CHAPTER VI

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.¹⁸ 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 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 40 °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 115 °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 18 °Cent. 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.

[Experiment 1.

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;¹⁹ 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.