The Power of Movement in Plants - Charles Darwin (top reads txt) 📗

they had better be at least temporarily included. This same source of doubt applies in some few other cases; for at the commencement of our observations we did not always attend sufficiently to whether the cotyledons stood nearly horizontally in the middle of the day. With several seedlings, the cotyledons assume a highly inclined position at night during so short a period of their life, that a doubt naturally arises whether this can be of any service to the plant. Nevertheless, in most of the cases given in the following list, the cotyledons may be as certainly said to sleep as may the leaves of any plant. In two cases, namely with the cabbage and radish, the cotyledons of which rise almost vertically during the few first nights of their life, it was ascertained by placing young seedlings in the klinostat, that the upward movement was not due to apogeotropism.

 

The names of the plants, the cotyledons of which stand at night at an angle of at least 60o 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 [page 300]

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 Oxalidae 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 60o 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. Oxalidae (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.

— Jacobaeus.

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. Cucurbitaceae (Fam. 106).

— aurantia.

Lagenaria vulgaris. Cucurbitaceae.

Cucumis dudaim. Cucurbitaceae.

Apium petroselinum. Umbelliferae (Fam. 113).

— graveolens.

Lactuca scariola. Compositae (Fam. 122).

Helianthus annuus (?). Compositae.

Ipomoea caerulea. Convolvulaceae (Fam. 151).

— purpurea.

— bona-nox.

— coccinea.

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List of Seedling Plants (continued).

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, [page 302]

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 75o above the horizon; at night they moved upwards, each through an angle of 15o so as to stand quite vertical and in contact with one another. On the second day they stood at noon at 59o above the horizon, and again at night were completely closed, each having risen 31o. 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 35o 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.

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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 62o 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 70o 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. 2 � 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.

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Cucurbitaceae.—The cotyledons of Cucurbita aurantia and ovifera, and of Lagenaria vulgaris, stand from the 1st to the 3rd day of their life at about 60o 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 45o 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* 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