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Infraspecific Morphological and Genome Size Variations in Linum Glaucum in Iran

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Infraspecific Morphological and Genome Size Variations in Linum Glaucum in Iran BIODIVERSITAS ISSN: 1412-033X Volume 16, Number 1, April 2015 E-ISSN: 2085-4722 Pages: 69-78 DOI: 10.13057/biodiv/d160109 Infraspecific morphological and genome size variations in Linum glaucum in Iran SEYED MEHDI TALEBI1,♥, MASOUD SHEIDAI2, MORTEZA ATRI3, FARIBA SHARIFNIA4, ZAHRA NOORMOHAMMADI5 1Department of Biology, Faculty of Sciences, Arak University, Arak, 38156-8-8349 Iran. Tel. +98-863-4173317. ♥email:[email protected]. 2Shahid Beheshti University, GC, Faculty of Biological Sciences, Tehran, Iran. 3Bu-Ali Sina Hamedan University, Hamedan, Iran. 4Department of Biology, North Tehran Branch, Islamic Azad University, Tehran Iran. 5Department of Biology, School of Sciences, Tehran Science and Research Branch, Islamic Azad University, Tehran, Iran Manuscript received: 27 December 2014. Revision accepted: 20 January 2015. Abstract. Talebi SM, Sheidai M, Atri M, Sharifnia F, Noormohammadi Z. 2015. Infraspecific morphological and genome size variations in Linum glaucum in Iran. Biodiversitas 16: 69-78. There are many discussions about taxonomic position of Linum glaucum in different Flora. In this study, the morphological traits associated with the nuclear genome size were used for identification of infraspecific variations in the nine populations of L. glaucum. Twenty three qualitative and quantitative morphological characteristics were investigated. The Analysis of variance tests showed significant difference for some morphological features. The Canonical Correspondence Analysis of habitat ecological factors showed that each of the habitats had prominent characteristics, and also Pearson’s coefficient of correlation confirmed the significant correlations between the morphological features in relation to ecological factors. In the morphological Unweighted Paired Group using Average method tree, populations were separated from each other, so that population No. 4 was separated from others. In the flow cytometery investigations, variation of about 1.19 times was presented between the maximum and minimum averages of the genome sizes in the populations. Minimum amount of genome size was found in populations No. 4. Significant correlations occurred between the nuclear genome size with habitat elevation as well as some of the quantitative morphological features. The mentioned variations caused to difference between the populations and lead to creation of ecotype and ecophene in the studied populations. Key words: Ecology, genome size, morphology, population. INTRODUCTION strile (Rechinger 1974). In 1967, Davis named it as L. austriacum L. subsp. glaucescens, following Rechinger Biodiversity is mostly investigated at the species level (1974) recognized this taxon as a synonym of L. glaucum. as well as higher taxonomic ranks. However, the basis for Therefore there are three different synonyms for L. biodiversity and organic evolution is the genetic difference glaucum in different Flora such as Flora Iranica (Rechinger within species (Ramel 1998). Linhart and Grant (1996) 1974) and Flora of Iran (Sharifnia and Assadi 2001). suggested that Comparable developmental variation Although L. glaucum is morphologically very similar to presents between different populations of a species L. austriacum, the other member of this section, but in (intraspecific), which likely resounds verifications to different taxonomical treatments such as palynology different natural environments and it is the source of plant (Talebi et al. 2012a), seed micromorphology (Talebi et al. species differentiation. Therefore, humans have used these 2012b), anatomy (Rashnou-Taei 2014) and also molecular infraspecefic variations for the domestication and genetic investigation (ISSR) (Talebi 2013) these species placed improvement of many of plant species (Diamond 2002). separately. The obtained results of these studies showed Linum glaucum Boiss. & Nöe belongs to section Linum that L. glaucum and L. austriacum are separate species. of the genus Linum L. This species is an Irano-Turanian In addition to morphological characteristics, variation element which naturally spread in the slops of the Zagrous between populations can be investigated with using 2C- Mountains in the different provinces of Iran such as value or nuclear genome size. One of the most important Kurdistan, Kermanshah, Zanjan, Hamadan, Lurestan as features of all living organisms is nuclear DNA amount. well as Markazi. In addition this species are found in the Swift (1950) definite C-value as DNA content in the neighboring countries as Iraq and Turkey (Rechinger 1974; unreplicated haploid nucleus. Nuclear DNA amount is Mobayen 1996; Sharifnia and Assadi 2001). positively related to the number of cellular traits such as: There are many discussions about taxonomic position time of mitotic cell cycle, nuclear DNA synthesis rate, the of L. glaucum. This taxon was described for the first time size of cell and its nuclear, through to the whole plant traits as L. alpinum Jacq, var. glaucescens Boiss. in Flora such as minimum generation time and geographic Orientalis (Boissier 1867), then Stapf recognized it as L. distribution (Bennett 1972, 1987; Kidd et al. 1987). Studies 70 BIODIVERSITAS 16 (1): 69-78, April 2015 confirmed that infraspecific variation in nuclear genome measured in picograms (pg) and the nuclei status described size amount is most significant for plant taxonomy as an in terms of ‘C’ value (Doležel et al. 2007). Both of the C- indicator of taxonomic heterogeneity (Murray 2005). value as well as the genome size can be expressed either in Because, the lacks of previously infraspecific study for DNA picograms (=10-9 g) or mega base pairs (1 pg = 978 L. glaucum in Iran as well as the world, we have used for Mbp). The nuclear DNA amount of the studied samples the first time in the present work morphological calculated on the basis of the values of the G1/G2 peak characteristics as well as nuclear genome size for means (Doležel and Bartoš 2005). identification and determining kind and level of infraspecific variations between the nine populations of this species in Iran. Ecological factors MATERIAL AND METHODS In order to compare the effect of different environmental factors on the nuclear genome size and Plant material morphological characteristics of L. glaucum populations, Nine populations of L. glaucum were randomly five factors were examined for each population’s habitat, collected from western regions of Iran during spring 2011 such as: longitude (E˚), latitude (N˚), altitude (in meters), (Table 1). Plant samples were identified based on the average of annual minimum and maximum temperature (in descriptions provided in accessible references such as Flora C˚). Longitude, latitude and altitude were calculated with Iranica (Rechinger 1974) and Flora of Iran (Sharifnia and Garmin GPS map76CSx, and the averages annual Assadi 2001). The voucher specimens were deposited in minimum and maximum temperature for each population the herbarium of Shahid Beheshti University (HSBU). were taken from the web site of the meteorological From each population four samples were selected randomly organization of Iran. and then examined for their nuclear DNA content and morphological features. Statistical analysis The mean and standard deviation of the studied Morphometry morphological characteristics were determined. In order to Twenty three quantitative and qualitative morphological group the studied populations on the basis of characteristics from the both of vegetative and reproductive morphological features, data were standardized (mean = 0, organs of this species were examined. The studied variance = 1) and used for multivariate analyses including morphological characteristics include: stem lengths and its Unweighted Paired Group using Average method branch number, stem diameter, shape, length and width of (UPGMA) as well as Principal Coordinate Analysis basal and floral leaves, the shape of apex, margin and base (PCoA) (Podani 2000). Analysis of variance (ANOVA) test of basal and floral leaf blade, basal leaf diameter, calyx was employed to assess the significant quantitative dimensions as well as corolla color and dimensions. Four morphological characteristics difference among the studied replications were used for quantitative characters populations, and Pearson’s coefficient of correlation was measurements and the used terminologies for qualitative used to determine significant correlations of nuclear morphological traits were on the basis of descriptive genome size and quantitative morphological traits in terminology provided by Stearn (1983). relation to ecological factors, as well as, longitude, latitude, altitude, average of annual minimum and maximum Flow cytometry temperature, so as to show possible relationship between In order to preparation of nuclei suspensions, small populations. The multivariate method, Canonical amounts of mature fresh leaf tissue together with an equal Correspondence Analysis (CCA), was used to elucidate the weight of mature leaf tissue of the standard reference were relationship between the studied populations and their used. In this study Allium cepa L. was used as standard environmental factors. NTSYS ver. 2 (1998) and SPSS ver. references, which has a 2C DNA value of 33.5 pg 9 (1998) softwares were used for statistical analyses. (Greilhuber and Ebert 1994). For preparation of the nuclear suspension two-stepped method was employed. Plant materials were chopped with RESULTS AND DISCUSSION a sharp scalpel with adding 400 μl nuclei isolation buffer in the plastic
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