1911 Encyclopædia Britannica/Algae/Rhodophyceae

IV. Rhodophyceae, or Florideae.—The members of this group are characterized by the possession of a red colouring matter, phycoerythrin, in addition to chlorophyll. There is, however, a considerable amount of difference in the shades of red which mark different species. The brightest belongs to those species which grow near low-water mark, or under the shade of larger algae at higher levels; species which grow near high-water mark are usually of so dark a hue that they are easily mistaken for brown seaweeds. Rhodophyceae are mostly marine, but not exclusively so. Thorea, Lemanea, Tuomeya, Stenocladia, Batrachospermum, Balbiania are genera belonging entirely to fresh water; and Bangia, Chantransia, Caloglossa, Bostrychia and Delesseria contain each one or more freshwater species. Most of the larger species of marine Rhodophyceae are attached by means of a disc to rocks, stones or shells. Many are epiphytic on other algae, more especially the larger Phaeophyceae and Rhodophyceae. As in the case of epiphytic brown seaweeds, the rhizoids of the epiphyte often penetrate the substance of the supporting alga. Some Red Algae find a home in the gelatinous substance of Flustra, Alcyonidium and other polyzoa, only emerging for the formation of the reproductive organs. Some are perforating algae and burrow into the substance of molluscan shells, in company with certain Green and Blue-green Algae. Some species belonging to the families Squamariaceae and Corallinaceae grow attached through their whole length and breadth, and are often encrusted with lime. The forms which grow away from the substratum vary greatly in external configuration. In point of size the largest cannot rival the larger Brown Algae, while the majority require the aid of the microscope for their investigation.

No unicellular Rhodophyceae are known, although a flagellate organism, Rhodomonas, has recently been described as possessed of the same red colouring matter. If the sub-group, Bangiaceae, be excluded, they may be said to consist exclusively of branched filaments. Growth in these cases takes place by means of an apical cell, from which successive segments are cut off by means of a transverse wall. The segment so cut off does not usually divide again by means of a transverse wall, nor indeed by a longitudinal wall which passes through the organic axis of the cell. New cells may be cut off laterally, which become the apical cells of branches. When the new cells grow no further, but constitute a palisading round the central cell covering its whole length, the condition is reached which characterizes the species of Polysiphonia, the “siphons” of which may be regarded as one-celled branches. To the law that no subsequent transverse division takes place in segments cut off from the apical cell, there seem to be two exceptions: first, the calcareous genus Corallina, in the pliable joints of which intercalated division occurs; and, second, the Nitophylleae, in which, moreover, median longitudinal division of axial cells is said to occur. Like the Fungi, therefore, the Red Algae consist for the most part of branched filaments, even where the thallus appears massive to the eye, and, as in the case of Fungi, this fact is not inconsistent with a great variety of external morphology. In the great majority the thallus is obviously filamentous, as in some species of Callithamnion. In other species of that genus an apparent cortication arises by the downward growth of rhizoids, which are retained within the gelatinous wall of the axial cells. In Batrachospermum the whole system of branches are retained within a diffluent gelatinous substance derived from the outer layers of the cell-walls. In other cases the mucilage is denser and the branches more closely compacted (Helminthora). In such cases as Lemanea, the terminal cells of the lateral branches form a superficial layer which has all the appearance of a parenchyma when viewed from the surface. In Champia and allied genera, the cylindrical axis is due not to the derivatives of one axial filament, but of several, the growth of which is co-ordinated to form a septated tube. The branching of the thallus, which meets the eye in all these cases, is due to the unlimited growth of a few branches. When such a lateral branch overtops the main axis whose growth has become limited, as in Plocamium and Dasya, a sympodium is formed. For the most part the branching is monopodial. Besides the differentiation into holdfast and shoot, and into branches of limited and branches of unlimited growth, there appear superficial structures of the nature of hairs. These are for the most part long, thin-walled, unicellular and colourless, and arise from the outer cells of the pseudo-cortex, or from the terminal cells of branches when the filaments are free. Among Rhodomelaceae, hair-like structures of a higher order are known. These arise from the axial cell, and are multicellular and branched. They soon fall off, and it is from the persistent basal cell that the branches of unlimited growth arise. Upon them also the reproductive organs arise in this family. It is not surprising, therefore, that they have been regarded as the rudiments of leaves. In Iridaea the thallus is an entire lamina; in Callophyllis a lobed lamina; in Delesseria it is provided with midrib and veins, simulating the appearance of a leaf of the higher plants; in Constantinea the axis remains cylindrical, and the lateral branches assume the form of leaves. In the compact thalli a secondary development often takes place by the growth of rhizoid-like internal filaments. They present a hypha-like appearance, running longitudinally for considerable distances. It is not difficult in such compact species to distinguish between superficial cells, whose chief function is assimilation, subjacent cells charged with reserve material, and a core of tissue engaged in the convection of elaborated material from part to part.

An interesting feature of the minute anatomy of Euflorideae, as the Red Algae, exclusive of the Bangiaceae, have been termed, is the existence of the so-called Floridean pit. When a cell divides it is found that there remains in the middle of the new wall a single large circular pit, which persists throughout the life of the cells, becoming more and more conspicuous with the progress of the thickening of the wall. These pits serve to indicate the genetic relationship of adjacent cells, when they form a compact pesudo-parenchyma, notwithstanding the fact that somewhat smaller secondary pits appear later between any contiguous cells. Protoplasmic continuity has been observed in the delicate membrane closing the pit.

Vegetative multiplication occurs only sparingly in Rhodophyceae. Melobesia callithamnioides gives rise to multicellular propagula; Griffithsia corallina is said to give rise to new individuals, by detaching portions of the thallus from the base of which new attachment organs have already arisen. The spores of Monospora are by some regarded as unicellular propagula. Reproduction is both asexual and sexual. It is noteworthy that although all the members of the group are aquatic no zoospores are produced, a negative character common to them and the Blue-green Algae. As a rule the asexual cells, and the male and female sexual cells arise upon different plants, so that the species may be said to be trioecious. Numerous exceptions, however, occur. Thus in Lemaneaceae asexual spores are unknown; in Batracho-spermum, Bonnemaisonia and Polysiphonia byssoides both kinds of sexual cells appear on the same plant; and in some cases the asexual cells may occur in conjunction with either the male or female sexual cells. The asexual cells are termed tetraspores on account of the usual occurrence of four in each sporangium. What may be termed monospores, bispores and octospores, however, are not unknown. The sporangia may be terminal or intercalated. When they are confined to special branches such branches are spoken of as stichidia. The tetraspores may arise by the simultaneous division of the contents of a sporangium, when they are arranged tetrahedrally, or they may arise by two successive divisions, in which case the arrangement may be zonate when the spores are in a row, or cruciate when the second divisions are at right angles to the first, or tetrahedral when the second divisions are at right angles to the first and also to one another. Tetraspores are at first naked, but soon acquire a cell-wall and germinate without a period of rest. The male sexual cells are produced singly in the terminal cells of branches. They are spoken of as spermatia. Great numbers of antheridia are usually crowded together, when the part is distinguishable by the absence of the usual red colour. In Polysiphonia they cover the joints of the so-called leaves; in Chondria they arise on flattened disks; in the more massive forms they arise in patches on the ordinary surface; in a few cases (Gracilaria, Corallina, Galaxaura) they line the walls of conceptacle-like depressions. The female sexual cell is represented by the contents of a cell which is terminal on ordinary or specialized branches. This is the carpogonium; it consists of a ventral portion which contains a nucleus, but in which no oosphere is differentiated, and an elongated tubular portion known as the trichogyne, into which the cytoplasm extends. Fertilization is effected by the passive convection of a spermatium from the antheridium to the trichogyne, to which it adheres, and to which it passes over its nucleus through an open communication set up at the point of contact. The nucleus then passes down the trichogyne and fuses with that of the egg. This fusion has been observed by Wille in Nemalion multifidim, and by Schmidle in Batrachospermum. It is singular that in the last-named species two nuclei occur regularly in the spermatium. The ventral portion of the carpogonium may be imbedded deep in the thallus in the massive species; the trichogyne, however, always reaches the surface. The first effect of fertilization is the occlusion of the trichogyne from the fertilized carpogonium. The subsequent course of development is characteristic of the Florideae. The carpogonium germinates forthwith, drawing its nourishment almost wholly from the parent plant. The ultimate product in all cases is a number of carpospores, but before this stage is reached the development is different in different subgroups. In Batrachospermum filaments arise from the carpogonium on all sides; in Chantransia and Scinaia on one side only; in Helminthora the filaments are enclosed in a dense mucilage; in Nemalion, prior to the formation of the filaments, a sterile segment is cut off below. In all these cases, however, the end-cells of the filaments each give rise to a carpospore, and the aggregate of such sporiferous filaments is a cystocarp. Again, in the family of the Gelidiaceae, the single filament arising from the carpogonium grows back into the tissue and preys upon the cells of the axis and larger branches, after which the end-cells give rise to carpospores and a diffused cystocarp is formed. In the whole group of the Cryptonemiales the parasitism becomes more marked still. The filaments arising from the carpogonia grow into long thin tubes, which fuse with special cells rich in protoplasm contents; and from these points issue isolated tufts of sporogenous filaments, several of which may form the product of one fertilized female cell. In Naccaria, one of the Gelidiaceae, it is observable that the ooblastema filament, as the tube arising from the fertilized carpogonium has been called, fuses completely with a cell contiguous to the carpogonium before giving rise to the foraging filaments already refered to. This is also the case among Cryptonemiales. In a whole series of Red Algae, the existence of a highly specialized auxiliary cell in the neighbourhood of the carpogonium is a characteristic feature. In the Gigartinales it is already differentiated previous to fertilization; in Rhodymeniales it arises subsequent to fertilization. In the Gigartinales, the filaments which arise from the auxiliary cell may spread and give rise to isolated tufts of sporogenous filaments, as in the Cryptonemiales. In the Rhodymeniales a single tuft arises directly from' the auxiliary cell. The carpospores are in all cases bright red naked masses of protoplasm when first discharged. They soon acquire a cell-wall, and germinate without a period of rest. When the cystocarps or segments of cystocarps are formed in the substance of a thallus, the site is marked merely by a swelling of the substance. When the cystocarp is produced externally, it may form a berry-like mass without an envelope, in which case it is known as a favella. In Rhodomelaceae there is a special urn-shaped envelope surrounding the sporogenous filaments. This is a ceramidium.

The attachment of the cell of an ooblastema filament to a cell of the thallus may be effected by means of a minute pore, or the two cells may fuse their contents into one protoplasmic mass. In the latter case, and especially where the union is with a special auxiliary cell, it is of importance to know what happens to the nuclei of the fusing cells. Schmitz was of opinion that in the cases of open union there occurred a fusion of nuclei similar to that which occurs in the sexual union of two cells. He founded his generalization to a large extent upon the observation that in Gloeosiphonia capillaris two cells completely fuse, and that only one nucleus can be detected in the fused mass. Oltmanns has recently re-investigated the phenomena in this plant, among others, and has shown that the nucleus of the cell which is being preyed upon recedes to the wall and gradually atrophies. The nucleus of the ooblastema filament dominates the

mass and from it all the nuclei of the carpospores are thus derived. There thus seems to be no justification for believing, as Schmitz taught, that a second sexual act occurs in the life-cycle of these Florideae.

Fig. 5.—Rhodophyceae, variously magnified.
A. Polysiphonia sp., apical region showing leading cell and cutting	K. Nemalian sp., carpogonial and antheridial branches.
off of pericentral cell.	L. Batrachospermum sp., trichogyne with spermatia attached;
B. Polysiphonia sp., transverse section through a branch, and at	carpospores arising from fertilized carpogonium.
a, mother-cell of tetraspores.	M. Polysiphonia sp., antheridium. .
C. Lomentaria sp., apex showing growth in length through	N. Constantinea sp., with flattened leaf-like appendages.
coordinated growth of many filaments.	O. Dudresnaya coccinea, fusion of ooblastema filaments with auxiliary
D. Delesseria sp., showing apical region with leading cell.	cells; a is an axial cell in transverse section with four appendages.
E. Chrysymenia uvaria, axis with swollen leaf-like appendages.	P. Callithamnion corymbosum, a joint cell with carpogonial branch
F. Polyzonia sp., branch with leaf-like branches of limited growth.	and a, b, two auxiliary cells.
G. Callithamnion sp., tetrasporangium with spores arranged in a tetrad.	Q. Callithamnion corymbosum, fusion of products of fertilization with
H. Corallina sp., tetrasporangia with zonate arrangement of	auxiliary cells, the nuclei of which a and b retire to the wall.
tetraspores.	R. Polysiphonia, sp., section through young cystocarp.
(A, C, D, E, F, G, H, K, L, M, P, Q, from Oltmanns, by permission of Gustav Fischer; B, N, O, R, from Engler and Prantl, by permission of Wilhelm Engelmann.)

The Bangiales are a relatively small group of Red Algae, to which much of the description now given does not apply. Structurally they are either a plate of cells, as in Porphyra, or filaments, as in Bangia. There is no exclusive apical growth, and the cells divide in all directions. The characteristic pit is also absent. Sexual and asexual reproduction prevail. The male cell is a spermatium, but the female cell bears no such receptive trichogyne as occurs in other Rhodophyceae. After fertilization the equivalent of the oospore divides directly to form a group of carpospores. There is thus a certain resemblance to Euflorideae, but sufficient difference to necessitate their being grouped apart. Fertilization by means of non-motile spermatia and a trichogyne are known among the Fungi in the families Collemaceae and Laboulbeniaceae.

A census of Rhodophyceae is furnished below: —

(1) Bangiaceae—4 families, 9 genera, 58 species.

(2) Nemalioninae—4 families, 33 genera, 343 species.

(3) Gigartininae—3 families, 54 genera, 409 species.

(4) Rhodymeninae—4 families, 92 genera, 602 species.

(De Toni’s Sylloge Algarum, 1897.)