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- Insectivorous Plants - 60/80 -


BYBLIS.

Byblis gigantea (Western Australia).--A dried specimen, about 18 inches in height, with a strong stem, was sent me from Kew. The leaves are some inches in length, linear, slightly flattened, with a small projecting rib on the lower surface. They are covered on all sides by glands of two kinds [page 344] --sessile ones arranged in rows, and others supported on moderately long pedicels. Towards the narrow summits of the leaves the pedicels are longer than elsewhere, and here equal the diameter of the leaf. The glands are purplish, much flattened, and formed of a single layer of radiating cells, which in the larger glands are from forty to fifty in number. The pedicels consist of single elongated cells, with colourless, extremely delicate walls, marked with the finest intersecting spiral lines. Whether these lines are the result of contraction from the drying of the walls, I do not know, but the whole pedicel was often spirally rolled up. These glandular hairs are far more simple in structure than the so-called tentacles of the preceding genera, and they do not differ essentially from those borne by innumerable other plants. The flower-peduncles bear similar glands. The most singular character about the leaves is that the apex is enlarged into a little knob, covered with glands, and about a third broader than the adjoining part of the attenuated leaf. In two places dead flies adhered to the glands. As no instance is known of unicellular structures having any power of movement,* Byblis, no doubt, catches insects solely by the aid of its viscid secretion. These probably sink down besmeared with the secretion and rest on the small sessile glands, which, if we may judge by the analogy of Drosophyllum, then pour forth their secretion and afterwards absorb the digested matter.

Supplementary Observations on the Power of Absorption by the Glandular Hairs of other Plants.--A few observations on this subject may be here conveniently introduced. As the glands of many, probably of all,

* Sachs, 'Trait de Bot.,' 3rd edit. 1874, p. 1026. [page 345]

the species of Droseraceae absorb fluids or at least allow them readily to enter,* it seemed desirable to ascertain how far the glands of other plants which are not specially adapted for capturing insects, had the same power. Plants were chosen for trial at hazard, with the exception of two species of saxifrage, which were selected from belonging to a family allied to the Droseraceae. Most of the experiments were made by immersing the glands either in an infusion of raw meat or more commonly in a solution of carbonate of ammonia, as this latter substance acts so powerfully and rapidly on protoplasm. It seemed also particularly desirable to ascertain whether ammonia was absorbed, as a small amount is contained in rain-water. With the Droseraceae the secretion of a viscid fluid by the glands does not prevent their absorbing; so that the glands of other plants might excrete superfluous matter, or secrete an odoriferous fluid as a protection against the attacks of insects, or for any other purpose, and yet have the power of absorbing. I regret that in the following cases I did not try whether the secretion could digest or render soluble animal substances, but such experiments would have been difficult on account of the small size of the glands and the small amount of secretion. We shall see in the next chapter that the secretion from the glandular hairs of Pinguicula certainly dissolves animal matter.

[Saxifraga umbrosa.--The flower-peduncles and petioles of the leaves are clothed with short hairs, bearing pink-coloured glands, formed of several polygonal cells, with their pedicels divided by partitions into distinct cells, which are generally colourless, but sometimes pink. The glands secrete a yellowish viscid fluid, by

*The distinction between true absorption and mere permeation, or imbibition, is by no means clearly understood: see Mller's 'Physiology,' Eng. translat. 1838, vol. i. p. 280. [page 346]

which minute Diptera are sometimes, though not often, caught.* The cells of the glands contain bright pink fluid, charged with granules or with globular masses of pinkish pulpy matter. This matter must be protoplasm, for it is seen to undergo slow but incessant changes of form if a gland be placed in a drop of water and examined. Similar movements were observed after glands had been immersed in water for 1, 3, 5, 18, and 27 hrs. Even after this latter period the glands retained their bright pink colour; and the protoplasm within their cells did not appear to have become more aggregated. The continually changing forms of the little masses of protoplasm are not due to the absorption of water, as they were seen in glands kept dry.

A flower-stem, still attached to a plant, was bent (May 29) so as to remain immersed for 23 hrs. 30 m. in a strong infusion of raw meat. The colour of the contents of the glands was slightly changed, being now of a duller and more purple tint than before. The contents also appeared more aggregated, for the spaces between the little masses of protoplasm were wider; but this latter result did not follow in some other and similar experiments. The masses seemed to change their forms more rapidly than did those in water; so that the cells had a different appearance every four or five minutes. Elongated masses became in the course of one or two minutes spherical; and spherical ones drew themselves out and united with others. Minute masses rapidly increased in size, and three distinct ones were seen to unite. The movements were, in short, exactly like those described in the case of Drosera. The cells of the pedicels were not affected by the infusion; nor were they in the following experiment.

Another flower-stem was placed in the same manner and for the same length of time in a solution of one part of nitrate of ammonia to 146 of water (or 3 grs. to 1 oz.), and the glands were discoloured in exactly the same manner as by the infusion of raw meat.

Another flower-stem was immersed, as before, in a solution of one part of carbonate of ammonia to 109 of water. The glands, after 1 hr. 30 m., were not discoloured, but after 3 hrs. 45 m. most of them had become dull purple, some of them blackish-

*In the case of Saxifraga tridactylites, Mr. Druce says ('Pharmaceutical Journal, ' May 1875) that he examined some dozens of plants, and in almost every instance remnants of insects adhered to the leaves. So it is, as I hear from a friend, with this plant in Ireland. [page 347]

green, a few being still unaffected. The little masses of protoplasm within the cells were seen in movement. The cells of the pedicels were unaltered. The experiment was repeated, and a fresh flower-stem was left for 23 hrs. in the solution, and now a great effect was produced; all the glands were much blackened, and the previously transparent fluid in the cells of the pedicels, even down to their bases, contained spherical masses of granular matter. By comparing many different hairs, it was evident that the glands first absorb the carbonate, and that the effect thus produced travels down the hairs from cell to cell. The first change which could be observed is a cloudy appearance in the fluid, due to the formation of very fine granules, which afterwards aggregate into larger masses. Altogether, in the darkening of the glands, and in the process of aggregation travelling down the cells of the pedicels, there is the closest resemblance to what takes place when a tentacle of Drosera is immersed in a weak solution of the same salt. The glands, however, absorb very much more slowly than those of Drosera. Besides the glandular hairs, there are star-shaped organs which do not appear to secrete, and which were not in the least affected by the above solutions.

Although in the case of uninjured flower-stems and leaves the carbonate seems to be absorbed only by the glands, yet it enters a cut surface much more quickly than a gland. Strips of the rind of a flower-stem were torn off, and the cells of the pedicels were seen to contain only colourless transparent fluid; those of the glands including as usual some granular matter. These strips were then immersed in the same solution as before (one part of the carbonate to 109 of water), and in a few minutes granular matter appeared in the lowercells of all the pedicels. The action invariably commenced (for I tried the experiment repeatedly) in the lowest cells, and therefore close to the torn surface, and then gradually travelled up the hairs until it reached the glands, in a reversed direction to what occurs in uninjured specimens. The glands then became discoloured, and the previously contained granular matter was aggregated into larger masses. Two short bits of a flower-stem were also left for 2 hrs. 40 m. in a weaker solution of one part of the carbonate to 218 of water; and in both specimens the pedicels of the hairs near the cut ends now contained much granular matter; and the glands were completely discoloured.

Lastly, bits of meat were placed on some glands; these were examined after 23 hrs., as were others, which had apparently not long before caught minute flies; but they did not present any [page 348] difference from the glands of other hairs. Perhaps there may not have been time enough for absorption. I think so as some glands, on which dead flies had evidently long lain, were of a pale dirty purple colour or even almost colourless, and the granular matter within them presented an unusual and somewhat peculiar appearance. That these glands had absorbed animal matter from the flies, probably by exosmose into the viscid secretion, we may infer, not only from their changed colour, but because, when placed in a solution of carbonate of ammonia, some of the cells in their pedicels become filled with granular matter; whereas the cells of other hairs, which had not caught flies, after being treated with the same solution for the same length of time, contained only a small quantity of granular matter. But more evidence is necessary before we fully admit that the glands of this saxifrage can absorb, even with ample time allowed, animal matter from the minute insects which they occasionally and accidentally capture.

Saxifraga rotundifolia (?).--The hairs on the flower-stems of this species are longer than those just described, and bear pale brown glands. Many were examined, and the cells of the pedicels were quite transparent. A bent stem was immersed for 30 m. in a solution of one part of carbonate of ammonia to 109 of water, and two or three of the uppermost cells in the pedicels now contained granular or aggregated matter; the glands having become of a bright yellowish-green. The glands of this species therefore absorb the carbonate much more quickly than do those of Saxifraga umbrosa, and the upper cells of the pedicels are likewise affected much more quickly. Pieces of the stem were cut off and immersed in the same solution; and now the process of aggregation travelled up the hairs in a reversed direction; the cells close to the cut surfaces being first affected.

Primula sinensis.--The flower-stems, the upper and lower surfaces of the leaves and their footstalks, are all clothed with a multitude of longer and shorter hairs. The pedicels of the longer hairs are divided by transverse partitions into eight or nine cells. The enlarged terminal cell is globular, forming a gland which secretes a variable amount of thick, slightly viscid, not acid, brownish-yellow matter.

A piece of a young flower-stem was first immersed in distilled water for 2 hrs. 30 m., and the glandular hairs were not at all affected. Another piece, bearing twenty-five short and nine long hairs, was carefully examined. The glands of the latter contained no solid or semi-solid matter; and those of only two [page 349] of the twenty-five short hairs contained some globules. This piece was then immersed for 2 hrs. in a solution of one part of carbonate of ammonia to 109 of water, and now the glands of the twenty-five shorter hairs, with two or three exceptions, contained either one large or from two to five smaller


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