Insectivorous Plants

with No. 5 that bits of meat had to be added thrice before all the short tentacles on the opposite side of the disc were inflected. The result was widely different when bits of meat were placed in a medial line at the distal or proximal ends of the disc. In three of the seventeen experi- ments thus made, owing either to the state of the leaf or to the smallness of the bit of meat, only the im- mediately adjoining tentacles were affected ; but in the other fourteen cases the tentacles at the opposite end of the leaf were inflected, though these were as distant from where the meat lay as were those on one side of the disc from the meat on the opposite side. In some of the present cases the tentacles on the sides were not at all affected, or in a less degree, or after a longer interval of time, than those at the opposite end. One pet of experiments is worth giving in fuller detail. Cubes of meat, not quite so small as those usually em- ployed, were placed on one side of the discs of four leaves, and cubes of the same size at the proximal or distal end of four other leaves. Now, when these two sets of leaves were compared after an interval of 24 hrs., they presented a striking difference. Those having the cubes on one side were very slightly affected on the opposite side ; whereas those with the cubes at either end had almost every tentacle at the opposite end, even the marginal ones, closely in- flected. After 48 hrs. the contrast in the state of the two sets was still great ; yet those with the meat 011 one side now had their discal and submarginal ten- tacles on the opposite side somewhat inflected, this being due to the large size of the cubes. Finally we may conclude from these thirty-five experiments, not to mention the six or seven previous ones, that the motor impulse is transmitted from any single gland

or small group of glands through the bade to the other tentacles more readily and effectually in a longitudinal than in a transverse direction. 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 nitro- genous 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 ex- cited. 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 in- flected, 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 subrnarginal 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 dis- charged at first with much greater force than after- wards. 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 impulso 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 con- tinued healthy for ten days after the operation. I therefore cut the glands off twenty-five tentacles, at different times and on different leaves, and seven- teen 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, how- ever, 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 under- gone 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 aggrega- tion, which in certain cases may be called reflex, and it is the only known instance in the vegetable king- dom. 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 papilla?, 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 aifected in exactly the same manner : the aggregation always com- mences in their glands, though these have not been directly excited, but have only received some influ- ence from the disc, as shown by their increased acid secretion. The protoplasm within the cells immedi- ately 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 aggre- gated, 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 011 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 con- tact 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,* and since read- ing 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 circum- ference of the disc, the

(Drotera rotundtfolia.) Leaf (enlarged) with the tentacles inflected directed movement of the well exhibited. An accu- rate drawing of a leaf with meat in this position is here reproduced (fig. 10), and we see the tentacles, in- cluding 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 180° 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 appear- ance of the above four leaves, each with their ten- tacles 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 remark- able ; 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 to- wards 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 accom- panying sketch (fig. 11) of a small leaf. Little vessels from the neigh- bouring 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

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 inoscu- lation between the two chief branches of the lateral bundle. The course of the vessels is very complex at the larger inosculation ; and here vessels, retain- ing 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 inoscula- tion. 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 ex- clusively 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 in. 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 dis- tinct 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 cer- tainly 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.*

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 ex- tremities paralysed. Excepting that the line between the two halves was here transverse instead of longitu- dinal, 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) * M. Ziegler made similar ex- '"'Comptes rendus,' 1874, p. 1417), {•eriments by cutting the spiral but arrived at conclusions \\idclj Teasels of Drosera intermedia different from mine.

proves that the spiral vessels of the cen tral 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 ten- tacles; so that the marginal ones are more closely connected together by spiral vessels than are tho others, and yet have much less power of communi- cating 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 stir- 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, pos- sible 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 ten- tacles ; it cannot, therefore, be sent along the fibro- vascular bundles. The effect of cutting the central vessels, in the above cases, in preventing the transmis- sion 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 Dionasa, that this same conclusion, namely that the motor impulse is not transmitted by the fibro-vascular bundles, is plainly confirmed ; and Professor 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 apace 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 desig- nated aggregation ; but to this subject I shall return. We further know that in the transmission of the aggre- gating 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 suc- cessive 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 ten- tacles 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 ten- tacles given by Dr. Warming,* 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 par- titions ; but rather fewer if down the inner parenchy- matous tissue. In either case it is remarkable that the impulse is able to pass through so many par- titions 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 T'^ 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 ex- terior rows of tentacles tends to travel laterally and towards the centre of the leaf, but not centrifugal! y, ia 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 in- flected tentacle, which was somewhat thinner than the bristle. The amount or extent, also, of the movement is great. Fully expanded tentacles in becoming in- flected SAveep through an angle of 180°; 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 half- way 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

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 con- tract* 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.t In the case of Drosera there is certainly much movement of the fluid throughout the tentacles whilst they are under- going 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 arc 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 oi

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 in- flection 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 im- mersed 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 some- times 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 180°. Professor Cohn, in his interesting paper* on the movements of the stamens of certain Composite, 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