all but nine inflected; and of the inflected tentacles all excepting four closely; blade slightly inflected. After 8 hrs. blade quite doubled up, and now all the tentacles excepting [page 162] eight closely inflected. The leaf remained in this state for two days.
(4) After 2 hrs. 20 m. only fifty-nine tentacles inflected; but after 5 hrs. all the tentacles closely inflected excepting two which were not affected, and eleven which were only sub-inflected; after 7 hrs. blade considerably inflected; after 12 hrs. much re-expansion.
(5) After 4 hrs. all the tentacles but fourteen inflected; after 9 hrs.
30 m. beginning to re-expand.
(6) After 1 hr. thirty-six tentacles inflected; after 5 hrs. all but fifty-four inflected; after 12 hrs. considerable re-expansion.
(7) After 4 hrs. 30 m. only thirty-five tentacles inflected or sub-inflected, and this small amount of inflection never increased.
Now for the seven corresponding leaves in water:--
(1) After 4 hrs. thirty-eight tentacles inflected; but after 7 hrs.
these, with the exception of six, re-expanded.
(2) After 4 hrs. 20 m. twenty inflected; these after 9 hrs. partially re-expanded.
(3) After 4 hrs. five inflected, which began to re-expand after 7 hrs.
(4) After 24 hrs. one inflected.
(5), (6) and (7) Not at all affected, though observed for 24 hrs., excepting the short tentacles on the borders of the disc, which as usual formed a ring.
A comparison of the leaves in the solution, especially of the first five or even six on the list, with those in the water, after 1 hr. or after 4 hrs., and in a still more marked degree after 7 hrs. or 8 hrs., could not leave the least doubt that the solution had produced a great effect. This was shown not only by the vastly greater number of inflected tentacles, but by the degree or closeness of their inflection, and by that of their blades. Yet each gland on leaf No. 1 (which bore 255 glands, all of which, excepting five, were inflected in 30 m.) could not have received more than one-four-millionth of a grain (.0000162 mg.) of the salt. Again, each gland on leaf No. 3 (which bore 233 glands, all of which, except nine, were inflected in 2 hrs. 30 m.) could have received at most only the 1/3584000 of a grain, or .0000181 mg.
Four leaves were immersed as before in a solution of one part to 656,250 of water (1 gr. to 1500 oz.); but on this occasion I happened to select leaves which were very little sensitive, as on other occasions I chanced to select unusually sensitive leaves. The leaves were not more affected after 12 hrs. than [page 163] the four corresponding ones in water; but after 24 hrs. they were slightly more inflected. Such evidence, however, is not at all trustworthy.
Twelve leaves were immersed, each in thirty minims of a solution of one part to 1,312,500 of water (1 gr. to 3000 oz.); so that each leaf received 1/48000 of a grain (.00135 mg.). The leaves were not in very good condition; four of them were too old and of a dark red colour; four were too pale, yet one of these latter acted well; the four others, as far as could be told by the eye, seemed in excellent condition. The result was as follows:--
(1) This was a pale leaf; after 40 m. about thirty-eight tentacles inflected; after 3 hrs. 30 m. the blade and many of the outer tentacles inflected; after 10 hrs. 15 m. all the tentacles but seventeen inflected, and the blade quite doubled up; after 24 hrs. all the tentacles but ten more or less inflected. Most of them were closely inflected, but twenty-five were only sub-inflected.
(2) After 1 hr. 40 m. twenty-five tentacles inflected; after 6 hrs. all but twenty-one inflected; after 10 hrs. all but sixteen more or less inflected; after 24 hrs. re-expanded.
(3) After 1 hr. 40 m. thirty-five inflected; after 6 hrs. "a large number" (to quote my own memorandum) inflected, but from want of time they were not counted; after 24 hrs. re-expanded.
(4) After 1 hr. 40 m. about thirty inflected; after 6 hrs. "a large number all round the leaf" inflected, but they were not counted; after 10 hrs. began to re-expand.
(5) to (12) These were not more inflected than leaves often are in water, having respectively 16, 8, 10, 8, 4, 9, 14, and 0 tentacles inflected. Two of these leaves, however, were remarkable from having their blades slightly inflected after 6 hrs.
With respect to the twelve corresponding leaves in water, (1) had, after 1 hr. 35 m., fifty tentacles inflected, but after 11 hrs. only twenty-two remained so, and these formed a group, with the blade at this point slightly inflected. It appeared as if this leaf had been in some manner accidentally excited, for instance by a particle of animal matter which was dissolved by the water. (2) After 1 hr. 45 m.
thirty-two tentacles inflected, but after 5 hrs. 30 m. only twenty-five inflected, and these after 10 hrs. all re-expanded; (3) after 1 hr.
twenty-five inflected, which after 10 hrs. 20 m. were all re-expanded; (4) and (5) after 1 hr. 35 m. six and seven tentacles inflected, which re-expanded after 11 hrs.; (6), (7) and (8) from one to three inflected, which [page 164] soon re-expanded; (9), (10), (11) and (12) none inflected, though observed for twenty-four hours.
Comparing the states of the twelve leaves in water with those in the solution, there could be no doubt that in the latter a larger number of tentacles were inflected, and these to a greater degree; but the evidence was by no means so clear as in the former experiments with stronger solutions. It deserves attention that the inflection of four of the leaves in the solution went on increasing during the first 6 hrs., and with some of them for a longer time; whereas in the water the inflection of the three leaves which were the most affected, as well as of all the others, began to decrease during this same interval. It is also remarkable that the blades of three of the leaves in the solution were slightly inflected, and this is a most rare event with leaves in water, though it occurred to a slight extent in one (No. 1), which seemed to have been in some manner accidentally excited. All this shows that the solution produced some effect, though less and at a much slower rate than in the previous cases. The small effect produced may, however, be accounted for in large part by the majority of the leaves having been in a poor condition.
Of the leaves in the solution, No. 1 bore 200 glands and received 1/48000 of a grain of the salt. Subtracting the seventeen tentacles which were not inflected, each gland could have absorbed only the 1/8784000 of a grain (.00000738 mg.). This amount caused the tentacle bearing each gland to be greatly inflected. The blade was also inflected.
Lastly, eight leaves were immersed, each in thirty minims of a solution of one part of the phosphate to 21,875,000 of water (1 gr. to 5000 oz.). Each leaf thus received 1/80000 of a grain of the salt, or .00081 mg. I took especial pains in selecting the finest leaves from the hot-house for immersion, both in the solution and the water, and almost all proved extremely sensitive. Beginning as before with those in the solution:--
(1) After 2 hrs. 30 m. all the tentacles but twenty-two inflected, but some only sub-inflected; the blade much inflected; after 6 hrs. 30 m.
all but thirteen inflected, with the blade immensely inflected; and remained so for 48 hrs.
(2) No change for the first 12 hrs., but after 24 hrs. all the tentacles inflected, excepting those of the outermost row, of which only eleven were inflected. The inflection continued to increase, and after 48 hrs. all the tentacles except three were inflected, [page 165]
and most of them rather closely, four or five being only sub-inflected.
(3) No change for the first 12 hrs.; but after 24 hrs. all the tentacles excepting those of the outermost row were sub-inflected, with the blade inflected. After 36 hrs. blade strongly inflected, with all the tentacles, except three, inflected or sub-inflected. After 48 hrs.
in the same state.
(4) to (8) These leaves, after 2 hrs. 30 m., had respectively 32, 17, 7, 4, and 0 tentacles inflected, most of which, after a few hours, re-expanded, with the exception of No. 4, which retained its thirty-two tentacles inflected for 48 hrs.
Now for the eight corresponding leaves in water:--
(1) After 2 hrs. 40 m. this had twenty of its outer tentacles inflected, five of which re-expanded after 6 hrs. 30 m. After 10 hrs.
15 m. a most unusual circ.u.mstance occurred, namely, the whole blade became slightly bowed towards the footstalk, and so remained for 48 hrs. The exterior tentacles, excepting those of the three or four outermost rows, were now also inflected to an unusual degree.
(2) to (8) These leaves, after 2 hrs. 40 m., had respectively 42, 12, 9, 8, 2, 1, and 0 tentacles inflected, which all re-expanded within 24 hrs., and most of them within a much shorter time.
When the two lots of eight leaves in the solution and in the water were compared after the lapse of 24 hrs., they undoubtedly differed much in appearance. The few tentacles on the leaves in water which were inflected had after this interval re-expanded, with the exception of one leaf; and this presented the very unusual case of the blade being somewhat inflected, though in a degree hardly approaching that of the two leaves in the solution. Of these latter leaves, No. 1 had almost all its tentacles, together with its blade, inflected after an immersion of 2 hrs. 30 m. Leaves No. 2 and 3 were affected at a much slower rate; but after from 24 hrs. to 48 hrs. almost all their tentacles were closely inflected, and the blade of one quite doubled up. We must therefore admit, incredible as the fact may at first appear, that this extremely weak solution acted on the more sensitive leaves; each of which received only the 1/80000 of a grain (.00081 mg.) of the phosphate. Now, leaf No. 3 bore 178 tentacles, and subtracting the three which were not inflected, each gland could have absorbed only the 1/14000000 of a grain, or .00000463 mg. Leaf No. 1, which was strongly acted on within 2 hrs. 30 m., and had all its outer tentacles, except thirteen, inflected within 6 hrs. 30 m., bore 260 tentacles; and on the same principle as before, each gland could have [page 166]
absorbed only 1/19760000 of a grain, or .00000328 mg.; and this excessively minute amount sufficed to cause all the tentacles bearing these glands to be greatly inflected. The blade was also inflected.]
Summary of the Results with Phosphate of Ammonia.--The glands of the disc, when excited by a half-minim drop (.0296 ml.), containing 1/3840 of a grain (.0169 mg.) of this salt, transmit a motor impulse to the exterior tentacles, causing them to bend inwards. A minute drop, containing 1/153600 of a grain (.000423 mg.), if held for a few seconds in contact with a gland, causes the tentacle bearing this gland to be inflected. If a leaf is left immersed for a few hours, and sometimes for a shorter time, in a solution so weak that each gland can absorb only the 1/9760000 of a grain (.00000328 mg.), this is enough to excite the tentacle into movement, so that it becomes closely inflected, as does sometimes the blade. In the general summary to this chapter a few remarks will be added, showing that the efficiency of such extremely minute doses is not so incredible as it must at first appear.
[Sulphate of Ammonia.--The few trials made with this and the following five salts of ammonia were undertaken merely to ascertain whether they induced inflection. Half-minims of a solution of one part of the sulphate of ammonia to 437 of water were placed on the discs of seven leaves, so that each received 1/960 of a grain, or .0675 mg. After 1 hr. the tentacles of five of them, as well as the blade of one, were strongly inflected. The leaves were not afterwards observed.
Citrate of Ammonia.--Half-minims of a solution of one part to 437 of water were placed on the discs of six leaves. In 1 hr. the short outer tentacles round the discs were a little inflected, with the glands on the discs blackened. After 3 hrs. 25 m. one leaf had its blade inflected, but none of the exterior tentacles. All six leaves remained in nearly the same state during the day, the submarginal tentacles, however, [page 167] becoming more inflected. After 23 hrs. three of the leaves had their blades somewhat inflected; and the submarginal tentacles of all considerably inflected, but in none were the two, three, or four outer rows affected. I have rarely seen cases like this, except from the action of a decoction of gra.s.s. The glands on the discs of the above leaves, instead of being almost black, as after the first hour, were now after 23 hrs. very pale. I next tried on four leaves half-minims of a weaker solution, of one part to 1312 of water (1 gr.
to 3 oz.); so that each received 1/2880 of a grain (.0225 mg.). After 2 hrs. 18 m. the glands on the disc were very dark-coloured; after 24 hrs. two of the leaves were slightly affected; the other two not at all.
Acetate of Ammonia.--Half-minims of a solution of about one part to 109 of water were placed on the discs of two leaves, both of which were acted on in 5 hrs. 30 m., and after 23 hrs. had every single tentacle closely inflected.
Oxalate of Ammonia.--Half-minims of a solution of one part to 218 of water were placed on two leaves, which, after 7 hrs., became moderately, and after 23 hrs. strongly, inflected. Two other leaves were tried with a weaker solution of one part to 437 of water; one was strongly inflected in 7 hrs.; the other not until 30 hrs. had elapsed.
Tartrate of Ammonia.--Half-minims of a solution of one part to 437 of water were placed on the discs of five leaves. In 31 m. there was a trace of inflection in the exterior tentacles of some of the leaves, and this became more decided after 1 hr. with all the leaves; but the tentacles were never closely inflected. After 8 hrs. 30 m. they began to re-expand. Next morning, after 23 hrs., all were fully re-expanded, excepting one which was still slightly inflected. The shortness of the period of inflection in this and the following case is remarkable.
Chloride of Ammonium.--Half-minims of a solution of one part to 437 of water were placed on the discs of six leaves. A decided degree of inflection in the outer and submarginal tentacles was perceptible in 25 m.; and this increased during the next three or four hours, but never became strongly marked. After only 8 hrs. 30 m. the tentacles began to re-expand, and by the next morning, after 24 hrs., were fully re-expanded on four of the leaves, but still slightly inflected on two.]
General Summary and Concluding Remarks on the Salts of Ammonia.--We have now seen that the nine [page 168] salts of ammonia which were tried, all cause the inflection of the tentacles, and often of the blade of the leaf. As far as can be ascertained from the superficial trials with the last six salts, the citrate is the least powerful, and the phosphate certainly by far the most. The tartrate and chloride are remarkable from the short duration of their action. The relative efficiency of the carbonate, nitrate, and phosphate, is shown in the following table by the smallest amount which suffices to cause the inflection of the tentacles.
Column 1 : Solutions, how applied. Column 2 : Carbonate of Ammonia.
Column 3 : Nitrate of Ammonia. Column 4 : Phosphate of Ammonia.
Placed on the glands of the disc, so as to act indirectly on the outer tentacles : 1/960 of a grain, or 0675 mg. : 1/2400 of a grain, or .027 mg. : 1/3840 of a grain, or .0169 mg.
Applied for a few seconds directly to the gland of an outer tentacle : 1/14400 of a grain, or .00445 mg. : 1/28800 of a grain, or .0025 mg.
grain, 1/153600 of a grain, or .000423 mg.
Leaf immersed, with time allowed for each gland to absorb all that it can : 1/268800 of a grain, or .00024 mg. : 1/691200 of a grain, or .0000937 mg. : 1/19760000 of a grain, or .00000328 mg.
Amount absorbed by a gland which suffices to cause the aggregation of the protoplasm in the adjoining cells of the tentacles. 1/134400 of a grain, or .00048 mg.