Author: Kahraman, Ahmet
Date published: July 1, 2010
Colchicaceae, which is a family with a complicated distribution pattern, is made up of 19 genera distributed in Africa, Asia , Australia, Eurasia and North America (Vinnersten and Reeves, 2003). The pattern indicates an early Gondwanan distribution, however a previous investigation (Vinnersten and Bremer, 2001) revealed that the family is much younger. The taxonomic history of the family began in 1805 when de Candolle was the first to use the family name Colchicaceae. The taxonomic treatment of several genera within Colchicaceae is still ambiguous. For instance, Colchicum is often taken to include the genera Bulbocodium L. and Merendera Ramond, but some authors separate three genera based on style and tepal characters. Recently, Persson (2007) listed the subgeneric taxa of the genus Colchicum including a few genera such as Bulbocodium, Fouha, Merendera, Monocaryum and Synsiphon. In this study, Merendera kurdica Bornm. was treated as a synonym of Colchicum kurdicum (Bornm.) Stef. The genus Colchicum L. (Colchicaceae) in its most inclusive sense (including Merendera Ramond and Bulbocodium L.) includes nearly 100 species, which are very unevenly distributed (Dusen and Sumbul, 2007). Most species are confined to limited regions, but some of which are very rich in species. The high frequencies of species and of endemics in Turkey and the Balkans indicate that these regions are major centres of diversity and speciation. Colchicum is represented by 39 taxa, of which 18 are endemic in Turkey (Brickell, 1984; Persson, 2000, 2005, 2007; Akan and Satil 2005; Dusen and Sumbul, 2007). Colchicum kurdicum (Bornm.) Stef. (syn. Merendera kurdica Bornm.), an Irano-Turanian element, is a perennial stemless geophyte with three lanceolate leaves, mid-brown and membranous outher corm tunics, yellow anthers, and ellipsoid capsules. C. kurdicum grows in alpine steppe by melting snow ranging from 1800 to 3000 m in Hakkari, Şırnak and Van vicinities in the Southeast Anatolia. Moreover, the species is distributed in Iran and Iraq. It is not also related to any of the Colchicum species. Geophytes have an important commercial value especially in ornamental, food and medicinal industries (Celik et al., 2004). C. kurdicum may be used as an ornamental plant because of its beautiful purple flowers. It is known as "Karçiçeği" in Turkey. Colchicine is the main alkaloid of some genera of the family Colchicaceae such as Colchicum, Merendera and Bulbocodium (Mróz, 2008). Apart from antimitotic, antitumor, antiflammatory features, colchicine is well known for its cytotoxic effects and causes vomiting, diarrhea, and miscarriage in vertebrates (Paris and Moyse, 1967; Gómez et al., 2003). Sutlupinar (2002) studied the alkoloid content several organs of C. kurdicum (syn. M. kurdica) and determined colchicine and demecolcine in its corms and some homoaporphine such as baytopine, behuanine and kreysigine in its leaves and flowers. He studied its phenolics and identified benzoic and vanillic acids as aromatic acids.The genus Colchicum in Turkey has been subject to a number of morphological studies (Brickell, 1984; Kucuker, 1990; Persson, 2000, 2005, 2007; Akan and Eker, 2005). Anatomy of Colchicum and some genera in the Liliaceae was investigated by Vaikos et al. (1989), Akan and Satil (2005), Kaviani (2008) and Kahraman et al. (2010). However, the anatomical structures of roots and leaves of Colchicum kurdicum have not been studied previously. Thus, we aims to examine anatomical properties of C. kurdicum for the first time and to discuss our recent findings with previous studies.
Material and methods
The specimens Colchicum kurdicum were collected from Van (Van-Bahçesaray road, Yukarı Narlıca to Karlıgeçit, 2620 m, 38o 08' 178'' N - 420 55' 903'' E, 02.06.2006, AG14220) and were cultivated in Nezahat Gökyiğit Botanic Garden (NGBB) in İstanbul. The specimens for anatomical investigations were kept in 70% alcohol untill the sections were prepared. The paraffin wax method was used for the cross sections of roots and leaves. The samples were embedded in the paraffin wax, sectioned at 8-15 ìm thickness with a Leica RM2125RT rotary microtome, stained with safranin-fast green solution and mounted using Canada Balsam. Surface sections of the leaves were also taken manually. The sections were examined and photographed using a Leica DM1000 binocular ligth microscope with a Leica DFC280 camera.
The epidermis is composed of a single layer of cells of various shapes at the surface and it is covered with a thin cuticle. 2-layered (rarely one-layered) exodermis is located underneath the epidermis. The cortex is covered with 4-6 layers of parenhymatous cells. These cells are filled with elongated needle-shaped crystals of calcium oxalate. Underneath the cortex, there is a single layered endodermis consisting of oval or rectangular cells. The pericycle has a single layer of parenchymatous cells that lies just inside the endodermis. The xylem consists of 4 sets of protoxylem, the narrow cells at the point of the arms, and one central and larger metaxylem. Phloem strands are located between protoxylem strands (Figs. 1-2).
The upper epidermis is covered by a the thinner cuticle than the lower epidermis. Both epidermises are made up of uniseriate, isodiametric or rectangular cells. Cells of the upper epidermis is larger than those of the lower epidermis. Stomata are anomocytic. They are found more abundant on the upper surface. They are also located slighly higher from the epidermal cells. Stomata on the upper surface are 30-35 µm in length and 29-36 µm in width whereas stomata on the lower surface are 31-37 µm in length and 22-26 µm in width. The leaf is of the equifacial type. The mesophyll is composed of 2 layers (sometimes 3) of elongated rectangular palisade parenchymatous cells and 2 to 3 layers of irregular spongy parenchymatous cells. The vascular bundle is of the collateral type and of nealy equal sizes. The midrib do not constitute a projecting part (Figs. 3-5).
In this study, root and leaf anatomy of C. kurdicum are reported for the first time. The cortex in the root is 4-6- layered parenchymatic cells. The first row of the cortex includes abundant raphides, elongated needle forms of calcium oxalate crystals. Akan and Satil (2005), who studied the anatomy of five species of Colchicum, determined raphides in only the C. crocifolium roots. They also recognized that protoxylem arms were triarch in C. szovitsii and C. cilicicum, tetrarch in C. serpentinum and C. persicum, and pentarch in C. crocifolium. Our present study showed that the protoxylem arms of C. kurdicum were found to be tetrarch. Thus, presence or absence of raphide chrystals in the root and number of the protoxylem arms can represent useful taxonomic characters. The leaf of C. kurdicum are equifacial and amphistomatic. Stomata are anomocytic and are located more abundant on the upper surface. Moreover, stomata on the lower epidermis are narrower. Our results reveal that in many respects the leaves of C. kurdicum is similar to those of the species previously investigated by Akan and Satil (2005). There are 2-layered (sometimes 3) palisade parenchyma and 2-3 layers of spongy parenchyma. In C. kurdicum the upper epidermal cells are larger than the lower epidermal ones. Akan and Satil (2005) reported that the palisade parenchyma was 2 or 3 in four species studied except C. cilicicum with the one-layered palisade and also in C. serpentinum and C. crocifolium the upper epidermal cells are larger than the lower epidermal cells. However, they documented that both epidermal cells were similar size in C. szovitsii, C. cilicicum and C. persicum. This present study shows that the root and leaf anatomical characteristics of C. kurdicum are different from those of the species previously examined. At the same time, the study may serve as a significant reference for future studies on Colchicum.
We sincerely thank Prof. Dr. Adil Güner and other staff at the Nezahat Gökyiğit Botanic Garden (NGBB) for fresh plant material.
Akan H, Eker I (2005) A new record for Turkey: Colchicum crocifolium Boiss., with some contributions to the description of species. Belgian J Bot 138(1):93-96
Akan H, Satil F (2005) Morphological and anatomical investigations on some species of Colchicum L. occuring in South East Anatolia, Turkey. Journal of Biological Sciences 5(4):402-410
Brickell CD (1984) Colchicum L. In: Davis PH (ed) Flora of Turkey and the East Aegean Islands. Edinburgh: Edinburgh University Press, Edinburgh 8:329-351
Celik A, Cicek M, Semiz G, Karincali M (2004) Taxonomical and ecological investigations on some geophytes growing around Denizli province (Turkey). Turk J Bot 28:205-211
De Candolle AP (1805) Colchicaceae. In: Lamarck MM, De Candolle AP (eds), Flore française. Stoupe, Paris, France 3:192-193
Dusen OD, Sumbul H (2007) A Morphological Investigation of Colchicum L. (Liliaceae) Species in the Mediterranean Region in Turkey. Turk J Bot 31:373-419
Gómez D, Azorín J, Bastida J, Viladomat F, Conida C (2003) Seasonal and spatial variations of alkaloids in Merendera montana in relation to chemical defense and phenology. Journal of Chemical Ecology 29(5):1117-1126
Kahraman A, Celep F, Dogan M, Koyuncu M (2010) Morpho-anatomical studies on Bellevalia paradoxa Boiss. belonging to Liliaceae. Aust J Crop Sci 4(3) 150- 154.
Kucuker O (1990) Studies on the endemic Colchicum taxa of Turkey: C. linulatum Boiss et Spruner. Doğa-Tr J Bot 14:178-189
Mróz L (2008) Variation of colchicine and metal content in Colchicum autumnale L. (Liliaceae) corms in relation to edaphic environment. Polish Journal of Ecology 56(3):431-441
Paris RR, Moyse H (1967) Précis de Matiere Médicale. Vol. 3. Masson and Comp. Editeurs, Paris
Persson K (2000) Colchicum L. - In: Güner A, Ozhatay N, Ekim T, Baser KHC (eds) Flora of Turkey and the East Aegean Islands (Supplement II). Edinburgh: Edinburgh University Press, Edinburgh 11:246-265
Persson K (2005) A new Turkish species of Colchicum (Colchicaceae) related to C. boissieri. Edinburg J Bot 62:181-192
Persson K (2007) Nomenclatural synopsis of the genus Colchicum (Colchicaceae), with some new species and combinations. Bot Jahrb Syst 127(2):165-242
Sütlüpınar N (2002) Investigations on Turkish Colchicum and Merendera species. Acta Pharmaceutica Turcica 44:175- 181
Vinnersten A, Bremer K (2001) Age and biogeography of major clades in Liliales. American Journal of Botany 88:1695-1703
Vinnersten A, Reeves G (2003) Phylogenetic relationships within Colchicaceae. American Journal of Botany 90(10):1455-1462
Ahmet Kahraman*, Ferhat Celep
Middle East Technical University, Department of Biological Sciences, Ankara, Turkey
*Corresponding author: firstname.lastname@example.org, email@example.com