Insectivorous Plants

Chapter 20

If an insect is placed on the central glands, or has been naturally caught there, the apex of the leaf curls inwards. For instance, dead flies were placed on three leaves near their bases, and after 24 hrs.

the previously straight apices were curled completely over, so as to embrace and conceal the flies; they had therefore moved through an angle of 180o. After three days the apex of one leaf, together with the tentacles, began to re-expand. But as far as I have seen-- and I made many trials--the sides of the leaf are never inflected, and this is the one functional difference between this species and Drosera rotundifolia.

Drosera intermedia (Hayne).--This species is quite as common in some parts of England as Drosera rotundifolia. It differs from Drosera anglica, as far as the leaves are concerned, only in their smaller size, and in their tips being generally a little reflexed. They capture a large number of insects. The tentacles are excited into movement by all the causes above specified; and aggregation ensues, with movement of the protoplasmic ma.s.ses. I have seen, through a lens, a tentacle beginning to bend in less than a minute after a particle of raw meat had been placed on the gland. The apex of the leaf curls over an exciting object as in the case of Drosera anglica. Acid secretion is copiously poured over captured insects. A leaf which had embraced a fly with all its tentacles re-expanded after nearly three days.

Drosera capensis.--This species, a native of the Cape of Good Hope, was sent to me by Dr. Hooker. The leaves are elongated, slightly concave along the middle and taper towards the apex, [page 280] which is bluntly pointed and reflexed. They rise from an almost woody axis, and their greatest peculiarity consists in their foliaceous green footstalks, which are almost as broad and even longer than the gland-bearing blade. This species, therefore, probably draws more nourishment from the air, and less from captured insects, than the other species of the genus. Nevertheless, the tentacles are crowded together on the disc, and are extremely numerous; those on the margins being much longer than the central ones. All the glands have the same form; their secretion is extremely viscid and acid.

The specimen which I examined had only just recovered from a weak state of health. This may account for the tentacles moving very slowly when particles of meat were placed on the glands, and perhaps for my never succeeding in causing any movement by repeatedly touching them with a needle. But with all the species of the genus this latter stimulus is the least effective of any. Particles of gla.s.s, cork, and coal-cinders, were placed on the glands of six tentacles; and one alone moved after an interval of 2 hrs. 30 m. Nevertheless, two glands were extremely sensitive to very small doses of the nitrate of ammonia, namely to about 1/20 of a minim of a solution (one part to 5250 of water), containing only 1/115200 of a grain (.000562 mg.) of the salt.

Fragments of flies were placed on two leaves near their tips, which became incurved in 15 hrs. A fly was also placed in the middle of the leaf; in a few hours the tentacles on each side embraced it, and in 8 hrs. the whole leaf directly beneath the fly was a little bent transversely. By the next morning, after 23 hrs., the leaf was curled so completely over that the apex rested on the upper end of the footstalk. In no case did the sides of the leaves become inflected. A crushed fly was placed on the foliaceous footstalk, but produced no effect.

Drosera spathulata (sent to me by Dr. Hooker).--I made only a few observations on this Australian species, which has long, narrow leaves, gradually widening towards their tips. The glands of the extreme marginal tentacles are elongated and differ from the others, as in the case of Drosera rotundifolia. A fly was placed on a leaf, and in 18 hrs. it was embraced by the adjoining tentacles. Gum-water dropped on several leaves produced no effect. A fragment of a leaf was immersed in a few drops of a solution of one part of carbonate of ammonia to 146 of water; all the glands were instantly blackened; the process of aggregation could be seen travelling rapidly down the cells of the tentacles; and the granules of protoplasm soon united into spheres and variously shaped ma.s.ses, which displayed the usual move- [page 281]

ments. Half a minim of a solution of one part of nitrate of ammonia to 146 of water was next placed on the centre of a leaf; after 6 hrs. some marginal tentacles on both sides were inflected, and after 9 hrs. they met in the centre. The lateral edges of the leaf also became incurved, so that it formed a half-cylinder; but the apex of the leaf in none of my few trials was inflected. The above dose of the nitrate (viz. 1/320 of a grain, or .202 mg.) was too powerful, for in the course of 23 hrs.

the leaf died.

Drosera filiformis.--This North American species grows in such abundance in parts of New Jersey as almost to cover the ground. It catches, according to Mrs. Treat,* an extraordinary number of small and large insects, even great flies of the genus Asilus, moths, and b.u.t.terflies. The specimen which I examined, sent me by Dr. Hooker, had thread-like leaves, from 6 to 12 inches in length, with the upper surface convex and the lower flat and slightly channelled. The whole convex surface, down to the roots--for there is no distinct footstalk--is covered with short gland-bearing tentacles, those on the margins being the longest and reflexed. Bits of meat placed on the glands of some tentacles caused them to be slightly inflected in 20 m.; but the plant was not in a vigorous state. After 6 hrs. they moved through an angle of 90o, and in 24 hrs. reached the centre. The surrounding tentacles by this time began to curve inwards. Ultimately a large drop of extremely viscid, slightly acid secretion was poured over the meat from the united glands. Several other glands were touched with a little saliva, and the tentacles became incurved in under 1 hr., and re-expanded after 18 hrs. Particles of gla.s.s, cork, cinders, thread, and gold-leaf, were placed on numerous glands on two leaves; in about 1 hr. four tentacles became curved, and four others after an additional interval of 2 hrs. 30 m. I never once succeeded in causing any movement by repeatedly touching the glands with a needle; and Mrs.

Treat made similar trials for me with no success. Small flies were placed on several leaves near their tips, but the thread-like blade became only on one occasion very slightly bent, directly beneath the insect. Perhaps this indicates that the blades of vigorous plants would bend over captured insects, and Dr. Canby informs me that this is the case; but the movement cannot be strongly p.r.o.nounced, as it was not observed by Mrs. Treat.

Drosera binata (or dichotoma).--I am much indebted to Lady

* "American Naturalist," December 1873, page 705. [page 282]

Dorothy Nevill for a fine plant of this almost gigantic Australian species, which differs in some interesting points from those previously described. In this specimen the rush-like footstalks of the leaves were 20 inches in length. The blade bifurcates at its junction with the footstalk, and twice or thrice afterwards, curling about in an irregular manner. It is narrow, being only 3/20 of an inch in breadth.

One blade was 7 1/2 inches long, so that the entire leaf, including the footstalk, was above 27 inches in length. Both surfaces are slightly hollowed out. The upper surface is covered with tentacles arranged in alternate rows; those in the middle being short and crowded together, those towards the margins longer, even twice or thrice as long as the blade is broad. The glands of the exterior tentacles are of a much darker red than those of the central ones. The pedicels of all are green. The apex of the blade is attenuated, and bears very long tentacles. Mr. Copland informs me that the leaves of a plant which he kept for some years were generally covered with captured insects before they withered.

The leaves do not differ in essential points of structure or of function from those of the previously described species. Bits of meat or a little saliva placed on the glands of the exterior tentacles caused well-marked movement in 3 m., and particles of gla.s.s acted in 4 m. The tentacles with the latter particles re-expanded after 22 hrs. A piece of leaf immersed in a few drops of a solution of one part of carbonate of ammonia to 437 of water had all the glands blackened and all the tentacles inflected in 5 m. A bit of raw meat, placed on several glands in the medial furrow, was well clasped in 2 hrs. 10 m.

by the marginal tentacles on both sides. Bits of roast meat and small flies did not act quite so quickly; and alb.u.men and fibrin still less quickly. One of the bits of meat excited so much secretion (which is always acid) that it flowed some way down the medial furrow, causing the inflection of the tentacles on both sides as far as it extended.

Particles of gla.s.s placed on the glands in the medial furrow did not stimulate them sufficiently for any motor impulse to be sent to the outer tentacles. In no case was the blade of the leaf, even the attenuated apex, at all inflected.

On both the upper and lower surface of the blade there are numerous minute, almost sessile glands, consisting of four, eight, or twelve cells. On the lower surface they are pale purple, on the upper greenish. Nearly similar organs occur on the foot-stalks, but they are smaller and often in a shrivelled condition. The minute glands on the blade can absorb rapidly: thus, a piece of leaf was immersed in a solution of one part of carbonate [page 283] of ammonia to 218 of water (1 gr. to 2 oz.), and in 5 m. they were all so much darkened as to be almost black, with their contents aggregated. They do not, as far as I could observe, secrete spontaneously; but in between 2 and 3 hrs. after a leaf had been rubbed with a bit of raw meat moistened with saliva, they seemed to be secreting freely; and this conclusion was afterwards supported by other appearances. They are, therefore, h.o.m.ologous with the sessile glands hereafter to be described on the leaves of Dionaea and Drosophyllum. In this latter genus they are a.s.sociated, as in the present case, with glands which secrete spontaneously, that is, without being excited.

Drosera binata presents another and more remarkable peculiarity, namely, the presence of a few tentacles on the backs of the leaves, near their margins. These are perfect in structure; spiral vessels run up their pedicels; their glands are surrounded by drops of viscid secretion, and they have the power of absorbing. This latter fact was shown by the glands immediately becoming black, and the protoplasm aggregated, when a leaf was placed in a little solution of one part of carbonate of ammonia to 437 of water. These dorsal tentacles are short, not being nearly so long as the marginal ones on the upper surface; some of them are so short as almost to graduate into the minute sessile glands. Their presence, number, and size, vary on different leaves, and they are arranged rather irregularly. On the back of one leaf I counted as many as twenty-one along one side.

These dorsal tentacles differ in one important respect from those on the upper surface, namely, in not possessing any power of movement, in whatever manner they may be stimulated. Thus, portions of four leaves were placed at different times in solutions of carbonate of ammonia (one part to 437 or 218 of water), and all the tentacles on the upper surface soon became closely inflected; but the dorsal ones did not move, though the leaves were left in the solution for many hours, and though their glands from their blackened colour had obviously absorbed some of the salt. Rather young leaves should be selected for such trials, for the dorsal tentacles, as they grow old and begin to wither, often spontaneously incline towards the middle of the leaf. If these tentacles had possessed the power of movement, they would not have been thus rendered more serviceable to the plant; for they are not long enough to bend round the margin of the leaf so as to reach an insect caught on the upper surface, Nor would it have been of any use if these tentacles could have [page 284] moved towards the middle of the lower surface, for there are no viscid glands there by which insects can be caught. Although they have no power of movement, they are probably of some use by absorbing animal matter from any minute insect which may be caught by them, and by absorbing ammonia from the rain-water. But their varying presence and size, and their irregular position, indicate that they are not of much service, and that they are tending towards abortion. In a future chapter we shall see that Drosophyllum, with its elongated leaves, probably represents the condition of an early progenitor of the genus Drosera; and none of the tentacles of Drosophyllum, neither those on the upper nor lower surface of the leaves, are capable of movement when excited, though they capture numerous insects, which serve as nutriment. Therefore it seems that Drosera binata has retained remnants of certain ancestral characters--namely a few motionless tentacles on the backs of the leaves, and fairly well developed sessile glands--which have been lost by most or all of the other species of the genus.]

Concluding Remarks.--From what we have now seen, there can be little doubt that most or probably all the species of Drosera are adapted for catching insects by nearly the same means. Besides the two Australian species above described, it is said* that two other species from this country, namely Drosera pallida and Drosera sulphurea, "close their leaves upon insects with "great rapidity: and the same phenomenon is mani-"fested by an Indian species, D. lunata, and by several "of those of the Cape of Good Hope, especially by "D. trinervis." Another Australian species, Drosera heterophylla (made by Lindley into a distinct genus, Sondera) is remarkable from its peculiarly shaped leaves, but I know nothing of its power of catching insects, for I have seen only dried specimens. The leaves form minute flattened cups, with the footstalks attached not to one margin, but to the bottom. The

* "Gardener"s Chronicle," 1874, p. 209. [page 285]

inner surface and the edges of the cups are studded with tentacles, which include fibro-vascular bundles, rather different from those seen by me in any other species; for some of the vessels are barred and punctured, instead of being spiral. The glands secrete copiously, judging from the quant.i.ty of dried secretion adhering to them. [page 286]

CHAPTER XIII.

DIONAEA MUSCIPULA.

Structure of the leaves--Sensitiveness of the filaments--Rapid movement of the lobes caused by irritation of the filaments--Glands, their power of secretion--Slow movement caused by the absorption of animal matter--Evidence of absorption from the aggregated condition of the glands--Digestive power of the secretion--Action of chloroform, ether, and hydrocyanic acid- -The manner in which insects are captured--Use of the marginal spikes--Kinds of insects captured--The transmission of the motor impulse and mechanism of the movements-- Re-expansion of the lobes.

THIS plant, commonly called Venus" fly-trap, from the rapidity and force of its movements, is one of the most wonderful in the world.* It is a member of the small family of the Droseraceae, and is found only in the eastern part of North Carolina, growing in damp situations. The roots are small; those of a moderately fine plant which I examined consisted of two branches about 1 inch in length, springing from a bulbous enlargement. They probably serve, as in the case of Drosera, solely for the absorption of water; for a gardener, who has been very successful in the cultivation of this plant, grows it, like an epiphytic orchid, in well-drained damp moss without any soil. The form of the bilobed leaf, with its foliaceous footstalk, is shown in the accompanying drawing (fig. 12).

* Dr. Hooker, in his address to the British a.s.sociation at Belfast, 1874, has given so full an historical account of the observations which have been published on the habits of this plant, that it would be superfluous on my part to repeat them.

"Gardener"s Chronicle," 1874, p. 464. [page 287]

The two lobes stand at rather less than a right angle to each other.

Three minute pointed processes or filaments, placed triangularly, project from the upper surfaces of both; but I have seen two leaves with four filaments on each side, and another with only two. These filaments are remarkable from their extreme sensitiveness to a touch, as shown not by their own movement, but by that of the lobes. The margins of the leaf are prolonged into sharp rigid projections which I will call spikes, into each of which a bundle

FIG. 12. (Dionaea muscipula.) Leaf viewed laterally in its expanded state.

of spiral vessels enters. The spikes stand in such a position that, when the lobes close, they inter-lock like the teeth of a rat-trap. The midrib of the leaf, on the lower side, is strongly developed and prominent.

The upper surface of the leaf is thickly covered, excepting towards the margins, with minute glands of a reddish or purplish colour, the rest of the leaf being green. There are no glands on the spikes, or on the foliaceous footstalk, The glands are formed of from [page 288] twenty to thirty polygonal cells, filled with purple fluid. Their upper surface is convex. They stand on very short pedicels, into which spiral vessels do not enter, in which respect they differ from the tentacles of Drosera. They secrete, but only when excited by the absorption of certain matters; and they have the power of absorption. Minute projections, formed of eight divergent arms of a reddish-brown or orange colour, and appearing under the microscope like elegant little flowers, are scattered in considerable numbers over the foot-stalk, the backs of the leaves, and the spikes, with a few on the upper surface of the lobes. These octofid projections are no doubt h.o.m.ologous with the papillae on the leaves of Drosera rotundifolia. There are also a few very minute, simple, pointed hairs, about 7/12000 (.0148 mm.) of an inch in length on the backs of the leaves.

The sensitive filaments are formed of several rows of elongated cells, filled with purplish fluid. They are a little above the 1/20 of an inch in length; are thin and delicate, and taper to a point. I examined the bases of several, making sections of them, but no trace of the entrance of any vessel could be seen. The apex is sometimes bifid or even trifid, owing to a slight separation between the terminal pointed cells. Towards the base there is constriction, formed of broader cells, beneath which there is an articulation, supported on an enlarged base, consisting of differently shaped polygonal cells. As the filaments project at right angles to the surface of the leaf, they would have been liable to be broken whenever the lobes closed together, had it not been for the articulation which allows them to bend flat down.

These filaments, from their tips to their bases, are exquisitely sensitive to a momentary touch. It is scarcely [page 289] possible to touch them ever so lightly or quickly with any hard object without causing the lobes to close. A piece of very delicate human hair, 2 1/2 inches in length, held dangling over a filament, and swayed to and fro so as to touch it, did not excite any movement. But when a rather thick cotton thread of the same length was similarly swayed, the lobes closed. Pinches of fine wheaten flour, dropped from a height, produced no effect. The above-mentioned hair was then fixed into a handle, and cut off so that 1 inch projected; this length being sufficiently rigid to support itself in a nearly horizontal line. The extremity was then brought by a slow movement laterally into contact with the tip of a filament, and the leaf instantly closed. On another occasion two or three touches of the same kind were necessary before any movement ensued. When we consider how flexible a fine hair is, we may form some idea how slight must be the touch given by the extremity of a piece, 1 inch in length, moved slowly.

Although these filaments are so sensitive to a momentary and delicate touch, they are far less sensitive than the glands of Drosera to prolonged pressure. Several times I succeeded in placing on the tip of a filament, by the aid of a needle moved with extreme slowness, bits of rather thick human hair, and these did not excite movement, although they were more than ten times as long as those which caused the tentacles of Drosera to bend; and although in this latter case they were largely supported by the dense secretion. On the other hand, the glands of Drosera may be struck with a needle or any hard object, once, twice, or even thrice, with considerable force, and no movement ensues.

This singular difference in the nature of the sensitiveness of the filaments of Dionaea and of [page 290] the glands of Drosera evidently stands in relation to the habits of the two plants. If a minute insect alights with its delicate feet on the glands of Drosera, it is caught by the viscid secretion, and the slight, though prolonged pressure, gives notice of the presence of prey, which is secured by the slow bending of the tentacles. On the other hand, the sensitive filaments of Dionaea are not viscid, and the capture of insects can be a.s.sured only by their sensitiveness to a momentary touch, followed by the rapid closure of the lobes.

As just stated, the filaments are not glandular, and do not secrete.

Nor have they the power of absorption, as may be inferred from drops of a solution of carbonate of ammonia (one part to 146 of water), placed on two filaments, not producing any effect on the contents of their cells, nor causing the lobes to close, When, however, a small portion of a leaf with an attached filament was cut off and immersed in the same solution, the fluid within the basal cells became almost instantly aggregated into purplish or colourless, irregularly shaped ma.s.ses of matter. The process of aggregation gradually travelled up the filaments from cell to cell to their extremities, that is in a reverse course to what occurs in the tentacles of Drosera when their glands have been excited. Several other filaments were cut off close to their bases, and left for 1 hr. 30 m. in a weaker solution of one part of the carbonate to 218 of water, and this caused aggregation in all the cells, commencing as before at the bases of the filaments.

Long immersion of the filaments in distilled water likewise causes aggregation. Nor is it rare to find the contents of a few of the terminal cells in a spontaneously aggregated condition. The aggregated [page 291] ma.s.ses undergo incessant slow changes of form, uniting and again separating; and some of them apparently revolve round their own axes. A current of colourless granular protoplasm could also be seen travelling round the walls of the cells. This current ceases to be visible as soon as the contents are well aggregated; but it probably still continues, though no longer visible, owing to all the granules in the flowing layer having become united with the central ma.s.ses. In all these respects the filaments of Dionaea behave exactly like the tentacles of Drosera.

Notwithstanding this similarity there is one remarkable difference. The tentacles of Drosera, after their glands have been repeatedly touched, or a particle of any kind has been placed on them, become inflected and strongly aggregated. No such effect is produced by touching the filaments of Dionaea; I compared, after an hour or two, some which had been touched and some which had not, and others after twenty-five hours, and there was no difference in the contents of the cells. The leaves were kept open all the time by clips; so that the filaments were not pressed against the opposite lobe.

Drops of water, or a thin broken stream, falling from a height on the filaments, did not cause the blades to close; though these filaments were afterwards proved to be highly sensitive. No doubt, as in the case of Drosera, the plant is indifferent to the heaviest shower of rain.

Drops of a solution of a half an ounce of sugar to a fluid ounce of water were repeatedly allowed to fall from a height on the filaments, but produced no effect, unless they adhered to them. Again, I blew many times through a fine pointed tube with my utmost force against the filaments without any effect; such blowing being received [page 292]

with as much indifference as no doubt is a heavy gale of wind. We thus see that the sensitiveness of the filaments is of a specialised nature, being related to a momentary touch rather than to prolonged pressure; and the touch must not be from fluids, such as air or water, but from some solid object.

Although drops of water and of a moderately strong solution of sugar, falling on the filaments, does not excite them, yet the immersion of a leaf in pure water sometimes caused the lobes to close. One leaf was left immersed for 1 hr. 10 m., and three other leaves for some minutes, in water at temperatures varying between 59o and 65o (15o to 18o.3 Cent.) without any effect. One, however, of these four leaves, on being gently withdrawn from the water, closed rather quickly. The three other leaves were proved to be in good condition, as they closed when their filaments were touched. Nevertheless two fresh leaves on being dipped into water at 75o and 62 1/2o (23o.8 and 16o.9 Cent.) instantly closed.

These were then placed with their footstalks in water, and after 23 hrs. partially re-expanded; on touching their filaments one of them closed. This latter leaf after an additional 24 hrs. again re-expanded, and now, on the filaments of both leaves being touched, both closed. We thus see that a short immersion in water does not at all injure the leaves, but sometimes excites the lobes to close. The movement in the above cases was evidently not caused by the temperature of the water.

It has been shown that long immersion causes the purple fluid within the cells of the sensitive filaments to become aggregated; and the tentacles of Drosera are acted on in the same manner by long immersion, often being somewhat inflected. In both cases the result is probably due to a slight degree of exosmose. [page 293]

I am confirmed in this belief by the effects of immersing a leaf of Dionaea in a moderately strong solution of sugar; the leaf having been previously left for 1 hr. 10 m. in water without any effect; for now the lobes closed rather quickly, the tips of the marginal spikes crossing in 2 m. 30 s., and the leaf being completely shut in 3 m.

Three leaves were then immersed in a solution of half an ounce of sugar to a fluid ounce of water, and all three leaves closed quickly. As I was doubtful whether this was due to the cells on the upper surface of the lobes, or to the sensitive filaments, being acted on by exosmose, one leaf was first tried by pouring a little of the same solution in the furrow between the lobes over the midrib, which is the chief seat of movement. It was left there for some time, but no movement ensued.

The whole upper surface of leaf was then painted (except close round the bases of the sensitive filaments, which I could not do without risk of touching them) with the same solution, but no effect was produced.

So that the cells on the upper surface are not thus affected. But when, after many trials, I succeeded in getting a drop of the solution to cling to one of the filaments, the leaf quickly closed. Hence we may, I think, conclude that the solution causes fluid to pa.s.s out of the delicate cells of the filaments by exosmose; and that this sets up some molecular change in their contents, a.n.a.logous to that which must be produced by a touch.

The immersion of leaves in a solution of sugar affects them for a much longer time than does an immersion in water, or a touch on the filaments; for in these latter cases the lobes begin to re-expand in less than a day. On the other hand, of the three leaves which were immersed for a short time in the solution, and were then washed by means of a syringe inserted [page 294] between the lobes, one re-expanded after two days; a second after seven days; and the third after nine days. The leaf which closed, owing to a drop of the solution having adhered to one of the filaments, opened after two days.

I was surprised to find on two occasions that the heat from the rays of the sun, concentrated by a lens on the bases of several filaments, so that they were scorched and discoloured, did not cause any movement; though the leaves were active, as they closed, though rather slowly, when a filament on the opposite side was touched. On a third trial, a fresh leaf closed after a time, though very slowly; the rate not being increased by one of the filaments, which had not been injured, being touched. After a day these three leaves opened, and were fairly sensitive when the uninjured filaments were touched. The sudden immersion of a leaf into boiling water does not cause it to close.

Judging from the a.n.a.logy of Drosera, the heat in these several cases was too great and too suddenly applied. The surface of the blade is very slightly sensitive; It may be freely and roughly handled, without any movement being caused. A leaf was scratched rather hard with a needle, but did not close; but when the triangular s.p.a.ce between the three filaments on another leaf was similarly scratched, the lobes closed. They always closed when the blade or midrib was deeply p.r.i.c.ked or cut. Inorganic bodies, even of large size, such as bits of stone, gla.s.s, &c.--or organic bodies not containing soluble nitrogenous matter, such as bits of wood, cork, moss,--or bodies containing soluble nitrogenous matter, if perfectly dry, such as bits of meat, alb.u.men, gelatine, &c., may be long left (and many were tried) on the lobes, and no movement is excited. The result, however, is widely different, as we [page 295] shall presently see, if nitrogenous organic bodies which are at all damp, are left on the lobes; for these then close by a slow and gradual movement, very different from that caused by touching one of the sensitive filaments. The footstalk is not in the least sensitive; a pin may be driven through it, or it may be cut off, and no movement follows.

The upper surface of the lobes, as already stated, is thickly covered with small purplish, almost sessile glands. These have the power both of secretion and absorption; but unlike those of Drosera, they do not secrete until excited by the absorption of nitrogenous matter. No other excitement, as far as I have seen, produces this effect. Objects, such as bits of wood, cork, moss, paper, stone, or gla.s.s, may be left for a length of time on the surface of a leaf, and it remains quite dry. Nor does it make any difference if the lobes close over such objects. For instance, some little b.a.l.l.s of blotting paper were placed on a leaf, and a filament was touched; and when after 24 hrs. the lobes began to re-open, the b.a.l.l.s were removed by the aid of thin pincers, and were found perfectly dry. On the other hand, if a bit of damp meat or a crushed fly is placed on the surface of an expanded leaf, the glands after a time secrete freely. In one such case there was a little secretion directly beneath the meat in 4 hrs.; and after an additional 3 hrs. there was a considerable quant.i.ty both under and close round it.

In another case, after 3 hrs. 40 m., the bit of meat was quite wet. But none of the glands secreted, excepting those which actually touched the meat or the secretion containing dissolved animal matter.

If, however, the lobes are made to close over a bit of meat or an insect, the result is different, for the glands over the whole surface of the leaf now secrete copiously. [page 296] As in this case the glands on both sides are pressed against the meat or insect, the secretion from the first is twice as great as when a bit of meat is laid on the surface of one lobe; and as the two lobes come into almost close contact, the secretion, containing dissolved animal matter, spreads by capillary attraction, causing fresh glands on both sides to begin secreting in a continually widening circle. The secretion is almost colourless, slightly mucilaginous, and, judging by the manner in which it coloured litmus paper, more strongly acid than that of Drosera. It is so copious that on one occasion, when a leaf was cut open, on which a small cube of alb.u.men had been placed 45 hrs. before, drops rolled off the leaf. On another occasion, in which a leaf with an enclosed bit of roast meat spontaneously opened after eight days, there was so much secretion in the furrow over the midrib that it trickled down. A large crushed fly (Tipula) was placed on a leaf from which a small portion at the base of one lobe had previously been cut away, so that an opening was left; and through this, the secretion continued to run down the footstalk during nine days,--that is, for as long a time as it was observed. By forcing up one of the lobes, I was able to see some distance between them, and all the glands within sight were secreting freely.