Diversity of Cypselar Anatomy in Five Species of Crepis L. of the Tribe Lactuceae (Asteraceae)
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1 THE JOURNAL OF ECONOMY, ENVIRONMENT AND SOCIETY a multidisciplinary journal of advanced studies Journal homepage: Diversity of Cypselar Anatomy in Five Species of Crepis L. of the Tribe Lactuceae (Asteraceae) Abhijit Shil* and Sobhan Kumar Mukherjee Department of Botany, Taxonomy and Biosystematics Laboratory, University of Kalyani, West Bengal, India abhijitbotanys@gmail.com Abstract: Article history: The paper deals with detail anatomical characters of cypselas of 5 species of the genus Original paper received on: Crepis L. [ Crepis biennis L., Crepis occidentalis Nutt, Crepis pyrenaica (L.) Greuter, Revised paper received on: Crepis schimperi (Sch. Bip. ex A. Rich.) Schweinf., Crepis turcica Degen and Bald] Paper accepted on: belonging to tribe Lactuceae. The anatomical variations of the above five taxa have been examined thoroughly to establish their possible utility in taxonomic study. The significant Keywords: Cypselar anatomy, anatomical features of cypsela, viz. cypselar shape in transverse section, cypselar ribs or Crepis L., elevations number, thickness of cypselar wall at ribs and furrow, nature of epicarpic cells, Lactuceae, Asteraceae. tissue differentiation of mesocarp, mesocarpic sclerotic braces, testal cell shape, orientation of testal cells, nature of mature endosperms, embryo and resin ducts in each cotyledon etc., have been studied. Presence of mesocarpic crystals (in Crepis occidentalis) is one of the important reliable taxonomic markers in this study. Based on all observed anatomical characters of cypselas, an artificial key to the studied species has been constructed. 1. Introduction Lactuceae represents 98 genera and more than 1550 species worldwide (Bremer, 1994). The genus Crepis L. (Lactuceae) including 200 species, distributed throughout the world (Mabberley, 2008) except in Australia and some parts of America. In India, only 9 species and 4 sub species (Hajra et al., 1995) are found. Most of the earlier workers paid little attention to discover internal features of cypselas. Lavialle (1912), Borthwick and Robbins (1928), Jeffrey (1966), Singh et al. (1973), Pandey, Singh and Chopra (1978), Beliaeva and Boyko (1980), Beliaeva (1982), Uyar and Cireli (1982), Pak and Kawano (1990a, b), Pak (1993), Bremer (1994), Mukherjee and Sarkar (1995), Mukherjee and Nordenstam (2004), Das and Mukherjee (2008), Das and Mukherjee (2012), Jana and Mukherjee (2012), Jana and Mukherjee (2013), Jana and Mukherjee (2014) have been attracted with cypsela structure of the tribe Lactuceae (Asteraceae). But, earlier information available in these respects may not be 1 sufficient to utilize as significant taxonomic parameter for systematic study of Crepis L. Therefore, the present study has been carried out to investigate the anatomical variation of the cypselas of Crepis biennis L., Crepis occidentalis Nutt, Crepis pyrenaica (L.) Greuter, Crepis schimperi (Sch. Bip. ex A. Rich.) Schweinf., Crepis turcica Degen and Bald in details of 5 species of the genus Crepis L., with the help of light microscope to realize the correlations among the taxa and to build an artificial key to the species on the basis of observed anatomical features of cypselas. 2. Materials and Methods The studied materials were obtained from the 4 herbaria of the world on request. Specimens listed in below in table I, were stored within desiccators with fused Calcium Chloride, in the Herbarium of the Department of Botany, University of Kalyani (Acronym: KAL).
2 Sl. No Name of the studied Taxa 1 Crepis biennis L Crepis occidentalis Nutt. Crepis pyrenaica (L.) Greuter Crepis schimperi (Sch. Bip. ex A. Rich.) Schweinf. Crepis turcica Degen and Bald. Source of the Specimens Bereich Botanik und Arboretum des Museums fur. Naturkunde der Humboldt Universitat Zu Berlin, DDR Berlin Spathstr. 80/81 (BHU), No. 229/735 Gray Herbarium (GH), Harvard University Herbaria & Libraries, A. D. Elmer, No Botanischer Garten der Universitat Zurich, Switzerland (Z),Alt St. Johann SG, Churfirsten 1680m asl, 2011, No. CHOZ Hortus Universitatis Hauniensis, Botanic Garden of Copenhagen (DK), 170GE , Denmark, Anno Hortus Universitatis Hauniensis, Botanic Garden of Copenhagen (DK), No. 172 Methods applied step by step to reveal the internal features of cypselas are as follows- 1. Cypselas were placed in 2N NaOH solution for hours or 70% phenol-glycerine solution depending on the hardness of the fruits. 2. Very thin handmade transverse section of the cypselas from the middle part of mature fruit have been prepared to know the internal features and then followed the method of Johanson (1940) for dehydration using various grades of alcohols. 3. Various internal features were observed under compound light microscope, viz. type of tissue, their arrangement, thickness, shape of the cells, orientation of the cells and presence or absence of cuticle etc. Other important features like, location of vascular trace, cavity, deposition of chemicals, distribution of crystals and number of secretory ducts etc. were also recorded. 4. Observed above noted characters were documented with the help of camera lucida drawings. 3. Result Comparative anatomical features of 5 studied species belonging to the genus Crepis of the tribe are given in Table II. Sl. No. Characteristic Features Crepis biennis L Table-II Crepis occidentalis Nutt. Crepis pyrenaica (L.) Greuter Crepis schimperi (Sch. Bip. ex A. Rich.) Schweinf. Crepis turcica Degen and Bald. 1 Cypselar shape in T.S. Oval Ellipsoid Oval- Circular Oval Oval (slightly pentagona) 2 Papillate hairs Present Present Absent Present Present 3 Cuticle Present Present (Thick) Present Present Present (Thick) 4 Ribs or Elevations Present Present Present Present Present 5 No. of ribs Size of the ribs Conspicuous Conspicuous Conspicuous Conspicuous Inconspic- uous 7 Thickness of cypselar wall , , , , at ribs and furrows (in µm) Thickness of pericarp at 62-75, , , ribs and furrows(in µm) Epicarpic hairs Absent Absent Absent Present Absent 10 Epicarpic cell shape Rectangular- Oval Rectangular Rectangular- Oval Rectangular- Oval Rectangular- Oval 11 Epicarpic cell wall Thick Thick Thick Thick Thick 12 Orientation of epicarpic cell Tangential Tangential Tangential Tangential Tangential 13 Mesocarpic tissue Present Present Present Present Present differentiation 14 Mesocarpic Present at Collapsed, Present at Present at furrows, Present below the parenchymatous cell furrows, irregular formed cavity furrows, irregular irregular epicarp 15 Mesocarpic crystal Absent Present Absent Absent Absent 16 Mesocarpic sclerotic braces Present at the rib Present at the rib Present at the rib Present at the rib Present continuously 2
3 17 Mesocarpic vascular bundle (vb) Present Present, (vb also present Present Present Present within testa ) 18 Mesocarpic cavity Absent Present Absent Present Absent 19 Testa thickness (in µm) Testal attachment with pericarp Attached Detached Attached Attached Attached 21 Tissue differen-tiation of testa Absent Present Present Absent Absent 22 Layers in testa Nature of testa Non- cellular Non- cellular Non- cellular Non- cellular Cellular 24 Testal cell shape Disorganized Disorganized Disorganized Disorganized Rectangular 25 Thick walled cells of testa Nil Nil Nil Nil Present 26 Orientation of testal cells NA NA NA NA Tangential 27 Endosperm in mature cypsela Persistent Persistent Persistent Persistent Persistent 28 Endosperm layer Biseriate Biseriate Biseriate Biseriate Biseriate 29 Mature embryo occupied Major part Major part Major part Major part Major part 30 Resin ducts in each cotyledon Relative size of the secretory Equal Unequal Unequal Unequal Unequal ducts 32 Relative size of the cotyledons Equal Equal Equal Unequal Equal Fig: 1. A-Diagramatic View, B- Part of Cypsela; 1A- 1B : Crepis biennis L. Fig: 2. A-Diagramatic View, B- Part of Cypsela; 2A- 2B: Crepis occidentalis Nutt. 3
4 Fig: 3. A-Diagramatic View, B- Part of Cypsela; 3A- 3B: Crepis pyrenaica (L.) Greuter. Fig: 4. A-Diagramatic View, B- Part of Cypsela; 4A- 4B: Crepis schimperi (Sch. Bip. ex A. Rich.) Schweinf. 4 Fig: 5. A-Diagramatic View, B- Part of Cypsela; 5A- 5B: Crepis turcica Degen and Bald. (LIST OF ABBREVIATION USED: C- Cuticle, COT- Cotyledon, CPA- Collapsed / Crusted Parenchyma, CR- Crystal, EN- Endosperm, EP- Epidermis, LLlarge lumen, ME- Mesocarp, MI- Inner mesocarp, MO- Outer mesocarp, NA- Not applicable, PH- Papillate hairy structure, RD- Resin Duct, SCL- Sclerenchyma, SCV- Secretary duct / cavity, T- Testa, TE- Testa epidermis,, TI- Testa inner zone/ Inner Testa, TPA- Thin wall Parenchyma, VC- Cavity, V.T- Vascular Trace.) 4. Discussion Fully adult cypselas of studied species are mostly round- oval but ellipsoid shaped in Crepis occidentalis, showing pericarp, testa, endosperm and cotyledons in cross section. Again, epicarp and mesocarp donate pericarp. Epicarpic cells; uniseriate, parenchymatous, thick-walled, rectangular to oval shaped, tangentially oriented, cuticularised, contain dark brown substances. Pandey, Singh and Chopra (1978), Jana and Mukherjee (2013) also have been illustrated the presence of dark brown substance in the epicarpic cells. All the studied taxa grown papillate hairs from the epicarp but C. schimperi also developed epicarpic hairs in addition with papillate hairs. Mesocarp of the
5 studied cypselas has shown minor variations within the studied taxa. It is made up of sclerenchymatous and parenchymatous tissue in all the studied taxa. Mesocarpic sclerenchyma multiseriate, discontinuous, present at each rib, just beneath the epicarp in all the studied taxa except in C. turcica, but sclerenchyma formed a continuous layer in Crepis occidentalis and Crepis turcica. Sclerenchymatous cells are thicker at each rib of Crepis biennis, which is also demonstrated by Pandey, Singh and Chopra (1978). Parenchymatous cells thin-walled, cellular mainly present at the furrows but secondarily crusted formed vallicular cavity when present at the ribs just below the sclerenchymatous brace. Out of the five studied taxa, vallicular cavity developed in Crepis occidentalis and C. schimperi. Presence of vallicular cavity within mesocarp have been reported by Beliaeva and Boyko (1980), Jana and Mukherjee (2014). Mesocarpic calcium oxalate crystal present in Crepis occidentalis as advocated by Jana and Mukherjee (2012). Testa primarily attached to pericarp although secondarily separated testa has also been found in C. occidentalis. Vascular traces are frequently originated at the base of sclerenchymatous brace. Testa biseriate in C. occidentalis and C. pyrenaica, but rest of the taxa shown uniseriate testa. Testa is represented by a crusted parenchymatous layer mainly with yellow brown substance as have been reported by Das and Mukherjee (2008), but cellular; cells thick-walled in C. turcica that also have been observed by Jana and Mukherjee (2012). Inner testa of C. occidentalis and outer testa of C. pyrenaica composed of collapsed thin walled cells. The above nature of testa has been reported by Pandey et al. (1978); Beliaeva (1982); Pak and Kawano (1990).Characteristic nature of cellular seed coat (testa) also reported by Lavialle (1912) and Borthwick and Robbins (1928) in the member of the tribe Lactuceae. Endosperm is consistently persistent and biseriate in mature cypsela. Beliaeva and Boyko (1980), Uyar and Cireli (1982), Pak (1993), Mukherjee and Sarkar (1995), Das and Mukherjee (2012) have promoted the same. Embryo occupied a major part of the cypsela with 2, plano-convex cotyledons. Cotyledons of the studied taxa showed very little variation among them, as they are unequal as in Crepis schimperi. Each cotyledon bears 1-8, equal secretory ducts but unequal in Crepis schimperi. An Artificial Key to the Studied Species: 1a. Testa bilayer; ribs always conspicuous.....(2) 1b. Testa unilayerd; ribs conspicuous or inconspicuous. (3) 2a. Cypsela oval to circular in cross section; mesocarpic parenchymatous cellular, not collapsed, number of ribs and mesocarpic crystal absent Crepis pyrenaica. 2b. Cypsela ellipsoid in cross section; mesocarpic parenchyma non- cellular, became collapsed, number of ribs 10 and mesocarpic crystal present... Crepis occidentalis. 3a. Ribs inconspicuous; testa cellular; resin ducts- 5 in each cotyledon.. Crepis turcica. 3b. Ribs conspicuous; testa non- cellular; resin ducts not 5 in each cotyledon. (4) 4a. Cotyledons unequal; resin ducts- 8 in each cotyledon....crepis schimperi. 4b. Cotyledons equal; resin ducts- 1 in each cotyledon.... Crepiss biennis. References Beliaeva, T.M. and Boyko, E.V. (1980). Morfologoanatomiceskoe stroenie semjanok pal nevostic ih Lactuca s.l. (Aster- aceae) v svjazi s ih sistematikoj. Botanicheskii Zhurnal. 65: Beliaeva, T.M. (1982). The genus Youngia (Asteraceae) from the Soviet fareast (USSR).Bot. Zh. (Leningr.) 67 (1): Borthwick, H. A. and Robbins, W. W. (1928). Lettuce seed and its germination. Hilgardia 3: Bremer, K. (1994). Asteraceae. Cladistics and Classification. Timber Press, Portland. Das, Debashis and Mukherjee, S. K. (2008). Diversity of cypselar features in seven species of the tribe Lactuceae. J. Econ. Taxon. Bot. 32(2): Das, Debashis and Mukherjee, S. K. (2012). Diversity of Cypselar Features in Six Species of the tribe Lactuceae (Asteraceae). In: "Multidisciplinary Approaches in Angiosperm Systematics"(Eds. Maiti, G.G. and Mukherjee, S. K.). Publication Cell, University of Kalyani, Kalyani, West Bengal, , India. ISBN: :
6 Hajra, P.K., Rao, R.R., and Uniyal, B. P. (1995). Flora of India. Vol. 12, 13. BSI. Calcutta. Jana, Bidyut Kumar and Mukherjee, S. K. (2012). Diversity of cypselar features of seven species of the genus Crepis L. in Compositae. Indian Journal of Fundamental and Applied Life Sciences. ISSN: Index Copernicus Value (ICV) = 5.01, 2 (1): Jana, B. K. and Mukherjee, S. K. (2013). Exomorphic and histological characters of fruits in some taxa of the tribe Lactuceae (Asteraceae). J. Econ. Tax. Bot. 37 (2): ISSN NO: Jana, B. K. and Mukherjee, S. K. (2014). Marker characters of cypselas in three taxa of Lactuceae (Compositae). ISSN: J. Botan. Soc. Bengal, 68(1): Jeffrey, C. (1966). Notes on Compositae I. The Cichorieae in East Tropical Africa. Kew Bull. 18: Lavialle, P. (1912). Researches sur le development de l ovarie en fruit chezles Composees. Ann. Sci. Nat. Bot. Ser. 9(15): Mabberley, D. J. (2008). Mabberley s Plant Book. Third Edition. Cambridge University Press, UK. Mukherjee, S. K. and Sarkar, A. K. (1995). Micromorphological and anatomical structures of cypselas in some members of the tribe Lactuceae (Compositae). J. natn. Bot. Soc. 49: Mukherjee, S.K. and Nordenstam, B. (2004). Diversity of carpopodial structure in the Asteraceae and its Taxonomic significance. Compositeae News letter. 41: Pak, J. H. & Kawano, S. (1990a). Biosystematic studies on the genus Ixeris (Compositae- Lactuceae) I. Fruit wall anatomy and its taxonomic implications. Acta Phytotax. Geobot. 41: Pak, J. H. & Kawano, S. (1990b). Biosystematic studies on the genus Ixeris and its allied genera (Compositae-Lactuceae) III. Fruit wall anatomy and karyology of Crepidiastrum and Paraixeris, and their taxonomic implications. Acta Phytotax. Geobot. 41 : Pak, J. H. (1993). Taxonomic implication of fruit wall anatomy and karyology of Crepis Sect.Ixeridopsis (Compositae; Lactuceae). Kor. J. Plant Tax. 23(1): Pandey, A. K., Singh, R. P. and Chopra, S. (1978). Development and structure of seeds and fruits in Compositae : Cichorieae. Phytomorph. 28(2): Singh, et al. (1973). Achene morphology : an aid to taxonomy of Indian plants. I. Compositae, Liguliferae. J.Indian Bot. Soc. 51: Uyar, N. and Cireli, B. (1982). Observations on the morphology, anatomy and ecology of Sonchus aspera And S. oleraceus growing in the Aegean region.doga.bilm.dergisi. 6 (1): Acknowledgement The authors are extremely thankful to Directors and Curators of Bereich Botanik und Arboretum des Museums fur. Naturkunde der Humboldt Universitat Zu Berlin (BHU), Botanic Garden of Copenhagen (DK), Gray Herbarium, Harvard University Herbaria & Libraries (GH) and Botanischer Garten der Universitat Zurich, Switzerland, Hortus Universitatis Hauniensis(Z) for supplying identified mature cypselas for this work. 6
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