THE DIGESTIVE POWER OF THE SECRETION OF DROSERA.
The secretion rendered acid by the direct and indirect excitement of the glands--Nature of the acid--Digestible substances--Albumen, its digestion arrested by alkalies, recommences by the addition of an acid--Meat--Fibrin--Syntonin--Areolar tissue--Cartilage--Fibro-cartilage-- Bone--Enamel and dentine--Phosphate of lime--Fibrous basis of bone--Gelatine--Chondrin--Milk, casein and cheese--Gluten--Legumin--Pollen--Globulin--Haematin--Indigestible substances--Epidermic productions--Fibro-elastic tissue--Mucin--Pepsin--Urea--Chitine--Cellulose--Gun-cotton--Chlorophyll--Fat and oil--Starch--Action of the secretion on living seeds--Summary and concluding remarks.
AS we have seen that nitrogenous fluids act very differently on the leaves of Drosera from non-nitrogenous fluids, and as the leaves remain clasped for a much longer time over various organic bodies than over inorganic bodies, such as bits of glass, cinder, wood, &c., it becomes an interesting inquiry, whether they can only absorb matter already in solution, or render it soluble,--that is, have the power of digestion. We shall immediately see that they certainly have this power, and that they act on albuminous compounds in exactly the same manner as does the gastric juice of mammals; the digested matter being afterwards absorbed. This fact, which will be clearly proved, is a wonderful one in the physiology of plants. I must here state that I have been aided throughout all my later experiments by many valuable suggestions and assistance given me with the greatest kindness by Dr. Burdon Sanderson.
It may be well to premise for the sake of any reader who knows nothing about the digestion of albuminous compounds by animals that this is effected by means of a ferment, pepsin, together with weak hydrochloric acid, though almost any acid will serve. Yet neither pepsin nor an acid by itself has any such power.* We have seen that when the glands of the disc are excited by the contact of any object, especially of one containing nitrogenous matter, the outer tentacles and often the blade become inflected; the leaf being thus converted into a temporary cup or stomach. At the same time the discal glands secrete more copiously, and the secretion becomes acid. Moreover, they transmit some influence to the glands of the exterior tentacles, causing them to pour forth a more copious secretion, which also becomes acid or more acid than it was before.
As this result is an important one, I will give the evidence. The secretion of many glands on thirty leaves, which had not been in any way excited, was tested with litmus paper; and the secretion of twenty-two of these leaves did not in the least affect the colour, whereas that of eight caused an exceedingly feeble and sometimes doubtful tinge of red. Two other old leaves, however, which appeared to have been inflected several times, acted much more decidedly on the paper. Particles of clean glass were then placed on five of the leaves, cubes of albumen on six, and bits of raw meat on three, on none of which was the secretion at this time in the least acid. After an interval of 24 hrs., when almost all the tentacles on
* It appears, however, according to Schiff, and contrary to the opinion of some physiologists, that weak hydrochloric dissolves, though slowly, a very minute quantity of coagulated albumen. Schiff, 'Phys. de la Digestion,' tom. ii. 1867, p. 25.
these fourteen leaves had become more or less inflected, I again tested the secretion, selecting glands which had not as yet reached the centre or touched any object, and it was now plainly acid. The degree of acidity of the secretion varied somewhat on the glands of the same leaf. On some leaves, a few tentacles did not, from some unknown cause, become inflected, as often happens; and in five instances their secretion was found not to be in the least acid; whilst the secretion of the adjoining and inflected tentacles on the same leaf was decidedly acid. With leaves excited by particles of glass placed on the central glands, the secretion which collects on the disc beneath them was much more strongly acid than that poured forth from the exterior tentacles, which were as yet only moderately inflected. When bits of albumen (and this is naturally alkaline), or bits of meat were placed on the disc, the secretion collected beneath them was likewise strongly acid. As raw meat moistened with water is slightly acid, I compared its action on litmus paper before it was placed on the leaves, and afterwards when bathed in the secretion; and there could not be the least doubt that the latter was very much more acid. I have indeed tried hundreds of times the state of the secretion on the discs of leaves which were inflected over various objects, and never failed to find it acid. We may, therefore, conclude that the secretion from unexcited leaves, though extremely viscid, is not acid or only slightly so, but that it becomes acid, or much more strongly so, after the tentacles have begun to bend over any inorganic or organic object; and still more strongly acid after the tentacles have remained for some time closely clasped over any object.
I may here remind the reader that the secretion appears to be to a certain extent antiseptic, as it checks the appearance of mould and infusoria, thus preventing for a time the discoloration and decay of such substances as the white of an egg, cheese, &c. It therefore acts like the gastric juice of the higher animals, which is known to arrest putrefaction by destroying the microzymes.
[As I was anxious to learn what acid the secretion contained, 445 leaves were washed in distilled water, given me by Prof. Frankland; but the secretion is so viscid that it is scarcely possible to scrape or wash off the whole. The conditions were also unfavourable, as it was late in the year and the leaves were small. Prof. Frankland with great kindness undertook to test the fluid thus collected. The leaves were excited by clean particles of glass placed on them 24 hrs. previously. No doubt much more acid would have been secreted had the leaves been excited by animal matter, but this would have rendered the analysis more difficult. Prof. Frankland informs me that the fluid contained no trace of hydrochloric, sulphuric, tartaric, oxalic, or formic acids. This having been ascertained, the remainder of the fluid was evaporated nearly to dryness, and acidified with sulphuric acid; it then evolved volatile acid vapour, which was condensed and digested with carbonate of silver. "The weight of the silver salt thus produced was only .37 gr., much too small a quantity for the accurate determination of the molecular weight of the acid. The number obtained, however, corresponded nearly with that of propionic acid; and I believe that this, or a mixture of acetic and butyric acids, were present in the liquid. The acid doubtless belongs to the acetic or fatty series."
Prof. Frankland, as well as his assistant, observed (and this is an important fact) that the fluid, "when acidified with sulphuric acid, emitted a powerful odour like that of pepsin." The leaves from which the secretion had been washed were also sent to Prof. Frankland; they were macerated for some hours, then acidified with sulphuric acid and distilled, but no acid passed over. Therefore the acid which fresh leaves contain, as shown by their discolouring litmus paper when crushed, must be of a different nature from that present in the secretion. Nor was any odour of pepsin emitted by them.
Although it has long been known that pepsin with acetic acid has the power of digesting albuminous compounds, it appeared advisable to ascertain whether acetic acid could be replaced, without the loss of digestive power, by the allied acids which are believed to occur in the secretion of Drosera, namely, propionic, butyric, or valerianic. Dr. Burdon Sanderson was so kind as to make for me the following experiments, the results of which are valuable, independently of the present inquiry. Prof. Frankland supplied the acids.
"1. The purpose of the following experiments was to determine the digestive activity of liquids containing pepsin, when acidulated with certain volatile acids belonging to the acetic series, in comparison with liquids acidulated with hydrochloric acid, in proportion similar to that in which it exists in gastric juice.
"2. It has been determined empirically that the best results are obtained in artificial digestion when a liquid containing two per thousand of hydrochloric acid gas by weight is used. This corresponds to about 6.25 cubic centimetres per litre of ordinary strong hydrochloric acid. The quantities of propionic, butyric, and valerianic acids respectively which are required to neutralise as much base as 6.25 cubic centimetres of HCl, are in grammes 4.04 of propionic acid, 4.82 of butyric acid, and 5.68 of valerianic acid. It was therefore judged expedient, in comparing the digestive powers of these acids with that of hydrochloric acid, to use them in these proportions.
"3. Five hundred cub. cent. of a liquid containing about 8 cub. cent. of a glycerine extract of the mucous membrane of the stomach of a dog killed during digestion having been prepared, 10 cub. cent. of it were evaporated and dried at 110o. This quantity yielded 0.0031 of residue.
"4. Of this liquid four quantities were taken which were severally acidulated with hydrochloric, propionic, butyric, and valerianic acids, in the proportions above indicated. Each liquid was then placed in a tube, which was allowed to float in a water bath, containing a thermometer which indicated a temperature of 38o to 40o Cent. Into each, a quantity of unboiled fibrin was introduced, and the whole allowed to stand for four hours, the temperature being maintained during the whole time, and care being taken that each contained throughout an excess of fibrin. At the end of the period each liquid was filtered. Of the filtrate, which of course contained as much of the fibrin as had been digested during the four hours, 10 cub. cent. were measured out and evaporated, and dried at 110o as before. The residues were respectively--
"In the liquid containing hydrochloric acid 0.4079 " " propionic acid 0.0601 " " butyric acid 0.1468 " " valerianic acid 0.1254
"Hence, deducting from each of these the above-mentioned residue, left when the digestive liquid itself was evaporated, viz. 0.0031, we have,
"For propionic acid 0.0570 " butyric acid 0.1437 " valerianic acid 0.1223
as compared with 0.4048 for hydrochloric acid; these several numbers expressing the quantities of fibrin by weight digested in presence of equivalent quantities of the respective acids under identical conditions.
"The results of the experiment may be stated thus:--If 100 represent the digestive power of a liquid containing pepsin with the usual proportion of hydrochloric acid, 14.0, 35.4, and 30.2, will represent respectively the digestive powers of the three acids under investigation.
"5. In a second experiment in which the procedure was in every respect the same, excepting that all the tubes were plunged into the same water-bath, and the residues dried at 115o C., the results were as follows:--
"Quantity of fibrin dissolved in four hours by 10 cub. cent. of the liquid:--
"Propionic acid 0.0563 Butyric acid 0.0835 Valerianic acid 0.0615
"The quantity digested by a similar liquid containing hydrochloric acid was 0.3376. Hence, taking this as 100, the following numbers represent the relative quantities digested by the other acids:--
"Propionic acid 16.5 Butyric acid 24.7 Valerianic acid 16.1
"6. A third experiment of the same kind gave:
"Quantity of fibrin digested in four hours by 10 cub. cent. of the liquid:--
"Hydrochloric acid 0.2915 Propionic acid 0.1490 Butyric acid 0.1044 Valerianic acid 0.0520
"Comparing, as before, the three last numbers with the first taken as 100, the digestive power of propionic acid is represented by 16.8; that of butyric acid by 35.8; and that of valerianic by 17.8.
"The mean of these three sets of observations (hydrochloric acid being taken as 100) gives for
"Propionic acid 15.8 Butyric acid 32.0 Valerianic acid 21.4
"7. A further experiment was made to ascertain whether the digestive activity of butyric acid (which was selected as being apparently the most efficacious) was relatively greater at ordinary temperatures than at the temperature of the body. It was found that whereas 10 cub. cent. of a liquid containing the ordinary proportion of hydrochloric acid digested 0.1311 gramme, a similar liquid prepared with butyric acid digested 0.0455 gramme of fibrin.
"Hence, taking the quantities digested with hydrochloric acid at the temperature of the body as 100, we have the digestive power of hydrochloric acid at the temperature of 16o to 18oCent. represented by 44.9; that of butyric acid at the same temperature being 15.6."
We here see that at the lower of these two temperatures, hydrochloric acid with pepsin digests, within the same time, rather less than half the quantity of fibrin compared with what it digests at the higher temperature; and the power of butyric acid is reduced in the same proportion under similar conditions and temperatures. We have also seen that butyric acid, which is much more efficacious than propionic or valerianic acids, digests with pepsin at the higher temperature less than a third of the fibrin which is digested at the same temperature by hydrochloric acid.
I will now give in detail my experiments on the digestive power of the secretion of Drosera, dividing the substances tried into two series, namely those which are digested more or less completely, and those which are not digested. We shall presently see that all these substances are acted on by the gastric juice of the higher animals in the same manner. I beg leave to call attention to the experiments under the head albumen, showing that the secretion loses its power when neutralised by an alkali, and recovers it when an acid is added.
Substances which are completely or partially digested by the Secretion of Drosera.
Albumen.--After having tried various substances, Dr. Burdon Sanderson suggested to me the use of cubes of coagulated albumen or hard-boiled egg. I may premise that five cubes of the same size as those used in the following experiments were placed for the sake of comparison at the same time on wet moss close to the plants of Drosera. The weather was hot, and after four days some of the cubes were discoloured and mouldy, with their angles a little rounded; but they were not surrounded by a zone of transparent fluid as in the case of those undergoing digestion. Other cubes retained their angles and white colour. After eight days all were somewhat reduced in size, discoloured, with their angles much rounded. Nevertheless in four out of the five specimens, the central parts were still white and opaque. So that their state differed widely, as we shall see, from that of the cubes subjected to the action of the secretion.
Rather large cubes of albumen were first tried; the tentacles were well inflected in 24 hrs.; after an additional day the angles of the cubes were dissolved and rounded;* but the cubes were too large, so that the leaves were injured, and after seven days one died and the others were dying. Albumen which has been kept for four or five days, and which, it may be presumed, has begun to decay slightly, seems to act more quickly than freshly boiled eggs. As the latter were generally used, I often moistened them with a little saliva, to make the tentacles close more quickly.
Experiment 2.--A cube of 1/10 of an inch (i.e. with each side 1/10 of an inch, or 2.54 mm. in length) was placed on a leaf, and after 50 hrs. it was converted into a sphere about 3/40 of an inch (1.905 mm.) in diameter, surrounded by perfectly transparent fluid. After ten days the leaf re-expanded, but there was still left on the disc a minute bit of albumen now rendered transparent. More albumen had been given to this leaf than could be dissolved or digested.
Experiment 3.--Two cubes of albumen of 1/20 of an inch (1.27 mm.) were placed on two leaves. After 46 hrs. every atom of one was dissolved, and most of the liquefied matter was absorbed, the fluid which remained being in this, as in all other cases, very acid and viscid. The other cube was acted on at a rather slower rate.
Experiment 4.--Two cubes of albumen of the same size as the last were placed on two leaves, and were converted in 50 hrs. into two large drops of transparent fluid; but when these were removed from beneath the inflected tentacles, and viewed by reflected light under the microscope, fine streaks of white opaque matter could be seen in the one, and traces of similar streaks in the other. The drops were replaced on the leaves, which re-expanded after 10 days; and now nothing was left except a very little transparent acid fluid.
Experiment 5.--This experiment was slightly varied, so that the albumen might be more quickly exposed to the action of the secretion. Two cubes, each of about 1/40 of an inch (.635 mm.), were placed on the same leaf, and two similar cubes on another
* In all my numerous experiments on the digestion of cubes of albumen, the angles and edges were invariably first rounded. Now, Schiff states ('Leons phys. de la Digestion,' vol. ii. 1867, page 149) that this is characteristic of the digestion of albumen by the gastric juice of animals. On the other hand, he remarks "les dissolutions, en chimie, ont lieu sur toute la surface des corps en contact avec l'agent dissolvant."
leaf. These were examined after 21 hrs. 30 m., and all four were found rounded. After 46 hrs. the two cubes on the one leaf were completely liquefied, the fluid being perfectly transparent; on the other leaf some opaque white streaks could still be seen in the midst of the fluid. After 72 hrs. these streaks disappeared, but there was still a little viscid fluid left on the disc; whereas it was almost all absorbed on the first leaf. Both leaves were now beginning to re-expand.]
The best and almost sole test of the presence of some ferment analogous to pepsin in the secretion appeared to be to neutralise the acid of the secretion with an alkali, and to observe whether the process of digestion ceased; and then to add a little acid and observe whether the process recommenced. This was done, and, as we shall see, with success, but it was necessary first to try two control experiments; namely, whether the addition of minute drops of water of the same size as those of the dissolved alkalies to be used would stop the process of digestion; and, secondly, whether minute drops of weak hydrochloric acid, of the same strength and size as those to be used, would injure the leaves. The two following experiments were therefore tried:--
Experiment 6.--Small cubes of albumen were put on three leaves, and minute drops of distilled water on the head of a pin were added two or three times daily. These did not in the least delay the process; for, after 48 hrs., the cubes were completely dissolved on all three leaves. On the third day the leaves began to re-expand, and on the fourth day all the fluid was absorbed.
Experiment 7.--Small cubes of albumen were put on two leaves, and minute drops of hydrochloric acid, of the strength of one part to 437 of water, were added two or three times. This did not in the least delay, but seemed rather to hasten, the process of digestion; for every trace of the albumen disappeared in 24 hrs. 30 m. After three days the leaves partially re-expanded, and by this time almost all the viscid fluid on their discs was absorbed. It is almost superfluous to state that cubes of albumen of the same size as those above used, left for seven days in a little hydrochloric acid of the above strength, retained all their angles as perfect as ever.
Experiment 8.--Cubes of albumen (of 1/20 of an inch, or 2.54 mm.) were placed on five leaves, and minute drops of a solution of one part of carbonate of soda to 437 of water were added at intervals to three of them, and drops of carbonate of potash of the same strength to the other two. The drops were given on the head of a rather large pin, and I ascertained that each was equal to about 1/10 of a minim (.0059 ml.), so that each contained only 1/4800 of a grain (.0135 mg.) of the alkali. This was not sufficient, for after 46 hrs. all five cubes were dissolved.
Experiment 9.--The last experiment was repeated on four leaves, with this difference, that drops of the same solution of carbonate of soda were added rather oftener, as often as the secretion became acid, so that it was much more effectually neutralised. And now after 24 hrs. the angles of three of the cubes were not in the least rounded, those of the fourth being so in a very slight degree. Drops of extremely weak hydrochloric acid (viz. one part to 847 of water) were then added, just enough to neutralise the alkali which was still present; and now digestion immediately recommenced, so that after 23 hrs. 30 m. three of the cubes were completely dissolved, whilst the fourth was converted into a minute sphere, surrounded by transparent fluid; and this sphere next day disappeared.
Experiment 10.--Stronger solutions of carbonate of soda and of potash were next used, viz. one part to 109 of water; and as the same-sized drops were given as before, each drop contained 1/1200 of a grain (.0539 mg.) of either salt. Two cubes of albumen (each about 1/40 of an inch, or .635 mm.) were placed on the same leaf, and two on another. Each leaf received, as soon as the secretion became slightly acid (and this occurred four times within 24 hrs.), drops either of the soda or potash, and the acid was thus effectually neutralised. The experiment now succeeded perfectly, for after 22 hrs. the angles of the cubes were as sharp as they were at first, and we know from experiment 5 that such small cubes would have been completely rounded within this time by the secretion in its natural state. Some of the fluid was now removed with blotting-paper from the discs of the leaves, and minute drops of hydrochloric acid of the strength of the one part to 200 of water was added. Acid of this greater strength was used as the solutions of the alkalies were stronger. The process of digestion now commenced, so that within 48 hrs. from the time when the acid was given the four cubes were not only completely dissolved, but much of the liquefied albumen was absorbed.
Experiment 11.--Two cubes of albumen (1/40 of an inch, or .635 mm.) were placed on two leaves, and were treated with alkalies as in the last experiment, and with the same result; for after 22 hrs. they had their angles perfectly sharp, showing that the digestive process had been completely arrested. I then wished to ascertain what would be the effect of using stronger hydrochloric acid; so I added minute drops of the strength of 1 per cent. This proved rather too strong, for after 48 hrs. from the time when the acid was added one cube was still almost perfect, and the other only very slightly rounded, and both were stained slightly pink. This latter fact shows that the leaves were injured,* for during the normal process of digestion the albumen is not thus coloured, and we can thus understand why the cubes were not dissolved.]
From these experiments we clearly see that the secretion has the power of dissolving albumen, and we further see that if an alkali is added, the process of digestion is stopped, but immediately recommences as soon as the alkali is neutralised by weak hydrochloric acid. Even if I had tried no other experiments than these, they would have almost sufficed to prove that the glands of Drosera secrete some ferment analogous to pepsin, which in presence of an acid gives to the secretion its power of dissolving albuminous compounds.
Splinters of clean glass were scattered on a large number of leaves, and these became moderately inflected. They were cut off and divided into three lots; two of them, after being left for some time in a little distilled water, were strained, and some dis-
* Sachs remarks ('Trait de Bot.' 1874, p. 774), that cells which are killed by freezing, by too great heat, or by chemical agents, allow all their colouring matter to escape into the surrounding water.
coloured, viscid, slightly acid fluid was thus obtained. The third lot was well soaked in a few drops of glycerine, which is well known to dissolve pepsin. Cubes of albumen (1/20 of an inch) were now placed in the three fluids in watch-glasses, some of which were kept for several days at about 90o Fahr. (32o.2 Cent.), and others at the temperature of my room; but none of the cubes were dissolved, the angles remaining as sharp as ever. This fact probably indicates that the ferment is not secreted until the glands are excited by the absorption of a minute quantity of already soluble animal matter,--a conclusion which is supported by what we shall hereafter see with respect to Dionaea. Dr. Hooker likewise found that, although the fluid within the pitchers of Nepenthes possesses extraordinary power of digestion, yet when removed from the pitchers before they have been excited and placed in a vessel, it has no such power, although it is already acid; and we can account for this fact only on the supposition that the proper ferment is not secreted until some exciting matter is absorbed.
On three other occasions eight leaves were strongly excited with albumen moistened with saliva; they were then cut off, and allowed to soak for several hours or for a whole day in a few drops of glycerine. Some of this extract was added to a little hydrochloric acid of various strengths (generally one to 400 of water), and minute cubes of albumen were placed in the mixture.* In two of these trials the cubes were not in the least acted on; but in the third
* As a control experiment bits of albumen were placed in the same glycerine with hydrochloric acid of the same strength; and the albumen, as might have been expected, was not in the least affected after two days.
the experiment was successful. For in a vessel containing two cubes, both were reduced in size in 3 hrs.; and after 24 hrs. mere streaks of undissolved albumen were left. In a second vessel, containing two minute ragged bits of albumen, both were likewise reduced in size in 3 hrs., and after 24 hrs. completely disappeared. I then added a little weak hydrochloric acid to both vessels, and placed fresh cubes of albumen in them; but these were not acted on. This latter fact is intelligible according to the high authority of Schiff,* who has demonstrated, as he believes, in opposition to the view held by some physiologists, that a certain small amount of pepsin is destroyed during the act of digestion. So that if my solution contained, as is probable, an extremely small amount of the ferment, this would have been consumed by the dissolution of the cubes of albumen first given; none being left when the hydrochloric acid was added. The destruction of the ferment during the process of digestion, or its absorption after the albumen had been converted into a peptone, will also account for only one out of the three latter sets of experiments having been successful.
Digestion of Roast Meat.--Cubes of about 1/20 of an inch (1.27 mm.) of moderately roasted meat were placed on five leaves which became in 12 hrs. closely inflected. After 48 hrs. I gently opened one leaf, and the meat now consisted of a minute central sphere, partially digested and surrounded by a thick envelope of transparent viscid fluid. The whole, without being much disturbed, was removed and placed under the microscope. In the central part the transverse striae on the muscular fibres were quite distinct; and it was
* 'Leons phys. de la Digestion,' 1867, tom. ii. pp. 114-126.
interesting to observe how gradually they disappeared, when the same fibre was traced into the surrounding fluid. They disappeared by the striae being replaced by transverse lines formed of excessively minute dark points, which towards the exterior could be seen only under a very high power; and ultimately these points were lost. When I made these observations, I had not read Schiff's account* of the digestion of meat by gastric juice, and I did not understand the meaning of the dark points. But this is explained in the following statement, and we further see how closely similar is the process of digestion by gastric juice and by the secretion of Drosera.
["On a dit le suc gastrique faisait perdre la fibre musculaire ses stries transversales. Ainsi nonce, cette proposition pourrait donner lieu une quivoque, car ce qui se perd, ce n'est que l'aspect extrieur de la striature et non les lments anatomiques qui la composent. On sait que les stries qui donnent un aspect si caractristique la fibre musculaire, sont le rsultat de la juxtaposition et du paralllisme des corpuscules lmentaires, placs, distances gales, dans l'intrieur des fibrilles contigus. Or, ds que le tissu connectif qui relie entre elles les fibrilles lmentaires vient se gonfler et se dissoudre, et que les fibrilles elles-mmes se dissocient, ce paralllisme est dtruit et avec lui l'aspect, le phnomne optique des stries. Si, aprs la dsagrgation des fibres, on examine au microscope les fibrilles lmentaires, on distingue encore trs-nettement leur intrieur les corpuscules, et on continue les voir, de plus en plus ples, jusqu'au moment o les fibrilles elles-mmes se liqufient et disparaissent dans le suc gastrique. Ce qui constitue la striature, proprement parler, n'est donc pas dtruit, avant la liqufaction de la fibre charnue elle-mme."]
In the viscid fluid surrounding the central sphere of undigested meat there were globules of fat and little bits of fibro-elastic tissue; neither of which were in
* 'Leons phys. de la Digestion,' tom. ii. p. 145.
the least digested. There were also little free parallelograms of yellowish, highly translucent matter. Schiff, in speaking of the digestion of meat by gastric juice, alludes to such parallelograms, and says:--
["Le gonflement par lequel commence la digestion de la viande, rsulte de l'action du suc gastrique acide sur le tissu connectif qui se dissout d'abord, et qui, par sa liqufaction, dsagrge les fibrilles. Celles-ci se dissolvent ensuite en grande partie, mais, avant de passer l'tat liquide, elles tendent se briser en petits fragments transversaux. Les 'sarcous elements' de Bowman, qui ne sont autre chose que les produits de cette division transversale des fibrilles lmentaires, peuvent tre prpars et isols l'aide du suc gastrique, pourvu qu'on n'attend pas jusqu' la liqufaction complte du muscle."]
After an interval of 72 hrs., from the time when the five cubes were placed on the leaves, I opened the four remaining ones. On two nothing could be seen but little masses of transparent viscid fluid; but when these were examined under a high power, fat-globules, bits of fibro-elastic tissue, and some few parallelograms of sarcous matter, could be distinguished, but not a vestige of transverse striae. On the other two leaves there were minute spheres of only partially digested meat in the centre of much transparent fluid.
Fibrin.--Bits of fibrin were left in water during four days, whilst the following experiments were tried, but they were not in the least acted on. The fibrin which I first used was not pure, and included dark particles: it had either not been well prepared or had subsequently undergone some change. Thin portions, about 1/10 of an inch square, were placed on several leaves, and though the fibrin was soon liquefied, the whole was never dissolved. Smaller particles were then placed on four leaves, and minute drops of hydrochloric acid (one part to 437 of water) were added; this seemed to hasten the process of digestion, for on one leaf all was liquified and absorbed after 20 hrs.; but on the three other leaves some undissolved residue was left after 48 hrs. It is remarkable that in all the above and following experiments, as well as when much larger bits of fibrin were used, the leaves were very little excited; and it was sometimes necessary to add a little saliva to induce complete inflection. The leaves, moreover, began to re-expand after only 48 hrs., whereas they would have remained inflected for a much longer time had insects, meat, cartilage, albumen, &c., been placed on them.
I then tried some pure white fibrin, sent me by Dr. Burdon Sanderson.
[Experiment 1.--Two particles, barely 1/20 of an inch (1.27 mm.) square, were placed on opposite sides of the same leaf. One of these did not excite the surrounding tentacles, and the gland on which it rested soon dried. The other particle caused a few of the short adjoining tentacles to be inflected, the more distant ones not being affected. After 24 hrs. both were almost, and after 72 hrs. completely, dissolved.
Experiment 2.--The same experiment with the same result, only one of the two bits of fibrin exciting the short surrounding tentacles. This bit was so slowly acted on that after a day I pushed it on to some fresh glands. In three days from the time when it was first placed on the leaf it was completely dissolved.
Experiment 3.--Bits of fibrin of about the same size as before were placed on the discs of two leaves; these caused very little inflection in 23 hrs., but after 48 hrs. both were well clasped by the surrounding short tentacles, and after an additional 24 hrs. were completely dissolved. On the disc of one of these leaves much clear acid fluid was left.
Experiment 4.--Similar bits of fibrin were placed on the discs of two leaves; as after 2 hrs. the glands seemed rather dry, they were freely moistened with saliva; this soon caused strong inflection both of the tentacles and blades, with copious secretion from the glands. In 18 hrs. the fibrin was completely liquefied, but undigested atoms still floated in the liquid; these, however, disappeared in under two additional days.]
From these experiments it is clear that the secretion completely dissolves pure fibrin. The rate of dissolution is rather slow; but this depends merely on this substance not exciting the leaves sufficiently, so that only the immediately adjoining tentacles are inflected, and the supply of secretion is small.
Syntonin.--This substance, extracted from muscle, was kindly prepared for me by Dr. Moore. Very differently from fibrin, it acts quickly and energetically. Small portions placed on the discs of three leaves caused their tentacles and blades to be strongly inflected within 8 hrs.; but no further observations were made. It is probably due to the presence of this substance that raw meat is too powerful a stimulant, often injuring or even killing the leaves.
Areolar Tissue.--Small portions of this tissue from a sheep were placed on the discs of three leaves; these became moderately well inflected in 24 hrs., but began to re-expand after 48 hrs., and were fully re-expanded in 72 hrs., always reckoning from the time when the bits were first given. This substance, therefore, like fibrin, excites the leaves for only a short time. The residue left on the leaves, after they were fully re-expanded, was examined under a high power and found much altered, but, owing to the presence of a quantity of elastic tissue, which is never acted on, could hardly be said to be in a liquefied condition.
Some areolar tissue free from elastic tissue was next procured from the visceral cavity of a toad, and moderately sized, as well as very small, bits were placed on five leaves. After 24 hrs. two of the bits were completely liquefied; two others were rendered transparent, but not quite liquefied; whilst the fifth was but little affected. Several glands on the three latter leaves were now moistened with a little saliva, which soon caused much inflection and secretion, with the result that in the course of 12 additional hrs. one leaf alone showed a remnant of undigested tissue. On the discs of the four other leaves (to one of which a rather large bit had been given) nothing was left except some transparent viscid fluid. I may add that some of this tissue included points of black pigment, and these were not at all affected. As a control experiment, small portions of this tissue were left in water and on wet moss for the same length of time, and remained white and opaque. From these facts it is clear that areolar tissue is easily and quickly digested by the secretion; but that it does not greatly excite the leaves.
Cartilage.--Three cubes (1/20 of an inch or 1.27 mm.) of white, translucent, extremely tough cartilage were cut from the end of a slightly roasted leg-bone of a sheep. These were placed on three leaves, borne by poor, small plants in my greenhouse during November; and it seemed in the highest degree improbable that so hard a substance would be digested under such unfavourable circumstances. Nevertheless, after 48 hrs., the cubes were largely dissolved and converted into minute spheres, surrounded by transparent, very acid fluid. Two of these spheres were completely softened to their centres; whilst the third still contained a very small irregularly shaped core of solid cartilage. Their surfaces were seen under the microscope to be curiously marked by prominent ridges, showing that the cartilage had been unequally corroded by the secretion. I need hardly say that cubes of the same cartilage, kept in water for the same length of time, were not in the least affected.
During a more favourable season, moderately sized bits of the skinned ear of a cat, which includes cartilage, areolar and elastic tissue, were placed on three leaves. Some of the glands were touched with saliva, which caused prompt inflection. Two of the leaves began to re-expand after three days, and the third on the fifth day. The fluid residue left on their discs was now examined, and consisted in one case of perfectly transparent, viscid matter; in the other two cases, it contained some elastic tissue and apparently remnants of half digested areolar tissue.
Fibro-cartilage (from between the vertebrae of the tail of a sheep). Moderately sized and small bits (the latter about 1/20 of an inch) were placed on nine leaves. Some of these were well and some very little inflected. In the latter case the bits were dragged over the discs, so that they were well bedaubed with the secretion, and many glands thus irritated. All the leaves re-expanded after only two days; so that they were but little excited by this substance. The bits were not liquefied, but were certainly in an altered condition, being swollen, much more transparent, and so tender as to disintegrate very easily. My son Francis prepared some artificial gastric juice, which was proved efficient by quickly dissolving fibrin, and suspended portions of the fibro-cartilage in it. These swelled and became hyaline, exactly like those exposed to the secretion of Drosera, but were not dissolved. This result surprised me much, as two physiologists were of opinion that fibro-cartilage would be easily digested by gastric juice. I therefore asked Dr. Klein to examine the specimens; and he reports that the two which had been subjected to artificial gastric juice were "in that state of digestion in which we find connective tissue when treated with an acid, viz. swollen, more or less hyaline, the fibrillar bundles having become homogeneous and lost their fibrillar structure." In the specimens which had been left on the leaves of Drosera, until they re-expanded, "parts were altered, though only slightly so, in the same manner as those subjected to the gastric juice as they had become more transparent, almost hyaline, with the fibrillation of the bundles indistinct." Fibro-cartilage is therefore acted on in nearly the same manner by gastric juice and by the secretion of Drosera.
Bone.--Small smooth bits of the dried hyoidal bone of a fowl moistened with saliva were placed on two leaves, and a similarly moistened splinter of an extremely hard, broiled mutton-chop bone on a third leaf. These leaves soon became strongly inflected, and remained so for an unusual length of time; namely, one leaf for ten and the other two for nine days. The bits of bone were surrounded all the time by acid secretion. When examined under a weak power, they were found quite softened, so that they were readily penetrated by a blunt needle, torn into fibres, or compressed. Dr. Klein was so kind as to make sections of both bones and examine them. He informs me that both presented the normal appearance of decalcified bone, with traces of the earthy salts occasionally left. The corpuscles with their processes were very distinct in most parts; but in some parts, especially near the periphery of the hyoidal bone, none could be seen. Other parts again appeared amorphous, with even the longitudinal striation of bone not distinguishable. This amorphous structure, as Dr. Klein thinks, may be the result either of the incipient digestion of the fibrous basis or of all the animal matter having been removed, the corpuscles being thus rendered invisible. A hard, brittle, yellowish substance occupied the position of the medulla in the fragments of the hyoidal bone.
As the angles and little projections of the fibrous basis were not in the least rounded or corroded, two of the bits were placed on fresh leaves. These by the next morning were closely inflected, and remained so,--the one for six and the other for seven days,--therefore for not so long a time as on the first occasion, but for a much longer time than ever occurs with leaves inflected over inorganic or even over many organic bodies. The secretion during the whole time coloured litmus paper of a bright red; but this may have been due to the presence of the acid super-phosphate of lime. When the leaves re-expanded, the angles and projections of the fibrous basis were as sharp as ever. I therefore concluded, falsely as we shall presently see, that the secretion cannot touch the fibrous basis of bone. The more probable explanation is that the acid was all consumed in decomposing the phosphate of lime which still remained; so that none was left in a free state to act in conjunction with the ferment on the fibrous basis.
Enamel and Dentine.--As the secretion decalcified ordinary bone, I determined to try whether it would act on enamel and dentine, but did not expect that it would succeed with so hard a substance as enamel. Dr. Klein gave me some thin transverse slices of the canine tooth of a dog; small angular fragments of which were placed on four leaves; and these were examined each succeeding day at the same hour. The results are, I think, worth giving in detail.]
[Experiment 1.--May 1st, fragment placed on leaf; 3rd, tentacles but little inflected, so a little saliva was added; 6th, as the tentacles were not strongly inflected, the fragment was transferred to another leaf, which acted at first slowly, but by the 9th closely embraced it. 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 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 canaliculi of the bone-corpuscles had become larger. Otherwise the corpuscles and their canaliculi 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. 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 masses. 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 masses 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 quantity 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 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 quantity of phosphate given to the above four leaves was extremely small, much was left in every case undissolved. A larger quantity 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 circumstance 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 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 masses, 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.
prove that gelatine is far from acting energetically on Drosera.
In the last chapter it was shown that a solution of isinglass 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 isinglass were much more strongly inflected than the other four. I conclude therefore that isinglass contains some, though perhaps very little, soluble albuminous 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 isinglass. 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.
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 isinglass; but I am assured by good authorities that it is most difficult, or impossible, to know whether chondrin is pure, and if it contained any albuminous 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 albumen, I know not. Rather large drops of milk excite so much secretion (which is very acid) that it sometimes trickles down 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 scraped 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.
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 masses, 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 albuminous, with
* Dr. Lauder Brunton, 'Handbook for Phys. Lab.' p. 529.
a non-albuminous, substance; and the absorption of a very small quantity of the former would excite the leaves, and yet not decrease the casein to a perceptible degree. Schiff asserts*--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 masses 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.
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, grasses, 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 albuminoids, 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 canal 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.
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 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 albumen), 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 scraped 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 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 albuminous 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.
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.
All the substances hitherto mentioned cause prolonged inflection of the tentacles, and are either completely or at least partially dissolved by the secretion. But there are many other substances, some of them containing nitrogen, which are not in the least acted on by the secretion, and do not induce inflection for a longer time than do inorganic and insoluble objects. These unexciting and indigestible substances are, as far as I have observed, epidermic productions (such as bits of human nails, balls of hair, the quills of feathers), fibro-elastic tissue, mucin, pepsin, urea, chitine, chlorophyll, cellulose, gun-cotton, fat, oil, and starch.
To these may be added dissolved sugar and gum, diluted alcohol, and vegetable infusions not containing albumen, for none of these, as shown in the last chapter, excite inflection. Now, it is a remarkable fact, which affords additional and important evidence, that the ferment of Drosera is closely similar to or identical with pepsin, that none of these same substances are, as far as it is known, digested by the gastric juice of animals, though some of them are acted on by the other secretions of the alimentary canal. Nothing more need be said about some of the above enumerated substances, excepting that they were repeatedly tried on the leaves of Drosera, and were not in the least affected by the secretion. About the others it will be advisable to give my experiments.
[Fibro-elastic Tissue.--We have already seen that when little cubes of meat, &c., were placed on leaves, the muscles, areolar tissue, and cartilage were completely dissolved, but the fibro-elastic tissue, even the most delicate threads, were left without the least signs of having been attacked. And it is well known that this tissue cannot be digested by the gastric juice of animals.*
Mucin.--As this substance contains about 7 per cent. of nitrogen, I expected that it would have excited the leaves greatly and been digested by the secretion, but in this I was mistaken. From what is stated in chemical works, it appears extremely doubtful whether mucin can be prepared as a pure principle. That which I used (prepared by Dr. Moore) was dry and hard. Particles moistened with water were placed on four leaves, but after two days there was only a trace of inflection in the immediately adjoining tentacles. These leaves were then tried with bits of meat, and all four soon became strongly inflected. Some of the dried mucin was then soaked in water for two days, and little cubes of the proper size were placed on three leaves. After four days the tentacles
* See, for instance, Schiff, 'Phys. de la Digestion,' 1867, tom. ii., p. 38.
round the margins of the discs were a little inflected, and the secretion collected on the disc was acid, but the exterior tentacles were not affected. One leaf began to re-expand on the fourth day, and all were fully re-expanded on the sixth. The glands which had been in contact with the mucin were a little darkened. We may therefore conclude that a small amount of some impurity of a moderately exciting nature had been absorbed. That the mucin employed by me did contain some soluble matter was proved by Dr. Sanderson, who on subjecting it to artificial gastric juice found that in 1 hr. some was dissolved, but only in the proportion of 23 to 100 of fibrin during the same time. The cubes, though perhaps rather softer than those left in water for the same time, retained their angles as sharp as ever. We may therefore infer that the mucin itself was not dissolved or digested. Nor is it digested by the gastric juice of living animals, and according to Schiff* it is a layer of this substance which protects the coats of the stomach from being corroded during digestion.
Pepsin.--My experiments are hardly worth giving, as it is scarcely possible to prepare pepsin free from other albuminoids; but I was curious to ascertain, as far as that was possible, whether the ferment of the secretion of Drosera would act on the ferment of the gastric juice of animals. I first used the common pepsin sold for medicinal purposes, and afterwards some which was much purer, prepared for me by Dr. Moore. Five leaves to which a considerable quantity of the former was given remained inflected for five days; four of them then died, apparently from too great stimulation. I then tried Dr. Moore's pepsin, making it into a paste with water, and placing such small particles on the discs of five leaves that all would have been quickly dissolved had it been meat or albumen. The leaves were soon inflected; two of them began to re-expand after only 20 hrs., and the other three were almost completely re-expanded after 44 hrs. Some of the glands which had been in contact with the particles of pepsin, or with the acid secretion surrounding them, were singularly pale, whereas others were singularly dark-coloured. Some of the secretion was scraped off and examined under a high power; and it abounded with granules undistinguishable from those of pepsin left in water for the same length of time. We may therefore infer, as highly probable (remembering what small quantities were given), that the ferment of Drosera does not act on or digest
* 'Leons phys. de la Digestion,' 1867, tom. ii., p. 304.
pepsin, but absorbs from it some albuminous impurity which induces inflection, and which in large quantity is highly injurious. Dr. Lauder Brunton at my request endeavoured to ascertain whether pepsin with hydrochloric acid would digest pepsin, and as far as he could judge, it had no such power. Gastric juice, therefore, apparently agrees in this respect with the secretion of Drosera.
Urea.--It seemed to me an interesting inquiry whether this refuse of the living body, which contains much nitrogen, would, like so many other animal fluids and substances, be absorbed by the glands of Drosera and cause inflection. Half-minim drops of a solution of one part to 437 of water were placed on the discs of four leaves, each drop containing the quantity usually employed by me, namely 1/960 of a grain, or .0674 mg.; but the leaves were hardly at all affected. They were then tested with bits of meat, and soon became closely inflected. I repeated the same experiment on four leaves with some fresh urea prepared by Dr. Moore; after two days there was no inflection; I then gave them another dose, but still there was no inflection. These leaves were afterwards tested with similarly sized drops of an infusion of raw meat, and in 6 hrs. there was considerable inflection, which became excessive in 24 hrs. But the urea apparently was not quite pure, for when four leaves were immersed in 2 dr. (7.1 ml.) of the solution, so that all the glands, instead of merely those on the disc, were enabled to absorb any small amount of impurity in solution, there was considerable inflection after 24 hrs., certainly more than would have followed from a similar immersion in pure water. That the urea, which was not perfectly white, should have contained a sufficient quantity of albuminous matter, or of some salt of ammonia, to have caused the above effect, is far from surprising, for, as we shall see in the next chapter, astonishingly small doses of ammonia are highly efficient. We may therefore conclude that urea itself is not exciting or nutritious to Drosera; nor is it modified by the secretion, so as to be rendered nutritious, for, had this been the case, all the leaves with drops on their discs assuredly would have been well inflected. Dr. Lauder Brunton informs me that from experiments made at my request at St. Bartholomew's Hospital it appears that urea is not acted on by artificial gastric juice, that is by pepsin with hydrochloric acid.
Chitine.--The chitinous coats of insects naturally captured by the leaves do not appear in the least corroded. Small square pieces of the delicate wing and of the elytron of a Staphylinus were placed on some leaves, and after these had re-expanded, the pieces were carefully examined. Their angles were as sharp as ever, and they did not differ in appearance from the other wing and elytron of the same insect which had been left in water. The elytron, however, had evidently yielded some nutritious matter, for the leaf remained clasped over it for four days; whereas the leaves with bits of the true wing re-expanded on the second day. Any one who will examine the excrement of insect-eating animals will see how powerless their gastric juice is on chitine.
Cellulose.--I did not obtain this substance in a separate state, but tried angular bits of dry wood, cork, sphagnum moss, linen, and cotton thread. None of these bodies were in the least attacked by the secretion, and they caused only that moderate amount of inflection which is common to all inorganic objects. Gun-cotton, which consists of cellulose, with the hydrogen replaced by nitrogen, was tried with the same result. We have seen that a decoction of cabbage-leaves excites the most powerful inflection. I therefore placed two little square bits of the blade of a cabbage-leaf, and four little cubes cut from the midrib, on six leaves of Drosera. These became well inflected in 12 hrs., and remained so for between two and four days; the bits of cabbage being bathed all the time by acid secretion. This shows that some exciting matter, to which I shall presently refer, had been absorbed; but the angles of the squares and cubes remained as sharp as ever, proving that the framework of cellulose had not been attacked. Small square bits of spinach-leaves were tried with the same result; the glands pouring forth a moderate supply of acid secretion, and the tentacles remaining inflected for three days. We have also seen that the delicate coats of pollen grains are not dissolved by the secretion. It is well known that the gastric juice of animals does not attack cellulose.
Chlorophyll.--This substance was tried, as it contains nitrogen. Dr. Moore sent me some preserved in alcohol; it was dried, but soon deliquesced. Particles were placed on four leaves; after 3 hrs. the secretion was acid; after 8 hrs. there was a good deal of inflection, which in 24 hrs. became fairly well marked. After four days two of the leaves began to open, and the other two were then almost fully re-expanded. It is therefore clear that this chlorophyll contained matter which excited the leaves to a moderate degree; but judging by the eye, little or none was dissolved; so that in a pure state it would not probably have been attacked by the secretion. Dr. Sanderson tried that which I used, as well as some freshly prepared, with artificial digestive liquid, and found that it was not digested. Dr. Lauder Brunton likewise tried some prepared by the process given in the British Pharmacopoeia, and exposed it for five days at the temperature of 37o Cent. to digestive liquid, but it was not diminished in bulk, though the fluid acquired a slightly brown colour. It was also tried with the glycerine extract of pancreas with a negative result. Nor does chlorophyll seem affected by the intestinal secretions of various animals, judging by the colour of their excrement.
It must not be supposed from these facts that the grains of chlorophyll, as they exist in living plants, cannot be attacked by the secretion; for these grains consist of protoplasm merely coloured by chlorophyll. My son Francis placed a thin slice of spinach leaf, moistened with saliva, on a leaf of Drosera, and other slices on damp cotton-wool, all exposed to the same temperature. After 19 hrs. the slice on the leaf of Drosera was bathed in much secretion from the inflected tentacles, and was now examined under the microscope. No perfect grains of chlorophyll could be distinguished; some were shrunken, of a yellowish-green colour, and collected in the middle of the cells; others were disintegrated and formed a yellowish mass, likewise in the middle of the cells. On the other hand, in the slices surrounded by damp cotton-wool, the grains of chlorophyll were green and as perfect as ever. My son also placed some slices in artificial gastric juice, and these were acted on in nearly the same manner as by the secretion. We have seen that bits of fresh cabbage and spinach leaves cause the tentacles to be inflected and the glands to pour forth much acid secretion; and there can be little doubt that it is the protoplasm forming the grains of chlorophyll, as well as that lining the walls of the cells, which excites the leaves.
Fat and Oil.--Cubes of almost pure uncooked fat, placed on several leaves, did not have their angles in the least rounded. We have also seen that the oil-globules in milk are not digested. Nor does olive oil dropped on the discs of leaves cause any inflection; but when they are immersed in olive oil, they become strongly inflected; but to this subject I shall have to recur. Oily substances are not digested by the gastric juice of animals.
Starch.--Rather large bits of dry starch caused well-marked inflection, and the leaves did not re-expand until the fourth day; but I have no doubt that this was due to the prolonged irritation of the glands, as the starch continued to absorb the secretion. The particles were not in the least reduced in size; and we know that leaves immersed in an emulsion of starch are not at all affected. I need hardly say that starch is not digested by the gastric juice of animals.
Action of the Secretion on Living Seeds.
The results of some experiments on living seeds, selected by hazard, may here be given, though they bear only indirectly on our present subject of digestion.
Seven cabbage seeds of the previous year were placed on the same number of leaves. Some of these leaves were moderately, but the greater number only slightly inflected, and most of them re-expanded on the third day. One, however, remained clasped till the fourth, and another till the fifth day. These leaves therefore were excited somewhat more by the seeds than by inorganic objects of the same size. After they re-expanded, the seeds were placed under favourable conditions on damp sand; other seeds of the same lot being tried at the same time in the same manner, and found to germinate well. Of the seven seeds which had been exposed to the secretion, only three germinated; and one of the three seedlings soon perished, the tip of its radicle being from the first decayed, and the edges of its cotyledons of a dark brown colour; so that altogether five out of the seven seeds ultimately perished.
Radish seeds (Raphanus sativus) of the previous year were placed on three leaves, which became moderately inflected, and re-expanded on the third or fourth day. Two of these seeds were transferred to damp sand; only one germinated, and that very slowly. This seedling had an extremely short, crooked, diseased, radicle, with no absorbent hairs; and the cotyledons were oddly mottled with purple, with the edges blackened and partly withered.
Cress seeds (Lepidum sativum) of the previous year were placed on four leaves; two of these next morning were moderately and two strongly inflected, and remained so for four, five, and even six days. Soon after these seeds were placed on the leaves and had become damp, they secreted in the usual manner a layer of tenacious mucus; and to ascertain whether it was the absorption of this substance by the glands which caused so much inflection, two seeds were put into water, and as much of the mucus as possible scraped off. They were then placed on leaves, which became very strongly inflected in the course of 3 hrs., and were still closely inflected on the third day; so that it evidently was not the mucus which excited so much inflection; on the contrary, this served to a certain extent as a protection to the seeds. Two of the six seeds germinated whilst still lying on the leaves, but the seedlings, when transferred to damp sand, soon died; of the other four seeds, only one germinated.
Two seeds of mustard (Sinapis nigra), two of celery (Apium graveolens)--both of the previous year, two seeds well soaked of caraway (Carum carui), and two of wheat, did not excite the leaves more than inorganic objects often do. Five seeds, hardly ripe, of a buttercup (Ranunculus), and two fresh seeds of Anemone nemorosa, induced only a little more effect. On the other hand, four seeds, perhaps not quite ripe, of Carex sylvatica caused the leaves on which they were placed to be very strongly inflected; and these only began to re-expand on the third day, one remaining inflected for seven days.
It follows from these few facts that different kinds of seeds excite the leaves in very different degrees; whether this is solely due to the nature of their coats is not clear. In the case of the cress seeds, the partial removal of the layer of mucus hastened the inflection of the tentacles. Whenever the leaves remain inflected during several days over seeds, it is clear that they absorb some matter from them. That the secretion penetrates their coats is also evident from the large proportion of cabbage, raddish, and cress seeds which were killed, and from several of the seedlings being greatly injured. This injury to the seeds and seedlings may, however, be due solely to the acid of the secretion, and not to any process of digestion; for Mr. Traherne Moggridge has shown that very weak acids of the acetic series are highly injurious to seeds. It never occurred to me to observe whether seeds are often blown on to the viscid leaves of plants growing in a state of nature; but this can hardly fail sometimes to occur, as we shall hereafter see in the case of Pinguicula. If so, Drosera will profit to a slight degree by absorbing matter from such seeds.]
Summary and Concluding Remarks on the Digestive Power of Drosera.
When the glands on the disc are excited either by the absorption of nitrogenous matter or by mechanical irritation, their secretion increases in quantity and becomes acid. They likewise transmit some influence to the glands of the exterior tentacles, causing them to secrete more copiously; and their secretion likewise becomes acid. With animals, according to Schiff,* mechanical irritation excites the glands of the stomach to secrete an acid, but not pepsin. Now, I have every reason to believe (though the fact is not fully established), that although the glands of Drosera are continually secreting viscid fluid to replace that lost by evaporation, yet they do not secrete the ferment proper for digestion when mechanically irritated, but only after absorbing certain matter, probably of a nitrogenous nature. I infer that this is the case, as the secretion from a large number of leaves which had been irritated by particles of glass placed on their discs did not digest albumen; and more especially from the analogy of Dionaea and Nepenthes. In like manner, the glands of the stomach of animals secrete pepsin, as Schiff asserts, only after they have absorbed certain soluble substances, which he designates as peptogenes. There is, therefore, a remarkable parallelism between the glands of Drosera and those of the stomach in the secretion of their proper acid and ferment.
The secretion, as we have seen, completely dissolves albumen, muscle, fibrin, areolar tissue, cartilage, the fibrous basis of bone, gelatine, chondrin, casein in the state in which it exists in milk, and gluten which has been subjected to weak hydrochloric acid. Syntonin and legumin excite the leaves so powerfully and quickly that there can hardly be a doubt that both would be dissolved by the secretion. The secretion
* 'Phys. de la Digestion,' 1867, tom. ii. pp. 188, 245.
failed to digest fresh gluten, apparently from its injuring the glands, though some was absorbed. Raw meat, unless in very small bits, and large pieces of albumen, &c., likewise injure the leaves, which seem to suffer, like animals, from a surfeit. I know not whether the analogy is a real one, but it is worth notice that a decoction of cabbage leaves is far more exciting and probably nutritious to Drosera than an infusion made with tepid water; and boiled cabbages are far more nutritious, at least to man, than the uncooked leaves. The most striking of all the cases, though not really more remarkable than many others, is the digestion of so hard and tough a substance as cartilage. The dissolution of pure phosphate of lime, of bone, dentine, and especially enamel, seems wonderful; but it depends merely on the long-continued secretion of an acid; and this is secreted for a longer time under these circumstances than under any others. It was interesting to observe that as long as the acid was consumed in dissolving the phosphate of lime, no true digestion occurred; but that as soon as the bone was completely decalcified, the fibrous basis was attacked and liquefied with the greatest ease. The twelve substances above enumerated, which are completely dissolved by the secretion, are likewise dissolved by the gastric juice of the higher animals; and they are acted on in the same manner, as shown by the rounding of the angles of albumen, and more especially by the manner in which the transverse striae of the fibres of muscle disappear.
The secretion of Drosera and gastric juice were both able to dissolve some element or impurity out of the globulin and haematin employed by me. The secretion also dissolved something out of chemically prepared casein, which is said to consist of two substances; and although Schiff asserts that casein in this state is not attacked by gastric juice, he might easily have overlooked a minute quantity of some albuminous matter, which Drosera would detect and absorb. Again, fibro-cartilage, though not properly dissolved, is acted on in the same manner, both by the secretion of Drosera and gastric juice. But this substance, as well as the so-called haematin used by me, ought perhaps to have been classed with indigestible substances.
That gastric juice acts by means of its ferment, pepsin, solely in the presence of an acid, is well established; and we have excellent evidence that a ferment is present in the secretion of Drosera, which likewise acts only in the presence of an acid; for we have seen that when the secretion is neutralised by minute drops of the solution of an alkali, the digestion of albumen is completely stopped, and that on the addition of a minute dose of hydrochloric acid it immediately recommences.
The nine following substances, or classes of substances, namely, epidermic productions, fibro-elastic tissue, mucin, pepsin, urea, chitine, cellulose, gun-cotton, chlorophyll, starch, fat and oil, are not acted on by the secretion of Drosera; nor are they, as far as is known, by the gastric juice of animals. Some soluble matter, however, was extracted from the mucin, pepsin, and chlorophyll, used by me, both by the secretion and by artificial gastric juice.
The several substances, which are completely dissolved by the secretion, and which are afterwards absorbed by the glands, affect the leaves rather differently. They induce inflection at very different rates and in very different degrees; and the tentacles remain inflected for very different periods of time. Quick inflection depends partly on the quantity of the substance given, so that many glands are simultaneously affected, partly on the facility with which it is penetrated and liquefied by the secretion, partly on its nature, but chiefly on the presence of exciting matter already in solution. Thus saliva, or a weak solution of raw meat, acts much more quickly than even a strong solution of gelatine. So again leaves which have re-expanded, after absorbing drops of a solution of pure gelatine or isinglass (the latter being the more powerful of the two), if given bits of meat, are inflected much more energetically and quickly than they were before, notwithstanding that some rest is generally requisite between two acts of inflection. We probably see the influence of texture in gelatine and globulin when softened by having been soaked in water acting more quickly than when merely wetted. It may be partly due to changed texture, and partly to changed chemical nature, that albumen, which had been kept for some time, and gluten which had been subjected to weak hydrochloric acid, act more quickly than these substances in their fresh state.
The length of time during which the tentacles remain inflected largely depends on the quantity of the substance given, partly on the facility with which it is penetrated or acted on by the secretion, and partly on its essential nature. The tentacles always remain inflected much longer over large bits or large drops than over small bits or drops. Texture probably plays a part in determining the extraordinary length of time during which the tentacles remain inflected over the hard grains of chemically prepared casein. But the tentacles remain inflected for an equally long time over finely powdered, precipitated phosphate of lime; phosphorus in this latter case evidently being the attraction, and animal matter in the case of casein. The leaves remain long inflected over insects, but it is doubtful how far this is due to the protection afforded by their chitinous integuments; for animal matter is soon extracted from insects (probably by exosmose from their bodies into the dense surrounding secretion), as shown by the prompt inflection of the leaves. We see the influence of the nature of different substances in bits of meat, albumen, and fresh gluten acting very differently from equal-sized bits of gelatine, areolar tissue, and the fibrous basis of bone. The former cause not only far more prompt and energetic, but more prolonged, inflection than do the latter. Hence we are, I think, justified in believing that gelatine, areolar tissue, and the fibrous basis of bone, would be far less nutritious to Drosera than such substances as insects, meat, albumen, &c. This is an interesting conclusion, as it is known that gelatine affords but little nutriment to animals; and so, probably, would areolar tissue and the fibrous basis of bone. The chondrin which I used acted more powerfully than gelatine, but then I do not know that it was pure. It is a more remarkable fact that fibrin, which belongs to the great class of Proteids,* including albumen in one of its sub-groups, does not excite the tentacles in a greater degree, or keep them inflected for a longer time, than does gelatine, or
* See the classification adopted by Dr. Michael Foster in Watts' 'Dictionary of Chemistry,' Supplement 1872, page 969.
areolar tissue, or the fibrous basis of bone. It is not known how long an animal would survive if fed on fibrin alone, but Dr. Sanderson has no doubt longer than on gelatine, and it would be hardly rash to predict, judging from the effects on Drosera, that albumen would be found more nutritious than fibrin. Globulin likewise belongs to the Proteids, forming another sub-group, and this substance, though containing some matter which excited Drosera rather strongly, was hardly attacked by the secretion, and was very little or very slowly attacked by gastric juice. How far globulin would be nutritious to animals is not known. We thus see how differently the above specified several digestible substances act on Drosera; and we may infer, as highly probable, that they would in like manner be nutritious in very different degrees both to Drosera and to animals.
The glands of Drosera absorb matter from living seeds, which are injured or killed by the secretion. They likewise absorb matter from pollen, and from fresh leaves; and this is notoriously the case with the stomachs of vegetable-feeding animals. Drosera is properly an insectivorous plant; but as pollen cannot fail to be often blown on to the glands, as will occasionally the seeds and leaves of surrounding plants, Drosera is, to a certain extent, a vegetable-feeder.
Finally, the experiments recorded in this chapter show us that there is a remarkable accordance in the power of digestion between the gastric juice of animals with its pepsin and hydrochloric acid and the secretion of Drosera with its ferment and acid belonging to the acetic series. We can, therefore, hardly doubt that the ferment in both cases is closely similar, if not identically the same. That a plant and an animal should pour forth the same, or nearly the same, complex secretion, adapted for the same purpose of digestion, is a new and wonderful fact in physiology. But I shall have to recur to this subject in the fifteenth chapter, in my concluding remarks on the Droseraceae.