Insectivorous Plants · Charles Darwin
ACIDS, MUCH DILUTED, WHICH CAUSE INFLECTION.
1. Nitric, strong inflection; poisonous. 2. Hydrochloric, moderate and slow inflection; not poisonous. 3. Hydriodic, strong inflection; poisonous. 4. Iodic, strong inflection; poisonous. 5. Sulphuric, strong inflection; somewhat poisonous. 6. Phosphoric, strong inflection; poisonous. 7. Boracic; moderate and rather slow inflection; not poisonous. 8. Formic, very slight inflection; not poisonous. 9. Acetic, strong and rapid inflection; poisonous. 10. Propionic, strong but not very rapid inflection; poisonous. 11. Oleic, quick inflection; very poisonous. 12. Carbolic, very slow inflection; poisonous. 13. Lactic, slow and moderate inflection; poisonous. 14. Oxalic, moderately quick inflection; very poisonous. 15. Malic, very slow but considerable inflection; not poisonous. 16. Benzoic, rapid inflection; very poisonous. 17. Succinic, moderately quick inflection: moderately poisonous. 18. Hippuric, rather slow inflection; poisonous. 19. Hydrocyanic, rather rapid inflection; very poisonous. ACIDS, DILUTED TO THE SAME DEGREE, WHICH DO NOT CAUSE INFLECTION. 1. Gallic; not poisonous. 2. Tannic; not poisonous. 3. Tartaric; not poisonous. 4. Citric; not poisonous. 5. Uric; (?) not poisonous.
Nitric Acid.—Four leaves were placed, each in thirty minims of one part by weight of the acid to 437 of water, so that each received 1/16 of a grain, or 4.048 mg. This strength was chosen for this and most of the following experiments, as it is the same [page 190] as that of most of the foregoing saline solutions. In 2 hrs. 30 m. some of the leaves were considerably, and in 6 hrs. 30 m. all were immensely, inflected, as were their blades. The surrounding fluid was slightly coloured pink, which always shows that the leaves have been injured. They were then left in water for three days; but they remained inflected and were evidently killed. Most of the glands had become colourless. Two leaves were then immersed, each in thirty minims of one part to 1000 of water; in a few hours there was some inflection; and after 24 hrs. both leaves had almost all their tentacles and blades inflected; they were left in water for three days, and one partially re-expanded and recovered. Two leaves were next immersed, each in thirty minims of one part to 2000 of water; this produced very little effect, except that most of the tentacles close to the summit of the petiole were inflected, as if the acid had been absorbed by the cut-off end.
Hydrochloric Acid.—One part to 437 of water; four leaves were immersed as before, each in thirty minims. After 6 hrs. only one leaf was considerably inflected. After 8 hrs. 15 m. one had its tentacles and blade well inflected; the other three were moderately inflected, and the blade of one slightly. The surrounding fluid was not coloured at all pink. After 25 hrs. three of these four leaves began to re-expand, but their glands were of a pink instead of a red colour; after two more days they fully re-expanded; but the fourth leaf remained inflected, and seemed much injured or killed, with its glands white. Four leaves were then treated, each with thirty minims of one part to 875 of water; after 21 hrs. they were moderately inflected; and on being transferred to water, fully re-expanded in two days, and seemed quite healthy. Hydriodic Acid.—One to 437 of water; three leaves were immersed as before, each in thirty minims. After 45 m. the glands were discoloured, and the surrounding fluid became pinkish, but there was no inflection. After 5 hrs. all the tentacles were closely inflected; and an immense amount of mucus was secreted, so that the fluid could be drawn out into long ropes. The leaves were then placed in water, but never re-expanded, and were evidently killed. Four leaves were next immersed in one part to 875 of water; the action was now slower, but after 22 hrs. all four leaves were closely inflected, and were affected in other respects as above described. These leaves did not re-expand, though left for four days in water. This acid acts far more powerfully than hydrochloric, and is poisonous.
Iodic Acid.—One to 437 of water; three leaves were immersed, [page 191] each in thirty minims; after 3 hrs. strong inflection; after 4 hrs. glands dark brown; after 8 hrs. 30 m. close inflection, and the leaves had become flaccid; surrounding fluid not coloured pink. These leaves were then placed in water, and next day were evidently dead. Sulphuric Acid.—One to 437 of water; four leaves were immersed, each in thirty minims; after 4 hrs. great inflection; after 6 hrs. surrounding fluid just tinged pink; they were then placed in water, and after 46 hrs. two of them were still closely inflected, two beginning to re-expand; many of the glands colourless. This acid is not so poisonous as hydriodic or iodic acids. Phosphoric Acid.—One to 437 of water; three leaves were immersed together in ninety minims; after 5 hrs. 30 m. some inflection, and some glands colourless; after 8 hrs. all the tentacles closely inflected, and many glands colourless; surrounding fluid pink. Left in water for two days and a half, remained in the same state and appeared dead. Boracic Acid.—One to 437 of water; four leaves were immersed together in 120 minims; after 6 hrs. very slight inflection; after 8 hrs. 15 m. two were considerably inflected, the other two slightly. After 24 hrs. one leaf was rather closely inflected, the second less closely, the third and fourth moderately. The leaves were washed and put into water; after 24 hrs. they were almost fully re-expanded and looked healthy. This acid agrees closely with hydrochloric acid of the same strength in its power of causing inflection, and in not being poisonous.
Formic Acid.—Four leaves were immersed together in 120 minims of one part to 437 of water; after 40 m. slight, and after 6 hrs. 30 m. very moderate inflection; after 22 hrs. only a little more inflection than often occurs in water. Two of the leaves were then washed and placed in a solution (1 gr. to 20 oz.) of phosphate of ammonia; after 24 hrs. they were considerably inflected, with the contents of their cells aggregated, showing that the phosphate had acted, though not to the full and ordinary degree. Acetic Acid.—Four leaves were immersed together in 120 minims of one part to 437 of water. In 1 hr. 20 m. the tentacles of all four and the blades of two were greatly inflected. After 8 hrs. the leaves had become flaccid, but still remained closely inflected, the surrounding fluid being coloured pink. They were then washed and placed in water; next morning they were still inflected and of a very dark red colour, but with their glands colourless. After another day they were dingy-coloured, and [page 192] evidently dead. This acid is far more powerful than formic, and is highly poisonous. Half-minim drops of a stronger mixture (viz. one part by measure to 320 of water) were placed on the discs of five leaves; none of the exterior tentacles, only those on the borders of the disc which actually absorbed the acid, became inflected. Probably the dose was too strong and paralysed the leaves, for drops of a weaker mixture caused much inflection; nevertheless the leaves all died after two days.
Propionic Acid.—Three leaves were immersed in ninety minims of a mixture of one part to 437 of water; in 1 hr. 50 m. there was no inflection; but after 3 hrs. 40 m. one leaf was greatly inflected, and the other two slightly. The inflection continued to increase, so that in 8 hrs. all three leaves were closely inflected. Next morning, after 20 hrs., most of the glands were very pale, but some few were almost black. No mucus had been secreted, and the surrounding fluid was only just perceptibly tinted of a pale pink. After 46 hrs. the leaves became slightly flaccid and were evidently killed, as was afterwards proved to be the case by keeping them in water. The protoplasm in the closely inflected tentacles was not in the least aggregated, but towards their bases it was collected in little brownish masses at the bottoms of the cells. This protoplasm was dead, for on leaving the leaf in a solution of carbonate of ammonia, no aggregation ensued. Propionic acid is highly poisonous to Drosera, like its ally acetic acid, but induces inflection at a much slower rate.
Oleic Acid (given me by Prof. Frankland).—Three leaves were immersed in this acid; some inflection was almost immediately caused, which increased slightly, but then ceased, and the leaves seemed killed. Next morning they were rather shrivelled, and many of the glands had fallen off the tentacles. Drops of this acid were placed on the discs of four leaves; in 40 m. all the tentacles were greatly inflected, excepting the extreme marginal ones; and many of these after 3 hrs. became inflected. I was led to try this acid from supposing that it was present (which does not seem to be the case)* in olive oil, the action of which is anomalous. Thus drops of this oil placed on the disc do not cause the outer tentacles to be inflected; yet when minute drops were added to the secretion surrounding the glands of the outer tentacles, these were occasionally, but by no means always, inflected. Two leaves were also immersed in this oil, and there * See articles on Glycerine and Oleic Acid in Watts’ ‘Dict. of Chemistry.’ [page 193] was no inflection for about 12 hrs.; but after 23 hrs. almost all the tentacles were inflected. Three leaves were likewise immersed in unboiled linseed oil, and soon became somewhat, and in 3 hrs. greatly, inflected. After 1 hr. the secretion round the glands was coloured pink. I infer from this latter fact that the power of linseed oil to cause inflection cannot be attributed to the albumin which it is said to contain.
Carbolic Acid.—Two leaves were immersed in sixty minims of a solution of 1 gr. to 437 of water; in 7 hrs. one was slightly, and in 24 hrs. both were closely, inflected, with a surprising amount of mucus secreted. These leaves were washed and left for two days in water; they remained inflected; most of their glands became pale, and they seemed dead. This acid is poisonous, but does not act nearly so rapidly or powerfully as might have been expected from its known destructive power on the lowest organisms. Half-minims of the same solution were placed on the discs of three leaves; after 24 hrs. no inflection of the outer tentacles ensued, and when bits of meat were given them, they became fairly well inflected. Again half-minims of a stronger solution, of one part to 218 of water, were placed on the discs of three leaves; no inflection of the outer tentacles ensued; bits of meat were then given as before; one leaf alone became well inflected, the discal glands of the other two appearing much injured and dry. We thus see that the glands of the discs, after absorbing this acid, rarely transmit any motor impulse to the outer tentacles; though these, when their own glands absorb the acid, are strongly acted on.
Lactic Acid.—Three leaves were immersed in ninety minims of one part to 437 of water. After 48 m. there was no inflection, but the surrounding fluid was coloured pink; after 8 hrs. 30 m. one leaf alone was a little inflected, and almost all the glands on all three leaves were of a very pale colour. The leaves were then washed and placed in a solution (1 gr. to 20 oz.) of phosphate of ammonia; after about 16 hrs. there was only a trace of inflection. They were left in the phosphate for 48 hrs., and remained in the same state, with almost all their glands discoloured. The protoplasm within the cells was not aggregated, except in a very few tentacles, the glands of which were not much discoloured. I believe, therefore, that almost all the glands and tentacles had been killed by the acid so suddenly that hardly any inflection was caused. Four leaves were next immersed in 120 minims of a weaker solution, of one part to 875 of water; after 2 hrs. 30 m. the surrounding fluid was quite pink; the glands were pale, but [page 194] there was no inflection; after 7 hrs. 30 m. two of the leaves showed some inflection, and the glands were almost white; after 21 hrs. two of the leaves were considerably inflected, and a third slightly; most of the glands were white, the others dark red. After 45 hrs. one leaf had almost every tentacle inflected; a second a large number; the third and fourth very few; almost all the glands were white, excepting those on the discs of two of the leaves, and many of these were very dark red. The leaves appeared dead. Hence lactic acid acts in a very peculiar manner, causing inflection at an extraordinarily slow rate, and being highly poisonous. Immersion in even weaker solutions, viz. of one part to 1312 and 1750 of water, apparently killed the leaves (the tentacles after a time being bowed backwards), and rendered the glands white, but caused no inflection.
Gallic, Tannic, Tartaric, and Citric Acids.—One part to 437 of water. Three or four leaves were immersed, each in thirty minims of these four solutions, so that each leaf received 1/16 of a grain, or 4.048 mg. No inflection was caused in 24 hrs., and the leaves did not appear at all injured. Those which had been in the tannic and tartaric acids were placed in a solution (1 gr. to 20 oz.) of phosphate of ammonia, but no inflection ensued in 24 hrs. On the other hand, the four leaves which had been in the citric acid, when treated with the phosphate, became decidedly inflected in 50 m. and strongly inflected after 5 hrs., and so remained for the next 24 hrs. Malic Acid.—Three leaves were immersed in ninety minims of a solution of one part to 437 of water; no inflection was caused in 8 hrs. 20 m., but after 24 hrs. two of them were considerably, and the third slightly, inflected—more so than could be accounted for by the action of water. No great amount of mucus was secreted. They were then placed in water, and after two days partially re-expanded. Hence this acid is not poisonous. Oxalic Acid.—Three leaves were immersed in ninety minims of a solution of 1 gr. to 437 of water; after 2 hrs. 10 m. there was much inflection; glands pale; the surrounding fluid of a dark pink colour; after 8 hrs. excessive inflection. The leaves were then placed in water; after about 16 hrs. the tentacles were of a very dark red colour, like those of the leaves in acetic acid. After 24 additional hours, the three leaves were dead and their glands colourless.
Benzoic Acid.—Five leaves were immersed, each in thirty minims of a solution of 1 gr. to 437 of water. This solution was so weak that it only just tasted acid, yet, as we shall see, was highly poisonous to Drosera. After 52 m. the submarginal [page 195] tentacles were somewhat inflected, and all the glands very pale-coloured; the surrounding fluid was coloured pink. On one occasion the fluid became pink in the course of only 12 m., and the glands as white as if the leaf had been dipped in boiling water. After 4 hrs. much inflection; but none of the tentacles were closely inflected, owing, as I believe, to their having been paralysed before they had time to complete their movement. An extraordinary quantity of mucus was secreted. Some of the leaves were left in the solution; others, after an immersion of 6 hrs. 30 m., were placed in water. Next morning both lots were quite dead; the leaves in the solution being flaccid, those in the water (now coloured yellow) of a pale brown tint, and their glands white. Succinic Acid.—Three leaves were immersed in ninety minims of a solution of 1 gr. to 437 of water; after 4 hrs. 15 m. considerable and after 23 hrs. great inflection; many of the glands pale; fluid coloured pink. The leaves were then washed and placed in water; after two days there was some re-expansion, but many of the glands were still white. This acid is not nearly so poisonous as oxalic or benzoic.
Uric Acid.—Three leaves were immersed in 180 minims of a solution of 1 gr. to 875 of warm water, but all the acid was not dissolved; so that each received nearly 1/16 of a grain. After 25 m. there was some slight inflection, but this never increased; after 9 hrs. the glands were not discoloured, nor was the solution coloured pink; nevertheless much mucus was secreted. The leaves were then placed in water, and by next morning fully re-expanded. I doubt whether this acid really causes inflection, for the slight movement which at first occurred may have been due to the presence of a trace of albuminous matter. But it produces some effect, as shown by the secretion of so much mucus. Hippuric Acid.—Four leaves were immersed in 120 minims of a solution of 1 gr. to 437 of water. After 2 hrs. the fluid was coloured pink; glands pale, but no inflection. After 6 hrs. some inflection; after 9 hrs. all four leaves greatly inflected; much mucus secreted; all the glands very pale. The leaves were then left in water for two days; they remained closely inflected, with their glands colourless, and I do not doubt were killed.
Hydrocyanic Acid.—Four leaves were immersed, each in thirty minims of one part to 437 of water; in 2 hrs. 45 m. all the tentacles were considerably inflected, with many of the glands pale; after 3 hrs. 45 m. all strongly inflected, and the surrounding fluid coloured pink; after 6 hrs. all closely inflected. After [page 196] an immersion of 8 hrs. 20 m. the leaves were washed and placed in water; next morning, after about 16 hrs., they were still inflected and discoloured; on the succeeding day they were evidently dead. Two leaves were immersed in a stronger mixture, of one part to fifty of water; in 1 hr. 15 m. the glands became as white as porcelain, as if they had been dipped in boiling water; very few of the tentacles were inflected; but after 4 hrs. almost all were inflected. These leaves were then placed in water, and next morning were evidently dead. Half-minim drops of the same strength (viz. one part to fifty of water) were next placed on the discs of five leaves; after 21 hrs. all the outer tentacles were inflected, and the leaves appeared much injured. I likewise touched the secretion round a large number of glands with minute drops (about 1/20 of a minim, or .00296 ml.) of Scheele’s mixture (6 per cent.); the glands first became bright red, and after 3 hrs. 15 m. about two-thirds of the tentacles bearing these glands were inflected, and remained so for the two succeeding days, when they appeared dead.]
Similar drops of a solution of one part to 218 of water (2 grs. to 1 oz.) quickly blackened the glands; some few tentacles thus treated moved, whilst others did not. The latter, however, on being subsequently moistened with saliva or given bits of meat, became incurved, though with extreme slowness; and this shows that they had been injured. Stronger solutions (but the strength was not ascertained) sometimes arrested all power of movement very quickly; thus bits of meat were placed on the glands of several exterior tentacles, and as soon as they began to move, minute drops of the strong solution were added. They continued for a short time to go on bending, and then suddenly stood still; other tentacles on the same leaves, with meat on their glands, but not wetted with the strychnine, continued to bend and soon reached the centre of the leaf.
Citrate of Strychnine.—Half-minims of a solution of one part to 437 of water were placed on the discs of six leaves; after 24 hrs. the outer tentacles showed only a trace of inflection. Bits of meat were then placed on three of these leaves, but in 24 hrs. only slight and irregular inflection occurred, proving that the leaves had been greatly injured. Two of the leaves to which meat had not been given had their discal glands dry and much injured. Minute drops of a strong solution of one part to 109 of water (4 grs. to 1 oz.) were added to the secretion round several glands, but did not produce nearly so plain an effect as the drops of a much weaker solution of the acetate. Particles of the dry citrate were placed on six glands; two of these moved some way towards the centre, and then stood still, being no doubt killed; three others curved much farther inwards, and were then fixed; one alone reached the centre. Five leaves were immersed, each in thirty minims of a solution of one part to 437 of water; so that each received 1/16 of a grain; after about 1 hr. some of the outer tentacles became inflected, and the glands were oddly mottled with black and white.
These glands, in from 4 hrs. to 5 hrs., became whitish and opaque, and the protoplasm in the cells of the tentacles was well aggregated. By this time two of the leaves were greatly inflected, but the three others not much more inflected than they were before. Nevertheless two fresh leaves, after an immersion respectively for 2 hrs. and 4 hrs. in the solution, were not killed; for on being left for 1 hr. 30 m. in a solution of one part of carbonate of ammonia to 218 of water, their tentacles became more inflected, and there was much aggregation. The glands [page 201] of two other leaves, after an immersion for 2 hrs. in a stronger solution, of one part of the citrate to 218 of water, became of an opaque, pale pink colour, which before long disappeared, leaving them white. One of these two leaves had its blade and tentacles greatly inflected; the other hardly at all; but the protoplasm in the cells of both was aggregated down to the bases of the tentacles, with the spherical masses in the cells close beneath the glands blackened. After 24 hrs. one of these leaves was colourless, and evidently dead.
Nicotine.—The secretion round several glands was touched with a minute drop of the pure fluid, and the glands were instantly blackened; the tentacles becoming inflected in a few minutes. Two leaves were immersed in a weak solution of two drops to 1 oz., or 437 grains, of water. When examined after 3 hrs. 20 m., only twenty-one tentacles on one leaf were closely inflected, and six on the other slightly so; but all the glands were blackened, or very dark-coloured, with the protoplasm in all the cells of all the tentacles much aggregated and dark-coloured. The leaves were not quite killed, for on being placed in a little solution of carbonate of ammonia (2 grs. to 1 oz.) a few more tentacles became inflected, the remainder not being acted on during the next 24 hrs. Half-minims of a stronger solution (two drops to 1/2 oz. of water) were placed on the discs of six leaves, and in 30 m. all those tentacles became inflected; the glands of which had actually touched the solution, as shown by their blackness; but hardly any motor influence was transmitted to the outer tentacles. After 22 hrs. most of the glands on the discs appeared dead; but this could not have been the case, as when bits of meat were placed on three of them, some few of the outer tentacles were inflected in 24 hrs. Hence nicotine has a great tendency to blacken the glands and to induce aggregation [page 204] of the protoplasm, but, except when pure, has very moderate power of inducing inflection, and still less power of causing a motor influence to be transmitted from the discal glands to the outer tentacles. It is moderately poisonous.
Text from Project Gutenberg, public domain in the United States.