The Power of Movement in Plants, Charles Darwin [the reading list book TXT] 📗

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 90o, so that each had risen 45o. A single cotyledon of another seedling was tapped in a like manner for 1 m., and it rose 27o in 9 m.; and after eight additional minutes it had risen 10o 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

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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 31o 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 8o. 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 60o. Several cotyledons of C. neglecta (likewise from S. Brazil) rose in from 5 m. to 15 m. to various angles between 16o and 34o, 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. 50o 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 34o; a second only 6o; a third 13o; and a fourth 17o. A cotyledon of C. florida similarly treated rose 9o; one of C. corymbosa rose 7 1/2o, and one of the very distinct C. mimosoides only 6o. 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 suborder 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 32o; they

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remained in this position for 15 m., but when looked at again 40 m. after the rubbing, each had fallen 14o. 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 30o. 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.*

 

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 48o; when looked at again in 35 m.

after being rubbed they had risen 4o 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 56o, a second 45o, a third almost 90o, and a fourth 90o. 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 sole notice which we have met with on the sensitiveness of cotyledons, relates to Mimosa; for Aug. P. De Candolle says (‘Phys. V�g.,’

1832, tom. ii. p. 865), “les cotyledons du M. pudica tendent � se raprocher par leurs faces sup�rieures lorsqu’on les irrite.”

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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 1 � 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.

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CHAPTER III.

 

SENSITIVENESS OF THE APEX OF THE RADICLE TO CONTACT AND TO OTHER IRRITANTS.

 

Manner in which radicles bend when they encounter an obstacle in the soil—

Vicia faba, tips of radicles highly sensitive to contact and other irritants—Effects of too high a temperature—Power of discriminating between objects attached on opposite sides—Tips of secondary radicles sensitive—Pisum, tips of radicles sensitive—Effects of such sensitiveness in overcoming geotropism—Secondary radicles—Phaseolus, tips of radicles hardly sensitive to contact, but highly sensitive to caustic and to the removal of a slice—Tropaeolum—Gossypium—Cucurbita—Raphanus—Aesculus, tip not sensitive to slight contact, highly sensitive to caustic—Quercus, tip highly sensitive to contact—Power of discrimination—Zea, tip highly sensitive, secondary radicles—Sensitiveness of radicles to moist air—

Summary of chapter.

 

IN order to see how the radicles of seedlings would pass over stones, roots, and other obstacles, which they must incessantly encounter in the soil, germinating beans (Vicia faba) were so placed that the tips of the radicles came into contact, almost rectangularly or at a high angle, with underlying plates of glass. In other cases the beans were turned about whilst their radicles were growing, so that they descended nearly vertically on their own smooth, almost flat, broad upper surfaces. The delicate root-cap, when it first touched any directly opposing surface, was a little flattened transversely; the flattening soon became oblique, and in a few hours quite disappeared, the apex now pointing at right angles, or at nearly right angles, to its former course. The radicle then seemed to glide in its new direction over the surface which had opposed [page 130]

it, pressing on it with very little force. How far such abrupt changes in its former course are aided by the circumnutation of the tip must be left doubtful. Thin slips of wood were cemented on more or less steeply inclined glass-plates, at right angles to the radicles which were gliding down them.

Straight lines had been painted along the growing terminal part of some of these radicles, before they met the opposing slip of wood; and the lines became sensibly curved in 2 h. after the apex had come into contact with the slips. In one case of a radicle, which was growing rather slowly, the root-cap, after encountering a rough slip of wood at right angles, was at first slightly flattened transversely: after an interval of 2 h. 30 m. the flattening became oblique; and after an additional 3 hours the flattening had wholly disappeared, and the apex now pointed at right angles to its former course. It then continued to grow in its new direction alongside the slip of wood, until it came to the end of it, round which it bent rectangularly. Soon afterwards when coming to the edge of the plate of glass, it was again bent at a large angle, and descended perpendicularly into the damp sand.

 

When, as