For instance, the cotyledons of Beta vulgaris, Solanum lycopersicum, Cerinthe major, and Lupinus luteus, when placed in darkness, moved down during the afternoon and early night, instead of rising as they would have done if they had been exposed to the light. All the individuals of the Solanum did not behave in the same manner, for the cotyledons of one circumnutated about the same spot between 2.30 and 10 P.M. The cotyledons of a seedling of Oxalis corniculata, which was feebly illuminated from above, moved downwards during the first morning in the normal manner, but on the second morning it moved upwards. The cotyledons of Lotus Jacoboeus were not affected by 4 h. of complete darkness, but when placed under a double skylight and thus feebly illuminated, they quite lost their periodical movements on the third morning. On the other hand, the cotyledons of Cucurbita ovifera moved in the normal manner during a whole day in darkness.
Seedlings of Githago segetum were feebly illuminated from above in the morning before their cotyledons had expanded, and they remained closed for the next 40 h. Other seedlings were placed in the dark after their cotyledons had opened in the morning and these did not begin to close until about 4 h. had elapsed. The cotyledons of Oxalis rosea sank vertically downwards after being left for 1 h. 20 m. in darkness; but those of some other species of Oxalis were not affected by several hours of darkness. The cotyledons of several species of Cassia are eminently susceptible to changes in the degree of light to which they are exposed: thus seedlings of an unnamed S. Brazilian species (a large and beautiful tree) were brought out of the hot-house and placed on a table in the middle of a room with two north-east and one north-west window, so that they were fairly well illuminated, though of course less so than in the hot-house, the day being moderately bright; and after 36 m. the cotyledons which had been horizontal rose up vertically and closed together as when asleep; after thus remaining on the table for 1 h. 13 m. they began to open. The cotyledons of young seedlings of another Brazilian species and of C. neglecta, treated in the same manner, behaved similarly, excepting that they did not rise up quite so much: they again became horizontal after about an hour.
Here is a more interesting case: seedlings of Cassia tora in two pots, which had stood for some time on the table in the room just described, had their cotyledons horizontal. One pot was now exposed for 2 h. to dull sunshine, and the cotyledons remained horizontal; it was then brought back to the table, and after 50 m. the cotyledons had risen 68° above the horizon. The other pot was placed during the same 2 h. behind a screen in the room, where the light was very obscure, and the cotyledons rose 63° above the horizon; the pot was then replaced on the table, and after 50 m. the cotyledons had fallen 33°. These two pots with seedlings of the same age stood close together, and were exposed to exactly the same amount of light, yet the cotyledons in the one pot were rising, whilst those in the other pot were at the same time sinking. This fact illustrates in a striking manner that their movements are not governed by the actual amount, but by a change in the intensity or degree of the light. A similar experiment was tried with two sets of seedlings, both exposed to a dull light, but different in degree, and the result was the same. The movements of the cotyledons of this Cassia are, however, determined (as in many other cases) largely by habit or inheritance, independently of light; for seedlings which had been moderately illuminated during the day, were kept all night and on the following morning in complete darkness; yet the cotyledons were partially open in the morning and remained open in the dark for about 6 h. The cotyledons in another pot, similarly treated on another occasion, were open at 7 A.M. and remained open in the dark for 4 h. 30 m., after which time they began to close. Yet these same seedlings, when brought in the middle of the day from a moderately bright into only a moderately dull light raised, as we have seen, their cotyledons high above the horizon.
Sensitiveness of Cotyledons to contact.—This subject does not possess much interest, as it is not known that sensitiveness of this kind is of any service to seedling plants. We have observed cases in only four genera, though we have vainly observed the cotyledons of many others. The genus cassia seems to be pre-eminent in this respect: thus, the cotyledons of C. tora, when extended horizontally, were both lightly tapped with a very thin twig for 3 m. and in the course of a few minutes they formed together an angle of 90°, so that each had risen 45°. A single cotyledon of another seedling was tapped in a like manner for 1 m., and it rose 27° in 9 m.; and after eight additional minutes it had risen 10° more; the opposite cotyledon, which was not tapped, hardly moved at all. The cotyledons in all these cases became horizontal again in less than half an hour. The pulvinus is the most sensitive part, for on slightly pricking three cotyledons with a pin in this part, they rose up vertically; but the blade was found also to be sensitive, care having been taken that the pulvinus was not touched. Drops of water placed quietly on these cotyledons produced no effect, but an extremely fine stream of water, ejected from a syringe, caused them to move upwards. When a pot of seedlings was rapidly hit with a stick and thus jarred, the cotyledons rose slightly. When a minute drop of nitric acid was placed on both pulvini of a seedling, the cotyledons rose so quickly that they could easily be seen to move, and almost immediately afterwards they began to fall; but the pulvini had been killed and became brown.
The cotyledons of an unnamed species of Cassia (a large tree from S. Brazil) rose 31° in the course of 26 m. after the pulvini and the blades had both been rubbed during 1 m. with a twig; but when the blade alone was similarly rubbed the cotyledons rose only 8°. The remarkably long and narrow cotyledons, of a third unnamed species from S. Brazil, did not move when their blades were rubbed on six occasions with a pointed stick for 30 s. or for 1 m.; but when the pulvinus was rubbed and slightly pricked with a pin, the cotyledons rose in the course of a few minutes through an angle of 6°o. Several cotyledons of C. neglecta (likewise from S. Brazil) rose in from 5 m. to 15 m. to various angles between 16v and 34°, after being rubbed during 1 m. with a twig. Their sensitiveness is retained to a somewhat advanced age, for the cotyledons of a little plant of C. neglecta, 34 days old and bearing three true leaves, rose when lightly pinched between the finger and thumb. Some seedlings were exposed for 30 m. to a wind (temp. 50° F.) sufficiently strong to keep the cotyledons vibrating, but this to our surprise did not cause any movement. The cotyledons of four seedlings of the Indian C. glauca were either rubbed with a thin twig for 2 m. or were lightly pinched: one rose 34°; a second only 6°; a third 13°; and a fourth 17°. A cotyledon of C. florida similarly treated rose 9°; one of C. corymbosa rose 7½°, and one of the very distinct C. mimosoides only 6°. Those of C. pubescens did not appear to be in the least sensitive; nor were those of C. nodosa, but these latter are rather thick and fleshy, and do not rise at night or go to sleep.
Smithia sensitiva.—This plant belongs to a distinct sub-order of the Leguminosae from Cassia. Both cotyledons of an oldish seedling, with the first true leaf partially unfolded, were rubbed for 1 m. with a fine twig, and in 5 m. each rose 32°; they remained in this position for 15 m., but when looked at again 40 m. after the rubbing, each had fallen 14°. Both cotyledons of another and younger seedling were lightly rubbed in the same manner for 1 m., and after an interval of 32 m. each had risen 30°. They were hardly at all sensitive to a fine jet of water. The cotyledons of S. Pfundii, an African water plant, are thick and fleshy; they are not sensitive and do not go to sleep.
Mimosa pudica and albida.—The blades of several cotyledons of both these plants were rubbed or slightly scratched with a needle during 1 m. or 2 m.; but they did not move in the least. When, however, the pulvini of six cotyledons of M. pudica were thus scratched, two of them were slightly raised. In these two cases perhaps the pulvinus was accidentally pricked, for on pricking the pulvinus of another cotyledon it rose a little. It thus appears that the cotyledons of Mimosa are less sensitive than those of the previously mentioned plants.[[29]]
[29] The sole notice which we have met with on the sensitiveness of cotyledons, relates to Mimosa; for Aug. P. De Candolle says (‘Phys. Vég.,’ 1832, tom. ii. p. 865), “les cotyledons du M. pudica tendent à se raprocher par leurs faces supérieures lorsqu’on les irrite.”
Oxalis sensitiva.—The blades and pulvini of two cotyledons, standing horizontally, were rubbed or rather tickled for 30 s. with a fine split bristle, and in 10 m. each had risen 48°; when looked at again in 35 m. after being rubbed they had risen 4° more; after 30 additional minutes they were again horizontal. On hitting a pot rapidly with a stick for 1 m., the cotyledons of two seedlings were considerably raised in the course of 11 m. A pot was carried a little distance on a tray and thus jolted; and the cotyledons of four seedlings were all raised in 10 m.; after 17 m. one had risen 56°, a second 45°, a third almost 90°, and a fourth 90°. After an additional interval of 40 m. three of them had re-expanded to a considerable extent. These observations were made before we were aware at what an extraordinarily rapid rate the cotyledons circumnutate, and are therefore liable to error. Nevertheless it is extremely improbable that the cotyledons in the eight cases given, should all have been rising at the time when they were irritated. The cotyledons of Oxalis Valdiviana and rosea were rubbed and did not exhibit any sensitiveness.
Finally, there seems to exist some relation between the habit of cotyledons rising vertically at night or going to sleep, and their sensitiveness, especially that of their pulvini, to a touch; for all the above-named plants sleep at night. On the other hand, there are many plants the cotyledons of which sleep, and are not in the least sensitive. As the cotyledons of several species of Cassia are easily affected both by slightly diminished light and by contact, we thought that these two kinds of sensitiveness might be connected; but this is not necessarily the case, for the cotyledons of Oxalis sensitiva did not rise when kept on one occasion for 1½ h., and on a second occasion for nearly 4 h., in a dark closet. Some other cotyledons, as those of Githago segetum, are much affected by a feeble light, but do not move when scratched by a needle. That with the same plant there is some relation between the sensitiveness of its cotyledons and leaves seems highly probable, for the above described Smithia and Oxalis have been called sensitiva, owing to their leaves being sensitive; and though the leaves of the several species of Cassia are not sensitive to a touch, yet if a branch be shaken or syringed with water, they partially assume their nocturnal dependent position. But the relation between the sensitiveness to contact of the cotyledons and of the leaves of the same plant is not very close, as may be inferred from the cotyledons of Mimosa pudica being only slightly sensitive, whilst the leaves are well known to be so in the highest degree. Again, the leaves of Neptunia oleracea are very sensitive to a touch, whilst the cotyledons do not appear to be so in any degree.