Insectivorous Plants Part 16

Two plants bearing four or five leaves, and with their roots in a little cup of water, were exposed to the vapour of some bits of camphor (about as large as a filbert-nut), under a vessel holding ten fluid oz.

After 10 hrs. no inflection ensued; but the glands appeared to be secreting more copiously. The leaves were in a narcotised condition, for on bits of meat being placed on two of them, there was no inflection in 3 hrs. 15 m., and even after 13 hrs. 15 m. only a few of the outer tentacles were slightly inflected; but this degree of movement shows that the leaves had not been killed by an exposure during 10 hrs. to the vapour of camphor.

Oil of Caraway.--Water is said to dissolve about a thousandth part of its weight of this oil. A drop was added to an ounce of water and the bottle occasionally shaken during a day; but many minute globules remained undissolved. Five leaves were immersed in this mixture; in from 4 m. to 5 m. there was some inflection, which became moderately p.r.o.nounced in two or [page 212] three additional minutes. After 14 m.

all five leaves were well, and some of them closely, inflected. After 6 hrs. the glands were white, and much mucus had been secreted. The leaves were now flaccid, of a peculiar dull-red colour, and evidently dead. One of the leaves, after an immersion of 4 m., was brushed, like the leaves in the camphor, but this produced no effect. A plant with its roots in water was exposed under a 10-oz. vessel to the vapour of this oil, and in 1 hr. 20 m. one leaf showed a trace of inflection.

After 5 hrs. 20 m. the cover was taken off and the leaves examined; one had all its tentacles closely inflected, the second about half in the same state; and the third all sub-inflected. The plant was left in the open air for 42 hrs., but not a single tentacle expanded; all the glands appeared dead, except here and there one, which was still secreting. It is evident that this oil is highly exciting and poisonous to Drosera.

Oil of Cloves.--A mixture was made in the same manner as in the last case, and three leaves were immersed in it. After 30 m. there was only a trace of inflection which never increased. After 1 hr. 30 m. the glands were pale, and after 6 hrs. white. No doubt the leaves were much injured or killed.

Turpentine.--Small drops placed on the discs of some leaves killed them, as did likewise drops of creosote. A plant was left for 15 m.

under a 12-oz. vessel, with its inner surface wetted with twelve drops of turpentine; but no movement of the tentacles ensued. After 24 hrs.

the plant was dead.

Glycerine.--Half-minims were placed on the discs of three leaves: in 2 hrs. some of the outer tentacles were irregularly inflected; and in 19 hrs. the leaves were flaccid and apparently dead; the glands which had touched the glycerine were colourless. Minute drops (about 1/20 of a minim) were applied to the glands of several tentacles, and in a few minutes these moved and soon reached the centre. Similar drops of a mixture of four dropped drops to 1 oz. of water were likewise applied to several glands; but only a few of the tentacles moved, and these very slowly and slightly. Half-minims of this same mixture placed on the discs of some leaves caused, to my surprise, no inflection in the course of 48 hrs. Bits of meat were then given them, and next day they were well inflected; notwithstanding that some of the discal glands had been rendered almost colourless. Two leaves were immersed in the same mixture, but only for 4 hrs.; they were not inflected, and on being afterwards left for 2 hrs. 30 m. in a solution (1 gr. to 1 oz.) of carbonate of ammonia, their glands were blackened, their tentacles inflected, and the protoplasm within their cells aggregated. It appears [page 213] from these facts that a mixture of four drops of glycerine to an ounce of water is not poisonous, and excites very little inflection; but that pure glycerine is poisonous, and if applied in very minute quant.i.ties to the glands of the outer tentacles causes their inflection.

The Effects of Immersion in Water and in various Solutions on the subsequent Action of Phosphate and Carbonate of Ammonia.--We have seen in the third and seventh chapters that immersion in distilled water causes after a time some degree of aggregation of the protoplasm, and a moderate amount of inflection, especially in the case of plants which have been kept at a rather high temperature. Water does not excite a copious secretion of mucus. We have here to consider the effects of immersion in various fluids on the subsequent action of salts of ammonia and other stimulants. Four leaves which had been left for 24 hrs. in water were given bits of meat, but did not clasp them. Ten leaves, after a similar immersion, were left for 24 hrs. in a powerful solution (1 gr. to 20 oz.) of phosphate of ammonia, and only one showed even a trace of inflection. Three of these leaves, on being left for an additional day in the solution, still remained quite unaffected. When, however, some of these leaves, which had been first immersed in water for 24 hrs., and then in the phosphate for 24 hrs. were placed in a solution of carbonate of ammonia (one part to 218 of water), the protoplasm in the cells of the tentacles became in a few hours strongly aggregated, showing that this salt had been absorbed and taken effect.

A short immersion in water for 20 m. did not r.e.t.a.r.d the subsequent action of the phosphate, or of splinters of gla.s.s placed on the glands; but in two instances an immersion for 50 m. prevented any effect from a solution of camphor. Several leaves which had been left for 20 m. in a solution of one part of white sugar to 218 of water were placed in the phosphate solution, the action of which was delayed; whereas a mixed solution of sugar and the phosphate did not in the least interfere with the effects of the latter. Three leaves, after being immersed for 20 m. in the sugar solution, were placed in a solution of carbonate of ammonia (one part to 218 of water); in 2 m. or 3 m. the glands were blackened, and after 7 m. the tentacles were considerably inflected, so that the solution of sugar, though it delayed the action of the phosphate, did not delay that of the carbonate. Immersion in a similar solution of gum arabic for 20 m. had no r.e.t.a.r.ding action on the phosphate. Three leaves were left for 20 m. in a mixture of one part of alcohol to seven parts of water, [page 214] and then placed in the phosphate solution: in 2 hrs. 15 m. there was a trace of inflection in one leaf, and in 5 hrs. 30 m. a second was slightly affected; the inflection subsequently increased, though slowly. Hence diluted alcohol, which, as we shall see, is hardly at all poisonous, plainly r.e.t.a.r.ds the subsequent action of the phosphate.

It was shown in the last chapter that leaves which did not become inflected by nearly a day's immersion in solutions of various salts and acids behaved very differently from one another when subsequently placed in the phosphate solution. I here give a table summing up the results.

Column 1 : Name of the Salts and Acids in Solution. Column 2 : Period of Immersion of the Leaves in Solutions of one part to 437 of water.

Column 3 : Effects produced on the Leaves by their subsequent Immersion for stated periods in a Solution of one part of phosphate of ammonia to 8750 of water, or 1 gr. to 20 oz.

Rubidium chloride. : 22 hrs. : After 30 m. strong inflection of the tentacles.

Pota.s.sium carbonate : 20 m. : Scarcely any inflection until 5 hrs. had elapsed.

Calcium acetate. : 24 hrs. : After 24 hrs. very slight inflection.

Calcium nitrate. : 24 hrs. : Do. do.

Magnesium acetate. : 22 hrs. : Some slight inflection, which became well p.r.o.nounced in 24 hrs.

Magnesium nitrate. : 22 hrs. : After 4 hrs. 30 m. a fair amount of inflection, which never increased.

Magnesium chloride : 22 hrs. : After a few minutes great inflection; after 4 hrs. all four leaves with almost every tentacle closely inflected.

Barium acetate. : 22 hrs. : After 24 hrs. two leaves out of four slightly inflected.

Barium nitrate. : 22 hrs. : After 30 m. one leaf greatly, and two others moderately, inflected; they remained thus for 24 hrs.

Strontium acetate. : 22 hrs. : After 25 m. two leaves greatly inflected; after 8 hrs. a third leaf moderately, and the fourth very slightly, inflected. All four thus remained for 24 hrs.

Strontium nitrate. : 22 hrs. : After 8 hrs. three leaves out of five moderately inflected; after 24 hrs. all five in this state; but not one closely inflected.

Aluminium chloride : 24 hrs. : Three leaves which had either been slightly or not at all affected by the chloride became after 7 hrs. 30 m. rather closely inflected. [page 215]

Column 1 : Name of the Salts and Acids in Solution. Column 2 : Period of Immersion of the Leaves in Solutions of one part to 437 of water.

Column 3 : Effects produced on the Leaves by their subsequent Immersion for stated periods in a Solution of one part of phosphate of ammonia to 8750 of water, or 1 gr. to 20 oz.

Aluminium nitrate. : 24 hrs. : After 25 hrs. slight and doubtful effect.

Lead chloride. : 23 hrs. : After 24 hrs. two leaves somewhat inflected, the third very little; and thus remained.

Manganese chloride : 22 hrs. : After 48 hrs. not the least inflection.

Lactic acid. : 48 hrs. : After 24 hrs. a trace of inflection in a few tentacles, the glands of which had not been killed by the acid.

Tannic acid. : 24 hrs. : After 24 hrs. no inflection.

Tartaric acid. : 24 hrs. : Do. do.

Citric acid. : 24 hrs. : After 50 m. tentacles decidedly inflected, and after 5 hrs. strongly inflected; so remained for the next 24 hrs.

Formic acid. : 22 hrs. : Not observed until 24 hrs. had elapsed; tentacles considerably inflected, and protoplasm aggregated.

In a large majority of these twenty cases, a varying degree of inflection was slowly caused by the phosphate. In four cases, however, the inflection was rapid, occurring in less than half an hour or at most in 50 m. In three cases the phosphate did not produce the least effect. Now what are we to infer from these facts? We know from ten trials that immersion in distilled water for 24 hrs. prevents the subsequent action of the phosphate solution. It would, therefore, appear as if the solutions of chloride of manganese, tannic and tartaric acids, which are not poisonous, acted exactly like water, for the phosphate produced no effect on the leaves which had been previously immersed in these three solutions. The majority of the other solutions behaved to a certain extent like water, for the phosphate produced, after a considerable interval of time, only a slight effect.

On the other hand, the leaves which had been immersed in the solutions of the chloride of rubidium and magnesium, of acetate of strontium, nitrate of barium, and citric acid, were quickly acted on by the phosphate. Now was water absorbed from these five weak solutions, and yet, owing to the presence of the salts, did not prevent the subsequent action of the phosphate? Or [page 216] may we not suppose* that the interstices of the walls of the glands were blocked up with the molecules of these five substances, so that they were rendered impermeable to water; for had water entered, we know from the ten trials that the phosphate would not afterwards have produced any effect? It further appears that the molecules of the carbonate of ammonia can quickly pa.s.s into glands which, from having been immersed for 20 m. in a weak solution of sugar, either absorb the phosphate very slowly or are acted on by it very slowly. On the other hand, glands, however they may have been treated, seem easily to permit the subsequent entrance of the molecules of carbonate of ammonia. Thus leaves which had been immersed in a solution (of one part to 437 of water) of nitrate of pota.s.sium for 48 hrs.--of sulphate of pota.s.sium for 24 hrs.--and of the chloride of pota.s.sium for 25 hrs.--on being placed in a solution of one part of carbonate of ammonia to 218 of water, had their glands immediately blackened, and after 1 hr. their tentacles somewhat inflected, and the protoplasm aggregated. But it would be an endless task to endeavour to ascertain the wonderfully diversified effects of various solutions on Drosera.

Alcohol (one part to seven of water).--It has already been shown that half-minims of this strength placed on the discs of leaves do not cause any inflection; and that when two days afterwards the leaves were given bits of meat, they became strongly inflected. Four leaves were immersed in this mixture, and two of them after 30 m. were brushed with a camel-hair brush, like the leaves in the solution of camphor, but this produced no effect.

* See Dr. M. Traube's curious experiments on the production of artificial cells, and on their permeability to various salts, described in his papers: "Experimente zur Theorie der Zellenbildung und Endosmose," Breslau, 1866; and "Experimente zur physicalischen Erklrung der Bildung der Zellhaut, ihres Wachsthums durch Intussusception,"

Breslau, 1874. These researches perhaps explain my results. Dr. Traube commonly employed as a membrane the precipitate formed when tannic acid comes into contact with a solution of gelatine. By allowing a precipitation of sulphate of barium to take place at the same time, the membrane becomes "infiltrated" with this salt; and in consequence of the intercalation of molecules of sulphate of barium among those of the gelatine precipitate, the molecular interstices in the membrane are made smaller. In this altered condition, the membrane no longer allows the pa.s.sage through it of either sulphate of ammonia or nitrate of barium, though it retains its permeability for water and chloride of ammonia. [page 217]

Nor did these four leaves, on being left for 24 hrs. in the diluted alcohol, undergo any inflection. They were then removed; one being placed in an infusion of raw meat, and bits of meat on the discs of the other three, with their stalks in water. Next day one seemed a little injured, whilst two others showed merely a trace of inflection. We must, however, bear in mind that immersion for 24 hrs. in water prevents leaves from clasping meat. Hence alcohol of the above strength is not poisonous, nor does it stimulate the leaves like camphor does.

The vapour of alcohol acts differently. A plant having three good leaves was left for 25 m. under a receiver holding 19 oz. with sixty minims of alcohol in a watch-gla.s.s. No movement ensued, but some few of the glands were blackened and shrivelled, whilst many became quite pale. These were scattered over all the leaves in the most irregular manner, reminding me of the manner in which the glands were affected by the vapour of carbonate of ammonia. Immediately on the removal of the receiver particles of raw meat were placed on many of the glands, those which retained their proper colour being chiefly selected. But not a single tentacle was inflected during the next 4 hrs. After the first 2 hrs. the glands on all the tentacles began to dry; and next morning, after 22 hrs., all three leaves appeared almost dead, with their glands dry; the tentacles on one leaf alone being partially inflected.

A second plant was left for only 5 m. with some alcohol in a watch-gla.s.s, under a 12-oz. receiver, and particles of meat were then placed on the glands of several tentacles. After 10 m. some of them began to curve inwards, and after 55 m. nearly all were considerably inflected; but a few did not move. Some anaesthetic effect is here probable, but by no means certain. A third plant was also left for 5 m.

under the same small vessel, with its whole inner surface wetted with about a dozen drops of alcohol. Particles of meat were now placed on the glands of several tentacles, some of which first began to move in 25 m.; after 40 m. most of them were somewhat inflected, and after 1 hr. 10 m. almost all were considerably inflected. From their slow rate of movement there can be no doubt that the glands of these tentacles had been rendered insensible for a time by exposure during 5 m. to the vapour of alcohol.

Vapour of Chloroform.--The action of this vapour on Drosera is very variable, depending, I suppose, on the const.i.tution or age of the plant, or on some unknown condition. It sometimes causes the tentacles to move with extraordinary rapidity, and sometimes produces no such effect. The glands are sometimes [page 218] rendered for a time insensible to the action of raw meat, but sometimes are not thus affected, or in a very slight degree. A plant recovers from a small dose, but is easily killed by a larger one.

A plant was left for 30 m. under a bell-gla.s.s holding 19 fluid oz.

(539.6 ml.) with eight drops of chloroform, and before the cover was removed, most of the tentacles became much inflected, though they did not reach the centre. After the cover was removed, bits of meat were placed on the glands of several of the somewhat incurved tentacles; these glands were found much blackened after 6 hrs. 30 m., but no further movement ensued. After 24 hrs. the leaves appeared almost dead.

A smaller bell-gla.s.s, holding 12 fluid oz. (340.8 ml.), was now employed, and a plant was left for 90 s. under it, with only two drops of chloroform. Immediately on the removal of the gla.s.s all the tentacles curved inwards so as to stand perpendicularly up; and some of them could actually be seen moving with extraordinary quickness by little starts, and therefore in an unnatural manner; but they never reached the centre. After 22 hrs. they fully re-expanded, and on meat being placed on their glands, or when roughly touched by a needle, they promptly became inflected; so that these leaves had not been in the least injured.

Another plant was placed under the same small bell-gla.s.s with three drops of chloroform, and before two minutes had elapsed, the tentacles began to curl inwards with rapid little jerks. The gla.s.s was then removed, and in the course of two or three additional minutes almost every tentacle reached the centre. On several other occasions the vapour did not excite any movement of this kind.

There seems also to be great variability in the degree and manner in which chloroform renders the glands insensible to the subsequent action of meat. In the plant last referred to, which had been exposed for 2 m.

to three drops of chloroform, some few tentacles curved up only to a perpendicular position, and particles of meat were placed on their glands; this caused them in 5 m. to begin moving, but they moved so slowly that they did not reach the centre until 1 hr. 30 m. had elapsed. Another plant was similarly exposed, that is, for 2 m. to three drops of chloroform, and on particles of meat being placed on the glands of several tentacles, which had curved up into a perpendicular position, one of these began to bend in 8 m., but afterwards moved very slowly; whilst none of the other tentacles [page 219] moved for the next 40 m. Nevertheless, in 1 hr. 45 m. from the time when the bits of meat had been given, all the tentacles reached the centre. In this case some slight anaesthetic effect apparently had been produced. On the following day the plant had perfectly recovered.

Another plant bearing two leaves was exposed for 2 m. under the 19-oz.

vessel to two drops of chloroform; it was then taken out and examined; again exposed for 2 m. to two drops; taken out, and re-exposed for 3 m.

to three drops; so that altogether it was exposed alternately to the air and during 7 m. to the vapour of seven drops of chloroform. Bits of meat were now placed on thirteen glands on the two leaves. On one of these leaves, a single tentacle first began moving in 40 m., and two others in 54 m. On the second leaf some tentacles first moved in 1 hr.

11 m. After 2 hrs. many tentacles on both leaves were inflected; but none had reached the centre within this time. In this case there could not be the least doubt that the chloroform had exerted an anaesthetic influence on the leaves.

On the other hand, another plant was exposed under the same vessel for a much longer time, viz. 20 m., to twice as much chloroform. Bits of meat were then placed on the glands of many tentacles, and all of them, with a single exception, reached the centre in from 13 m. to 14 m. In this case, little or no anaesthetic effect had been produced; and how to reconcile these discordant results, I know not.