DROSOPHYLLUM--RORIDULA--BYBLIS--GLANDULAR HAIRS OF OTHER PLANTS--CONCLUDING REMARKS ON THE DROSERACEAE.
Drosophyllum--Structure of leaves--Nature of the secretion--Manner of catching insects-- Power of absorption--Digestion of animal substances--Summary on Drosophyllum--Roridula--Byblis--Glandular hairs of other plants, their power of absorption--Saxifraga--Primula--Pelargonium--Erica--Mirabilis--Nicotiana--Summary on glandular hairs--Concluding remarks on the Droseraceae.
DROSOPHYLLUM LUSITANICUM.--This rare plant has been found only in Portugal, and, as I hear from Dr. Hooker, in Morocco. I obtained living specimens through the great kindness of Mr. W.C. Tait, and afterwards from Mr. G. Maw and Dr. Moore. Mr. Tait informs me that it grows plentifully on the sides of dry hills near Oporto, and that vast numbers of flies adhere to the leaves. This latter fact is well-known to the villagers, who call the plant the "fly-catcher, " and hang it up in their cottages for this purpose. A plant in my hot-house caught so many insects during the early part of April, although the weather was cold and insects scarce, that it must have been in some manner strongly attractive to them. On four leaves of a young and small plant, 8, 10, 14, and 16 minute insects, chiefly Diptera, were found in the autumn adhering to them. I neglected to examine the roots, but I hear from Dr. Hooker that they are very small, as in the case of the previously mentioned members of the same family of the Droseraceae.
The leaves arise from an almost woody axis; they are linear, much attenuated towards their tips, and several inches in length. The upper surface is concave, the lower convex, with a narrow channel down the middle. Both surfaces, with the exception of the channel, are covered with glands, supported on pedicels and arranged in irregular longitudinal rows. These organs I shall call tentacles, from their close resemblance to those of Drosera, though they have no power of movement. Those on the same leaf differ much in length. The glands also differ in size, and are of a bright pink or of a purple colour; their upper surfaces are convex, and the lower flat or even concave, so that they resemble miniature mushrooms in appearance. They are formed of two (as I believe) layers of delicate angular cells, enclosing eight or ten larger cells with thicker, zigzag walls. Within these larger cells there are others marked by spiral lines, and apparently connected with the spiral vessels which run up the green multi-cellular pedicels. The glands secrete large drops of viscid secretion. Other glands, having the same general appearance, are found on the flower-peduncles and calyx.
FIG. 14. (Drosophyllum lusitanicum.) Part of leaf, enlarged seven times, showing lower surface.
Besides the glands which are borne on longer or shorter pedicels, there are numerous ones, both on the upper and lower surfaces of the leaves, so small as to be scarcely visible to the naked eye. They are colourless and almost sessile, either circular or oval in outline; the latter occurring chiefly on the backs of the leaves (fig. 14). Internally they have exactly the same structure as the larger glands which are supported on pedicels; and indeed the two sets almost graduate into one another. But the sessile glands differ in one important respect, for they never secrete spontaneously, as far as I have seen, though I have examined them under a high power on a hot day, whilst the glands on pedicels were secreting copiously. Nevertheless, if little bits of damp albumen or fibrin are placed on these sessile glands, they begin after a time to secrete, in the same manner as do the glands of Dionaea when similarly treated. When they were merely rubbed with a bit of raw meat, I believe that they likewise secreted. Both the sessile glands and the taller ones on pedicels have the power of rapidly absorbing nitrogenous matter.
The secretion from the taller glands differs in a remarkable manner from that of Drosera, in being acid before the glands have been in any way excited; and judging from the changed colour of litmus paper, more strongly acid than that of Drosera. This fact was observed repeatedly; on one occasion I chose a young leaf, which was not secreting freely, and had never caught an insect, yet the secretion on all the glands coloured litmus paper of a bright red. From the quickness with which the glands are able to obtain animal matter from such substances as well-washed fibrin and cartilage, I suspect that a small quantity of the proper ferment must be present in the secretion before the glands are excited, so that a little animal matter is quickly dissolved.
Owing to the nature of the secretion or to the shape of the glands, the drops are removed from them with singular facility. It is even somewhat difficult, by the aid of a finely pointed polished needle, slightly damped with water, to place a minute particle of any kind on one of the drops; for on withdrawing the needle, the drop is generally withdrawn; whereas with Drosera there is no such difficulty, though the drops are occasionally withdrawn. From this peculiarity, when a small insect alights on a leaf of Drosophyllum, the drops adhere to its wings, feet, or body, and are drawn from the gland; the insect then crawls onward and other drops adhere to it; so that at last, bathed by the viscid secretion, it sinks down and dies, resting on the small sessile glands with which the surface of the leaf is thickly covered. In the case of Drosera, an insect sticking to one or more of the exterior glands is carried by their movement to the centre of the leaf; with Drosophyllum, this is effected by the crawling of the insect, as from its wings being clogged by the secretion it cannot fly away.
There is another difference in function between the glands of these two plants: we know that the glands of Drosera secrete more copiously when properly excited. But when minute particles of carbonate of ammonia, drops of a solution of this salt or of the nitrate of ammonia, saliva, small insects, bits of raw or roast meat, albumen, fibrin or cartilage, as well as inorganic particles, were placed on the glands of Drosophyllum, the amount of secretion never appeared to be in the least increased. As insects do not commonly adhere to the taller glands, but withdraw the secretion, we can see that there would be little use in their having acquired the habit of secreting copiously when stimulated; whereas with Drosera this is of use, and the habit has been acquired. Nevertheless, the glands of Drosophyllum, without being stimulated, continually secrete, so as to replace the loss by evaporation. Thus when a plant was placed under a small bell-glass with its inner surface and support thoroughly wetted, there was no loss by evaporation, and so much secretion was accumulated in the course of a day that it ran down the tentacles and covered large spaces of the leaves.
The glands to which the above named nitrogenous substances and liquids were given did not, as just stated, secrete more copiously; on the contrary, they absorbed their own drops of secretion with surprising quickness. Bits of damp fibrin were placed on five glands, and when they were looked at after an interval of 1 hr. 12 m., the fibrin was almost dry, the secretion having been all absorbed. So it was with three cubes of albumen after 1 hr. 19 m., and with four other cubes, though these latter were not looked at until 2 hrs. 15 m. had elapsed. The same result followed in between 1 hr. 15 m. and 1 hr. 30 m. when particles both of cartilage and meat were placed on several glands. Lastly, a minute drop (about 1/20 of a minim) of a solution of one part of nitrate of ammonia to 146 of water was distributed between the secretion surrounding three glands, so that the amount of fluid surrounding each was slightly increased; yet when looked at after 2 hrs., all three were dry. On the other hand, seven particles of glass and three of coal-cinders, of nearly the same size as those of the above named organic substances, were placed on ten glands; some of them being observed for 18 hrs., and others for two or three days; but there was not the least sign of the secretion being absorbed. Hence, in the former cases, the absorption of the secretion must have been due to the presence of some nitrogenous matter, which was either already soluble or was rendered so by the secretion. As the fibrin was pure, and had been well washed in distilled water after being kept in glycerine, and as the cartilage had been soaked in water, I suspect that these substances must have been slightly acted on and rendered soluble within the above stated short periods.
The glands have not only the power of rapid absorption, but likewise of secreting again quickly; and this latter habit has perhaps been gained, inasmuch as insects, if they touch the glands, generally withdraw the drops of secretion, which have to be restored. The exact period of re-secretion was recorded in only a few cases. The glands on which bits of meat were placed, and which were nearly dry after about 1 hr. 30 m., when looked at after 22 additional hours, were found secreting; so it was after 24 hrs. with one gland on which a bit of albumen had been placed. The three glands to which a minute drop of a solution of nitrate of ammonia was distributed, and which became dry after 2 hrs., were beginning to re-secrete after only 12 additional hours.
Tentacles Incapable of Movement.--Many of the tall tentacles, with insects adhering to them, were carefully observed; and fragments of insects, bits of raw meat, albumen, &c., drops of a solution of two salts of ammonia and of saliva, were placed on the glands of many tentacles; but not a trace of movement could ever be detected. I also repeatedly irritated the glands with a needle, and scratched and pricked the blades, but neither the blade nor the tentacles became at all inflected. We may therefore conclude that they are incapable of movement.
On the Power of Absorption possessed by the Glands.--It has already been indirectly shown that the glands on pedicels absorb animal matter; and this is further shown by their changed colour, and by the aggregation of their contents, after they have been left in contact with nitrogenous substances or liquids. The following observations apply both to the glands supported on pedicels and to the minute sessile ones. Before a gland has been in any way stimulated, the exterior cells commonly contain only limpid purple fluid; the more central ones including mulberry-like masses of purple granular matter. A leaf was placed in a little solution of one part of carbonate of ammonia to 146 of water (3 grs. to 1 oz.), and the glands were instantly darkened and very soon became black; this change being due to the strongly marked aggregation of their contents, more especially of the inner cells. Another leaf was placed in a solution of the same strength of nitrate of ammonia, and the glands were slightly darkened in 25 m., more so in 50 m., and after 1 hr. 30 m. were of so dark a red as to appear almost black. Other leaves were placed in a weak infusion of raw meat and in human saliva, and the glands were much darkened in 25 m., and after 40 m. were so dark as almost to deserve to be called black. Even immersion for a whole day in distilled water occasionally induces some aggregation within the glands, so that they become of a darker tint. In all these cases the glands are affected in exactly the same manner as those of Drosera. Milk, however, which acts so energetically on Drosera, seems rather less effective on Drosophyllum, for the glands were only slightly darkened by an immersion of 1 hr. 20 m., but became decidedly darker after 3 hrs. Leaves which had been left for 7 hrs. in an infusion of raw meat or in saliva were placed in the solution of carbonate of ammonia, and the glands now became greenish; whereas, if they had been first placed in the carbonate, they would have become black. In this latter case, the ammonia probably combines with the acid of the secretion, and therefore does not act on the colouring matter; but when the glands are first subjected to an organic fluid, either the acid is consumed in the work of digestion or the cell-walls are rendered more permeable, so that the undecomposed carbonate enters and acts on the colouring matter. If a particle of the dry carbonate is placed on a gland, the purple colour is quickly discharged, owing probably to an excess of the salt. The gland, moreover, is killed.
Turning now to the action of organic substances, the glands on which bits of raw meat were placed became dark-coloured; and in 18 hrs. their contents were conspicuously aggregated. Several glands with bits of albumen and fibrin were darkened in between 2 hrs. and 3 hrs.; but in one case the purple colour was completely discharged. Some glands which had caught flies were compared with others close by; and though they did not differ much in colour, there was a marked difference in their state of aggregation. In some few instances, however, there was no such difference, and this appeared to be due to the insects having been caught long ago, so that the glands had recovered their pristine state. In one case, a group of the sessile colourless glands, to which a small fly adhered, presented a peculiar appearance; for they had become purple, owing to purple granular matter coating the cell-walls. I may here mention as a caution that, soon after some of my plants arrived in the spring from Portugal, the glands were not plainly acted on by bits of meat, or insects, or a solution of ammonia--a circumstance for which I cannot account.
Digestion of Solid Animal Matter.--Whilst I was trying to place on two of the taller glands little cubes of albumen, these slipped down, and, besmeared with secretion, were left resting on some of the small sessile glands. After 24 hrs. one of these cubes was found completely liquefied, but with a few white streaks still visible; the other was much rounded, but not quite dissolved. Two other cubes were left on tall glands for 2 hrs. 45 m., by which time all the secretion was absorbed; but they were not perceptibly acted on, though no doubt some slight amount of animal matter had been absorbed from them. They were then placed on the small sessile glands, which being thus stimulated secreted copiously in the course of 7 hrs. One of these cubes was much liquefied within this short time; and both were completely liquefied after 21 hrs. 15 m.; the little liquid masses, however, still showing some white streaks. These streaks disappeared after an additional period of 6 hrs. 30 m.; and by next morning (i.e. 48 hrs. from the time when the cubes were first placed on the glands) the liquefied matter was wholly absorbed. A cube of albumen was left on another tall gland, which first absorbed the secretion and after 24 hrs. poured forth a fresh supply. This cube, now surrounded by secretion, was left on the gland for an additional 24 hrs., but was very little, if at all, acted on. We may, therefore, conclude, either that the secretion from the tall glands has little power of digestion, though strongly acid, or that the amount poured forth from a single gland is insufficient to dissolve a particle of albumen which within the same time would have been dissolved by the secretion from several of the small sessile glands. Owing to the death of my last plant, I was unable to ascertain which of these alternatives is the true one.
Four minute shreds of pure fibrin were placed, each resting on one, two, or three of the taller glands. In the course of 2 hrs. 30 m. the secretion was all absorbed, and the shreds were left almost dry. They were then pushed on to the sessile glands. One shred, after 2 hrs. 30 m., seemed quite dissolved, but this may have been a mistake. A second, when examined after 17 hrs. 25 m., was liquefied, but the liquid as seen under the microscope still contained floating granules of fibrin. The other two shreds were completely liquefied after 21 hrs. 30 m.; but in one of the drops a very few granules could still be detected. These, however, were dissolved after an additional interval of 6 hrs. 30 m.; and the surface of the leaf for some distance all round was covered with limpid fluid. It thus appears that Drosophyllum digests albumen and fibrin rather more quickly than Drosera can; and this may perhaps be attributed to the acid, together probably with some small amount of the ferment, being present in the secretion, before the glands have been stimulated; so that digestion begins at once.
Concluding Remarks.--The linear leaves of Drosophyllum differ but slightly from those of certain species of Drosera; the chief differences being, firstly, the presence of minute, almost sessile, glands, which, like those of Dionaea, do not secrete until they are excited by the absorption of nitrogenous matter. But glands of this kind are present on the leaves of Drosera binata, and appear to be represented by the papillae on the leaves of Drosera rotundifolia. Secondly, the presence of tentacles on the backs of the leaves; but we have seen that a few tentacles, irregularly placed and tending towards abortion, are retained on the backs of the leaves of Drosera binata. There are greater differences in function between the two genera. The most important one is that the tentacles of Drosophyllum have no power of movement; this loss being partially replaced by the drops of viscid secretion being readily withdrawn from the glands; so that, when an insect comes into contact with a drop, it is able to crawl away, but soon touches other drops, and then, smothered by the secretion, sinks down on the sessile glands and dies. Another difference is, that the secretion from the tall glands, before they have been in any way excited, is strongly acid, and perhaps contains a small quantity of the proper ferment. Again, these glands do not secrete more copiously from being excited by the absorption of nitrogenous matter; on the contrary, they then absorb their own secretion with extraordinary quickness. In a short time they begin to secrete again. All these circumstances are probably connected with the fact that insects do not commonly adhere to the glands with which they first come into contact, though this does sometimes occur; and that it is chiefly the secretion from the sessile glands which dissolves animal matter out of their bodies.
Roridula dentata.--This plant, a native of the western parts of the Cape of Good Hope, was sent to me in a dried state from Kew. It has an almost woody stem and branches, and apparently grows to a height of some feet. The leaves are linear, with their summits much attenuated. Their upper and lower surfaces are concave, with a ridge in the middle, and both are covered with tentacles, which differ greatly in length; some being very long, especially those on the tips of the leaves, and some very short. The glands also differ much in size and are somewhat elongated. They are supported on multicellular pedicels.
This plant, therefore, agrees in several respects with Drosophyllum, but differs in the following points. I could detect no sessile glands; nor would these have been of any use, as the upper surface of the leaves is thickly clothed with pointed, unicellular hairs directed upwards. The pedicels of the tentacles do not include spiral vessels; nor are there any spiral cells within the glands. The leaves often arise in tufts and are pinnatifid, the divisions projecting at right angles to the main linear blade. These lateral divisions are often very short and bear only a single terminal tentacle, with one or two short ones on the sides. No distinct line of demarcation can be drawn between the pedicels of the long terminal tentacles and the much attenuated summits of the leaves. We may, indeed, arbitrarily fix on the point to which the spiral vessels proceeding from the blade extend; but there is no other distinction.
It was evident from the many particles of dirt sticking to the glands that they secrete much viscid matter. A large number of insects of many kinds also adhered to the leaves. I could nowhere discover any signs of the tentacles having been inflected over the captured insects; and this probably would have been seen even in the dried specimens, had they possessed the power of movement. Hence, in this negative character, Roridula resembles its northern representative, Drosophyllum.
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--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.
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.
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.
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 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 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 spherical masses of semi-solid matter. Three of the glands of the nine long hairs likewise included similar masses. In a few hairs there were also globules in the cells immediately beneath the glands. Looking to all thirty-four hairs, there could be no doubt that the glands had absorbed some of the carbonate. Another piece was left for only 1 hr. in the same solution, and aggregated matter appeared in all the glands. My son Francis examined some glands of the longer hairs, which contained little masses of matter, before they were immersed in any solution; and these masses slowly changed their forms, so that no doubt they consisted of protoplasm. He then irrigated these hairs for 1 hr. 15 m., whilst under the microscope, with a solution of one part of the carbonate to 218 of water; the glands were not perceptibly affected, nor could this have been expected, as their contents were already aggregated. But in the cells of the pedicels numerous, almost colourless, spheres of matter appeared, which changed their forms and slowly coalesced; the appearance of the cells being thus totally changed at successive intervals of time.
The glands on a young flower-stem, after having been left for 2 hrs. 45 m. in a strong solution of one part of the carbonate to 109 of water, contained an abundance of aggregated masses, but whether generated by the action of the salt, I do not know. This piece was again placed in the solution, so that it was immersed altogether for 6 hrs. 15 m., and now there was a great change; for almost all the spherical masses within the gland-cells had disappeared, being replaced by granular matter of a darker brown. The experiment was thrice repeated with nearly the same result. On one occasion the piece was left immersed for 8 hrs. 30 m., and though almost all the spherical masses were changed into the brown granular matter, a few still remained. If the spherical masses of aggregated matter had been originally produced merely by some chemical or physical action, it seems strange that a somewhat longer immersion in the same solution should so completely alter their character. But as the masses which slowly and spontaneously changed their forms must have consisted of living protoplasm, there is nothing surprising in its being injured or killed, and its appearance wholly changed by long immersion in so strong a solution of the carbonate as that employed. A solution of this strength paralyses all movement in Drosera, but does not kill the protoplasm; a still stronger solution prevents the protoplasm from aggregating into the ordinary full-sized globular masses, and these, though they do not disintegrate, become granular and opaque. In nearly the same manner, too hot water and certain solutions (for instance, of the salts of soda and potash) cause at first an imperfect kind of aggregation in the cells of Drosera; the little masses afterwards breaking up into granular or pulpy brown matter. All the foregoing experiments were made on flower-stems, but a piece of a leaf was immersed for 30 m. in a strong solution of the carbonate (one part to 109 of water), and little globular masses of matter appeared in all the glands, which before contained only limpid fluid.
I made also several experiments on the action of the vapour of the carbonate on the glands; but will give only a few cases. The cut end of the footstalk of a young leaf was protected with sealing-wax, and was then placed under a small bell-glass, with a large pinch of the carbonate. After 10 m. the glands showed a considerable degree of aggregation, and the protoplasm lining the cells of the pedicels was a little separated from the walls. Another leaf was left for 50 m. with the same result, excepting that the hairs became throughout their whole length of a brownish colour. In a third leaf, which was exposed for 1 hr. 50 m., there was much aggregated matter in the glands; and some of the masses showed signs of breaking up into brown granular matter. This leaf was again placed in the vapour, so that it was exposed altogether for 5 hrs. 30 m.; and now, though I examined a large number of glands, aggregated masses were found in only two or three; in all the others, the masses, which before had been globular, were converted into brown, opaque, granular matter. We thus see that exposure to the vapour for a considerable time produces the same effects as long immersion in a strong solution. In both cases there could hardly be a doubt that the salt had been absorbed chiefly or exclusively by the glands.
On another occasion bits of damp fibrin, drops of a weak infusion of raw meat and of water, were left for 24 hrs. on some leaves; the hairs were then examined, but to my surprise differed in no respect from others which had not been touched by these fluids. Most of the cells, however, included hyaline, motionless little spheres, which did not seem to consist of protoplasm, but, I suppose, of some balsam or essential oil.
Pelargonium zonale (var. edged with white).--The leaves are clothed with numerous multicellular hairs; some simply pointed; others bearing glandular heads, and differing much in length. The glands on a piece of leaf were examined and found to contain only limpid fluid; most of the water was removed from beneath the covering glass, and a minute drop of one part of carbonate of ammonia to 146 of water was added; so that an extremely small dose was given. After an interval of only 3 m. there were signs of aggregation within the glands of the shorter hairs; and after 5 m. many small globules of a pale brown tint appeared in all of them; similar globules, but larger, being found in the large glands of the longer hairs. After the specimen had been left for 1 hr. in the solution, many of the smaller globules had changed their positions; and two or three vacuoles or small spheres (for I know not which they were) of a rather darker tint appeared within some of the larger globules. Little globules could now be seen in some of the uppermost cells of the pedicels, and the protoplasmic lining was slightly separated from the walls of the lower cells. After 2 hrs. 30 m. from the time of first immersion, the large globules within the glands of the longer hairs were converted into masses of darker brown granular matter. Hence from what we have seen with Primula sinensis, there can be little doubt that these masses originally consisted of living protoplasm.
A drop of a weak infusion of raw meat was placed on a leaf, and after 2 hrs. 30 m. many spheres could be seen within the glands. These spheres, when looked at again after 30 m., had slightly changed their positions and forms, and one had separated into two; but the changes were not quite like those which the protoplasm of Drosera undergoes. These hairs, moreover, had not been examined before immersion, and there were similar spheres in some glands which had not been touched by the infusion.
Erica tetralix.--A few long glandular hairs project from the margins of the upper surfaces of the leaves. The pedicels are formed of several rows of cells, and support rather large globular heads, secreting viscid matter, by which minute insects are occasionally, though rarely, caught. Some leaves were left for 23 hrs. in a weak infusion of raw meat and in water, and the hairs were then compared, but they differed very little or not at all. In both cases the contents of the cells seemed rather more granular than they were before; but the granules did not exhibit any movement. Other leaves were left for 23 hrs. in a solution of one part of carbonate of ammonia to 218 of water, and here again the granular matter appeared to have increased in amount; but one such mass retained exactly the same form as before after an interval of 5 hrs., so that it could hardly have consisted of living protoplasm. These glands seem to have very little or no power of absorption, certainly much less than those of the foregoing plants.
Mirabilis longiflora.--The stems and both surfaces of the leaves bear viscid hairs. young plants, from 12 to 18 inches in height in my greenhouse, caught so many minute Diptera, Coleoptera, and larvae, that they were quite dusted with them. The hairs are short, of unequal lengths, formed of a single row of cells, surmounted by an enlarged cell which secretes viscid matter. These terminal cells or glands contain granules and often globules of granular matter. Within a gland which had caught a small insect, one such mass was observed to undergo incessant changes of form, with the occasional appearance of vacuoles. But I do not believe that this protoplasm had been generated by matter absorbed from the dead insect; for, on comparing several glands which had and had not caught insects, not a shade of difference could be perceived between them, and they all contained fine granular matter. A piece of leaf was immersed for 24 hrs. in a solution of one part of carbonate of ammonia to 218 of water, but the hairs seemed very little affected by it, excepting that perhaps the glands were rendered rather more opaque. In the leaf itself, however, the grains of chlorophyll near the cut surfaces had run together, or become aggregated. Nor were the glands on another leaf, after an immersion for 24 hrs. in an infusion of raw meat, in the least affected; but the protoplasm lining the cells of the pedicels had shrunk greatly from the walls. This latter effect may have been due to exosmose, as the infusion was strong. We may, therefore, conclude that the glands of this plant either have no power of absorption or that the protoplasm which they contain is not acted on by a solution of carbonate of ammonia (and this seems scarcely credible) or by an infusion of meat.
Nicotiana tabacum.--This plant is covered with innumerable hairs of unequal lengths, which catch many minute insects. The pedicels of the hairs are divided by transverse partitions, and the secreting glands are formed of many cells, containing greenish matter with little globules of some substance. Leaves were left in an infusion of raw meat and in water for 26 hrs., but presented no difference. Some of these same leaves were then left for above 2 hrs. in a solution of carbonate of ammonia, but no effect was produced. I regret that other experiments were not tried with more care, as M. Schloesing has shown* that tobacco plants supplied with the vapour of carbonate of ammonia yield on analysis a greater amount of nitrogen than other plants not thus treated; and, from what we have seen, it is probable that some of the vapour may be absorbed by the glandular hairs.]
Summary of the Observations on Glandular Hairs.--From the foregoing observations, few as they are, we see that the glands of two species of Saxifraga, of a Primula and Pelargonium, have the power of rapid absorption; whereas the glands of an Erica, Mirabilis, and Nicotiana, either have no such power, or the contents of the cells are not affected by the fluids employed, namely a solution of carbonate of ammonia and an infusion of raw meat. As the glands of the Mirabilis contain protoplasm, which did not become aggregated from exposure to the fluids just named, though the contents of the cells in the blade of the leaf were greatly affected by carbonate of ammonia, we may infer that they cannot absorb. We may further infer that the innumerable insects caught by this plant are of no more service to it than are those which adhere to the deciduous and sticky scales of the leaf-buds of the horse-chestnut.
The most interesting case for us is that of the two species of Saxifraga, as this genus is distantly allied to Drosera. Their glands absorb matter from an infusion of raw meat, from solutions of the nitrate and carbonate of ammonia, and apparently from decayed insects. This was shown by the changed dull purple colour of the protoplasm within the cells of the glands, by its state of aggregation, and apparently by its more rapid spontaneous movements.
* 'Comptes rendus,' June 15, 1874. A good abstract of this paper is given in the 'Gardener's Chronicle,' July 11, 1874.
The aggregating process spreads from the glands down the pedicels of the hairs; and we may assume that any matter which is absorbed ultimately reaches the tissues of the plant. On the other hand, the process travels up the hairs whenever a surface is cut and exposed to a solution of the carbonate of ammonia.
The glands on the flower-stalks and leaves of Primula sinensis quickly absorb a solution of the carbonate of ammonia, and the protoplasm which they contain becomes aggregated. The process was seen in some cases to travel from the glands into the upper cells of the pedicels. Exposure for 10 m. to the vapour of this salt likewise induced aggregation. When leaves were left from 6 hrs. to 7 hrs. in a strong solution, or were long exposed to the vapour, the little masses of protoplasm became disintegrated, brown, and granular, and were apparently killed. An infusion of raw meat produced no effect on the glands.
The limpid contents of the glands of Pelargonium zonale became cloudy and granular in from 3 m. to 5 m. when they were immersed in a weak solution of the carbonate of ammonia; and in the course of 1 hr. granules appeared in the upper cells of the pedicels. As the aggregated masses slowly changed their forms, and as they suffered disintegration when left for a considerable time in a strong solution, there can be little doubt that they consisted of protoplasm. It is doubtful whether an infusion of raw meat produced any effect.
The glandular hairs of ordinary plants have generally been considered by physiologists to serve only as secreting or excreting organs, but we now know that they have the power, at least in some cases, of absorbing both a solution and the vapour of ammonia. As rain-water contains a small percentage of ammonia, and the atmosphere a minute quantity of the carbonate, this power can hardly fail to be beneficial. Nor can the benefit be quite so insignificant as it might at first be thought, for a moderately fine plant of Primula sinensis bears the astonishing number of above two millions and a half of glandular hairs,* all of which are able to absorb ammonia brought to them by the rain. It is moreover probable that the glands of some of the above named plants obtain animal matter from the insects which are occasionally entangled by the viscid secretion.
CONCLUDING REMARKS ON THE DROSERACEAE.
The six known genera composing this family have now been described in relation to our present subject, as far as my means have permitted. They all capture insects. This is effected by Drosophyllum, Roridula, and Byblis, solely by the viscid fluid secreted from their glands; by Drosera, through the same means, together with the movements of the tentacles; by Dionaea and Aldrovanda, through the closing of the blades of the leaf. In these two last genera rapid
* My son Francis counted the hairs on a space measured by means of a micrometer, and found that there were 35,336 on a square inch of the upper surface of a leaf, and 30,035 on the lower surface; that is, in about the proportion of 100 on the upper to 85 on the lower surface. On a square inch of both surfaces there were 65,371 hairs. A moderately fine plant bearing twelve leaves (the larger ones being a little more than 2 inches in diameter) was now selected, and the area of all the leaves, together with their foot-stalks (the flower-stems not being included), was found by a planimeter to be 39.285 square inches; so that the area of both surfaces was 78.57 square inches. Thus the plant (excluding the flower-stems) must have borne the astonishing number of 2,568,099 glandular hairs. The hairs were counted late in the autumn, and by the following spring (May) the leaves of some other plants of the same lot were found to be from one-third to one-fourth broader and longer than they were before; so that no doubt the glandular hairs had increased in number, and probably now much exceeded three millions.
movement makes up for the loss of viscid secretion. In every case it is some part of the leaf which moves. In Aldrovanda it appears to be the basal parts alone which contract and carry with them the broad, thin margins of the lobes. In Dionaea the whole lobe, with the exception of the marginal prolongations or spikes, curves inwards, though the chief seat of movement is near the midrib. In Drosera the chief seat is in the lower part of the tentacles, which, homologically, may be considered as prolongations of the leaf; but the whole blade often curls inwards, converting the leaf into a temporary stomach.
There can hardly be a doubt that all the plants belonging to these six genera have the power of dissolving animal matter by the aid of their secretion, which contains an acid, together with a ferment almost identical in nature with pepsin; and that they afterwards absorb the matter thus digested. This is certainly the case with Drosera, Drosophyllum, and Dionaea; almost certainly with Aldrovanda; and, from analogy, very probable with Roridula and Byblis. We can thus understand how it is that the three first-named genera are provided with such small roots, and that Aldrovanda is quite rootless; about the roots of the two other genera nothing is known. It is, no doubt, a surprising fact that a whole group of plants (and, as we shall presently see, some other plants not allied to the Droseraceae) should subsist partly by digesting animal matter, and partly by decomposing carbonic acid, instead of exclusively by this latter means, together with the absorption of matter from the soil by the aid of roots. We have, however, an equally anomalous case in the animal kingdom; the rhizocephalous crustaceans do not feed like other animals by their mouths, for they are destitute of an alimentary canal; but they live by absorbing through root-like processes the juices of the animals on which they are parasitic.*
Of the six genera, Drosera has been incomparably the most successful in the battle for life; and a large part of its success may be attributed to its manner of catching insects. It is a dominant form, for it is believed to include about 100 species, which range in the Old World from the Arctic regions to Southern India, to the Cape of Good Hope, Madagascar, and Australia; and in the New World from Canada to Tierra del Fuego. In this respect it presents a marked contrast with the five other genera, which appear to be failing groups. Dionaea includes only a single species, which is confined to one district in Carolina. The three varieties or closely allied species of Aldrovanda, like so many water-plants, have a wide range from Central Europe to Bengal and Australia. Drosophyllum includes only one species, limited to Portugal and Morocco. Roridula and Byblis each have (as I
* Fritz Mller, 'Facts for Darwin, ' Eng. trans. 1869, p. 139. The rhizocephalous crustaceans are allied to the cirripedes. It is hardly possible to imagine a greater difference than that between an animal with prehensile limbs, a well-constructed mouth and alimentary canal, and one destitute of all these organs and feeding by absorption through branching root-like processes. If one rare cirripede, the Anelasma squalicola, had become extinct, it would have been very difficult to conjecture how so enormous a change could have been gradually effected. But, as Fritz Mller remarks, we have in Anelasma an animal in an almost exactly intermediate condition, for it has root-like processes embedded in the skin of the shark on which it is parasitic, and its prehensile cirri and mouth (as described in my monograph on the Lepadidae, 'Ray Soc.' 1851, p. 169) are in a most feeble and almost rudimentary condition. Dr. R. Kossmann has given a very interesting discussion on this subject in his 'Suctoria and Lepadidae,' 1873. See also, Dr. Dohrn, 'Der Ursprung der Wirbelthiere,' 1875, p. 77.
Bentham and Hooker, 'Genera Plantarum.' Australia is the metropolis of the genus, forty-one species having been described from this country, as Prof. Oliver informs me.
hear from Prof. Oliver) two species; the former confined to the western parts of the Cape of Good Hope, and the latter to Australia. It is a strange fact that Dionaea, which is one of the most beautifully adapted plants in the vegetable kingdom, should apparently be on the high-road to extinction. This is all the more strange as the organs of Dionaea are more highly differentiated than those of Drosera; its filaments serve exclusively as organs of touch, the lobes for capturing insects, and the glands, when excited, for secretion as well as for absorption; whereas with Drosera the glands serve all these purposes, and secrete without being excited.
By comparing the structure of the leaves, their degree of complication, and their rudimentary parts in the six genera, we are led to infer that their common parent form partook of the characters of Drosophyllum, Roridula, and Byblis. The leaves of this ancient form were almost certainly linear, perhaps divided, and bore on their upper and lower surfaces glands which had the power of secreting and absorbing. Some of these glands were mounted on pedicels, and others were almost sessile; the latter secreting only when stimulated by the absorption of nitrogenous matter. In Byblis the glands consist of a single layer of cells, supported on a unicellular pedicel; in Roridula they have a more complex structure, and are supported on pedicels formed of several rows of cells; in Drosophyllum they further include spiral cells, and the pedicels include a bundle of spiral vessels. But in these three genera these organs do not possess any power of movement, and there is no reason to doubt that they are of the nature of hairs or trichomes. Although in innumerable instances foliar organs move when excited, no case is known of a trichome having such power.* We are thus led to inquire how the so-called tentacles of Drosera, which are manifestly of the same general nature as the glandular hairs of the above three genera, could have acquired the power of moving. Many botanists maintain that these tentacles consist of prolongations of the leaf, because they include vascular tissue, but this can no longer be considered as a trustworthy distinction. The possession of the power of movement on excitement would have been safer evidence. But when we consider the vast number of the tentacles on both surfaces of the leaves of Drosophyllum, and on the upper surface of the leaves of Drosera, it seems scarcely possible that each tentacle could have aboriginally existed as a prolongation of the leaf. Roridula, perhaps, shows us how we may reconcile these difficulties with respect to the homological nature of the tentacles. The lateral divisions of the leaves of this plant terminate in long tentacles; and these include spiral vessels which extend for only a short distance up them, with no line of demarcation between what is plainly the prolongation of the leaf and the pedicel of a glandular hair. Therefore there would be nothing anomalous or unusual in the basal parts of these tentacles, which correspond with the marginal ones of Drosera, acquiring the power of movement; and we know that in Drosera it is only the lower part which becomes inflected. But in order to understand how in this latter genus not only the marginal but all the inner tentacles have become capable of movement, we must further assume, either that through the principle of correlated development this
* Sachs, 'Trait de Botanique' 3rd edit. 1874, p. 1026.
Dr. Warming 'Sur la Diffrence entres les Trichomes,' Copenhague, 1873, p. 6. 'Extrait des Videnskabelige Meddelelser de la Soc. d'Hist. nat. de Copenhague,' Nos. 10-12, 1872.
power was transferred to the basal parts of the hairs, or that the surface of the leaf has been prolonged upwards at numerous points, so as to unite with the hairs, thus forming the bases of the inner tentacles.
The above named three genera, namely Drosophyllum, Roridula, and Byblis, which appear to have retained a primordial condition, still bear glandular hairs on both surfaces of their leaves; but those on the lower surface have since disappeared in the more highly developed genera, with the partial exception of one species, Drosera binata. The small sessile glands have also disappeared in some of the genera, being replaced in Roridula by hairs, and in most species of Drosera by absorbent papillae. Drosera binata, with its linear and bifurcating leaves, is in an intermediate condition. It still bears some sessile glands on both surfaces of the leaves, and on the lower surface a few irregularly placed tentacles, which are incapable of movement. A further slight change would convert the linear leaves of this latter species into the oblong leaves of Drosera anglica, and these might easily pass into orbicular ones with footstalks, like those of Drosera rotundifolia. The footstalks of this latter species bear multicellular hairs, which we have good reason to believe represent aborted tentacles.
The parent form of Dionaea and Aldrovanda seems to have been closely allied to Drosera, and to have had rounded leaves, supported on distinct footstalks, and furnished with tentacles all round the circumference, with other tentacles and sessile glands on the upper surface. I think so because the marginal spikes of Dionaea apparently represent the extreme marginal tentacles of Drosera, the six (sometimes eight) sensitive filaments on the upper surface, as well as the more numerous ones in Aldrovanda, representing the central tentacles of Drosera, with their glands aborted, but their sensitiveness retained. Under this point of view we should bear in mind that the summits of the tentacles of Drosera, close beneath the glands, are sensitive.
The three most remarkable characters possessed by the several members of the Droseraceae consist in the leaves of some having the power of movement when excited, in their glands secreting a fluid which digests animal matter, and in their absorption of the digested matter. Can any light be thrown on the steps by which these remarkable powers were gradually acquired?
As the walls of the cells are necessarily permeable to fluids, in order to allow the glands to secrete, it is not surprising that they should readily allow fluids to pass inwards; and this inward passage would deserve to be called an act of absorption, if the fluids combined with the contents of the glands. Judging from the evidence above given, the secreting glands of many other plants can absorb salts of ammonia, of which they must receive small quantities from the rain. This is the case with two species of Saxifraga, and the glands of one of them apparently absorb matter from captured insects, and certainly from an infusion of raw meat. There is, therefore, nothing anomalous in the Droseraceae having acquired the power of absorption in a much more highly developed degree.
It is a far more remarkable problem how the members of this family, and Pinguicula, and, as Dr. Hooker has recently shown, Nepenthes, could all have acquired the power of secreting a fluid which dissolves or digests animal matter. The six genera of the Droseraceae very probably inherited this power from a common progenitor, but this cannot apply to Pinguicula or Nepenthes, for these plants are not at all closely related to the Droceraceae. But the difficulty is not nearly so great as it at first appears. Firstly, the juices of many plants contain an acid, and, apparently, any acid serves for digestion. Secondly, as Dr. Hooker has remarked in relation to the present subject in his address at Belfast (1874), and as Sachs repeatedly insists,* the embryos of some plants secrete a fluid which dissolves albuminous substances out of the endosperm; although the endosperm is not actually united with, only in contact with, the embryo. All plants, moreover, have the power of dissolving albuminous or proteid substances, such as protoplasm, chlorophyll, gluten, aleurone, and of carrying them from one part to other parts of their tissues. This must be effected by a solvent, probably consisting of a ferment together with an acid. Now, in the case of plants which are able to absorb already soluble matter from captured insects, though not capable of true digestion, the solvent just referred to, which must be occasionally present in the glands, would be apt to exude from the glands together with the viscid secretion, inasmuch as endosmose is accompanied by exosmose. If such exudation did ever occur, the solvent would act on the animal matter contained within the captured insects, and this would be an act of true digestion. As it cannot be doubted that this process would be of high service to plants
* 'Trait de Botanique' 3rd edit. 1874, p. 844. See also for following facts pp. 64, 76, 828, 831.
Since this sentence was written, I have received a paper by Gorup-Besanez ('Berichte der Deutschen Chem. Gesellschaft,' Berlin, 1874, p. 1478), who, with the aid of Dr. H. Will, has actually made the discovery that the seeds of the vetch contain a ferment, which, when extracted by glycerine, dissolves albuminous substances, such as fibrin, and converts them into true peptones.
growing in very poor soil, it would tend to be perfected through natural selection. Therefore, any ordinary plant having viscid glands, which occasionally caught insects, might thus be converted under favourable circumstances into a species capable of true digestion. It ceases, therefore, to be any great mystery how several genera of plants, in no way closely related together, have independently acquired this same power.
As there exist several plants the glands of which cannot, as far as is known, digest animal matter, yet can absorb salts of ammonia and animal fluids, it is probable that this latter power forms the first stage towards that of digestion. It might, however, happen, under certain conditions, that a plant, after having acquired the power of digestion, should degenerate into one capable only of absorbing animal matter in solution, or in a state of decay, or the final products of decay, namely the salts of ammonia. It would appear that this has actually occurred to a partial extent with the leaves of Aldrovanda; the outer parts of which possess absorbent organs, but no glands fitted for the secretion of any digestive fluid, these being confined to the inner parts.
Little light can be thrown on the gradual acquirement of the third remarkable character possessed by the more highly developed genera of the Droseraceae, namely the power of movement when excited. It should, however, be borne in mind that leaves and their homologues, as well as flower-peduncles, have gained this power, in innumerable instances, independently of inheritance from any common parent form; for instance, in tendril-bearers and leaf-climbers (i.e. plants with their leaves, petioles and flower-peduncles, &c., modified for prehension) belonging to a large number of the most widely distinct orders,--in the leaves of the many plants which go to sleep at night, or move when shaken,--and in the irritable stamens and pistils of not a few species. We may therefore infer that the power of movement can be by some means readily acquired. Such movements imply irritability or sensitiveness, but, as Cohn has remarked,* the tissues of the plants thus endowed do not differ in any uniform manner from those of ordinary plants; it is therefore probable that all leaves are to a slight degree irritable. Even if an insect alights on a leaf, a slight molecular change is probably transmitted to some distance across its tissue, with the sole difference that no perceptible effect is produced. We have some evidence in favour of this belief, for we know that a single touch on the glands of Drosera does not excite inflection; yet it must produce some effect, for if the glands have been immersed in a solution of camphor, inflection follows within a shorter time than would have followed from the effects of camphor alone. So again with Dionaea, the blades in their ordinary state may be roughly touched without their closing; yet some effect must be thus caused and transmitted across the whole leaf, for if the glands have recently absorbed animal matter, even a delicate touch causes them to close instantly. On the whole we may conclude that the acquirement of a high degree of sensitiveness and of the power of movement by certain genera of the Droseraceae presents no greater difficulty than that presented by the similar but feebler powers of a multitude of other plants.
* See the abstract of his memoir on the contractile tissues of plants, in the 'Annals and Mag. of Nat. Hist.' 3rd series, vol. xi. p. 188.)
The specialised nature of the sensitiveness possessed by Drosera and Dionaea, and by certain other plants, well deserves attention. A gland of Drosera may be forcibly hit once, twice, or even thrice, without any effect being produced, whilst the continued pressure of an extremely minute particle excites movement. On the other hand, a particle many times heavier may be gently laid on one of the filaments of Dionaea with no effect; but if touched only once by the slow movement of a delicate hair, the lobes close; and this difference in the nature of the sensitiveness of these two plants stands in manifest adaptation to their manner of capturing insects. So does the fact, that when the central glands of Drosera absorb nitrogenous matter, they transmit a motor impulse to the exterior tentacles much more quickly than when they are mechanically irritated; whilst with Dionaea the absorption of nitrogenous matter causes the lobes to press together with extreme slowness, whilst a touch excites rapid movement. Somewhat analogous cases may be observed, as I have shown in another work, with the tendrils of various plants; some being most excited by contact with fine fibres, others by contact with bristles, others with a flat or a creviced surface. The sensitive organs of Drosera and Dionaea are also specialised, so as not to be uselessly affected by the weight or impact of drops of rain, or by blasts of air. This may be accounted for by supposing that these plants and their progenitors have grown accustomed to the repeated action of rain and wind, so that no molecular change is thus induced; whilst they have been rendered more sensitive by means of natural selection to the rarer impact or pressure of solid bodies. Although the absorption by the glands of Drosera of various fluids excites movement, there is a great difference in the action of allied fluids; for instance, between certain vegetable acids, and between citrate and phosphate of ammonia. The specialised nature and perfection of the sensitiveness in these two plants is all the more astonishing as no one supposes that they possess nerves; and by testing Drosera with several substances which act powerfully on the nervous system of animals, it does not appear that they include any diffused matter analogous to nerve-tissue.
Although the cells of Drosera and Dionaea are quite as sensitive to certain stimulants as are the tissues which surround the terminations of the nerves in the higher animals, yet these plants are inferior even to animals low down in the scale, in not being affected except by stimulants in contact with their sensitive parts. They would, however, probably be affected by radiant heat; for warm water excites energetic movement. When a gland of Drosera, or one of the filaments of Dionaea, is excited, the motor impulse radiates in all directions, and is not, as in the case of animals, directed towards special points or organs. This holds good even in the case of Drosera when some exciting substance has been placed at two points on the disc, and when the tentacles all round are inflected with marvellous precision towards the two points. The rate at which the motor impulse is transmitted, though rapid in Dionaea, is much slower than in most or all animals. This fact, as well as that of the motor impulse not being specially directed to certain points, are both no doubt due to the absence of nerves. Nevertheless we perhaps see the prefigurement of the formation of nerves in animals in the transmission of the motor impulse being so much more rapid down the confined space within the tentacles of Drosera than elsewhere, and somewhat more rapid in a longitudinal than in a transverse direction across the disc. These 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.