SigPhi · Charles Darwin

The Power of Movement in Plants

Page 16 of 32

being produced before the leaves, are not protected by them from radiation, and consequently often fail to yield fruit. An excellent observer[10] has remarked that one variety of the cherry has the petals of its flowers much curled backwards, and after a severe frost all the stigmas were killed; whilst at the same time, in another variety with incurved petals, the stigmas were not in the least injured.

[3] This distinction was first pointed out (according to Pfeffer, ‘Die Periodischen Bewegungen der Blattorgane,’ 1875, p. 161) by Dassen in 1837.

[8] Martins in ‘Bull. Soc. Bot. de France,’ tom. xix. 1872. Wells, in his famous ‘Essay on Dew,’ remarks that an exposed thermometer rises as soon as even a fleecy cloud, high in the sky, passes over the zenith.

This view that the sleep of leaves saves them from being chilled at night by radiation, would no doubt have occurred to Linnæus, had the principle of radiation been then discovered; for he suggests in many parts of his ‘Somnus Plantarum’ that the position of the leaves at night protects the young stems and buds, and often the young inflorescence, against cold winds. We are far from doubting that an additional advantage may be thus gained; and we have observed with several plants, for instance, Desmodium gyrans, that whilst the blade of the leaf sinks vertically down at night, the petiole rises, so that the blade has to move through a greater angle in order to assume its vertical position than would otherwise have been necessary; but with the result that all the leaves on the same plant are crowded together as if for mutual protection.

We doubted at first whether radiation would affect in any important manner objects so thin as are many cotyledons and leaves, and more especially affect differently their upper and lower surfaces; for although the temperature of their upper surfaces would undoubtedly fall when freely exposed to a clear sky, yet we thought that they would so quickly acquire by conduction the temperature of the surrounding air, that it could hardly make any sensible difference to them, whether they stood horizontally and radiated into the open sky, or vertically and radiated chiefly in a lateral direction towards neighbouring plants and other objects. We endeavoured, therefore, to ascertain something on this head by preventing the leaves of several plants from going to sleep, and by exposing to a clear sky when the temperature was beneath the freezing-point, these, as well as the other leaves on the same plants which had already assumed their nocturnal vertical position. Our experiments show that leaves thus compelled to remain horizontal at night, suffered much more injury from frost than those which were allowed to assume their normal vertical position. It may, however, be said that conclusions drawn from such observations are not applicable to sleeping plants, the inhabitants of countries where frosts do not occur. But in every country, and at all seasons, leaves must be exposed to nocturnal chills through radiation, which might be in some degree injurious to them, and which they would escape by assuming a vertical position.

In our experiments, leaves were prevented from assuming their nyctitropic position, generally by being fastened with the finest entomological pins (which did not sensibly injure them) to thin sheets of cork supported on sticks. But in some instances they were fastened down by narrow strips of card, and in others by their petioles being passed through slits in the cork. The leaves were at first fastened close to the cork, for as this is a bad conductor, and as the leaves were not exposed for long periods, we thought that the cork, which had been kept in the house, would very slightly warm them; so that if they were injured by the frost in a greater degree than the free vertical leaves, the evidence would be so much the stronger that the horizontal position was injurious. But we found that when there was any slight difference in the result, which could be detected only occasionally, the leaves which had been fastened closely down suffered rather more than those fastened with very long and thin pins, so as to stand from ½ to 3/4 inch above the cork. This difference in the result, which is in itself curious as showing what a very slight difference in the conditions influences the amount of injury inflicted, may be attributed, as we believe, to the surrounding warmer air not circulating freely beneath the closely pinned leaves and thus slightly warming them. This conclusion is supported by some analogous facts hereafter to be given.

We will now describe in detail the experiments which were tried. These were troublesome from our not being able to predict how much cold the leaves of the several species could endure. Many plants had every leaf killed, both those which were secured in a horizontal position and those which were allowed to sleep—that is, to rise up or sink down vertically. Others again had not a single leaf in the least injured, and these had to be re-exposed either for a longer time or to a lower temperature.

Oxalis acetosella.—A very large pot, thickly covered with between 300 and 400 leaves, had been kept all winter in the greenhouse. Seven leaves were pinned horizontally open, and were exposed on March 16th for 2 h. to a clear sky, the temperature on the surrounding grass being –4° C. (24° to 25° F.). Next morning all seven leaves were found quite killed, so were many of the free ones which had previously gone to sleep, and about 100 of them, either dead or browned and injured were picked off. Some leaves showed that they had been slightly injured by not expanding during the whole of the next day, though they afterwards recovered. As all the leaves which were pinned open were killed, and only about a third or fourth of the others were either killed or injured, we had some little evidence that those which were prevented from assuming their vertically dependent position suffered most.

The following night (17th) was clear and almost equally cold (–3° to –4° C. on the grass), and the pot was again exposed, but this time for only 30 m. Eight leaves had been pinned out, and in the morning two of them were dead, whilst not a single other leaf on the many plants was even injured.

On the 23rd the pot was exposed for 1 h. 30 m., the temperature on the grass being only –2° C., and not one leaf was injured: the pinned open leaves, however, all stood from ½ to 3/4 of an inch above the cork.

On the 24th the pot was again placed on the ground and exposed to a clear sky for between 35 m. and 40 m. By a mistake the thermometer was left on an adjoining sun-dial 3 feet high, instead of being placed on the grass; it recorded 25° to 26° F. (–3.3° to –3.8° C.), but when looked at after 1 h. had fallen to 22° F. (–5.5° C.); so that the pot was perhaps exposed to rather a lower temperature than on the two first occasions. Eight leaves had been pinned out, some close to the cork and some above it, and on the following morning five of them (i.e. 63 per cent.) were found killed. By counting a portion of the leaves we estimated that about 250 had been allowed to go to sleep, and of these about 20 were killed (i.e. only 8 per cent.), and about 30 injured.

Considering these cases, there can be no doubt that the leaves of this Oxalis, when allowed to assume their normal vertically dependent position at night, suffer much less from frost than those (23 in number) which had their upper surfaces exposed to the zenith.

Oxalis carnosa.—A plant of this Chilian species was exposed for 30 m.

to a clear sky, the thermometer on the grass standing at –2° C., with some of its leaves pinned open, and not one leaf on the whole bushy plant was in the least injured. On the 16th of March another plant was similarly exposed for 30 m., when the temperature on the grass was only a little lower, viz., –3° to –4° C. Six of the leaves had been pinned open, and next morning five of them were found much browned. The plant was a large one, and none of the free leaves, which were asleep and depended vertically, were browned, excepting four very young ones. But three other leaves, though not browned, were in a rather flaccid condition, and retained their nocturnal position during the whole of the following day. In this case it was obvious that the leaves which were exposed horizontally to the zenith suffered most. This same pot was afterwards exposed for 35–40 m. on a slightly colder night, and every leaf, both the pinned open and the free ones, was killed. It may be added that two pots of O. corniculata (var.

Atro-purpurea) were exposed for 2 h. and 3 h. to a clear sky with the temp. on grass –2° C., and none of the leaves, whether free or pinned open, were at all injured.

Arachis hypogoea.—Some plants in a pot were exposed at night for 30 m.

to a clear sky, the temperature on the surrounding grass being –2° C., and on two nights afterwards they were again exposed to the same temperature, but this time during 1 h. 30 m. On neither occasion was a single leaf, whether pinned open or free, injured; and this surprised us much, considering its native tropical African home. Two plants were next exposed (March 16th) for 30 m. to a clear sky, the temperature of the surrounding grass being now lower, viz., between –3° and –4° C., and all four pinned-open leaves were killed and blackened. These two plants bore 22 other and free leaves (excluding some very young bud-like ones) and only two of these were killed and three somewhat injured; that is, 23 per cent. were either killed or injured, whereas all four pinned-open leaves were utterly killed.

On another night two pots with several plants were exposed for between 35 m. and 40 m. to a clear sky, and perhaps to a rather lower temperature, for a thermometer on a dial, 3 feet high, close by stood at –3.3° to –3.8° C. In one pot three leaves were pinned open, and all were badly injured; of the 44 free leaves, 26 were injured, that is, 59 per cent. In the other pot 3 leaves were pinned open and all were killed; four other leaves were prevented from sleeping by narrow strips of stiff paper gummed across them, and all were killed; of 24 free leaves, 10 were killed, 2 much injured, and 12 unhurt; that is, 50 per cent. of the free leaves were either killed or much injured. Taking the two pots together, we may say that rather more than half of the free leaves, which were asleep, were either killed or injured, whilst all the ten horizontally extended leaves, which had been prevented from going to sleep, were either killed or much injured.

Cassia floribunda.—A bush was exposed at night for 40 m. to a clear sky, the temperature on the surrounding grass being –2° C., and not a leaf was injured.[11] It was again exposed on another night for 1 h., when the temperature of the grass was –4° C.; and now all the leaves on a large bush, whether pinned flat open or free, were killed, blackened, and shrivelled, with the exception of those on one small branch, low down, which was very slightly protected by the leaves on the branches above. Another tall bush, with four of its large compound leaves pinned out horizontally, was afterwards exposed (temp. of surrounding grass exactly the same, viz., –4° C.), but only for 30 m. On the following morning every single leaflet on these four leaves was dead, with both their upper and lower surfaces completely blackened. Of the many free leaves on the bush, only seven were blackened, and of these only a single one (which was a younger and more tender leaf than any of the pinned ones) had both surfaces of the leaflets blackened. The contrast in this latter respect was well shown by a free leaf, which stood between two pinned-open ones; for these latter had the lower surfaces of their leaflets as black as ink, whilst the intermediate free leaf, though badly injured, still retained a plain tinge of green on the lower surface of the leaflets. This bush exhibited in a striking manner the evil effects of the leaves not being allowed to assume at night their normal dependent position; for had they all been prevented from doing so, assuredly every single leaf on the bush would have been utterly killed by this exposure of only 30 m.

The leaves whilst sinking downwards in the evening twist round, so that the upper surface is turned inwards, and is thus better protected than the outwardly turned lower surface. Nevertheless, it was always the upper surface which was more blackened than the lower, whenever any difference could be perceived between them; but whether this was due to the cells near the upper surface being more tender, or merely to their containing more chlorophyll, we do not know.

[11] Cassia laevigata was exposed to a clear sky for 35 m., and C.

calliantha (a Guiana species) for 60 m., the temperature on the surrounding grass being –2° C., and neither was in the least injured.

But when C. laevigata was exposed for 1 h., the temp. on the surrounding grass being between –3° and –4° C., every leaf was killed.

Melilotus officinalis.—A large pot with many plants, which had been kept during the winter in the greenhouse, was exposed during 5 h. at night to a slight frost and clear sky. Four leaves had been pinned out, and these died after a few days; but so did many of the free leaves.

Therefore nothing certain could be inferred from this trial, though it indicated that the horizontally extended leaves suffered most. Another large pot with many plants was next exposed for 1 h., the temperature on the surrounding grass being lower, viz., -3° to –4° C. Ten leaves had been pinned out, and the result was striking, for on the following morning all these were found much injured or killed, and none of the many free leaves on the several plants were at all injured, with the doubtful exception of two or three very young ones.

Melilotus Italica.—Six leaves were pinned out horizontally, three with their upper and three with their lower surfaces turned to the zenith.

The plants were exposed for 5 h. to a clear sky, the temperature on ground being about –1° C. Next morning the six pinned-open leaves seemed more injured even than the younger and more tender free ones on the same branches. The exposure, however, had been too long, for after an interval of some days many of the free leaves seemed in almost as bad a condition as the pinned-out ones. It was not possible to decide whether the leaves with their upper or those with their lower surfaces turned to the zenith had suffered most.

Melilotus suaveolens.—Some plants with 8 leaves pinned out were exposed to a clear sky during 2 h., the temperature on the surrounding grass being –2° C. Next morning 6 out of these 8 leaves were in a flaccid condition. There were about 150 free leaves on the plant, and none of these were injured, except 2 or 3 very young ones. But after two days, the plants having been brought back into the greenhouse, the 6 pinned-out leaves all recovered.

Melilotus Taurica.—Several plants were exposed for 5 h. during two nights to a clear sky and slight frost, accompanied by some wind; and 5 leaves which had been pinned out suffered more than those both above and below on the same branches which had gone to sleep. Another pot, which had likewise been kept in the greenhouse, was exposed for 35–40 m. to a clear sky, the temperature of the surrounding grass being between –3° and –4° C. Nine leaves had been pinned out, and all of these were killed. On the same plants there were 210 free leaves, which had been allowed to go to sleep, and of these about 80 were killed, i.e. only 38 per cent.

Melilotus Petitpierreana.—The plants were exposed to a clear sky for 35–40 m.: temperature on surrounding grass –3° to –4° C. Six leaves had been pinned out so as to stand about ½ inch above the cork, and four had been pinned close to it. These 10 leaves were all killed, but the closely pinned ones suffered most, as 4 of the 6 which stood above the cork still retained small patches of a green colour. A considerable number, but not nearly all, of the free leaves, were killed or much injured, whereas all the pinned out ones were killed.

Melilotus macrorrhiza.—The plants were exposed in the same manner as in the last case. Six leaves had been pinned out horizontally, and five of them were killed, that is, 83 percent. We estimated that there were 200 free leaves on the plants, and of these about 50 were killed and 20 badly injured, so that about 35 per cent of the free leaves were killed or injured.

Lotus aristata.—Six plants were exposed for nearly 5 h. to a clear sky; temperature on surrounding grass –1.5° C. Four leaves had been pinned out horizontally, and 2 of these suffered more than those above or below on the same branches, which had been allowed to go to sleep. It is rather a remarkable fact that some plants of Lotus Jacoboeus, an inhabitant of so hot a country as the Cape Verde Islands, were exposed one night to a clear sky, with the temperature of the surrounding grass –2° C., and on a second night for 30 m. with the temperature of the grass between –3° and –4° C., and not a single leaf, either the pinned-out or free ones, was in the least injured.

Marsilea quadrifoliata.—A large plant of this species—the only Cryptogamic plant known to sleep—with some leaves pinned open, was exposed for 1 h. 35 m. to a clear sky, the temperature on the surrounding ground being –2° C., and not a single leaf was injured.

After an interval of some days the plant was again exposed for 1 h. to a clear sky, with the temperature on the surrounding ground lower, viz., –4° C. Six leaves had been pinned out horizontally, and all of them were utterly killed. The plant had emitted long trailing stems, and these had been wrapped round with a blanket, so as to protect them from the frozen ground and from radiation; but a very large number of leaves were left freely exposed, which had gone to sleep, and of these only 12 were killed. After another interval, the plant, with 9 leaves pinned out, was again exposed for 1 h., the temperature on the ground being again –4° C. Six of the leaves were killed, and one which did not at first appear injured afterwards became streaked with brown. The trailing branches, which rested on the frozen ground, had one-half or three-quarters of their leaves killed, but of the many other leaves on the plant, which alone could be fairly compared with the pinned-out ones, none appeared at first sight to have been killed, but on careful search 12 were found in this state. After another interval, the plant with 9 leaves pinned out, was exposed for 35–40 m. to a clear sky and to nearly the same, or perhaps a rather lower, temperature (for the thermometer by an accident had been left on a sun-dial close by), and 8 of these leaves were killed. Of the free leaves (those on the trailing branches not being considered), a good many were killed, but their number, compared with the uninjured ones, was small. Finally, taking the three trials together, 24 leaves, extended horizontally, were exposed to the zenith and to unobstructed radiation, and of these 20 were killed and 1 injured; whilst a relatively very small proportion of the leaves, which had been allowed to go to sleep with their leaflets vertically dependent, were killed or injured.

The cotyledons of several plants were prepared for trial, but the weather was mild and we succeeded only in a single instance in having seedlings of the proper age on nights which were clear and cold. The cotyledons of 6 seedlings of Mimosa pudica were fastened open on cork and were thus exposed for 1 h. 45 m. to a clear sky, with the temperature on the surrounding ground at 29° F.; of these, 3 were killed. Two other seedlings, after their cotyledons had risen up and had closed together, were bent over and fastened so that they stood horizontally, with the lower surface of one cotyledon fully exposed to the zenith, and both were killed. Therefore of the 8 seedlings thus tried 5, or more than half, were killed. Seven other seedlings with their cotyledons in their normal nocturnal position, viz., vertical and closed, were exposed at the same time, and of these only 2 were killed.[12] Hence it appears, as far as these few trials tell anything, that the vertical position at night of the cotyledons of Mimosa pudica protects them to a certain degree from the evil effects of radiation and cold.

[12] We were surprised that young seedlings of so tropical a plant as Mimosa pudica were able to resist, as well as they did, exposure for 1 hr. 45 m. to a clear sky, the temperature on the surrounding ground being 29° F. It may be added that seedlings of the Indian ‘Cassia pubescens’ were exposed for 1 h. 30 m. to a clear sky, with the temp.

on the surrounding ground at –2° C., and they were not in the least injured.

Concluding Remarks on the Radiation from Leaves at Night.—We exposed on two occasions during the summer to a clear sky several pinned-open leaflets of Trifolium pratense, which naturally rise at night, and of Oxalis purpurea, which naturally sink at night (the plants growing out of doors), and looked at them early on several successive mornings, after they had assumed their diurnal positions. The difference in the amount of dew on the pinned-open leaflets and on those which had gone to sleep was generally conspicuous; the latter being sometimes absolutely dry, whilst the leaflets which had been horizontal were coated with large beads of dew.

This shows how much cooler the leaflets fully exposed to the zenith must have become, than those which stood almost vertically, either upwards or downwards, during the night.

From the several cases above given, there can be no doubt that the position of the leaves at night affects their temperature through radiation to such a degree, that when exposed to a clear sky during a frost, it is a question of life and death. We may therefore admit as highly probable, seeing that their nocturnal position is so well adapted to lessen radiation, that the object gained by their often complicated sleep movements, is to lessen the degree to which they are chilled at night. It should be kept in mind that it is especially the upper surface which is thus protected, as it is never directed towards the zenith, and is often brought into close contact with the upper surface of an opposite leaf or leaflet.

We failed to obtain sufficient evidence, whether the better protection of the upper surface has been gained from its being more easily injured than the lower surface, or from its injury being a greater evil to the plant. That there is some difference in constitution between the two surfaces is shown by the following cases. Cassia floribunda was exposed to a clear sky on a sharp frosty night, and several leaflets which had assumed their nocturnal dependent position with their lower surfaces turned outwards so as to be exposed obliquely to the zenith, nevertheless had these lower surfaces less blackened than the upper surfaces which were turned inwards and were in close contact with those of the opposite leaflets. Again, a pot full of plants of Trifolium resupinatum, which had been kept in a warm room for three days, was turned out of doors (Sept. 21st) on a clear and almost frosty night. Next morning ten of the terminal leaflets were examined as opaque objects under the microscope. These leaflets, in going to sleep, either turn vertically upwards, or more commonly bend a little over the lateral leaflets, so that their lower surfaces are more exposed to the zenith than their upper surfaces. Nevertheless, six of these ten leaflets were distinctly yellower on the upper than on the lower and more exposed surface. In the remaining four, the result was not so plain, but certainly whatever difference there was leaned to the side of the upper surface having suffered most.

It has been stated that some of the leaflets experimented on were fastened close to the cork, and others at a height of from ½ to 3/4 of an inch above it; and that whenever, after exposure to a frost, any difference could be detected in their states, the closely pinned ones had suffered most. We attributed this difference to the air, not cooled by radiation, having been prevented from circulating freely beneath the closely pinned leaflets. That there was really a difference in the temperature of leaves treated in these two different methods, was plainly shown on one occasion; for after the exposure of a pot with plants of Melilotus dentata for 2 h. to a clear sky (the temperature on the surrounding grass being –2° C.), it was manifest that more dew had congealed into hoar-frost on the closely pinned leaflets, than on those which stood horizontally a little above the cork. Again, the tips of some few leaflets, which had been pinned close to the cork, projected a little beyond the edge, so that the air could circulate freely round them. This occurred with six leaflets of Oxalis acetosella, and their tips certainly suffered rather less then the rest of the same leaflets; for on the following morning they were still slightly green. The same result followed, even still more clearly, in two cases with leaflets of Melilotus officinalis which projected a little beyond the cork; and in two other cases some leaflets which were pinned close to the cork were injured, whilst other free leaflets on the same leaves, which had not space to rotate and assume their proper vertical position, were not at all injured.

Another analogous fact deserves notice: we observed on several occasions that a greater number of free leaves were injured on the branches which had been kept motionless by some of their leaves having been pinned to the corks, than on the other branches. This was conspicuously the case with those of Melilotus Petitpierreana, but the injured leaves in this instance were not actually counted. With Arachis hypogaea, a young plant with 7 stems bore 22 free leaves, and of these 5 were injured by the frost, all of which were on two stems, bearing four leaves pinned to the cork-supports. With Oxalis carnosa, 7 free leaves were injured, and every one of them belonged to a cluster of leaves, some of which had been pinned to the cork. We could account for these cases only by supposing that the branches which were quite free had been slightly waved about by the wind, and that their leaves had thus been a little warmed by the surrounding warmer air. If we hold our hands motionless before a hot fire, and then wave them about, we immediately feel relief; and this is evidently an analogous, though reversed, case. These several facts—in relation to leaves pinned close to or a little above the cork-supports—to their tips projecting beyond it—and to the leaves on branches kept motionless—seem to us curious, as showing how a difference, apparently trifling, may determine the greater or less injury of the leaves. We may even infer as probable that the less or greater destruction during a frost of the leaves on a plant which does not sleep, may often depend on the greater or less degree of flexibility of their petioles and of the branches which bear them.

NYCTITROPIC OR SLEEP MOVEMENTS OF COTYLEDONS.

We now come to the descriptive part of our work, and will begin with cotyledons, passing on to leaves in the next chapter. We have met with only two brief notices of cotyledons sleeping. Hofmeister,[13] after stating that the cotyledons of all the observed seedlings of the Caryophylleae (Alsineae and Sileneae) bend upwards at night (but to what angle he does not state), remarks that those of Stellaria media rise up so as to touch one another; they may therefore safely be said to sleep. Secondly, according to Ramey,[14] the cotyledons of Mimosa pudica and of Clianthus Dampieri rise up almost vertically at night and approach each other closely. It has been shown in a previous chapter that the cotyledons of a large number of plants bend a little upwards at night, and we here have to meet the difficult question at what inclination may they be said to sleep? According to the view which we maintain, no movement deserves to be called nyctitropic, unless it has been acquired for the sake of lessening radiation; but this could be discovered only by a long series of experiments, showing that the leaves of each species suffered from this cause, if prevented from sleeping. We must therefore take an arbitrary limit. If a cotyledon or leaf is inclined at 60° above or beneath the horizon, it exposes to the zenith about one-half of its area; consequently the intensity of its radiation will be lessened by about half, compared with what it would have been if the cotyledon or leaf had remained horizontal. This degree of diminution certainly would make a great difference to a plant having a tender constitution. We will therefore speak of a cotyledon and hereafter of a leaf as sleeping, only when it rises at night to an angle of about 60°, or to a still higher angle, above the horizon, or sinks beneath it to the same amount. Not but that a lesser diminution of radiation may be advantageous to a plant, as in the case of Datura stramonium, the cotyledons of which rose from 31° at noon to 55° at night above the horizon. The Swedish turnip may profit by the area of its leaves being reduced at night by about 30 per cent., as estimated by Mr. A. S.

Wilson; though in this case the angle through which the leaves rose was not observed. On the other hand, when the angular rise of cotyledons or of leaves is small, such as less than 30°, the diminution of radiation is so slight that it probably is of no significance to the plant in relation to radiation. For instance, the cotyledons of Geranium Ibericum rose at night to 27° above the horizon, and this would lessen radiation by only 11 per cent.: those of Linum Berendieri rose to 33°, and this would lessen radiation by 16 per cent.

There are, however, some other sources of doubt with respect to the sleep of cotyledons. In certain cases, the cotyledons whilst young diverge during the day to only a very moderate extent, so that a small rise at night, which we know occurs with the cotyledons of many plants, would necessarily cause them to assume a vertical or nearly vertical position at night; and in this case it would be rash to infer that the movement was effected for any special purpose. On this account we hesitated long whether we should introduce several Cucurbitaceous plants into the following list; but from reasons, presently to be given, we thought that 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 60° with the horizon, are arranged in the appended list on the same system as previously followed. The numbers of the Families, and with the Leguminosae the numbers of the Tribes, have been added to show how widely the plants in question are distributed throughout the dicotyledonous series. A few remarks will have to be made about many of the plants in the list. In doing so, it will be convenient not to follow strictly any systematic order, but to treat of the Oxalidæ and the Leguminosae at the close; for in these two Families the cotyledons are generally provided with a pulvinus, and their movements endure for a much longer time than those of the other plants in the list.

List of Seedling Plants, the cotyledons of which rise or sink at night to an angle of at least 60° above or beneath the horizon.

Brassica oleracea. Cruciferae (Fam. 14). — napus (as we are informed by Prof. Pfeffer). Raphanus sativus. Cruciferae. Githago segetum.

Caryophylleae (Fam. 26). Stellaria media (according to Hofmeister, as quoted). Caryophylleae. Anoda Wrightii. Malvaceae (Fam. 36). Gossypium (var. Nankin cotton). Malvaceae. Oxalis rosea. Oxalidæ (Fam. 41). — floribunda. — articulata. — Valdiviana. — sensitiva. Geranium rotundifolium. Geraniaceae (Fam. 47). Trifolium subterraneum.

Leguminosae (Fam. 75, Tribe 3). — strictum. — leucanthemum. Lotus ornithopopoides. Leguminosae (Tribe 4). — peregrinus. — Jacobæus.

Clianthus Dampieri. Leguminosae (Tribe 5)—according to M. Ramey.

Smithia sensitiva. Leguminosae (Tribe 6). Haematoxylon Campechianum.

Leguminosae (Tribe 13)—according to Mr. R. I. Lynch. Cassia mimosoides.

Leguminosae (Tribe 14). — glauca. — florida. — corymbosa. — pubescens.

— tora. — neglecta. — 3 other Brazilian unnamed species. Bauhinia (sp.?. Leguminosae (Tribe 15). Neptunia oleracea. Leguminosae (Tribe 20). Mimosa pudica. Leguminosae (Tribe 21). — albida. Cucurbita ovifera. Cucurbitaceæ (Fam. 106). — aurantia. Lagenaria vulgaris.

Cucurbitaceæ. Cucumis dudaim. Cucurbitaceæ. Apium petroselinum.

Umbelliferae (Fam. 113). — graveolens. Lactuca scariola. Compositæ (Fam. 122). Helianthus annuus (?). Compositæ. Ipomœa caerulea.

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