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A Contribution to the Floral Anatomy and Embryology of Linaceae

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A Contribution to the Floral Anatomy and Embryology of Linaceae A CONTRIBUTION TO THE FLORAL ANATOMY AND EMBRYOLOGY OF LINACEAE By L. L. N a r a y a n a Department o f Botany, Osmania University, Hyderabad-! (Received for publication on November 29, 1962) I ntroduction T h e Linaceae have not received much attention from the point of view of floral anatomy, the only work being that of Saunders (1937). Information on the embryology of the family is meagre and this has been summarized by Schnarf (1931). Kappert (1933) reported poly- embryony in Linum usHatissimum. Soueges (1924, 1937) described the embryo development in Radiola linoides and Linum catharticum. Johan­ sen (1950) classified the embryo development in this family under the Linum variation of Solanad type. Subsequently, Mauritzon (1934) studied the embryology of Radiola linoides. Doraisami and Gopinath (1945) studied the development of the female gametophyte, endosperm and embryo in Linum mysorense. In view of the scanty information on the floral anatomy and embryology of Linaceae, this study was undertaken and the present account deals with the floral morphology of Durandea penlagyna (Warb.) K. Schym., Hugonia mystax Linn., Linum grandiflorum Desf., L. rubrum Rafin and Reinwardtio trigyna Planch. M aterials a n d M ethods The materials were fixed in formalin acetic alcohol. Customary methods of dehydration, infiltration and embedding were followed. Sections were cut at a thickness of 5-12 microns and stained in Eherlisch Haematoxylin. The herbarium material of Durandea was soaked in water for 24 hours. It was then boiled in a weak solution of KOH for 2-3 hours and later washed in running water for about 2 hours. It was fixed in formalin acetic alcohol for 24 hours and then treated in the usual way. F lo ra l A natom y The flower is pedicillate, regular, bisexual, heterochlamydeous and pentamerous. The sepals and petals show quincuncial and contorted aestivation respectively (Text-Figs. 4, 5, 9, 10, 14, 15, 17, 18, 24-27, 34 2 . m m II.U Text-Fios. 1-20. T.S. of flower from base upwards. F i^ . 1-10, Durane4a pentagyna. Figs. 11-20. Hugonia mystax, ^ 37, 44, 45, 49). The 10 stamens of Durandea and Hugonia are of two heights, the antepetaJous ones being shorter (Text-Fig. 10). In Linum species and Reinwardtia there are 5 fertile antesepalous stamens tmm It 2t Text-Figs. 21-39. T.S. of flower from base upwards. Figs. 21-29. Linum frandiflorum. Figs. 30-39, L, rubrum. VText-Fiqs. 40-55.^; Figs. 40-49, 55. Reinwardtia trigyna. Figs. 40-49. T.S. of flower from base upwards. Fig. 55. Mature pollen grain. Figs. 50, 52, 53. Linum gramUflorum. Fig, 50. T.S. of anther lobe showing microspore mother cells, tapctum and wall layers. Fig. 52. P.M.C. showing cytokinesis. Fig. 53. Pollen te tj^ . Figs. 51, 54. Hugonia mystax. Fig. 51. Anther lobe show ing 1-nucleate poUen grams and tapetal layer. Fig. 54. Pollen grain showing ‘O N Q ’. alternating with 5 sterile non-vascular filiform staminodes (Text-Figs. 27, 39, 47). The filaments in all the species are basally connate (Text- Figs. 5, 6, 8, 9, 18, 26. 35-37, 46). The syncarpous gynoecium consists of 5 carpels in Durandea and Linum species (Text-Figs. 7-9, 27 29, 37, 38). 5-3 in Hugonia (Text- Figs. 19, 20) and 3 in Rcinwardtia TText-Figs. 47, 48). The number of loculi corresponds with the number of carpels in Durandea and Hugonia (Text-Figs. 8, 9, 19, 20). Fn Linum species and Rcinwardtia as each locule is divided into two chambers by a sterile septum, the number of loculi at the base of the ovary is double the number of carpels (Text- Figs. 27, 36. 37, 47). But, the septum is incomplete towards the top of the ovary and at this level the number of loculi corresponds with the number of carpels (Text-Figs. 28, 29, 38, 48). Each locule bears two pendulous ovules (Text-Figs. 7, 9, 19, 20, 28, 29. 37, 38, 47, 48). The pedicel shows a ring of vascular bundles in Durandea and L. ruhrum (Text-Figs. 1, 30) and a siphonostele in the rest (Text-Figs. 11, 21, 40). Fn Durandea and L. grandiflorum the conjoint sepal laterals and sepal midribs arise in two alternating whorls (Text-Figs. 2, 22); in Hugonia they are close together (Texi-Fig. 12). In L. ruhrum and Reinwardtia the sepal traces (Text-Figs. 31, 41) divide radially forming a median and two lateral bundles (Text-Figs. 32, 42). The petals are single-traced and they arise independently from the main stele alter­ nating with the sepal midribs (Text-Figs. 3, 13, 23, 24, 33, 34, 43, 44). The traces supplying the perianth members divide further forming smaller bundles in the respective organs (Text-Figs. 3-6, 8-10, 13-15, 17, 18, 23-27, 33-37, 43-49). The traces for the 10 stamens in Hugonia arise in two alternating whorls, the antesepalous ones being organized first (Text-Figs. 14, 15). They emerge out and at the periphery of the thalamus they present a horse-shoe-shaped outline (Text-Fig. 16). In Durandea on the other hand the traces for the antepetalous stamens become demarcated first (Text-Fig. 4) and thus, the androecium is obdiplostemonous. Tn Linum species and Reinwardtia only 5 staminal traces arise on the sepal radii and these feed the fertile stamens (Text-Figs. 25. 27, 34-37, 44-47). The sterile filaments representing the antepetalous stamens are devoid of vascular supply (Text-Figs. 27, 39, 47). At about the level of separation of the staminal tube, the dorsal carpellary traces are organized (Text-Figs. 6, 18, 26, 36, 46). In Durandea, each dorsal carpellary trace divides to form two lateral branches, which again divide into two each (Text-Figs. 6-8). In Hugonia the dorsal carpellary traces divide into numerous branches in the ovary wall (Text-Figs. 18-20). Alternating with these the common median laterals are organized (Text-Figs. 18-20). The main stele then closes to form a ring which in turn splits into as many ventral bundles as there are carpels and they lie opposite the loculi (Text-Figs. 7, 19, 20). In Linum species the emerging staminal traces divide tangentially demarcating an inner ring of bundles, the common median laterals (Text-Figs. 25, 26, 35). At a higher level the dorsal carpellary traces are organized (Text-Figs. 26, 36). The common ventral bundles lie along the septal radii (Text-Figs. 27-29, 37, 38). In Reinwardiia the Text-Fios. 56-70. Figs. 56-58, 64, 67, 68, 70. Reinwardiia trigyna. Figs. 56-58. Stages in the development of ovule. Fig. 64. L.S. of ovules showing M.M.C. Fig. 67. 8-nucleate embryo-sac; note early differentiation of antipodals. Fig. 68. Organized- 8-nucleate embryo-sac. Fig. 70. Micropylar parts of embryo- sac showing egg apparatus and secondary nucleus surrounded by well-differentiated endothelium. Figs. 59, 65, 66. Hugonia mystax. Fig. 59. Mature ovule. Fig. 65. M.M.C. and parietal tissue ; note poorly differentiated endothelium. Fig. 66. Megaspore tetrad and parietal layers. Figs. 60, 69. Linum rubruni- Fig. 60, MLature ovule. Fig. 69. Micropylar parts of mature embryo-sac showing egg apparatus and secondary nucleus surrounded by well-differentiated endothelium. Figs. 61, 62, 63. Linum grandiflorum. Fig. 61. Mature ovule. Fig. 62. Andiesporium. Fig. 63, Two megaspore mother cells. median laterals arise as common bundles, but become split up radially as they enter the ovary wall ('Text-Figs. 45-48). The common ventral bundles lying on the sepal radii divide into two at the level where the ovular traces are given off (Text-Fig. 48). In all the members the ventral bundles are used up in the ovular supply. Only the dorsal car- pellary traces extend to the tips of the stylar branches (Text-Figs. 10, 39, 49). MlCROSPOROGENESlS AND M a LE GAMETOPHYTE , A fully differentiated anther shows an epidermis and four wall layers (Text-Figs. 50, 51). Of these, the innermost develops into the secretory tapetum, whose cells become binucleate (Text-Figs. 50, 51). II is absorbed as the microspores are formed in the anthers. In Hugonia, however, the tapetal cells persist in the anthers even after uni­ nucleate pollen grains are formed (Text-Fig. 51). The cells of the hypo- dermal wall layer undergo radial elongation, develop fibrous thicken­ ings and function as the endothecium. The middle layers get crushed during the development of the anther. Cytokinesis takes place by peripheral furrowing (Text-Fig. 52) and tetrahedral pollen tetrads are formed (Text-Fig. 53). The pollen grains are 3-colporate in Linum and Hugonia (Text-Fig. 54), while in Reinwardtia they arc non-aperturate (Text-Fig. 55). They are 3-celled at the shedding stage (Text-Fig. 55). The exine ornamentation is different in the genera studied. It is verrucose with the verrucae of the same size in Linum rubrum and Hugonia (Text-Fig. 54) and of different sizes in L. grandiflorum and Reinwardtia (Text-Fig. 55). Starch grains are present in the mature pollen grains. ‘ONCUS’-like structures are present in the pollen grains of Hugonia (Text-Fig. 54). O v u l e The ovule is crassinucellar, bitegminal and anatropous (Text- Figs. 56-61). The ovule primordium arises on the placenta as a small cushion-like outgrowth. The integumentary primordia are formed by the time the archesporium is differentiated. It soon bends upwards during development and the mature ovule becomes anatropous with the micropyle pointing upwards (Text-Figs. 58-61). The integuments are free (Text-Figs. 57-61). The outer integument consists of two layers of cells in Linum species and three layers of cells in Hugonia and Reinwardtia.
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