Karyological Studies on Oxytropis (Fabaceae) from Turkey

Caryologia International Journal of Cytology, Cytosystematics and Cytogenetics

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Karyological studies on Oxytropis (Fabaceae) from Turkey

Esra Martin, Seher Karaman Erkul & Zeki Aytaç

To cite this article: Esra Martin, Seher Karaman Erkul & Zeki Aytaç (2015) Karyological studies on Oxytropis (Fabaceae) from Turkey, Caryologia, 68:4, 357-362, DOI: 10.1080/00087114.2015.1109926 To link to this article: https://doi.org/10.1080/00087114.2015.1109926

Published online: 27 Nov 2015.

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Full Terms & Conditions of access and use can be found at https://www.tandfonline.com/action/journalInformation?journalCode=tcar20 Caryologia: International Journal of Cytology, Cytosystematics and Cytogenetics, 2015 Vol. 68, No. 4, 357–362, http://dx.doi.org/10.1080/00087114.2015.1109926

Karyological studies on Oxytropis (Fabaceae) from Turkey Esra Martina, Seher Karaman Erkulb* and Zeki Aytaçc aNecmettin Erbakan University, Faculty of Science, Department of Biotechnology, Konya, Turkey; bAksaray University, Faculty of Arts and Science, Department of Biology, Aksaray, Turkey; cGazi University, Faculty of Science, Department of Biology, Teknikokullar, Ankara, Turkey

The karyological analyses of 11 species, one taxon of which is endemic of Oxytropis (Fabaceae), distributed in Turkey were examined in this study. Among these taxa, the chromosome numbers were as follows: 2n =16inOxytropis kotschyana, O. pallasii, O. pilosa, O. savellanica, O. persica, O. albana, O. argyroleuca, O. aucheri, O. karjaginii, O. lupinoides; and 2n =96inO. lazica. The karyotype analysis of taxa belonging to the genus Oxytropis was performed using Image Analysis System (Bs200Pro). Keywords: karyotype; Image Analysis System; Oxytropis; Turkey

1. Introduction Krusheva 1986; Probatova and Sokolovskaya 1986; Oxytropis DC. is considered the most closely related Rudyka 1986; Jurtsev 1988; Yan et al. 1989; Zhang and genus to Astragalus L. in its gross morphology. These Ma 1989; Lavrenko et al. 1990; Zakharjeva 1990; Jahan two genera co-exist in the same environment and habitats. et al. 1994; Starlinger et al. 1994; Stepanov 1994; Wang Although it differs from Astragalus only in keel petal et al. 1994; Zhang et al. 1994; Gu and Sun 1996; Pavlova (pointed versus obtuse) and pod septum (arising from 1996; Gervais et al. 1997; Favarger 1997; Aedo et al. adaxial suture versus abaxial), it was never considered in 1998; Filippov et al. 1998; Gervais and Blondeau 1999; Astragalus after its separation by De Candolle (1802). Zhu and Ohashi 2000; Yan et al. 1995, 2000; Volkova Recent molecular studies based on nrDNA ITS and et al. 2003). Several cytological publications have been chloroplast trnL intron data (Wojciechowski et al. 1999; written about Fabaceae family in recent years (Martin Wojciechowski 2005) have demonstrated that Oxytropis is et al. 2008; Pedro and Delgado-Salinas 2009; Hejazi monophyletic and not nested within Astragalus sensu et al. 2010; Sepet et al. 2011). stricto, but forms a separate clade within the large Astra- Karyological knowledge needs to be used in galean clade. It has also been shown that Oxytropis is a conjunction with other sources of data to achieve a better sister group of Astragalus which has Eurasian origin understanding of the cytologic relationship of Oxytropis fi (Ranjbar et al. 2010). Based on cytological (Ledingham taxa, leading to their natural classi cation. In this study, 1957, 1960) and morphological evidence, it has been sug- meristematic root tips were obtained from all species of gested that Oxytropis evolved in Eurasia (Chaudhary et al. the genus for karyotype analysis. As well as the underly- 2008). The genus Oxytropis has been revised by Karaman ing somatic chromosome numbers, the chromosome data Erkul and Aytaç (2013) who reduced its species to 11 and obtained by Image Analysis System were used to considered only one species as endemic to Turkey. analyze chromosome morphology. Cytogenetic studies of the genus Oxytropis are mostly based on the chromosome number. The somatic chromosome number of the genus has been found to be 2n = 16, 2. Materials and methods 24, 32, 48, 56, 58, 64, 80, 94, 96, 98 (Gurzenkov 1973; The specimens were collected and identified by SKE and Belaeva and Siplivinsky 1976; Rostovtseva 1977; ZA. Localities of the plant material used in this study Krogulevich 1976, 1978; Kovanda 1978; Engelskjon and collectors are shown in Table 1. 1979; Moraldo and Valva 1980; Krasnoborov et al. 1980; The karyological study of the genus Oxytropis within Zhukova and Petrovsky 1980; Andreev 1981; Astanova the taxa was conducted on the meristematic cells of root and Abdusaljamova 1981; Dawe and Murray 1981a, tips. The root cells, grown in the dark at room tempera- 1981b; Strid and Franzen 1981; Petrovsky and Zhukova ture, were laid into α-mono-bromonaphthalene solvent 1981; Löve and Löve 1982; Pogan 1982; Yurtsev and and kept for 16 h in the refrigerator during the pretreat- Zhukova 1982; Zhukova 1983; Krasnikova et al. 1983, ment procedure. Afterwards, the root tips were fixed in 1984; Gurzenkov and Pavlova 1984; Rostovtseva 1984; 3:1 absolute alcohol, glacial acetic acid and stored in a Strid and Andersson 1985; Ashraf and Gohil 1986; refrigerator in 70% alcohol. The root tips were painted at

*Corresponding author. Email: [email protected]

Table 1. The studied taxa and their collection locations.

Taxa Location O. argyroleuca Bornm. Ankara: Polatlı-Sazılar, gypsum slopes, 748 m, 21 July 2008, S. Karaman 2104. O. pallasii Pers. Erzurum: 21 km to Oltu, steppe, 1750 m, 2 June 2008, S. Karaman 2188. O. pilosa (L.) Dc. Artvin: 5 km from Artvin to Şavşat, road sides, 400 m, 10 May 2008, 29 June 2008, S. Karaman 2200. O. lazica Boiss. Rize: İkizdere, Ballıköy, Anzer plateau, alpine meadows, 2658–2750 m, 3 September 2008, S. Karaman 2206. O. aucheri Boiss. Ağrı:Doğu Beyazıt, calcareous slopes, 1813–1905 m, 8 July 2007, S. Karaman 2140. O. savellanica Bunge ex Niğde: Aladağ, Meydan plateau, alpine meadows, 2900 m, 5 July 2007, S. Karaman 2129. Boiss. O. lupinoides Grossh. Erzincan: Sürek, steppe, 1115 m, 8 August 2007, S. Karaman 2143. O. albana Steven Kars: Susuz, Kiziroğlu Village, Kısır Mountain, alpine meadows, 2940–3165 m, 10 July 2007, S. Karaman 2028. O. karjagini Grossh. Erzurum: Hınıs, near Aras River, calcareous slopes, 8 July 2007, S. Karaman 2141. O. persica Boiss. Niğde: Kızıltepe, alpine meadows, 2760 m, 25 September 2007, S. Karaman 2165. O. kotschyana Boiss. et Hakkari: Nehil stream, 62 km from Hakkari to Yüksekova, steppe, 1750 m, 27 May 2008, S. Karaman Hohen. & Z. Aytaç 2183 room temperature in 1N HCl with 2% aceto-orcein for 2 The somatic chromosome number of Oxytropis pilosa h. The mitotic metaphase cells determined under a light was obtained as 2n=16 (Figure 2c). The chromosome microscope were transferred to a computer. Finally, kary- length ranges between 1.53 and 2.42 μm. Chromosome ological analyses of the taxa were conducted with Image arm ratios were measured as 1.00–2.16 μm and the kary- Analysis System (Bs200Pro; http://www.bab.com.tr/ otype formula is 1M+5m+2sm. The centromeric index prgdis.php?prog_id=bssito&dilsec=1). values varied between 3.98 and 5.83, and relative lengths Chromosomes were classiﬁed using the nomenclature ranged from 10.09 to 15.98. The haploid chromosome of Levan et al. (1964). Karyotype analyses of mitosis length is 15.14 μm. The asymmetry index is 0.0002. An metaphase were obtained for 10 different root tips from ideogram of the taxon was drawn using the Image Anal- each taxa. ysis System (Figure 2c). The somatic chromosome number of Oxytropis savel- 3. Results lanica was obtained as 2n=16 (Figure 2d). The chromosome length ranges between 1.63 and 3.21 μm. The chromosome number and karyomorphology of Oxy- Chromosome arm ratios were measured as 1.30–2.03 μm tropis kotschyana, O. pallasii, O. pilosa, O. savellanica, and the karyotype formula is 6m+2sm. The centromeric O. persica, O. albana, O. lazica, O. argyroleuca, O. index values varied between 3.50 and 6.52, and relative aucheri, O. karjaginii, and O. lupinoides were deter- lengths ranged from 8.83 to 17.37. The haploid chromo- mined. Detailed morphological characteristics of the some length is 18.49 μm. The asymmetry index is somatic chromosomes are given in Figure 1. The idio- 0.0005. An ideogram of the taxon was drawn using the gram of taxa is shown in Figure 2. The characteristics of Image Analysis System (Figure 2d). somatic chromosomes in studied taxa are given below. The somatic chromosome number of Oxytropis The somatic chromosome number of Oxytropis persica was obtained as 2n=16 (Figure 2e). The kotschyana was obtained as 2n = 16 (Figure. 2a). The chromosome length ranges between 2.83 and 4.25 μm. chromosome length ranges between 1.56 and 3.03 μm. Chromosome arm ratios were measured as 1.18–2.23 μm Chromosome arm ratios were measured as 1.29–1.65 μm and the karyotype formula is 6m+2sm. The centromeric and the karyotype formula is 8m. The centromeric index index values varied between 3.31 and 7.12, and relative values varied between 3.32 and 6.71, and relative lengths lengths ranged from 10.33 to 15.50. The haploid chro- ranged from 8.38 to 16.26. The haploid chromosome mosome length is 27.45 μm. The asymmetry index is length is 18.66 μm. The asymmetry index is 0.0004. An 0.0003. An ideogram of the taxon was drawn using the ideogram of the taxon was drawn using the Image Anal- Image Analysis System (Figure 2e). ysis System (Figure 2a). The somatic chromosome number of Oxytropis The somatic chromosome number of Oxytropis pal- albana was obtained as 2n=16 (Figure 1f). The lasii was obtained as 2n=16 (Figure 2b). The chromo- chromosome length ranges between 2.30 and 3.64 μm. some length ranges between 1.41 and 2.10 μm. Chromosome arm ratios were measured as 1.30–2.30 μm Chromosome arm ratios were measured as 1.31–2.13 μm and the karyotype formula is 7m+1sm. The centromeric and the karyotype formula is 6m+2sm. The centromeric index values varied between 3.75 and 6.71, and relative index values varied between 4.12 and 5.67, and relative lengths ranged from 9.87 to 15.59. The haploid chromo- lengths ranged from 10.04 to 14.91. The haploid chro- some length is 23.36 μm. The asymmetry index is mosome length is 14.07 μm. The asymmetry index is 0.0003. An ideogram of the taxon was drawn using the 0.0001. An ideogram of the taxon was drawn using the Image Analysis System (Figure 2f). Image Analysis System (Figure 2b). Caryologia: International Journal of Cytology, Cytosystematics and Cytogenetics 359

Figure 1. Mitotic metaphase chromosomes of Oxytropis. (a) O. kotschyana;(b)O. pallasii; (c) O. pilosa; (d) O. savellanica; (e) O. persica;(f)O. albana; (g) O. lazica;(h)O. argyroleuca; (i) O. aucheri;(j)O. karjaginii;(k)O. lupinoides. Bar: 10 μm.

The somatic chromosome number of Oxytropis lazica chromosome length ranges between 1.76 and 3.30 μm. was obtained as 2n=96 (Figure 2g). The chromosome Chromosome arm ratios were measured as 1.15–1.67 μm length ranges between 0.79 and 3.50 μm. Because the and the karyotype formula is 7m+1sm. The centromeric chromosomes of the taxon are very small, centromeres index values varied between 3.25 and 6.89, and relative and the type of chromosomes could not be determined lengths ranged from 9.13 to 17.10. The haploid chromo- and chromosome morphology of the mitotic metaphase some length is 19.29 μm. The asymmetry index is chromosomes of taxon is investigated according to their 0.0004. An ideogram of the taxon was drawn using the total length. The haploid chromosome length is 84.46 Image Analysis System (Figure 2i). μm. The asymmetry index is 0.0004. An ideogram of the The somatic chromosome number of Oxytropis kar- taxon was drawn using the Image Analysis System jaginii was obtained as 2n=16 (Figure 2j). The chro- (Figure 2g). mosome length ranges between 1.32 and 2.76 μm. The somatic chromosome number of Oxytropis Chromosome arm ratios were measured as 1.08–1.63 μm argyroleuca was obtained as 2n=16 (Figure 2h). The and the karyotype formula is 8m. The centromeric index chromosome length ranges between 1.65 and 2.52 μm. values varied between 3.36 and 6.93, and relative lengths Chromosome arm ratios were measured as 1.17–2.00 μm ranged from 7.60 to 15.82. The haploid chromosome and the karyotype formula is 7m+1. The centromeric length is 17.43 μm. The asymmetry index is 0.0006. An index values varied between 4.31 and 6.79, and relative ideogram of the taxon was drawn using the Image Anal- lengths ranged from 9.70 to 14.81. The haploid chromo- ysis System (Figure 2j). some length is 17.01 μm. The asymmetry index is The somatic chromosome number of Oxytropis lupi- 0.0002. An ideogram of the taxon was drawn using the noides was obtained as 2n=16 (Figure 1k). The chro- Image Analysis System (Figure 2h). mosome length ranges between 1.77 and 3.18 μm. The somatic chromosome number of Oxytropis Chromosome arm ratios were measured as 1.24–1.68 μm aucheri was obtained as 2n=16 (Figure 2i). The and the karyotype formula is 8m. The centromeric index 360 E. Martin et al.

Figure 2. Idiograms of Oxytropis taxa. (a) O. kotschyana;(b)O. pallasii; (c) O. pilosa; (d) O. savellanica; (e) O. persica;(f)O. albana; (g) O. lazica;(h)O. argyroleuca; (i) O. aucheri;(j)O. karjaginii;(k)O. lupinoides. Bar: 10 μm. values varied between 3.47 and 6.65, and relative lengths O. albana, O. argyroleuca, and O. aucher is 7m+1sm, ranged from 8.95 to 16.03. The haploid chromosome and O. pilosa is 1M+5m+2sm. This also indicates that length is 19.84 μm. The asymmetry index is 0.0003. An chromosome morphology between species is specificto ideogram of the taxon was drawn using the Image Anal- these taxa. ysis System (Figure 2k). The shortest chromosome was 0.79 μm(O. lazica), and the longest chromosome was 4.25 μm(O. persica). O. pilosa had the smallest arm ratio (1.00), and O. 4. Discussion albana had the largest arm ratio (2.30). According to the It is determined that the caryotype formule of Oxytropis centromeric index, the O. aucheri had the smallest index kotschyana, O. karjaginii, and O. lupinoides is 8m, value (3.25), and O. persica. O. karjaginii had the lar- O. pallasii, O. savellanica,andO. persica is 6m+2sm, gest (7.12). O. pallasii had the shortest total length of Caryologia: International Journal of Cytology, Cytosystematics and Cytogenetics 361 haploid complement (14.07 μm), and O. persica had the Engelskjon T. 1979. Chromosome numbers in vascular plants longest (27.45 μm). O. pallasii had the smallest asymme- from Norway, including Svalbard. Opera Bot. 52:1–38. try index (0.0001), and O. karjaginii had the largest Favarger C. 1997. Notes de caryologie alpienne VI. Bull Soc Neuchateloise Sci Nat. 120:19–33. asymmetry index (0.0006). Filippov EG, Kulikov PV, Knyazev MS. 1998. 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