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A handbook of systematic botany

Chapter 11: Class 4. Schizophyta, Fission-Algæ.
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A comprehensive manual lays out a morphological and comparative framework for plant classification, explaining principles that regard simpler, more complete forms as older and reduced or specialized forms as younger. It gives ordered treatments of Thallophyta, algae, fungi, and vascular plants, presenting diagnostic keys, structural descriptions, and taxonomic sequences that emphasise relationships and progressive reduction. Technical terminology for floral and vegetative organs is defined for consistent use, and recent revisions of algal and fungal groups are integrated. Numerous illustrations and appendices compare earlier classification systems and provide tabular keys to support identification and teaching.

Fig. 17.—Various Diatomeæ. A Synedra radians. B Epithemia turgida (from the two different sides). C Cymbella cuspidata. D Cocconeis pediculus (on the right several situated on a portion of a plant, on the left a single one more highly magnified).

Order 1. Diatomaceæ. This order may be divided into two sub-orders, viz.—

Sub-Order 1. Placochromaticæ. The chromatophores are discoid, large, 1 or 2 in each cell; the structure of the valves is bilateral and always without reticulate markings. The following groups belong to this sub-order: Gomphonemeæ, Cymbelleæ, Amphoreæ, Achnantheæ, Cocconeideæ, Naviculeæ, Amphipleureæ, Plagiotropideæ, Amphitropideæ, Nitzchieæ, Surirayeæ, and Eunotieæ.

Sub-Order 2. Coccochromaticæ. The chromatophores are granular, small and many in each cell. The structure of the cells is zygomorphic or centric, often with reticulate markings. The following groups belong to this sub-order: Fragilarieæ, Meridieæ, Tabellarieæ, Licmophoreæ, Biddulphieæ, Anguliferæ, Eupodisceæ, Coscinodisceæ, and Melosireæ.

Class 4. Schizophyta, Fission-Algæ.

The individuals are 1—many celled; the thallus consists in many of a single cell, in others of chains of cells, the cells dividing in only one definite direction (Figs. 18, 21). In certain Fission-Algæ the cell-chain branches (Fig. 30) and a difference between the anterior and the posterior ends of the chain is marked; in some, the cells may be united into the form of flat plates by the cell-division taking place in two directions; and in others into somewhat cubical masses, or rounded lumps of a less decided form, by the divisions taking place in three directions; or less defined masses may be formed by the divisions taking place in all possible directions.

The cell-walls rarely contain cellulose, they often swell considerably (Figs. 20, 22), and show distinct stratifications, or they are almost completely changed into a mucilaginous mass in which the protoplasts are embedded, e.g. in Nostoc (Fig. 22), and in the “Zooglœa” stage of the Bacteria (Fig. 27). Sexual reproduction is wanting. Vegetative reproduction by division and the separation of the divisional products by the splitting of the cell-wall or its becoming mucilaginous; among the Nostocaceæ, Lyngbyaceæ, Scytonemaceæ, etc., “Hormogonia” are found; in Chamæsiphon and others single reproductive akinetes are formed. Many Fission-Algæ conclude the growing period by the formation of resting akinetes or aplanospores.

The Schizophyta may be divided into 2 families:

1. Schizophyceæ.

2. Bacteria.

Family 1. Schizophyceæ,[5] Blue-Green Algæ.

All the Blue-green Algæ are able to assimilate carbon by means of a colouring material containing chlorophyll (cyanophyll); but the chlorophyll in this substance is masked by a blue (phycocyan), or red (phycoerythrin, e.g. in Trichodesmium erythræum in the Red Sea) colouring matter which may be extracted from them in cold water after death. The colouring matter, in most of them, permeates the whole of the protoplasm (excepting the cell-nucleus), but in a few (e.g. Glaucocystis, Phragmonema), slightly developed chromatophores are to be found. Where the cells are united into filaments (cell-chains) a differentiation into apex and base (Rivulariaceæ) may take place, and also between ordinary vegetative cells and heterocysts; these latter cannot divide, and are distinguished from the ordinary vegetative cells (Fig. 22 h) by their larger size, yellow colour, and poverty of contents. Branching sometimes occurs and is either true or spurious.

Fig. 18.Microcoleus lyngbyanus: a portion of a filament, the thick sheath encloses only one cell-chain; in one place a cell is drawn out by the movement of the cell-chain; b the cell-chain has divided into two parts (“hormongonia”) which commence to separate from each other.

The cell-chain in the spurious branching divides into two parts, of which either one or both grow beyond the place of division (Fig. 18) and often out to both sides (e.g. Scytonema), the divisions however, always take place transversely to the longitudinal direction of the cell-chain. In the true branching a cell elongates in the direction transverse to the cell-chain, and the division then takes place nearly at right angles to the former direction (Sirosiphoniaceæ).

Fig. 19.Cylindrospermum majus: a resting akinete with heterocyst; b-d germinating stages of a resting akinete; e filament with two heterocysts and the formation of new akinetes; f part of a filament with a heterocyst, and mature resting akinete.

Cilia are wanting, but the filaments are sometimes self-motile (e.g. hormogonia in Nostoc) and many partly turn round their axes, partly slide forward or backward (Oscillaria).

Reproduction takes place by spores and hormogonia in addition to simple cell-division. Hormogonia are peculiar fragments of a cell-chain capable of motion, and often exhibit a vigorous motion in the sheath, until at last they escape and grow into a new individual (Fig. 18). The spores are reproductive akinetes (Chamæsiphon, etc.) or resting akinetes; these latter arise by the vegetative cells enlarging and constructing a thick cell-wall (Fig. 19 e f). On germination, this cell-wall bursts and the new cell-chain elongates in the same longitudinal direction as before (Fig. 19 b c). Many (e.g. Oscillaria) may however winter in their ordinary vegetative stage. Aplanospores are wanting.

The Fission-Algæ are very prevalent in fresh water and on damp soil, less so in salt water; they also often occur in water which abounds in decaying matter. Some are found in warm springs with a temperature as high as 50° C.

The Family may be divided into 2 sub-families:

1. Homocysteæ (heterocysts are wanting): Chroococcaceæ, Lyngbyaceæ and Chamœsiphonaceæ.

2. Heterocysteæ (heterocysts present): Nostocaceæ, Rivulariaceæ, Scytonemaceæ and Sirosiphoniaceæ.

Order 1. Chroococcaceæ. The individuals are 1—many-celled, but all the cells are uniform, united to form plates or irregular masses, often surrounded by a mucilaginous cell-wall, but never forming cell-chains. Multiplication by division and sometimes by resting akinetes, but reproductive akinetes are wanting. Chroococcus, Aphanocapsa, Glœocapsa (Fig. 20), Cœlosphærium, Merismopedium, Glaucocystis, Oncobyrsa, Polycystis, Gomphosphæria.

Fig. 20.Glœocapsa atrata: A, B, C, D, E various stages of development.

Fig. 21.Oscillaria; a terminal, b central portion of a filament.

Order 2. Lyngbyaceæ (Oscillariaceæ). The cells are discoid (Fig. 21), united to straight or spirally twisted, free filaments, which are unbranched, or with spurious branching. The ends of the cell-chains are similar. Heterocysts absent. Reproduction by synakinetes, resting akinetes are wanting. Oscillaria (Fig. 21), Spirulina, Lyngbya, Microcoleus, Symploca, Plectonema.

Order 3. Chamæsiphonaceæ. The individuals are 1—many-celled, attached, unbranched filaments with differentiation into apex and base, without heterocysts. Multiplication by reproductive akinetes; resting akinetes are wanting. Dermocarpa, Clastidium, Chamæsiphon, Godlewskia, Phragmonema.

Order 4. Nostocaceæ. The individuals are formed of multicellular, unbranched filaments, without differentiation into apex and base; heterocysts present. Reproduction by synakinetes and resting akinetes.

Fig. 22.Nostoc verrucosum. A The plant in its natural size; an irregularly folded jelly-like mass. B One of the cell-chains enlarged, with its heterocysts (h), embedded in its mucilaginous sheath.

Some genera are not mucilaginous, e.g. Cylindrospermum (Fig. 19). The cell-chains in others, e.g. Nostoc, wind in between one another and are embedded in large structureless jelly-like masses, which may attain the size of a plum or even larger (Fig. 22); sometimes they are found floating in the water, sometimes attached to other bodies. Other genera as follows: Aphanizomenon and Anabæna (in lakes and smaller pieces of water); Nodularia is partly pelagic. Some occur in the intercellular spaces of higher plants, thus Nostoc-forms are found in Anthoceros, Blasia, Sphagnum, Lemna, and in the roots of Cycas and Gunnera; Anabæna in Azolla.

Order 5. Rivulariaceæ. The individuals are multicellular filaments, with differentiation into apex and base; spurious branching, and a heterocyst at the base of each filament, reproduction by synakinetes and resting akinetes, rarely by simple reproductive akinetes. Rivularia, Glœotrichia, Isactis, Calothrix.

Order 6. Scytonemaceæ. The individuals are formed of multicellular filaments with no longitudinal division; differentiation into apex and base very slight or altogether absent; branching spurious; heterocysts present. Reproduction by synakinetes, rarely by resting akinetes and ordinary reproductive akinetes. Tolypothrix, Scytonema, Hassalia, Microchæte.

Order 7. Sirosiphoniaceæ. The individuals are formed of multicellular threads with longitudinal divisions; true branching and heterocysts, and often distinct differentiation into apex and base. Reproduction by synakinetes, rarely by resting akinetes and ordinary reproductive akinetes. Hapalosiphon, Stigonema, Capsosira, Nostocopsis, Mastigocoleus.

Family 2. Bacteria.[6]

The Bacteria (also known as Schizomycetes, and Fission-Fungi) are the smallest known organisms, and form a parallel group to the Blue-green Algæ, but separated from these Algæ by the absence of their colouring material; chlorophyll is only found in a few Bacteria.

The various forms under which the vegetative condition of the Bacteria appear, are termed as follows:

1. Globular forms, cocci (Figs. 27, 30 c): spherical or ellipsoidal, single cells, which, however, are usually loosely massed together and generally termed “Micrococci.”

2. Rod-like forms: more or less elongated bodies; the shorter forms have been styled “Bacterium” (in the narrower sense of the word), and the term “Bacillus” has been applied to longer forms which are straight and cylindrical (Figs. 28, 29, 30 E).

Fig. 23.Spirillum sanguineum. Four specimens. One has two cilia at the same end, the sulphur grains are seen internally.

3. Thread-like forms: unbranched, long, round filaments, resembling those of Oscillaria, are possessed by Leptothrix (very thin, non-granular filaments; Fig. 30 A, the small filaments) and Beggiatoa (thicker filaments, with strong, refractile grains or drops of sulphur (Fig. 31); often self-motile). Branched filaments, with false branching like many Scytonemaceæ, are found in Cladothrix (Fig. 30 B, G).

4. Spiral forms: Rod-like or filamentous bodies, which more or less strongly resemble a corkscrew with a spiral rising to the left. In general these are termed Spirilla (Fig. 23); very attenuated spirals, Vibriones (standing next to Fig. 30 M); if the filaments are slender and flexible with a closely wound spiral, Spirochætæ (Fig. 24).

5. The Merismopedium-form, consisting of rounded cells arranged in one plane, generally in groups of four, and produced by divisions perpendicular to each other.

6. The Sarcina-form, consisting of roundish cells which are produced by cellular division in all the three directions of space, united into globular or ovoid masses (“parcels”) e.g. Sarcina ventriculi (Figs. 25, 26).

Fig. 24.Spirochæte obermeieri, in active motion (b) and shortly before the termination of the fever (c); a blood corpuscles.

All Bacteria are unicellular. In the case of the micrococci this is self-evident, but in the “rod,” “thread,” and “spiral” Bacteria, very often numerous cells remain united together and their individual elements can only be recognised by the use of special reagents.

Fig. 25.Sarcina ventriculi. One surface only is generally seen. Those cells which are drawn with double contour are seen with the correct focus, and more distinctly than those cells lying deeper drawn with single contour.

Fig. 26.Sarcina minuta: a-d successive stages of one individual (from 4–10 p.m.); f an individual of 32 cells.

The condition termed “Zooglœa,” which reminds us of Nostoc, is produced by the cells becoming strongly mucilaginous. A number of individuals in active division are found embedded in a mass of mucilage, which either contains only one, or sometimes more, of the above-named forms. The individuals may eventually swarm out and continue their development in an isolated condition. Such mucilaginous masses occur especially upon moist vegetables (potatoes, etc.), on the surface of fluids with decaying raw or cooked materials, etc. The mucilaginous envelope is thrown into folds when the Bacteria, with their mucilaginous cell-walls, multiply so rapidly that there is no more room on the surface of the fluid.

The cells of the Bacteria are constructed like other plant-cells in so far as their diminutive size has allowed us to observe them. The cell-wall only exceptionally shows the reactions of cellulose (in Sarcina, Leuconostoc; also in a Vinegar-bacterium, Bacterium xylinum); a mucilaginous external layer is always present. The body of the cell mostly appears to be an uniform or finely granulated protoplasm. Very few species (e.g. Bacillus virens) contain chlorophyll; others are coloured red (purple sulphur Bacteria); the majority are colourless. Bacillus amylobacter shows a reaction of a starch-like material when treated with iodine before the spore-formation. Some Bacteria contain sulphur (see p. 37). The body, which has been described as a cell-nucleus, is still of a doubtful nature.

Artificial colourings with aniline dyes (especially methyl-violet, gentian-violet, methylene-blue, fuchsin, Bismarck-brown and Vesuvin) play an important part in the investigations of Bacteria.

Movement. Many Bacteria are self-motile; the long filaments of Beggiatoa exhibit movements resembling those of Oscillaria. In many motile forms the presence of cilia or flagella has been proved by the use of stains; many forms have one, others several cilia attached at one or both ends (Fig. 23) or distributed irregularly over the whole body; the cilia are apparently elongations of the mucilaginous covering and not, as in the other Algæ of the protoplasm. In Spirochæte the movement is produced by the flexibility of the cell itself. Generally speaking, the motion resembles that of swarm-cells (i.e. rotation round the long axis and movement in irregular paths); but either end has an equal power of proceeding forwards.

The swarming motion must not be confounded with the hopping motion of the very minute particles under the microscope (Brownian movement).

Vegetative reproduction takes place by continued transverse division; hence the name “Fission-Fungi” or “Fission-Algæ,” has been applied to the Bacteria.

Spores. The spores are probably developed in two ways. In the ENDOSPOROUS species (Figs. 28, 29), the spore arises as a new cell inside the mother-cell. The spores are strongly refractile, smaller than the mother-cell, and may be compared to the aplanospores of other Algæ. In addition to these there are the ARTHROSPOROUS species in which the cells, just as in Nostoc and other Blue-green Algæ, assume the properties of spores without previously undergoing an endogenous new construction, and are able to germinate and form new vegetative generations (Fig. 27). The formation of spores very often commences when the vegetative development begins to be restricted.

Fig. 27.Leuconostoc mesenterioides: a a zooglœa, natural size; b cross section of zooglœa; c filaments with spores; d mature spores; e-i successive stages of germination; in e portions of the ruptured spore-wall are seen on the external side of the mucilaginous covering. (b-i magnified 520.)

The spores germinate as in Nostoc by the bursting of the external layer of the cell-wall, either by a transverse or longitudinal cleft, but always in the same way, in the same species (Fig. 28, example of transverse cleft).

Distribution. Bacteria and their germs capable of development, are found everywhere, in the air (dust), in surface water, and in the superficial layers of the soil. The number varies very much in accordance with the nature of the place, season, etc. They enter, together with air and food, into healthy animals and occur always in their alimentary tract.

Growth and reproduction depend upon the conditions of temperature. There is a certain minimum, optimum and maximum for each species; for instance (in degrees Centigrade)—

Minim. Opt. Maxim.
Bacillus subtilis + 6 c. 30 + 50
B. anthracis 15 20–25 43
Spirillum choleræ asiaticæ 8 37 40 (but grows only feebly if under 16°).
Bacterium tuberculosis 28 37–38 42

Fig. 28.Bacillus megaterium: a outline of a living, vegetative cell-rod; b a living, motile, pair of rods; p a similar 4-celled rod after the effects of iodine alcohol; c a 5-celled rod in the first stages of spore-formation; d-f successive stages of spore-formation in one and the same pair of rods (in the course of an afternoon); r a rod with mature spores; g1–g3 three stages of a 5-celled rod, with spores sown in nutritive solution; h1–h2, i, k, l stages of germination; m a rod in the act of transverse division, grown out from a spore which had been sown eight hours previously. (After de Bary; a mag. 250, the other figures 600 times).

Fig. 29.Bacillus amylobacter. Motile rods, partly cylindrical and without spores, partly swollen into various special shapes and with spore-formation in the swelling. s Mature spore, with thick mucilaginous envelope. (After de Bary; mag. 600 times, with the exception of s, which is more highly magnified.)

The functions of life cease on a slight excess of the maximum or minimum temperature, numbness setting in when either of these limits is passed. Crenothrix-threads provided with mucilaginous envelopes may, according to Zopf, sustain a temperature of-10°. Some Bacteria are said to be able to resist the exposure to as low a temperature as-110° for a short time. It is not known at what degree of cold the death of the Bacteria occurs: the greatest degree of heat which the vegetative cells can withstand is about the same as that for other vegetative plant-cells, namely, about 50–60° C. Certain Bacteria, e.g. B. thermophilus, grow and thrive vigorously at 70° C. Many spores, on the contrary, are able to bear far higher temperatures (in several species a temperature for some duration of above 100°, those of Bacillus subtilis, for instance, can withstand for hours a temperature of 100° in nutrient solutions; the spores remain capable of development after exposure to a dry heat of 123° C.).

The Desiccation of the air, if prolonged, kills many forms when in the vegetative condition. The spores however can bear a much longer period of dryness, some even several years.

Oxygen. Some species cannot live without a supply of free oxygen (Aerobic), e.g. the Vinegar-bacteria, the Hay-bacilli, the Anthrax-bacilli, the Cholera-Microspira. Other species again thrive vigorously without supply of free oxygen, and are even checked in their development by the admission of air (Anaerobic), e.g. the butyric acid Bacterium (Clostridium butyricium = Bacillus amylobacter). A distinction may be drawn between obligate and facultative aerobics and obligate and facultative anaerobics. Several Bacteria, producing fermentation, may grow without the aid of oxygen when they are living in a solution in which they can produce fermentation; but, if this is not the case, they can only grow when a supply of oxygen is available. A great number of the pathogenic Bacteria belong to the facultative anaerobics.

A luminous Bacterium (Bacillus phosphorescens) which in the presence of a supply of oxygen gives a bluish-white light, has been found in sea-water. Phosphorescent Bacteria have frequently been observed upon decaying sea-fish, as well as on the flesh of other animals; by transferring the Bacteria from cod fish to beef, etc., the latter may be made luminous.

Organic carbon compounds are indispensable for all Bacteria, (except, as it appears, for the nitrifying organisms), as they can only obtain the necessary supplies of carbon from this source. The supplies of nitrogen, which also they cannot do without, can be obtained equally as well from organic compounds as from inorganic salts, such as saltpetre or ammonia-compounds. The various “ash-constituents” are also essential for their nourishment.

While Moulds and Yeast-Fungi grow best in an acid substratum, the Bacteria, on the other hand, generally thrive best in a neutral or slightly alkaline one.

In sterilization, disinfection, and antisepsis, means are employed by which the Bacteria are killed, or checked in their development, for instance, by heat (ignition, cooking, hot vapours, hot air, etc.), or poisons (acids, corrosive sublimate). The process of preserving articles of food, in which they are boiled and then hermetically sealed, aims at destroying the Bacteria, or the spores of those which already may be present in them, and excluding all others.

As the Bacteria are unable to assimilate carbon from the carbonic acid of the air, but must obtain it from the carbon-compounds already in existence in the organic world, they are either saprophytes or parasites. Some are exclusively either the one or the other, obligate saprophytes or parasites. But there are transitional forms among them, some of which are at ordinary times saprophytes, but may, when occasion offers, complete their development wholly or partly as parasites—facultative parasites; others are generally parasitic, but may also pass certain stages of development as saprophytes—facultative saprophytes.

All chlorophyll-free organisms act in a transforming and disturbing manner on the organic compounds from which they obtain their nourishment, and while they themselves grow and multiply, they produce, each after its kind, compounds of a less degree of complexity, i.e. they produce fermentation, putrefaction, sometimes the formation of poisons, and in living beings often disease.

Those organisms which produce fermentation are called ferments; this word, however, is also employed for similar transformations in purely chemical materials (inorganic ferments or enzymes). Many organic (“living”) ferments, among which are Yeast-cells and Bacteria, give off during their development certain inorganic and soluble ferments (enzymes) which may produce other transformations without themselves being changed. Different organisms may produce in the same substratum different kinds of transformation; alcoholic fermentation may for instance be produced by different species of Fungi, but in different proportions, and the same species produces in different substrata, different transformations (e.g. the Vinegar-bacteria oxydize diluted alcohol to vinegar, and eventually to carbonic acid and water).

In the study of Bacteria it is absolutely necessary to sterilize the vessels employed in cultivation, the apparatus, and nutrient solutions, i.e. to free them from Bacteria germs and also to preserve the cultures from the intrusion of any foreign germs (“pure-cultures”). A firm, transparent, nutritive medium is frequently employed. This may be prepared by adding to the nutrient solutions (broth) either gelatine, or—when the Bacteria are to be cultivated at blood-heat—serum of sheep’s or calf’s blood, agar-agar or carragen; serum alone may in itself serve as a nutrient medium. The so-called “plate-cultures” are frequently employed, i.e. the germs are isolated by shaking them with the melted liquid nutrient gelatine, which is then spread on a glass plate and allowed to coagulate; when later on the individual germs grow into colonies, these remain separate in the solid substratum and it is easy to pursue their further development. Similar plate-cultures may also be cultivated in test-tubes and on microscopic slides. The slides and glass plates must be placed in “moist chambers” free from Bacteria. By sowing a few cells (if possible one) using a fine platinum wire, pure cultures for further investigation may be obtained.

In order to prove the relationship between pathogenic Bacteria and certain diseases, the experimental production of pathogenic Bacteria by the inoculation of Bacteria from pure cultures into healthy animals, is very important.

It has not so far been possible to establish a classification of the Bacteria, as the life-history of many species, has not yet been sufficiently investigated.[7] The opinions of botanists are at variance, in many cases, about the forms of growth of a particular kind. Some species are pleomorphic (many-formed) while others possess only one form.

The following Bacteria are Saprophytes:—

Cladothrix dichotoma is common in stagnant and running water which is impregnated with organic matter; the cell-chains have false branching. According to Zopf, Leptothrix ochracea is one of the forms of this species which, in water containing ferrous iron (e.g. as FeCO3), regularly embeds ferric-oxide in its sheath by means of the activity of the protoplasm. Leptothrix ochracea and other Iron-bacteria, according to Winogradsky (1888), do not continue their growth in water free from protoxide of iron; while they multiply enormously in water which contains this salt of iron. The large masses of ochre-coloured slime, found in meadows, bogs, and lakes, are probably due to the activity of the Iron-bacteria.

Fig. 30.Cladothrix dichotoma.

Those forms which, according to Zopf’s views, represent the forms of development of Cladothrix dichotoma are placed together in Fig. 30. A represents a group of plants, seventy times magnified, attached to a Vaucheria. The largest one is branched like a tree, with branches of ordinary form; a specimen with spirally twisted branches is seen to the right of the figure, at the lower part some small Leptothrix-like forms. B shows the manner of branching and an incipient Coccus-formation. C a Coccus-mass whose exit from the sheath has been observed. D the same mass as C after the course of a day, the Cocci having turned into rods. E a group of Cocci in which some have developed into shorter or longer rods. F one of these rods before and after treatment with picric acid, which causes the chain-like structure to become apparent. G a portion of a plant with conspicuous sheath, two lateral branches are being formed. H part of a plant, whose cells have divided and form Cocci. The original form of the cells in which the Cocci are embedded may still be recognised. I. Leptothrix-filaments with conspicuous mucilaginous sheath, from which a series of rods is about to emerge; the rod near the bottom is dead, and has remained lying in the sheath. K part of a plant which is forming Cocci, those at the top are in the zooglœa-stage, at the base they are elongating to form rods and Leptothrix-filaments. L a portion of a branched Cladothrix, which divides into motile Bacillus-forms; the rays at the free ends indicate the currents which the cilia produce in the water. M a spirally-twisted, swarming filament, before and after division into halves. N part of a tree-like zooglœa with Cocci and short rods.—All of these spirilla, zooglœa, etc., which Zopf has connected with Clad. dichotoma, are according to Winogradsky, independent organisms.

Micrococcus ureæ produces urinal fermentation (transformation of urinal matter into ammonium carbonate); aerobic; round cells generally united to form bent chains or a zooglœa.—Several other kinds of Bacteria have the same action as this one: in damp soil containing ammonia-compounds, saltpetre-formations are produced by M. nitrificans and several different kinds of Bacteria.

Micrococcus prodigiosus is found on articles of food containing starch; “bleeding bread” is caused by this Bacterium, which has the power of forming a red pigment; it also occurs in milk, and produces lactic acid.

Leuconostoc mesenterioides is the frog-spawn Bacterium (Fig. 27) which is found in sugar manufactories, and has the power of producing a viscous fermentation in saccharine solutions which have been derived from plants, e.g. in beetroot-sugar manufactories, where large accumulations of mucilage are formed at the expense of the sugar, with an evolution of carbonic acid. The cell-rows, resembling somewhat a pearl necklace, have thick mucilaginous cell-walls, and form white “Nostoc”-lumps. The mucilage eventually deliquesces and the cells separate from each other; arthrospores?—Similar viscous deteriorations occur in beer and wine, which may then be drawn out into long, string like filaments—“ropiness.”

Bacterium aceti, the Vinegar-bacterium, oxidizes alcohol into acetic acid (acetous-fermentation) and forms a greyish covering of Bacteria (“Vinegar-mother”) on the surface of the liquid; the acetic acid formed, becomes by continued oxidization by B. aceti, again transformed into carbonic acid and water. Aerobic; short cylindrical cells, often united into chains, or to form a zooglœa; sometimes also rod-and spindle-shaped. The Vinegar-bacteria and other kinds with ball- or rod-forms sometimes become swollen, spindle-shaped, or oval links; they are supposed to be diseased forms[8] (“Involution-forms”).

Bacillus lacticus (Bacterium acidi lactici, Zopf) is always found in milk which has stood for some time, and in sour foods (cabbage, cucumbers, etc.); it turns the milk sour by producing lactic acid fermentation in the sugar contained in the milk; the lactic acid formed, eventually causes the coagulation of the casein. It resembles the Vinegar-bacteria, occurring as small cylindrical cells, rarely in short rows; not self-motile.—Several other Bacteria appear to act in the same way, some occurring in the mouth of human beings; some of these Bacteria give to butter its taste and flavour.

The kefir-grains which are added to milk for the preparation of kefir, contain in large numbers a Bacterium (Dispora caucasica) in the zooglœa-form, a Yeast-fungus, and Bacillus lacticus. Kefir is a somewhat alcoholic sour milk, rich in carbonic acid; it is a beverage manufactured by the inhabitants of the Caucasus, from the milk of cows, goats, or sheep, and is sometimes used as a medicine. In the production of kefir, lactic acid fermentation takes place in one part of the sugar contained in the milk, and alcoholic fermentation in another part, and the casein which had become curdled is partially liquefied (peptonised) by an enzyme of a Zooglœa-bacterium.

Bacillus amylobacter (Bacillus butyricus), the Butyric-acid-bacterium (Fig. 29), is a very common anaerobic which produces fermentation in sugar and lactic-acid salts, and whose principal product is butyric acid. It destroys articles of food and (together with other species) plays a part in the butyric acid fermentation which is necessary in the making of cheese; it is very active wherever portions of plants are decaying, in destroying the cellulose in the cell-walls of herbaceous plants, and is thus useful in the preparation of flax and hemp. The cells are self-motile, generally cylindrical, sometimes united into short rows; endosporous; the spore-forming cells swell, assume very different forms, and show granulose reaction. The germ-tube grows out in the direction of the long axis of the spore.

Bacillus subtilis, the Hay-bacillus, is developed in all decoctions of hay; a slender, aerobic, self-motile Bacillus; endosporous (aplanospores); the spore-wall ruptures transversely on germination.

Crenothrix kuehniana occurs in the springs of many baths, in wells, in water or drain-pipes.

Fig. 31.Beggiatoa alba: a from a fluid containing abundance of sulphuretted hydrogen; b after lying 24 hours in a solution devoid of sulphuretted hydrogen; c after lying an additional 48 hours in a solution devoid of sulphuretted hydrogen, by this means the transverse walls and vacuoles have become visible.

Beggiatoa (parallel with the Blue-green Alga Oscillaria). Long filaments formed of cylindrical cells which are attached by one of the ends, but which are nearly always free when observed. The filaments, like those of Oscillaria, describe conical figures in their revolutions, the free filaments slide upwards and parallel with one another; sheaths are wanting; strongly refractive sulphur drops are found in the interior. The Beggiatoas are the most prevalent Sulphur-bacteria. They occur, very commonly in large numbers, wherever plant or animal remains are decaying in water in which sulphuretted hydrogen is being formed; thus, for example, B. alba (Fig. 31) occurs frequently as a white covering or slimy film on mud containing organic remains. B. mirabilis is remarkable for its size and its strong peristaltic movements. The Sulphur-bacteria oxidize the sulphuretted hydrogen, and accumulate sulphur in the shape of small granules of soft amorphic sulphur, which in the living cell never passes over into the crystalline state. They next oxidize this sulphur into sulphuric acid, which is immediately rendered neutral by absorbed salts of calcium, and is given off in the form of a sulphate, thus CaCO3 is principally changed into CaSO4. In the absence of sulphur the nutritive processes are suspended, and consequently death occurs either sooner or later. The Sulphur-bacteria may exist and multiply in a fluid which only contains traces of organic matter, in which organisms devoid of chlorophyll are not able to exist. The Beggiatoas very frequently form white, bulky masses in sulphur wells and in salt water, the traces of organic material which the sulphur water contains proving sufficient for them. The cellulose-fermentation, to which the sulphur wells in all probability owe their origin, mainly procures them suitable conditions for existence. The CaCO3 and H2S, formed during the cellulose fermentation by the reduction of CaSO4 is again changed into CaSO4 and CO2 by the Sulphur-bacteria (Winogradsky, 1887).—Other Sulphur-bacteria, the so-called purple Sulphur-bacteria, e.g. B. roseo-persicina, Spirillum sanguineum (Fig. 23), Bacterium sulfuratum, etc., have their protoplasm mixed with a red colouring matter (bacterio-purpurin) which, like chlorophyll, has the power, in the presence of light, of giving off oxygen (as proved by T. W. Englemann, 1888, in oxygen-sensitive Bacteria). The three purple Sulphur-bacteria mentioned, are, according to Winogradsky, not pleomorphic kinds but embrace numerous species.

Many Spirilli (Spirillum tenue, S. undula, S. plicatile, and others) are found prevalent in decaying liquids.

Bacteria (especially Bacilli) are the cause of many substances emitting a foul odour, and of various changes in milk.

Parasitic Bacteria live in other living organisms; but the relation between “host” and parasite may vary in considerable degree. Some parasites do no injury to their host, others produce dangerous contagious diseases; some choose only a special kind as host, others again live equally well in many different ones. There are further specific and individual differences with regard to the predisposition of the host, and every individual has not the same receptivity at all times.

The harmless parasites of human beings. Several of the above mentioned saprophytes may also occur in the alimentary canal of human beings; e.g., the Hay-bacillus, the Butyric-acid-bacillus, etc.; but the gastric juice prevents the development of others, at all events in their vegetative condition. Sarcina ventriculi, “packet-bacterium,” is only known to occur in the stomach and intestines of human beings, and makes its appearance in certain diseases of the stomach (dilation of the stomach, etc.) in great numbers, without, however, being the cause of the disease. It occurs in somewhat cubical masses of roundish cells (Fig. 25).

Less dangerous parasites. In the mouth, especially between and on the teeth, a great many Bacteria are to be found (more than fifty species are known), e.g. Leptothrix buccalis (long, brittle, very thin filaments which are united into bundles), Micrococci in large lumps, Spirochæte cohnii, etc. Some of them are known to be injurious, as they contribute in various ways to the decay of the teeth (caries dentium); a Micrococcus, for instance, forms lactic acid in materials containing sugar and starch, and the acid dissolves the lime salts in the external layers of the teeth: those parts of the teeth thus deprived of lime are attacked by other Bacteria, and become dissolved. Inflammation in the tissues at the root of a tooth, is probably produced by septic materials which have been formed by Bacteria in the root-canal.

Dangerous Parasites. In a large number of the infectious diseases of human beings and animals, it has been possible to prove that parasitic Bacteria have been the cause of the disease. Various pathogenic Bacteria of this nature, belonging to the coccus, rod, and spiral Bacteria groups, are mentioned in the following:—

Pathogenic Micrococci. Staphylococcus pyogenes aureus produces abscesses of various natures (boils, suppurative processes in internal organs). The same effects are produced by—

Streptococcus pyogenes, which is the most frequent cause of malignant puerperal fever; it is perhaps identical with—

Streptococcus erysipelatis, which is the cause of erysipelas in human beings.

Diplococcus pneumoniæ (A. Fränkel) is the cause of pneumonia, and of the epidemic cerebro-spinal meningitis.

Gonococcus (Neisser) is the cause of gonorrhea and inflammation of the eyes.

Pathogenic Rod-Bacteria. Bacterium choleræ gallinarum, an aerobic, facultative parasite which produces fowl-cholera among poultry; it is easily cultivated on various substrata as a saprophyte. The disease may be conveyed both through wounds and by food, and may also be communicated to mammals.

Bacillus anthracis, the Anthrax bacillus (Fig. 32), chiefly attacks mammals, especially herbivorous animals (house mice, guinea-pigs, rabbits, sheep, cattle), in a less degree omnivorous animals (including human beings), and in a still less degree the Carnivores. Aerobic. Cylindrical cells, 3–4 times as long as broad, united into long rod-like bodies, which may elongate into long, bent, and twisted filaments. Not self-motile. Endosporous. Germination takes place without the throwing off of any spore-membrane (compare Hay-bacillus p. 37 which resembles it). Contagion may take place both by introduction into wounds, and from the mucous membrane of the intestines or lungs, both by vegetative cells and by spores; in intestinal anthrax, however, only by spores. The Bacillus multiplies as soon as it has entered the blood, and the anthrax disease commences. The Bacilli not only give off poison, but also deprive the blood of its oxygen. Vegetative cells only occur in living animals. This species is a facultative parasite which in the first stage is a saprophyte, and only in this condition forms spores.