary condition for one of movement. And though I speak with hesitation on such a point, I am inclined to see in Haberlandt's and my own experiments a means of distinguishing between the pressure and statolith theories. Noll, however, considers that the shaking method is not essentially different from that of Knight's experiment, and adds that the result might have been foreseen.