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- The Power of Movement in Plants - 50/98 -


[page 307]

Mirabilis jalapa and longiflora (Nyctagineae).--The cotyledons, which are of unequal size, stand horizontally during the middle of the day, and at night rise up vertically and come into close contact with one another. But this movement with M. longiflora lasted for only the three first nights.

Beta vulgaris (Polygoneae).--A large number of seedlings were observed on three occasions. During the day the cotyledons sometimes stood sub-horizontally, but more commonly at an angle of about 50o above the horizon, and for the first two or three nights they rose up vertically so as to be completely closed. During the succeeding one or two nights they rose only a little, and afterwards hardly at all.

Amaranthus caudatus (Amaranthaceae).--At noon the cotyledons of many seedlings, which had just germinated, stood at about 45o above the horizon, and at 10.15 P.M. some were nearly and the others quite closed. On the following morning they were again well expanded or open.

Cannabis sativa (Cannabineae).--We are very doubtful whether this plant ought to be here included. The cotyledons of a large number of seedlings, after being well illuminated during the day, were curved downwards at night, so that the tips of some pointed directly to the ground, but the basal part did not appear to be at all depressed. On the following morning they were again flat and horizontal. the cotyledons of many other seedlings were at the same time not in any way affected. Therefore this case seems very different from that of ordinary sleep, and probably comes under the head of epinasty, as is the case with the leaves of this plant according to Kraus. The cotyledons are heliotropic, and so is the hypocotyl in a still stronger degree.

Oxalis.--We now come to cotyledons provided with a pulvinus, all of which are remarkable from the continuance of the nocturnal movements during several days or even weeks, and apparently after growth has ceased. The cotyledons of O. rosea, floribunda and articulata sink vertically down at night and clasp the upper part of the hypocotyl. Those of O. Valdiviana and sensitiva, on the contrary, rise vertically up, so that their upper surfaces come into close contact; and after the young leaves are developed these are clasped by the cotyledons. As in the daytime they stand horizontally, or are even a little deflected beneath the horizon, they move in the evening through an angle of at least 90o. Their complicated circumnutating movements during the day have [page 308] been described in the first chapter. The experiment was a superfluous one, but pots with seedlings of O. rosea and floribunda were turned upside down, as soon as the cotyledons began to show any signs of sleep, and this made no difference in their movements.

Leguminosae.--It may be seen in our list that the cotyledons of several species in nine genera, widely distributed throughout the Family, sleep at night; and this probably is the case with many others. The cotyledons of all these species are provided with a pulvinus; and the movement in all is continued during many days or weeks. In Cassia the cotyledons of the ten species in the list rise up vertically at night and come into close contact with one another. We observed that those of C. florida opened in the morning rather later than those of C. glauca and pubescens. The movement is exactly the same in C. mimosoides as in the other species, though its subsequently developed leaves sleep in a different manner. The cotyledons of an eleventh species, namely, C. nodosa, are thick and fleshy, and do not rise up at night. The circumnutation of the cotyledons during the day of C. tora has been described in the first chapter. Although the cotyledons of Smithia sensitiva rose from a horizontal position in the middle of the day to a vertical one at night, those of S. Pfundii, which are thick and fleshy, did not sleep. When Mimosa pudica and albida have been kept at a sufficiently high temperature during the day, the cotyledons come into close contact at night; otherwise they merely rise up almost vertically. The circumnutation of those of M. pudica has been described. The cotyledons of a Bauhinia from St. Catharina in Brazil stood during the day at an angle of about 50o above the horizon, and at night rose to 77o; but it is probable that they would have closed completely, if the seedlings had been kept in a warmer place.

Lotus.--In three species of Lotus the cotyledons were observed to sleep. Those of L. Jacoboeus present the singular case of not rising at night in any conspicuous manner for the first 5 or 6 days of their life, and the pulvinus is not well developed at this period. Afterwards the sleeping movement is well displayed, though to a variable degree, and is long continued. We shall hereafter meet with a nearly parallel case with the leaves of Sida rhombifolia. The cotyledons of L. Gebelii are only slightly raised at night, and differ much in this respect from the three species in our list. [page 309]

Trifolium.--The germination of 21 species was observed. In most of them the cotyledons rise hardly at all, or only slightly, at night; but those of T. glomeratum, striatum and incarnactum rose from 45o to 55o above the horizon. With T. subterraneum, leucanthemum and strictum, they stood up vertically; and with T. strictum the rising movement is accompanied, as we shall see, by another movement, which makes us believe that the rising is truly nyctitropic. We did not carefully examine the cotyledons of all the species for a pulvinus, but this organ was distinctly present in those of T. subterraneum and strictum; whilst there was no trace of a pulvinus in some species, for instance, in T. resupinatum, the cotyledons of which do not rise at night.

Trifolium subterraneum.--The blades of the cotyledons on the first day after germination (Nov. 21st) were not fully expanded, being inclined at about 35o above the horizon; at night they rose to about 75o. Two days afterwards the blades at noon were horizontal, with the petioles highly inclined upwards; and it is remarkable that the nocturnal movement is almost wholly confined to the blades, being effected by the pulvinus at their bases; whilst the petioles retain day and night nearly the same inclination. On this night (Nov. 23rd), and for some few succeeding nights, the blades rose from a horizontal into a vertical position, and then became bowed inwards at about an average angle of 10o; so that they had passed through an angle of 100o. Their tips now almost touched one another, their bases being slightly divergent. The two blades thus formed a highly inclined roof over the axis of the seedling. This movement is the same as that of the terminal leaflet of the tripartite leaves of many species of Trifolium. After an interval of 8 days (Nov. 29th) the blades were horizontal during the day, and vertical at night, and now they were no longer bowed inwards. They continued to move in the same manner for the following two months, by which time they had increased greatly in size, their petioles being no less than .8 of an inch in length, and two true leaves had by this time been developed.

Trifolium strictum.--On the first day after germination the cotyledons, which are provided with a pulvinus, stood at noon horizontally, and at night rose to only about 45o above the horizon. Four days afterwards the seedlings were again observed at night, and now the blades stood vertically and were in contact, excepting the tips, which were much deflexed, so that they faced the zenith. At this age the petioles are curved [page 310] upwards, and at night, when the bases of the blades are in contact, the two petioles together form a vertical ring surrounding the plumule. The cotyledons continued to act in nearly the same manner for 8 or 10 days from the period of germination; but the petioles had by this time become straight and had increased much in length. After from 12 to 14 days the first simple true leaf was formed, and during the ensuing fortnight a remarkable movement was repeatedly observed. At I. (Fig. 125) we have a sketch, made in the middle of the day, of a seedling about a fortnight old. The two cotyledons, of which Rc is the right and Lc the left one, stand directly opposite one another,

Fig. 125. Trifolium strictum: diurnal and nocturnal positions of the two cotyledons and of the first leaf. I. Seedling viewed obliquely from above, during the day: Rc, right cotyledon; Lc, left cotyledon; F, first true leaf. II. A rather younger seedling, viewed at night: Rc, right cotyledon raised, but its position not otherwise changed; Lc, left cotyledon raised and laterally twisted; F, first leaf raised and twisted so as to face the left twisted cotyledon. III. Same seedling viewed at night from the opposite side. The back of the first leaf, F, is here shown instead of the front, as in II.

and the first true leaf (F) projects at right angles to them. At night (see II. and III.) the right cotyledon (Rc) is greatly raised, but is not otherwise changed in position. The left cotyledon (Lc) is likewise raised, but it is also twisted so that its blade, instead of exactly facing the opposite one, now stands at nearly right angles to it. This nocturnal twisting movement is effected not by means of the pulvinus, but by the twisting of the whole length of the petiole, as could be seen by the curved line of its upper concave surface. At the same time the true leaf (F) rises up, so as to stand vertically, or it even passes the vertical and is inclined a little inwards. It also twists a little, by which means the upper surface of its blade fronts, and almost comes into contact with, the upper surface of the twisted [page 311] left cotyledon. This seems to be the object gained by these singular movements. Altogether 20 seedlings were examined on successive nights, and in 19 of them it was the left cotyledon alone which became twisted, with the true leaf always so twisted that its upper surface approached closely and fronted that of the left cotyledon. In only one instance was the right cotyledon twisted, with the true leaf twisted towards it; but this seedling was in an abnormal condition, as the left cotyledon did not rise up properly at night. This whole case is remarkable, as with the cotyledons of no other plant have we seen any nocturnal movement except vertically upwards or downwards. It is the more remarkable, because we shall meet with an analogous case in the leaves of the allied genus Melilotus, in which the terminal leaflet rotates at night so as to present one edge to the zenith and at the same time bends to one side, so that its upper surface comes into contact with that of one of the two now vertical lateral leaflets.]

Concluding Remarks on the Nyctitropic Movements of Cotyledons.--The sleep of cotyledons (though this is a subject which has been little attended to), seems to be a more common phenomenon than that of leaves. We observed the position of the cotyledons during the day and night in 153 genera, widely distributed throughout the dicotyledonous series, but otherwise selected almost by hazard; and one or more species in 26 of these genera placed their cotyledons at night so as to stand vertically or almost vertically, having generally moved through an angle of at least 60o. If we lay on one side the Leguminosae, the cotyledons of which are particularly liable to sleep, 140 genera remain; and out of these, the cotyledons of at least one species in 19 genera slept. Now if we were to select by hazard 140 genera, excluding the Leguminosae, and observed their leaves at night, assuredly not nearly so many as 19 would be found to include sleeping species. We here refer exclusively to the plants observed by ourselves. [page 312]

In our entire list of seedlings, there are 30 genera, belonging to 16 Families, the cotyledons of which in some of the species rise or sink in the evening or early night, so as to stand at least 60o above or beneath the horizon. In a large majority of the genera, namely, 24, the movement is a rising one; so that the same direction prevails in these nyctitropic movements as in the lesser periodic ones described in the second chapter. The cotyledons move downwards during the early part of the night in only 6 of the genera; and in one of them, Cannabis, the curving down of the tip is probably due to epinasty, as Kraus believes to be the case with the leaves. The downward movement to the amount of 90o is very decided in Oxalis Valdiviana and sensitiva, and in Geranium rotundifolium. It is a remarkable fact that with Anoda Wrightii, one species of Gossypium and at least 3


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