The Power of Movement in Plants, Charles Darwin [the reading list book TXT] 📗

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

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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.

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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

species of Ipomoea, the cotyledons whilst young and light sink at night very little or not at all; although this movement becomes well pronounced as soon as they have grown large and heavy. Although the downward movement cannot be attributed to the weight of the cotyledons in the several cases which were investigated, namely, in those of the Anoda, Ipomoea purpurea and bona-nox, nor in that of I. coccinea, yet bearing in mind that cotyledons are continually circumnutating, a slight cause might at first have determined whether the great nocturnal movement should be upwards or downwards. We may therefore suspect that in some aboriginal member of the groups in question, the weight of the cotyledons first determined the downward direction. The fact of the cotyledons of these species not sinking down much whilst they are young and tender, seems opposed to the belief that the greater movement when they are

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grown older, has been acquired for the sake of protecting them from radiation at night; but then we should remember that there are many plants, the leaves of which sleep, whilst the cotyledons do not; and if in some cases the leaves are protected from cold at night whilst the cotyledons are not protected, so in other cases it may be of more importance to the species that the nearly full-grown cotyledons should be better protected than the young ones.

 

In all the species of Oxalis observed by us, the cotyledons are provided with pulvini; but this organ has become more or less rudimentary in O.

corniculata, and the amount of upward movement of its cotyledons at night is very variable, but is never enough to be called sleep. We omitted to ascertain whether the cotyledons of Geranium rotundifolium possess pulvini.

In the Leguminosae all the cotyledons which sleep, as far as we have seen, are provided with pulvini. But with Lotus Jacobaeus, these are not fully developed during the first few days of the life of the seedling, and the cotyledons do not then rise much at night. With Trifolium strictum the blades of the cotyledons rise at night by the aid of their pulvini; whilst the petiole of one cotyledon twists half-round at the same time, independently of its pulvinus.

 

As a general rule, cotyledons which are provided with pulvini continue to rise or sink at night during a much longer period than those destitute of this organ. In this latter case the movement no doubt depends on alternately greater growth on the upper and lower side of the petiole, or of the blade, or of both, preceded probably by the increased turgescence of the growing cells. Such movements generally last for a very short period—

for instance, with Brassica and Githago for 4 or 5 nights, with Beta for 2

or 3, and with

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Raphanus for only a single night. There are, however, some strong exceptions to this rule, as the cotyledons of Gossypium, Anoda and Ipomoea do not possess pulvini, yet continue to move and to grow for a long time.

We thought at first that when the movement lasted for only 2 or 3 nights, it could hardly be of any service to the plant, and hardly deserved to be called sleep; but as many quickly-growing leaves sleep for only a few nights, and as cotyledons are rapidly developed and soon complete their growth, this doubt now seems to us not well-founded, more especially as these movements are in many instances so strongly pronounced. We may here mention another point of similarity between sleeping leaves and cotyledons, namely, that some of the latter (for instance, those of Cassia and Githago) are easily affected by the absence of light; and they then either close, or if closed do not open; whereas others (as with the cotyledons of Oxalis) are very little affected by light. In the next chapter it will be shown that the nyctitropic movements both of cotyledons and leaves consist of a modified form of circumnutation.

 

As in the Leguminosae and Oxalidae, the leaves and the cotyledons of the same species generally sleep, the idea at first naturally occurred to us, that the sleep of the cotyledons was merely an early development of a habit proper to a more advanced stage of life. But no such explanation can be admitted, although there