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 60° 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 90° 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 Ipomœa, 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, Ipomœa 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 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 Jacobæus, 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 Raphanus for only a single night. There are, however, some strong exceptions to this rule, as the cotyledons of Gossypium, Anoda and Ipomœa 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 Oxalidæ, 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 seems to be some connection, as might have been expected, between the two sets of cases. For the leaves of many plants sleep, whilst their cotyledons do not do so—of which fact Desmodium gyrans offers a good instance, as likewise do three species of Nicotiana observed by us; also Sida rhombifolia, Abutilon Darwinii, and Chenopodium album. On the other hand, the cotyledons of some plants sleep and not the leaves, as with the species of Beta, Brassica, Geranium, Apium, Solanum, and Mirabilis, named in our list. Still more striking is the fact that, in the same genus, the leaves of several or of all the species may sleep, but the cotyledons of only some of them, as occurs with Trifolium, Lotus, Gossypium, and partially with Oxalis. Again, when both the cotyledons and the leaves of the same plant sleep, their movements may be of a widely dissimilar nature: thus with Cassia the cotyledons rise vertically up at night, whilst their leaves sink down and twist round so as to turn their lower surfaces outwards. With seedlings of Oxalis Valdiviana, having 2 or 3 well-developed leaves, it was a curious spectacle to behold at night each leaflet folded inwards and hanging perpendicularly downwards, whilst at the same time and on the same plant the cotyledons stood vertically upwards.
These several facts, showing the independence of the nocturnal movements of the leaves and cotyledons on the same plant, and on plants belonging to the same genus, lead to the belief that the cotyledons have acquired their power of movement for some special purpose. Other facts lead to the same conclusion, such as the presence of pulvini, by the aid of which the nocturnal movement is continued during some weeks. In Oxalis the cotyledons of some species move vertically upwards, and of others vertically downwards at night; but this great difference within the same natural genus is not so surprising as it may at first appear, seeing that the cotyledons of all the species are continually oscillating up and down during the day, so that a small cause might determine whether they should rise or sink at night. Again, the peculiar nocturnal movement of the left-hand cotyledon of Trifolium strictum, in combination with that of the first true leaf. Lastly, the wide distribution in the dicotyledonous series of plants with cotyledons which sleep. Reflecting on these several facts, our conclusion seems justified, that the nyctitropic movements of cotyledons, by which the blade is made to stand either vertically or almost vertically upwards or downwards at night, has been acquired, at least in most cases, for some special purpose; nor can we doubt that this purpose is the protection of the upper surface of the blade, and perhaps of the central bud or plumule, from radiation at night.
CHAPTER VII.
MODIFIED CIRCUMNUTATION: NYCTITROPIC OR SLEEP MOVEMENTS OF LEAVES.
Conditions necessary for these movements—List of Genera and Families, which include sleeping plants—Description of the movements in the several Genera—Oxalis: leaflets folded at night—Averrhoa: rapid movements of the leaflets—Porlieria: leaflets close when plant kept very dry—Tropaeolum: leaves do not sleep unless well illuminated during day—Lupinus: various modes of sleeping—Melilotus: singular movements of terminal leaflet—Trifolium—Desmodium: rudimentary lateral leaflets, movements of, not developed on young plants, state of their pulvini—Cassia: complex movements of the leaflets—Bauhinia: leaves folded at night—Mimosa pudica: compounded movements of leaves, effect of darkness—Mimosa albida, reduced leaflets of—Schrankia: downward movement of the pinnae—Marsilea: the only cryptogam known to sleep—Concluding remarks and summary—Nyctitropism consists of modified circumnutation, regulated by the alternations of light and darkness—Shape of first true leaves.
We now come to the nyctitropic or sleep movements of leaves. It should be remembered that we confine this term to leaves which place their blades at night either in a vertical position or not more than 30° from the vertical,—that is, at least 60° above or beneath the horizon. In some few cases this is effected by the rotation of the blade, the petiole not being either raised or lowered to any considerable extent. The limit of 30° from the vertical is obviously an arbitrary one, and has been selected for reasons previously assigned, namely, that when the blade approaches the perpendicular as nearly as this, only half as much of the surface is exposed at night to the zenith and to free radiation as when the blade is horizontal. Nevertheless, in a few instances, leaves which seem to be prevented by their structure from moving to so great an extent as 60° above or beneath the horizon, have been included amongst sleeping plants.
It should be premised that the nyctitropic movements of leaves are easily affected by the conditions to which the plants have been subjected. If the ground is kept too dry, the movements are much delayed or fail: according to Dassen,[[1]] even if the air is very dry the leaves of Impatiens and Malva are rendered motionless. Carl Kraus has also lately insisted[[2]] on the great influence which the quantity of water absorbed has on the periodic movements of leaves; and he believes that this cause chiefly determines the variable amount of sinking of the leaves of Polygonum convolvulus at night; and if so, their movements are not in our sense strictly nyctitropic. Plants in order to sleep must have been exposed to a proper temperature: Erythrina crista-galli, out of doors and nailed against a wall, seemed in fairly good health, but the leaflets did not sleep, whilst those on another plant kept in a warm greenhouse were all vertically dependent at night. In a kitchen-garden the leaflets of Phaseolus vulgaris did not sleep during the early part of the summer. Ch. Royer says,[[3]] referring I suppose to the native plants in France, that they do not sleep when the temperature is below 5° C. or 41° F. In the case of several sleeping plants, viz., species of Tropaeolum, Lupinus, Ipomœa, Abutilon, Siegesbeckia, and probably other genera, it is indispensable that the leaves should be well illuminated during the day in order that they may assume at night a vertical position; and it was probably owing to this cause that seedlings of Chenopodium album and Siegesbeckia orientalis, raised by us during the middle of the winter, though kept at a proper temperature, did not sleep. Lastly, violent agitation by a strong wind, during a few minutes, of the leaves of Maranta arundinacea (which previously had not been disturbed in the hot-house), prevented their sleeping during the two next nights.
[1] Dassen,’Tijdschrift vor. Naturlijke Gesch. en Physiologie,’ 1837, vol. iv. p. 106. See also Ch. Royer on the importance of a proper state of turgescence of the cells, in ‘Annal. des Sc. Nat. Bot.’ (5th series), ix. 1868, p. 345.