Results of De Bary’s Investigations in Parasitism.
“When the idea of parasitism was rendered definite by the fundamental distinction drawn by De Bary between a parasite and a saprophyte, it soon became evident that some further distinction must be made between obligate facultative parasites and saprophytes respectively. De Bary, when he proposed these terms for adoption, was clearly alive to the existence of transitions which we now know to be numerous and so gradual in character that we can no longer define any such physiological groups. Twenty years ago penicillium and mucor would have been regarded as saprophytes of the most obligate type, but we now know that under certain circumstances these fungi can become parasites, and the borderland between facultative parasites and saprophytes on the one hand, and between the former and true parasites on the other, can no longer be recognised.”
In 1866 the germ of an idea was sown which has taken root and extended. De Bary pointed out that in the case of lichens we have either a fungus parasite on an algæ, or else certain organisms hitherto accepted as algæ are merely incomplete forms.
“In 1879 the same observer definitely launched the new hypothesis of symbiosis. The word itself is due to Frank, who, in a valuable paper on the biology of the thallus of certain lichens, very clearly set forth the existence of various stages of life in common among all the lower forms of plants. The details of these matters are now principally of historical interest. We now know that lichens are dual organisms, composed of various algæ, symbiotic with Ascomycetes, with Basidiomycetes, and, as Massee has shown, even with Gastromycetes. The soil contains also bacterio-lichens. Hence arose a new biological idea—that a fungus may be in such nicely-balanced relationship with the host from which it derives its sustenance, that it may be attended with nearly equal advantage to both.
“In the humus of forests we find the roots of beeches and other Cupuliferæ (willows, pines, and so forth) clothed with a dense mantle of hyphæ, and swollen into fleshlike masses of mycorhiza. In similar soils, and in moorlands, which abound in the slowly decomposing root-fibres and other vegetable remains so characteristic of these soils, the roots of orchids, heaths, gentians, &c., are similarly provided with fungi, the hyphæ of which penetrate further into the tissues, and even send haustoria into the living cells, but without injuring them. As observations multiplied it became clear that the mycorhiza, or fungus-root, was not to be dismissed as a mere case of roots affected by parasites, but that a symbiotic union, comparable to that of the lichens, exists, and we must assume that both tree and fungus derive benefit from the connection.
Fig. 280.—Fine Section through Truffle.
a. Asci filled with spores; b. Mycelia, × 250.
“Frank stated, as the result of his experimental research, that seedling forest-trees cannot be grown in sterilised soil, where their roots are prevented from forming mycorhiza; and he concluded that the fungus conveys organic materials to the roots, which it obtains by breaking down the leaf-mould and decaying plant remains, together with water and minerals from the soil, and plays the especial part of a nitrogen-catching apparatus. In return for this import service the root pays a tax to the fungus by sparing it certain of its tissue contents. It is a curious fact then that the mycorhiza is only formed where humus or vegetable mould abounds.”
These instructive investigations offer an intelligible explanation of the growth of that well-known subterranean fungus, the truffle (Tuber cibarium), the microscopic appearances of a section of which formed the subject of a paper I contributed to “The Popular Science Review” some years ago (1862). The fungus, as will be seen by the fine section cut through a truffle, [Fig. 280], consists of flocculent filaments, which in the first instance cover the ground at the fall of the leaf in autumn, under oak or beech trees, the hyphæ of which penetrate the ground, through the humid soil to the root-hairs of the tree. Filaments (mycelia) are again given off which terminate in asci or sacs filled with minute spores of about 1⁄2500th of an inch in size, while the interspaces are filled up by mycelia, that become consolidated into a firm nut-like body.
What happens, then, is this: Trees and plants with normal roots and root-hairs, when growing in ordinary soil, can adapt their roots to life in a soil heavily charged with humus only by contracting symbiotic association with the fungus and paying the tax demanded by the latter in return for its supplies and services. If this adaptation is impossible, and no other suitable variation is evolved, such trees cannot grow in such soils. The physiological relations of the root to the fungus must be different in details in the case of non-green, purely saprophytic, plants, Neottia, Monotropa, &c., and in that of green plants like Erica, Fagus, and Pinus. It is, however, a well-known fact that ordinary green plants cannot utilize vegetable débris directly, and forest trees do so in appearance only, for the fungi, yeasts and bacteria there are actively decomposing the leaves and other remains. A class of pseudo-symbiotic organisms are, however, being brought into the foreground, where the combined action of two symbionts results in the death of or injury to a third plant, each symbiont alone proving harmless. Some time ago Vuillemin showed that a disease in olives results from the invasion of a bacillus (B. oleæ), which can, however, only obtain its way into the tissues through the passages driven by the hyphæ of a fungus (Chætophoma). The resulting injury is a sort of burr. This observer also observed the same bacillus and fungus in the canker burrs of the ash.
Among many similar cases well worth further attention are the invasion of potato-tubers by bacteria, these making their way down the decaying hyphæ of pioneer fungi. Professor Marshall Ward has seen tomatoes infected by similar means, and other facts show that many bacteria which quicken the rotting of wood are thus led into the tissues by fungi.
Probably no subject in the whole domain of cryptogamic botany has wider bearings on agricultural science than the study of the flora and changes on and in manure and soil. Nitrifying bacteria play a very important part by providing plant life with a most necessary food. They occur in the soil, and two kinds have been described—the one kind converting ammonia into nitrous acid, and the other changing nitrous into nitric acid. We are principally indebted to Winogradsky for our knowledge of these bacteria; he furnishes instances of the bearing of bacteriological work on this department of science, and explains, not only the origin of nitre-beds and deposits, but also the way the ammonia compounds fixed by the soil in the neighbourhood of the root-hairs are nitrified, and so rendered directly available to plant life. The investigations of other observers show that the nitrifying organism is a much more highly-developed and complex form than had been suspected; that it can be grown on various media, and that it exhibits considerable polymorphism—i.e., it can be made to branch out and show other characteristics of a true fungus. “I have,” writes Professor Ward, “for some time insisted on the fact that river water contains reduced forms of bacteria—i.e., forms so altered by exposure to light, changes of temperature, and the low nutritive value of the water, that it is only after prolonged culture in richer food media that their true nature becomes apparent.” Strutzer and Hartleb show that the morphological form of the nitrifying organism can be profoundly altered by just such variations of the conditions described by Ward, and that it occurs as a branched mycelial form; as bacilli or bacteria; or as cocci of various dimensions, according to the conditions.
“These observations, and others made on variations in form (polymorphism) in other fungi and bacteria, open out a vast field for further work, and must lead to advancement in our knowledge of these puzzling organisms; they also help us to explain many inconsistencies in the existing systems of classification of the so-called ‘species’ of bacteria as determined by test-tube culture.”
Algæ.—The algals have a special charm for microscopists. I am free to confess my interest in these organisms, and for several reasons. In this humid climate of ours they are accessible during the greater part of the year; they can be found in any damp soil, in bog, moss, and in water—indeed, wherever the conditions for their existence seem to be at all favourable for development. Should the soil dry up for a time, when the rain returns algæ are seen to spring into life and give forth their dormant spores, which once more resume the circle of formation and propagation. In the earliest stage of development the spore or spore cell is so very small when in a desiccated state, that any number may be carried about by the slightest breath of air and borne away to a great distance. To all such organisms I originally gave the name of Ærozoa; now recognised as ærobic and anærobic organisms ([Fig. 281]).
Fig. 281.—Ærobic Spores × 200.
1. Ærobic fungi caught over a sewer; 2. Fragments of Penicillium spores; 3. Ærobic fungi taken in the time of the cholera visitation, 1854.
With reference to the ærobic bacteria I have only to add that in addition to the simple mode of taking them on glass slides smeared over with glycerine, special forms of æroscopes are now in use for the purpose, consisting of a small cylinder in which a current of air is produced by an aspirator and diffused through a glass vessel containing a sterilised fluid. These are in constant use in all bacteriological laboratories. The results obtained are transferred to sterilised flasks or tubes as those shown in a former chapter.
Miquel, who has given considerable attention to the subject of ærobic and anærobic bacteria, reckons that the number of spores that find their way into the human system by respiration, even should health be perfectly sound, may be estimated at 300,000 a day.
One of the most commonly met with forms of micro-organisms is Leptothrix buccalis. It chiefly finds its nutritive material in the interstices of the teeth, and is composed of short rods and tufted stems of vigorous growth, to which the name of Bacillus subtilis has been given ([Fig. 282]). Among numerous other fungoid bodies discovered in the mouth, Sarcinæ have been found. See [Plate IX]., No. 7.
Fig. 282.—Section of the Mucous Membrane of the Mouth, × 350.
Showing: a. The denser connective tissue; b. Teased out tissue; c. Muscular fibre; d. Leptothrix buccalis, together with minute forms of bacteria and micrococci; e. Ascomycetes and starch granules.
The Beggiatoa, a sewage fungus, found by me in the river Lea water of 1884 growing in great profusion, consists chiefly of mycelial threads and a number of globular, highly refractive bodies, and may be regarded as evidence of the presence in the water of an abnormal amount of sulphates which set free a gas, sulphuretted hydrogen, of a dangerous and offensive character. Another curious body closely allied to Beggiatoa alba is Cladothrix; this assumes a whitish pellicle on the surface of putrefying liquids.
These saprophytes obtain nourishment from organic matter; nevertheless they are not true parasites in the first stage of their existence, during which they live freely in the water or in damp soil; they, however, become pathogenic parasites when they penetrate into the tissues of animals, and necessarily live at the expense of their host.
Fungi, Algæ, Lichens, etc.
Tuffen West, del. Edmund Evans.
Bacteria, as I have said, were for a long time classed with fungi under the name of Schizomycetes. But the more recent researches into their organisation, and more especially into their mode of reproduction, show that they rather more resemble a group of algæ devoid of chlorophyll. Zopf asserts that the same species of algals may at one time be presented in the form of a plant living freely in water, or in damp ground, in association with chlorophyllaceous protoplasm, and at another in the form of a bacterium devoid of green colouring matter, and receiving nourishment from organic substances previously elaborated by plants or animals, thus accommodating itself, according to circumstances, to two very different modes of existence.
That widely-distributed single-cell plant, the Palmoglœa macrococca of Kützing, that spreads itself as a green slime over damp stones, walls, and other bodies, affords an example. If a small portion be scraped off and placed on a slip of glass, and examined with a half or a quarter-inch power, it will be seen to consist of a number of ovoid cells, having a transparent structureless envelope, nearly filled by granular matter of a greenish colour. At certain periods this mass divides into two parts, and ultimately the cell becomes two. Sometimes the cells are united end to end, just as we see them united in the actively-growing yeast plant; but in this case the growth is accelerated, apparently, by cold and damp. Another plant belonging to the same species, the Protococcus pluvialis, is found in every pool of water, the spores of which must be always floating in the air, since they appear after every shower of rain.
Protococcus pluvialis is furnished with motile organs—two or more vibratile flagella passing through perforations in the cell-wall—whereby, at certain stages, they move rapidly about. The flagella are distinctly seen on the application of the smallest drop of iodine. The more remarkable of the several forms presented by the plant is that of naked spores, termed by Flotow Hæmatococcus porphyrocephalus. These minute bodies are usually seen to consist of green, red, and colourless granules in equal proportions, and occupying different portions of the cell. They seem to have some share in the after subdivision of the cell ([Fig. 283]). There are also still-cells, which sub-divide into two, while the motile cells divide into four or eight. It is not quite clear what becomes of the motile zoospores, B, but as they have been seen to become encysted, they doubtless have a special function, or become still-cells under certain circumstances.
It appears that both longitudinal and transverse division of the primordial cell takes place; and that the vibratile flagella of the parent cell retain to the last their function and their motion after the primordial cell has become detached and transformed into an independent secondary cell ([Fig. 283], G).
Fig. 283.—Cell Development. (Protococcus pluvialis.)
Protococcus pluvialis, Kützing. Hæmatococcus pluvialis, Flotow. Chlamidococcus versatilis, A. Braun. Chlamidococcus pluvialis, Flotow and Braun.
A. Division of a simple cell into two, each primordial vesicle having developed a cellulose envelope; B. Zoospores, having escaped from a cell; C. Division of an encysted cell into segments; D. Division of another cell, with vibratile flagella projecting through cell-wall; E. An encysted flagellate cell; F. Division of an encysted nucleated cell into four parts, with vibratile filaments projecting; G. Fission of a young cell.
The most striking of the vital phenomena presented by Protococcus is that of periodicity. Certain forms—for instance, encysted zoospores, of a certain colour, appear in a given infusion, at first exclusively, then they gradually diminish, become more and more rare, and finally disappear altogether. After some time their number again increases, and this may be repeated. Thus, a cell which at one time presented only still forms at another contained only motile ones. The same may be said with respect to segmentation. If a number of motile cells be transferred from a larger vessel into a smaller one, in the course of a few hours most of them will have subsided to the bottom, and in the course of the day observed to be on the point of sub-division. On the following morning division will have become completed; on the next day the bottom of the vessel will be found covered with a new generation of self-dividing cells, which, again, will produce another generation. This regularity, however, is not always observed. The influence of every change in the external conditions of life upon the plant is very remarkable. It is only necessary to pour water from a smaller into a larger or shallower vessel to at once induce segmentation of cells. The same phenomenon occurs in other algals; thus Vaucheria almost always develops zoospores at whatever time of year they may be brought from their natural habitat into a warm room. Light is conducive to the manifestation of vital action in the motile spores; they usually collect in great numbers on the surface of the water, and at that part exposed to the strongest light.
But in the act of propagation, on the contrary, and when about to pass into the still condition, the motile Protococcus cell seems to shun light, and falls to the bottom of the vessel. Too strong sunlight, as when concentrated by a lens, quickly kills the young zoospores. A temperature of undue elevation is injurious to the development of their vital activity and the formation of the zoospores. Frost destroys motile, but not still zoospores.[55]
Stephanosphæra pluvialis is a conspicuous variety of the fresh-water algals, described by Cohn. It consists of a cell containing eight primordial cells filled with chlorophyll, uniformly arranged (see [Plate I]., No. 24 d). The globular mother-cell rotates, somewhat in the same way as the volvox, by vibratile flagella, two of which are seen projecting from each cell and piercing the transparent outer cell wall. Every cell divides first into two, then four, and lastly eight cells, each one of which again divides into a number of micro-gonidia, which have a motion within the mother-cell, and ultimately escape from it. Under certain circumstances each of the eight young cells is observed to change places in the interior of the cell; eventually they escape, lose their flagella, form a thicker membrane as at b, and for a time remain motionless, and sink to the bottom of the vessel in which they are contained. If the vessel is permitted to become thoroughly dry, and then again has water poured into it, motile cells reappear; from which circumstance it is probable that these represent the resting spores of the plant. When in the condition of greatest activity its division into eight is perfected during the night, and early in the morning light the young cells escape and pass through similar changes. It is calculated that in eight days, under favourable circumstances, 16,777,216 families may be formed from one resting-cell of Stephanosphæra. In certain of the cells, and at particular periods, remarkable amœboid bodies ([Plate I]., No. 24 c) make their appearance. There is a marked difference between Stephanosphæra and Chlamydococcus, for while in the latter the individual portions of a primordial cell separate entirely from one another, each developing its own enveloping membrane, and ultimately escaping as a unicellular individual; in the former, on the other hand, the eight portions remain for a time living in companionship.
Volvocineæ.—A fresh-water unicellular plant of singular beauty and interest to the microscopist is the Volvox globator ([Plate I]., No. 15). No. 16 represents a portion of another cell, with brownish amœboid bodies enclosed in the protoplasmic web. It is common to our fresh-water pools, and attains a diameter of about 1⁄20th or 1⁄30th of an inch. Its movement is peculiar, a continued roll onwards, or a rotation like that of a top; at other times it glides along smoothly. When examined under a sufficiently high power, it is seen to be a hollow sphere, studded with green spots, and traversed by green threads connecting each of the spots or spores with the maternal cell. From each of the spores proceed two long flagella, lashing filaments, which keep the globular body on the move. After a time the sphere bursts, and the contained sporules issue forth and speedily pass through a similar stage of development. These interesting cells were long taken to be animal bodies. Ehrenberg described them as Monads, possessing a mouth, stomach, and an eye.
The setting free of the young volvox is essentially a process of cell division, occurring during the warmer periods of the year, and, as Professor Cohn shows, is a considerable advance upon the simpler conjugation of two smaller cells in desmids; it more closely resembles that which prevails among the higher algæ and a large number of cryptogams. As autumn advances the volvox spherules usually cease to multiply by the formation of zoosporanges, and certain of their ordinary cells begin to undergo changes by which they are converted, some into male or sperm-cells, others into germ-cells, but the greater number appear to remain sterile. Both kinds of cells at first so nearly resemble each other that it is only when the sperm cells begin to undergo sub-division that they are seen to be about three times the size of the sterile cells. Then the primary cell resolves itself into a cluster of peculiar secondary cells, each consisting of an elongated body containing an orange-coloured endochrome and a pair of long flagella, as seen in the antherozoids of the higher cryptogams. As the sperm-cells approach maturity the clusters may be seen to move within them; the bundles then separate and show an independent active movement while still within the cavity of the primary cell, and finally escape through a rupture in the cell-wall, rapidly diffusing themselves as they pass through the cavity. The germ-cells continue to increase in size without undergoing sub-division, at first showing large vacuoles in their protoplasm, but subsequently becoming filled with a darker coloured endochrome. The form of the cell also changes from its flask-like shape to the globular, and at the same time seems to acquire a firmer envelope. Over this the swarming antherozoids diffuse themselves and penetrate the substance to the interior, and are then lost to view. The product of this fusion, Cohn tells us, is a reproductive cell, or “oospore,” which speedily becomes enveloped in another membrane with a thicker external coat, beset with conical-pointed processes; and now the chlorophyll of the young cell gives place, as in Palmoglæ, to starch and reddish or orange-coloured, and a more highly refractive, fluid. As many as forty of such oospores have been counted in a single sphere of volvox, which then acquires the peculiar appearance observed by Ehrenberg, and described by him under the name of Volvox stellatus. The further history of this wonderful spheroid unicellular plant has been traced out by Kirchner, who found that their germination commences in the early months of the year—in February—with the liberation of the spherical endospore from its envelope and its division into four cells. A remarkable phenomenon has been observed by Dr. Braxton Hicks—the conversion of an ordinary volvox cell into a moving mass of protoplasm that bears a striking resemblance to the well-known amœba. “Towards the end of the autumn the endochrome mass of the volvox increases to nearly double its ordinary size, but instead of undergoing the usual sub-division so as to produce a macrogonidium, it loses its colour and regularity of form, and becomes an irregular mass of colourless protoplasm, containing a number of brownish granules.” The final change and the ultimate destination of these curious amœboid bodies have not been satisfactorily made out, but from other observations on the protoplasmic contents of the cells of the roots of mosses, which in the course of two hours become changed into ciliated bodies, it is believed that this is the mode in which these fragile structures are enabled to retain life and to resist all the external conditions, such as damp, dryness, and the alternations of heat and cold.
It would be quite impossible to deny the great similarity there is between the structure of volvox and that of the motile cell of Protococcus pluvialis. The influence of reagents will sometimes cause the connecting processes of the young cells, as in Protococcus, to be drawn back into the central mass, and the connecting threads are sometimes seen as double lines, or tubular prolongations of the membrane. At other times they appear to be connected by star-like prolongations to the parent cell ([Plate I]., No. 15), presenting an almost identical appearance with Pediastrum pertusum. Another body designated by Ehrenberg Sphærosira volvox is an ordinary volvox in a different stage of development; its only features of dissimilarity being that a large proportion of the green cells, instead of being single, are double or quadruple, and that the groups of flagellate cells form by their aggregation discoid bodies, each furnished with a single flagellum. These clusters separate themselves from the parent cell, and swim off freely under the forms which have been designated Uvella and Syncrypta by Ehrenberg. Mr. Henry Carter, F.R.S., who made a careful investigation of unicellular plants, described Sphærosira as the male, or spermatic form of volvox.
Among other organisms closely allied to volvox and included in Volvocineæ, affording the microscopist many interesting transitional forms in their various modes of fructification, are the Eudorina, still-water organisms that pass through a similar process of reproduction as the volvox. In the Pandorina morum, its reproduction is curiously intermediate between the lower and the higher types; as within each cell is a mulberry-like mass, composed of cells possessing a definite number of swarm spores, sixteen usually, which rupture the mother cell, and swim off furnished with a pair of flagella. A similar process takes place in some of the Confervaceæ and other fresh-water algæ. The Palmella, again, consist of ([Plate I]., No. 21) minute organisms of very simple structure, which grow either on damp surfaces or in fresh water. The stonework of some of our churches is often seen to be covered with a species of Palmella, that take the form of an indefinite slimy film. The “red snow” of Arctic or Alpine regions, considered to be a species of Protococcus, is frequently placed among Palmella. A more characteristic form of the P. cruenta is the Hæmatococcus sanguinis, the whole mass of which is sub-divided by partitions enclosing a larger or smaller number of cells, which diffuse their granular contents through the gelatinous mass in which their several changes take place. The albuminoid envelope of these masses is seen to contain parasitic growths, which have given rise to some discussion, especially when their filaments are observed to radiate in various directions.
The Oscillariaceæ constitute a genus of Confervaceæ which have always had very great interest for the microscopist in consequence of their very remarkable animal-like movements, and from which they derive their generic name. For more than a century these Bacillaria have excited the curiosity of all observers without any one having derived more than an approximate idea of their remarkable rhythmical movements. The frustule consists of a number of very fine short threads attached together by a gelatinous sheath, in one species all of equal length. Their backward and forward movement is of a most singular character; the only other conferva in which I have seen a motion of a similar kind is the Schizonema. In this species the frustules are packed together in regular series, the front and side views being always in the same direction. These several bodies move along within the filamentous sheath without leaving their respective places. On carefully following the movement, it is seen at first much extended, and then more compressed, while the frustules become more linear in their arrangement, and present a closer resemblance to Bacillaria paradoxa, augmented by the circumstance that the frustules are seen to move in both directions. A frustule of Schizonema can move independently of the sheath, and so will a detached frustule of bacillaria. This peculiar and exceptionally anomalous phenomenon as that of the movements of bacillaria can hardly be confined to a solitary species. The movements of the frustules are much accelerated by warmth and light. The longer filaments of other minute species only slightly exhibit any motion of the kind, but have peculiar undulating motions.
Fig. 284.—Confervaceæ.
1. Volvox globator; 2. A section of volvox, showing the flagellate margin of the cell; 3. A portion more highly magnified, to show the young volvocina, with their nuclei and thread-like attachments; 4. Spirogyra, near which are spores in different stages of development; 5. Conferva floccosa; 6. Stigeoclonium protensum, jointed filaments and single zoospores; 7. Staurocarpus gracilis, conjugating filaments and spores.
Confervaceæ are a genus of algals. The species consist of unbranched filaments composed of cylindrical or moniliform cells, with starch granules. Many are vesicular, and all multiply by zoospores generated in the interior of the plant at the expense of the granular matter. They are, for the most part, found in fresh water attached or floating, some in salt water, and a few in both, in colour usually green, but occasionally olive, violet, and red. The Confervaceæ proper are often divided into four families: 1. Hydrodictidæ; 2. Zygnemidæ; 3. Confervidæ; 4. Chætophoridæ. To the microscopist all the plants of this genera are extremely interesting as subjects for the study of cell multiplication. The process usually takes place in the terminal cell, the first step in which is the division of the endochrome, and then follows a sort of hour-glass contraction across the cavity of the parent cell, whereby it is divided into two equal parts. This is better seen in some of the desmids than in [Fig. 284], Nos. 4, 5, and 6. Some species are characterised by a different mode of reproduction; these possess a number of nuclei, and multiply by zoospores of two kinds, the largest of which have either two or four cilia, which germinate directly the smaller are biciliated; conjugation has been seen to take place in a few instances.
Allied to the Confervaceæ is an interesting plant, Sphæroplea annulina, which has received careful attention from Cohn. The oospores of this plant are the product of a process partaking of a sexual nature, and when mature are filled with reddish fat vesicles which divide by segmentation.
The Ædogoniaceæ also closely resemble Confervaceæ in habits of life, but differ in some particulars, especially so in the mode of reproduction (only a single large zoospore being set free from each cell) and by the almost complete fission of the cell-wall or one of the rings which serve as a hinge. The zoospores are the largest known among algals, and each is described as having a red eye-spot. The Chætophoraceæ form an interesting group of confervoid plants, and are usually found in running streams, as they prefer pure water. One of the characteristics of the group is that the extremities of the branches are prolonged into an acute-shaped termination, as represented in [Fig. 284], No. 6. A very pretty object under the microscope is Draparnaldia glomerata, belonging to this species. It consists of an axis composed of a row of cells, and at regular intervals whorls of slender prolongations, containing chlorophyll or endochrome of a deeper green; these attain to an extraordinary length.
The Batrachospermæ bear a strong resemblance to frog-spawn, from which they derive their name, and are chiefly a marine group of algals allied to the Rhodespermeæ or red seaweeds. The late Dr. A. Hassall first described them; they have since received more careful attention from M. Sirodot. They are reddish-green, extremely flexible, and nothing can surpass the grace of their movements in water; but when removed from their element they lose all form, and resemble a jelly-like substance without a trace of organisation; but if allowed to remain quiet they regain their original shape.
The presence of the cell-membrane will be best demonstrated by breaking up the filaments, either by moving the thin glass cover, or by cutting through a mass of them in all directions with a fine dissecting knife. On now examining the slide, in most instances many detached empty pieces of the cell-membrane, with its striæ, will be seen, as well as filaments partly deprived of protoplasm. On the application of iodine all these appearances become more distinguishable in consequence of the filament turning red or brown, while the empty cells remain either unaffected, or present a slight yellowish tint, as is frequently the case with cellulose when old.
Fig. 285.—Mesoglia vermicularis.
With regard to the contents of the cell, the endochrome is coloured in the Oscillatoriæ, and is distinguishable by circular bands or rings around the axis of the cylindrical filament. Iodine stains them brown or red, and syrup and dilute sulphuric acid produce a beautiful rose colour. As to their mode of propagation, nothing positive is known. If kept for some time they gradually lose their green colour; a portion of the mass, becoming brown, sinks to the bottom of the vessel, and presents a granular layer.
Mesoglia vermicularis ([Fig. 285]) consists of strings of cells cohering and held together by their membranous covering. In the lowly organised plant Vaucheria ([Plate I]., No. 23, V. sessilis)—so named after its discoverer Vaucher, a German botanist—a genus of Siphonaceæ, we have an example of true processes of sexual generation. The branching filaments are often seen to bear at their sides peculiar globular bodies or oval protuberances, nipple-shaped buddings-out of the cell-wall, filled with a dark-coloured endochrome and distributed in pairs, one of which curves round to meet the other, when conjugation is seen to take place. Near these bodies others are found with pointed projections, which have been described as “horns,” but these, Pringshelm says, are “antherids which produce antherozoids in their interior,” while the capsule-like bodies constituting the oospores become, when fertilised, a new generation, which swarm out through a cavity or aperture in the parent cell-wall.
The fruit of fresh-water and most olive-green algals is enclosed in spherical cavities under the epidermis of the frond, termed conceptacles, and may be either male or female. The zoids are bottle-shaped and have flagella; the transparent vesicle in which they are contained is itself enclosed in a second of similar form. In monœcious and diœcious algals the female conceptacles are distinguished from the male by their olive colour. The spores, when developed, are borne on a pedicle emanating from the inner wall of the conceptacle. They rupture the outer wall at its apex; at first the spore appears simple, but soon after a series of changes takes place, consisting in a splitting up of the endochrome into six or eight masses of spheroidal bodies. A budding-out occurs in a few hours’ time, and ultimately elongates into a cylindrical thread. The Vaucheria present a double mode of reproduction, and their fronds consist of branching tubes resembling in their general character that of the Bryophyta, from which indeed they differ only in respect of the arrangement of their green contents. In that most remarkable plant Saprolegnia ferox, which is structurally so closely allied to Vaucheria, though separated from them by the absence of green colouring matter, a corresponding analogy in the processes of development takes place. In the formation of its zoospores, an intermediate step is presented between that of the algæ and a class of plants formally placed among fungi.
The Ulvaceæ.—The typical form of seaweeds is the Ulva lactuca, well known from its fronds of dark-green “laver” on every coast throughout the world. Ulvæ are seen to differ but little from the preceding group of fresh-water algals. The specific difference is that the cells, when multiplied by binary subdivision, not only remain in firm connection with each other but possess a more regular arrangement. The frond plane of the algal is either more simple or lobed, and is formed of a double layer of cells closely packed together and producing zoospores. The whole group is chiefly distinguished from Porphyra by their green colour, the latter being roseate or purple. Ulvæ are mostly marine, with one or two exceptions. One species (U. thermalis) grows in the hot springs of Gastein, in a temperature of about 117° Fahr. The development of Ulvæ is seen in [Fig. 286]. The isolated cells, A, resemble in some points those of the Protococcus; these give rise to successive subdivisions determining the clusters seen at B and C, and by their aggregation to the confervoid filament shown at D. These filaments increase in length and breadth by successive additions, and finally take the form of fronds, or rows of cells.
Fig. 286.—Successive Stages of Development of Ulvæ.
A. Isolated spores; B and C. Clusters of cells; D. Cells in the filamentous stage.
Fig. 287.—Sphacelaria cirrhosa, with spores borne at the sides of the branchlets.
The marine greenish-olive algæ present a general appearance which might at first sight be mistaken for plants of a higher order of cryptogams. Their fronds have no longer the form of a filament, but assume that of a membranous expansion of cells. The cells in which zoospores are found have an increased quantity of coloured protoplasm accumulated towards one point of the cell-wall; while the zoospores are observed to converge with their apices towards the same point. In some algæ, which seem to be closely related in form and structure to the Bryophyta, we notice this important difference, that the zoospores are developed in an organ specially destined for the purpose, presenting peculiarities of form and distinguishing it from other parts of the branching tubular frond. In the genus Derbesia distinct spore cases develop, a young branch of which, when destined to become a spore case, instead of elongating indefinitely, begins, after having arrived at a certain length, to swell out into an ovoid vesicle, in the cavity of which a considerable accumulation of protoplasm takes place. This is separated from the rest of the plant, and becomes an opaque mass, surrounded by a distinct membrane. After a time a division of the mass takes place, and a number of pyriform zoospores, each of which is furnished with flagella, are set free.
Desmidiaceæ, Diatomaceæ, Algæ.
Tuffen West, del. Edmund Evans.
Plate II.
Fig. 288.—Cutleria dichotoma. Section of lacinia of a frond, showing the stalked eight-chambered oosporanges growing on tufts with intercalated filaments. Magnified 50 diameters.
In Cutleria ([Fig. 288]) we have a special feature of interest with two kinds of organs, seemingly opposed to each other with regard to their reproductive functions. The sporangia not only differ from those of other species, but the frond consists of olive-coloured irregularly-divided flagella, on each side of which tufts (sori) consisting of the reproductive organs, intermixed with hair-like bodies, are scattered. The zoospores are divided by transverse partitions into four cavities, each of which is again bisected by a longitudinal median septum. When first thrown off they are in appearance so much like the spores of Puccinia that they may be mistaken for them, although so very much larger than those of other olive-coloured algæ.
Florideæ, the red algæ ([Plate II].), present many varieties of structure, although less appears to be known of their reproductive processes than of lower forms of cryptogamic plants. These are, however, of three kinds. The first, to which the term polyspore has been applied, is that of a gelatinous or membranous pericarp or conceptacle, in which an indefinite number of zoospores are contained. This organ may be either at the summit or base of a branch, or it may be concealed in or below the cortical layer of the stem. In some cases a number of spore-bearing filaments emanate from a kind of membrane at the base of a spheroidal cellular perisporangium, by the rupture of which the zoospores formed from the endochrome of the filaments make their escape. Other changes have been observed; however, they all agree in one particular, namely, that the zoospore is developed in the interior of a cell, the wall of which forms its perispore, and the internal protoplasmic membrane endochrome, the zoospore itself, for the escape of which the perispore opens out at its apex.
Fig. 289.—Dasya kutzingiana, with seed vessel and two rows of tetraspores. Magnified 50 diameters.
The second form is more simple, and consists of a globular or ovoid cell, containing a central granular mass; this ultimately divides into four quadrate-shaped spores; these, on attaining maturity, escape by rupture of the cell-wall. Another organ, called a tetraspore, takes its origin in the cortical layer. The tetraspores are arranged either in an isolated manner along the branches, or in numbers together; in some instances the branches that contain them are so modified in form they look like special organs, and have been called stichidia; as, for example, in Dasya ([Fig. 289]). Of the third kind of reproductive organ a difference of opinion exists as to the signification of their antheridia; although always produced in precisely the same situations as the tetraspores and polyspores, they are agglomerations of little colourless cells, either united in a bunch, as in Griffithsia, or enclosed in a transparent cylinder, as in Polysiphonia, or covering a kind of expanded disc of peculiar form, as in Laurencia. According to competent observers, the cells contain spermatozoids. Nägeli describes the spermatozoid as a spiral fibre, which, as it escapes, lengthens itself in the form of a screw. Thuret, on the contrary, says the contents are granular, and offer no trace of a spiral filament, but are expelled from the cells by a slow motion. The antheridia appear in their most simple form in Callithamnion ([Plate II]., Nos. 32 and 34), being reduced to a small mass of cells composed by numerous little bunches which are sessile on the bifurcations of the terminal branches. The spores are simpler structures than the tetraspores, and mostly occupy a more important position. They are not scattered through the frond, but grouped in definite masses, and generally enclosed in a special capsule or conceptacle, which may be mistaken for a tetraspore case. The simplest form of the spore fruit consists of spherical masses of spores attached to the wall of the frond, or imbedded in its substance, without a proper conceptacle; such a fruit is called a favellidium, and occurs in Halymenia; the same name is applied to the fruits of similar structures not perfectly immersed, as those of Gigartina, Gelidium, &c., where they form tubercular swellings on the lobes. In some, the tubercles present a pore at the summit, through which the spores emerge forth. In other cases, as in Ceramium ([Plate II]., Nos. 27 and 37), the spores occupy a more conspicuous place; a characteristic species is Delessaria ([Plate II]., No. 39), the coccidium either occurring on lateral branches, or is sessile on the face of the frond, when it consists of a case filled with angular-shaped spores attached to the wall of the case. The general external appearance of the red algæ is very varied, but it seems to attain to its deepest colouring in the Red Sea, which, it is said, is entirely due to the peculiarly vivid red seaweed. They are all exquisite objects for the microscope, as may be surmised from the few varieties presented in [Plate II]. The Florideæ of the warmer seas exhibit most elegantly formed fronds, as will be seen on reference to the “Phycologia Australica” of the late Dr. William Harvey, F.R.S.
The Characeæ may be placed among the highest of the algals, if only for the complexity of their reproductive organs, which certainly offer a contrast in their simplicity of structure. Chara vulgaris, stonewort, is a simple fresh-water plant, preferring still freshwater ponds or slow-moving rivers running over a chalky soil. It thus derives the calcareous matter found in the axis of the plant, together with a small portion of silica. Its filaments (or branches, as some botanists prefer to call them) are given off in whorls. The Characeæ are a small family of acrogens, consisting of only two or three at most. They are monœcious and diœcious, the two kinds of fruit being often placed close together. They may easily be grown in a tall glass jar for observation. All that is necessary is to put the jar occasionally under the house tap and let the water run slowly over the top for a short time, thus renewing the contents without disturbing the plant. The hard water supplied to London suits chara better than softer water. Both chara and nitella are objects of great interest to microscopists, since in the former the important fact of vegetable circulation was first observed. A portion of the plant of the natural size is shown in [Fig. 290], No. 1.
Characeæ.
Fig. 290.—Diagrammatic sketch of Chara.
1. A stem of Chara vulgaris, natural size; 2. Magnified view (arrows indicating the course taken by the chlorophyll); 3. A limb, with buds protruding; 4. Portion of a leaf of Vallisneria spiralis, showing cyclosis of chlorophyll granules.
Each plant is composed of an assemblage of long tubiform cells placed end to end, with fixed intervals, around which the branchlets are disposed with great regularity. In nitella the stem and branches are composed of simple cells, which sometimes attain to several inches in length. Each node, or zone, from which the branches spring, consists of a single plate, or layer, of small cells, which are a continuation of the cortical layer of the internode ([Fig. 290], No. 3) as an outgrowth. Each cell is partially filled with chlorophyll granules, and it is these that are seen under the microscope taking the course shown by the arrows ([Fig. 290], No. 2). The rate of movement of the granules is accelerated by moderate warmth and retarded by cold. It is in viewing the circulation in water plants that the warm stage of the microscope is brought into use. Borne along with the protoplasmic stream are a number of solid particles consisting of starch granules and other matters. The method of viewing the circulation is by cutting sections off a portion of the plant with a very sharp knife, and arranging them in a growing cell with a few drops of water, and covering over with a thin cover-glass.
Fig. 291.—The Fructification of Chara fragilis.
A. Portion of filament containing “antheroids”; B. A group of antheridial filaments, composed of a series of cells, within each of which antherozoids are formed; C. The escape of mature antherozoids, with whip-like prolongations, about to swim off; D. Antherid supported on flask-shaped pedicle; E. Nucule enlarging, and seen to contain oospores; F. Spores and elaters of Equisetum; G. Spores surrounded by elaters of Equisetum.
The reproductive process of Chara is effected by two sets of bodies, both of which are placed at the base of the branches ([Fig. 291], E and D) either on the same or different plants, one set known as globules or antherids, and the other as nucules, containing the oospores or archegones. These are often of a bright red colour, and have covering plates, or shields (B and E), curiously marked, and the central portion is composed of a number of filaments rolled up (as in E) or free (as seen at B), projecting out from the centre of the sphere. The antherid is supported on a short flask-shaped pedicle, which projects into the interior. At the apex of each of the eight manubria is a roundish hyaline cell, termed a capitulum, and at its apex again six smaller or secondary capitula. The long whip-shaped filaments are divided by transverse septa into a hundred or more compartments, every one of which is filled with an antherozoid (as at A), consisting of a spiral thread of protoplasm packed into two or three coils; these escape and become free (as seen at C), each having two long fine flagella. The young antherozoid swims off with a lashing action, and the whole field appears for a time filled with life. They swim about freely, but their motion gradually ceases, and soon they arrive at a state of inaction.
Nitella appears to have a somewhat different mode of fructification to that of its congener. It puts forth a long filamentous branch from one of its joints, which, on reaching the surface of the water, terminates in a whitish fruit-like cluster. It is even a more delicate and less robust algal than chara, and every care should be taken to imitate the still water in which it grows. It delights in shady woods and in calcareous open pools.
Similar care is requisite with regard to Vallisneria; and a more equal temperature is better suited to the growth of this aquatic plant. It should be planted in the middle of the jar or aquarium, about two inches deep in mould, closely pressed down, then gently fill the jar with water. When the water requires changing, a small portion only should be run off at a time. It appears to thrive in proportion to the frequency of changing the water, and taking care that the water added rather increases the temperature than lowers it.
The natural habitat of the Frog-bit, another water-plant of much interest, is found on the surface of ponds and ditches; in the autumn its seeds fall, and become buried in the mud at the bottom during the winter; in the spring these plants rise to the surface, produce flowers, and grow throughout the summer. Chara may be found in many places around London, and in the upper reaches of the Thames.
Anacharis alsinastrum.—This remarkable plant is so unlike any other water-plant that it may be at once recognised by its leaves growing in threes round a slender stem. It is also known as “Waterthyme,” from a resemblance it bears to that plant.
The colour of the plant is deep green; the leaves are nearly half an inch long, by an eighth wide, egg-shaped at the point, with serrated edges. Its powers of increase are prodigious, as every fragment is capable of becoming an independent plant, producing roots and stems, and extending itself indefinitely in every direction. The specific gravity of it is so nearly that of water, that it is more disposed to sink than float. A small branch of the plant is represented, with a hydra attached to it, in a subsequent chapter.
The special cells in which the circulation is most readily seen are the elongated cells around the margin of the leaf and those of the midrib. On examining the leaf with polarised light, the cells are observed to contain a large proportion of silica, and present a very interesting appearance. A bright band of light encircles the leaf, and traverses its centre. In fact, the leaf is set, as it were, in a framework of silica. By boiling the leaf for a short time in equal parts of nitric acid and water, a portion of the vegetable tissue is destroyed, and the silica rendered more distinct, without changing the form of the leaf.
It is necessary to make a thin section or strip from the leaf of Vallisneria for the purpose of exhibiting the circulation in the cells, as shown in [Fig. 290], No. 4. Among the cell granules, a few of a more transparent character than the rest, are seen to have a nucleolus within.
The phenomenon of cell cyclosis occurs in other plants beside those growing in water. The leaf of the common plantain or dock, Plantago, furnishes a good example, the movement being seen both in the cells of the plant and hairs of the cuticle torn from the midrib.
Cell-division.—In order to study the process of cell-division the hairs on the stamens of Tradescantia should be taken. Remove one from a bud on a warm day and let a drop of a one per cent. sugar solution fall upon it, and cover it with a thin glass cover. Place it for a short time in a moist-chamber ([Fig. 256]), and then examine it with a magnifying power of 500 diameters. The nucleus of the cell will be seen, near its terminal position, to gradually elongate in the direction of the longer axis of the cell and become more granular, while the protoplasm moves towards the extreme end; the nucleus at the same time will present a striated appearance, with the fibrilla arranged parallel to the longer axis of the nucleus, and at length approach each other at the poles. A nuclear spindle will now be produced, and the fibres ruptured in the equatorial plane, so that two nuclei will be found in place of the one. The best preparations of nuclei are obtained by making thin longitudinal sections of actively-growing plants (young rootlets of Pinus, for example), and staining them with hæmatoxylin in the manner described in a former chapter.