Chromosome numbers of species of the genus Jurinea Cass
Transkript
Chromosome numbers of species of the genus Jurinea Cass
CARYOLOGIA Vol. 62, no. 1: 16-23, 2009 Chromosome numbers of species of the genus Jurinea Cass. (Asteraceae) in Turkey Dogan1,* Bekir, Ahmet Duran2, Esra Martın3 and Erdogan Esref Hakkı4 Selcuk University, Faculty of Education, Department of Science Education, 42090, Meram, Konya, Turkey. Selcuk University, Faculty of Education, Department of Biology Education, 42090, Meram, Konya, Turkey. 3 Nigde University, Faculty of Science and Arts, Department of Biology, 51350, Nigde, Turkey. 4 Selcuk University, Faculty of Agriculture, Department of Field Crops, 42070, Selçuklu, Konya, Turkey. 1 2 Abstract — Chromosome numbers of 13 Jurinea species, collected from different localities in Turkey, were determined for the first time. All the Jurinea species investigated were diploid with chromosome numbers counted as 2n = 34. The basic chromosome number of the genus was x = 17. In this study, polyploidy was determined in nine taxa of Jurinea. Polyploid cells were 2n = 4x = 68 and they were tetraploid. The mitotic metaphase plates were given all the taxa of Jurinea. The research has made contribution to the cytotaxonomic revision of the genus Jurinea in Turkey. Key words: Cardueae, Carduinae, Compositae, Jurinea, Turkey. INTRODUCTION The Asteraceae with its approximately 1,620 genera and more than 23,600 species is the largest family of flowering plants (Stevens 2001). Asteraceae is widely distributed within diverse regions ranging from Southwest of US, Mexico, Southern Brazil, South Africa, Middle and Southwest Asia as well as Australia. South America is accepted to be phylogenetically the geographic origin of the family (Bremer 1994). Asteraceae is represented with the largest number of species in the Flora of Turkey with a total number of recorded species being 1209. From these species, 447 are endemic, with an endemism ratio of 37%. With a total number of 134 genera this family constitutes the second largest family of Turkish flora (Davıs et al. 1988; Özhatay and Kültür 2006). Jurinea Cass. is one of the most important genera within Asteraceae and it comprises about 200 species in total (Susanna et al. 2006). Native distribution of Jurinea is specifically involving Central Asia, Iran, Turkey and the Mediterranean basin. Jurinea is located within the monophyletic tribe Cardueae which is constituted of five sub- * Corresponding author: e-mail: doganbekir2000@yahoo. com tribes, namely, Carduinae, Centaureinae, Carlininae, Cardopatiinae Less., and Echinopinae. Jurinea is a genus located within the subtribe Carduinae (Susanna et al. 2006). The genus Jurinea is represented with 18 species within Mediterranean and Irano-Turanian phytogeographic regions of Turkey. Five of these species are endemic to Turkey resulting in an endemism ratio of 27.8% (Danin and Davıs 1975; Dogan 2007; Dogan et al. 2007). Jurinea taxa endemic to our country are; J. alpigena C.Koch, J. cadmea Boiss., J. pontica Hausskn. et Freyn ex Hausskn., J. brevicaulis Boiss., J. cataonica Boiss. et Hausskn. In the Flora of Turkey (Danin and Davıs 1975), it is also denoted that within the genus Jurinea there are some species which are not endemic to Anatolia but are found rather rarely. These species are; J. eriobasis DC., J. kilaea Azn., J. mesopotamica Hand.-Mazz., J. ramulosa Boiss. & Hausskn., J. multiflora (L.) B. Fedtsch. and J. stoechadifolia DC. Some of these species, namely J. eriobasis, J. mesopotamica and J. multiflora, have been placed to the category of data deficient (DD) (Ekim et al. 2000). Additionally, J. stoechadifolia and J. cypria Boiss. were defined, by Duman and Aytac (1999) and by Raus and Everest (2002) respectively, as new records for Turkey. Based on the studies of Rana and Sharma (2000), J. dolomiaea was determined as a me- chromosome numbers of species of the genus jurinea cass. (asteraceae) in turkey dicinal and aromatic plant species. Similarly J. macrocephala was described as an economically important species due to its potential usage in pharmaceutical industry (Dobrıyal et al. 1997). Negı et al. (2002) have also denoted that this species is equally important for the perfume and fragrance industries. There are many karyological studies on the genus Jurinea as Baksay (1956); Blanca & Cueto (1992); Borhıdı (1968); Chıchırıcco et al. (1979); Chouksanova et al. (1968); Dıaz De La Guardıa (1995); Gagnıdze and Gvınıashvılı (1997); Holub et al. (1971); Humphrıes et al. (1978); Jee et al. (1989); Kıeft and Van Loon (1978); Küpfer (1969a, b); Kuzmanov and Ancev (1973); Lungeanu (1973); Magulaev (1982); Sokolovskaıa and Probatova (1980); Vır Jee and Kachroo (1985); Vogt and Oberprıeler (1993). Related to the genus Jurinea, there aren`t any cytological studies on the species of the Flora of Turkey. This is the first record on the somatic chromosome numbers of 13 species of the genus that are natively distributed in Turkey. MATERIALS AND METHODS Samples of the genus Jurinea were collected from different localities of Turkey, with their akenes, during 2004-2006 vegetation periods. Collected samples are maintained as herbarium material at the KNYA and Selcuk University Faculty of Education herbaria. For somatic chromosome numbers, root tips were obtained from seeds germinated for ten days on wet filter paper in petri dishes. Root tips were pretreated with α-monobromonaphthalene for 16 h at 4ºC, fixed in ethanol: glacial acetic acid (3:1) for 24 h and stored at 4ºC until use. Root tips were washed in distilled water to remove the fixative, hydrolysed in 1N HCl for 13 min at room temperature and stained in 2% aceto-orcein for 2 h. Permanent slides were made with the standard liquid nitrogen method; slides were dried for 24 h at room temperature and mounted in depex. Chromosome counts were made using Bs200Pro Image Analysis Software (Martın et al. 2006). RESULTS AND DISCUSSION In the genus Jurinea, the chromosome number of all the species analysed were found as 2n = 34. Analyses were performed in the following species: J. consanguinea DC., J. alpigena C. Koch, 17 J. cadmea Boiss., J. mollis (L.) Reichb., J. macrocalathia C. Koch., J. pontica Hausskn. & Freyn ex Hausskn., J. pulchella DC., J. cypria Boiss., J. macrocephala DC., J. aucherana DC., J. ramulosa Boiss. & Hausskn., J. cataonica Boiss. & Hausskn., J. ancyrensis Bornm. In nine of the studied species polyploidy was detected in different cells of the preparate (2n = 4x = 68). Cytological results of the Jurinea species obtained from this study were given below. The orders of the species are according to their list given in the revision of the genus Jurinea in Turkey (Dogan 2007). Jurinea consanguinea - Konya: Sultan Daglari, North of Hidirlik dag hotel, 1130 m, 37º14.913’N, 038º49.427’E, 18.07.2006, BD 1001; its chromosome number was 2n = 34 (Fig. 1). According to our best knowledge, this is the first report on the chromosome number of the species. Jurinea alpigena - Karabuk: Yenice, Keltepe, 16301680 m, 41º03.796’N, 032º27.781’E, 21.07.2006, BD 1510; chromosome number was found as 2n = 34 (Fig. 2). This is the first report on the chromosome number of the species. J. alpigena is a species endemic to Turkey. Jurinea cadmea - Izmir: Odemis, Bozdag, 16501750 m, 38º20.671’N, 028º06.403’E, 19.07.2006, BD 1503; it presented a chromosome number of 2n = 34 (Fig. 3). This is the first report on the chromosome number of the species. The species is endemic to Turkey. Jurinea mollis - Kirklareli: 1 km after the crossroad for Vize-Sergen, Quercus sp. forest, 310-320 m, 41º37.312’N, 027º40.950’E, 07.07.2005, BD 1504; it exhibited 2n = 34 (Fig. 4). Jurinea macrocalathia - Tekirdag: Malkara, North of Kumbag, 5-25 m, 40º51.852’N, 027º27.604’E, BD 1507; it exhibited 2n = 34 (Fig. 5). With this study, the chromosome number of the species is given for the first time. Jurinea pontica - Kocaeli: Esme, 500 m before the signboard of the Sakarya province, 40-50 m, 40º44.118’N, 033º36.871’N, 09.07.2005, BD 1011; chromosome number was determined to be 2n = 34 (Fig. 6). This is the first report on the chromosome number of the species. This species is endemic to Turkey. Jurinea pulchella - Van: Yuzuncu Yil University campus, around the heating center, 1650 m, 05.08.2005, BD 1025; 2n = 34 (Fig. 7). We report the first chromosomal counting data of the species. Jurinea cypria - Mersin: Mut, plateau of Kozlar, edge of Ayı stream, 1330 m, 30.08.2006, BD 1521; 2n = 34 (Fig. 8). This is the first report on the chromosome number of the species. 18 dogan, duran, martın and hakkı Fig. 1 — J. consanguinea (2n = 34), Bar: 10 µm.; Fig. 2 — J. alpigena (2n = 34), Bar: 10 µm.; Fig. 3 — J. cadmea (2n = 34), Bar: 10 µm.; Fig. 4 — J. mollis (2n = 34), Bar: 10 µm.; Fig. 5 — J. macrocalathia (2n = 34), Bar: 10 µm.; Fig. 6 — J. pontica (2n = 34), Bar: 10 µm.; Fig. 7 — J. pulchella (2n = 34), Bar: 10 µm.; Fig. 8 — J. cypria (2n = 34), Bar: 10 µm. chromosome numbers of species of the genus jurinea cass. (asteraceae) in turkey Jurinea macrocephala - Konya: Halkapinar, Aydos mountain, Kayasaray village, 1 km East of Karasirt, 1700 m, 37º22.249’N, 034º17.308’E, 31.07.2005, BD 1017; 2n = 34 (Fig. 9). This is the first report on the chromosome number of the species. Jurinea aucherana - Erzincan: Old Cayirli road, 1-1.5 km apart from Botas petroleum pipeline, slopes of the hill on the left, 1760-2095 m, 39º53.429’N, 039º45.778’E, 22.08.2006, BD 1519; 2n = 34 (Fig. 10). We report the first, to our best knowledge, chromosomal counts of the species. 19 Jurinea ramulosa - Kahramanmaras: Ahirdagi, 1390 m, 37º37.057’N, 036º52.074’E, BD 1513; chromosome number as 2n = 34 (Fig. 11). In our study we report the chromosome number of the species for the first time. Jurinea cataonica - Erzincan: Old Cayirli road, 10 km, slopes aside the road, 1750 m, 39º47.954’ N, 039º30.343’ E, 07.08.2005, BD 1029; chromosome numbers were determined as 2n = 34 (Fig. 12). This is the first report on the chromosome number of the species. This species is endemic to Turkey. Fig. 9 — J. macrocephala (2n = 34), Bar: 10 µm.; Fig. 10 — J. aucherana (2n = 34), Bar: 10 µm.; Fig. 11 — J. ramulosa (2n = 34), Bar: 10 µm.; Fig. 12 — J. cataonica (2n = 34), Bar: 10 µm.; Fig. 13 — J. ancyrensis (2n = 34), Bar: 10 µm. 20 dogan, duran, martın and hakkı Table 1. Chromosome counts in taxa of the genus Jurinea in the world. Taxon J. ancyrensis Bornm. J. alpigena C. Koch J. arachnoidea Bunge J. arachnoidea Bunge J. aucherana DC. J. bellidioides Boiss. J. bocconii (Guss.) Guss. J. cadmea Boiss. J. cataonica Boiss. & Hausskn. J. ceratocarpa (Dcne.) Benth. J. ceratocarpa (Dcne.) Benth. J. ciscaucasica (Sosn.) Iljin J. ciscaucasica (Sosn.) Iljin J. consanguinea DC. J. consanguinea DC. subsp. bulgarica (Vel.) Koz. J. coronopofolia Somm. & Levier J. cyanoides Reichenb. J. cypria Boiss. J. elegans (Stev.) DC. J. filicifolia Boiss. J. fontqueri Cuatrec. J. humilis (Desf.) DC. J. humilis (Desf.) DC. J. humilis (Desf.) DC. J. humilis (Desf.) DC. J. humilis (Desf.) DC. J. humilis (Desf.) DC. J. iljinii Grossh. J. kilaea Azn. subsp. kilaea J. macrocalathia C. Koch. J. macrocephala DC. J. mollis (Torner) Reichenb. J. mollis (Torner) Reichenb. J. mollis (Torner) Reichenb. J. mollis (Torner) Reichenb. J. mollis (Torner) Reichenb. J. mollis (Torner) Reichenb. J. mollis (Torner) Reichenb. anatolica (Boiss.) Stoj. & Stef. J. mollis (Torner) Reichenb. moschata (DC.) Nyman J. mollis (Torner) Reichenb. transsilvanica (Spr.) Soo J. pontica Hausskn. & Freyn ex Hausskn. J. pulchella DC. J. ramulosa Boiss. & Hausskn. J. simonkaiana Nyar J. sintenisii Bornm. J. stoechadifolia (Bieb.) DC. J. stoechadifolia (Bieb.) DC. J. transchanica Regel et Schmalh. J. transsilvanica (Spr.) Schur J. turcica B.Dogan & A.Duran sp. nov. J. tzar-ferdinandii Dav. J. venusta Iljin 2n Locality 34 34 34 36 34 34 34 34 34 36 36 32 34 34 30 34 30 34 36 36 34 24, 58 34 34 34 34 34 34 30 34 34 30 34 34 34, 35 36 34 30 34 30 34 34 34 36 32 32 32 32 30 34 30 34 Turkey Turkey Russia Russia Turkey Russia Italy Turkey Turkey India India Russia Russia Turkey Bulgaria Russia Germany Turkey Russia U.S.A Spain Morocco Spain Spain Morocco Spain Spain Russia Bulgaria Turkey Turkey Hungary Greece Italy Slovakia Romania Turkey Bulgaria Italy Hungary Turkey Turkey Turkey Romania Russia Bulgaria Bulgaria Russia Hungary Turkey Bulgaria Russia Authors Present study Present study Magulaev, 1976 Spasskaya and Plaksına, 1995 Present study Magulaev, 1976 Mıcelı et al. 1978 Present study Present study Jee et al. 1989 Vır Jee and Kachroo, 1985 Sokolovskaıa and Probatova, 1980 Magulaev, 1976 Present study Kuzmanov and Georgıeva, 1977 Magulaev, 1976 Poddubnaja-Arnoldı, 1931 Present study Magulaev, 1986 Gagnıdze and Gvınıashvılı, 1997 Dıaz De La Guardıa, 1995 Humphrıes et al. 1978 Blanca and Cueto, 1992 Küpfer, 1974 Vogt and Oberprıeler, 1993 Küpfer, 1969b Küpfer, 1969a Magulaev, 1982 Kuzmanov and Ancev, 1973 Present study Present study Baksay, 1956 Snogerup, 1995 Kıeft and Van Loon, 1978 Holub et al. 1971 Lungeanu, 1973 Present study Kuzmanov and KOZUHAROV, 1967 Chıchırıcco et al. 1979 Borhıdı, 1968 Present study Present study Present study Lungeanu, 1973 Chouksanova et al. 1968 Kuzmanov et al. 1990 Kuzmanov et al. 1993 Sz.-Borsos, 1970 Kuzmanov et al. 1986 Present study Kuzmanov et al. 1986 Magulaev, 1976 chromosome numbers of species of the genus jurinea cass. (asteraceae) in turkey Jurinea ancyrensis - Elazig: Keban, KebanAgli road, 3th km, on the slopes, 800-850 m, 38º48.692’N, 038º 43.929’E, 21.08.2006, BD 1516; 2n = 34 (Fig. 13). This study presents the first report on the chromosome number of the species. This species is endemic to Turkey. In this study, the chromosome numbers of the natively distributed 13 species of the genus Jurinea were all determined as 2n = 34. The chromosomal numbers were remarkably consistent among the species studied. Our data confirmed the chromosomal counts for J. mollis (Kochjarova 1990) and they are the first reports for the remaining species studied. Chromosomal counting was not possible for the species J. kileae, J. brevicaulis and J. stoechadifolia due to the fact that we were unable to germinate the achenes of these species. Four of the species studied are endemic to Anatolia (J. alpigena, J. cadmea, J. pontica and J. cataonica). Although it was rare, during our cytogenetic analyses few cells revealed polyploidy in some of our preparates. This study has clearly indicated that the chromosomal counting is not a parameter that can be used in differentiation of the Jurinea species. Beside the chromosomal numbers, chromosomal morphologies should also be studied. However, the somatic chromosomes of the Jurinea species are very small and it is difficult to determine the exact positions of the centromer. Hence, it was not possible to determine the precise chromosomal morphologies. In the present study, all the Jurinea species studied were determined to be comprised of a basic chromosome number of x = 17. However, in nine of the species tetraploidy was evidenced with 2n = 4x = 68. Tetraploid cells were determined in the same preparates where diploid chromosomes were counted. In general, Jurinea presents low level of variation in chromosome number, with 2n = 34 in most of the species. Some species of Jurinea are disploids, with 2n = 24, 30, 32, 35, 36 and 58 (Missouri Botanical Garden, http://mobot. mobot.org/W3T/Search/ipcn.html and Index to Chromosome Numbers in the Asteraceae, http:// www-asteraceae.cla.kobe-u.ac.jp/index. html). In all the 13 Jurinea species collected from the natural Flora of Turkey cytological studies revealed an even number of chromosomes (2n = 34). According to literature survey, all the cytological studies on Jurinea are related to the chromosomal counts. Overall available data on these previous studies were given in Table 1. No study was found on the karyotypic evaluation in the genus Jurinea. 21 From the Jurinea species that are naturally grown in Turkey, only J. mollis was previously studied by different research groups and its somatic chromosome number was reported as 2n = 30, 34, 35 and 36 (Missouri Botanical Garden, http://mobot.mobot.org/W3T/Search/ipcn. html and Index to Chromosome Numbers in the Asteraceae, http://www-asteraceae.cla.kobe-u.ac.jp/ index.html). In accordance with the literature, our study also revealed the chromosome number of this species as 2n = 34. In the present study, somatic chromosome numbers of 13 Jurinea species from the family of Asteraceae were determined for the first time. We believe this study will play a positive role to enlighten this taxonomically complex genus with morphologically unsolved problems. Acknowledgements — We express our thanks to TUBITAK (Project no. 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