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gathered by chance a dozen plants, bearing fifty-six fully expanded leaves, and on thirty-one of these dead insects or remnants of them adhered; and, no doubt, many more would have been caught afterwards by these same leaves, and still more by those as yet not expanded. On one plant all six leaves had caught their prey; and on several plants very many leaves had caught more than a single insect. On one large leaf I found the remains of thirteen distinct insects. Flies (Diptera) are captured much oftener than other insects. The largest kind which I have seen caught was a small butterfly (Canonympha pamphilus); but the Rev. H. M. Wilkinson informs me that he found a large living dragon-fly with its body firmly held by two leaves. As this plant is extremely common in some districts, the number of insects thus annually slaughtered must be prodigious. Many plants cause the death of insects, for instance the sticky buds of the horse-chestnut (Esculus hippocastanum), without thereby receiving, as far as we can perceive, any advantage; but it was soon evident that Drosera was

which was published in the Gardener's Chronicle,' 1863, p. 30. Mr. Scott shows that gentle irritation of the hairs, as well as insects placed on the disc of the leaf, cause the hairs to bend inwards. Mr. A. W. Bennett also gave another interesting account of the movements of the leaves before the British Association for 1873. In this same year Dr. Warming published an essay, in which he describes the structure of the so-called hairs, entitled, "Sur la Différence entre les Trichomes," &c., extracted from the proceedings of the Soc. d'Hist. Nat. de Copenhague. I shall also have occasion hereafter to refer

to a paper by Mrs. Treat, of New Jersey, on some American species of Drosera. Dr. Burdon Sanderson delivered a lecture on Dionæa, before the Royal Institution (published in 'Nature,' June 14, 1874), in which a short account of my observations on the power of true digestion possessed by Drosera and Dionæa first appeared. Prof. Asa Gray has done good service by calling attention to Drosera, and to other plants having similar habits, in 'The Nation' (1874, pp. 261 and 232), and in other publications. Dr. Hooker, also, in his important address on Carnivorous Plants (Brit. Assoc., Belfast, 1874), has given a history of the subject

excellently adapted for the special purpose of catching insects, so that the subject seemed well worthy of investigation.

The results have proved highly remarkable; the more important ones being--firstly, the extraordinary

FIG. 1.*

(Drosera rotundifolia.)

Leaf viewed from above; enlarged four times.

sensitiveness of the glands to slight pressure and to minute doses of certain nitrogenous fluids, as shown by the movements of the so-called hairs or tentacles;

* The drawings of Drosera and Dionæa, given in this work, were made for me by my son George Darwin; those of Aldrovanda, and of the several species of Utri

cularia, by my son Francis. They have been excellently reproduced on wood by Mr. Cooper, 188 Strai d.

secondly, the power possessed by the leaves of rendering soluble or digesting nitrogenous substances, and of afterwards absorbing them; thirdly, the changes which take place within the cells of the tentacles, when the glands are excited in various ways.

It is necessary, in the first place, to describe briefly the plant. It bears from two or three to five or six leaves, generally extended more or less horizontally, but sometimes standing vertically upwards. The shape and general appearance of a leaf is shown, as seen from above, in fig. 1, and as seen laterally, in fig. 2. The leaves are commonly a little broader than long,

FIG. 2.

(Drosera rotundifolia.)

Old leaf viewed laterally; enlarged about five times.

but this was not the case in the one here figured. The whole upper surface is covered with gland-bearing filaments, or tentacles, as I shall call them, from their manner of acting. The glands were counted on thirtyone leaves, but many of these were of unusually large size, and the average number was 192; the greatest number being 260, and the least 130. The glands are each surrounded by large drops of extremely viscid secretion, which, glittering in the sun, have given rise to the plant's poetical name of the sun-dew.

The tentacles on the central part of the leaf or disc are short and stand upright, and their pedicels are green. Towards the margin they become longer and longer and more incline

outwards, with their pedicels of a purple colour. Those on the extreme margin project in the same plane with the leaf, or more commonly (see fig. 2) are considerably reflexed. A few tentacles spring from the base of the footstalk or petiole, and these are the longest of all, being sometimes nearly of an inch in length. On a leaf bearing altogether 252 tentacles, the short ones on the disc, having green pedicels, were in number to the longer submarginal and marginal tentacles, having purple pedicels, as nine to sixteen.

A tentacle consists of a thin, straight, hair-like pedicel, carrying a gland on the summit. The pedicel is somewhat flattened, and is formed of several rows of elongated cells, filled with purple fluid or granular matter.* There is, however, a narrow zone close beneath the glands of the longer tentacles, and a broader zone near their bases, of a green tint. Spiral vessels, accompanied by simple vascular tissue, branch off from the vascular bundles in the blade of the leaf, and run up all the tentacles into the glands.

Several eminent physiologists have discussed the homological nature of these appendages or tentacles, that is, whether they ought to be considered as hairs (trichomes) or prolongations of the leaf. Nitschke has shown that they include all the elements proper to the blade of a leaf; and the fact of their including vascular tissue was formerly thought to prove that they were prolongations of the leaf, but it is now known that vessels sometimes enter true hairs.† The power of movement which they possess is a strong argument against their being viewed as hairs. The conclusion which seems to me the most probable will be given in Chap. XV., namely that they existed primordially as glandular hairs, or mere epidermic formations, and that their upper part should still be so considered; but that their lower

*According to Nitschke ('Bot. Zeitung,' 1861, p. 224) the purple fluid results from the metamorphosis of chlorophyll. Mr. Sorby examined the colouring matter with the spectroscope, and informs me that it consists of the commonest species of erythrophyll, "which is often met with in leaves with low vitality, and in parts, like the petioles, which carry on leaf-functions in a very imperfect manner. All that can be said, therefore, is that the hairs

(or tentacles) are coloured like parts of a leaf which do not fulfil their proper office."

+ Dr. Nitschke has discussed this subject in 'Bot. Zeitung,' 1861, p. 241, &c. See also Dr. Warming (Sur la Différence entre les Trichomes,' &c., 1873), who gives references to various publications. See also Groenland and Trécul, Annal. des Sc. nat. bot.' (4th series), tom. iii. 1855, pp. 297 and 303.

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part, which alone is capable of movement, consists of a prolongation of the leaf; the spiral vessels being extended from this to the uppermost part. We shall hereafter see that the terminal tentacles of the divided leaves of Roridula are still in an intermediate condition.

The glands, with the exception of those borne by the extreme

[graphic][merged small][merged small]

Longitudinal section of a gland; greatly magnified. From Dr. Warming.

marginal tentacles, are oval, and of nearly uniform size, viz. about of an inch in length. Their structure is remarkable, and their functions complex, for they secrete, absorb, and are acted on by various stimulants. They consist of an outer layer of small polygonal cells, containing purple granular matter or fluid, and with the walls thicker than those of the pedicels.

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