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.
[2] ‘Beiträge zur Kentniss der Bewegungen,’ etc., in ‘Flora,’ 1879, pp. 42, 43, 67, etc.
[3] ‘Annal. des Sc. Nat. Bot.’ (5th Series), ix. 1868, p. 366.
We will now give our observations on sleeping plants, made in the manner described in the Introduction. The stem of the plant was always secured (when not stated to the contrary) close to the base of the leaf, the movements of which were being observed, so as to prevent the stem from circumnutating. As the tracings were made on a vertical glass in front of the plant, it was obviously impossible to trace its course as soon as the leaf became in the evening greatly inclined either upwards or downwards; it must therefore be understood that the broken lines in the diagrams, which represent the evening and nocturnal courses, ought always to be prolonged to a much greater distance, either upwards or downwards, than appears in them. The conclusions which may be deduced from our observations will be given near the end of this chapter.
In the following list all the genera which include sleeping plants are given, as far as known to us. The same arrangement is followed as in former cases, and the number of the Family is appended. This list possesses some interest, as it shows that the habit of sleeping is common to some few plants throughout the whole vascular series. The greater number of the genera in the list have been observed by ourselves with more or less care; but several are given on the authority of others (whose names are appended in the list), and about these we have nothing more to say. No doubt the list is very imperfect, and several genera might have been added from the ‘Somnus Plantarum’ by Linnæus; but we could not judge in some of his cases, whether the blades occupied at night a nearly vertical position. He refers to some plants as sleeping, for instance, Lathyrus odoratus and Vicia faba, in which we could observe no movement deserving to be called sleep, and as no one can doubt the accuracy of Linnæus, we are left in doubt.
[List of Genera, including species the leaves of which sleep.
CLASS I. DICOTYLEDONS.
Sub-class I. ANGIOSPERMS.
Genus Family.
Githago Caryophylleae (26). Stellaria (Batalin). ” Portulaca (Ch.Royer). Portulaceae (27). Sida Malvaceae (36). Abutilon. ” Malva (Linnæus and Pfeffer). ” Hibiscus (Linnæus). ” Anoda. ” Gossypium. ” Ayenia (Linnæus). Sterculaceae (37). Triumfetta (Linnæus). Tiliaceae (38). Linum (Batalin). Lineae (39). Oxalis. Oxalidæ (41). Averrhoa. ” Porlieria. Zygophylleæ (45). Guiacum. ” Impatiens (Linnæus, Pfeffer, Batalin). Balsamineae (48). Tropaeolum. Tropaeoleae (49). Crotolaria (Thiselton Dyer). Leguminosae (75) Tribe II. Lupinus. ” ” Cytisus. ” ” Trigonella. ” Tr. III. Medicago. ” Melilotus. ” ” Trifolium. ” ” Securigera. ” Tr. IV. Lotus. ” ” Psoralea. ” Tr. V. Amorpha (Cuchartre). ” ” Daelea. ” ” Indigofera. ” ” Tephrosia. ” ” Wistaria. ” ” Robinia. ” ” Sphaerophysa. ” ” Colutea. ” ” Astragalus. ” ” Glycyrrhiza. ” ” Coronilla. ” Tr. VI. Hedysarum. ” ” Genus Family. Onobrychis. Leguminosae (75) Tr. VI. Smithia. ” ” Arachis. ” ” Desmodium. ” ” Urania. ” ” Vicia. ” Tr. VII. Centrosema. ” Tr. VIII. Amphicarpæa. ” ” Glycine. ” ” Erythrina. ” ” Apios. ” ” Phaseolus. ” ” Sophora. ” Tr. X. Caesalpinia. ” Tr. XIII. Haematoxylon. ” ” Gleditschia (Duchartre). ” ” Poinciana. ” ” Cassia. ” Tr. XIV. Bauhinia. ” Tr. XV. Tamarindus. ” Tr. XVI. Adenanthera. ” Tr. XX. Prosopis. ” ” Neptunia. ” ” Mimosa. ” ” Schrankia. ” ” Acacia. ” Tr. XXII. Albizzia. ” Tr. XXIII. Melaleuca (Bouché). Myrtaceae (94). Genus Family. Aenothera (Linnæus). Omagrarieae (100). Passiflora. Passifloracea (105). Siegesbeckia. Compositæ (122). Ipomœa. Convolvulacea (151). Nicotiana. Solaneae (157). Mirabilis. Nyctagineae (177). Polygonum (Batalin). Polygoneae (179). Amaranthus. Amaranthaceae (180). Chenopodium. Chenopodieae (181). Pimelia (Bouché). Thymeteae (188). Euphorbia. Euphorbiaceae (202) Phyllanthus (Pfeffer). ”
Sub-class II. GYMNOSPERMS. Aies (Chatin).
CLASS II. MONOCOTYLEDONS.
Thalia. Cannaceae (21). Maranta. ” Colocasia. Aroideae (30). Strephium. Gramineæ (55).
CLASS III. ACOTYLEDONS.
Marsilea. Marsileaceae (4).
Githago segetum (Caryophylleae).—The first leaves produced by young seedlings, rise up and close together at night. On a rather older seedling, two young leaves stood at noon at 55° above the horizon, and at night at 86°, so each had risen 31°. The angle, however, was less in some cases. Similar observations were occasionally made on young leaves (for the older ones moved very little) produced by nearly full-grown plants. Batalin says (‘Flora,’ Oct. 1st, 1873, p. 437) that the young leaves of Stellaria close up so completely at night that they form together great buds.
Sida (Malvaceae).—the nyctitropic movements of the leaves in this genus are remarkable in some respects. Batalin informs us (see also ‘Flora,’ Oct. 1st, 1873, p. 437) that those of S. napaea fall at night, but to what angle he cannot remember. The leaves of S. rhombifolia and retusa, on the other hand, rise up vertically, and are pressed against the stem. We have therefore here within the same genus, directly opposite movements. Again, the leaves of S. rhombifolia are furnished with a pulvinus, formed of a mass of small cells destitute of chlorophyll, and with their longer axes perpendicular to the axis of the petiole. As measured along this latter line, these cells are only 1/5th of the length of those of the petiole; but instead of being abruptly separated from them (as is usual with the pulvinus in most plants), they graduate into the larger cells of the petiole. On the other hand, S. napaea, according to Batalin, does not possess a pulvinus; and he informs us that a gradation may be traced in the several species of the genus between these two states of the petiole. Sida rhombifolia presents another peculiarity, of which we have seen no other instance with leaves that sleep: for those on very young plants, though they rise somewhat in the evening, do not go to sleep, as we observed on several occasions; whilst those on rather older plants sleep in a conspicuous manner. For instance a leaf (.85 of an inch in length) on a very young seedling 2 inches high, stood at noon 9° above the horizon, and at 10 P.M. at 28°, so it had risen only 19°; another leaf (1.4 inch in length) on a seedling of the same height, stood at the same two periods at 7° and 32°, and therefore had risen 25°. These leaves, which moved so little, had a fairly well-developed pulvinus. After an interval of some weeks, when the same seedlings were 2½ and 3 inches in height, some of the young leaves stood up at night quite vertically, and others were highly inclined; and so it was with bushes which were fully grown and were flowering.
Fig. 126. Sida rhombifolia: circumnutation and nyctitropic (or sleep) movements of a leaf on a young plant, 9½ inches high; filament fixed to midrib of nearly full-grown leaf, 2 3/8 inches in length; movement traced under a sky-light. Apex of leaf 5 5/8 inches from the vertical glass, so diagram not greatly enlarged.
The movement of a leaf was traced from 9.15 A.M. on May 28th to 8.30 A.M. on the 30th. The temperature was too low (15°–16° C.), and the illumination hardly sufficient; consequently the leaves did not become quite so highly inclined at night, as they had done previously and as they did subsequently in the hot-house: but the movements did not appear otherwise disturbed. On the first day the leaf sank till 5.15 P.M.; it then rose rapidly and greatly till 10.5 P.M., and only a little higher during the rest of the night (Fig. 126). Early on the next day (29th) it fell in a slightly zigzag line rapidly until 9 A.M., by which time it had reached nearly the same place as on the previous morning. During the remainder of the day it fell slowly, and zigzagged laterally. The evening rise began after 4 P.M. in the same manner as before, and on the second morning it again fell rapidly. The ascending and descending lines do not coincide, as may be seen in the diagram. On the 30th a new tracing was made (not here given) on a rather enlarged scale, as the apex of the leaf now stood 9 inches from the vertical glass. In order to observe more carefully the course pursued at the time when the diurnal fall changes into the nocturnal rise, dots were made every half-hour between 4 P.M. and 10.30 P.M. This rendered the lateral zigzagging movement during the evening more conspicuous than in the diagram given, but it was of the same nature as there shown. The impression forced on our minds was that the leaf was expending superfluous movement, so that the great nocturnal rise might not occur at too early an hour.
Abutilon Darwinii (Malvaceae).—The leaves on some very young plants stood almost horizontally during the day, and hung down vertically at night. Very fine plants kept in a large hall, lighted only from the roof, did not sleep at night for in order to do so the leaves must be well illuminated during the day. The cotyledons do not sleep. Linnæus says that the leaves of his Sida abutilon sink perpendicularly down at night, though the petioles rise. Prof. Pfeffer informs us that the leaves of a Malva, allied to M. sylvestris, rise greatly at night; and this genus, as well as that of Hibiscus, are included by Linnæus in his list of sleeping plants.
Anoda Wrightii (Malvaceae).—The leaves, produced by very young plants, when grown to a moderate size, sink at night either almost vertically down or to an angle of about 45° beneath the horizon; for there is a considerable degree of variability in the amount of sinking at night, which depends in part on the degree to which they have been illuminated during the day. But the leaves, whilst quite young, do not sink down at night, and this is a very unusual circumstance. The summit of the petiole, where it joins the blade, is developed into a pulvinus, and this is present in very young leaves which do not sleep; though it is not so well defined as in older leaves.
Gossypium (var. Nankin cotton, Malvaceae).—Some young leaves, between 1 and 2 inches in length, borne by two seedlings 6 and 7½ inches in height, stood horizontally, or were raised a little above the horizon at noon on July 8th and 9th; but by 10 P.M. they had sunk down to between 68° and 90° beneath the horizon. When the same plants had grown to double the above height, their leaves stood at night almost or quite vertically dependent. The leaves on some large plants of G. maritimum and Brazilense, which were kept in a very badly lighted hot-house, only occasionally sank much downwards at night, and hardly enough to be called sleep.
Oxalis (Oxalidæ).—In most of the species in this large genus the three leaflets sink vertically down at night; but as their sub-petioles are short the blades could not assume this position from the want of space, unless they were in some manner rendered narrower; and this is effected by their becoming more or less folded (Fig. 127). The angle formed by the two halves of the same leaflet was found to vary in different individuals of several species between 92° and 150°; in three of the best folded leaflets of O. fragrans it was 76°, 74°, and 54°. The angle is often different in the three leaflets of the same leaf. As the leaflets sink down at night and become folded, their lower surfaces are brought near together (see B), or even into close contact; and from this circumstance it might be thought that the object of the folding was the protection of their lower surfaces. If this had been the case, it would have formed a strongly marked exception to the rule, that when there is any difference in the degree of protection from radiation of the two surfaces of the leaves, it is always the upper surface which is the best protected. But that the folding of the leaflets, and consequent mutual approximation of their lower surfaces, serves merely to allow them to sink down vertically, may be inferred from the fact that when the leaflets do not radiate from the summit of a common petiole, or, again, when there is plenty of room from the sub-petioles not being very short, the leaflets sink down without becoming folded. This occurs with the leaflets of O. sensitiva, Plumierii, and bupleurifolia.
Fig. 127. Oxalis acetosella: A, leaf seen from vertically above; B, diagram of leaf asleep, also seen from vertically above.
There is no use in giving a long list of the many species which sleep in the above described manner. This holds good with species having rather fleshy leaves, like those of O. carnosa, or large leaves like those of O. Ortegesii, or four leaflets like those of O. variabilis. There are, however, some species which show no signs of sleep, viz., O. pentaphylla, enneaphylla, hirta, and rubella. We will now describe the nature of the movements in some of the species.
Oxalis acetosella.—The movement of a leaflet, together with that of the main petiole, are shown in the following diagram (Fig. 128), traced between 11 A.M. on October 4th and 7.45 A.M. on the 5th. After 5.30 P.M. on the 4th the leaflet sank rapidly, and at 7 P.M. depended vertically. for some time before it assumed this latter position, its movements could, of course, no longer be traced on the vertical glass, and the broken line in the diagram ought to be extended much further down in this and all other cases. By 6.45 A.M. on the following morning it had risen considerably, and continued to rise for the next hour; but, judging from other observations, it would soon have begun to fall again. Between 11 A.M. and 5.30 P.M. the leaflet moved at least four times up and four times down before the great nocturnal fall commenced; it reached its highest point at noon. Similar observations were made on two other leaflets, with nearly the same results. Sachs and Pfeffer have also described briefly[4] the autonomous movements of the leaves of this plant.
[4] Sachs in ‘Flora,’ 1863, p. 470, etc; Pfeffer, ‘Die Period. Bewegungen,’ etc., 1875, p. 53.
Fig 128. Oxalis acetosella: circumnutation and nyctitropic movements of a nearly full-grown leaf, with filament attached to the midrib of one of the leaflets; traced on vertical glass during 20 h. 45m.
On another occasion the petiole of a leaf was secured to a little stick close beneath the leaflets, and a filament tipped with a bead of sealing-wax was affixed to the mid-rib of one of them, and a mark was placed close behind. At 7 P.M., when the leaflets were asleep, the filament depended vertically down, and the movements of the bead were then traced till 10.40 P.M., as shown in the following diagram (Fig. 129). We here see that the leaflet moved a little from side to side, as well as a little up and down, whilst asleep.
Oxalis Valdiviana.—The leaves resemble those of the last species, and the movements of two leaflets (the main petioles of both having been secured) were traced during two days; but the tracings are not given, as they resembled that of O. acetosella, with the exception that the up and down oscillations were not so frequent during the day, and there was more lateral movement, so that broader ellipses were described. The leaves awoke early in the morning, for by 6.45 A.M. on June 12th and 13th they had not only risen to their full height, but had already begun to fall, that is, they were circumnutating. We have seen in the last chapter that the cotyledons, instead of sinking, rise up vertically at night.
Fig 129. Oxalis acetosella: circumnutation of leaflet when asleep; traced on vertical glass during 3 h. 40 m.
Oxalis Ortegesii.—The large leaves of this plant sleep like those of the previous species. The main petioles are long, and that of a young leaf rose 20° between noon and 10 P.M., whilst the petiole of an older leaf rose only 13°. Owing to this rising of the petioles, and the vertical sinking of the large leaflets, the leaves become crowded together at night, and the whole plant then exposes a much smaller surface to radiation than during the day.
Oxalis Plumierii.—In this species the three leaflets do not surround the summit of the petiole, but the terminal leaflet projects in the line of the petiole, with a lateral leaflet on each side. They all sleep by bending vertically downwards, but do not become at all folded. The petiole is rather long, and, one having been secured to a stick, the movement of the terminal leaflet was traced during 45 h. on a vertical glass. It moved in a very simple manner, sinking rapidly after 5 P.M., and rising rapidly early next morning. During the middle of the day it moved slowly and a little laterally. Consequently the ascending and descending lines did not coincide, and a single great ellipse was formed each day. There was no other evidence of circumnutation, and this fact is of interest, as we shall hereafter see.
Oxalis sensitiva.—The leaflets, as in the last species, bend vertically down at night, without becoming folded. The much elongated main petiole rises considerably in the evening, but in some very young plants the rise did not commence until late at night. We have seen that the cotyledons, instead of sinking like the leaflets, rise up vertically at night.
Oxalis bupleurifolia.—This species is rendered remarkable by the petioles being foliaceous, like the phyllodes of many Acacias. The leaflets are small, of a paler green and more tender consistence than the foliaceous petioles. The leaflet which was observed was .55 inch in length, and was borne by a petiole 2 inches long and .3 inch broad. It may be suspected that the leaflets are on the road to abortion or obliteration, as has actually occurred with those of another Brazilian species, O. rusciformis. Nevertheless, in the present species the nyctitropic movements are perfectly performed. The foliaceous petiole was first observed during 48 h., and found to be in continued circumnutation, as shown in the accompanying figure (Fig. 130). It rose during the day and early part of the night, and fell during the remainder of the night and early morning; but the movement was not sufficient to be called sleep. The ascending and descending lines did not coincide, so that an ellipse was formed each day. There was but little zigzagging; if the filament had been fixed longitudinally, we should probably have seen that there was more lateral movement than appears in the diagram.
Fig. 130. Oxalis bupleurifolia: circumnutation of foliaceous petiole, filament fixed obliquely across end of petiole; movements traced on vertical glass from 9 A.M. June 26th to 8.50 A.M. 28th. Apex of leaflet 4½ inches from the glass, so movement not much magnified. Plant 9 inches high, illuminated from above. Temp. 23½°–24½° C.
A terminal leaflet on another leaf was next observed (the petiole being secured), and its movements are shown in Fig. 131. During the day the leaflets are extended horizontally, and at night depend vertically; and as the petiole rises during the day the leaflets have to bend down in the evening more than 90°, so as to assume at night their vertical position. On the first day the leaflet simply moved up and down; on the second day it plainly circumnutated between 8 A.M. and 4.30 P.M., after which hour the great evening fall commenced.
Fig. 131. Oxalis bupleurifolia: circumnutation and nyctitropic movement of terminal leaflet, with filament affixed along the midrib; traced on a vertical glass from 9 A.M. on June 26th to 8.45 A.M. 28th. Conditions the same as in the last case.
Averrhoa bilimbi (Oxalidæ).—It has long been known,[5] firstly, that the leaflets in this genus sleep; secondly, that they move spontaneously during the day; and thirdly, that they are sensitive to a touch; but in none of these respects do they differ essentially from the species of Oxalis. They differ, however, as Mr. R. I. Lynch[6] has lately shown, in their spontaneous movements being strongly marked. In the case of A. bilimbi, it is a wonderful spectacle to behold on a warm sunny day the leaflets one after the other sinking rapidly downwards, and again ascending slowly. Their movements rival those of Desmodium gyrans. At night the leaflets hang vertically down; and now they are motionless, but this may be due to the opposite ones being pressed together (Fig. 132). The main petiole is in constant movement during the day, but no careful observations were made on it. The following diagrams are graphic representations of the variations in the angle, which a given leaflet makes with the vertical. The observations were made as follows. The plant growing in a pot was kept in a high temperature, the petiole of the leaf to be observed pointing straight at the observer, being separated from him by a vertical pane of glass. The petiole was secured so that the basal joint, or pulvinus, of one of the lateral leaflets was at the centre of a graduated arc placed close behind the leaflet. A fine glass filament was fixed to the leaf, so as to project like a continuation of the midrib. This filament acted as an index; and as the leaf rose and fell, rotating about its basal joint, its angular movement could be recorded by reading off at short intervals of time the position of the glass filament on the graduated arc. In order to avoid errors of parallax, all readings were made by looking through a small ring painted on the vertical glass, in a line with the joint of the leaflet and the centre of the graduated arc. In the following diagrams the ordinates represent the angles which the leaflet made with the vertical at successive instants.[7] It follows that a fall in the curve represents an actual dropping of the leaf, and that the zero line represents a vertically dependent position. Fig. 133 represents the nature of the movements which occur in the evening, as soon as the leaflets begin to assume their nocturnal position. At 4.55 P.M. the leaflet formed an angle of 85° with the vertical, or was only 5° below the horizontal; but in order that the diagram might get into our page, the leaflet is represented falling from 75° instead of 85°. Shortly after 6 P.M. it hung vertically down, and had attained its nocturnal position. Between 6.10 and 6.35 P.M. it performed a number of minute oscillations of about 2° each, occupying periods of 4 or 5 m. The complete state of rest of the leaflet which ultimately followed is not shown in the diagram. It is manifest that each oscillation consists of a gradual rise, followed by a sudden fall. Each time the leaflet fell, it approached nearer to the nocturnal position than it did on the previous fall. The amplitude of the oscillations diminished, while the periods of oscillation became shorter.
[5] Dr. Bruce, ‘Philosophical Trans.,’ 1785, p. 356.
[6] ‘Journal Linn. Soc.,’ vol. xvi. 1877, p. 231.
[7] In all the diagrams 1 mm. in the horizontal direction represents one minute of time. Each mm. in the vertical direction represents one degree of angular movement. In Figs. 133 and 134 the temperature is represented (along the ordinates) in the scale of 1 mm. to each 0.1 degree C. In Fig. 135 each mm. equals 0.2° F.
Fig. 132. Averrhoa bilimbi: leaf asleep; drawing reduced.
Fig. 133. Averrhoa bilimbi: angular movements of a leaflet during its evening descent, when going to sleep. Temp. 78°–81° F.
In bright sunshine the leaflets assume a highly inclined dependent position. A leaflet in diffused light was observed rising for 25 m. A blind was then pulled up so that the plant was brightly illuminated (BR in Fig. 134), and within a minute it began to fall, and ultimately fell 47°, as shown in the diagram. This descent was performed by six descending steps, precisely similar to those by which the nocturnal fall is effected. The plant was then again shaded (SH), and a long slow rise occurred until another series of falls commenced at BR’, when the sun was again admitted. In this experiment cool air was allowed to enter by the windows being opened at the same time that the blinds were pulled up, so that in spite of the sun shining on the plant the temperature was not raised.
The effect of an increase of temperature in diffused light is shown in Fig. 135. The temperature began to rise at 11.35 A.M. (in consequence of the fire being lighted), but by 12.42 a marked fall had occurred. It may be seen in the diagram that when the temperature was highest there were rapid oscillations of small amplitude, the mean position of the leaflet being at the time nearer the vertical. When the temperature began to fall, the oscillations became slower and larger, and the mean position of the leaf again approached the horizontal. The rate of oscillation was sometimes quicker than is represented in the above diagram. Thus, when the temperature was between 31° and 32° C., 14 oscillations of a few degrees occurred in 19 m. On the other hand, an oscillation may be much slower; thus a leaflet was observed (temperature 25° C.) to rise during 40 m. before it fell and completed its oscillation.
Fig. 134. Averrhoa bilimbi: angular movements of leaflet during a change from bright illumination to shade; temperature (broken line) remaining nearly the same.
Fig. 135. Averrhoa bilimbi: angular movement of leaflet during a change of temperature; light remaining the same. The broken line shows the change of temperature.
Fig. 136. Porlieria hygrometrica: circumnutation and nyctitropic movements of petiole of leaf, traced from 9.35 A.M. July 7th to about midnight on the 8th. Apex of leaf 7½ inches from the vertical glass. Temp. 19½°–20½° C.
Porlieria hygrometrica (Zygophylleæ).—The leaves of this plant (Chilian form) are from 1 to 1½ inch in length, and bear as many as 16 or 17 small leaflets on each side, which do not stand opposite one another. They are articulated to the petiole, and the petiole to the branch by a pulvinus. We must premise that apparently two forms are confounded under the same name: the leaves on a bush from Chili, which was sent to us from Kew, bore many leaflets, whilst those on plants in the Botanic Garden at Würzburg bore only 8 or 9 pairs; and the whole character of the bushes appeared somewhat different. We shall also see that they differ in a remarkable physiological peculiarity. On the Chilian plant the petioles of the younger leaves on upright branches, stood horizontally during the day, and at night sank down vertically so as to depend parallel and close to the branch beneath. The petioles of rather older leaves did not become at night vertically depressed, but only highly inclined. In one instance we found a branch which had grown perpendicularly downwards, and the petioles on it moved in the same direction relatively to the branch as just stated, and therefore moved upwards. On horizontal branches the younger petioles likewise move at night in the same direction as before, that is, towards the branch, and are consequently then extended horizontally; but it is remarkable that the older petioles on the same branch, though moving a little in the same direction, also bend downwards; they thus occupy a somewhat different position, relatively to the centre of the earth and to the branch, from that of the petioles on the upright branches. With respect to the leaflets, they move at night towards the apex of the petiole until their midribs stand nearly parallel to it; and they then lie neatly imbricated one over the other. Thus half of the upper surface of each leaflet is in close contact with half of the lower surface of the one next in advance; and all the leaflets, excepting the basal ones, have the whole of their upper surfaces and half of their lower surfaces well protected. Those on the opposite sides of the same petiole do not come into close contact at night, as occurs with the leaflets of so many Leguminosae but are separated by an open furrow; nor could they exactly coincide, as they stand alternately with respect to one another.
The circumnutation of the petiole of a leaf 3/4 of an inch in length, on an upright branch, was observed during 36h., and is shown in the preceding diagram (Fig. 136). On the first morning, the leaf fell a little and then rose until 1 P.M., and this was probably due to its being now illuminated through a skylight from above; it then circumnutated on a very small scale round the same spot until about 4 P.M., when the great evening fall commenced. During the latter part of the night or very early on the next morning the leaf rose again. On the second day it fell during the morning till 1 P.M., and this no doubt is its normal habit. From 1 to 4 P.M. it rose in a zigzag line, and soon afterwards the great evening fall commenced. It thus completed a double oscillation during the 24 h.
The specific name given to this plant by Ruiz and Pavon, indicates that in its native arid home it is affected in some manner by the dryness or dampness of the atmosphere.[8] In the Botanic Garden at Würzburg, there was a plant in a pot out of doors which was daily watered, and another in the open ground which was never watered. After some hot and dry weather there was a great difference in the state of the leaflets on these two plants; those on the unwatered plant in the open ground remaining half, or even quite, closed during the day. But twigs cut from this bush, with their ends standing in water, or wholly immersed in it, or kept in damp air under a bell-glass, opened their leaves though exposed to a blazing sun; whilst those on the plant in the ground remained closed. The leaves on this same plant, after some heavy rain, remained open for two days; they then became half closed during two days, and after an additional day were quite closed. This plant was now copiously watered, and on the following morning the leaflets were fully expanded. The other plant growing in a pot, after having been exposed to heavy rain, was placed before a window in the Laboratory, with its leaflets open, and they remained so during the daytime for 48 h.; but after an additional day were half closed. The plant was then watered, and the leaflets on the two following days remained open. On the third day they were again half closed, but on being again watered remained open during the two next days. From these several facts we may conclude that the plant soon feels the want of water; and that as soon as this occurs, it partially or quite closes its leaflets, which in their then imbricated condition expose a small surface to evaporation. It is therefore probable that this sleep-like movement, which occurs only when the ground is dry, is an adaptation against the loss of moisture.
[8] ‘Systema Veg. Florae Peruvianae et Chilensis,’ tom. i. p. 95, 1798. We cannot understand the account given by the authors of the behaviour of this plant in its native home. There is much about its power of foretelling changes in the weather; and it appears as if the brightness of the sky largely determined the opening and closing of the leaflets.
A bush about 4 feet in height, a native of Chili, which was thickly covered with leaves, behaved very differently, for during the day it never closed its leaflets. On July 6th the earth in the small pot in which it grew appeared extremely dry, and it was given a very little water. After 21 and 22 days (on the 27th and 28th), during the whole of which time the plant did not receive a drop of water, the leaves began to droop, but they showed no signs of closing during the day. It appeared almost incredible that any plant, except a fleshy one, could have kept alive in soil so dry, which resembled the dust on a road. On the 29th, when the bush was shaken, some leaves fell off, and the remaining ones were unable to sleep at night. It was therefore moderately watered, as well as syringed, late in the evening. On the next morning (30th) the bush looked as fresh as ever, and at night the leaves went to sleep. It may be added that a small branch while growing on the bush was enclosed, by means of a curtain of bladder, during 13 days in a large bottle half full of quicklime, so that the air within must have been intensely dry; yet the leaves on this branch did not suffer in the least, and did not close at all during the hottest days. Another trial was made with the same bush on August 2nd and 6th (the soil appearing at this latter date extremely dry), for it was exposed out of doors during the whole day to the wind, but the leaflets showed no signs of closing. The Chilian form therefore differs widely from the one at Würzburg, in not closing its leaflets when suffering from the want of water; and it can live for a surprisingly long time without water.
Tropaeolum majus (?) (cultivated var.) (Tropaeoleae).—Several plants in pots stood in the greenhouse, and the blades of the leaves which faced the front-lights were during the day highly inclined and at night vertical; whilst the leaves on the back of the pots, though of course illuminated through the roof, did not become vertical at night. We thought, at first, that this difference in their positions was in some manner due to heliotropism, for the leaves are highly heliotropic. The true explanation, however, is that unless they are well illuminated during at least a part of the day they do not sleep at night; and a little difference in the degree of illumination determines whether or not they shall become vertical at night. We have observed no other so well-marked a case as this, of the influence of previous illumination on nyctitropic movements. The leaves present also another peculiarity in their habit of rising or awaking in the morning, being more strongly fixed or inherited than that of sinking or sleeping at night. The movements are caused by the bending of an upper part of the petiole, between ½ and 1 inch in length; but the part close to the blade, for about 1/4 of an inch in length, does not bend and always remains at right angles to the blade. The bending portion does not present any external or internal difference in structure from the rest of the petiole. We will now give the experiments on which the above conclusions are founded.
A large pot with several plants was brought on the morning of Sept. 3rd out of the greenhouse and placed before a north-east window, in the same position as before with respect to the light, as far as that was possible. On the front of the plants, 24 leaves were marked with thread, some of which had their blades horizontal, but the greater number were inclined at about 45°, beneath the horizon; at night all these, without exception, became vertical. Early on the following morning (4th) they reassumed their former positions, and at night again became vertical. On the 5th the shutters were opened at 6.15 A.M., and by 8.18 A.M., after the leaves had been illuminated for 2 h. 3 m. and had acquired their diurnal position, they were placed in a dark cupboard. They were looked at twice during the day and thrice in the evening, the last time at 10.30 P.M., and not one had become vertical. At 8 A.M. on the following morning (6th) they still retained the same diurnal position, and were now replaced before the north-east window. At night all the leaves which had faced the light had their petioles curved and their blades vertical; whereas none of the leaves on the back of the plants, although they had been moderately illuminated by the diffused light of the room, were vertical. They were now at night placed in the same dark cupboard; at 9 A.M. on the next morning (7th) all those which had been asleep had reassumed their diurnal position. The pot was then placed for 3 h. in the sunshine, so as to stimulate the plants; at noon they were placed before the same north-east window, and at night the leaves slept in the usual manner and awoke on the following morning. At noon on this day (8th) the plants, after having been left before the north-east window for 5 h. 45 m. and thus illuminated (though not brightly, as the sky was cloudy during the whole time), were replaced in the dark cupboard, and at 3 P.M. the position of the leaves was very little, if at all, altered, so that they are not quickly affected by darkness; but by 10.15 P.M. all the leaves which had faced the north-east sky during the 5 h. 45 m. of illumination stood vertical, whereas those on the back of the plant retained their diurnal position. On the following morning (9th) the leaves awoke as on the two former occasions in the dark, and they were kept in the dark during the whole day; at night a very few of them became vertical, and this was the one instance in which we observed any inherited tendency or habit in this plant to sleep at the proper time. That it was real sleep was shown by these same leaves reassuming their diurnal position on the following morning (10th) whilst still kept in the dark.
The pot was then (9.45 A.M. 10th) replaced, after having been kept for 36 h. in darkness, before the north-east window; and at night the blades of all the leaves (excepting a few on the back of the plants) became conspicuously vertical.
At 6.45 A.M. (11th) after the plants had been illuminated on the same side as before during only 25 m., the pot was turned round, so that the leaves which had faced the light now faced the interior of the room, and not one of these went to sleep at night; whilst some, but not many, of those which had formerly stood facing the back of the room and which had never before been well illuminated or gone to sleep, now assumed a vertical position at night. On the next day (12th) the plant was turned round into its original position, so that the same leaves faced the light as formerly, and these now went to sleep in the usual manner. We will only add that with some young seedlings kept in the greenhouse, the blades of the first pair of true leaves (the cotyledons being hypogean) stood during the day almost horizontally and at night almost vertically.
A few observations were subsequently made on the circumnutation of three leaves, whilst facing a north-east window; but the tracings are not given, as the leaves moved somewhat towards the light. It was, however, manifest that they rose and fell more than once during the daytime, the ascending and descending lines being in parts extremely zigzag. The nocturnal fall commenced about 7 P.M., and the leaves had risen considerably by 6.45 A.M. on the following morning.
Leguminosae.—This Family includes many more genera with sleeping species than all the other families put together. The number of the tribes to which each genus belongs, according to Bentham and Hooker’s arrangement, has been added.
Crotolaria (sp.?) (Tribe 2).—This plant is monophyllous, and we are informed by Mr. T. Thiselton Dyer that the leaves rise up vertically at night and press against the stem.
Lupinus (Tribe 2).—The palmate or digitate leaves of the species in this large genus sleep in three different manners. One of the simplest, is that all the leaflets become steeply inclined downwards at night, having been during the day extended horizontally. This is shown in the accompanying figures (Fig. 137), of a leaf of L. pilosus, as seen during the day from vertically above, and of another leaf asleep with the leaflets inclined downwards. As in this position they are crowded together, and as they do not become folded like those in the genus Oxalis, they cannot occupy a vertically dependent position; but they are often inclined at an angle of 50° beneath the horizon. In this species, whilst the leaflets are sinking, the petioles rise up, in two instances when the angles were measured to the extent of 23°. The leaflets of L. sub-carnosus and arboreus, which were horizontal during the day, sank down at night in nearly the same manner; the former to an angle of 38° and the latter of 36°, beneath the horizon; but their petioles did not move in any plainly perceptible degree. It is, however, quite possible, as we shall presently see, that if a large number of plants of the three foregoing and of the following species were to be observed at all seasons, some of the leaves would be found to sleep in a different manner.
Fig. 137. Lupinus pilosus: A, leaf seen from vertically above in daytime; B, leaf asleep, seen laterally at night.
In the two following species the leaflets, instead of moving downwards, rise at night. With L. Hartwegii some stood at noon at a mean angle of 36° above the horizon, and at night at 51°, thus forming together a hollow cone with moderately steep sides. The petiole of one leaf rose 14° and of a second 11° at night. With L. luteus a leaflet rose from 47° at noon to 65° above the horizon at night, and another on a distinct leaf rose from 45° to 69°. The petioles, however, sink at night to a small extent, viz., in three instances by 2°, 6°, and 9° 30 seconds. Owing to this movement of the petioles, the outer and longer leaflets have to bend up a little more than the shorter and inner ones, in order that all should stand symmetrically at night. We shall presently see that some leaves on the same individual plants of L. luteus sleep in a very different manner.
We now come to a remarkable position of the leaves when asleep, which is common to several species of Lupines. On the same leaf the shorter leaflets, which generally face the centre of the plant, sink at night, whilst the longer ones on the opposite side rise; the intermediate and lateral ones merely twisting on their own axes. But there is some variability with respect to which leaflets rise or fall. As might have been expected from such diverse and complicated movements, the base of each leaflet is developed (at least in the case of L. luteus) into a pulvinus. The result is that all the leaflets on the same leaf stand at night more or less highly inclined, or even quite vertically, forming in this latter case a vertical star. This occurs with the leaves of a species purchased under the name of L. pubescens; and in the accompanying figures we see at A (Fig. 138) the leaves in their diurnal position; and at B the same plant at night with the two upper leaves having their leaflets almost vertical. At C another leaf, viewed laterally, is shown with the leaflets quite vertical. It is chiefly or exclusively the youngest leaves which form at night vertical stars. But there is much variability in the position of the leaves at night on the same plant; some remaining with their leaflets almost horizontal, others forming more or less highly inclined or vertical stars, and some with all their leaflets sloping downwards, as in our first class of cases. It is also a remarkable fact, that although all the plants produced from the same lot of seeds were identical in appearance, yet some individuals at night had the leaflets of all their leaves arranged so as to form more or less highly inclined stars; others had them all sloping downwards and never forming a star; and others, again, retained them either in a horizontal position or raised them a little.
Fig. 138. Lupinus pubescens: A, leaf viewed laterally during the day; B, same leaf at night; C, another leaf with the leaflet forming a vertical star at night. Figures reduced.
We have as yet referred only to the different positions of the leaflets of L. pubescens at night; but the petioles likewise differ in their movements. That of a young leaf which formed a highly inclined star at night, stood at noon at 42° above the horizon, and during the night at 72°, so had risen 30°. The petiole of another leaf, the leaflets of which occupied a similar position at night, rose only 6°. On the other hand, the petiole of a leaf with all its leaflets sloping down at night, fell at this time 4°. The petioles of two rather older leaves were subsequently observed; both of which stood during the day at exactly the same angle, viz., 50° above the horizon, and one of these rose 7°–8°, and the other fell 3°–4° at night. We meet with cases like that of L. pubescens with some other species. On a single plant of L. mutabilis some leaves, which stood horizontally during the day, formed highly inclined stars at night, and the petiole of one rose 7°. Other leaves which likewise stood horizontally during the day, had at night all their leaflets sloping downwards at 46° beneath the horizon, but their petioles had hardly moved. Again, L. luteus offered a still more remarkable case, for on two leaves, the leaflets which stood at noon at about 45° above the horizon, rose at night to 65° and 69°, so that they formed a hollow cone with steep sides. Four leaves on the same plant, which had their leaflets horizontal at noon, formed vertical stars at night; and three other leaves equally horizontal at noon, had all their leaflets sloping downwards at night. So that the leaves on this one plant assumed at night three different positions. Though we cannot account for this fact, we can see that such a stock might readily give birth to species having widely different nyctitropic habits.
Little more need be said about the sleep of the species of Lupinus; several, namely, L. polyphyllus, nanus, Menziesii, speciosus, and albifrons, though observed out of doors and in the greenhouse, did not change the position of their leaves sufficiently at night to be said to sleep. From observations made on two sleeping species, it appears that, as with Tropaeolum majus, the leaves must be well illuminated during the day in order to sleep at night. For several plants, kept all day in a sitting-room with north-east windows, did not sleep at night; but when the pots were placed on the following day out of doors, and were brought in at night, they slept in the usual manner. the trial was repeated on the following day and night with the same result.
Some observations were made on the circumnutation of the leaves of L. luteus and arboreus. It will suffice to say that the leaflets of the latter exhibited a double oscillation in the course of 24 h.; for they fell from the early morning until 10.15 A.M., then rose and zigzagged greatly till 4 P.M., after which hour the great nocturnal fall commenced. By 8 A.M. on the following morning the leaflets had risen to their proper height. We have seen in the fourth chapter, that the leaves of Lupinus speciosus, which do not sleep, circumnutate to an extraordinary extent, making many ellipses in the course of the day.
Cytisus (Tribe 2), Trigonella and Medicago (Tribe 3).—Only a few observations were made on these three genera. The petioles on a young plant, about a foot in height, of Cytisus fragrans rose at night, on one occasion 23° and on another 33°. The three leaflets also bend upwards, and at the same time approach each other, so that the base of the central leaflet overlaps the bases of the two lateral leaflets. They bend up so much that they press against the stem; and on looking down on one of these young plants from vertically above, the lower surfaces of the leaflets are visible; and thus their upper surfaces, in accordance with the general rule, are best protected from radiation. Whilst the leaves on these young plants were thus behaving, those on an old bush in full flower did not sleep at night.
Fig. 139. Medicago marina: A, leaves during the day; B, leaves asleep at night.
Trigonella Cretica resembles a Melilotus in its sleep, which will be immediately described. According to M. Royer,[9] the leaves of Medicago maculata rise up at night, and “se renversent un peu de manière à presenter obliquement au ciel leur face inférieure.” A drawing is here given (Fig. 139) of the leaves of M. marina awake and asleep; and this would almost serve for Cytisus fragrans in the same two states.
[9] ‘Annales des Sc. Nat. Bot.’ (5th series), ix. 1868, p. 368.
Melilotus (Tribe 3).—The species in this genus sleep in a remarkable manner. The three leaflets of each leaf twist through an angle of 90°, so that their blades stand vertically at night with one lateral edge presented to the zenith (Fig. 140). We shall best understand the other and more complicated movements, if we imagine ourselves always to hold the leaf with the tip of the terminal leaflet pointed to the north. The leaflets in becoming vertical at night could of course twist so that their upper surfaces should face to either side; but the two lateral leaflets always twist so that this surface tends to face the north, but as they move at the same time towards the terminal leaflet, the upper surface of the one faces about N.N.W., and that of the other N.N.E. The terminal leaflet behaves differently, for it twists to either side, the upper surface facing sometimes east and sometimes west, but rather more commonly west than east. The terminal leaflet also moves in another and more remarkable manner, for whilst its blade is twisting and becoming vertical, the whole leaflet bends to one side, and invariably to the side towards which the upper surface is directed; so that if this surface faces the west the whole leaflet bends to the west, until it comes into contact with the upper and vertical surface of the western lateral leaflet. Thus the upper surface of the terminal and of one of the two lateral leaflets is well protected.
The fact of the terminal leaflet twisting indifferently to either side and afterwards bending to the same side, seemed to us so remarkable, that we endeavoured to discover the cause. We imagined that at the commencement of the movement it might be determined by one of the two halves of the leaflet being a little heavier than the other. Therefore bits of wood were gummed on one side of several leaflets, but this produced no effect; and they continued to twist in the same direction as they had previously done. In order to discover whether the same leaflet twisted permanently in the same direction, black threads were tied to 20 leaves, the terminal leaflets of which twisted so that their upper surfaces faced west, and 14 white threads to leaflets which twisted to the east. These were observed occasionally during 14 days, and they all continued, with a single exception, to twist and bend in the same direction; for one leaflet, which had originally faced east, was observed after 9 days to face west. The seat of both the twisting and bending movement is in the pulvinus of the sub-petioles.
Fig. 140. Melilotus officinalis: A, leaf during the daytime. B, another leaf asleep. C, a leaf asleep as viewed from vertically above; but in this case the terminal leaflet did not happen to be in such close contact with the lateral one, as is usual.
We believe that the leaflets, especially the two lateral ones, in performing the above described complicated movements generally bend a little downwards; but we are not sure of this, for, as far as the main petiole is concerned, its nocturnal movement is largely determined by the position which the leaf happens to occupy during the day. Thus one main petiole was observed to rise at night 59°, whilst three others rose only 7° and 9°. The petioles and sub-petioles are continually circumnutating during the whole 24 h., as we shall presently see.
The leaves of the following 15 species, M. officinalis, suaveolens, parviflora, alba, infesta, dentata, gracilis, sulcata, elegans, coerulea, petitpierreana, macrorrhiza, Italica, secundiflora, and Taurica, sleep in nearly the same manner as just described; but the bending to one side of the terminal leaflet is apt to fail unless the plants are growing vigorously. With M. petitpierreana and secundiflora the terminal leaflet was rarely seen to bend to one side. In young plants of M. Italica it bent in the usual manner, but with old plants in full flower, growing in the same pot and observed at the same hour, viz., 8.30 P.M., none of the terminal leaflets on several scores of leaves had bent to one side, though they stood vertically; nor had the two lateral leaflets, though standing vertically, moved towards the terminal one. At 10.30 P.M., and again one hour after midnight, the terminal leaflets had become very slightly bent to one side, and the lateral leaflets had moved a very little towards the terminal one, so that the position of the leaflets even at this late hour was far from the ordinary one. Again, with M. Taurica the terminal leaflets were never seen to bend towards either of the two lateral leaflets, though these, whilst becoming vertical, had bent towards the terminal one. The sub-petiole of the terminal leaflet in this species is of unusual length, and if the leaflet had bent to one side, its upper surface could have come into contact only with the apex of either lateral leaflet; and this, perhaps, is the meaning of the loss of the lateral movement.
The cotyledons do not sleep at night. the first leaf consists of a single orbicular leaflet, which twists at night so that the blade stands vertically. It is a remarkable fact that with M. Taurica, and in a somewhat less degree with M. macrorrhiza and petitpierreana, all the many small and young leaves produced during the early spring from shoots on some cut-down plants in the greenhouse, slept in a totally different manner from the normal one; for the three leaflets, instead of twisting on their own axes so as to present their lateral edges to the zenith, turned upwards and stood vertically with their apices pointing to the zenith. They thus assumed nearly the same position as in the allied genus Trifolium; and on the same principle that embryological characters reveal the lines of descent in the animal kingdom, so the movements of the small leaves in the above three species of Melilotus, perhaps indicate that this genus is descended from a form which was closely allied to and slept like a Trifolium. Moreover, there is one species, M. messanensis, the leaves of which, on full-grown plants between 2 and 3 feet in height, sleep like the foregoing small leaves and like those of a Trifolium. We were so much surprised at this latter case that, until the flowers and fruit were examined, we thought that the seeds of some Trifolium had been sown by mistake instead of those of a Melilotus. It appears therefore probable that M. messanensis has either retained or recovered a primordial habit.
The circumnutation of a leaf of M. officinalis was traced, the stem being left free; and the apex of the terminal leaflet described three laterally extended ellipses, between 8 A.M. and 4 P.M.; after the latter hour the nocturnal twisting movement commenced. It was afterwards ascertained that the above movement was compounded of the circumnutation of the stem on a small scale, of the main petiole which moved most, and of the sub-petiole of the terminal leaflet. The main petiole of a leaf having been secured to a stick, close to the base of the sub-petiole of the terminal leaflet, the latter described two small ellipses between 10.30 A.M., and 2 P.M. At 7.15 P.M., after this same leaflet (as well as another) had twisted themselves into their vertical nocturnal position, they began to rise slowly, and continued to do so until 10.35 P.M., after which hour they were no longer observed.
As M. messanensis sleeps in an anomalous manner, unlike that of any other species in the genus, the circumnutation of a terminal leaflet, with the stem secured, was traced during two days. On each morning the leaflet fell, until about noon, and then began to rise very slowly; but on the first day the rising movement was interrupted between 1 and 3 P.M. by the formation of a laterally extended ellipse, and on the second day, at the same time, by two smaller ellipses. The rising movement then recommenced, and became rapid late in the evening, when the leaflet was beginning to go to sleep. The awaking or sinking movement had already commenced by 6.45 A.M. on both mornings.
Trifolium (Tribe 3).—The nyctitropic movements of 11 species were observed, and were found to be closely similar. If we select a leaf of T. repens having an upright petiole, and with the three leaflets expanded horizontally, the two lateral leaflets will be seen in the evening to twist and approach each other, until their upper surfaces come into contact. At the same time they bend downwards in a plane at right angles to that of their former position, until their midribs form an angle of about 45° with the upper part of the petiole. This peculiar change of position requires a considerable amount of torsion in the pulvinus. The terminal leaflet merely rises up without any twisting and bends over until it rests on and forms a roof over the edges of the now vertical and united lateral leaflets. Thus the terminal leaflet always passes through an angle of at least 90°, generally of 130° or 140°, and not rarely—as was often observed with T. subterraneum—of 180°. In this latter case the terminal leaflet stands at night horizontally (as in Fig. 141), with its lower surface fully exposed to the zenith. Besides the difference in the angles, at which the terminal leaflets stand at night in the individuals of the same species, the degree to which the lateral leaflets approach each other often likewise differs.
Fig. 141. Trifolium repens: A, leaf during the day; B, leaf asleep at night.
We have seen that the cotyledons of some species and not of others rise up vertically at night. The first true leaf is generally unifoliate and orbicular; it always rises, and either stands vertically at night or more commonly bends a little over so as to expose the lower surface obliquely to the zenith, in the same manner as does the terminal leaflet of the mature leaf. But it does not twist itself like the corresponding first simple leaf of Melilotus. With T. Pannonicum the first true leaf was generally unifoliate, but sometimes trifoliate, or again partially lobed and in an intermediate condition.
Circumnutation.—Sachs described in 1863[10] the spontaneous up and down movements of the leaflets of T. incarnatum, when kept in darkness. Pfeffer made many observations on the similar movements in T. pratense.[11] He states that the terminal leaflet of this species, observed at different times, passed through angles of from 30° to 120° in the course of from 1½ to 4 h. We observed the movements of T. subterraneum, resupinatum, and repens.
[10] ‘Flora,’ 1863, p. 497.
[11] ‘Die Period. Bewegungen,’ 1875, pp. 35, 52.
Trifolium subterraneum.—A petiole was secured close to the base of the three leaflets, and the movement of the terminal leaflet was traced during 26½ h., as shown in the figure on the next page.
Between 6.45 A.M. and 6 P.M. the apex moved 3 times up and 3 times down, completing 3 ellipses in 11 h. 15 m. The ascending and descending lines stand nearer to one another than is usual with most plants, yet there was some lateral motion. At 6 P.M. the great nocturnal rise commenced, and on the next morning the sinking of the leaflet was continued until 8.30 A.M., after which hour it circumnutated in the manner just described. In the figure the great nocturnal rise and the morning fall are greatly abbreviated, from the want of space, and are merely represented by a short curved line. The leaflet stood horizontally when at a point a little beneath the middle of the diagram; so that during the daytime it oscillated almost equally above and beneath a horizontal position. At 8.30 A.M. it stood 48° beneath the horizon, and by 11.30 A.M. it had risen 50° above the horizon; so that it passed through 98° in 3 h. By the aid of the tracing we ascertained that the distance travelled in the 3 h. by the apex of this leaflet was 1.03 inch. If we look at the figure, and prolong upwards in our mind’s eye the short curved broken line, which represents the nocturnal course, we see that the latter movement is merely an exaggeration or prolongation of one of the diurnal ellipses. The same leaflet had been observed on the previous day, and the course then pursued was almost identically the same as that here described.
Fig. 142. Trifolium subterraneum: circumnutation and nyctitropic movement of terminal leaflet (.68 inch in length), traced from 6.45 A.M. July 4th to 9.15 A.M. 5th. Apex of leaf 3 7/8 inches from the vertical glass, and movement, as here shown, magnified 5 1/4 times, reduced to one-half of original scale. Plant illuminated from above; temp. 16°–17° C.
Trifolium resupinatum.—A plant left entirely free was placed before a north-east window, in such a position that a terminal leaflet projected at right angles to the source of the light, the sky being uniformly clouded all day. The movements of this leaflet were traced during two days, and on both were closely similar. Those executed on the second day are shown in Fig. 143. The obliquity of the several lines is due partly to the manner in which the leaflet was viewed, and partly to its having moved a little towards the light. From 7.50 A.M. to 8.40 A.M. the leaflet fell, that is, the awakening movement was continued. It then rose and moved a little laterally towards the light. At 12.30 it retrograded, and at 2.30 resumed its original course, having thus completed a small ellipse during the middle of the day. In the evening it rose rapidly, and by 8 A.M. on the following morning had returned to exactly the same spot as on the previous morning. The line representing the nocturnal course ought to be extended much higher up, and is here abbreviated into a short, curved, broken line. The terminal leaflet, therefore, of this species described during the daytime only a single additional ellipse, instead of two additional ones, as in the case of T. subterraneum. But we should remember that it was shown in the fourth chapter that the stem circumnutates, as no doubt does the main petiole and the sub-petioles; so that the movement represented in Fig. 143 is a compounded one. We tried to observe the movements of a leaf kept during the day in darkness, but it began to go to sleep after 2 h. 15 m., and this was well pronounced after 4 h. 30 m.
Fig 143. Trifolium resupinatum: circumnutation and nyctitropic movements of the terminal leaflet during 24 hours.
Trifolium repens.—A stem was secured close to the base of a moderately old leaf, and the movement of the terminal leaflet was observed during two days. This case is interesting solely from the simplicity of the movements, in contrast with those of the two preceding species. On the first day the leaflet fell between 8 A.M. and 3 P.M., and on the second between 7 A.M. and 1 P.M. On both days the descending course was somewhat zigzag, and this evidently represents the circumnutating movement of the two previous species during the middle of the day. After 1 P.M., Oct. 1st (Fig. 144), the leaflet began to rise, but the movement was slow on both days, both before and after this hour, until 4 P.M. The rapid evening and nocturnal rise then commenced. Thus in this species the course during 24 h. consists of a single great ellipse; in T. resupinatum of two ellipses, one of which includes the nocturnal movement and is much elongated; and in T. subterraneum of three ellipses, of which the nocturnal one is likewise of great length.
Securigera coronilla (Tribe 4).—The leaflets, which stand opposite one another and are numerous, rise up at night, come into close contact, and bend backwards at a moderate angle towards the base of the petiole.
Fig. 144. Trifolium repens: circumnutation and nyctitropic movements of a nearly full-grown terminal leaflet, traced on a vertical glass from 7 A.M. Sept. 30th to 8 A.M. Oct. 1st. Nocturnal course, represented by curved broken line, much abbreviated.
Lotus (Tribe 4).—The nyctitropic movements of 10 species in this genus were observed, and found to be the same. The main petiole rises a little at night, and the three leaflets rise till they become vertical, and at the same time approach each other. This was conspicuous with L. Jacoboeus, in which the leaflets are almost linear. In most of the species the leaflets rise so much as to press against the stem, and not rarely they become inclined a little inwards with their lower surfaces exposed obliquely to the zenith. This was clearly the case with L. major, as its petioles are unusually long, and the leaflets are thus enabled to bend further inwards. The young leaves on the summits of the stems close up at night so much, as often to resemble large buds. The stipule-like leaflets, which are often of large size, rise up like the other leaflets, and press against the stem (Fig. 145). All the leaflets of L. Gebelii, and probably of the other species, are provided at their bases with distinct pulvini, of a yellowish colour, and formed of very small cells. The circumnutation of a terminal leaflet of L. peregrinus (with the stem secured) was traced during two days, but the movement was so simple that it is not worth while to give the diagram. The leaflet fell slowly from the early morning till about 1 P.M. It then rose gradually at first, but rapidly late in the evening. It occasionally stood still for about 20 m. during the day, and sometimes zigzagged a little. The movement of one of the basal, stipule-like leaflets was likewise traced in the same manner and at the same time, and its course was closely similar to that of the terminal leaflet.
Fig. 145. Lotus Creticus: A, stem with leaves awake during the day; B, with leaves asleep at night. SS, stipule-like leaflets.
In Tribe 5 of Bentham and Hooker, the sleep-movements of species in 12 genera have been observed by ourselves and others, but only in Robinia with any care. Psoralea acaulis raises its three leaflets at night; whilst Amorpha fruticosa,[12] Dalea alopecuroides, and Indigofera tinctoria depress them. Ducharte[13] states that Tephrosia caribaea is the sole example of “folioles couchées le long du pétiole et vers la base;” but a similar movement occurs, as we have already seen, and shall again see in other cases. Wistaria Sinensis, according to Royer,[14] “abaisse les folioles qui par une disposition bizarre sont inclinées dans la même feuille, les supérieures vers le sommet, les inférieures vers la base du petiole commun;” but the leaflets on a young plant observed by us in the greenhouse merely sank vertically downwards at night. The leaflets are raised in Sphaerophysa salsola, Colutea arborea, and Astragalus uliginosus, but are depressed, according to Linnæus, in Glycyrrhiza. The leaflets of Robinia pseudo-acacia likewise sink vertically down at night, but the petioles rise a little, viz., in one case 3°, and in another 4°. The circumnutating movements of a terminal leaflet on a rather old leaf were traced during two days, and were simple. The leaflet fell slowly, in a slightly zigzag line, from 8 A.M. to 5 P.M., and then more rapidly; by 7 A.M. on the following morning it had risen to its diurnal position. There was only one peculiarity in the movement, namely, that on both days there was a distinct though small oscillation up and down between 8.30 and 10 A.M., and this would probably have been more strongly pronounced if the leaf had been younger.
[12] Ducharte, ‘Eléments de Botanique’, 1867, p. 349.
[13] Ibid., p. 347.
[14] ‘Ann. des Sciences Nats. Bot.’ (5th series), ix. 1868.
Coronilla rosea (Tribe 6).—the leaves bear 9 or 10 pairs of opposite leaflets, which during the day stand horizontally, with their midribs at right angles to the petiole. At night they rise up so that the opposite leaflets come nearly into contact, and those on the younger leaves into close contact. At the same time they bend back towards the base of the petiole, until their midribs form with it angles of from 40° to 50° in a vertical plane, as here figured (Fig. 146). The leaflets, however, sometimes bend so much back that their midribs become parallel to and lie on the petiole. They thus occupy a reversed position to what they do in several Leguminosae, for instance, in Mimosa pudica; but, from standing further apart, they do not overlap one another nearly so much as in this latter plant. The main petiole is curved slightly downwards during the day, but straightens itself at night. In three cases it rose from 3° above the horizon at noon, to 9° at 10 P.M.; from 11° to 33°; and from 5° to 33°—the amount of angular movement in this latter case amounting to 28°. In several other species of Coronilla the leaflets showed only feeble movements of a similar kind.