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As the quadrifids are developed from papillae which at first closely resemble those on the outside of the bladders and on the surfaces of the leaves, I may here state that the two hemispherical cells with which these latter papillae are crowned, and which in their natural state are perfectly transparent, likewise absorb carbonate and nitrate of ammonia; for, after an immersion of 23 hrs. in solutions of one part of both these salts to 437 of water, their primordial utricles were a little shrunk and of a pale brown tint, and sometimes finely granular.
The same result followed from the immersion of a whole branch for nearly three days in a solution of one part of the carbonate to 1750 of water. The grains of chlorophyll, also, in the cells of the leaves on this branch became in many places aggregated into little green masses, which were often connected together by the finest threads.
On the Absorption of certain Fluids by the Glands on the Valve and Collar.--The glands round the orifices of bladders which are still young, or which have been [page 417] long kept in moderately pure water, are colourless; and their primordial utricles are only slightly or hardly at all granular. But in the greater number of plants in a state of nature--and we must remember that they generally grow in very foul water--and with plants kept in an aquarium in foul water, most of the glands were of a pale brownish tint; their primordial utricles were more or less shrunk, sometimes ruptured, with their contents often coarsely granular or aggregated into little masses. That this state of the glands is due to their having absorbed matter from the surrounding water, I cannot doubt; for, as we shall immediately see, nearly the same results follow from their immersion for a few hours in various solutions. Nor is it probable that this absorption is useless, seeing that it is almost universal with plants growing in a state of nature, excepting when the water is remarkably pure.
The pedicels of the glands which are situated close to the slit-like orifice, both those on the valve and on the collar, are short; whereas the pedicels of the more distant glands are much elongated and project inwards. The glands are thus well placed so to be washed by any fluid coming out of the bladder through the orifice. The valve fits so closely, judging from the result of immersing uninjured bladders in various solutions, that it is doubtful whether any putrid fluid habitually passes outwards. But we must remember that a bladder generally captures several animals; and that each time a fresh animal enters, a puff of foul water must pass out and bathe the glands.
Moreover, I have repeatedly found that, by gently pressing bladders which contained air, minute bubbles were driven out through the orifice; and if a bladder is laid on blotting paper and gently pressed, water oozes out. [page 418] In this latter case, as soon as the pressure is relaxed, air is drawn in, and the bladder recovers its proper form. If it is now placed under water and again gently pressed, minute bubbles issue from the orifice and nowhere else, showing that the walls of the bladder have not been ruptured. I mention this because Cohn quotes a statement by Treviranus, that air cannot be forced out of a bladder without rupturing it. We may therefore conclude that whenever air is secreted within a bladder already full of water, some water will be slowly driven out through the orifice. Hence I can hardly doubt that the numerous glands crowded round the orifice are adapted to absorb matter from the putrid water, which will occasionally escape from bladders including decayed animals.
[In order to test this conclusion, I experimented with various solutions on the glands. As in the case of the quadrifids, salts of ammonia were tried, since these are generated by the final decay of animal matter under water. Unfortunately the glands cannot be carefully examined whilst attached to the bladders in their entire state. Their summits, therefore, including the valve, collar, and antennae, were sliced off, and the condition of the glands observed; they were then irrigated, whilst beneath a covering glass, with the solutions, and after a time re-examined with the same power as before, namely No. 8 of Hartnack. The following experiments were thus made.
As a control experiment solutions of one part of white sugar and of one part of gum to 218 of water were first used, to see whether these produced any change in the glands. It was also necessary to observe whether the glands were affected by the summits of the bladders having been cut off. The summits of four were thus tried; one being examined after 2 hrs. 30 m., and the other three after 23 hrs.; but there was no marked change in the glands of any of them.
Two summits bearing quite colourless glands were irrigated with a solution of carbonate of ammonia of the same strength (viz. one part to 218 of water) , and in 5 m. the primordial utricles of most of the glands were somewhat contracted; they were also thickened in specks or patches, and had assumed a pale [page 419] brown tint. When looked at again after 1 hr. 30 m., most of them presented a somewhat different appearance. A third specimen was treated with a weaker solution of one part of the carbonate to 437 of water, and after 1 hr. the glands were pale brown and contained numerous granules.
Four summits were irrigated with a solution of one part of nitrate of ammonia to 437 of water. One was examined after 15 m., and the glands seemed affected; after 1 hr. 10 m. there was a greater change, and the primordial utricles in most of them were somewhat shrunk, and included many granules. In the second specimen, the primordial utricles were considerably shrunk and brownish after 2 hrs. Similar effects were observed in the two other specimens, but these were not examined until 21 hrs. had elapsed. The nuclei of many of the glands apparently had increased in size. Five bladders on a branch, which had been kept for a long time in moderately pure water, were cut off and examined, and their glands found very little modified. The remainder of this branch was placed in the solution of the nitrate, and after 21 hrs. two bladders were examined, and all their glands were brownish, with their primordial utricles somewhat shrunk and finely granular.
The summit of another bladder, the glands of which were in a beautifully clear condition, was irrigated with a few drops of a mixed solution of nitrate and phosphate of ammonia, each of one part to 437 of water. After 2 hrs. some few of the glands were brownish. After 8 hrs. almost all the oblong glands were brown and much more opaque than they were before; their primordial utricles were somewhat shrunk and contained a little aggregated granular matter. The spherical glands were still white, but their utricles were broken up into three or four small hyaline spheres, with an irregularly contracted mass in the middle of the basal part. These smaller spheres changed their forms in the course of a few hours and some of them disappeared. By the next morning, after 23 hrs. 30 m., they had all disappeared, and the glands were brown; their utricles now formed a globular shrunken mass in the middle. The utricles of the oblong glands had shrunk very little, but their contents were somewhat aggregated. Lastly, the summit of a bladder which had been previously irrigated for 21 hrs. with a solution of one part of sugar to 218 of water without being affected, was treated with the above mixed solution; and after 8 hrs. 30 m. all the glands became brown, with their primordial utricles slightly shrunk.
Four summits were irrigated with a putrid infusion of raw [page 420]
meat. No change in the glands was observable for some hours, but after 24 hrs. most of them had become brownish, and more opaque and granular than they were before. In these specimens, as in those irrigated with the salts of ammonia, the nuclei seemed to have increased both in size and solidity, but they were not measured. Five summits were also irrigated with a fresh infusion of raw meat; three of these were not at all affected in 24 hrs., but the glands of the other two had perhaps become more granular. One of the specimens which was not affected was then irrigated with the mixed solution of the nitrate and phosphate of ammonia, and after only 25 m. the glands contained from four or five to a dozen granules. After six additional hours their primordial utricles were greatly shrunk.
The summit of a bladder was examined, and all the glands found colourless, with their primordial utricles not at all shrunk; yet many of the oblong glands contained granules just resolvable with No. 8 of Hartnack. It was then irrigated with a few drops of a solution of one part of urea to 218 of water. After 2 hrs. 25 m. the spherical glands were still colourless; whilst the oblong and two-armed ones were of a brownish tint, and their primordial utricles much shrunk, some containing distinctly visible granules. After 9 hrs. some of the spherical glands were brownish, and the oblong glands were still more changed, but they contained fewer separate granules; their nuclei, on the other hand, appeared larger, as if they had absorbed the granules.
After 23 hrs. all the glands were brown, their primordial utricles greatly shrunk, and in many cases ruptured.
A bladder was now experimented on, which was already somewhat affected by the surrounding water; for the spherical glands, though colourless, had their primordial utricles slightly shrunk; and the oblong glands were brownish, with their utricles much, but irregularly, shrunk. The summit was treated with the solution of urea, but was little affected by it in 9 hrs.; nevertheless, after 23 hrs. the spherical glands were brown, with their utricles more shrunk; several of the other glands were still browner, with their utricles contracted into irregular little masses.
Two other summits, with their glands colourless and their utricles not shrunk, were treated with the same solution of urea. After 5 hrs. many of the glands presented a shade of brown, with their utricles slightly shrunk. After 20 hrs. 40 m. some few of them were quite brown, and contained [page 421] irregularly aggregated masses; others were still colourless, though their utricles were shrunk; but the greater number were not much affected. This was a good instance of how unequally the glands on the same bladder are sometimes affected, as likewise often occurs with plants growing in foul water. Two other summits were treated with a solution which had been kept during several days in a warm room, and their glands were not at all affected when examined after 21 hrs.
A weaker solution of one part of urea to 437 of water was next tried on six summits, all carefully examined before being irrigated. The first was re-examined after 8 hrs. 30 m., and the glands, including the spherical ones, were brown; many of the oblong glands having their primordial utricles much shrunk and including granules. The second summit, before being irrigated, had been somewhat affected by the surrounding water, for the spherical glands were not quite uniform in appearance; and a few of the oblong ones were brown, with their utricles shrunk. Of the oblong glands, those which were before colourless, became brown in 3 hrs. 12 m. after irrigation, with their utricles slightly shrunk. The spherical glands did not become brown, but their contents seemed changed in appearance, and after 23 hrs.
still more changed and granular. Most of the oblong glands were now dark brown, but their utricles were not greatly shrunk. The four other specimens were examined after 3 hrs. 30 m., after 4 hrs., and 9 hrs.; a brief account of their condition will be sufficient. The spherical glands were not brown, but some of them were finely granular. Many of the oblong glands were brown, and these, as well as others which still remained colourless, had their utricles more or less shrunk, some of them including small aggregated masses of matter.]
Summary of the Observations on Absorption.--From the facts now given there can be no doubt that the variously shaped glands on the valve and round the collar have the power of absorbing matter from weak solutions of certain salts of ammonia and urea, and from a putrid infusion of raw meat. Prof. Cohn believes that they secrete slimy matter; but I was not able to perceive any trace of such action, excepting that, after immersion in alcohol, extremely fine lines could sometimes be seen radiating from their [page 422] surfaces. The glands are variously affected by absorption; they often become of a brown colour; sometimes they contain very fine granules, or moderately sized grains, or irregularly aggregated little masses; sometimes the nuclei appear to have increased in size; the primordial utricles are generally more or less shrunk and sometimes ruptured. Exactly the same changes may be observed in the glands of plants growing and flourishing in foul water. The spherical glands are generally affected rather differently from the oblong and two-armed ones. The former do not so commonly become brown, and are acted on more slowly. We may therefore infer that they differ somewhat in their natural functions.
It is remarkable how unequally the glands on the bladders on the same branch, and even the glands of the same kind on the same bladder, are affected by the foul water in which the plants have grown, and by the solutions which were employed. In the former case I presume that this is due either to little currents bringing matter to some glands and not to others, or to unknown differences in their constitution. When the glands on the same bladder are differently affected by a solution, we may suspect that some of them had previously absorbed a small amount of matter from the water. However this may be, we have seen that the glands on the same leaf of Drosera are sometimes very unequally affected, more especially when exposed to certain vapours.
If glands which have already become brown, with their primordial utricles shrunk, are irrigated with one of the effective solutions, they are not acted on, or only slightly and slowly. If, however, a gland contains merely a few coarse granules, this does not prevent a solution from acting. I have never seen [page 423] any appearance making it probable that glands which have been strongly affected by absorbing matter of any kind are capable of recovering their pristine, colourless, and homogeneous condition, and of regaining the power of absorbing.
From the nature of the solutions which were tried, I presume that nitrogen is absorbed by the glands; but the modified, brownish, more or less shrunk, and aggregated contents of the oblong glands were never seen by me or by my son to undergo those spontaneous changes of form characteristic of protoplasm. On the other hand, the contents of the larger spherical glands often separated into small hyaline globules or irregularly shaped masses, which changed their forms very slowly and ultimately coalesced, forming a central shrunken mass. Whatever may be the nature of the contents of the several kinds of glands, after they have been acted on by foul water or by one of the nitrogenous solutions, it is probable that the matter thus generated is of service to the plant, and is ultimately transferred to other parts.
The glands apparently absorb more quickly than do the quadrifid and bifid processes; and on the view above maintained, namely that they absorb matter from putrid water occasionally emitted from the bladders, they ought to act more quickly than the processes; as these latter remain in permanent contact with captured and decaying animals.
Finally, the conclusion to which we are led by the foregoing experiments and observations is that the bladders have no power of digesting animal matter, though it appears that the quadrifids are somewhat affected by a fresh infusion of raw meat. It is certain that the processes within the bladders, and the glands outside, absorb matter from salts of [page 424] ammonia, from a putrid infusion of raw meat, and from urea. The glands apparently are acted on more strongly by a solution of urea, and less strongly by an infusion of raw meat, than are the processes. The case of urea is particularly interesting, because we have seen that it produces no effect on Drosera, the leaves of which are adapted to digest fresh animal matter. But the most important fact of all is, that in the present and following species the quadrifid and bifid processes of bladders containing decayed animals generally include little masses of spontaneously moving protoplasm; whilst such masses are never seen in perfectly clean bladders.
Development of the Bladders.--My son and I spent much time over this subject with small success. Our observations apply to the present species and to Utricularia vulgaris, but were made chiefly on the latter, as the bladders are twice as large as those of Utricularia neglecta. In the early part of autumn the stems terminate in large buds, which fall off and lie dormant during the winter at the bottom.
The young leaves forming these buds bear bladders in various stages of early development. When the bladders of Utricularia vulgaris are about 1/100 inch (.254 mm.) in diameter (or 1/200 in the case of Utricularia neglecta), they are circular in outline, with a narrow, almost closed, transverse orifice, leading into a hollow filled with water; but the bladders are hollow when much under 1/100 of an inch in diameter. The orifices face inwards or towards the axis of the plant. At this early age the bladders are flattened in the plane in which the orifice lies, and therefore at right angles to that of the mature bladders. They are covered exteriorly with papillae of different sizes, many of which have an elliptical outline. A bundle of vessels, formed of [page 425] simple elongated cells, runs up the short footstalk, and divides at the base of the bladder. One branch extends up the middle of the dorsal surface, and the other up the middle of the ventral surface. In full-grown bladders the ventral bundle divides close beneath the collar, and the two branches run on each side to near where the corners of the valve unite with the collar; but these branches could not be seen in very young bladders.
FIG. 23. (Utricularia vulgaris.) Longitudinal section through a young bladder, 1/100 of an inch in length, with the orifice too widely open.
The accompanying figure (fig. 23) shows a section, which happened to be strictly medial, through the footstalk and between the nascent antennae of a bladder of Utricularia vulgaris, 1/100 inch in diameter. The specimen was soft, and the young valve became separated from the collar to a greater degree than is natural, and is thus represented. We here clearly see that the valve and collar are infolded prolongations of the walls of the bladder. Even at this early age, glands could be detected on the valve. The state of the quadrifid processes will presently be described. The antennae at this period consist of minute cellular projections (not shown in the above figure, as they do not lie in the medial plane), which soon bear incipient bristles. In five instances the young antennae were not of quite equal length; and this fact is intelligible if I am right in believing that they represent two divisions of the leaf, rising from the end of the bladder; for, with the true leaves, whilst very young, the divisions are never, as far as I have seen, strictly opposite; they [page 426] must therefore be developed one after the other, and so it would be with the two antennae.
At a much earlier age, when the half formed bladders are only 1/300 inch (.0846 mm.) in diameter or a little more, they present a totally different appearance. One is represented on the left side of the accompanying drawing (fig. 24). The young leaves
FIG. 24. (Utricularia vulgaris.) Young leaf from a winter bud, showing on the left side a bladder in its earliest stage of development.
at this age have broad flattened segments, with their future divisions represented by prominences, one of which is shown on the right side.
Now, in a large number of specimens examined by my son, the young bladders appeared as if formed by the oblique folding over of the apex and of one margin with a prominence, against the opposite margin. The circular hollow between the infolded apex and infolded prominence apparently contracts into the narrow orifice, wherein the valve and collar will be developed; the bladder itself being formed by the confluence of the opposed [page 427] margins of the rest of the leaf.
But strong objections may be urged against this view, for we must in this case suppose that the valve and collar are developed asymmetrically from the sides of the apex and prominence. Moreover, the bundles of vascular tissue have to be formed in lines quite irrespective of the original form of the leaf. Until gradations can be shown to exist between this the earliest state and a young yet perfect bladder, the case must be left doubtful.
As the quadrifid and bifid processes offer one of the greatest peculiarities in the genus, I carefully observed their development in Utricularia neglecta. In bladders about 1/100 of an inch in diameter, the inner surface is studded with papillae, rising from small cells at the junctions of the larger ones. These papillae consist of a delicate conical protuberance, which narrows into a very short footstalk, surmounted by two minute cells. They thus occupy the same relative position, and closely resemble, except in being smaller and rather more prominent, the papillae on the outside of the bladders, and on the surfaces of the leaves. The two terminal cells of the papillae first become much elongated in a line parallel to the inner surface of the bladder. Next, each is divided by a longitudinal partition. Soon the two half-cells thus formed separate from one another; and we now have four cells or an incipient quadrifid process. As there is not space for the two new cells to increase in breadth in their original plane, the one slides partly under the other. Their manner of growth now changes, and their outer sides, instead of their apices, continue to grow. The two lower cells, which have slid partly beneath the two upper ones, form the longer and more upright pair of processes; whilst the two upper cells form the shorter [page 428] and more horizontal pair; the four together forming a perfect quadrifid. A trace of the primary division between the two cells on the summits of the papillae can still be seen between the bases of the longer processes. The development of the quadrifids is very liable to be arrested. I have seen a bladder 1/50 of an inch in length including only primordial papillae; and another bladder, about half its full size, with the quadrifids in an early stage of development.
As far as I could make out, the bifid processes are developed in the same manner as the quadrifids, excepting that the two primary terminal cells never become divided, and only increase in length. The glands on the valve and collar appear at so early an age that I could not trace their development; but we may reasonably suspect that they are developed from papillae like those on the outside of the bladder, but with their terminal cells not divided into two. The two segments forming the pedicels of the glands probably answer to the conical protuberance and short footstalk of the quadrifid and bifid processes.
I am strengthened in the belief that the glands are developed from papillae like those on the outside of the bladders, from the fact that in Utricularia amethystina the glands extend along the whole ventral surface of the bladder close to the footstalk.
Living plants from Yorkshire were sent me by Dr. Hooker. This species differs from the last in the stems and leaves being thicker or coarser; their divisions form a more acute angle with one another; the notches on the leaves bear three or four short bristles instead of one; and the bladders are twice as large, or about 1/5 of an inch (5.08 mm.) in diameter. In all essential respects the bladders resemble those of Utricularia neglecta, but the sides of the peristome are perhaps a little more [page 429] prominent, and always bear, as far as I have seen, seven or eight long multicellular bristles. There are eleven long bristles on each antenna, the terminal pair being included. Five bladders, containing prey of some kind, were examined. The first included five Cypris; a large copepod and a Diaptomus; the second, four Cypris; the third, a single rather large crustacean; the fourth, six crustaceans; and the fifth, ten. My son examined the quadrifid processes in a bladder containing the remains of two crustaceans, and found some of them full of spherical or irregularly shaped masses of matter, which were observed to move and to coalesce. These masses therefore consisted of protoplasm.
FIG. 25. (Utricularia minor.) Quadrifid process, greatly enlarged.
This rare species was sent me in a living state from Cheshire, through the kindness of Mr. John Price. The leaves and bladders are much smaller than those of Utricularia neglecta. The leaves bear fewer and shorter bristles, and the bladders are more globular. The antennae, instead of projecting in front of the bladders, are curled under the valve, and are armed with twelve or fourteen extremely long multicellular bristles, generally arranged in pairs. These, with seven or eight long bristles on both sides of the peristome, form a sort of net over the valve, which would tend to prevent all animals, excepting very small ones, entering the bladder. The valve and collar have the same essential structure as in the two previous species; but the glands are not quite so numerous; the oblong ones are rather more elongated, whilst the two-armed ones are rather less elongated. The four bristles which project obliquely from the lower edge of the valve are short.
Their shortness, compared with those on the valves of the foregoing species, is intelligible if my view is correct that they serve to prevent too large animals forcing an entrance through the valve, thus injuring it; for the valve is already protected to a certain extent by the incurved antennae, together with the lateral bristles. The bifid processes are like those in the previous species; but the quadrifids differ in the four arms (fig. 25) [page 430] being directed to the same side; the two longer ones being central, and the two shorter ones on the outside.
The plants were collected in the middle of July; and the contents of five bladders, which from their opacity seemed full of prey, were examined. The first contained no less than twenty-four minute fresh-water crustaceans, most of them consisting of empty shells, or including only a few drops of red oily matter; the second contained twenty; the third, fifteen; the fourth, ten, some of them being rather larger than usual; and the fifth, which seemed stuffed quite full, contained only seven, but five of these were of unusually large size.
The prey, therefore, judging from these five bladders, consists exclusively of fresh-water crustaceans, most of which appeared to be distinct species from those found in the bladders of the two former species. In one bladder the quadrifids in contact with a decaying mass contained numerous spheres of granular matter, which slowly changed their forms and positions.
This North American species, which is aquatic like the three foregoing ones, has been described by Mrs. Treat, of New Jersey, whose excellent observations have already been largely quoted. I have not as yet seen any full description by her of the structure of the bladder, but it appears to be lined with quadrifid processes. A vast number of captured animals were found within the bladders; some being crustaceans, but the greater number delicate, elongated larvae, I suppose of Culicidae. On some stems, "fully nine out of every ten bladders contained these larvae or their remains." The larvae "showed signs of life from twenty-four to thirty-six hours after being imprisoned," and then perished. [page 431]
Utricularia montana--Description of the bladders on the subterranean rhizomes--Prey captured by the bladders of plants under culture and in a state of nature--Absorption by the quadrifid processes and glands--Tubers serving as reservoirs for water--Various other species of Utricularia--Polypompholyx--Genlisea, different nature of the trap for capturing prey-- Diversified methods by which plants are nourished.
FIG. 26. (Utricularia montana.) Rhizome swollen into a tuber; the branches bearing minute bladders; of natural size.
UTRICULARIA MONTANA.--This species inhabits the tropical parts of South America, and is said to be epiphytic; but, judging from the state of the roots (rhizomes) of some dried specimens from the herbarium at Kew, it likewise lives in earth, probably in crevices of rocks. In English hothouses it is grown in peaty soil. Lady Dorothy Nevill was so kind as to give me a fine plant, and I received another from Dr. Hooker. The leaves are entire, instead of being much divided, as in the foregoing aquatic species. They are elongated, about 1 1/2 inch in breadth, and furnished with a distinct footstalk. The plant produces numerous colourless rhizomes, as thin as threads, which bear minute bladders, and occasionally swell into tubers, as will [page 432] hereafter be described. These rhizomes appear exactly like roots, but occasionally throw up green shoots. They penetrate the earth sometimes to the depth of more than 2 inches; but when the plant grows as an epiphyte, they must creep amidst the mosses, roots, decayed bark, &c., with which the trees of these countries are thickly covered.
As the bladders are attached to the rhizomes, they are necessarily subterranean. They are produced in extraordinary numbers. One of my plants, though young, must have borne several hundreds; for a single branch out of an entangled mass had thirty-two, and another branch, about 2 inches in length (but with its end and one side branch broken off), had seventy- three bladders.* The bladders are compressed and rounded, with the ventral surface, or that between the summit of the long delicate footstalk and valve, extremely short (fig. 27). They are colourless and almost as transparent as glass, so that they appear smaller than they really are, the largest being under the 1/20 of an inch (1.27 mm.) in its longer diameter. They are formed of rather large angular cells, at the junctions of which oblong papillae project, corresponding with those on the surfaces of the bladders of the previous species. Similar papillae abound on the rhizomes, and even on the entire leaves, but they are rather broader on the latter. Vessels, marked with parallel bars instead of by a spiral line, run up the footstalks, and
* Prof. Oliver has figured a plant of Utricularia Jamesoniana ('Proc.
Linn. Soc.' vol. iv. p. 169) having entire leaves and rhizomes, like those of our present species; but the margins of the terminal halves of some of the leaves are converted into bladders. This fact clearly indicates that the bladders on the rhizomes of the present and following species are modified segments of the leaf; and they are thus brought into accordance with the bladders attached to the divided and floating leaves of the aquatic species. [page 433]
just enter the bases of the bladders; but they do not bifurcate and extend up the dorsal and ventral surfaces, as in the previous species.