Chapter 8. Modified Circumnutation: Movements Excited By Light

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_ CHAPTER VIII. MODIFIED CIRCUMNUTATION: MOVEMENTS EXCITED BY LIGHT

Distinction between heliotropism and the effects of light on the periodicity of the movements of leaves--Heliotropic movements of Beta, Solanum, Zea, and Avena--Heliotropic movements towards an obscure light in Apios, Brassica, Phalaris, Tropaeolum, and Cassia--Apheliotropic movements of tendrils of Bignonia--Of flower-peduncles of Cyclamen--Burying of the pods--Heliotropism and apheliotropism modified forms of circumnutation-- Steps by which one movement is converted into the other-- Transversal-heliotropismus or diaheliotropism influenced by epinasty, the weight of the part and apogeotropism--Apogeotropism overcome during the middle of the day by diaheliotropism--Effects of the weight of the blades of cotyledons--So called diurnal sleep--Chlorophyll injured by intense light--Movements to avoid intense light

SACHS first clearly pointed out the important difference between the action of light in modifying the periodic movements of leaves, and in causing them to bend towards its source.* The latter, or heliotropic movements are determined by the direction of the light, whilst periodic movements are affected by changes in its intensity and not by its direction. The periodicity of the circumnutating movement often continues for some time in darkness, as we have seen in the last chapter; whilst heliotropic bending ceases very quickly when the light fails. Nevertheless, plants which have ceased through long-continued darkness to move periodically, if re-exposed to the light are still, according to Sachs, heliotropic.

Apheliotropism, or, as usually designated, negative

* 'Physiologie Veg.' (French Translation), 1868, pp. 42, 517, etc.

heliotropism, implies that a plant, when unequally illuminated on the two sides, bends from the light, instead of, as in the last sub-class of cases, towards it; but apheliotropism is comparatively rare, at least in a well-marked degree. There is a third and large sub-class of cases, namely, those of "transversal-Heliotropismus" of Frank, which we will here call diaheliotropism. Parts of plants, under this influence, place themselves more or less transversely to the direction whence the light proceeds, and are thus fully illuminated. There is a fourth sub-class, as far as the final cause of the movement is concerned; for the leaves of some plants when exposed to an intense and injurious amount of light direct themselves, by rising or sinking or twisting, so as to be less intensely illuminated. Such movements have sometimes been called diurnal sleep. If thought advisable, they might be called paraheliotropic, and this term would correspond with our other terms.

It will be shown in the present chapter that all the movements included in these four sub-classes, consist of modified circumnutation. 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. Any kind of movement in relation to light will obviously be much facilitated by each part circumnutating or bending successively in all directions, so that an already existing movement has only to be increased in some one direction, and to be lessened or stopped in the other directions, in order that it should become heliotropic, apheliotropic, etc., as the case may be. In the next chapter some observations on the sensitiveness of plants to light, their rate of bending towards it, and the accuracy with which they point towards its source, etc., will be given. Afterwards it will be shown--and this seems to us a point of much interest--that sensitiveness to light is sometimes confined to a small part of the plant; and that this part when stimulated by light, transmits an influence to distant parts, exciting them to bend.

Heliotropism.--When a plant which is strongly heliotropic (and species differ much in this respect) is exposed to a bright lateral light, it bends quickly towards it, and the course pursued by the stem is quite or nearly straight. But if the light is much dimmed, or occasionally interrupted, or admitted in only a slightly oblique direction, the course pursued is more or less zigzag; and as we have seen and shall again see, such zigzag movement results from the elongation or drawing out of the ellipses, loops, etc., which the plant would have described, if it had been illuminated from above. On several occasions we were much struck with this fact, whilst observing the circumnutation of highly sensitive seedlings, which were unintentionally illuminated rather obliquely, or only at successive intervals of time.

Fig. 168. Beta vulgaris: circumnutation of hypocotyl, deflected by the light being slightly lateral, traced on a horizontal glass from 8.30 A.M. to 5.30 P.M. Direction of the lighted taper by which it was illuminated shown by a line joining the first and penultimate dots. Figure reduced to one-third of the original scale.

[For instance two young seedlings of Beta vulgaris were placed in the middle of a room with north-east windows, and were kept covered up, except during each observation which lasted for only a minute or two; but the result was that their hypocotyls bowed themselves to the side, whence some light occasionally entered, in lines which were only slightly zigzag. Although not a single ellipse was even approximately formed, we inferred from the zigzag lines - and, as it proved, correctly-- that their hypocotyls were circumnutating, for on the following day these same seedlings were placed in a completely darkened room, and were observed each time by the aid of a small wax taper held almost directly above them, and their movements were traced on a horizontal glass above; and now their hypocotyls clearly circumnutated (Fig. 168, and Fig. 39, formerly given, p. 52); yet they moved a short distance towards the side where the taper was held up. If we look at these diagrams, and suppose that the taper had been held more on one side, and that the hypocotyls, still circumnutating, had bent themselves within the same time much more towards the light, long zigzag lines would obviously have been the result.

Fig. 169. Avena sativa: heliotropic movement and circumnutation of sheath-like cotyledon (1 ½ inch in height) traced on horizontal glass from 8 A.M. to 10.25 P.M. Oct. 16th.

Again, two seedlings of Solanum lycopersicum were illuminated from above, but accidentally a little more light entered on one than on any other side, and their hypocotyls became slightly bowed towards the brighter side; they moved in a zigzag line and described in their course two little triangles, as seen in Fig. 37 (p. 50), and in another tracing not given. The sheath-like cotyledons of Zea mays behaved, under nearly similar circumstances, in a nearly similar manner as described in our first chapter (p. 64), for they bowed themselves during the whole day towards one side, making, however, in their course some conspicuous flexures. Before we knew how greatly ordinary circumnutation was modified by a lateral light, some seedling oats, with rather old and therefore not highly sensitive cotyledons, were placed in front of a north-east window, towards which they bent all day in a strongly zigzag course. On the following day they continued to bend in the same direction (Fig. 169), but zigzagged much less. The sky, however, became between 12.40 and 2.35 P.M. overcast with extraordinarily dark thunder-clouds, and it was interesting to note how plainly the cotyledons circumnutated during this interval.

The foregoing observations are of some value, from having been made when we were not attending to heliotropism; and they led us to experiment on several kinds of seedlings, by exposing them to a dim lateral light, so as to observe the gradations between ordinary circumnutation and heliotropism. Seedlings in pots were placed in front of, and about a yard from, a north-east window; on each side and over the pots black boards were placed; in the rear the pots were open to the diffused light of the room, which had a second north-east and a north-west window. By hanging up one or more blinds before the window where the seedlings stood, it was easy to dim the light, so that very little more entered on this side than on the opposite one, which received the diffused light of the room. Late in the evening the blinds were successively removed, and as the plants had been subjected during the day to a very obscure light, they continued to bend towards the window later in the evening than would otherwise have occurred. Most of the seedlings were selected because they were known to be highly sensitive to light, and some because they were but little sensitive, or had become so from having grown old. The movements were traced in the usual manner on a horizontal glass cover; a fine glass filament with little triangles of paper having been cemented in an upright position to the hypocotyls. Whenever the stem or hypocotyl became much bowed towards the light, the latter part of its course had to be traced on a vertical glass, parallel to the window, and at right angles to the horizontal glass cover. Fig. 170. Apios graveolens: heliotropic movement of hypocotyl (.45 of inch in height) towards a moderately bright lateral light, traced on a horizontal glass from 8.30 A.M. to 11.30 A.M. Sept. 18th. Figure reduced to one-third of original scale.

Apios graveolens.--The hypocotyl bends in a few hours rectangularly towards a bright lateral light. In order to ascertain how straight a course it would pursue when fairly well illuminated on one side, seedlings were first placed before a south-west window on a cloudy and rainy morning; and the movement of two hypocotyls were traced for 3 h., during which time they became greatly bowed towards the light. One of these tracings is given on p. 422 (Fig. 170), and the course may be seen to be almost straight. But the amount of light on this occasion was superfluous, for two seedlings were placed before a north-east window, protected by an ordinary linen and two muslin blinds, yet their hypocotyls moved towards this rather dim light in only slightly zigzag lines; but after 4 P.M., as the light waned, the lines became distinctly zigzag. One of these seedlings, moreover, described in the afternoon an ellipse of considerable size, with its longer axis directed towards the window.

We now determined that the light should be made dim enough, so we began by exposing several seedlings before a north-east window, protected by one linen blind, three muslin blinds, and a towel. But so little light entered that a pencil cast no perceptible shadow on a white card, and the hypocotyls did not bend at all towards the window. During this time, from 8.15 to 10.50 A.M., the hypocotyls zigzagged or circumnutated near the same spot, as may be seen at A, in Fig. 171. The towel, therefore, was removed at 10.50 A.M., and replaced by two muslin blinds, and now the light passed through one ordinary linen and four muslin blinds. When a pencil was held upright on a card close to the seedlings, it cast a shadow (pointing from the window) which could only just be distinguished. Yet this very slight excess of light on one side sufficed to cause the hypocotyls of all the seedlings immediately to begin bending in zigzag lines towards the window. The course of one is shown at A (Fig. 171): after moving towards the window from 10.50 A.M. to 12.48 P.M. it bent from the window, and then returned in a nearly parallel line; that is, it almost completed between 12.48 and 2 P.M. a narrow ellipse. Late in the evening, as the light waned, the hypocotyl ceased to bend towards the window, and circumnutated on a small scale round the same spot; during the night it moved considerably backwards, that is, became more upright, through the action of apogeotropism. At B, we have a tracing of the movements of another seedling from the hour (10.50 A.M.) when the towel was removed; and it is in all essential respects similar to the previous one. In these two cases there could be no doubt that the ordinary circumnutating movement of the hypocotyl was modified and rendered heliotropic.

Fig. 171. Apios graveolens: heliotropic movement and circumnutation of the hypocotyls of two seedlings towards a dim lateral light, traced on a horizontal glass during the day. The broken lines show their return nocturnal courses. Height of hypocotyl of A .5, and of B .55 inch. Figure reduced to one-half of original scale.

Brassica oleracea.--The hypocotyl of the cabbage, when not disturbed by a lateral light, circumnutates in a complicated manner over nearly the same space, and a figure formerly given is here reproduced (Fig. 172). If the hypocotyl is exposed to a moderately strong lateral light it moves quickly towards this side, travelling in a straight, or nearly straight, line. But when the lateral light is very dim its course is extremely tortuous, and evidently consists of modified circumnutation. Seedlings were placed before a north-east window, protected by a linen and muslin blind and by a towel. The sky was cloudy, and whenever the clouds grew a little lighter an additional muslin blind was temporarily suspended. The light from the window was

Fig. 172. Brassica oleracea: ordinary circumnutating movement of the hypocotyl of a seedling plant.

thus so much obscured that, judging by the unassisted eye, the seedlings appeared to receive more light from the interior of the room than from the window; but this was not really the case, as was shown by a very faint shadow cast by a pencil on a card. Nevertheless, this extremely small excess of light on one side caused the hypocotyls, which in the morning had stood upright, to bend at right angles towards the window, so that in the evening (after 4.23 P.M.) their course had to be traced on a vertical glass parallel to the window. It should be stated that at 3.30 P.M., by which time the sky had become darker, the towel was removed and replaced by an additional muslin blind, which itself was removed at 4 P.M., the other two blinds being left suspended. In Fig. 173 the course pursued, between 8.9 A.M. and 7.10 P.M., by one of the hypocotyls thus

Fig. 173. Brassica oleracea: heliotropic movement and circumnutation of a hypocotyl towards a very dim lateral light, traced during 11 hours, on a horizontal glass in the morning, and on a vertical glass in the evening. Figure reduced to one-third of the original scale.

exposed is shown. It may be observed that during the first 16 m. the hypocotyl moved obliquely from the light, and this, no doubt, was due to its then circumnutating in this direction. Similar cases were repeatedly observed, and a dim light rarely or never produced any effect until from a quarter to three-quarters of an hour had elapsed. After 5.15 P.M., by which time the light had become obscure, the hypocotyl began to circumnutate about the same spot. The contrast between the two figures (172 and 173) would have been more striking, if they had been originally drawn on the same scale, and had been equally reduced. But the movements shown in Fig. 172 were at first more magnified, and have been reduced to only one-half of the original scale; whereas those in Fig. 173 were at first less magnified, and have been reduced to a one-third scale. A tracing made at the same time with the last of the movements of a second hypocotyl, presented a closely analogous appearance; but it did not bend quite so much towards the light, and it circumnutated rather more plainly.

Fig. 174. Phalaris Canariensis: heliotropic movement and circumnutation of a rather old cotyledon, towards a dull lateral light, traced on a horizontal glass from 8.15 A.M. Sept. 16th to 7.45 A.M. 17th. Figure reduced to one-third of original scale.

Phalaris Canariensis.--The sheath-like cotyledons of this monocotyledonous plant were selected for trial, because they are very sensitive to light and circumnutate well, as formerly shown (see Fig. 49, p. 63). Although we felt no doubt about the result, some seedlings were first placed before a south-west window on a moderately bright morning, and the movements of one were traced. As is so common, it moved for the first 45 m. in a zigzag line; it then felt the full influence of the light, and travelled towards it for the next 2 h. 30 m. in an almost straight line. The tracing has not been given, as it was almost identical with that of Apios under similar circumstances (Fig. 170). By noon it had bowed itself to its full extent; it then circumnutated about the same spot and described two ellipses; by 5 P.M. it had retreated considerably from the light, through the action of apogeotropism. After some preliminary trials for ascertaining the right degree of obscurity, some seedlings were placed (Sept. 16th) before a north-east window, and light was admitted through an ordinary linen and three muslin blinds. A pencil held close by the pot now cast a very faint shadow on a white card, pointing from the window. In the evening, at 4.30 and again at 6 P.M., some of the blinds were removed. In Fig. 174 we see the course pursued under these circumstances by a rather old and not very sensitive cotyledon, 1.9 inch in height, which became much bowed, but was never rectangularly bent towards the light. From 11 A.M., when the sky became rather duller, until 6.30 P.M., the zigzagging was conspicuous, and evidently consisted of drawn-out ellipses. After 6.30 P.M. and during the night, it retreated in a crooked line from the window. Another and younger seedling moved during the same time much more quickly and to a much greater distance, in an only slightly zigzag line towards the light; by 11 A.M. it was bent almost rectangularly in this direction, and now circumnutated about the same place.

Tropaeolum majus.--Some very young seedlings, bearing only two leaves, and therefore not as yet arrived at the climbing stage of growth, were first tried before a north-east window without any blind. The epicotyls bowed themselves towards the light so rapidly that in little more than 3 h. their tips pointed rectangularly towards it. The lines traced were either nearly straight or slightly zigzag; and in this latter case we see that a trace of circumnutation was retained even under the influence of a moderately bright light. Twice whilst these epicotyls were bending towards the window, dots were made every 5 or 6 minutes, in order to detect any trace of lateral movement, but there was hardly any; and the lines formed by their junction were nearly straight, or only very slightly zigzag, as in the other parts of the figures. After the epicotyls had bowed themselves to the full extent towards the light, ellipses of considerable size were described in the usual manner.

After having seen how the epicotyls moved towards a moderately bright light, seedlings were placed at 7.48 A.M. (Sept. 7th) before a north-east window, covered by a towel, and shortly afterwards by an ordinary linen blind, but the epicotyls still moved towards the window. At 9.13 A.M. two additional muslin blinds were suspended, so that the seedlings received very little more light from the window than from the interior of the room. The sky varied in brightness, and the seedlings occasionally

Fig. 175. Tropaeolum majus: heliotropic movement and circumnutation of the epicotyl of a young seedling towards a dull lateral light, traced on a horizontal glass from 7.48 A.M. to 10.40 P.M. Figure reduced to one-half of the original scale.

received for a short time less light from the window than from the opposite side (as ascertained by the shadow cast), and then one of the blinds was temporarily removed. In the evening the blinds were taken away, one by one. the course pursued by an epicotyl under these circumstances is shown in Fig. 175. During the whole day, until 6.45 P.M., it plainly bowed itself towards the light; and the tip moved over a considerable space. After 6.45 P.M. it moved backwards, or from the window, till 10.40 P.M., when the last dot was made. Here, then, we have a distinct heliotropic movement, effected by means of six elongated figures (which if dots had been made every few minutes would have been more or less elliptic) directed towards the light, with the apex of each successive ellipse nearer to the window than the previous one. Now, if the light had been only a little brighter, the epicotyl would have bowed itself more to the light, as we may safely conclude from the previous trials; there would also have been less lateral movement, and the ellipses or other figures would have been drawn out into a strongly marked zigzag line, with probably one or two small loops still formed. If the light had been much brighter, we should have had a slightly zigzag line, or one quite straight, for there would have been more movement in the direction of the light, and much less from side to side.

Fig. 176. Tropaeolum majus: heliotropic movement and circumnutation of an old internode towards a lateral light, traced on a horizontal glass from 8 A.M. Nov. 2nd to 10.20 A.M. Nov. 4th. Broken lines show the nocturnal course.

Sachs states that the older internodes of this Tropaeolum are apheliotropic; we therefore placed a plant, 11 3/4 inches high, in a box, blackened within, but open on one side in front of a north-east window without any blind. A filament was fixed to the third internode from the summit on one plant, and to the fourth internode of another. These internodes were either not old enough, or the light was not sufficiently bright, to induce apheliotropism, for both plants bent slowly towards, instead of from the window during four days. The course, during two days of the first-mentioned internode, is given in Fig. 176; and we see that it either circumnutated on a small scale, or travelled in a zigzag line towards the light. We have thought this case of feeble heliotropism in one of the older internodes of a plant, which, whilst young, is so extremely sensitive to light, worth giving.

Fig. 177. Cassia tora: heliotropic movement and circumnutation of a hypocotyl (1 ½ inch in height) traced on a horizontal glass from 8 A.M. to 10.10 P.M. Oct. 7th. Also its circumnutation in darkness from 7 A.M. Oct. 8th to 7.45 A.M. Oct. 9th.

Cassia tora.--The cotyledons of this plant are extremely sensitive to light, whilst the hypocotyls are much less sensitive than those of most other seedlings, as we had often observed with surprise. It seemed therefore worth while to trace their movements. They were exposed to a lateral light before a north-east window, which was at first covered merely by a muslin blind, but as the sky grew brighter about 11 A.M., an additional linen blind was suspended. After 4 P.M. one blind and then the other was removed. The seedlings were protected on each side and above, but were open to the diffused light of the room in the rear. Upright filaments were fixed to the hypocotyls of two seedlings, which stood vertically in the morning. The accompanying figure (Fig. 177) shows the course pursued by one of them during two days; but it should be particularly noticed that during the second day the seedlings were kept in darkness, and they then circumnutated round nearly the same small space. On the first day (Oct. 7th) the hypocotyl moved from 8 A.M. to 12.23 P.M., toward the light in a zigzag line, then turned abruptly to the left and afterwards described a small ellipse. Another irregular ellipse was completed between 3 P.M. and about 5.30 P.M., the hypocotyl still bending towards the light. The hypocotyl was straight and upright in the morning, but by 6 P.M. its upper half was bowed towards the light, so that the chord of the arc thus formed stood at an angle of 20o with the perpendicular. After 6 P.M. its course was reversed through the action of apogeotropism, and it continued to bend from the window during the night, as shown by the broken line. On the next day it was kept in the dark (excepting when each observation was made by the aid of a taper), and the course followed from 7 A.M. on the 8th to 7.45 A.M. on the 9th is here likewise shown. The difference between the two parts of the figure (177), namely that described during the daytime on the 7th, when exposed to a rather dim lateral light, and that on the 8th in darkness, is striking. The difference consists in the lines during the first day having been drawn out in the direction of the light. The movements of the other seedling, traced under the same circumstances, were closely similar.

Apheliotropism.--We succeeded in observing only two cases of apheliotropism, for these are somewhat rare; and the movements are generally so slow that they would have been very troublesome to trace.

Fig. 178. Bignonia capreolata: apheliotropic movement of a tendril, traced on a horizontal glass from 6.45 A.M. July 19th to 10 A.M. 20th. Movements as originally traced, little magnified, here reduced to two-thirds of the original scale.

Bignonia capreolata.--No organ of any plant, as far as we have seen, bends away so quickly from the light as do the tendrils of this Bignonia. They are also remarkable from circumnutating much less regularly than most other tendrils, often remaining stationary; they depend on apheliotropism for coming into contact with the trunks of trees.* The stem of a young plant was tied to a stick at the base of a pair of fine tendrils, which projected almost vertically upwards; and it was placed in front of a north-east window, being protected on all other sides from the light. The first dot was made at 6.45 A.M., and by 7.35 A.M. both tendrils felt the full influence of the light, for they moved straight away from it until 9.20 A.M., when they circumnutated for a time, still moving, but only a little, from the light (see Fig. 178 of the left-hand tendril). After 3 P.M. they again moved rapidly away from the light in zigzag lines. By a late hour in the evening both had moved so far, that they pointed in a direct line from the light. During the night they returned a little in a nearly opposite direction. On the following morning they again moved from the light and converged, so that by the evening they had become interlocked, still pointing from the light. The right-hand tendril, whilst converging, zigzagged much more than the one figured. Both tracings showed that the apheliotropic movement was a modified form of circumnutation.

Cyclamen Persicum.--Whilst this plant is in flower the peduncles stand upright, but their uppermost part is hooked so that the flower itself hangs downwards. As soon as the pods begin to swell, the peduncles increase much in length and slowly curve downwards, but the short, upper, hooked part straightens itself. Ultimately the pods reach the ground, and if this is covered with moss or dead leaves, they bury themselves. We have often seen saucer-like depressions formed by the pods in damp sand or sawdust; and one pod (.3 of inch in diameter) buried itself in sawdust for three-quarters of its length.** We shall have occasion hereafter to consider the object gained by this burying process. The peduncles can change the direction of their curvature, for if a pot, with plants having their peduncles already bowed downwards, be placed horizontally, they slowly bend at right angles to their former direction towards the centre of the earth. We therefore at first attributed the movement to geotropism; but a pot which had lain horizontally with the pods

* 'The Movements and Habits of Climbing Plants,' 1875, p. 97.

** The peduncles of several other species of Cyclamen twist themselves into a spire, and according to Erasmus Darwin ('Botanic Garden,' Canto., iii. p. 126), the pods forcibly penetrate the earth. See also Grenier and Godron, 'Flore de France,' tom. ii. p. 459.

all pointing to the ground, was reversed, being still kept horizontal, so that the pods now pointed directly upwards; it was then placed in a dark cupboard, but the pods still pointed upwards after four days and nights. The pot, in the same position, was next brought back into the light, and after two days there was some bending downwards of the peduncles, and on the fourth day two of them pointed to the centre of the earth, as did the others after an additional day or two. Another plant, in a pot which had always stood upright, was left in the dark cupboard for six days; it bore 3 peduncles, and only one became within this

Fig. 179. Cyclamen Persicum: downward apheliotropic movement of a flower-peduncle, greatly magnified (about 47 times?), traced on a horizontal glass from 1 P.M. Feb. 18th to 8 A.M. 21st.

time at all bowed downwards, and that doubtfully. The weight, therefore, of the pods is not the cause of the bending down. This pot was then brought back into the light, and after three days the peduncles were considerably bowed downwards. We are thus led to infer that the downward curvature is due to apheliotropism; though more trials ought to have been made.

In order to observe the nature of this movement, a peduncle bearing a large pod which had reached and rested on the ground, was lifted a little up and secured to a stick. A filament was fixed across the pod with a mark beneath, and its movement, greatly magnified, was traced on a horizontal glass during 67 h. The plant was illuminated during the day from above. A copy of the tracing is given on p. 434 (Fig. 179); and there can be no doubt that the descending movement is one of modified circumnutation, but on an extremely small scale. The observation was repeated on another pod, which had partially buried itself in sawdust, and which was lifted up a quarter of an inch above the surface; it described three very small circles in 24 h. Considering the great length and thinness of the peduncles and the lightness of the pods, we may conclude that they would not be able to excavate saucer-like depressions in sand or sawdust, or bury themselves in moss, etc., unless they were aided by their continued rocking or circumnutating movement.]

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. Now, in the case of Tropaeolum (Fig. 175) the stem or epicotyl obviously circumnutated during the whole day, and yet it continued at the same time to move heliotropically; this latter movement being effected by the apex of each successive elongated figure or ellipse standing nearer to the light than the previous one. In the case of Cassia (Fig. 177) the comparison of the movement of the hypocotyl, when exposed to a dim lateral light and to darkness, is very instructive; as is that between the ordinary circumnutating movement of a seedling Brassica (Figs. 172, 173), or that of Phalaris (Figs. 49, 174), and their heliotropic movement towards a window protected by blinds. In both these cases, and in many others, it was interesting to notice how gradually the stems began to circumnutate as the light waned in the evening. We have therefore many kinds of gradations from a movement towards the light, which must be considered as one of circumnutation very slightly modified and still consisting of ellipses or circles,--though a movement more or less strongly zigzag, with loops or ellipses occasionally formed,--to a nearly straight, or even quite straight, heliotropic course.

A plant, when exposed to a lateral light, though this may be bright, commonly moves at first in a zigzag line, or even directly from the light; and this no doubt is due to its circumnutating at the time in a direction either opposite to the source of the light, or more or less transversely to it. As soon, however, as the direction of the circumnutating movement nearly coincides with that of the entering light, the plant bends in a straight course towards the light, if this is bright. The course appears to be rendered more and more rapid and rectilinear, in accordance with the degree of brightness of the light--firstly, by the longer axes of the elliptical figures, which the plant continues to describe as long as the light remains very dim, being directed more or less accurately towards its source, and by each successive ellipse being described nearer to the light. Secondly, if the light is only somewhat dimmed, by the acceleration and increase of the movement towards it, and by the retardation or arrestment of that from the light, some lateral movement being still retained, for the light will interfere less with a movement at right angles to its direction, than with one in its own direction.*

* In his paper, 'Ueber orthotrope und plagiotrope Pflanzentheile' ('Arbeiten des Bot. Inst. in Würzburg,' Band ii. Heft ii. 1879), Sachs has discussed the manner in which geotropism and heliotropism are affected by differences in the angles at which the organs of plants stand with respect to the direction of the incident force.

The result is that the course is rendered more or less zigzag and unequal in rate. Lastly, when the light is very bright all lateral movement is 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 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.

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 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.

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.

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 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 altered, and their regular growth is so much disturbed, that it is perhaps rash to infer from their movements what would occur under normal conditions. (See Godlewski, 'Bot. Zeitung,' Feb. 14th, 1879.)

the action of apogeotropism was quite eliminated. Nevertheless, they did not rise nearly so much at night, as when subjected to apogeotropism. Is it not possible, or even probable, that leaves and cotyledons, which have moved upwards in the evening through the action of apogeotropism during countless generations, may inherit a tendency to this movement? We have seen that the hypocotyls of several Leguminous plants have from a remote period inherited a tendency to arch themselves; and we know that the sleep-movements of leaves are to a certain extent inherited, independently of the alternations of light and darkness.

In our observations on the circumnutation of those cotyledons and leaves which do not sleep at night, we met with hardly any distinct cases of their sinking a little in the evening, and rising again in the morning,--that is, of movements the reverse of those just discussed. We have no doubt that such cases occur, inasmuch as the leaves of many plants sleep by sinking vertically downwards. How to account for the few cases which were observed must be left doubtful. The young leaves of Cannabis sativa sink at night between 30o and 40o beneath the horizon; and Kraus attributes this to epinasty in conjunction with the absorption of water. Whenever epinastic growth is vigorous, it might conquer diaheliotropism in the evening, at which time it would be of no importance to the plant to keep its leaves horizontal. The cotyledons of Anoda Wrightii, of one variety of Gossypium, and of several species of Ipomoea, remain horizontal in the evening whilst they are very young; as they grow a little older they curve a little downwards, and when large and heavy sink so much that they come under our definition of sleep. In the case of the Anoda and of some species of Ipomoea, it was proved that the downward movement did not depend on the weight of the cotyledons; but from the fact of the movement being so much more strongly pronounced after the cotyledons have grown large and heavy, we may suspect that their weight aboriginally played some part in determining that the modification of the circumnutating movement should be in a downward direction.

The so-called Diurnal Sleep of Leaves, Or Paraheliotropism.--This is another class of movements, dependent on the action of light, which supports to some extent the belief that the movements above described are only indirectly due to its action. We refer to the movements of leaves and cotyledons which when moderately illuminated are diaheliotropic; but which change their positions and present their edges to the light, when the sun shines brightly on them. These movements have sometimes been called diurnal sleep, but they differ wholly with respect to the object gained from those properly called nyctitropic; and in some cases the position occupied during the day is the reverse of that during the night.

[It has long been known* that when the sun shines brightly on the leaflets of Robinia, they rise up and present their edges to the light; whilst their position at night is vertically downwards. We have observed the same movement, when the sun shone brightly on the leaflets of an Australian Acacia. Those of Amphicarpaea monoica turned their edges to the sun; and an analogous movement of the little almost rudimentary basal leaflets of Mimosa albida was on one occasion so rapid that it could be distinctly seen through a lens. the elongated, unifoliate, first leaves of Phaseolus Roxburghii stood at 7 A.M. at 20o above the horizon, and no doubt they afterwards sank a little lower. At noon, after having been exposed for about 2 h. to

* Pfeffer gives the names and dates of several ancient writers in his 'Die Periodischen Bewegungen,' 1875, p. 62.

a bright sun, they stood at 56o above the horizon; they were then protected from the rays of the sun, but were left well illuminated from above, and after 30 m. they had fallen 40o, for they now stood at only 16o above the horizon. Some young plants of Phaseolus Hernandesii had been exposed to the same bright sunlight, and their broad, unifoliate, first leaves now stood up almost or quite vertically, as did many of the leaflets on the trifoliate secondary leaves; but some of the leaflets had twisted round on their own axes by as much as 90o without rising, so as to present their edges to the sun. The leaflets on the same leaf sometimes behaved in these two different manners, but always with the result of being less intensely illuminated. These plants were then protected from the sun, and were looked at after 1 ½ h.; and now all the leaves and leaflets had reassumed their ordinary sub-horizontal positions. The copper-coloured cotyledons of some seedlings of Cassia mimosoides were horizontal in the morning, but after the sun had shone on them, each had risen 45 1/2o above the horizon. the movement in these several cases must not be confounded with the sudden closing of the leaflets of Mimosa pudica, which may sometimes be noticed when a plant which has been kept in an obscure place is suddenly exposed to the sun; for in this case the light seems to act, as if it were a touch.

From Prof. Wiesner's interesting observations, it is probable that the above movements have been acquired for a special purpose. the chlorophyll in leaves is often injured by too intense a light, and Prof. Wiesner* believes that it is protected by the most diversified means, such as the presence of hairs, colouring matter, etc., and amongst other means by the leaves presenting their edges to the sun, so that the blades then receive much less light. He experimented on the young leaflets of Robinia, by fixing them in such a position that they could not escape being intensely illuminated, whilst others were allowed to place themselves obliquely; and the former began to suffer from the light in the course of two days.

In the cases above given, the leaflets move either upwards

* 'Die Näturlichen Einrichtungen zum Schutze des Chlorophylls,' etc., 1876. Pringsheim has recently observed under the microscope the destruction of chlorophyll in a few minutes by the action of concentrated light from the sun, in the presence of oxygen. See, also, Stahl on the protection of chlorophyll from intense light, in 'Bot. Zeitung,' 1880.

or twist laterally, so as to place their edges in the direction of the sun's light; but Cohn long ago observed that the leaflets of Oxalis bend downwards when fully exposed to the sun. We witnessed a striking instance of this movement in the very large leaflets of O. Ortegesii. A similar movement may frequently be observed with the leaflets of Averrhoa bilimbi (a member of the Oxalidae); and a leaf is here represented (Fig. 180) on which the sun had shone. A diagram (Fig. 134) was given in the last chapter, representing the oscillations by which a leaflet rapidly descended under these circumstances; and the movement may be seen closely to resemble that (Fig. 133) by

Fig. 180. Averrhoa bilimbi: leaf with leaflets depressed after exposure to sunshine; but the leaflets are sometimes more depressed than is here shown. Figure much reduced.

which it assumed its nocturnal position. It is an interesting fact in relation to our present subject that, as Prof. Batalin informs us in a letter, dated February, 1879, the leaflets of Oxalis acetosella may be daily exposed to the sun during many weeks, and they do not suffer if they are allowed to depress themselves; but if this be prevented, they lose their colour and wither in two or three days. Yet the duration of a leaf is about two months, when subjected only to diffused light; and in this case the leaflets never sink downwards during the day.]

As the upward movements of the leaflets of Robinia, and the downward movements of those of Oxalis, have been proved to be highly beneficial to these plants when subjected to bright sunshine, it seems probable that they have been acquired for the special purpose of avoiding too intense an illumination. As it would have been very troublesome in all the above cases to have watched for a fitting opportunity and to have traced the movement of the leaves whilst they were fully exposed to the sunshine, we did not ascertain whether paraheliotropism always consisted of modified circumnutation; but this certainly was the case with the Averrhoa, and probably with the other species, as their leaves were continually circumnutating. _

Read next: Chapter 9. Sensitiveness Of Plants To Light: Its Transmitted Effects

Read previous: Chapter 7. Modified Circumnutation: Nyctitropic Or Sleep Movements Of Leaves

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