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SARGASSUM TENERRIMUM J.Ag.' by C A CONTRIBUTION TO THE MORPHOLOGY AND LIFE-HISTORY OF SARGASSUM TENERRIMUM J.Ag.' BY C. SURYA PRAKASA RAO, M,Sc. (Lecturer in Botany, Andhra University, Guntar, lndta) Received November 8, 1945) (Communicated by Prof Y. Bharadwajl, Ms-'., P'l D (Lond), r.L.S, r s l, F.A.S:. INTRODUCTION NO contribution has yet been made towards the life-history of Indian Sargassums in spite of their relative abundance. The genus is one of the most conspicuous and common of the sea-weeds which one can see either washed ashore or growing attached to rocks below low-tide level along the east and west coasts of India. In 1848 J. Agardh described several species of Sarga~sum from this country (Specie~ Alga urn, Vol. 1, A:ga; F.tco/de; co,~,,p!,cte s, Land 1848) and in the following year R. K. G:eviile gave art account of fourtcen sp~'c,es ("Algm Orientalis--Desc'iption of new species belong;rig to th~ Ge:ms Sargassum.", Ann. and Mag. Nat. Hzst., S~r. 2. Vol. 3). B~rgesen (1933) also listed six of these species from Bombay in 1933 and one species from C~yIon in 1936. Th~,ywere: Sargassum tenerrimum J. Ag. (which h~ thinks syaony- mous with S. Campbellianum Greville), S. cinerum J. Ag. var. barberifolia Grun., S. Wightii (Grev. mscr.) J. Ag., S. ilici[olium (Ttlrn~ C. Ag., S. plagio- phyllum (Mort) J. Ag., S. myriocystum J. Ag., S. crist~,efolium C. Ag. Apart from a taxonomic description of Indian species of Sargassum, no attempt has been made to give a detailed account of any of them. At the suggesUon of Professor Y. Bharadwaja, the present work s was, therefore, undertaken by the writer to fill up this gap. HISTORICAL While a fair amount of anatomical and cytological work has been done on other members of the Fucales, very little is known about Sargassum. Miss S'mons (1906) studied the development of conceptacles and embryo in Sargassum filipendula, while Nienberg (1910) described the oogonium and * From the Department of Botany, Benares Hindu University. 2 This is a portion of the work so~bmltted in part fulfilment for the M.Sc. Degreeof the Be.nares Hmdu Untversatyin 1939 B~ 39 40 C. Surya Prakasa Rao sporeling development in Sargassum liaifolium. The later work is largely from Japanese workers of ~vhom Tahara (1909) and Kunzida (1928) contri- buted tte most. The former author records definite periodicity in the liberation of oogonia of Sargassum corresponding to that in Dictyota, while the l~tter gives a thorough account of the cytological details in the oogenesls and spermatogenests of Sargassum Horneri. The present paper, however. deals with Sargassum tenerrimum J. Ag. MATERIAL AND METHODS The material for this investigation was collected from Bombay in January 1939.~ It was killed and fixed in formalin-acetic-alcohol. Other fixatives such as Land's formalin-alcohol, Dr. Lynds Jones' formalin-alcohol, and chromo-acetic solution were also tried and found useful on material collected lately by the writer from Vizagapatam. The usual paraffin method was followed in sectioning the material, sections being cut from 8-14/~ thick. Staining was done with Heidenhain's iron-alum ha~matoxylin and Delafield's ha~matoxylin. The latter, owing to its staining the young walls, proved more useful than the former. MORPHOLOGY The plant under investigation agrees in taxonomic characters with the Sargassum tenerrimum described and figured by Boergesen (1933) and Setchell (1935). J. Agardh (1848) described the plant originally from the coast of Bombay under the present name, while Greville (1849) described the same as Sargassum Campbelliamum. Grunow (1916) later referred to this form as a variety of S. tennerimum under the name var. campbellianum. The plant has a-basal adhesive disc or hold-fast from which arises a short primary axis bearing several branches together. The axis is rounded, glabrous, scarcely 2 ram. across. The secondary branches fork repeatedly in their turn. "Leaves" are 2-6 cm. long and 89to 189cm. wide, gradually decreasing in size from the base upwards. They are thin, translucent and linear-lanceolate, with toothed margin" and rather indistinct midrib. Crypto- stomata are freely scattered on the "leaves" and receptacles whi!e they are rare on the axis. The receptacles are freely-branched structures being attached to the upper portion of tile stipe of the leaf. They are androgynous, spinose and show various cases of transformation of the branches., Cases have been observed where a portion of the receptacular branch may be a t I am indebted to Captain S. C. Dtxat, M.^., u.sc., formerly of Wilson College, Bombay, or the supply of the material. Contribution to JCorpholo#y & Life.History o[ S. tenerrimum 41 leaf, or a bladder, or may be surmounted by a leafy expanse. The floats are either borne separately on short stalks or assoicated with the very much branched receptacle branches. They are spherical or egg-shaped, the tip b~ing commonly terminated by a small leafy appendage. T,~is description coincides more with that given by Boergesen than that by S~tcheU who describes under the same name a plant from Hong-Kong in which the "leav:s" are much smaller. ,~ATOMY S'mons (1906) summariscs the histological work done on the genus up to 1906. The work of Reinke (1876) on S. Boryanum, of O'.tmanns (1889) on S. linifolium and S. varians, of Hansteen (1892) on S. bacciferum, and the investigations of S mons herself on S. filipendula show that the growth of the axis and lateral organs takes place by means of a single 3-sided apical cell. In the ' stem' th-ee kinds of tissues are recognised :--:he assimilatory system, constituted by the epidermal cells; the storage system which is between the assimilatory and the innermost third system called the conducting system. According to Hansteen, the cells of the three systems are co1~unicating with each other by pores, but S mons (1906) could not see such pores in bet material of S. filipendula. In spite of b-~st ~fforts, the writer also could not see pores in S. tennerimum. In S. tennerimum also. both in the main axls and the laterals, there is a three-sided apical ceU which contributes along with its neighbourmg cells to the d,ffcrentiation of all subsequent systems. Tae apical cell (FLgs. I A and B) is situated in a pit or cavity which is its own creation, fo reed due to the quick succession in the production of cells in a tangentml plane and tt:e consequent lack of space for the elbowing of the resulting cells. It cuts eft cells parallel to the three sides and the base and thus maintains its ~dentlty throughout. The cells of the al3ove-mmtioned three systems get d~fferentiated at some distance from th~ apex. Th~ transverse section of the stem distmc~.!y shows the three regions, n. m2y, assimilato~y, storage and conducting. Tae outer- most assimilatory system consists of compactly-placed cells wim.h convex outer walls, covered by mucilage. Taey are rich in ch-ematophores and reserve material. Tae ceils of this system maintain their merist~matic activity throughout their life and always d,wde at right angles to the surface. They are also responsible for the development of male and fema!e concept- acles and cryptostomata. The cells of the second system, which also possess reserve material, are larger and possess wavy walls. They enclose many intercellular spaces. The cells of the conducting system are elongated, 42 C. Surya Prakasa Rao ]~t~. I A--transverse ann B---tongitudtnal section of growing point showing three-sided apical cell ; C-G--transverse secuons of receptacles showing mlhal stages m t!ze development of conceptacles C--showing conceptacle-~mtlal; D--showing two-celled stage; E--showing three-celled stage, F--showing five-celled stage, G--showing tongue-cell ; H-J--stages in the development of cryptostomata //--showing two-celled stage, l---showing five-celled stage, J--sho~mg tongue-cell of the same d~dmg Uansversely t, tongue.ce,ll, o, oogomum initial, m, rn'uellage. All • 1650. ContriSutzon 'o ,PlorpAolo#y & Li[e.H~story o/'S. tenerrimum r compactly arranged and possess oblique cross-waUs. They lack in reserve material. In a transverse section of the leaf also three regions can be recognized. The thickness of the leaf decreases as we pass from the m~drib to sides, this being due to the reduction in the number of layers of cells that compose the storage tissue. RECEPTACLES AND CONCEPTACLES Contrary to what we find in Fucus, the function of reproduction is relegated to specialized lateral branch systems. This character is also found in other Sargassaceous plants, such as Bifurcaria, Cystophyllum, Marginaria, Carpophyllum, etc. The receptacles are usually bisexual, although unisexual female receptacles are also met with. The conceptacles may be separated from one another by many layers of cells, or there may be only a little tissue between them. Their ostioles can be seen a bit raised over the general surface of the receptacle. DEVELOPMENT OF MaLE AND FEMALE CONCEPTACLES The d~velopmental history of conceptacles given by Bower (1880) for six plants of Fucaceae, that the initial cell does not contribute to the forma- tion of the conceptacle, was supported by subsequent workers like Valiante (1883) (in Cystoseira), Oltmanns (1889) (in Halydris siliquosa, Himanthalia lorea and Ascophyllurn modosum), and Grauber (1896) (in Seiroccus~, till Simons (1906) in her account of Sargassumfilipendula contradicted this view. She was supported by Nienburg (1910). In early stages, the genesis of male and female conceptacles in S. tennerrimum is identical, so that a generahsed description will hold good for both.
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