On the 11th this second leaf began to re-expand; the fragment was manifestly softened, and Dr. Klein reports, "a great deal of enamel and the greater part of the dentine decalcified."
Experiment 2.--May 1st, fragment placed on leaf; 2nd, tentacles fairly well inflected, with much secretion on the disc, and remained so until the 7th, when the leaf re-expanded. The fragment was now transferred to a fresh leaf, which next day (8th) was inflected in the strongest manner, and thus remained until the 11th, when it re-expanded. Dr.
Klein reports, "a great deal of enamel and the greater part of the dentine decalcified."
Experiment 3.--May 1st, fragment moistened with saliva and placed on a leaf, which remained well inflected until 5th, when it re-expanded. The enamel was not at all, and the dentine only slightly, softened. The fragment was now transferred to a fresh leaf, which next morning (6th) was strongly inflected, and remained so until the 11th. The enamel and dentine both now somewhat softened; and Dr. Klein reports, "less than half the enamel, but the greater part of the dentine decalcified."
Experiment 4.--May 1st, a minute and thin bit of dentine, moistened with saliva, was placed on a leaf, which was soon inflected, and re-expanded on the 5th. The dentine had become as flexible as thin paper. It was then transferred to a fresh leaf, which next morning (6th) was strongly inflected, and reopened on the 10th. The decalcified dentine was now so tender that it was torn into shreds merely by the force of the re-expanding tentacles.]
From these experiments it appears that enamel is attacked by the secretion with more difficulty than dentine, as might have been expected from its extreme hardness; and both with more difficulty than ordinary bone. After the process of dissolution has once commenced, it is carried on with greater ease; this may be inferred from the leaves, to which the fragments were transferred, becoming in all four cases strongly inflected in the course of a single day; whereas the first set of leaves acted much less quickly and [page 108] energetically. The angles or projections of the fibrous basis of the enamel and dentine (except, perhaps, in No. 4, which could not be well observed) were not in the least rounded; and Dr. Klein remarks that their microscopical structure was not altered. But this could not have been expected, as the decalcification was not complete in the three specimens which were carefully examined.
Fibrous Basis of Bone.--I at first concluded, as already stated, that the secretion could not digest this substance. I therefore asked Dr.
Burdon Sanderson to try bone, enamel, and dentine, in artificial gastric juice, and he found that they were after a considerable time completely dissolved. Dr. Klein examined some of the small lamellae, into which part of the skull of a cat became broken up after about a week"s immersion in the fluid, and he found that towards the edges the "matrix appeared rarefied, thus producing the appearance as if the ca.n.a.liculi of the bone-corpuscles had become larger. Otherwise the corpuscles and their ca.n.a.liculi were very distinct." So that with bone subjected to artificial gastric juice complete decalcification precedes the dissolution of the fibrous basis. Dr. Burdon Sanderson suggested to me that the failure of Drosera to digest the fibrous basis of bone, enamel, and dentine, might be due to the acid being consumed in the decomposition of the earthy salts, so that there was none left for the work of digestion. Accordingly, my son thoroughly decalcified the bone of a sheep with weak hydrochloric acid; and seven minute fragments of the fibrous basis were placed on so many leaves, four of the fragments being first damped with saliva to aid prompt inflection. All seven leaves became inflected, but only very moderately, in the course of a day. [page 109] They quickly began to re-expand; five of them on the second day, and the other two on the third day. On all seven leaves the fibrous tissue was converted into perfectly transparent, viscid, more or less liquefied little ma.s.ses. In the middle, however, of one, my son saw under a high power a few corpuscles, with traces of fibrillation in the surrounding transparent matter. From these facts it is clear that the leaves are very little excited by the fibrous basis of bone, but that the secretion easily and quickly liquefies it, if thoroughly decalcified. The glands which had remained in contact for two or three days with the viscid ma.s.ses were not discoloured, and apparently had absorbed little of the liquefied tissue, or had been little affected by it.
Phosphate of Lime.--As we have seen that the tentacles of the first set of leaves remained clasped for nine or ten days over minute fragments of bone, and the tentacles of the second set for six or seven days over the same fragments, I was led to suppose that it was the phosphate of lime, and not any included animal matter, which caused such long continued inflection. It is at least certain from what has just been shown that this cannot have been due to the presence of the fibrous basis. With enamel and dentine (the former of which contains only 4 per cent. of organic matter) the tentacles of two successive sets of leaves remained inflected altogether for eleven days. In order to test my belief in the potency of phosphate of lime, I procured some from Prof.
Frankland absolutely free of animal matter and of any acid. A small quant.i.ty moistened with water was placed on the discs of two leaves.
One of these was only slightly affected; the other remained closely inflected for ten days, when a few of the tentacles began to [page 110]
re-expand, the rest being much injured or killed. I repeated the experiment, but moistened the phosphate with saliva to insure prompt inflection; one leaf remained inflected for six days (the little saliva used would not have acted for nearly so long a time) and then died; the other leaf tried to re-expand on the sixth day, but after nine days failed to do so, and likewise died. Although the quant.i.ty of phosphate given to the above four leaves was extremely small, much was left in every case undissolved. A larger quant.i.ty wetted with water was next placed on the discs of three leaves; and these became most strongly inflected in the course of 24 hrs. They never re-expanded; on the fourth day they looked sickly, and on the sixth were almost dead. Large drops of not very viscid fluid hung from their edges during the six days. This fluid was tested each day with litmus paper, but never coloured it; and this circ.u.mstance I do not understand, as the superphosphate of lime is acid. I suppose that some superphosphate must have been formed by the acid of the secretion acting on the phosphate, but that it was all absorbed and injured the leaves; the large drops which hung from their edges being an abnormal and dropsical secretion.
Anyhow, it is manifest that the phosphate of lime is a most powerful stimulant. Even small doses are more or less poisonous, probably on the same principle that raw meat and other nutritious substances, given in excess, kill the leaves. Hence the conclusion, that the long continued inflection of the tentacles over fragments of bone, enamel, and dentine, is caused by the presence of phosphate of lime, and not of any included animal matter, is no doubt correct.
Gelatine.--I used pure gelatine in thin sheets given [page 111] me by Prof. Hoffmann. For comparison, squares of the same size as those placed on the leaves were left close by on wet moss. These soon swelled, but retained their angles for three days; after five days they formed rounded, softened ma.s.ses, but even on the eighth day a trace of gelatine could still be detected. Other squares were immersed in water, and these, though much swollen, retained their angles for six days.
Squares of 1/10 of an inch (2.54 mm.), just moistened with water, were placed on two leaves; and after two or three days nothing was left on them but some acid viscid fluid, which in this and other cases never showed any tendency to regelatinise; so that the secretion must act on the gelatine differently to what water does, and apparently in the same manner as gastric juice.* Four squares of the same size as before were then soaked for three days in water, and placed on large leaves; the gelatine was liquefied and rendered acid in two days, but did not excite much inflection. The leaves began to re-expand after four or five days, much viscid fluid being left on their discs, as if but little had been absorbed. One of these leaves, as soon as it re-expanded, caught a small fly, and after 24 hrs. was closely inflected, showing how much more potent than gelatine is the animal matter absorbed from an insect. Some larger pieces of gelatine, soaked for five days in water, were next placed on three leaves, but these did not become much inflected until the third day; nor was the gelatine completely liquefied until the fourth day. On this day one leaf began to re-expand; the second on the fifth; and third on the sixth. These several facts
* Dr. Lauder Brunton, "Handbook for the Phys. Laboratory," 1873, pp.
477, 487; Schiff, "Leons phys. de la Digestion," 1867, p. 249. [page 112]
prove that gelatine is far from acting energetically on Drosera.
In the last chapter it was shown that a solution of isingla.s.s of commerce, as thick as milk or cream, induces strong inflection. I therefore wished to compare its action with that of pure gelatine.
Solutions of one part of both substances to 218 of water were made; and half-minim drops (.0296 ml.) were placed on the discs of eight leaves, so that each received 1/480 of a grain, or .135 mg. The four with the isingla.s.s were much more strongly inflected than the other four. I conclude therefore that isingla.s.s contains some, though perhaps very little, soluble alb.u.minous matter. As soon as these eight leaves re-expanded, they were given bits of roast meat, and in some hours all became greatly inflected; again showing how much more meat excites Drosera than does gelatine or isingla.s.s. This is an interesting fact, as it is well known that gelatine by itself has little power of nourishing animals.*
Chondrin.--This was sent me by Dr. Moore in a gelatinous state. Some was slowly dried, and a small chip was placed on a leaf, and a much larger chip on a second leaf. The first was liquefied in a day; the larger piece was much swollen and softened, but was not completely liquefied until the third day. The undried jelly was next tried, and as a control experiment small cubes were left in water for four days and retained their angles. Cubes of the same size were placed on two leaves, and larger cubes on two other leaves. The tentacles and laminae of the latter were closely inflected after 22 hrs., but those of the
* Dr. Lauder Brunton gives in the "Medical Record," January 1873, p.
36, an account of Voit"s view of the indirect part which gelatine plays in nutrition. [page 113]
two leaves with the smaller cubes only to a moderate degree. The jelly on all four was by this time liquefied, and rendered very acid. The glands were blackened from the aggregation of their protoplasmic contents. In 46 hrs. from the time when the jelly was given, the leaves had almost re-expanded, and completely so after 70 hrs.; and now only a little slightly adhesive fluid was left unabsorbed on their discs.
One part of chondrin jelly was dissolved in 218 parts of boiling water, and half-minim drops were given to four leaves; so that each received about 1/480 of a grain (.135 mg.) of the jelly; and, of course, much less of dry chondrin. This acted most powerfully, for after only 3 hrs.
30 m. all four leaves were strongly inflected. Three of them began to re-expand after 24 hrs., and in 48 hrs. were completely open; but the fourth had only partially re-expanded. All the liquefied chondrin was by this time absorbed. Hence a solution of chondrin seems to act far more quickly and energetically than pure gelatine or isingla.s.s; but I am a.s.sured by good authorities that it is most difficult, or impossible, to know whether chondrin is pure, and if it contained any alb.u.minous compound, this would have produced the above effects.
Nevertheless, I have thought these facts worth giving, as there is so much doubt on the nutritious value of gelatine; and Dr. Lauder Brunton does not know of any experiments with respect to animals on the relative value of gelatine and chondrin.
Milk.--We have seen in the last chapter that milk acts most powerfully on the leaves; but whether this is due to the contained casein or alb.u.men, I know not. Rather large drops of milk excite so much secretion (which is very acid) that it sometimes trickles down [page 114] from the leaves, and this is likewise characteristic of chemically prepared casein. Minute drops of milk, placed on leaves, were coagulated in about ten minutes. Schiff denies* that the coagulation of milk by gastric juice is exclusively due to the acid which is present, but attributes it in part to the pepsin; and it seems doubtful whether with Drosera the coagulation can be wholly due to the acid, as the secretion does not commonly colour litmus paper until the tentacles have become well inflected; whereas the coagulation commences, as we have seen, in about ten minutes. Minute drops of skimmed milk were placed on the discs of five leaves; and a large proportion of the coagulated matter or curd was dissolved in 6 hrs. and still more completely in 8 hrs. These leaves re-expanded after two days, and the viscid fluid left on their discs was then carefully sc.r.a.ped off and examined. It seemed at first sight as if all the casein had not been dissolved, for a little matter was left which appeared of a whitish colour by reflected light. But this matter, when examined under a high power, and when compared with a minute drop of skimmed milk coagulated by acetic acid, was seen to consist exclusively of oil-globules, more or less aggregated together, with no trace of casein. As I was not familiar with the microscopical appearance of milk, I asked Dr. Lauder Brunton to examine the slides, and he tested the globules with ether, and found that they were dissolved. We may, therefore, conclude that the secretion quickly dissolves casein, in the state in which it exists in milk.
Chemically Prepared Casein.--This substance, which
* "Leons," &c. tom. ii. page 151. [page 115]
is insoluble in water, is supposed by many chemists to differ from the casein of fresh milk. I procured some, consisting of hard globules, from Messrs. Hopkins and Williams, and tried many experiments with it.
Small particles and the powder, both in a dry state and moistened with water, caused the leaves on which they were placed to be inflected very slowly, generally not until two days had elapsed. Other particles, wetted with weak hydrochloric acid (one part to 437 of water) acted in a single day, as did some casein freshly prepared for me by Dr. Moore.
The tentacles commonly remained inflected for from seven to nine days; and during the whole of this time the secretion was strongly acid. Even on the eleventh day some secretion left on the disc of a fully re-expanded leaf was strongly acid. The acid seems to be secreted quickly, for in one case the secretion from the discal glands, on which a little powdered casein had been strewed, coloured litmus paper, before any of the exterior tentacles were inflected.
Small cubes of hard casein, moistened with water, were placed on two leaves; after three days one cube had its angles a little rounded, and after seven days both consisted of rounded softened ma.s.ses, in the midst of much viscid and acid secretion; but it must not be inferred from this fact that the angles were dissolved, for cubes immersed in water were similarly acted on. After nine days these leaves began to re-expand, but in this and other cases the casein did not appear, as far as could be judged by the eye, much, if at all, reduced in bulk.
According to Hoppe-Seyler and Lubavin* casein consists of an alb.u.minous, with
* Dr. Lauder Brunton, "Handbook for Phys. Lab." p. 529. [page 116]
a non-alb.u.minous, substance; and the absorption of a very small quant.i.ty of the former would excite the leaves, and yet not decrease the casein to a perceptible degree. Schiff a.s.serts*--and this is an important fact for us--that "la casine purifie des chemistes est un corps presque compltement inattaquable par le suc gastrique." So that here we have another point of accordance between the secretion of Drosera and gastric juice, as both act so differently on the fresh casein of milk, and on that prepared by chemists.
A few trials were made with cheese; cubes of 1/20 of an inch (1.27 mm.) were placed on four leaves, and these after one or two days became well inflected, their glands pouring forth much acid secretion. After five days they began to re-expand, but one died, and some of the glands on the other leaves were injured. Judging by the eye, the softened and subsided ma.s.ses of cheese, left on the discs, were very little or not at all reduced in bulk. We may, however, infer from the time during which the tentacles remained inflected,--from the changed colour of some of the glands,--and from the injury done to others, that matter had been absorbed from the cheese.
Legumin.--I did not procure this substance in a separate state; but there can hardly be a doubt that it would be easily digested, judging from the powerful effect produced by drops of a decoction of green peas, as described in the last chapter. Thin slices of a dried pea, after being soaked in water, were placed on two leaves; these became somewhat inflected in the course of a single hour, and most strongly so in 21 hrs. They re-expanded after three or four days.
* "Leons" &c. tom. ii. page 153. [page 117]
The slices were not liquefied, for the walls of the cells, composed of cellulose, are not in the least acted on by the secretion.
Pollen.--A little fresh pollen from the common pea was placed on the discs of five leaves, which soon became closely inflected, and remained so for two or three days.
The grains being then removed, and examined under the microscope, were found discoloured, with the oil-globules remarkably aggregated. Many had their contents much shrunk, and some were almost empty. In only a few cases were the pollen-tubes emitted. There could be no doubt that the secretion had penetrated the outer coats of the grains, and had partially digested their contents. So it must be with the gastric juice of the insects which feed on pollen, without masticating it.* Drosera in a state of nature cannot fail to profit to a certain extent by this power of digesting pollen, as innumerable grains from the carices, gra.s.ses, rumices, fir-trees, and other wind-fertilised plants, which commonly grow in the same neighbourhood, will be inevitably caught by the viscid secretion surrounding the many glands.
Gluten.--This substance is composed of two alb.u.minoids, one soluble, the other insoluble in alcohol. Some was prepared by merely washing wheaten flour in water. A provisional trial was made with rather large pieces placed on two leaves; these, after 21 hrs., were closely inflected, and remained so for four days, when one was killed and the other had its glands extremely blackened, but was not afterwards observed.
* Mr. A.W. Bennett found the undigested coats of the grains in the intestinal ca.n.a.l of pollen-eating Diptera; see "Journal of Hort. Soc.
of London," vol. iv. 1874, p. 158.
Watts" "Dict. of Chemistry," vol. ii. 1872, p. 873. [page 118]
Smaller bits were placed on two leaves; these were only slightly inflected in two days, but afterwards became much more so. Their secretion was not so strongly acid as that of leaves excited by casein.
The bits of gluten, after lying for three days on the leaves, were more transparent than other bits left for the same time in water. After seven days both leaves re-expanded, but the gluten seemed hardly at all reduced in bulk. The glands which had been in contact with it were extremely black. Still smaller bits of half putrid gluten were now tried on two leaves; these were well inflected in 24 hrs., and thoroughly in four days, the glands in contact being much blackened.
After five days one leaf began to re-expand, and after eight days both were fully re-expanded, some gluten being still left on their discs.
Four little chips of dried gluten, just dipped in water, were next tried, and these acted rather differently from fresh gluten. One leaf was almost fully re-expanded in three days, and the other three leaves in four days. The chips were greatly softened, almost liquefied, but not nearly all dissolved. The glands which had been in contact with them, instead of being much blackened, were of a very pale colour, and many of them were evidently killed.
In not one of these ten cases was the whole of the gluten dissolved, even when very small bits were given. I therefore asked Dr. Burdon Sanderson to try gluten in artificial digestive fluid of pepsin with hydrochloric acid; and this dissolved the whole. The gluten, however, was acted on much more slowly than fibrin; the proportion dissolved within four hours being as 40.8 of gluten to 100 of fibrin. Gluten was also tried in two other digestive fluids, in which hydrochloric acid was replaced by propionic [page 119] and butyric acids, and it was completely dissolved by these fluids at the ordinary temperature of a room. Here, then, at last, we have a case in which it appears that there exists an essential difference in digestive power between the secretion of Drosera and gastric juice; the difference being confined to the ferment, for, as we have just seen, pepsin in combination with acids of the acetic series acts perfectly on gluten. I believe that the explanation lies simply in the fact that gluten is too powerful a stimulant (like raw meat, or phosphate of lime, or even too large a piece of alb.u.men), and that it injures or kills the glands before they have had time to pour forth a sufficient supply of the proper secretion. That some matter is absorbed from the gluten, we have clear evidence in the length of time during which the tentacles remain inflected, and in the greatly changed colour of the glands.
At the suggestion of Dr. Sanderson, some gluten was left for 15 hrs. in weak hydrochloric acid (.02 per cent.), in order to remove the starch.
It became colourless, more transparent, and swollen. Small portions were washed and placed on five leaves, which were soon closely inflected, but to my surprise re-expanded completely in 48 hrs. A mere vestige of gluten was left on two of the leaves, and not a vestige on the other three. The viscid and acid secretion, which remained on the discs of the three latter leaves, was sc.r.a.ped off and examined by my son under a high power; but nothing could be seen except a little dirt, and a good many starch grains which had not been dissolved by the hydrochloric acid. Some of the glands were rather pale. We thus learn that gluten, treated with weak hydrochloric acid, is not so powerful or so enduring a [page 120] stimulant as fresh gluten, and does not much injure the glands; and we further learn that it can be digested quickly and completely by the secretion.
[Globulin or Crystallin.--This substance was kindly prepared for me from the lens of the eye by Dr. Moore, and consisted of hard, colourless, transparent fragments. It is said* that globulin ought to "swell up in water and dissolve, for the most part forming a gummy liquid;" but this did not occur with the above fragments, though kept in water for four days. Particles, some moistened with water, others with weak hydrochloric acid, others soaked in water for one or two days, were placed on nineteen leaves. Most of these leaves, especially those with the long soaked particles, became strongly inflected in a few hours. The greater number re-expanded after three or four days; but three of the leaves remained inflected during one, two, or three additional days. Hence some exciting matter must have been absorbed; but the fragments, though perhaps softened in a greater degree than those kept for the same time in water, retained all their angles as sharp as ever. As globulin is an alb.u.minous substance, I was astonished at this result; and my object being to compare the action of the secretion with that of gastric juice, I asked Dr. Burdon Sanderson to try some of the globulin used by me. He reports that "it was subjected to a liquid containing 0.2 per cent. of hydrochloric acid, and about 1 per cent. of glycerine extract of the stomach of a dog. It was then ascertained that this liquid was capable of digesting 1.31 of its weight of unboiled fibrin in 1 hr.; whereas, during the hour, only 0.141 of the above globulin was dissolved. In both cases an excess of the substance to be digested was subjected to the liquid." We thus see that within the same time less than one-ninth by weight of globulin than of fibrin was dissolved; and bearing in mind that pepsin with acids of the acetic series has only about one-third of the digestive power of pepsin with hydrochloric acid, it is not surprising that the fragments of
* Watts" "Dictionary of Chemistry," vol. ii. page 874.
I may add that Dr. Sanderson prepared some fresh globulin by Schmidt"s method, and of this 0.865 was dissolved within the same time, namely, one hour; so that it was far more soluble than that which I used, though less soluble than fibrin, of which, as we have seen, 1.31 was dissolved. I wish that I had tried on Drosera globulin prepared by this method. [page 121]
globulin were not corroded or rounded by the secretion of Drosera, though some soluble matter was certainly extracted from them and absorbed by the glands.
Haematin.--Some dark red granules, prepared from bullock"s blood, were given me; these were found by Dr. Sanderson to be insoluble in water, acids, and alcohol, so that they were probably haematin, together with other bodies derived from the blood. Particles with little drops of water were placed on four leaves, three of which were pretty closely inflected in two days; the fourth only moderately so. On the third day the glands in contact with the haematin were blackened, and some of the tentacles seemed injured. After five days two leaves died, and the third was dying; the fourth was beginning to re-expand, but many of its glands were blackened and injured. It is therefore clear that matter had been absorbed which was either actually poisonous or of too stimulating a nature. The particles were much more softened than those kept for the same time in water, but, judging by the eye, very little reduced in bulk. Dr. Sanderson tried this substance with artificial digestive fluid, in the manner described under globulin, and found that whilst 1.31 of fibrin, only 0.456 of the haematin was dissolved in an hour; but the dissolution by the secretion of even a less amount would account for its action on Drosera. The residue left by the artificial digestive fluid at first yielded nothing more to it during several succeeding days.]
Substances which are not Digested by the Secretion.