Notes on Indian Red Alg^—I
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NOTES ON INDIAN RED ALG^—I B y M . s . B alakrishnan University Botany Laboratory, Madras-5 (With 21 Text-Figurcs) (Received for publication on December 16, 1957) D u r in g a special study of Indian red algae, the writer came across two interesting epiphytic red algae belonging to the family Ceramiaceae. These are described in the present paper. Gymnothamnion elegans (Schousboe) J. Agardh This alga was collected growing on Corynomorpha pnsmatica (Ag.) J. Ag. from Cape Comorin, South India. It forms small bright red tufts 1-Hcm. high on the basal portions of the host. The pro strate creeping portion consists of branched filaments fixed to the host by means of multicellular rhizoids at intervals (Text-Fig. 2). The cells of the creeping filaments are cylindrical, 10-12 x 30-45/x and thick walled. The cells giving rise to rhizoids are much shorter than the others and these also give rise to erect branches oiT the upper side (Text-Fig. 1). The erect branches are 10-15 mm. in height, plumose, usually simple, but occasionally pinnately branched; cells of the erect branches are up to 25 ^ long and 10-15/x wide. Generally they are slightly broader at the top (Text-Fig. 3). Each cell bears at the top a pair of opposite branchlets (pinnules) six to eight cells long, the cells measuring 10-12 n x 10-18 ft. The branchlets are distichously arranged and except in the youngest parts, almost of uniform length so that the erect branches are characteristically plumose and oblong or linear lanceolate in outline (Text-Fig. 1). Production of the branchlets starts right from the base (Text-Fig. 1); it is only in the older stages that the lower parts of the erect branches become bare, evidently due to the dropping off of older branchlets. Both the prostrate and erect filaments are totally ecorticate. The tetrasporangia are borne terminally on the branchlets. They are ellipsoid to ovate, 25-30/x wide and 3 0 - ^ /x long. The tetraspores are usually pyramidally arranged (Text-Fig. 3), but cruciate arrange ment is also frequently met with (Text-Fig. 4) (see also Bornet and Thuret, 1876, Plate X, Fig. 2). • The alga is dioecious. Spermatangia are borne in clusters on short multicellular accessory branches at the tips of the branchlets (Text- Figs. 7, 8). Each cell of these lateral branches functions as a sper- matangial mother cell and bears two to three spermatangia at the top, (Text-Fig. 11) (see also Bornet and Thuret, 1876, Plate X, Fig. 3). Text-Fios. 1-4. Gymnothamnion elegans. Fig. 1. Part of plant showing general habit and branching. Fig. 2. Part of creeping filament to show branching and rhizoids. Figs. 3-4. Portions of erect branches to show tetrasporangia. Development o f the procarp up to feriilizaiion The procarps are borne on the branchlets near ihe tips, usually on the subterminal cell. The sequence of development is as follows: The subterminal cell first cuts off a pericentral cell on the abaxial side. (Text-Fig. 5), and a little Jater, one more pericentral cell is cut off aterally (Text-Fig. 6, pc). The abaxial pericentral cell divideis further Tbxt-Fiqs. 5-12. Gymnothanwion elegans. Fig. 5. First stage in the for mation of the procarp. Fig. 6. A later stage showing the carpogonial branch initiated and the lateral pericentral cell formed. Figs. 7-8. Tips of branchlcts from a male plant showing early stages in the formation of the spermatangia. Fig. 9. Young procarp showing the four<elled carpogonial branch with the trichogyr.e just developing and the lateral sterile branch with its terwiral hair. Fig. 10. Early stage in gonimoblast formation. Fig. 11. Tip of branchlet from male plant show ing fully formed spermatangia. Fig. 12. Early post-fertilization stage showing division of the carpogonium into three ceUs and the auxiliary cell cut off from the supporting cell, (awe, auxiliary cell; c6r, carpogonial branch; cc, connecting cell; cpg; carpogonium; gon, gonimolobes; pc, lateral pericentral cell; sc, supporting cell; St, lateral sterile-branch.) to produce a four-celled carpogonial branch and a two to three-celled lateral sterile branch (Text-Fig. 9, S(). This lateral sterile branch is initiated when the carpogonial branch has started development and is two to three cells long. The terminal cell of the carpogonial branch develops into the carpogonium with a long trichogyne while that ol the lateral sterile branch develops into a long unicellular hair (Text-Figs. 9, 12). The lateral pericentral cell of the fertile segment usually remains undivided during later stages of development (Text-Figs. 10, pc\ 12, 13, 14), but it occasionally grows into a two to three-celled branch sometimes terminating in a hair (Text-Fig. 9, pd). It has also been observed that the lateral sterile branch is always formed on the side away from the lateral pericentral ceil (Text-Figs. 9, 12). Thus, at the time of fertilization, the supporting cell (Text-Fig. 9, 5c)carries; (1) a four-celled carpogonial branch on one side and (2) a two to three- celled lateral sterile branch on the other. The sequence of development described agrees with that described for the Mediterranean form of this alga (as Callithamnion eleganx Schousboe) by Bornet and Thuret (1876) and for Ptilota (Kylin, 1923) and Plumaria (Suneson, 1938; Drew, 1939). In all these cases the first formed abaxial pericentral cell produces a four-celled carpogonial branch and a two-celled lateral sterile branch. However, in Ptilota and Plumaria the fertile segment has two lateral pericentral cells, one on either side of the supporting cell, whereas in Gymnoihamnion there is only one lateral pericentral cell {cf. Bornet and Thuret, 1876. p. 33). Again, in Ptilota (Kylin, 1923) the terminal cell of the fertile branch- let, the two lateral pericentral cells of the procarpic segment and the lateral sterile cell of the procarp all develop into branches ending in terniinal hairs so that the fertile branchlet is crowned by a group of four hairs. In Plumaria on the other hand (Suneson, 1938; Drew, 1939) only the lateral sterile branch of the procarp always bears a ter minal hair; hair formation by one or more of the other branches is generally suppressed. It is, however, interesting to observe that Bornet and Thuret (1876, Plate X, Fig. 4), show a fertile branchlet of Gymnothamnion elegans from the Mediterranean crowned with five hairs. Development of the procarp after fertilization After fertilization, the trichogyne is first cut off. The fertilized carpogonium then divides to form three cells, the lowest of which functions as a connecting cell (sporogenous cell) and fuses with the auxiliary cell (Text-Fig. 12, cc). As in all Ceramiales, the auxiliary cell is cut off from the supporting cell, only after fertilization and lies in close proximity to the carpogonium (Text-Fig. 12, aux). After fusion of the carpogonium and the auxiliary cell is effected, the tricho gyne seems to vdther away. The carpogonium and the carpogonial branch cells also start degenerating. The auxiliary cell now divides into an upper larger and lower smaller cell. The former is the goni- moblast initial while the latter forms the so called ‘ foot cell.’ The gonimoblast initial, in its turn, gives rise to three to five gonimolobes (Text-Fig. 10, gon). Each gonimolobe consists of a compact, branched stout gonimoblast cluster, the lowermost cell of which forms a stalk while all other cells produce carpospores (Text-Figs. 13, 14). The Text-Figs 13-14. Gymnothamnion elegans. Figs. 13-14. Young and old cystocarps showing structure and the persistent lateral sterile branch with its terminal hair, {sc, supporting cell.) * carpospores are quite large, bright red, and measure 15-20p in dia meter. After the liberation of carpospores, the empty outer sheath persists as a transparent sac for sometime (Text-Fig. 14). Vestiges of the carpogonial branch can be made out even in the ripe cystocarps. The lateral sterile branch with its terminal hair always persists and is prominently seen in the fructifications (Text-Figs. 10, 13, 14). ■ Bornet and Thuret (1876) have given an account of the asexual and sexual reproduction of this alga. The present account agrees in general with their observations. In this paper the writer has given details of early post-fertilization development also. The details of early post-fertilization development are in agreement with what has been described for the related genera Ptilota (Kylin, 1923) and Plumaria (Suneson, 1938; Drew, 1939). In these two genera, the carpogonium after tertilization divides into two cells, the lower of which functions as the connecting cell (sporogenous cell) and fuses with the auxiliary cell. The present observations indicate that the Indian Gymnothamnion differs from these and shows similarity to genera like Antithanmion in the division of the fertilized carpogonium into three cells, the lowermost of which establishes connection with the auxi liary cell. Callithamnion elegans Schousboe ex C. Agardh (1828, p. 162) was transferred to Plumaria Schmitz {non Plumaria Stackhouse) by Schmitz in 1889 as P. schousboei (Bornet) Schmitz. J. Agardh (1892), however, segregated it as a new genus, Gymnothamnion [type species, G. elegans (Schousboe) J. Ag.]. Gymnothamnion is distinguished from Plumaria Schmitz by its small size, absence of cortication and simple unbranched nature of the branchlets. The present study indicates that the formation of only one lateral pericentral cell in the fertile segment and the division of the fertilized carpogonium into three parts may serve as additional distinguishing features. Collins and Hervey (1917) added two more species to Gymno thamnion, G. sericeum (Harvey) Collins et Hervey and G. bipinnatum Collins et Hervey. Feldmann-Mazoyer (1940, p. 353) included G.