The Power of Movement in Plants · Charles Darwin
NYCTITROPIC OR SLEEP MOVEMENTS OF COTYLEDONS.
We now come to the descriptive part of our work, and will begin with cotyledons, passing on to leaves in the next chapter. We have met with only two brief notices of cotyledons sleeping. Hofmeister,[13] after stating that the cotyledons of all the observed seedlings of the Caryophylleae (Alsineae and Sileneae) bend upwards at night (but to what angle he does not state), remarks that those of Stellaria media rise up so as to touch one another; they may therefore safely be said to sleep. Secondly, according to Ramey,[14] the cotyledons of Mimosa pudica and of Clianthus Dampieri rise up almost vertically at night and approach each other closely. It has been shown in a previous chapter that the cotyledons of a large number of plants bend a little upwards at night, and we here have to meet the difficult question at what inclination may they be said to sleep? According to the view which we maintain, no movement deserves to be called nyctitropic, unless it has been acquired for the sake of lessening radiation; but this could be discovered only by a long series of experiments, showing that the leaves of each species suffered from this cause, if prevented from sleeping. We must therefore take an arbitrary limit.
If a cotyledon or leaf is inclined at 60° above or beneath the horizon, it exposes to the zenith about one-half of its area; consequently the intensity of its radiation will be lessened by about half, compared with what it would have been if the cotyledon or leaf had remained horizontal. This degree of diminution certainly would make a great difference to a plant having a tender constitution. We will therefore speak of a cotyledon and hereafter of a leaf as sleeping, only when it rises at night to an angle of about 60°, or to a still higher angle, above the horizon, or sinks beneath it to the same amount. Not but that a lesser diminution of radiation may be advantageous to a plant, as in the case of Datura stramonium, the cotyledons of which rose from 31° at noon to 55° at night above the horizon. The Swedish turnip may profit by the area of its leaves being reduced at night by about 30 per cent., as estimated by Mr. A. S. Wilson; though in this case the angle through which the leaves rose was not observed. On the other hand, when the angular rise of cotyledons or of leaves is small, such as less than 30°, the diminution of radiation is so slight that it probably is of no significance to the plant in relation to radiation. For instance, the cotyledons of Geranium Ibericum rose at night to 27° above the horizon, and this would lessen radiation by only 11 per cent.: those of Linum Berendieri rose to 33°, and this would lessen radiation by 16 per cent.
The names of the plants, the cotyledons of which stand at night at an angle of at least 60° with the horizon, are arranged in the appended list on the same system as previously followed. The numbers of the Families, and with the Leguminosae the numbers of the Tribes, have been added to show how widely the plants in question are distributed throughout the dicotyledonous series. A few remarks will have to be made about many of the plants in the list. In doing so, it will be convenient not to follow strictly any systematic order, but to treat of the Oxalidæ and the Leguminosae at the close; for in these two Families the cotyledons are generally provided with a pulvinus, and their movements endure for a much longer time than those of the other plants in the list. List of Seedling Plants, the cotyledons of which rise or sink at night to an angle of at least 60° above or beneath the horizon.
Brassica oleracea. Cruciferae (Fam. 14). — napus (as we are informed by Prof. Pfeffer). Raphanus sativus. Cruciferae. Githago segetum. Caryophylleae (Fam. 26). Stellaria media (according to Hofmeister, as quoted). Caryophylleae. Anoda Wrightii. Malvaceae (Fam. 36). Gossypium (var. Nankin cotton). Malvaceae. Oxalis rosea. Oxalidæ (Fam. 41). — floribunda. — articulata. — Valdiviana. — sensitiva. Geranium rotundifolium. Geraniaceae (Fam. 47). Trifolium subterraneum. Leguminosae (Fam. 75, Tribe 3). — strictum. — leucanthemum. Lotus ornithopopoides. Leguminosae (Tribe 4). — peregrinus. — Jacobæus. Clianthus Dampieri. Leguminosae (Tribe 5)—according to M. Ramey. Smithia sensitiva. Leguminosae (Tribe 6). Haematoxylon Campechianum. Leguminosae (Tribe 13)—according to Mr. R. I. Lynch. Cassia mimosoides. Leguminosae (Tribe 14). — glauca. — florida. — corymbosa. — pubescens. — tora. — neglecta. — 3 other Brazilian unnamed species. Bauhinia (sp.?. Leguminosae (Tribe 15). Neptunia oleracea. Leguminosae (Tribe 20). Mimosa pudica. Leguminosae (Tribe 21). — albida. Cucurbita ovifera. Cucurbitaceæ (Fam. 106). — aurantia. Lagenaria vulgaris. Cucurbitaceæ. Cucumis dudaim. Cucurbitaceæ. Apium petroselinum. Umbelliferae (Fam. 113). — graveolens. Lactuca scariola. Compositæ (Fam. 122). Helianthus annuus (?). Compositæ. Ipomœa caerulea. Convolvulaceae (Fam. 151). — purpurea. — bona-nox. — coccinea. Solanum lycopersicum. Solaneae (Fam. 157.) Mimulus, (sp. ?) Scrophularineae (Fam. 159)—from information given us by Prof. Pfeffer. Mirabilis jalapa. Nyctagineae (Fam. 177). Mirabilis longiflora. Beta vulgaris. Polygoneae (Fam. 179). Amaranthus caudatus. Amaranthaceae (Fam. 180). Cannabis sativa (?). Cannabineae (Fam. 195).
Brassica oleracea (Cruciferae).—It was shown in the first chapter that the cotyledons of the common cabbage rise in the evening and stand vertically up at night with their petioles in contact. But as the two cotyledons are of unequal height, they frequently interfere a little with each other’s movements, the shorter one often not standing quite vertically. They awake early in the morning; thus at 6.45 A.M. on Nov. 27th, whilst if was still dark, the cotyledons, which had been vertical and in contact on the previous evening, were reflexed, and thus presented a very different appearance. It should be borne in mind that seedlings in germinating at the proper season, would not be subjected to darkness at this hour in the morning. The above amount of movement of the cotyledons is only temporary, lasting with plants kept in a warm greenhouse from four to six days; how long it would last with seedlings growing out of doors we do not know. Raphanus sativus.—In the middle of the day the blades of the cotyledons of 10 seedlings stood at right angles to their hypocotyls, with their petioles a little divergent; at night the blades stood vertically, with their bases in contact and with their petioles parallel. Next morning, at 6.45 A.M., whilst it was still dark, the blades were horizontal. On the following night they were much raised, but hardly stood sufficiently vertical to be said to be asleep, and so it was in a still less degree on the third night. Therefore the cotyledons of this plant (kept in the greenhouse) go to sleep for even a shorter time than those of the cabbage. Similar observations were made, but only during a single day and night, on 13 other seedlings likewise raised in the greenhouse, with the same result.
The petioles of the cotyledons of 11 young seedlings of Sinapis nigra were slightly divergent at noon, and the blades stood at right angles to the hypocotyls; at night the petioles were in close contact, and the blades considerably raised, with their bases in contact, but only a few stood sufficiently upright to be called asleep. On the following morning, the petioles diverged before it was light. The hypocotyl is slightly sensitive, so that if rubbed with a needle it bends towards the rubbed side. In the case of Lepidium sativum, the petioles of the cotyledons of young seedlings diverge during the day and converge so as to touch each other during the night, by which means the bases of the tripartite blades are brought into contact; but the blades are so little raised that they cannot be said to sleep. The cotyledons of several other cruciferous plants were observed, but they did not rise sufficiently during the night to be said to sleep. Githago segetum (Caryophylleae).—On the first day after the cotyledons had burst through the seed-coats, they stood at noon at an angle of 75° above the horizon; at night they moved upwards, each through an angle of 15° so as to stand quite vertical and in contact with one another. On the second day they stood at noon at 59° above the horizon, and again at night were completely closed, each having risen 31°. On the fourth day the cotyledons did not quite close at night. The first and succeeding pairs of young true leaves behaved in exactly the same manner. We think that the movement in this case may be called nyctitropic, though the angle passed through was small. The cotyledons are very sensitive to light and will not expand if exposed to an extremely dim one.
Anoda Wrightii (Malvaceae).—The cotyledons whilst moderately young, and only from .2 to .3 inch in diameter, sink in the evening from their mid-day horizontal position to about 35° beneath the horizon. But when the same seedlings were older and had produced small true leaves, the almost orbicular cotyledons, now .55 inch in diameter, moved vertically downwards at night. This fact made us suspect that their sinking might be due merely to their weight; but they were not in the least flaccid, and when lifted up sprang back through elasticity into their former dependent position. A pot with some old seedlings was turned upside down in the afternoon, before the nocturnal fall had commenced, and at night they assumed in opposition to their own weight (and to any geotropic action) an upwardly directed vertical position. When pots were thus reversed, after the evening fall had already commenced, the sinking movement appeared to be somewhat disturbed; but all their movements were occasionally variable without any apparent cause. This latter fact, as well as that of the young cotyledons not sinking nearly so much as the older ones, deserves notice. Although the movement of the cotyledons endured for a long time, no pulvinus was exteriorly visible; but their growth continued for a long time. The cotyledons appear to be only slightly heliotropic, though the hypocotyl is strongly so.
Gossypium arboreum (?) (var. Nankin cotton) (Malvaceae).—The cotyledons behave in nearly the same manner as those of the Anoda. On June 15th the cotyledons of two seedlings were .65 inch in length (measured along the midrib) and stood horizontally at noon; at 10 P.M. they occupied the same position and had not fallen at all. On June 23rd, the cotyledons of one of these seedlings were 1.1 inch in length, and by 10 P.M. they had fallen from a horizontal position to 62° beneath the horizon. The cotyledons of the other seedling were 1.3 inch in length, and a minute true leaf had been formed; they had fallen at 10 P.M. to 70° beneath the horizon. On June 25th, the true leaf of this latter seedling was .9 inch in length, and the cotyledons occupied nearly the same position at night. By July 9th the cotyledons appeared very old and showed signs of withering; but they stood at noon almost horizontally, and at 10 P.M. hung down vertically.
Gossypium herbaceum.—It is remarkable that the cotyledons of this species behave differently from those of the last. They were observed during 6 weeks from their first development until they had grown to a very large size (still appearing fresh and green), viz. 2½ inches in breadth. At this age a true leaf had been formed, which with its petiole was 2 inches long. During the whole of these 6 weeks the cotyledons did not sink at night; yet when old their weight was considerable and they were borne by much elongated petioles. Seedlings raised from some seed sent us from Naples, behaved in the same manner; as did those of a kind cultivated in Alabama and of the Sea-island cotton. To what species these three latter forms belong we do not know. We could not make out in the case of the Naples cotton, that the position of the cotyledons at night was influenced by the soil being more or less dry; care being taken that they were not rendered flaccid by being too dry. The weight of the large cotyledons of the Alabama and Sea-island kinds caused them to hang somewhat downwards, when the pots in which they grew were left for a time upside down. It should, however, be observed that these three kinds were raised in the middle of the winter, which sometimes greatly interferes with the proper nyctitropic movements of leaves and cotyledons.
Cucurbitaceæ.—The cotyledons of Cucurbita aurantia and ovifera, and of Lagenaria vulgaris, stand from the 1st to the 3rd day of their life at about 60° above the horizon, and at night rise up so as to become vertical and in close contact with one another. With Cucumis dudaim they stood at noon at 45° above the horizon, and closed at night. The tips of the cotyledons of all these species are, however, reflexed, so that this part is fully exposed to the zenith at night; and this fact is opposed to the belief that the movement is of the same nature as that of sleeping plants. After the first two or three days the cotyledons diverge more during the day and cease to close at night. Those of Trichosanthes anguina are somewhat thick and fleshy, and did not rise at night; and they could perhaps hardly be expected to do so. On the other hand, those of _Acanthosicyos horrida_[15] present nothing in their appearance opposed to their moving at night in the same manner as the preceding species; yet they did not rise up in any plain manner. This fact leads to the belief that the nocturnal movements of the above-named species has been acquired for some special purpose, which may be to protect the young plumule from radiation, by the close contact of the whole basal portion of the two cotyledons.
Apium graveolens.—The cotyledons at noon were horizontal, and at 10 P.M. stood at an angle of 61° above the horizon. Lactuca scariola (Compositæ).—The cotyledons whilst young stood sub-horizontally during the day, and at night rose so as to be almost vertical, and some were quite vertical and closed; but this movement ceased when they had grown old and large, after an interval of 11 days. Helianthus annuus (Compositæ).—This case is rather doubtful; the cotyledons rise at night, and on one occasion they stood at 73° above the horizon, so that they might then be said to have been asleep. Ipomœa caerulea vel Pharbitis nil (Convolvulaceae).—The cotyledons behave in nearly the same manner as those of the Anoda and Nankin cotton, and like them grow to a large size. Whilst young and small, so that their blades were from .5 to .6 of an inch in length, measured along the middle to the base of the central notch, they remained horizontal both during the middle of the day and at night. As they increased in size they began to sink more and more in the evening and early night; and when they had grown to a length (measured in the above manner) of from 1 to 1.25 inch, they sank between 55° and 70° beneath the horizon. They acted, however, in this manner only when they had been well illuminated during the day. Nevertheless, the cotyledons have little or no power of bending towards a lateral light, although the hypocotyl is strongly heliotropic. They are not provided with a pulvinus, but continue to grow for a long time.
Ipomœa purpurea (vel Pharbitis hispida).—The cotyledons behave in all respects like those of I. caerulea. A seedling with cotyledons .75 inch in length (measured as before) and 1.65 inch in breadth, having a small true leaf developed, was placed at 5.30 P.M. on a klinostat in a darkened box, so that neither weight nor geotropism could act on them. At 10 P.M. one cotyledon stood at 77° and the other at 82° beneath the horizon. Before being placed in the klinostat they stood at 15° and 29° beneath the horizon. The nocturnal position depends chiefly on the curvature of the petiole close to the blade, but the whole petiole becomes slightly curved downwards. It deserves notice that seedlings of this and the last-named species were raised at the end of February and another lot in the middle of March, and the cotyledons in neither case exhibited any nyctitropic movement. Ipomœa bona-nox.—The cotyledons after a few days grow to an enormous size, those on a young seedling being 3 1/4 inches in breadth. They were extended horizontally at noon, and at 10 P.M. stood at 63° beneath the horizon. five days afterwards they were 4½ inches in breadth, and at night one stood at 64° and the other 48° beneath the horizon. Though the blades are thin, yet from their great size and from the petioles being long, we imagined that their depression at night might be determined by their weight; but when the pot was laid horizontally, they became curved towards the hypocotyl, which movement could not have been in the least aided by their weight, at the same time they were somewhat twisted upwards through apogeotropism. Nevertheless, the weight of the cotyledons is so far influential, that when on another night the pot was turned upside down, they were unable to rise and thus to assume their proper nocturnal position.
Ipomœa coccinea.—The cotyledons whilst young do not sink at night, but when grown a little older, but still only .4 inch in length (measured as before) and .82 in breadth, they became greatly depressed. In one case they were horizontal at noon, and at 10 P.M. one of them stood at 64° and the other at 47° beneath the horizon. The blades are thin, and the petioles, which become much curved down at night, are short, so that here weight can hardly have produced any effect. With all the above species of Ipomœa, when the two cotyledons on the same seedling were unequally depressed at night, this seemed to depend on the position which they had held during the day with reference to the light. Solanum lycopersicum (Solaneae).—The cotyledons rise so much at night as to come nearly in contact. Those of ‘S. palinacanthum’ were horizontal at noon, and by 10 P.M. had risen only 27° 30 minutes; but on the following morning before it was light they stood at 59° above the horizon, and in the afternoon of the same day were again horizontal. The behaviour of the cotyledons of this latter species seems, therefore, to be anomalous. 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 50° 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 45° 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 90°. Their complicated circumnutating movements during the day have 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 50° above the horizon, and at night rose to 77°; 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. 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 45° to 55° 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.
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