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Picea Smithiana (Wall) Boiss* Proc. Indian Acad. Sci., Vol. 81 B, No. 3, 1975, pp. 101-110 Anatomy of the mature embryo and seedling of Picea smithiana (Wall) Boiss* K. VENKATARATNAM,B. CHACKO, B. D. DESHPANDE AND S. K. PILLAI Department of Biological Sciences, Birla Institute of Technology and Science, Pilani 333031 MS received 27 September 1973 (Communicated by Prof. V. Puri, F.A.Sc.) ABSTRACT The anatomy of the mature embryo and M-day old seedling of Picea smithiana (Wall) Boiss. has been described. The organization of the shoot and root apices in both is similar. The shoot apex shows five cytohistological zones, where the entire surface layer shows periclina[ divisions and a small subapical initials zone. In the radicular and root apices there is a common initiating zone for the stele and columella in the centre surrounded by another common initiating zone for the cortex and peripheral region of the root cap. In the mature embryo the kappe divisions of the protoderm resulting in the peripheral region of the cap can be noted, which, however, disappear in the root of the seedling. The 34-day old seedling shows a root-hypocotyl-cotyledon vasculature without any connection with the shoot, where no vasculature ha~ yet developed. INTRODUCTION REPORTS on the structure of the mature embryo and seedling of gymnosperms are rare (Buchholz and Old 1933; Schopf 1943; Alien 1947 a, b; Tepper 1963, 1964; Gregory and Romberger 1972; Riding 1972). The shoot apex of the adult plant of Picea smithiana has been studied by Shah and Thulasi (1967). This report presents information about the anatomy of the mature embryo and 34-day old seedling of Picea smithiana. MATERIALS AND METHODS Seeds of Picea smithiana (kindly supplied by Prof. R. V. Singh, Depart- ment of Forestry, Himachal Pradesh University, Solan, Simla Hills) were soaked in water for an hour, the seedcoat removed and the embryo fixed. Seeds germinated only after a cold treatment at about 5°C for 25 days or * This work has boon financed in part by a grant made by the United States Department of Agriculture, Agricultural Re~mrch Servioz, authoris~d by Public Law 480. 10I B 1--March 75 102 K. VENKATARATNAM et al. more. The treated seeds were sown in the soil in September 1972, and they germinated in about 15 days. All materials were fixed in formalin-acetic acid-alcohol, processed through the alcohol-xylol series and embedded in paraffin. Serial sections, transverse and longitudinal, of the mature embryo and shoot and root apices of the seedlings, were cut at 8 ~, thickness. Trans- verse sections of the seedlings were taken at 15/~ thickness for studying the root-shoot transition. Northan's variation of Foster's schedule (Johansen 1940) for tannic acid-iron chloride and safranin was used in most cases. Chlorazol Black E also gave good results. OBSERVATIONS The shoot apex of the mature embryo is dome-shaped initially (figures ] and 22), and conical after soaking in water (figure 2). The shoot apex of the 34-day old seedling, though bigger, exhibits the same shape, and the first needle primordia become evident (figures 3 and 23). The average height and width of the shoot apex of the mature embryo are 91.32~ and 205.02ft and of the seedling 139 ~ and 250-2 f~ respectively. Tanniferous contents are noticed in the seedling apices. In the embryonic apex many granular contents are noticed in the cells, which disappear with germination and advancement of growth (figure 22). Zonation of tke skoot apex--The following zones are evident. Zone 1. The apical initials--These cells occupy the summit of the apex and in the mature embryo range from 6 to 8 (figure 22). In the 34-day old seedling apex their number is only 4 or 5 (figures 23). Shah and Thulasi (1967) have reported 2 to 6 apical initials in the adult plant apices. The apical initials are large with spherical, deeply staining nuclei and cell contents. In the cells of the mature embryo the nuclei are more prominent and occupy three-quarters of the cell volume. The cell walls are more or less uniformly thickened and the corner thickenings reported by Shah and Thulasi (1967) were not observed. Both anticlinal and periclinal divisions occur with the former predominating (figures 22). Periclinal divisions in the apical initials contribute to the subapical initials proximally, the distal derivatives persisting as the apical initials. Periclinal divisions in the shoot apical cells of the young embryo at cotyledonary initiation was reported by Gregory and Romberger (1972) in Picea abies. Periclines have been reported in the apical cells of the mature shoot of Torreya ealifornica by Kemp (1943), four species of Cupressaceae by AI Sheriff (1952), five species of Conifers by Jackman (1960), Cephalotaxus drupaeea by Singh (196I), Cupressus species by Pillai (1963 a), Thuja oriental& Thuja eompaeta, Juni- pcrus chinensis and Callitris robusta by Pillai (A. 1963 a). Anatomy of the mature embryo .. 103 @ @@ @ © U JJ " @ ab @ 6.,°.,° J . FOR I)2)3,S)2) )oo~ FOR 7-9 Figures 1-21. 1. Median L.s. of shoot apex of dormant mature embryo. 2. Median L.s, of shoot apex of embryo 1 hr after seed-wetting. 3. Median L.s. of shoot apex of 34-day old seedling. 4. Three-dimensional picture of seedling showing vasculature. 5. K6rper divisions in pecipheral region of root cap. 6. Median L.s. of radicle of mature embryo. 7. Median L.s. of root of 34--day old seedling. 8. Transection of root 2,050~ above root initials. 9. Transec- tion of root 4,500~ above root initials. 10. Transection of root 59,900p above root initials, 11. T.s. of hypoeotyl. 12. T.s. of hypocotyl 825 p below the shoot apex. 13. T.s. of hypocotyl 555~ below the shoot apex. 14. T.s. of hypocotyl 465p below the shoot apex. 15. T.s. of hypocotyl 375 p below the shoot apex. 16. T.s. of cotyledonary node. 17. T.s. of hypocotyl 225p below the shoot apex. 18. T.s. of hypocotyl 120p below the shoot apex. 19. T.s. of hypocotyl 80~ below the shoot apex. 20. T.s. of hypocotyl 30p below the shoot apex. 21. Vascular bundle in the cotyledon. i04 K. VENKATARATNAM et a~. Zone 2. The surface layer on the flanks--This zone arises by anticlinal divisions of the apical initials and is not treated as part of the flanking zone. In the mature embryo this zone has elongated cells with small nuclei. In the embryo and seedling apices periclinal divisions are common (figures 22 and 23), the inner derivatives contributing to the flanking zone. Increased frequency of periclines is observed during the initiation of needle primordia, the cells becoming wider at the locus of initiation (figures 3 and 23). Similar periclines are also reported in Abiesconcolor, Piceaexcelsaand Pinus montana (Korody 1937). Zone 3. The sub-apical initials--This zone, comprised of the inner derivatives of periclines in the apical initials (figures 1, 2, 3, 22 and 23), has cells with prominent, deeply stained nuclei and lightly stained cell contents. There are no corner wall thickenings. As the seedling grows, the cells enlarge (figure 23). Gregory and Romberger (1972) reported variation in the volume of this zone during growth up to 148 days in Picea abies. They also reported vacuolation in the precotyledonary shoot apex. Periclines in the peripheral cells of this zone form the flanking zone and anticlinal divisions in the basal cells, the rib meristem. Zone 4. The rib meristem--The highly vacuolated cells elongated parallel to the long axis of the shoot are arranged in regular vertical rows. The number of rows depends on the number of the subapical initials present. The width of this zone increases basipetally by periclinal divisions followed by cell enlargement and this is evident during seedling growth also. Anticlinal divisions also occur. Pith mother cells are absent, as reported by Pillai (1964) in Araucaria spp. In the shoot apex of the embryo the maturation of pith is nearer to the apical initials. The formation of pith is a gradual process as in the case of Sequoia (Cross 1943), Araucaria spp. (Pillai 1964). As the rib meristem extends into the hypocotyl, the vacuolation and length of the cells increase (figure 3). Zone 5. The flanking Zone or peripheral rneristem--This zone is a combined product of the periclinal divisions of the surface layer on the flanks and the anticlinal divisions of the subapical initials. The cells are longer than broad and densely cytoplasmic. Distally this zone is very narrow but becomes wider due to subsequent T-divisions. During the initiation of needle primordia divisions occur in all planes. The flanking zone forms a cylinder around the rib meristem and pith and is the site of development of procambium, cortex of the axis, and the subepidermal tissues of the needle. Root apex--The structure of the root apex in the mature embryo and seedling is fundamentally the same (figures 6, 7, 24, 25). In the mature embryo a disc-shaped group of 5 to 6 initials is present, while in the seedling root apices, a transverse plate of about 4 ceils form the initials Anatomy of the mature embryo .... 105 (figures 24, 25). These initials produce the stele and pith proximally and columella distally. The radicular stele is broader than the stele of the seedling root, evidently due to frequent kOrper divisions (figures 24, 25). In the radicle of the mature embryo the deeply stained narrow cells of the stele can be easily distinguished from those of the cortex and also by the colourless endodermis separating the stele from the cortex (figure 24).
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