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Embryological Investigations in <Emphasis Type="Italic">Centipeda

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Embryological Investigations in <Emphasis Type= Proc. indian Acad. Sci. (Plant Sci.), Vol. 96, No. 2, June 1986, pp. 141-146. Printed in India. Embryologicai investigations in Centipeda mŸ (Asteraceae) P SWARAJYA LAKSHMI and T PULLAIAH Department of Botany, Sri Krishnadevaraya University, Anantapur 515 003, India MS received 17 February 1986: revised 30 May 1986 Almtraet. Anthersare tetrasporangiate. Anther tapetum is of the periplasmodialtype and its cells become polyploid. Microspore tetrads are tetrahedral. Pollen grains are shed at 3-celled stage. Ovule is anatropous, unitegmic and tenuinucellate. Embryo sac development is of the Polygonum type. Antipodals are either 2 of 3 in number. They become polyploid and are persistent. Synergids are hooked. Endosperm developmentis of the cellular type and embryo development conforms to the Grand period 1, Megarchetype II, series A and sub series A2 in the first embryonic group and it keys out to the Calendula subtype. Keywords. Embryology; Centipeda minima; Asteraceae. 1. Introduction Although embryological investigations in Angiosperms are quite extensive, a large number of genera remained uninvestigated. Maheshwari (1963) in this connection says 'future descriptive work in plant embryology should either be on such families and genera which have so far escaped attention'. Centipeda is one such genus where embryological information is not available (Pullaiah 1984). Hence the present investigation has been undertaken. 2. Material and methods Material of Centipeda minima (L.) A. Br. and Aschers was collected from Nilgiris and Kodaikanal in Tamiinadu. Flower heads at different stages of development were fixed in FAA. Dehydration was carried out in tertiary butyl alcohol and embedded in paraffin wax ofmeiting point 58-60~ Serial longitudinal and transverse sections were cut at a thickness of 5-6 ~tm and stained in Delafield's haematoxylin. Voucher specimen No. 2944 has been deposited in the Herbarium of the University and Madras Herbarium, Coimbatore. 3. Observations 3.1 Microsporangium The earliest stage that we could get in the anther was microspore tetrad stage. Anther wall at this stage consists of epidermis, hypodermis, degenerating middle layer and anther tapetum. Anther tapetum is of the Amoeboid type and the cells show nuclear divisions and fusions resuiting in polyploid nuclei. The hypodermal layer develops fibrous thickenings and functions as fibrous endothecium. The microspore tetrads are in a tetrahedral manner (figure 1B). The microspore after its sepamtion from the tetrad 141 142 P Swarajya Lakshmi and T Pullaiah ,...~ ~~ Q C |.,,t" ª .... o.o3ml A,H'K B-F. o.ozmm Figure 1. A. LS part of anther lobe showing periplasmodium and one-nucleate pollen grains. B. Microspore tetrad. C. One-nucleate pollen grain. D. 2-celled pollen grain. E. 3- celled pollen grain. F. Mature pollen grain. G. LS ovule. H. Megaspore tetrad. I. Micropylar part of the embryo sac showing egg apparatus and polar nuclei. J. Antipodal cells. K. Organised embryo sac. enlarges and acquires a thick exine (figure 1C). Its nucleus divides and two cells are formed. The smaller generative cell separates itself and enters into the cytoplasm of the vegetative cell (figure 1D). It divides to form two sperms (figure lE). The sperms which are oval become linear and filiform at maturity (figure 1F). Embryological investigations in C. minima 143 3.2 Ovule The ovule is anatropous, unitegmic (figure 1G) and tenuinucellate. The cells of inner epidermis of the integument develop into an integumentary tapetal layer at the time of megaspore tetrad formation in the ovule. It remains uniseriate with uninucleate cells throughout its further growth till it is completely absorbed by the growing embryo. 3.3 Megasporogenesis and female gametophyte The hypodermal archesporial cell functions directly as the megaspore mother cell and undergoes meiotic division resulting in a linear tetrad of megaspores (figure 1H). The chalazal megaspore is functional which after 3 mitotic divisions develops into a polygonum type of embryo sac (figures 1I, K). Antipodal cells are either 2 or 3 in number, if 2 are present the upper is binucleate. The antipodal cells later on become polyploid (figures I J, K). The antipodals are persistent and they are seen till the initiation of cotyledons in the embryo (figures 2D-F). The synergids are hooked (figure 1I) and they are ephemeral. 3.4 Fertilisation, endosperm and embryo Entry of pollen tube is porogamous (figure 2A). Triple fusion completes earlier than syngamy. Endosperm development is of the cellular type. Primary endosperm nucleus divides earlier than the zygote (figures 2B, C). The first and subsequent divisions are followed by ceil wall formation (figures 2B-F). The endosperm is completely consumed by the growing embryo but for one or two layers of cells. The zygote divides transversely resulting in a terminal cell (ca) anda basal cell (cb) (figures 3A-C). The terminal cell divides vertically and the basal cell transversely resulting in a 4-celled T-shaped first proembryonal tetrad. Each of the terminal cells divide by a vertical wall at right angles to the first giving rise to the quadrants (figure 3E). At this stage the cell m divides vertically forming two juxtaposed cells while the cell ci undergoes a transverse division forming cells n and n' (figure 3E). The destination of individual tiers and the development of embryo (figures 3E-N) are represented schematically in the following manner. (i) First cell generation. Proembryo consists of 2 cells disposed in 2 tiers: ca = pco + pvt; cb = phy +icc + iec + co + s. (ii) Second cell generation. 4-Celled proembryo, cells disposed in 3 tiers: q =pco +pvt; m =phy + icc; ci = iec + co + s. (iii) Third cell generation. 8-Celled proembryo, cells disposed in 4 tiers: q=pco+pvt; m=phy+ icc; n=iec+ co; n'= s. (iv) Fourth cell generation. 16-Celled proembryo, cells disposed in 5 tiers: q=pco+pvt; m=phy+icc; n=iec+co; o = 1/2 s; p= 1/2 s. 144 P Swarajya Lakshmi and T Pullaiah ~pt ,,' i..'.v....-- t 1 A m Figure 2. A. Micropylar part of the embryo sac showing the entry of the pollen tube. B--F. Depict the development of endosperm. Embryo development thus follows Grand period I, Megarchetype II, sedes A and subseries A2 in the first embryonic group according to Sou~ges (1939) system. Further both the cells o and p contribute to the suspensor which is the characteristic feature of Calendula type of Mestre (1964). Embryological investigations in C. minima 145 ~q .m mg tn .m M Figure 3. A. Zygote.B--N. Show stages in the developmentof embryo. 4. Discussion Sood and Thakur (1985) have recently reported that anther tapetum in Prenanthes brunoniana, also belonging to this family Asteraceae, is of the secretory type. In the present study, the tapetum is found to be Amoeboid. In the family Asteraceae occurrence of Amoeboid tapetum is characteristic feature and hence the report of Sood and Thakur (1985) deserves reinvestigation. In fact figure 7 of them (Sood and Thakur 1985) represents the breakdown of cell walls of anther tapetum which represents the incipient stage of the periplasmodium. Further, the life of periplasmodium in many members of the Asteraceae is short lived and hence they (Sood and Thakur 1985) might have missed the periplasmodium in their study. Glandular anther tapetum has also been reported earlier in a few members like Chrysothamnus (Snow 1945; Anderson 1970), Ainsliaea aptera (Kapil and Sethi 1962), 146 P Swarajya Lakshmi and T Pullaiah Vernonia cinerea, V. cinerascens (Tiagi and Taimni 1963), Sonchus oleraceus (Walter and Kuta 1971), S. arvensis, S. asper (Kaul et al 1975), Hypochoeris radicata (Kaul 1972), Youngiajaponica (Kaul 1973) and Tragopogon gracile (Singh and Kaul 1974)o As pointed out by Pullaiah (1984) periplasmodial tapetum is the characteristic feature of Asteraceae and all those reports where a glandular tapetum has been reported seem to be erroneus. We reinvestigated the anther tapetum in Vernonia cinerea, Sonchus oleraceus and Youngia japonica and found a periplasmodial tapetum. The suspensor in Centipeda minima is formed from o and p while in a majority of Asteraceae it is formed by the tier p only (Mestre 1964; PuUaiah 1984). The type of embryogeny where suspensor is formed by o and pis known as the Calendula type and it is also met with in other members of Asteraceae like Eupatorium ayapana, Calendula officinalis, Dimorphotheca pluvialis and Cnicus benedictus (Mestre 1964). References Anderson L C 1970 Embryology of Chrysothamnus (Astereae, Compositae); Madrono 20 337-342 Kapil R N and Sethi S B 1962 Gametogenesis and seed development in Ainsliaea aptera; Phytomorphology 12 222-234 Kaul V 1972 Embryology and development of fruit in Cichoreae II. Hypochoeris radicata; Proc. 59th lndian Sci. Congr. Part 3, p 338 Kaul V 1973 Embryology and development of fruit in Cichoreae III. Youngiajaponica; Proc. 60th Indian Sci. Congr. Part 3, p 328 Kaul V, Dathan A S R and Singh D 1975 Embryological studies in the genus Sonchus; J. lndian Bot. Soc. 54 238-245 Maheshwari P 1963 History and present status of angiosperm embryology; in Recent Advances in the embryology of Angiosperms (ed) P Maheswari (Delhi: lnt. Soc. Plant Morphologlsts) pp 1-13 Mestre J C 1964 Recerches d'embryogenie comparee: Les rapports phylogenetiques des Composees, Ph.D. thesis, University of Pa¡ Pa¡ PuUaiah T 1984 Embryology of Compositae (New Delhi: Today and Tomorrow). Singh D and Kaul V 1974 Cytoembryological studies on Indian Tragopogon, Cytologia 40 35-44 Snow E 1945 Floral morphology of Chrysothamnus nauseosus-specious; Bot. Gaz. 106 451--463 Sood S K and Thakur T S 1985 Development of male and female gametophyte and integument initiation in Prenanthes brunoniana (Compositae); Proc. Indian Acad. Sci. (Plant Sci.) 94 553-559 Sou+ges R 1939 Embryogenie et Classil~cation (Paris: Herman et Cie) Tiagi B and Taimni S 1963 Floral morphology and embryology of Vernonia cinerascensand V.
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