Чарльз Дарвин
Insectivorous Plants
As long as the glands remain excited, and this may last for many days, even for eleven, as when in contact with phosphate of lime, they continue to transmit a motor impulse to the basal and bending parts of their own pedicels, for otherwise they would re-expand. The great difference in the length of time during which tentacles remain inflected over inorganic objects, and over objects of the same size containing soluble nitrogenous matter, proves the same fact. But the intensity of the impulse transmitted from an excited gland, which has begun to pour forth its acid secretion and is at the same time absorbing, seems to be very small compared with that which it transmits when first excited. Thus, when moderately large bits of meat were placed on one side of the disc, and the discal and sub-marginal tentacles on the opposite side became inflected, so that their glands at last touched the meat and absorbed matter from it, they did not transmit any motor influence to the exterior rows of tentacles on the same side, for these never became inflected. If, however, meat had been placed on the glands of these same tentacles before they had begun to secrete copiously and to absorb, they undoubtedly would have affected the exterior rows. Nevertheless, when I gave some phosphate of lime, which is a most powerful stimulant, to several submarginal tentacles already considerably inflected, but not yet in contact with some phosphate previously placed on two glands in the centre of the disc, the exterior tentacles on the same side were acted on.
When a gland is first excited, the motor impulse is discharged within a few seconds, as we know from the bending of the tentacle; and it appears to be discharged at first with much greater force than afterwards. Thus, in the case above given of a small fly naturally caught by a few glands on one side of a leaf, an impulse was slowly transmitted from them across the whole breadth of the leaf, causing the opposite tentacles to be temporarily inflected, but the glands which remained in contact with the insect, though they continued for several days to send an impulse down their own pedicels to the bending place, did not prevent the tentacles on the opposite side from quickly re-expanding; so that the motor discharge must at first have been more powerful than afterwards.
When an object of any kind is placed on the disc, and the surrounding tentacles are inflected, their glands secrete more copiously and the secretion becomes acid, so that some influence is sent to them from the discal glands. This change in the nature and amount of the secretion cannot depend on the bending of the tentacles, as the glands of the short central tentacles secrete acid when an object is placed on them, though they do not themselves bend. Therefore I inferred that the glands of the disc sent some influence up the surrounding tentacles to their glands, and that these reflected back a motor impulse to their basal parts; but this view was soon proved erroneous. It was found by many trials that tentacles with their glands closely cut off by sharp scissors often become inflected and again re-expand, still appearing healthy. One which was observed continued healthy for ten days after the operation. I therefore cut the glands off twenty-five tentacles, at different times and on different leaves, and seventeen of these soon became inflected, and afterwards re-expanded. The re-expansion commenced in about 8 hrs. or 9 hrs., and was completed in from 22 hrs. to 30 hrs. from the time of inflection. After an interval of a day or two, raw meat with saliva was placed on the discs of these seventeen leaves, and when observed next day, seven of the headless tentacles were inflected over the meat as closely as the uninjured ones on the same leaves; and an eighth headless tentacle became inflected after three additional days. The meat was removed from one of these leaves, and the surface washed with a little stream of water, and after three days the headless tentacle re-expanded for the second time. These tentacles without glands were, however, in a different state from those provided with glands and which had absorbed matter from the meat, for the protoplasm within the cells of the former had undergone far less aggregation. From these experiments with headless tentacles it is certain that the glands do not, as far as the motor impulse is concerned, act in a reflex manner like the nerve-ganglia of animals.
But there is another action, namely that of aggregation, which in certain cases may be called reflex, and it is the only known instance in the vegetable kingdom. We should bear in mind that the process does not depend on the previous bending of the tentacles, as we clearly see when leaves are immersed in certain strong solutions. Nor does it depend on increased secretion from the glands, and this is shown by several facts, more especially by the papillae, which do not secrete, yet undergoing aggregation, if given carbonate of ammonia or an infusion of raw meat. When a gland is directly stimulated in any way, as by the pressure of a minute particle of glass, the protoplasm within the cells of the gland first becomes aggregated, then that in the cells immediately beneath the gland, and so lower and lower down the tentacles to their bases; – that is, if the stimulus has been sufficient and not injurious. Now, when the glands of the disc are excited, the exterior tentacles are affected in exactly the same manner: the aggregation always commences in their glands, though these have not been directly excited, but have only received some influence from the disc, as shown by their increased acid secretion. The protoplasm within the cells immediately beneath the glands are next affected, and so downwards from cell to cell to the bases of the tentacles. This process apparently deserves to be called a reflex action, in the same manner as when a sensory nerve is irritated, and carries an impression to a ganglion which sends back some influence to a muscle or gland, causing movement or increased secretion; but the action in the two cases is probably of a widely different nature. After the protoplasm in a tentacle has been aggregated, its redissolution always begins in the lower part, and slowly travels up the pedicel to the gland, so that the protoplasm last aggregated is first redissolved. This probably depends merely on the protoplasm being less and less aggregated, lower and lower down in the tentacles, as can be seen plainly when the excitement has been slight. As soon, therefore, as the aggregating action altogether ceases, redissolution naturally commences in the less strongly aggregated matter in the lowest part of the tentacle, and is there first completed.
Direction of the Inflected Tentacles. – When a particle of any kind is placed on the gland of one of the outer tentacles, this invariably moves towards the centre of the leaf; and so it is with all the tentacles of a leaf immersed in any exciting fluid. The glands of the exterior tentacles then form a ring round the middle part of the disc, as shown in a previous figure (fig. 4, p. 10). The short tentacles within this ring still retain their vertical position, as they likewise do when a large object is placed on their glands, or when an insect is caught by them. In this latter case we can see that the inflection of the short central tentacles would be useless, as their glands are already in contact with their prey.
The result is very different when a single gland on one side of the disc is excited, or a few in a group. These send an impulse to the surrounding tentacles, which do not now bend towards the centre of the leaf, but to the point of excitement. We owe this capital observation to Nitschke,53 and since reading his paper a few years ago, I have repeatedly verified it. If a minute bit of meat be placed by the aid of a needle on a single gland, or on three or four together, halfway between the centre and the circumference of the disc, the directed movement of the surrounding tentacles is well exhibited. An accurate drawing of a leaf with meat in this position is here reproduced (fig. 10), and we see the tentacles, including some of the exterior ones, accurately directed to the point where the meat lay. But a much better plan is to place a particle of the phosphate of lime moistened with saliva on a single gland on one side of the disc of a large leaf, and another particle on a single gland on the opposite side. In four such trials the excitement was not sufficient to affect the outer tentacles, but all those near the two points were directed to them, so that two wheels were formed on the disc of the same leaf; the pedicels of the tentacles forming the spokes, and the glands united in a mass over the phosphate representing the axles. The precision with which each tentacle pointed to the particle was wonderful; so that in some cases I could detect no deviation from perfect accuracy. Thus, although the short tentacles in the middle of the disc do not bend when their glands are excited in a direct manner, yet if they receive a motor impulse from a point on one side, they direct themselves to the point equally well with the tentacles on the borders of the disc.
In these experiments, some of the short tentacles on the disc, which would have been directed to the centre, had the leaf been immersed in an exciting fluid, were now inflected in an exactly opposite direction, viz. towards the circumference. These tentacles, therefore, had deviated as much as 180o from the direction which they would have assumed if their own glands had been stimulated, and which may be considered as the normal one. Between this, the greatest possible and no deviation from the normal direction, every degree could be observed in the tentacles on these several leaves. Notwithstanding the precision with which the tentacles generally were directed, those near the circumference of one leaf were not accurately directed towards some phosphate of lime at a rather distant point on the opposite side of the disc. It appeared as if the motor impulse in passing transversely across nearly the whole width of the disc had departed somewhat from a true course. This accords with what we have already seen of the impulse travelling less readily in a transverse than in a longitudinal direction. In some other cases, the exterior tentacles did not seem capable of such accurate movement as the shorter and more central ones.
Nothing could be more striking than the appearance of the above four leaves, each with their tentacles pointing truly to the two little masses of the phosphate on their discs. We might imagine that we were looking at a lowly organised animal seizing prey with its arms. In the case of Drosera the explanation of this accurate power of movement, no doubt, lies in the motor impulse radiating in all directions, and whichever side of a tentacle it first strikes, that side contracts, and the tentacle consequently bends towards the point of excitement. The pedicels of the tentacles are flattened, or elliptic in section. Near the bases of the short central tentacles, the flattened or broad face is formed of about five longitudinal rows of cells; in the outer tentacles of the disc it consists of about six or seven rows; and in the extreme marginal tentacles of above a dozen rows. As the flattened bases are thus formed of only a few rows of cells, the precision of the movements of the tentacles is the more remarkable; for when the motor impulse strikes the base of a tentacle in a very oblique direction relatively to its broad face, scarcely more than one or two cells towards one end can be affected at first, and the contraction of these cells must draw the whole tentacle into the proper direction. It is, perhaps, owing to the exterior pedicels being much flattened that they do not bend quite so accurately to the point of excitement as the more central ones. The properly directed movement of the tentacles is not an unique case in the vegetable kingdom, for the tendrils of many plants curve towards the side which is touched; but the case of Drosera is far more interesting, as here the tentacles are not directly excited, but receive an impulse from a distant point; nevertheless, they bend accurately towards this point.
On the Nature of the Tissues through which the Motor Impulse is Transmitted. – It will be necessary first to describe briefly the course of the main fibro-vascular bundles. These are shown in the accompanying sketch (fig. 11) of a small leaf. Little vessels from the neighbouring bundles enter all the many tentacles with which the surface is studded; but these are not here represented. The central trunk, which runs up the footstalk, bifurcates near the centre of the leaf, each branch bifurcating again and again according to the size of the leaf. This central trunk sends off, low down on each side, a delicate branch, which may be called the sublateral branch. There is also, on each side, a main lateral branch or bundle, which bifurcates in the same manner as the others. Bifurcation does not imply that any single vessel divides, but that a bundle divides into two. By looking to either side of the leaf, it will be seen that a branch from the great central bifurcation inosculates with a branch from the lateral bundle, and that there is a smaller inosculation between the two chief branches of the lateral bundle. The course of the vessels is very complex at the larger inosculation; and here vessels, retaining the same diameter, are often formed by the union of the bluntly pointed ends of two vessels, but whether these points open into each other by their attached surfaces, I do not know. By means of the two inosculations all the vessels on the same side of the leaf are brought into some sort of connection. Near the circumference of the larger leaves the bifurcating branches also come into close union, and then separate again, forming a continuous zigzag line of vessels round the whole circumference. But the union of the vessels in this zigzag line seems to be much less intimate than at the main inosculation. It should be added that the course of the vessels differs somewhat in different leaves, and even on opposite sides of the same leaf, but the main inosculation is always present.
Now in my first experiments with bits of meat placed on one side of the disc, it so happened that not a single tentacle was inflected on the opposite side; and when I saw that the vessels on the same side were all connected together by the two inosculations, whilst not a vessel passed over to the opposite side, it seemed probable that the motor impulse was conducted exclusively along them.
In order to test this view, I divided transversely with the point of a lancet the central trunks of four leaves, just beneath the main bifurcation; and two days afterwards placed rather large bits of raw meat (a most powerful stimulant) near the centre of the disc above the incision – that is, a little towards the apex – with the following results: —
[(1) This leaf proved rather torpid: after 4 hrs. 40 m. (in all cases reckoning from the time when the meat was given) the tentacles at the distal end were a little inflected, but nowhere else; they remained so for three days, and re-expanded on the fourth day. The leaf was then dissected, and the trunk, as well as the two sublateral branches, were found divided.
(2) After 4 hrs. 30 m. many of the tentacles at the distal end were well inflected. Next day the blade and all the tentacles at this end were strongly inflected, and were separated by a distinct transverse line from the basal half of the leaf, which was not in the least affected. On the third day, however, some of the short tentacles on the disc near the base were very slightly inflected. The incision was found on dissection to extend across the leaf as in the last case.
(3) After 4 hrs. 30 m. strong inflection of the tentacles at the distal end, which during the next two days never extended in the least to the basal end. The incision as before.
(4) This leaf was not observed until 15 hrs. had elapsed, and then all the tentacles, except the extreme marginal ones, were found equally well inflected all round the leaf. On careful examination the spiral vessels of the central trunk were certainly divided; but the incision on one side had not passed through the fibrous tissue surrounding these vessels, though it had passed through the tissue on the other side.54]
The appearance presented by the leaves (2) and (3) was very curious, and might be aptly compared with that of a man with his backbone broken and lower extremities paralysed. Excepting that the line between the two halves was here transverse instead of longitudinal, these leaves were in the same state as some of those in the former experiments, with bits of meat placed on one side of the disc. The case of leaf (4) proves that the spiral vessels of the central trunk may be divided, and yet the motor impulse be transmitted from the distal to the basal end; and this led me at first to suppose that the motor force was sent through the closely surrounding fibrous tissue; and that if one half of this tissue was left undivided, it sufficed for complete transmission. But opposed to this conclusion is the fact that no vessels pass directly from one side of the leaf to the other, and yet, as we have seen, if a rather large bit of meat is placed on one side, the motor impulse is sent, though slowly and imperfectly, in a transverse direction across the whole breadth of the leaf. Nor can this latter fact be accounted for by supposing that the transmission is effected through the two inosculations, or through the circumferential zigzag line of union, for had this been the case, the exterior tentacles on the opposite side of the disc would have been affected before the more central ones, which never occurred. We have also seen that the extreme marginal tentacles appear to have no power to transmit an impulse to the adjoining tentacles; yet the little bundle of vessels which enters each marginal tentacle sends off a minute branch to those on both sides, and this I have not observed in any other tentacles; so that the marginal ones are more closely connected together by spiral vessels than are the others, and yet have much less power of communicating a motor impulse to one another.
But besides these several facts and arguments we have conclusive evidence that the motor impulse is not sent, at least exclusively, through the spiral vessels, or through the tissue immediately surrounding them. We know that if a bit of meat is placed on a gland (the immediately adjoining ones having been removed) on any part of the disc, all the short sur- rounding tentacles bend almost simultaneously with great precision towards it. Now there are tentacles on the disc, for instance near the extremities of the sublateral bundles (fig. 11), which are supplied with vessels that do not come into contact with the branches that enter the surrounding tentacles, except by a very long and extremely circuitous course. Nevertheless, if a bit of meat is placed on the gland of a tentacle of this kind, all the surrounding ones are inflected towards it with great precision. It is, of course, possible that an impulse might be sent through a long and circuitous course, but it is obviously impossible that the direction of the movement could be thus communicated, so that all the surrounding tentacles should bend precisely to the point of excitement. The impulse no doubt is transmitted in straight radiating lines from the excited gland to the surrounding tentacles; it cannot, therefore, be sent along the fibro-vascular bundles. The effect of cutting the central vessels, in the above cases, in preventing the transmission of the motor impulse from the distal to the basal end of a leaf, may be attributed to a considerable space of the cellular tissue having been divided. We shall hereafter see, when we treat of Dionaea, that this same conclusion, namely that the motor impulse is not transmitted by the fibro-vascular bundles, is plainly confirmed; and Prof. Cohn has come to the same conclusion with respect to Aldrovanda – both members of the Droseraceae.
As the motor impulse is not transmitted along the vessels, there remains for its passage only the cellular tissue; and the structure of this tissue explains to a certain extent how it travels so quickly down the long exterior tentacles, and much more slowly across the blade of the leaf. We shall also see why it crosses the blade more quickly in a longitudinal than in a transverse direction; though with time it can pass in any direction. We know that the same stimulus causes movement of the tentacles and aggregation of the protoplasm, and that both influences originate in and proceed from the glands within the same brief space of time. It seems therefore probable that the motor impulse consists of the first commencement of a molecular change in the protoplasm, which, when well developed, is plainly visible, and has been designated aggregation; but to this subject I shall return. We further know that in the transmission of the aggregating process the chief delay is caused by the passage of the transverse cell-walls; for as the aggregation travels down the tentacles, the contents of each successive cell seem almost to flash into a cloudy mass. We may therefore infer that the motor impulse is in like manner delayed chiefly by passing through the cell-walls.
The greater celerity with which the impulse is transmitted down the long exterior tentacles than across the disc may be largely attributed to its being closely confined within the narrow pedicel, instead of radiating forth on all sides as on the disc. But besides this confinement, the exterior cells of the tentacles are fully twice as long as those of the disc; so that only half the number of transverse partitions have to be traversed in a given length of a tentacle, compared with an equal space on the disc; and there would be in the same proportion less retardation of the impulse. Moreover, in sections of the exterior tentacles given by Dr. Warming,55 the parenchymatous cells are shown to be still more elongated; and these would form the most direct line of communication from the gland to the bending place of the tentacle. If the impulse travels down the exterior cells, it would have to cross from between twenty to thirty transverse partitions; but rather fewer if down the inner parenchymatous tissue. In either case it is remarkable that the impulse is able to pass through so many partitions down nearly the whole length of the pedicel, and to act on the bending place, in ten seconds. Why the impulse, after having passed so quickly down one of the extreme marginal tentacles (about 1/20 of an inch in length), should never, as far as I have seen, affect the adjoining tentacles, I do not understand. It may be in part accounted for by much energy being expended in the rapidity of the transmission.
Most of the cells of the disc, both the superficial ones and the larger cells which form the five or six underlying layers, are about four times as long as broad. They are arranged almost longitudinally, radiating from the footstalk. The motor impulse, therefore, when transmitted across the disc, has to cross nearly four times as many cell-walls as when transmitted in a longitudinal direction, and would consequently be much delayed in the former case. The cells of the disc converge towards the bases of the tentacles, and are thus fitted to convey the motor impulse to them from all sides. On the whole, the arrangement and shape of the cells, both those of the disc and tentacles, throw much light on the rate and manner of diffusion of the motor impulse. But why the impulse proceeding from the glands of the exterior rows of tentacles tends to travel laterally and towards the centre of the leaf, but not centrifugally, is by no means clear.
Mechanism of the Movements, and Nature of the Motor Impulse. – Whatever may be the means of movement, the exterior tentacles, considering their delicacy, are inflected with much force. A bristle, held so that a length of 1 inch projected from a handle, yielded when I tried to lift with it an inflected tentacle, which was somewhat thinner than the bristle. The amount or extent, also, of the movement is great. Fully expanded tentacles in becoming inflected sweep through an angle of 180o; and if they are beforehand reflexed, as often occurs, the angle is considerably greater. It is probably the superficial cells at the bending place which chiefly or exclusively contract; for the interior cells have very delicate walls, and are so few in number that they could hardly cause a tentacle to bend with precision to a definite point. Though I carefully looked, I could never detect any wrinkling of the surface at the bending place, even in the case of a tentacle abnormally curved into a complete circle, under circumstances hereafter to be mentioned.
All the cells are not acted on, though the motor impulse passes through them. When the gland of one of the long exterior tentacles is excited, the upper cells are not in the least affected; about halfway down there is a slight bending, but the chief movement is confined to a short space near the base; and no part of the inner tentacles bends except the basal portion. With respect to the blade of the leaf, the motor impulse may be transmitted through many cells, from the centre to the circumference, without their being in the least affected, or they may be strongly acted on and the blade greatly inflected. In the latter case the movement seems to depend partly on the strength of the stimulus, and partly on its nature, as when leaves are immersed in certain fluids.
The power of movement which various plants possess, when irritated, has been attributed by high authorities to the rapid passage of fluid out of certain cells, which, from their previous state of tension, immediately contract.56 Whether or not this is the primary cause of such movements, fluid must pass out of closed cells when they contract or are pressed together in one direction, unless they at the same time expand in some other direction. For instance, fluid can be seen to ooze from the surface of any young and vigorous shoot if slowly bent into a semi-circle. In the case of Drosera there is certainly much movement of the fluid throughout the tentacles whilst they are undergoing inflection. Many leaves can be found in which the purple fluid within the cells is of an equally dark tint on the upper and lower sides of the tentacles, extending also downwards on both sides to equally near their bases. If the tentacles of such a leaf are excited into movement, it will generally be found after some hours that the cells on the concave side are much paler than they were before, or are quite colourless, those on the convex side having become much darker. In two instances, after particles of hair had been placed on glands, and when in the course of 1 hr. 10 m. the tentacles were incurved halfway towards the centre of the leaf, this change of colour in the two sides was conspicuously plain. In another case, after a bit of meat had been placed on a gland, the purple colour was observed at intervals to be slowly travelling from the upper to the lower part, down the convex side of the bending tentacle. But it does not follow from these observations that the cells on the convex side become filled with more fluid during the act of inflection than they contained before; for fluid may all the time be passing into the disc or into the glands which then secrete freely.
The bending of the tentacles, when leaves are immersed in a dense fluid, and their subsequent re-expansion in a less dense fluid, show that the passage of fluid from or into the cells can cause movements like the natural ones. But the inflection thus caused is often irregular; the exterior tentacles being sometimes spirally curved. Other unnatural movements are likewise caused by the application of dense fluids, as in the case of drops of syrup placed on the backs of leaves and tentacles. Such movements may be compared with the contortions which many vegetable tissues undergo when subjected to exosmose. It is therefore doubtful whether they throw any light on the natural movements.
If we admit that the outward passage of fluid is the cause of the bending of the tentacles, we must suppose that the cells, before the act of inflection, are in a high state of tension, and that they are elastic to an extraordinary degree; for otherwise their contraction could not cause the tentacles often to sweep through an angle of above 180o. Prof. Cohn, in his interesting paper57 on the movements of the stamens of certain Compositae, states that these organs, when dead, are as elastic as threads of india-rubber, and are then only half as long as they were when alive. He believes that the living protoplasm within their cells is ordinarily in a state of expansion, but is paralysed by irritation, or may be said to suffer temporary death; the elasticity of the cell-walls then coming into play, and causing the contraction of the stamens. Now the cells on the upper or concave side of the bending part of the tentacles of Drosera do not appear to be in a state of tension, nor to be highly elastic; for when a leaf is suddenly killed, or dies slowly, it is not the upper but the lower sides of the tentacles which contract from elasticity. We may, therefore, conclude that their movements cannot be accounted for by the inherent elasticity of certain cells, opposed as long as they are alive and not irritated by the expanded state of their contents.
A somewhat different view has been advanced by other physiologists – namely that the protoplasm, when irritated, contracts like the soft sarcode of the muscles of animals. In Drosera the fluid within the cells of the tentacles at the bending place appears under the microscope thin and homogeneous, and after aggregation consists of small, soft masses of matter, undergoing incessant changes of form and floating in almost colourless fluid. These masses are completely redissolved when the tentacles re-expand. Now it seems scarcely possible that such matter should have any direct mechanical power; but if through some molecular change it were to occupy less space than it did before, no doubt the cell-walls would close up and contract. But in this case it might be expected that the walls would exhibit wrinkles, and none could ever be seen. Moreover, the contents of all the cells seem to be of exactly the same nature, both before and after aggregation; and yet only a few of the basal cells contract, the rest of the tentacle remaining straight.
