Чарльз Дарвин
Insectivorous Plants
Experiment 9. – A cube of 1/10 of an inch of very compact roasted beef was placed on a leaf, which opened spontaneously after twelve days; so much feebly acid secretion was left on the leaf that it trickled off. The meat was completely disintegrated, but not all dissolved; there was no mould. The little mass was placed under the microscope; some of the fibrillae in the middle still exhibited transverse striae; others showed not a vestige of striae; and every gradation could be traced between these two states. Globules, apparently of fat, and some undigested fibro-elastic tissue remained. The meat was thus in the same state as that formerly described, which was half digested by Drosera. Here, again, as in the case of albumen, the digestive process seems slower than in Drosera. At the opposite end of the same leaf, a firmly compressed pellet of bread had been placed; this was completely disintegrated, I suppose, owing to the digestion of the gluten, but seemed very little reduced in bulk.
Experiment 10. – A cube of 1/20 of an inch of cheese and another of albumen were placed at opposite ends of the same leaf. After nine days the lobes opened spontaneously a little at the end enclosing the cheese, but hardly any or none was dissolved, though it was softened and surrounded by secretion. Two days subsequently the end with the albumen also opened spontaneously (i.e. eleven days after it was put on), a mere trace in a blackened and dry condition being left.
Experiment 11. – The same experiment with cheese and albumen repeated on another and rather torpid leaf. The lobes at the end with the cheese, after an interval of six days, opened spontaneously a little; the cube of cheese was much softened, but not dissolved, and but little, if at all, reduced in size. Twelve hours afterwards the end with the albumen opened, which now consisted of a large drop of transparent, not acid, viscid fluid.
Experiment 12. – Same experiment as the two last, and here again the leaf at the end enclosing the cheese opened before the opposite end with the albumen; but no further observations were made.
Experiment 13. – A globule of chemically prepared casein, about 1/10 of an inch in diameter, was placed on a leaf, which spontaneously opened after eight days. The casein now consisted of a soft sticky mass, very little, if at all, reduced in size, but bathed in acid secretion.]
These experiments are sufficient to show that the secretion from the glands of Dionaea dissolves albumen, gelatine, and meat, if too large pieces are not given. Globules of fat and fibro-elastic tissue are not digested. The secretion, with its dissolved matter, if not in excess, is subsequently absorbed. On the other hand, although chemically prepared casein and cheese (as in the case of Drosera) excite much acid secretion, owing, I presume, to the absorption of some included albuminous matter, these substances are not digested, and are not appreciably, if at all, reduced in bulk.
[Effects of the Vapours of Chloroform, Sulphuric Ether, and Hydrocyanic Acid. – A plant bearing one leaf was introduced into a large bottle with a drachm (3.549 ml.) of chloroform, the mouth being imperfectly closed with cotton-wool. The vapour caused in 1 m. the lobes to begin moving at an imperceptibly slow rate; but in 3 m. the spikes crossed, and the leaf was soon completely shut. The dose, however, was much too large, for in between 2 and 3 hrs. the leaf appeared as if burnt, and soon died.
Two leaves were exposed for 30 m. in a 2-oz: vessel to the vapour of 30 minims (1.774 ml.) of sulphuric ether. One leaf closed after a time, as did the other whilst being removed from the vessel without being touched. Both leaves were greatly injured. Another leaf, exposed for 20 m. to 15 minims of ether, closed its lobes to a certain extent, and the sensitive filaments were now quite insensible. After 24 hrs. this leaf recovered its sensibility, but was still rather torpid. A leaf exposed in a large bottle for only 3 m. to ten drops was rendered insensible. After 52 m. it recovered its sensibility, and when one of the filaments was touched, the lobes closed. It began to reopen after 20 hrs. Lastly another leaf was exposed for 4 m. to only four drops of the ether; it was rendered insensible, and did not close when its filaments were repeatedly touched, but closed when the end of the open leaf was cut off. This shows either that the internal parts had not been rendered insensible, or that an incision is a more powerful stimulus than repeated touches on the filaments. Whether the larger doses of chloroform and ether, which caused the leaves to close slowly, acted on the sensitive filaments or on the leaf itself, I do not know.
Cyanide of potassium, when left in a bottle, generates prussic or hydrocyanic acid. A leaf was exposed for 1 hr. 35 m. to the vapour thus formed; and the glands became within this time so colourless and shrunken as to be scarcely visible, and I at first thought that they had all dropped off. The leaf was not rendered insensible; for as soon as one of the filaments was touched it closed. It had, however, suffered, for it did not reopen until nearly two days had passed, and was not even then in the least sensitive. After an additional day it recovered its powers, and closed on being touched and subsequently reopened. Another leaf behaved in nearly the same manner after a shorter exposure to this vapour.]
On the Manner in which Insects are caught. – We will now consider the action of the leaves when insects happen to touch one of the sensitive filaments. This often occurred in my greenhouse, but I do not know whether insects are attracted in any special way by the leaves. They are caught in large numbers by the plant in its native country. As soon as a filament is touched, both lobes close with astonishing quickness; and as they stand at less than a right angle to each other, they have a good chance of catching any intruder. The angle between the blade and footstalk does not change when the lobes close. The chief seat of movement is near the midrib, but is not confined to this part; for, as the lobes come together, each curves inwards across its whole breadth; the marginal spikes however, not becoming curved. This move- ment of the whole lobe was well seen in a leaf to which a large fly had been given, and from which a large portion had been cut off the end of one lobe; so that the opposite lobe, meeting with no resistance in this part, went on curving inwards much beyond the medial line. The whole of the lobe, from which a portion had been cut, was afterwards removed, and the opposite lobe now curled completely over, passing through an angle of from 120o to 130o, so as to occupy a position almost at right angles to that which it would have held had the opposite lobe been present.
From the curving inwards of the two lobes, as they move towards each other, the straight marginal spikes intercross by their tips at first, and ultimately by their bases. The leaf is then completely shut and encloses a shallow cavity. If it has been made to shut merely by one of the sensitive filaments having been touched, or if it includes an object not yielding soluble nitrogenous matter, the two lobes retain their inwardly concave form until they re-expand. The re-expansion under these circumstances – that is when no organic matter is enclosed – was observed in ten cases. In all of these, the leaves re-expanded to about two-thirds of the full extent in 24 hrs. from the time of closure. Even the leaf from which a portion of one lobe had been cut off opened to a slight degree within this same time. In one case a leaf re-expanded to about two-thirds of the full extent in 7 hrs., and completely in 32 hrs.; but one of its filaments had been touched merely with a hair just enough to cause the leaf to close. Of these ten leaves only a few re-expanded completely in less than two days, and two or three required even a little longer time. Before, however, they fully re-expand, they are ready to close instantly if their sensitive filaments are touched. How many times a leaf is capable of shutting and opening if no animal matter is left enclosed, I do not know; but one leaf was made to close four times, reopening afterwards, within six days, On the last occasion it caught a fly, and then remained closed for many days.
This power of reopening quickly after the filaments have been accidentally touched by blades of grass, or by objects blown on the leaf by the wind, as occasionally happens in its native place,63 must be of some importance to the plant; for as long as a leaf remains closed, it cannot of course capture an insect.
When the filaments are irritated and a leaf is made to shut over an insect, a bit of meat, albumen, gelatine, casein, and, no doubt, any other substance containing soluble nitrogenous matter, the lobes, instead of remaining concave, thus including a concavity, slowly press closely together throughout their whole breadth. As this takes place, the margins gradually become a little everted, so that the spikes, which at first intercrossed, at last project in two parallel rows. The lobes press against each other with such force that I have seen a cube of albumen much flattened, with distinct impressions of the little prominent glands; but this latter circumstance may have been partly caused by the corroding action of the secretion. So firmly do they become pressed together that, if any large insect or other object has been caught, a corresponding projection on the outside of the leaf is distinctly visible. When the two lobes are thus completely shut, they resist being opened, as by a thin wedge driven between them, with astonishing force, and are generally ruptured rather than yield. If not ruptured, they close again, as Dr. Canby informs me in a letter, "with quite a loud flap." But if the end of a leaf is held firmly between the thumb and finger, or by a clip, so that the lobes cannot begin to close, they exert, whilst in this position, very little force.
I thought at first that the gradual pressing together of the lobes was caused exclusively by captured insects crawling over and repeatedly irritating the sensitive filaments; and this view seemed the more probable when I learnt from Dr. Burdon Sanderson that whenever the filaments of a closed leaf are irritated, the normal electric current is disturbed. Nevertheless, such irritation is by no means necessary, for a dead insect, or a bit of meat, or of albumen, all act equally well; proving that in these cases it is the absorption of animal matter which excites the lobes slowly to press close together. We have seen that the absorption of an extremely small quantity of such matter also causes a fully expanded leaf to close slowly; and this movement is clearly analogous to the slow pressing together of the concave lobes. This latter action is of high functional importance to the plant, for the glands on both sides are thus brought into contact with a captured insect, and consequently secrete. The secretion with animal matter in solution is then drawn by capillary attraction over the whole surface of the leaf, causing all the glands to secrete and allowing them to absorb the diffused animal matter. The movement, excited by the absorption of such matter, though slow, suffices for its final purpose, whilst the movement excited by one of the sensitive filaments being touched is rapid, and this is indis- pensable for the capturing of insects. These two movements, excited by two such widely different means, are thus both well adapted, like all the other functions of the plant, for the purposes which they subserve.
There is another wide difference in the action of leaves which enclose objects, such as bits of wood, cork, balls of paper, or which have had their filaments merely touched, and those which enclose organic bodies yielding soluble nitrogenous matter. In the former case the leaves, as we have seen, open in under 24 hrs. and are then ready, even before being fully-expanded, to shut again. But if they have closed over nitrogen-yielding bodies, they remain closely shut for many days; and after re-expanding are torpid, and never act again, or only after a considerable interval of time. In four instances, leaves after catching insects never reopened, but began to wither, remaining closed – in one case for fifteen days over a fly; in a second, for twenty-four days, though the fly was small; in a third for twenty-four days over a woodlouse; and in a fourth, for thirty-five days over a large Tipula. In two other cases leaves remained closed for at least nine days over flies, and for how many more I do not know. It should, however, be added that in two instances in which very small insects had been naturally caught the leaf opened as quickly as if nothing had been caught; and I suppose that this was due to such small insects not having been crushed or not having excreted any animal matter, so that the glands were not excited. Small angular bits of albumen and gelatine were placed at both ends of three leaves, two of which remained closed for thirteen and the other for twelve days. Two other leaves remained closed over bits of meat for eleven days, a third leaf for eight days, and a fourth (but this had been cracked and injured) for only six days. Bits of cheese, or casein, were placed at one end and albumen at the other end of three leaves; and the ends with the former opened after six, eight, and nine days, whilst the opposite ends opened a little later. None of the above bits of meat, albumen, &c., exceeded a cube of 1/10 of an inch (2.54 mm.) in size, and were sometimes smaller; yet these small portions sufficed to keep the leaves closed for many days. Dr. Canby informs me that leaves remain shut for a longer time over insects than over meat; and from what I have seen, I can well believe that this is the case, especially if the insects are large.
In all the above cases, and in many others in which leaves remained closed for a long but unknown period over insects naturally caught, they were more or less torpid when they reopened. Generally they were so torpid during many succeeding days that no excitement of the filaments caused the least movement. In one instance, however, on the day after a leaf opened which had clasped a fly, it closed with extreme slowness when one of its filaments was touched; and although no object was left enclosed, it was so torpid that it did not re-open for the second time until 44 hrs. had elapsed. In a second case, a leaf which had expanded after remaining closed for at least nine days over a fly, when greatly irritated, moved one alone of its two lobes, and retained this unusual position for the next two days. A third case offers the strongest exception which I have observed; a leaf, after remaining clasped for an unknown time over a fly, opened, and when one of its filaments was touched, closed, though rather slowly. Dr. Canby, who observed in the United States a large number of plants which, although not in their native site, were probably more vigorous than my plants, informs me that he has "several times known vigorous leaves to devour their prey several times; but ordinarily twice, or, quite often, once was enough to render them unserviceable." Mrs. Treat, who cultivated many plants in New Jersey, also informs me that "several leaves caught successively three insects each, but most of them were not able to digest the third fly, but died in the attempt. Five leaves, however, digested each three flies, and closed over the fourth, but died soon after the fourth capture. Many leaves did not digest even one large insect." It thus appears that the power of digestion is somewhat limited, and it is certain that leaves always remain clasped for many days over an insect, and do not recover their power of closing again for many subsequent days. In this respect Dionaea differs from Drosera, which catches and digests many insects after shorter intervals of time.
We are now prepared to understand the use of the marginal spikes, which form so conspicuous a feature in the appearance of the plant (fig. 12, p. 287), and which at first seemed to me in my ignorance useless appendages. From the inward curvature of the lobes as they approach each other, the tips of the marginal spikes first intercross, and ultimately their bases. Until the edges of the lobes come into contact, elongated spaces between the spikes, varying from the 1/15 to the 1/10 of an inch (1.693 to 2.54 mm.) in breadth, according to the size of the leaf, are left open. Thus an insect, if its body is not thicker than these measurements, can easily escape between the crossed spikes, when disturbed by the closing lobes and in- creasing darkness; and one of my sons actually saw a small insect thus escaping. A moderately large insect, on the other hand, if it tries to escape between the bars will surely be pushed back again into its horrid prison with closing walls, for the spikes continue to cross more and more until the edges of the lobes come into contact. A very strong insect, however, would be able to free itself, and Mrs. Treat saw this effected by a rose-chafer (Macrodactylus subspinosus) in the United States. Now it would manifestly be a great disadvantage to the plant to waste many days in remaining clasped over a minute insect, and several additional days or weeks in afterwards recovering its sensibility; inasmuch as a minute insect would afford but little nutriment. It would be far better for the plant to wait for a time until a moderately large insect was captured, and to allow all the little ones to escape; and this advantage is secured by the slowly intercrossing marginal spikes, which act like the large meshes of a fishing-net, allowing the small and useless fry to escape.
As I was anxious to know whether this view was correct – and as it seems a good illustration of how cautious we ought to be in assuming, as I had done with respect to the marginal spikes, that any fully developed structure is useless – I applied to Dr. Canby. He visited the native site of the plant, early in the season, before the leaves had grown to their full size, and sent me fourteen leaves, containing naturally captured insects. Four of these had caught rather small insects, viz. three of them ants, and the fourth a rather small fly, but the other ten had all caught large insects, namely, five elaters, two chrysomelas, a curculio, a thick and broad spider, and a scolopendra. Out of these ten insects, no less than eight were beetles,64 and out of the whole fourteen there was only one, viz. a dipterous insect, which could readily take flight. Drosera, on the other hand, lives chiefly on insects which are good flyers, especially Diptera, caught by the aid of its viscid secretion. But what most concerns us is the size of the ten larger insects. Their average length from head to tail was .256 of an inch, the lobes of the leaves being on an average .53 of an inch in length, so that the insects were very nearly half as long as the leaves within which they were enclosed. Only a few of these leaves, therefore, had wasted their powers by capturing small prey, though it is probable that many small insects had crawled over them and been caught, but had then escaped through the bars.
The Transmission of the Motor Impulse, and Means of Movement. – It is sufficient to touch any one of the six filaments to cause both lobes to close, these becoming at the same time incurved throughout their whole breadth. The stimulus must therefore radiate in all directions from any one filament. It must also be transmitted with much rapidity across the leaf, for in all ordinary cases both lobes close simultaneously, as far as the eye can judge. Most physiologists believe that in irritable plants the excitement is transmitted along, or in close connection with, the fibro-vascular bundles. In Dionaea, the course of these vessels (composed of spiral and ordinary vascular tissue) seems at first sight to favour this belief; for they run up the midrib in a great bundle, sending off small bundles almost at right angles on each side. These bifurcate occasionally as they extend towards the margin, and close to the margin small branches from adjoining vessels unite and enter the marginal spikes. At some of these points of union the vessels form curious loops, like those described under Drosera. A continuous zigzag line of vessels thus runs round the whole circumference of the leaf, and in the midrib all the vessels are in close contact; so that all parts of the leaf seem to be brought into some degree of communication. Nevertheless, the presence of vessels is not necessary for the transmission of the motor impulse, for it is transmitted from the tips of the sensitive filaments (these being about the 1/20 of an inch in length), into which no vessels enter; and these could not have been overlooked, as I made thin vertical sections of the leaf at the bases of the filaments.
On several occasions, slits about the 1/10 of an inch in length were made with a lancet, close to the bases of the filaments, parallel to the midrib, and, therefore, directly across the course of the vessels. These were made sometimes on the inner and sometimes on the outer sides of the filaments; and after several days, when the leaves had reopened, these filaments were touched roughly (for they were always rendered in some degree torpid by the operation), and the lobes then closed in the ordinary manner, though slowly, and sometimes not until after a considerable interval of time. These cases show that the motor impulse is not transmitted along the vessels, and they further show that there is no necessity for a direct line of communication from the filament which is touched towards the midrib and opposite lobe, or towards the outer parts of the same lobe.
Two slits near each other, both parallel to the midrib, were next made in the same manner as before, one on each side of the base of a filament, on five distinct leaves, so that a little slip bearing a filament was connected with the rest of the leaf only at its two ends. These slips were nearly of the same size; one was carefully measured; it was .12 of an inch (3.048 mm.) in length, and .08 of an inch (2.032 mm.) in breadth; and in the middle stood the filament. Only one of these slips withered and perished. After the leaf had recovered from the operation, though the slits were still open, the filaments thus circumstanced were roughly touched, and both lobes, or one alone, slowly closed. In two instances touching the filament produced no effect; but when the point of a needle was driven into the slip at the base of the filament, the lobes slowly closed. Now in these cases the impulse must have proceeded along the slip in a line parallel to the midrib, and then have radiated forth, either from both ends or from one end alone of the slip, over the whole surface of the two lobes.
Again, two parallel slits, like the former ones, were made, one on each side of the base of a filament, at right angles to the midrib. After the leaves (two in number) had recovered, the filaments were roughly touched, and the lobes slowly closed; and here the impulse must have travelled for a short distance in a line at right angles to the midrib, and then have radiated forth on all sides over both lobes. These several cases prove that the motor impulse travels in all directions through the cellular tissue, independently of the course of the vessels.
With Drosera we have seen that the motor impulse is transmitted in like manner in all directions through the cellular tissue; but that its rate is largely governed by the length of the cells and the direction of their longer axes. Thin sections of a leaf of Dionaea were made by my son, and the cells, both those of the central and of the more superficial layers, were found much elongated, with their longer axes directed towards the midrib; and it is in this direction that the motor impulse must be sent with great rapidity from one lobe to the other, as both close simultaneously. The central parenchymatous cells are larger, more loosely attached together, and have more delicate walls than the more superficial cells. A thick mass of cellular tissue forms the upper surface of the midrib over the great central bundle of vessels.
When the filaments were roughly touched, at the bases of which slits had been made, either on both sides or on one side, parallel to the midrib or at right angles to it, the two lobes, or only one, moved. In one of these cases, the lobe on the side which bore the filament that was touched moved, but in three other cases the opposite lobe alone moved; so that an injury which was sufficient to prevent a lobe moving did not prevent the transmission from it of a stimulus which excited the opposite lobe to move. We thus also learn that, although normally both lobes move together, each has the power of independent movement. A case, indeed, has already been given of a torpid leaf that had lately re-opened after catching an insect, of which one lobe alone moved when irritated. Moreover, one end of the same lobe can close and re- expand, independently of the other end, as was seen in some of the foregoing experiments.
When the lobes, which are rather thick, close, no trace of wrinkling can be seen on any part of their upper surfaces, It appears therefore that the cells must contract. The chief seat of the movement is evidently in the thick mass of cells which overlies the central bundle of vessels in the midrib. To ascertain whether this part contracts, a leaf was fastened on the stage of the microscope in such a manner that the two lobes could not become quite shut, and having made two minute black dots on the midrib, in a transverse line and a little towards one side, they were found by the micrometer to be 17/1000 of an inch apart. One of the filaments was then touched and the lobes closed; but as they were prevented from meeting, I could still see the two dots, which now were 15/1000 of an inch apart, so that a small portion of the upper surface of the midrib had contracted in a transverse line 2/1000 of an inch (.0508 mm.).
We know that the lobes, whilst closing, become slightly incurved throughout their whole breadth. This movement appears to be due to the contraction of the superficial layers of cells over the whole upper surface. In order to observe their contraction, a narrow strip was cut out of one lobe at right angles to the midrib, so that the surface of the opposite lobe could be seen in this part when the leaf was shut. After the leaf had recovered from the operation and had re-expanded, three minute black dots were made on the surface opposite to the slit or window, in a line at right angles to the midrib. The distance between the dots was found to be 40/1000 of an inch, so that the two extreme dots were 80/1000 of an inch apart. One of the filaments was now touched and the leaf closed. On again measuring the distances between the dots, the two next to the midrib were nearer together by 1 to 2/1000 of an inch, and the two further dots by 3 to 4/1000 of an inch, than they were before; so that the two extreme dots now stood about 5/1000 of an inch (.127 mm.) nearer together than before. If we suppose the whole upper surface of the lobe, which was 400/1000 of an inch in breadth, to have contracted in the same proportion, the total contraction will have amounted to about 25/1000 or 1/40 of an inch (.635 mm.); but whether this is sufficient to account for the slight inward curvature of the whole lobe, I am unable to say.
Finally, with respect to the movement of the leaves, the wonderful discovery made by Dr. Burdon Sanderson65 is now universally known; namely that there exists a normal electrical current in the blade and footstalk; and that when the leaves are irritated, the current is disturbed in the same manner as takes place during the contraction of the muscle of an animal.
The Re-expansion of the Leaves. – This is effected at an insensibly slow rate, whether or not any object is enclosed. One lobe can re-expand by itself, as occurred with the torpid leaf of which one lobe alone had closed. We have also seen in the experiments with cheese and albumen that the two ends of the same lobe can re-expand to a certain extent independently of each other. But in all ordinary cases both lobes open at the same time. The re-expansion is not determined by the sensitive filaments; all three filaments on one lobe were cut off close to their bases; and the three Nuttall, in his 'Gen. American Plants,' p. 277 (note), says that, whilst collecting this plant in its native home, "I had occasion to observe that a detached leaf would make repeated efforts towards disclosing itself to the influence of the sun; these attempts consisted in an undulatory motion of the marginal ciliae, accompanied by a partial opening and succeeding collapse of the lamina, which at length terminated in a complete expansion and in the destruction of sensibility." I am indebted to Prof. Oliver for this reference; but I do not understand what took place.
leaves thus treated re-expanded, – one to a partial extent in 24 hrs., – a second to the same extent in 48 hrs., and the third, which had been previously injured, not until the sixth day. These leaves after their re-expansion closed quickly when the filaments on the other lobe were irritated. These were then cut off one of the leaves, so that none were left. This mutilated leaf, notwithstanding the loss of all its filaments, re-expanded in two days in the usual manner. When the filaments have been excited by immersion in a solution of sugar, the lobes do not expand so soon as when the filaments have been merely touched; and this, I presume, is due to their having been strongly affected through exosmose, so that they continue for some time to transmit a motor impulse to the upper surface of the leaf.
