1911 Encyclopædia Britannica/Algae/Chlorophyceae

II. Chlorophyceae.—This group includes those algae in which the green colouring matter, chlorophyll, is not accompanied by a second colouring matter, as it is in other groups. It consists of three subdivisions—Conjugatae, Euchlorophyceae and Characeae. Of these the first and last are relatively small and sharply defined families, distinguished from the second family, which forms the bulk of the group, by characters so diverse that their inclusion with them in one larger group can only be justified on the ground of convenience. Chlorophyceae include both marine and freshwater plants.

Euchlorophyceae in their turn have been until recently regarded as made up of the three series of families—Protococcales, Confervales and Siphonales. As the result of recent investigations by two Swedish algologists, Bohlin and Luther, it has been proposed to make a re-classification of a far-reaching nature. Algae are withdrawn from each of the three series enumerated above and consolidated into an entirely new group. In these algae, the colouring matter is said to be yellowish-green, not strictly green, and contained in numerous small discoid chromatophores which are devoid of pyrenoids. The products of assimilation are stored up in the form of a fatty substance and not starch. A certain inequality in the character of the two cilia of the zoospores of some of the members of the group has earned for it the title Heterokontae, from the Greek κοντός, a punting-pole. In consonance with this name, its authors propose to re-name the Conjugatae; Akontae and Oedogoniaceae with a chaplet of cilia become Stephanokontae, and the algae remaining over in the three series from which the Heterokontae and Stephanokontae are withdrawn become Isokontae. Conjugatae, Protococcales and Characeae are exclusively freshwater; Confervales and Siphonales are both freshwater and marine, but the latter group attains its greatest development in the sea. Some Chlorophyceae are terrestrial in habit, usually growing on a damp substratum, however. Trentepohlia grows on rocks and can survive considerable desiccation. Phycopeltis grows on the surface of leaves, Phyllobium and Phyllosiphon in their tissues. Gomontia is a shell-boring alga,

Dermatophyton grows on the carapace of the tortoise and Trichophilus in the hairs of the sloth. Certain Protococcales and Confervales exist as the gonidia of the lichenthallus.

A. Chlamydomonas sp., unicellular; chr., chromatophore; p., pyrenoid;
n., nucleus; p.v., pulsating vacuoles; e.s., eyespot.
B₁. Volvox sp., with a, antheridia, and o, oogonia.
B₂. Volvox sp., surface view of a single cell showing connexions.
C. Pandorina sp., a 16-celled colony.
D. Hydrodictyon, a single mesh surrounded by 6 cells.
E. Microspora sp., showing H-pieces in the wall.
F.Entoderma sp., endophytic in Ectocarpus.
G. Coleochaete sp., growing as a plate.
H. Oedogonium sp., intercalated growth by insertion of new piece (a) leaving caps.
K. Struvea sp., showing branches forming a net-work.
L.Caulerpa sp., showing portion of axis with leaf-like and root-like appendages.
M₁. Chara sp., axis with leaf-like appendages and a branch.
M₂. Chara sp., apical region.
N. Botrydium, a simple siphonaceous alga with root-like attachment.
O. Acetabularia Mediterranea, mushroom-like calcareous siphonaceous alga.

A, C, E, F, G, H, K, L, M₁, M₂ from Engler and Prantl, Pflanzenfamilien, by permission of William Engelmann; B₁, N from Vines, Students’ Text Book of Botany,
by permission of Swan Sonnenschein and Co.; B₂, D, O from Oltmanns, Morphologie u. Biologie der Algen, by permission of Gustav Fischer.)

The thallus is of more varied structure in this group than in any other. In the simplest case it may consist of a single cell, which may remain free during the whole of the greater part of its existence, or be loosely aggregated together within a common mucilage, or be held together by the adhesion of the cell-walls at the surface of contact. The aggregations or colonies, as they are termed, may assume the form of a plate, a ring, a solid sphere, a hollow sphere, a perforate sphere, a closed net, or a simple or branched filament. It is not easy in all cases to draw a distinction between a colony of plants and a multicellular individual. In a Volvox sphere, for example, there is a marked protoplasmic continuity between all the cells of the colony. The Ulvaceae, the thallus of which consists of laminae, one or more cells thick, or hollow tubes, probably represent a still more advanced stage in the passage of a colony into a multicellular plant. Here there is some amount of localization of growth and distinction of parts. It is only in such cases as Volvox and Ulvaceae that there is any pretension to the formation of a true parenchyma within the limits of the Chlorophyceae. In the whole series of the Confervales, the thallus consists of filaments branched or unbranched, attached at one extremity, and growing almost wholly at the free end. The branches end in fine hairs in Chaetophoraceae. In Coleochaetaceae the branches are often welded into a plate, simulating a parenchyma. In all Conjugatae and most Protococcales, and in the bulk of the Confervales, the thallus consists of a cell or cells, the protoplast of which contains a single nucleus. In Hydrodictyaceae, Cladophoraceae, Sphaeropleaceae and Gomontiaceae this is no longer the case. Instead of a single relatively large nucleus, each cell is found to contain many small nuclei, and is spoken of as a coenocyte. This character becomes still more pronounced in the large group of the Siphonales. Valoniaceae and Dasycladaceae are partially septate, but elsewhere no cellulose partitions occur, and the thallus is more or less the continuous tube from which the group is named. Yet the siphonaceous algae may assume great variety of form and reach a high degree of differentiation. Protosiphon and Botrydium, on the one hand, are minute vesicles attached to muddy surfaces by rhizoids; Caulerpa, on the other, presents a remarkable instance of the way in which much the same external morphology as that of cormophytes has been reached by a totally different internal structure. Many Siphonales are encrusted with lime like Corallina among Red Algae. Penicillus is brush-like, Halimeda and Cymopolia are jointed, Acetabularia has much the same external form as an expanded Coprinus, Neomeris simulates the fertile shoot of Equisetum with its densely packed whorled branches, and in Microdictyon, Anadyomene, Struvea and Boodlea the branches, spreading in one plane become bound together in a more or less close network. Characeae are separated from other Chlorophyceae by a long interval, and present the highest degree of differentiation of parts known among Green Algae. Attached to the bottom of pools by means of rhizoids, the thallus of Characeae grows upwards by means of an apical cell, giving off whorled appendages at regular intervals. The appendages have a limited growth; but in connexion with each whorl there arise, singly or in pairs, branches which have the same unlimited growth as the main axis. There is thus a close approach to the external morphology of the higher plants. The streaming of the protoplasm, known elsewhere among Chlorophyceae, is a conspicuous feature of the cells of Characeae.

The Chlorophyceae excel all other groups of algae in the magnitude and variety of form of the chlorophyll-bodies. In Ulva and Mesocarpus the chromatophore is a single plate, which in the latter genus places its edge towards the incident light; in Spirogyra they are spiral bands embedded in the primordial utricle; in Zygnema they are a pair of stellate masses, the rays of which branch peripherally; in Oedogonium they are longitudinally-disposed anastomosing bands; in Desmids plates with irregular margins; in Cladophora polyhedral plates; in Vaucheria minute elliptical bodies occurring in immense numbers. Embedded in the chromatophore, much in the same way as the nucleus is embedded in the cytoplasm, are the pyrenoids. Unknown in Cyanophyceae and Phoeophyceae, known only in Bangiaceae and Nemalion among Rhodophyceae, they are of frequent occurrence among Chlorophyceae, excepting Characeae. Sometimes several pyrenoids occur in each chloroplast, as in Mesocarpus and Spirogyra; sometimes only an occasional chloroplast contains pyrenoid at all, as in Cladophora. The pyrenoid seems to be of proteid nature and gelatinous consistency, and to arise as a new formation or by division of pre-existing pyrenoids. When carbon-assimilation is active, starch-granules crowd upon the surface of the pyrenoid and completely obscure it from view.

Special provision for vegetative multiplication is not common among Chlorophyceae. Valonia and Caulerpa among Siphonales detach portions of their thallus, which are capable of independent growth. In Caulerpa no other means of multiplication is as yet known. In Characeae no fewer than four methods of vegetative reproduction have been described, and the facility with which buds and branches are in these cases detached has been adduced as an evidence of affinity with Bryophyta, which, as a class, are distinguished by their ready resort to vegetative reproduction.

With regard to true reproduction, which is characterized by the formation of special cells, the group Euchlorophyceae is characterized by the production of zoospores (Gr. ζῷον, animal, σπορά, seed); that is to say, cells capable of motility through the agency of cilia. Such ciliary motion is known in the adult condition of the cells of Volvocaceae, but where this is not the case the reproductive cells are endowed with motility for a brief period. The zoospore is usually a pyriform mass of naked protoplasm, the beaked end of which where the cilia arise is devoid of colouring matter. A reddish-brown body, known as the eyespot, is usually situated near the limits of the hyaline portion, and in the protoplasm contractile vacuoles similar to those of lower animals have been occasionally detected. The movement of the zoospore is effected by the lashing of the cilia and is in the direction of the beak, while the zoospore slowly rotates on its long axis at the same time. Usually two cilia are present; in Botrydium and Hydrodictyon only one is present; in certain species of Cladophora four; in Dasycladus a chaplet, and in Oedogonium a ring of many cilia. The so-called zoospore of Vaucheria is a coenocyte covered over with paired cilia corresponding in position to nuclei lying below. In all other cases, zoospores are uninucleate bodies. Zoospores arise in cells of ordinary size and form termed zoosporangia. In unicellular forms (Sphaerella) the thallus becomes transformed into a zoosporangium at the reproductive stage. In the zoosporangia of Oedogonium, Tetraspora and Coleochaete the contents become transformed into a single zoospore. In most cases repeated division seems to take place, and the final number is represented by some power of two. In coenocytic forms the zoospores would seem to arise simultaneously, probably because many nuclei are already present. The escape of zoospores is effected by the degeneration of the sporangial wall (Chaetophora), or by a pore (Cladophora), a slit (Pediastrum), or a circular fracture (Oedogonium). Zoospores are of two kinds: (1) Those which come to rest and germinate to form a new plant; these are asexual and are zoospores proper. (2) Those which are unable to germinate of themselves, but fuse with another cell, the product giving rise to a new individual; these are sexual and are zoogametes (Gr. ζῷον, animal, and γαμέτης, γαμετή, husband, wife). When two similar zoogametes fuse, the process is conjugation, and the product a zygospore (Gr. ζυγόν, yoke). Usually, however, only one of the fusing cells is a zoogamete, the other gamete being a much larger resting cell. In such a case the zoogamete is male, is called an antherozoid or spermatozoid, and arises in an antheridium; the larger gamete is an oosphere and arises in an oogonium. The fusion is now known as fertilization and the product is an oospore. Reproduction by conjugation is also known as isogamy, by fertilization as oogamy. When zoospores come to rest, a new cell is formed and germination ensues at once. When zygospores and oospores are produced a new cell-wall is also formed, but a long period of rest ensues. All investigation goes to show that an essential part of sexual union is the fusion of the two nuclei concerned. It is interesting to know, on the authority of Oltmanns, that when the oosphere is forming in the oogonium of Vaucheria, there is a retrocession of all the included nuclei but one. That the antherozoid of Vaucheria contains a single nucleus had been inferred before.

From a comparison of those Euchlorophyceae which have been most closely investigated, it appears probable that sexual reproductive cells have in the course of evolution arisen as the result of specialization among asexual reproductive cells, and that in turn oogamous reproduction has arisen as the result of differentiation of the two conjugating cells into the smaller male gamete and the larger male gamete. It would further appear that oogamous reproduction has arisen independently in each of the three main groups of Euchlorophyceae, viz. Protococcales, Siphonales and Confervales. Thus among Volvocaceae, a family of Protococcales, while in some of the genera (Chloraster, Sphondylomorum) no sexual union has as yet been observed, in others (Pandorina, Chlorogonium, Stephanosphaera, Sphaerella) conjugation of similar gametes takes place, in others still (Phacotus, Eudorina, Volvox) the union is of the nature of fertilization. No other family of Protococcales has advanced beyond the stage of isogamous reproduction. Again, among Siphonales only one family (Vaucheriaceae) had reached the stage of oogamy, although an incipient heterogamy is said to occur in two other families (Codiaceae, Bryopsidaceae). Elsewhere among Siphonales, in those cases where reproductive cells are known, the reproduction is either isogamous or asexual. Among Confervales there is no family in which sexual reproduction—isogamy or oogamy—is not known to occur among some of the component species, and as many as four families (Cylindrocapsaceae, Sphaeropleaceae, Oedogoniaceae, Coleochaetaceae) are oogamous. On these, as well as other grounds, Confervales are regarded as having attained to the highest rank among Euchlorophyceae. Although the phenomena attending isogamous and oogamous reproduction respectively are essentially the same in all cases, slight variations in both instances appear in different families, attributable doubtless to the independent origin of the process in different groups. Thus, although isogamy consists in typical cases of a union of naked motile gametes by a fusion which begins at the beaked ends, and results in the formation of an immotile spherical zygote surrounded by a cell-wall, in Leptosira it is noticeable that the fusion begins at the blunt end; in a species of Chlamydomonas the two gametes are each included in a cell-wall before fusion; and in many cases the zygote retains for some time its motility with the double number of cilia. Again, in oogamous reproduction, while in general only one oosphere is differentiated in the oogonium, in Sphaeroplea several oospheres arise in each oogonium; and while the oospheres usually contract away from the oogonial wall, acquiring for themselves a new cell-wall after fertilization, in Coleochaete the oosphere remains throughout in contact with the oogonial wall. The oosphere is in all cases fertilized while still within the oogonium, the antherozoids being admitted by means of a pore. There is usually distinguishable upon the surface of the oosphere an area free from chlorophyll, known as the receptive spot, at which the fusion with the antherozoid takes place; and in many cases, before fertilization, a small mucilaginous mass has been observed to separate itself off from the oosphere at this point and to escape through the pore. In Coleochaete the oogonial wall is drawn out into a considerable tube, which is provided with an apical pore, and this tube has a somewhat similar appearance to the imperforate trichogyne of Florideae to be hereafter described. In certain species of Oedogonium minute male plantlets, known as dwarf males, become attached to the female plant in the neighbourhood of the oogonia, thus facilitating fertilization. Indeed the genus Oedogonium exhibits a high degree of specialization in its reproductive system, considering that its thallus has not advanced beyond the stage of an unbranched filament.

Many Euchlorophyceae are endowed with both asexual and sexual reproduction. Such are Coleochaete, Oedogonium, Cylindrocapsa, Ulothrix, Vaucheria, Volvox, &c. In others only the asexual method is yet known. When a species resorts to both methods, it is generally found that the asexual method prevails in the early part of the vegetative period and the sexual towards the close of that period. This is in consonance with the facts already mentioned that zoospores germinate forthwith, and that the sexually-produced cell or zygote enters upon a period of rest. It is known that zoogametes, which usually conjugate, may, when conjugation fails, germinate directly (Sphaerella). In rare cases the oosphere has been known to germinate without fertilization (Oedogonium, Cylindrocapsa). The germination of a zygospore or oospore is effected by the rupture of an outer cuticularized exosporium; then the cell may protrude an inner wall, the endosporium, and grow out into the new plant (Vaucheria), or the contents may break up into a first brood of zoospores. It is held that in Coleochaete a parenchyma results from the division of the oospore, from each cell of which a zoospore arises.

Reproduction is also effected among Euchlorophyceae by means of aplanospores and akinetes. Aplanospores would seem to represent zoospores arrested in their development; without reaching the stage of motility, they germinate within the sporangium. Akinetes are ordinary thallus cells, which on account of their acquisition of a thick wall are capable of surviving unfavourable conditions. Both aplanospores and akinetes may germinate with or without the formation of zoospores at the initial stage.

Among Conjugatae reproduction is effected solely by means of conjugation of what are literally aplanospores. Among those Desmidiaceae which live a free life, two plants become surrounded by a common mucilage, in which they lie either parallel (Closterium) or crosswise (Cosmarium). Gaps then appear in the apposed surfaces, usually at the isthmus; the entire protoplasts either pass out to melt into one another clear of the old walls, or partly pass out and fuse without complete detachment from the old walls. Among colonial Desmidiaceae, the break-up of the filament is a preliminary to this conjugation; otherwise the process is the same. The zygospore becomes surrounded with its own wall, consisting finally of three layers, the outer of which is furnished with spicular prominences of various forms. In Zygnemaceae there is no dissolution of the filaments, but the whole contents of one cell pass over by means of a conjugation-tube into the cavity of a cell of a neighbouring filament, where the zygospore is formed by the fusion of the two protoplasts.

Fig. 3.—Chlorophyceae, variously magnified.

A. Spirogyra sp., in conjugation.	E3. Coleochaete sp., zoospore.
B. Zoospore of Pandorina. B2 3 4,    stages of conjugation.	F1 2 3. Protosiphon, conjugation of    zoogametes.
C. Ulothriz sp., zoospores escaping.    C2 3, stages of conjugation.	G. Derbesia sp., zoospore with    chaplet of cilia
D1. Oedogonium sp., oogonium    at a moment of fertilization    with dwarf male attached.	H1 Chara sp., oogonium and    antheridium at a node on    a lateral appendage.
D2. Oedogonium sp., zoospore    with crown of cilia.	H2. Chara sp., antherozoid.
E1. Coleochaete sp., with anthe-    ridia and an oogonium.	K1. Vaucheria sp., oogonium and    antheridium before fertilization.
E2. Coleochaete sp., fertilized egg    with investment of filaments.	K2. Vaucheria sp., after fertilization.
(A from Cooke, British Freshwater Algae, by permission of Kegan Paul, Trench, Trübner and Co.; C, E, F, G, H, K from Engler and Prantl, by permission of Wilhelm Engelmann; B1 from Vines, by permission of Swan Sonnenschein and Co.; B2, D from Oltmanns, by permission of Gustav Fischer.)

In these cases the activity of one of the gametes, and the passivity of the other, is regarded as evidence of incipient sex. In Sirogonium there is cell-division in the parent-cell prior to conjugation; and as two segments are cut off in the case of the active gamete, and only one in the case of the passive gamete, there is a corresponding difference of size, marking another step in the sexual differentiation. In Zygogonium, although no cell-division takes place, the gametes consist of a portion only of the contents of a cell, and this is regularly the case in Mesocarpaceae, which occupy the highest grade among Conjugatae. Some Zygnemaceae and Mesocarpaceae form either a short conjugating tube, or none at all, but the filaments approach each other by a knee-like bend, and the zygospore is formed at the point of contact, often being partially contained within the walls of the parent-cell. It would seem that in some cases the nuclei of the gametes remain distinct in the zygospore for a considerable time after conjugation. It is probable that in all cases nuclear fusion takes place sooner or later. In Zygnemaceae and Mesocarpaceae the zygospore, after a period of rest, germinates, to form a new filamentous colony; in Desmidiaceae its contents divide on germination, and thus give rise to two or more Desmids. Gametes which fail to conjugate sometimes assume the appearance of zygospores and germinate in due course. They are known as azygospores.

The reproduction of Characeae is characterized by a pronounced oogamy, the reproductive organs being the most highly differentiated among Chlorophyceae. The antheridia and oogonia are formed at the nodes of the appendages. The oogonium, seated on a stalk cell, is surrounded by an investment consisting of five spirally-wound cells, from the projecting ends of which segments are cut off, constituting the so-called stigma. The oosphere is not differentiated within the wall of the oogonium, but certain cells known as wendungszellen, the significance of which has given rise to much speculation, are cut off from the basal portion of the parent-cell during its development. The antheridia are spherical orange-coloured bodies of very complex structure. The antherozoid is a spirally-coiled thread of protoplasm, furnished at one end with a pair of cilia. It much more resembles the antherozoids of Bryophyta and certain Pteridophyta than any known among other algae. The fertilized egg charged with food reserves rests for a considerable period, surrounded by its cortex, the whole having assumed a reddish-brown colour. On germination it gives rise to a row of cells in which short (nodal) and long (internodal) cells alternate. From the first node arise rhizoids; from the second a lateral bud, which becomes the new plant. This peculiar product of germination, which intervenes between the oospore and the adult form, is the proembryo. It will be remembered that in Musci, the asexual spore somewhat similarly gives rise to a protonema, from which the adult plant is produced as a lateral bud. The proembryonic branches of Characeae, one of the means of vegetative reproduction already referred to, are so called because they repeat the characters of the proembryo.

Before leaving the Chlorophyceae, it should be mentioned that the genus Volvox has been included by some zoologists (Bütschli, for example) among Flagellata; on the other hand, certain green Flagellata, such as Euglena, are by some botanists (for example, van Tieghem) among unicellular plants. A similar uncertainty exists with reference to certain groups of Phaeophyceae, and the matter will thus arise again.

A census of the Chlorophyceae is furnished below:—

1. Confervoideae—12 families, 77 genera, 1021 species.
2. Siphoneae—9 families, 26 genera, 271 species.
3. Protococcoideae—2 families, 90 genera, 342 species.
4. Conjugateae—2 families, 33 genera, 1296 species.
   (De Toni’s Sylloge Algarum, 1889.)
5. Characeae—2 families, 6 genera, 181 species.
   (Engler and Prantl’s Pflanzenfamilien, 1897.)

III. Phaeophyceae.—The Phaeophyceae are distinguished by the possession of a brown colouring matter, phycophaein, in addition to chlorophyll. They consist of the following groups:—Fucaceae, Phaeosporeae, Dictyotaceae, Cryptomonadaceae, Peridiniaceae and Diatomaceae. Of these the first three include multicellular plants, some of them of great size; the last three are unicellular organisms, with little in common with the rest excepting the possession of a brown colouring matter. Fucaceae and Phaeosporeae are doubtless closely allied, and to these Dictyotaceae may be joined, though the relationship is less close. They constitute the Euphaeophyceae, and will be dealt with in the first place.

Euphaeophyceae are almost exclusively marine, growing on rocks and stones on the coast, or epiphytic upon other algae. In tidal seas they range from the limits of high water to some distance beyond the low-water line. On the British coasts zones are observable in passing from high to low water mark, characterized by the prevalence of different species, thus:—Pelvetia canaliculata, Fucus platycarpus, Fucus vesiculosus, Ascophyllum nodosum, Fucus serratus, Laminaria digitata. Some species are minute filamentous plants, requiring the microscope for their detection; others, like Lessonia, are of considerable bulk, or, like Macrocystis, of enormous length. In Fucaceae, Dictyotacea, and in Laminariaceae and Sphacelariaceae, among Phaeosporeae, the thallus consists of a true parenchyma; elsewhere it consists of free filaments, or filaments so compacted together, as in Cutleriaceae and Desmarestiaceae, as to form a false parenchyma. In Fucaceae and Laminariaceae the inner tissue is differentiated into a conducting system. In Laminariaceae the inflation of the ends of conducting cells gives rise to the so-called