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CONTEMPORARY INTER-SPECIFIC HYBRIDIZATION BETWEEN Cirsium Aduncum and C

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CONTEMPORARY INTER-SPECIFIC HYBRIDIZATION BETWEEN Cirsium Aduncum and C UDC 575:630 DOI: 10.2298/GENSR1602497S Original scientific paper CONTEMPORARY INTER-SPECIFIC HYBRIDIZATION BETWEEN Cirsium aduncum AND C. Haussknechtii (Asteraceae): EVIDENCE FROM MOLECULAR AND MORPHOLOGICAL DATA Masoud SHEIDAI*1, Mona NAJI1, Zahra NOORMOHAMMADI2, Maryam NOUROOZI3, Somayeh GHASEMZADEH-BARAKI1 1Faculty of Biological Sciences, Shahid Beheshti University, Tehran, Iran 2 Department of Biology, School of Sciences, Sciences and Research Branch, Islamic Azad University, Tehran, Iran 3Department of Horticulture, collage of Aburaihan, University of Tehran, Tehran Sheidai M., Ma Naji, Z. Noormohammadi, M.Nouroozi, S. Ghasemzadeh-Baraki (2016): Contemporary inter-specific hybridization between Cirsium aduncum and C. haussknechtii (Asteraceae): evidence from molecular and morphological data- Genetika, Vol 48, No. 2,497-514. Cirsium aduncum Fisch. & C.A.Mey. Ex DC. and C. haussknechtii Boiss., (Asteraceae) are important medicinal plant species that grow in different geographical regions of Iran. We had no knowledge about population genetic structure, intra-specific and inter-specific gene flow and the presence of hybrid zone for this two species in Iran. Therefore, in order to provide data for conservation of these two medicinally important species, the population genetic analysis and morphometric studies were performed in 18 geographical populations of these species. ANOVA and MDS analyses revealed significant morphological difference among the studied populations in either species, while MDS plot showed morphological overlap in plants of these two species. AMOVA test revealed significant genetic difference among the studied populations. Mantel test showed positive significant correlation between genetic and geographical distances and the occurrence of isolation by distance. Population assignment test and STRUCTURE plot of genetic data revealed inter-specific introgression between these species. Keywords: AMOVA, Circium, gene flow, population assignment, STRUCTURE analysis. ___________________________ Corresponding author: Masoud Sheidai, Faculty of Biological Sciences, Shahid Beheshti University, Tehran, Iran, email: [email protected], Tel: +98 9122593378 498 GENETIKA, Vol. 48, No.2, 497-514, 2016 INTRODUCTION The genus Cirsium Mill. (Asteracea) contains about 250 perennial, biennial or rarely annual spiny species. These species grow in the Northern hemisphere, Europe, North Africa, Siberia, Central Asia, West and East Africa, as well as Central America (ZOMLEFER, 1994; BURES et al., 2004; SEGARRA-MORAGUES et al., 2007). Cirsium species show wide ranges of ecological adaptations in places they grow and occupy different localities with regard to elevation, temperature and edaphic factors due to their genetic adaptability and plasticity. They also are well known for interspecific hybrid formation and wide gene flow, which helps their adaptation to various environmental conditions (BUREŠ et al., 2004). Most of Cirsium species are of medicinal values and have been used to reduce fever, blood cholesterol and triglyceride level and also used against jaundice. Some of the Cirsium species have been used as food plants (ORHAN et al., 2013; ABBET et al., 2014). Population genetics study particularly analysis of genetic diversity is important in plant conservation and molecular ecology studies (FREELAND et al., 2011). It provides data on the genetic variability, gene flow versus population genetic isolation, population genetic fragmentation, the role of genetic drift, the bottleneck and any other evolutionary forces acting on populations, divergence (SHEIDAI et al., 2012; 2013; 2014). Population genetic studies can also trough light on hybridization and inter-specific gene flow between the species that grow in overlapping geographical regions (ALBALADEJO and APARICIO, 2007, IJBARI et al., 2014). Cirsium aduncum Fisch. & C.A.Mey. Ex DC. and C. haussknechtii Boiss., have medicinal and food values and are widely used by locals. We have no information available about genetic diversity, population genetic structure and gene flow among these medicinal species in Iran. Therefore, the present investigation was carried out to provide data about population genetic structure and morphological variability of 18 geographical populations that may be of help in conservation of these species. Multilocus molecular markers such as simple sequence repeat (SSR) markers and inter- simple sequence repeat (ISSR) markers are useful in studying introgressive hybridization (GASKIN and KAZMER, 2009). Therefore, for population genetic study and to illustrate of inter-species gene flow, we used ISSR markers that are reproducible, simple to work and not expensive and were shown to be informative for genetic diversity and population structure studies in the genus Cirsium (SEIF et al., 2012; NOUROOZI et al., 2013; SHEIDAI et al., 2013). The genus Cirsium is well known for production of inter-specific hybrids and the occurrence of morphological variability and intergradation of diagnostic characters (DABEYDEEN, 1980). Therefore, we studied morphological variability in the studied species and populations too. The intergradation of diagnostic morphological characters along with genetic admixture have been considered as strong evidence for inter-specific genetic introgression (DABEYDEEN, 1980; GALBANY-CASALS et al., 2012; IJBARI et al., 2014). MATERIALS AND METHODS Plant materials Sixty-one plant accessions were collected from 18 geographical populations of Cirsium aduncum Fisch. & C.A.Mey. Ex DC. and C. haussknechtii Boiss. Details of localities are provided in Table 1. Voucher specimens are deposited in Herbarium of Shahid Beheshti M.SHEIDAI et al: INTERSPECIFIC INTROGRESSION IN CIRCIUM 499 University (HSBU). Fresh leaves were collected and used for DNA extraction and molecular study. Table 1. Circium species and populations studied, their locality and voucher No. Population Species Latitude Longitude Localities and voucher No. East Azarbaijan:Mianeh-Bostanabad, 1844 m, 1 C. aduncum 37°43'31.95" 46°58'0.19" Nouroozi, 2010500-HSBU. East Azarbaijan: Tabriz-Ahar,1794m, 2 C. aduncum 38°21'39.77" 46°51'42.79" Naji2010506HSBU. East Azarbaijan : Mianeh-Bostanabad , 1830 m , 3 C. aduncum 37°42'33.90" 46°58'50.34" Naji , 2010507-HSBU. East Azarbaijan : Ahar-Meshkinshahr , 1257 m , 4 C. aduncum 38°25'41.09" 47°15'5.17" Naji , 2010508-HSBU. East Azarbaijan : Marana-Tabriz , 1736 m , Naji , 5 C. aduncum 38°20'40.23" 45°50'1.95" 2010509-HSBU. East Azarbaijan : Bostanabad-Mianeh , 1778 m , 6 C. aduncum 37°51'13.29" 46°47'9.22" Nouroozi , 2010501-HSBU. Ardebil: Meshkinshahr, Mazraejahan, 1169 m, 7 C. aduncum 38°22'47.45" 47°29'28.83" Nouroozi & Fathollahi, 8600170-HSBU. Ardabil : Meshkinshahr-Ardebil , 1400 m , 8 C. aduncum 38°29'17.29" 48° 0'0.59" Nouroozi , 8600172-HSBU. East Azarbaijan : Ahar-Meshkinshahr , 1300 m , 9 C. aduncum 38°26'50.12" 47°13'26.53" Nouroozi , 2010502-HSBU. West Azarbaijan : Qatur-khoy , 1500 m , Nourooz , 10 C. aduncum 38°27'1.91" 44°40'25.73" 8600171-HSBU. East Azarbaijan : Bostanabad-Mianeh , 1675 m , 11 C. aduncum 37°38'33.40" 47° 4'46.38" Nouroozi , 2010503-HSBU. East Azarbaijan : Ahar - tabriz , Goijabel , 1850 m , 12 C. aduncum 38°23'1.50" 46°49'45.26" Nouroozi , 2010504-HSBU. East Azarbaijan : Kandavan, Hilevar village, 1600 13 C. haussknechtii 37°54'28.96" 46° 9'47.88" m, Nouroozi, 8600187-HSBU. East Azarbaijan : Ahar - kaleibar , 1200 m , 14 C. haussknechtii 38°49'56.61" 47° 3'15.91" Nouroozi , 8600190-HSBU. East Azarbaijan : Jolfa , kiamaki , Daran village , 15 C. haussknechtii 38°49'20.98" 45°49'16.88" 1455 m , Nouroozi , 8600188-HSBU. West Azarbaijan : Urmia-Salmas , 1380 m , 16 C. haussknechtii 38°10'58.80" 44°48'11.91" Nouroozi , 8600191-HSBU. Ardabil : Ardebil - khalkhal , 1615 m , Nouroozi , 17a C. haussknechtii 37°42'19.04" 48°28'11.95" 2010505-HSBU. Ardabil : Ardebil - khalkhal , 1650 m , Nourooz , 17b C. haussknechtii 37°53'29.04" 48°23'36.84" Nouroozi , 8600189-HSBU. Ardabil : Ardebil-Namin , 1353 m , Naji , 2010510- 18 C. haussknechtii 38°23'28.89" 48°26'42.05" HSBU. Morphological studies Morphological characters studied included quantitative and qualitative characters that are given in Table 2. 500 GENETIKA, Vol. 48, No.2, 497-514, 2016 Table 2. Morphological characters studied. 1 Pappus length 2 Capitulum length (With floret) 3 Involucre length 4 Involucre width 5 Length of the involucre bract 6 Width of the involucre bract 7 Length the terminal spine in Involucre Bract 8 Length of the middle lobe of leaf 9 Width of the middle lobe of leaf 10 Length of spine in the middle lobe leaf 11 Length of the terminal spine in Leaf 12 Length of the spine in leaf margin 13 Corolla length 14 Corolla tube length 15 Lamina length 16 Length of the shortest corolla lobe 17 Length of the longest corolla lobe 18 Ratio of length to width of involucre 19 Ratio of length to width of involucre bract 20 Ratio of lamina to corolla tube 21 Ratio of corolla tube to corolla 22 Ratio of lamina to corolla 23 Ratio of tallest to shortest corolla lobe 24 Ratio of length to width of middle lobe in leaf 25 Ratio of length to width of leaf 26 Ratio of middle lobe spine to terminal spine in leaf 27 Cover of the upper surface of leaf 28 Cover of the lower surface of leaf 29 Stem cover 30 Position of the last leaf to capitulum 31 Cover of the involucre bract M.SHEIDAI et al: INTERSPECIFIC INTROGRESSION IN CIRCIUM 501 DNA extraction and ISSR assay Fresh leaves were collected randomly in each of the studied populations and dried in silica gel powder. Genomic DNA was extracted using CTAB activated charcoal protocol (SHEIDAI et al., 2013). The quality of extracted DNA was examined by running on 0.8% agarose gel. Ten ISSR primers; (AGC)5GT, (CA)7GT, (AGC)5GG, UBC810, (CA)7AT, (GA)9C, UBC807, UBC811, (GA)9T and (GT)7CA commercialized by UBC (the University of British Columbia) were used. PCR reactions were performed in a 25μl volume containing 10 mM Tris- HCl buffer at pH 8; 50 mM KCl; 1.5 mM MgCl2; 0.2 mM of each dNTP (Bioron, Germany); 0.2 μM of a single primer; 20 ng genomic DNA and 3 U of Taq DNA polymerase (Bioron, Germany).
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