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elsewhere, and somewhat more rapid in a longitudinal than in a transverse direction across the disc. plants exhibit still more plainly their inferiority to animals in the absence of any reflex action, except in so far as the glands of Drosera, when excited from a distance, send back some influence which causes the contents of the cells to become aggregated down to the bases of the tentacles. But the greatest inferiority of all is the absence of a central organ, able to receive impressions from all points, to transmit their effects in any definite direction, to store them up and reproduce them.



Pinguicula vulgaris - Structure of leaves - Number of insects and other objects caught - Movement of the margins of the leaves — Uses of this movement - Secretion, digestion, and absorptionAction of the secretion on various animal and vegetable substances -The effects of substances not containing soluble nitrogenous matter on the glands - Pinguicula grandiflora — Pinguicula lusitanica, catches insects-Movement of the leaves, secretion and digestion.

PINGUICULA VULGARIS.-This plant grows in moist places, generally on mountains. It bears on an average eight, rather thick, oblong, light green leaves, having scarcely any footstalk. A full-sized leaf is about 1 inch in length and inch in breadth. The young central leaves are deeply concave, and project upwards; the older ones towards the outside are flat or convex, and lie close to the ground, forming a rosette from 3 to 4 inches in diameter. The margins of the leaves are incurved. Their upper surfaces are thickly covered with two sets of glandular hairs, differing in the size of the glands and in the length of their pedicels. The larger glands have a circular outline as seen from above, and are of moderate thickness; they are divided by radiating partitions into sixteen cells, containing light-green, homogeneous fluid. They are supported on elongated, unicellular pedicels (containing a nucleus with a nucleolus) which rest on slight prominences. The small glands differ only in being formed of about half the number of cells, containing much paler fluid, and supported on much shorter pedicels. Near the midrib, towards the base of the leaf, the

pedicels are multicellular, are longer than elsewhere, and bear smaller glands. All the glands secrete a colourless fluid, which is so viscid that I have seen a fine thread drawn out to a length of 18 inches; but the fluid in this case was secreted by a gland which had been excited. The edge of the leaf is translucent, and does not bear any glands; and here the spiral vessels, proceeding from the midrib, terminate in cells marked by a spiral line, somewhat like those within the glands of Drosera.

The roots are short. Three plants were dug up in North Wales on June 20, and carefully washed; each bore five or six unbranched roots, the longest of which was only 1-2 of an inch. Two rather young plants were examined on September 28; these had a greater number of roots, namely eight and eighteen, all under 1 inch in length, and very little branched.

I was led to investigate the habits of this plant by being told by Mr. W. Marshall that on the mountains of Cumberland many insects adhere to the leaves.

A friend sent me on June 23 thirty-nine leaves from North Wales, which were selected owing to objects of some kind adhering to them. Of these leaves, thirty-two had caught 142 insects, or on an average 44 per leaf, minute fragments of insects not being included. Besides the insects, small leaves belonging to four different kinds of plants, those of Erica tetralix being much the commonest, and three minute seedling plants, blown by the wind, adhered to nineteen of the leaves. One had caught as many as ten leaves of the Erica. Seeds or fruits, commonly of Carex and one of Juncus, besides bits of moss and other rubbish, likewise adhered to six of the thirty-nine leaves. The same friend, on June 27, collected nine plants bearing seventy-four leaves, and all of these, with the exception of three young leaves, had caught insects; thirty insects were counted on one leaf, eighteen on a second, and sixteen on a third. Another friend examined on August 22 some plants in Donegal, Leland, and found insects on 70 out of 157 leaves; fifteen of

these leaves were sent me, each having caught on an average 2.4 insects. To nine of them, leaves (mostly of Erica tet alix) adhered; but they had been specially selected on this latter account. I may add that early in August my son found leaves of this same Erica and the fruits of a Carex on the leaves of a Pinguicula in Switzerland, probably Pinguicula alpina ; some insects, but no great number, also adhered to the leaves of this plant, which had much better developed roots than those of Pinguicula vulgaris. In Cumberland, Mr. Marshall, on September 3, carefully examined for me ten plants bearing eighty leaves; and on sixtythree of these (i.e. on 79 per cent.) he found insects, 143 in number; so that each leaf had on an average 2-27 insects. A few days later he sent me some plants with sixteen seeds or fruits adhering to fourteen leaves. There was a seed on three leaves on the same plant. The sixteen seeds belonged to nine different kinds, which could not be recognised, excepting one of Ranunculus, and several belonging to three or four distinct species of Carex. It appears that fewer insects are caught late in the year than earlier; thus in Cumberland from twenty to twenty-four insects were observed in the middle of July on several leaves, whereas in the beginning of September the average number was only 2.27. Most of the insects, in all the foregoing cases, were Diptera, but with many minute Hymenoptera, including some ants, a few small Coleoptera, larvæ, spiders, and even small moths.

We thus see that numerous insects and other objects are caught by the viscid leaves; but we have no right to infer from this fact that the habit is beneficial to the plant, any more than in the before given case of the Mirabilis, or of the horse-chestnut. But it will presently be seen that dead insects and other nitrogenous bodies excite the glands to increased secretion; and that the secretion then becomes acid and has the power of digesting animal substances, such as albumen, fibrin, &c. Moreover, the dissolved nitrogenous matter is absorbed by the glands, as shown by their limpid contents being aggregated into slowly moving granular masses of protoplasm. The same results follow when insects are naturally captured, and as the plant lives in poor soil and has small roots, there can be no

doubt that it profits by its power of digesting and absorbing matter from the prey which it habitually captures in such large numbers. It will, however, be convenient first to describe the movements of the leaves.

Movements of the Leaves.--That such thick, large leaves as those of Pinguicula vulgaris should have the power of curving inwards when excited has never even been suspected. It is necessary to select for experiment leaves with their glands secreting freely, and which have been prevented from capturing many insects; as old leaves, at least those growing in a state of nature, have their margins already curled so much inwards that they exhibit little power of movement, or move very slowly. I will first give in detail the more important experiments which were tried, and then make some concluding remarks.

Experiment 1.-A young and almost upright leaf was selected, with its two lateral edges equally and very slightly incurved. A row of small flies was placed along one margin. When looked at next day, after 15 hrs., this margin, but not the other, was found folded inwards, like the helix of the human ear, to the breadth of of an inch, so as to lie partly over the row of flies (fig. 15). The glands on which the flies rested, as well as those on the overlapping margin which had been brought into contact with the flies, were all secreting copiously.

FIG. 15.

Experiment 2.-A row of flies was placed on one margin of a rather old leaf, which lay flat on the ground; and in this case the margin, after the same interval as before, namely 15 hrs., had only just begun to curl inwards; but so much secretion had been poured forth that the spoonshaped tip of the leaf was filled with it. row of small flies. Experiment 3.-Fragments of a large fly were placed close to the apex of a vigorous leaf, as well as along half one margin.

(Pinguicula vulgaris.) Outline of leaf with left

margin inflected over a

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