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A handbook of systematic botany

Chapter 15: Class 6. Chlorophyceæ (Green 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. 32.Anthrax bacillus (Bacillus anthracis) with red (b) and white (a) blood-corpuscles.

Fig. 33.Anthrax bacillus. The formation of the spores; magnified 450 times.

Bacillus tuberculosis produces tuberculosis in human beings, also in domestic animals (perlsucht). It is a distinct parasite, but may also live saprophytically. It is rod-formed, often slightly bent, and is recognised principally by its action with stains (when stained with an alkaline solution of methyl-blue or carbolic fuchsin, it retains the colour for a long time even in solutions of mineral acids, in contrast with the majority of well-known Bacteria): it probably forms spores which are able to resist heat, dryness, etc.

Bacillus lepræ produces leprosy; Bacillus mallei produces glanders; Bacillus tetani, tetanus (the tetanus bacillus is very common in soil; anaerobic); Bacillus diphtheriæ, diphtheria; Bacillus typhosus, typhoid fever, etc.

Pathogenic Spiral Bacteria. Spirochæte obermeieri (Fig. 24) produces intermittent fever (febris recurrens); it makes its appearance in the blood during the attacks of fever, but it is not to be found during intervals when there is no fever. Obligate parasite.

Spirillum choleræ asiaticæ (Microspira comma) without doubt produces Asiatic cholera; an exceedingly motile spirillum, which is also found in short, bent rods (known as the “Comma-bacillus”), it lives in the intestines of those attacked by the disease, and gives off a strong poison which enters the body. It is easily cultivated as a saprophyte.

A great many circumstances seem to show that a number of other infectious diseases (syphilis, small-pox, scarlet-fever, measles, yellow-fever, etc.) owe their origin to parasitic Bacteria, but this has not been proved with certainty in all cases.

It has been possible by means of special cultivations (ample supply of oxygen, high temperature, antiseptic materials) to produce from the parasitic Bacteria described above (e.g. the fowl-cholera and the anthrax Bacteria) physiological varieties which are distinct from those appearing in nature and possess a less degree of “virulence,” i.e. produce fever and less dangerous symptoms in those animals which are inoculated with them. The production of such physiological varieties has come to be of great practical importance from the fact that they are used as vaccines, i.e. these harmless species produce in the animals inoculated with them immunity from the malignant infectious Bacteria from which they were derived. This immunity is effected by the change of the products of one or more of the Bacteria, but we do not yet know anything about the way in which they act on the animal organism. The white blood corpuscles, according to the Metschnikoff, play the part of “Phagocytes” by absorbing and destroying the less virulent Bacteria which have entered the blood, and by so doing they are gradually enabled to overcome those of a more virulent nature.

Fig. 34.a and b The same blood-cell of a Frog: a in the act of engulfing an anthrax-bacillus; b after an interval of a few minutes when the bacillus has been absorbed.

Class 5. Conjugatæ.

The Algæ belonging to this class have chlorophyll, and pyrenoids round which starch is formed. The cells divide only in one direction, they live solitarily, or united to form filaments which generally float freely (seldom attached). Swarm-cells are wanting. The fertilisation is isogamous (conjugation) and takes place by means of aplanogametes. The zygote, after a period of rest, produces, immediately on germination, one or more new vegetative individuals; sometimes akinetes or aplanospores are formed in addition. They only occur in fresh or slightly brackish water.

Order 1. Desmidiaceæ. The cells generally present markings on the outer wall, and are mostly divided into two symmetrical halves by a constriction in the middle, or there is at least a symmetrical division of the protoplasmic cell-contents. The cell-wall consists nearly always of two layers, the one overlapping the other (Fig. 35 C). The cells either live solitarily or are united into unbranched filaments. The mass of protoplasm formed by the fusion of the two conjugating cells becomes the zygote, which on germination produces one (or after division 2, 4 or 8) new vegetative individual. The chromatophores are either star-, plate-, or band-shaped, and regularly arranged round the long axis of the cell.

Fig. 35.—A Cell of Gymnozyga brebissonii, external view showing the distribution of the pores. B A portion of the membrane of Staurastrum bicorne with pores containing protoplasmic projections. C Cell-wall of Hyalotheca mucosa during cell-division: the central part, being already formed, shows the connection with the divisional wall.

The Desmidiaceæ are not able to swim independently, many species, however, show movements of different kinds by rising and sliding forward on the substratum. These movements, which are partly dependent upon, and partly independent of light and the force of gravitation, are connected with the protrusion of a mucilaginous stalk. The mucilage, which sometimes surrounds the whole individual, may acquire a prismatic structure, it is secreted by the protoplasmic threads which project through certain pores definitely situated in the walls (Fig. 35 A, B).

Vegetative multiplication takes places by division. A good example of this is found in Cosmarium botrytis (Fig. 36 A-D). The nucleus and chromatophores divide, and simultaneously the central indentation becomes deeper, the outer wall is then ruptured making a circular aperture through which the inner wall protrudes forming a short, cylindrical canal between the two halves to which it is attached (Fig. 36 C). After elongation the canal is divided by a central transverse wall, which commences as a ring round its inner surface and gradually forms a complete septum. The dividing wall gradually splits, and the two individuals separate from each other, each one having an old and a new half. The two daughter-cells bulge out, receive a supply of contents from the parent-cells, and gradually attain their mature size and development (Fig. 36 B-D). Exceptions to this occur in some forms.

Fig. 36.Cosmarium botrytis. A-D Different stages of cell-division.

Fig. 37.Cosmarium meneghinii: a-c same individual seen from the side, from the end, and from the edge; d-f stages of conjugation; g-i germination of the zygote.

Conjugation takes place in the simplest way in Mesotænium, where the two conjugating cells unite by a short tube (conjugation-canal), which is not developed at any particular point. The aplanogametes merge together after the dissolution of the dividing wall, like two drops of water, almost without any trace of preceding contraction, so that the cell-wall of the zygote generally lies in close contact with the conjugating cells. The conjugating cells in the others lie either transversely (e.g. Cosmarium, Fig. 37 d; Staurastrum, etc.), or parallel to one another (e.g. Penium, Closterium, etc.), and emit a short conjugation-canal (Fig. 37 d) from the centre of that side of each cell which is turned towards the other one. These canals touch, become spherical, and on the absorption of the dividing wall the aplanogametes coalesce in the swollen conjugation-canal (Fig. 37 e), which is often surrounded by a mucilaginous envelope. The zygote, which is often spherical, is surrounded by a thick cell-wall, consisting of three layers; the outermost of these sometimes bears thorn-like projections, which in some species are simple (Fig. 37 f), in others branched or variously marked; in some, however, it remains always smooth (e.g. Tetmemorus, Desmidium). Deviation from this mode of conjugation may occur within certain genera (e.g. Closterium, Penium). Upon germination the contents of the zygote emerge, surrounded by the innermost layers of the wall (Fig. 37 g, h) and generally divide into two parts which develop into two new individuals, placed transversely to each other (Fig. 37 i); these may have a somewhat more simple marking than is generally possessed by the species.

Fig. 38.—Desmidiaceæ. A Closterium moniliferum; B Penium crassiusculum; C Micrasterias truncata (front and end view); D Euastrum elegans; E Staurastrum muticum (end view).

The most frequent genera are:—

A. Solitary cells: Mesotænium, Penium (Fig. 38 B), Cylindrocystis, Euastrum (Fig. 38 D), Micrasterias (Fig. 38 C), Cosmarium (Fig. 36, 37), Xanthidium, Staurastrum (Fig. 38 E), Pleurotænium, Docidium, Tetmemorus, Closterium (Fig. 38 A), Spirotænia.

B. Cells united into filaments: Sphærozosma, Desmidium, Hyalotheca, Gymnozyga, Ancylonema, Gonatozygon.

Order 2. Zygnemaceæ. Cell-wall without markings. The cells are cylindrical, not constricted in the centre, and (generally) united into simple, unbranched filaments. The whole contents of the conjugating cells take part in the formation of the zygote, which on germination grows out directly into a new filament.

Spirogyra is easily recognised by its spiral chlorophyll band; Zygnema has two star-like chromatophores in each cell (Fig. 40); both these genera are very common Algæ in ponds and ditches.

Fig. 39.Spirogyra longata. A At the commencement of conjugation, the conjugation-canals begin to protrude at a and touch one another at b; the spiral chlorophyll band and cell-nuclei (k) are shown. B A more advanced stage of conjugation; a, a’ the rounded female and male aplanogametes: in b’ the male aplanogamete is going over to and uniting with the female aplanogamete (b).

Fig. 40.—A cell of Zygnema. S Pyrenoid.

Fig. 41.Zygnema insigne, with zygote.

Fig. 42.—Germinating zygote of Spirogyra jugalis: the young plant is still unicellular; the end which is still in the wall of the zygote is elongated and root-like; the chromatophore divides and forms the spiral band.

The conjugation among the Zygnemaceæ takes place in the following manner: the cells of two filaments, lying side by side, or two cells, the one being situated above the other in the same filament (Fig. 41), push out small protuberances opposite each other (Fig. 39 A, a, b); these finally meet, and the dividing wall is absorbed so that a tube is formed connecting one cell with the other; the protoplasmic contents round off, and the whole of these contents of one of the cells glides through the conjugation-tube and coalesces with that of the other (Fig. 39 B), the aggregate mass then rounds off, surrounds itself with a cell-wall, and becomes a zygote. A distinct difference may be found between the cells in the two filaments, those in the one whose protoplasmic contents pass over being cylindrical, while those of the recipient one are more barrel-shaped, and of a larger diameter. The former may be regarded as a male, the latter as a female plant. The zygote germinates after a period of rest, and grows out into a new filament (Fig. 42).

Order 3. Mesocarpaceæ. The cell-walls are glabrous, unconstricted in the centre, and united into simple unbranched filaments. The chromatophore consists of an axial chlorophyll-plate, with several pyrenoids. The zygote is formed by the coalescence of two cells (Fig. 43) (sometimes three or four), but the whole protoplasmic contents of the cells do not take part in this process, a portion always remaining behind; the aplanogametes coalesce in the conjugation-canal. The zygote thus formed appears incapable of germination until after 3–5 divisions. Of the cells so formed, only one is fertile, the sterile cells, according to Pringsheim, constituting a rudimentary sporocarp. The germinating cells grow out into a new filament. In this order, conjugation has been observed between two cells of the same filament. The Mesocarpaceæ thrive best in water which contains lime.

Fig. 43.—Mougeotia calcarea. Cells showing various modes of conjugation: at m tripartition; pg quadripartition; s quinquipartilion of the zygote.

Class 6. Chlorophyceæ (Green Algæ).

These Algæ are coloured green by chlorophyll, seldom in combination with other colouring matter, and then especially with red. The product of assimilation is frequently starch, which generally accumulates round certain specially formed portions of protoplasm termed pyrenoids. The thallus is uni- or multicellular; in the higher forms (certain Siphoneæ) the organs of vegetation attain differentiation into stem and leaf. The asexual reproduction takes place in various ways; the sexual reproduction is effected by conjugation of motile gametes, or by oogamous fertilisation. The swarm-cells (zoospores, gametes, and spermatozoids) are constructed symetrically, and have true protoplasmic cilia, these generally being attached to the front end of the swarm-cells. Most of these Algæ live in water (fresh or salt); some are found upon damp soil, stones, or tree-stems, and some live enclosed in other plants.

The Class is divided into three families:—

1. Protococcoideæ: Volvocaceæ, Tetrasporaceæ, Chlorosphæraceæ, Pleurococcaceæ, Protococcaceæ, Hydrodictyaceæ.

2. Confervoideæ: Ulvaceæ, Ulothricaceæ, Chætophoraceæ, Mycoideaceæ, Cylindrocapsaceæ, Œdogoniaceæ, Coleochætaceæ, Cladophoraceæ, Gomontiaceæ, Sphæropleaceæ.

3. Siphoneæ: Botrydiaceæ, Bryopsidaceæ, Derbesiaceæ, Vaucheriaceæ, Phyllosiphonaceæ, Caulerpaceæ, Codiaceæ, Valoniaceæ, Dasycladaceæ.

Family 1. Protococcoideæ.

The Algæ which belong to this group are uni- or multicellular with the cells more or less firmly connected, sometimes in a definite, sometimes in an indefinite form (Fig. 47). Colonies are formed either by division or by small unicellular individuals becoming united in a definite manner; the colonies formed in this latter way are termed Cœnobia. Apical cells and branching are absent. Multiplication by division; asexual reproduction by zoospores, rarely by akinetes. Sexual reproduction may be wanting, or it takes place by isogamous, rarely by oogamous fertilisation.

Some are attached by means of a stalk to other objects (Characium, Fig. 49), others occur as “Endophytes” in the tissues of certain Mosses or Phanerogams, e.g. Chlorochytrium lemnæ, in Lemna trisulca; Endosphæra, in the leaves of Potamogeton, Mentha aquatica, and Peplis portula; Phyllobium, in the leaves of Lysimachia nummularia, Ajuga, Chlora, and species of Grasses; Scotinosphæra in the leaves of Hypnum and Lemna trisulca; the majority, however, live free in water and in damp places. Many species which were formerly considered to belong to this family have been proved to be higher Algæ in stages of development.

Order 1. Volvocaceæ. The individuals in this order are either uni- or multicellular, and during the essential part of their life are free-swimming organisms. They are generally encased in a mucilaginous envelope, through which 2–6 cilia project from every cell. The vegetative reproduction takes place by the division of all, or a few, of the cells of the individual; in some a palmella-stage is found in addition. The sexual reproduction takes place by isogamous or oogamous fertilisation.

The Volvocaceæ may be considered to include the original forms of the Chlorophyceæ, because, among other reasons, the motile stage is here the most prominent; they also form the connecting link between the animal Flagellata, and forms intermediate to the Syngeneticæ may perhaps be found amongst them. Three series of green Algæ may be supposed to have taken their origin from the Volvocaceæ: Conjugateæ (Desmidiaceæ) which have lost the swarming stage, but whose conjugation is the nearest to the fertilisation in Chlamydomonas pulvisculus: the Protococcaceæ in which the vegetative divisions have disappeared, while the swarming stage continues to be present, though of shorter duration; and Tetrasporaceæ, in which the vegetative divisions are more prominent, whilst the swarming stage is less so.

A. Unicellular Individuals. The principle genera are: Chlamydomonas, Sphærella, Phacotus.—Sphærella nivalis is the Alga which produces the phenomenon of “Red Snow,” well known on high mountains and on ice and snow fields in the polar regions. The red colouring matter which appears in this and other green Algæ, especially in the resting cells, is produced by the alteration of the chlorophyll.

Phacotus lenticularis has an outer covering incrusted with lime, which, at death, or after division, opens out into two halves. Species may be found among Chlamydomonas, in which conjugation takes place between gametes of similar size without cell-wall, but in C. pulvisculus conjugation takes place between male and female aplanogametes which are surrounded by a mucilaginous envelope.

Fig. 44.Gonium pectorale.

Fig. 45.Pandorina morum.

B. Multicellular Individuals. The most important genera are Gonium, Stephanosphæra, Pandorina, Eudorina, Volvox.—Gonium has 4 or 16 cells arranged in a definite pattern in a flat plate (Fig. 44). Pandorina (Fig. 45), has 16 cells arranged in a sphere (Fig. 45 A). The vegetative reproduction takes place in this way: each cell, after having rounded off, and after the withdrawal of the cilia, divides itself into 16 new ones (Fig. 45 B), each forming a new individual, which soon grows to the size of the mother-individual. It was in this Alga that the conjugation of self-motile gametes was first discovered by Pringsheim, 1869. When conjugation is about to take place, each cell divides into sixteen, as in vegetative reproduction, but the 16 × 16 cells all separate from one another (Fig. 45 C, female gametes, and D, male gametes), and swarm solitarily in the water. The male are, most frequently, smaller than the female, but otherwise they are exactly alike; they are more or less pear-shaped, with a colourless anterior end, 2 cilia, a red “eye-spot,” etc. After swarming for some time they approach each other, two and two, generally a large and a smaller one, and come into contact at their colourless end; in a few moments they coalesce and become one cell (Fig. 45 E, F), this has at first a large colourless anterior end, 4 cilia, and 2 “eye-spots” (Fig. 45 G), but these soon disappear and the cell becomes uniformly dark-green and spherical, and surrounds itself with a thick cell-wall, losing at the same time its power of motion: the zygote (Fig. 45 H) is formed, and becomes later on a deep red colour. On the germination of the zygote, the protoplasmic cell-contents burst open the wall (Fig. 45 J), and emerge as a large swarmspore (Fig. 45 K) which divides into 16 cells, and the first small individual is formed (Fig. 45 L, M).

Eudorina is like Pandorina in structure, but stands somewhat higher, since the contrast between the conjugating sexual cells is greater, the female one being a motionless oosphere.

Fig. 46.Volvox globator, sexual individual: a antheridia which have formed spermatozoids; b oogonia.

The highest stage of development is found in Volvox (Fig. 46). The cells are here arranged on the circumference of a sphere, and enclose a cavity filled with mucilage. The number of these cells may vary from 200–22,000, of which the majority are vegetative and not reproductive, but some become large, motionless oospheres (Fig. 46 b); others, which may appear as solitary individuals, divide and form disc-shaped masses of from 8–256 small spermatozoids (Fig. 46 a). After the oosphere has been fertilised by these, the oospore surrounds itself by a thick, sometimes thorny cell-wall, and on germination becomes a new individual of few cells. A few cells conspicuous by their larger size may be found (1–9, but generally 8) in certain individuals, and these provide the vegetative reproduction, each forming by division a new individual.

Order 2. Tetrasporaceæ reproduce both by vegetative divisions and swarmspores, some have also gamete-conjugation. The principal genera are: Tetraspora, Apiocystis, Dactylococcus, Dictyosphærium, Chlorangium.

Order 3. Chlorosphæraceæ. Chlorosphæra.

Order 4. Pleurococcaceæ. In this order the swarm-stages and sexual reproduction are entirely absent. Vegetative reproduction by division. The principal genera are: Pleurococcus (Fig. 47), Scenedesmus (Fig. 48), Raphidium, Oocystis, Schizochlamys, Crucigenia, Selenastrum.—Pleurococcus vulgaris (Fig. 47) is one of the most common Algæ throughout the world, occurring as green coverings on tree-stems, and damp walls, and it is one of the most common lichen-gonidia.

Fig. 47.Pleurococcus vulgaris.

Fig. 48.Scenedesmus quadricauda.

Order 5. Protococcaceæ. The cells are motionless, free or affixed on a stalk (e.g. Characium, Fig. 49), either separate or loosely bound to one another; they never form multicellular individuals. Multiplication by division is nearly always wanting. Reproduction takes place by swarmspores, which have 1 or 2 cilia, and sexual reproduction in some by gamete-conjugation. The principal genera are: Chlorococcum, Chlorochytrium, Chlorocystis, Scotinosphæra, Endosphæra, Phyllobium, Characium, Ophiocytium, Sciadium.

Fig. 49.Characium strictum. A The cell-contents have divided into many swarmspores. B Swarmspores escaping.

Order 6. Hydrodictyaceæ. The individuals are unicellular but several unite after the zoospore-stage into definitely formed families (cœnobia). Ordinary vegetative division is wanting, but asexual reproduction takes place by zoospores (or by motionless cells without cilia), which unite and form a family similar to the mother-family, inside the mother-cell, or in a mucilaginous envelope. Where sexual reproduction is found it takes place by gamete-conjugation. The principal genera are: Pediastrum (Fig. 50), Cœlastrum, Hydrodictyon (Fig. 51).

Fig. 5O.Pediastrum asperum.

Fig. 51.Hydrodictyou reticulatum. A A cell where the zoospores are on the point of arranging themselves to form a net. B A cell with gametes swarming out.

The cœnobium of Hydrodictyon reticulatum (Water-net) is formed of a large number of cells which are cylindrical, and attached to one another by the ends (Fig. 51). The asexual reproduction takes place by zoospores, which are formed in large numbers (7,000–20,000) in each mother-cell, within which they move about for a time, and then come to rest and arrange themselves into a new net (Fig. 51 A) which is set free by the dissolution of the wall of the mother-cell, grows, and becomes a new cœnobium. The sexual reproduction takes place by gamete-conjugation. The gametes are formed in the same manner as the zoospores, but in larger numbers (30,000–100,000), and swarm out of the mother-cell (Fig. 51 B). The zygote forms, on germination, 2–5 large zoospores, each with one or two cilia, these generally swarm about for a time, and after a period of rest become irregular thorny bodies (polyhedra); their contents again divide into zoospores, the thorny external coating of the polyhedra is cast off, and the zoospores, surrounded by the dilated internal coating, unite to form a small family, which produces several others in the manner described.

Family 2. Confervoideæ.

The individuals are always multicellular, the cells firmly bound together and united into unbranched or branched filaments, expansions, or masses of cells which grow by intercallary divisions or have apical growth. In the first seven orders the cells are uninuclear, but the cells of the remaining three orders contain several nuclei. Asexual reproduction by zoospores, akinetes or aplanospores. Sexual reproduction by isogamous or oogamous fertilisation.

The Confervoideæ, through the Ulvaceæ, are connected with the Tetrasporaceæ, and from the Coleochætaceæ, which is the most highly developed order, there are the best reasons for supposing that the Mosses have taken their origin. The Cladophoraceæ show the nearest approach to the Siphoneæ.

Order 1. Ulvaceæ. The thallus consists of one or two layers of parenchymatous cells, connected together to form either a flat membrane (Monostroma, Ulva) or a hollow tube (Enteromorpha), and may be either simple, lobed, or branched. Reproduction takes place by detached portions of the thallus; or asexually by zoospores or akinetes. Gamete-conjugation is known to take place in some members of this order, the zygote germinating without any resting-stage. The majority are found in salt or brackish water.

Fig. 52.Ulothrix zonata: a portion of a filament with zoospores, which are formed two in each cell (zoosporangium); the dark spots are the red “eye-spots”; 1, 2, 3, 4, denote successive stages in the development of the zoospores; b a single zoospore, v the pulsating vacuole; c portion of a filament with gametes, sixteen are produced in each gametangium; d free gametes, solitary or in the act of conjugation; e the conjugation is completed, and the formed zygote has assumed the resting-stage.

Order 2. Ulothricaceæ. The thallus consists normally of a simple unbranched filament (sometimes a small expansion consisting of one layer of cells is formed, as in Schizomeris and Prasiola which were formerly described as separate genera). Asexual reproduction takes place by means of zoospores (with 1, 2, or 4 cilia), akinetes or aplanospores; the last named may germinate immediately, or only after a period of rest. Sexual reproduction takes place by the conjugation of gametes of about the same size, each having two cilia (Fig. 52 d). The zygote of Ulothrix, on germination, produces a brood of zoospores which swarm for a time and then elongate to become Ulothrix-filaments (alternation of generations). The gametes may also germinate without conjugation in the same manner as the zoospores. The principal genera are: Ulothrix, Hormidium, Conferva, Microspora.—Ulothrix zonata is very common in running fresh water. Nearly all the species of Hormidium occur on damp soil, tree-stems and stones.

Order 3. Chætophoraceæ. The thallus consists of a single, branched, erect or creeping filament of cells, often surrounded by mucilage. The cells have only one nucleus. Asexual reproduction by zoospores with 2 or 4 cilia, by akinetes, or aplanospores. In many, conjugation between gametes with 2 cilia may be found. They approach on one side, Ulothricaceæ, and on the other, Mycoideaceæ. The principal genera are: Stigeoclonium, Draparnaldia, Chætophora, Entoderma, Aphanochæte, Herposteiron, Phæothamnion, Chlorotylium, Trichophilus, Gongrosira, Trentepohlia. Most of the species of Trentepohlia are coloured red by the presence of a red colouring material, which occurs in addition to the chlorophyll. They are aerial Algæ which live on stones (T. jolithus, “violet stone,” so named on account of its violet-like odour in rainy weather), on bark and old wood (T. umbrina), or on damp rocks (T. aurea). Trichophilus welckeri lives in the hair of Bradypus.

Order 4. Mycoideaceæ. The thallus is discoid, consisting of one or more cell-layers, and is always attached. Asexual reproduction by zoospores with 2 or 4 cilia. Sexual reproduction in some species by the conjugation of gametes with 2 cilia. This order forms the connecting link between Chætophoraceæ and Coleochætaceæ. The species occur in fresh water (Chætopeltis) as well as in salt (Pringsheimia), on the carapace of tortoises (Dermatophyton = Epiclemmydia), or endophytic between the cuticle and the epidermal cells of the leaves of tropical plants, destroying the leaf-tissue (Mycoidea).

Order 5. Cylindrocapsaceæ. The thallus consists of a simple (rarely, in parts, formed of many rows) unbranched filament, attached in the young condition, which has short cells with a single nucleus, and is enveloped in a thick envelope with a laminated structure. Asexual reproduction by zoospores with 2 cilia, which are formed 1, 2, or 4 in each vegetative cell. The antheridia are produced by a single cell, or a group of cells, in a filament, dividing several times without increasing in size. Two egg-shaped spermatozoids, each with 2 cilia (Fig. 53 D), are formed in each antheridium, and escape through an aperture in the side; in the first stages they are enclosed in a bladder-like membrane (Fig. 53 B, C). Other cells of the filament swell out and form oogonia (Fig. 53 A), which resemble those of Œdogonium. After fertilisation, the oospore surrounds itself with a thick wall, and assumes a reddish colour. The germination is unknown. The unfertilised oospheres remain green, divide often into 2–4 daughter-cells, and grow into new filaments.

Fig. 53.Cylindrocopsa involuta. A Oogonium with oosphere (o) surrounded by spermatozoids (s). B Two antheridia, each with two spermatozoids. C Spermatozoids surrounded by their bladder-like membrane. D Free spermatozoid.

This order, which only includes one genus, Cylindrocapsa, forms the connecting link between Ulothricaceæ and Œdogoniaceæ. The few species (4) occur only in fresh water.

Order 6. Œdogoniaceæ. The thallus consists of branched (Bulbochæte) or unbranched (Œdogonium) filaments, attached in the early stages. The cells may be longer or shorter, and have one nucleus. Asexual reproduction by zoospores, which have a chaplet of cilia round the base of the colourless end (Fig. 6 a). Sexual reproduction takes place by oogamous fertilisation. On the germination of the oospore, 4 zoospores are formed (Fig. 54 F). They occur only in fresh or slightly brackish water. The division of the cells takes place in quite a peculiar and unusual manner. At the upper end of the cell which is about to divide, a ring-shaped thickening of soft cellulose is formed transversely round the wall; the cell-nucleus of the mother-cell and the protoplasm then divide by a transverse wall into two portions of similar size, and the cell-wall bursts transversely along the central line of the thickened ring. The cell-wall thus divides into two parts—the upper one short, the “cap,” and the lower one much longer, the “sheath.” The portions of the original cell-wall now separate from each other, the cellulose ring extending, and supplying an additional length of cell-wall between them. The cap and sheath will project a little in front of the piece thus inserted. The dividing wall between the two new cells is formed near to the uppermost edge of the sheath, and gradually becomes thicker and firmer. The inserted piece of wall forms the larger part of the wall of the upper cell: the remainder is formed by the cap. This mode of division is repeated exactly in the same way, and new caps are formed close below the first one, one for every division.

Fig. 54.A Œdogonium ciliatum. A Female plant with three oogonia (og) and dwarf-males (m). B An oogonium with spermatozoid (z) seen entering the oosphere (o) having passed through an aperture near the summit of the oogonium; m dwarf-male. C Ripe oospore. D Œdogonium gemelliparum. F Portion of a male filament from which spermatozoids (z) are emerging. E Portion of filament of Bulbochæte; the upper oogonium still encloses the oospore, in the central one the oospore is escaping while the lower one is empty. F Four zoospores developed from an oospore. G Zoospore germinating.

Fertilisation takes place in the following way. The oogonium is a large ellipsoidal, swollen cell (og, in Fig. 54 A), whose contents are rounded off into an oosphere with a colourless receptive-spot (see B); an aperture is formed in the wall of the oogonium, through which the spermatozoids are enabled to enter (B). The spermatozoids are produced either directly, as in D (in pairs), in basal cells of the filament, or indirectly. In the latter case a swarmspore (androspore) is formed which comes to rest, attaches itself to an oogonium, germinates, and gives rise to a filament of a very few cells—dwarf-male (A, B, m). The spermatozoids are formed in the upper cell of the dwarf-male (m), and are set free by the summit of the antheridium lifting off like a lid. On the germination of the oospore (C), which takes place in the following spring, 4 zoospores are produced (F) (i.e. the sexual generation); these swarm about for a time, and ultimately grow into new filaments.

Fig. 55.Coleochæte pulvinata. A A portion of a thallus with organs of reproduction; a oogonium before, b after fertilisation; c an antheridium, closed; d open, with emerging spermatozoid. B Ripe oogonium, with envelope. C Germination of the oospore. D Zoospore. E Spermatozoid.

Order 7. Coleochætaceæ. The thallus is always attached, and of a disc- or cushion-shape, formed by the dichotomous branching of filaments of cells united in a pseudo-parenchymatous manner. Each cell has only one nucleus. Asexual reproduction by zoospores with 2 cilia (Fig. 55 D), which may arise in all the cells. Sexual reproduction by oogamous fertilisation. The spermatozoids resemble the swarmspores, but are smaller (E), and originate singly (in the species figured) in small conical cells (c, d in A). The oogonia are developed at the extremities of certain branches: they are bottle-shaped cells with very long and thin necks (trichogyne), open at the end (a in A); at the base of each oogonium is a spherical oosphere. The spermatozoids reach the oosphere through the trichogyne, or through an aperture in the wall when the trichogyne is absent, and fertilisation having taken place, the oogonium becomes surrounded by a cell-layer (envelope), which grows out from the cells near its base (b in A), and in this way a kind of fruit is formed (B) (spermocarp, cystocarp).

The oospore, next spring, divides and forms a parenchymatous tissue (homologous with the Moss-sporophyte); this bursts open the envelope (C), and a zoospore (homologous with the spores of the Moss-capsule) arises in each of the cells, and produces a new Coleochæte. We have then, in this case, a still more distinct alternation of generations than in Œdogonium. Only one genus, Coleochæte, is known, but it contains several species, all living in fresh water.

Order 8. Cladophoraceæ. This order is probably derived from the Ulothricaceæ. The thallus consists of a single, unbranched or branched filament, generally with an apical cell. The cells have each 2 or more nuclei. Asexual reproduction by zoospores with 2 or 4 cilia, and by akinetes. Conjugation of gametes with 2 cilia is found in some genera. They occur in salt as well as in fresh water. The principal genera are: Urospora, Chætomorpha, Rhizoclonium, Cladophora; of the last named genus the species C. lanosa and C. rupestris are common in salt water; C. fracta and C. glomerata in fresh water.