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Cytogenetic Studies in Some Species of Melica L. and Milium L. in Iran

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Cytogenetic Studies in Some Species of Melica L. and Milium L. in Iran © 2009 The Japan Mendel Society Cytologia 74(2): 185–193, 2009 Cytogenetic Studies in Some Species of Melica L. and Milium L. in Iran Masoud Sheidai* and Khaled Moghaddam Shahid Beheshti University, GC, Faculty of Biological Sciences, Tehran, Iran Received August 22, 2008; accepted July 20, 2009 Summary Cytogenetical studies were performed on 15 populations of 7 species and subspecies of the genus Melica as well as 4 populations of 2 species of the genus Milium growing wild in Iran. The meiotic analysis of Melica and Milium species studied showed the occurrence of a post pachytene diffuse stage possibly due to adaptation to adverse environmental conditions. Melica species studied possess 2nϭ2xϭ18 chromosome number. The chromosome numbers of M. persica subsp. persica and M. persica subsp. inaequiglumis have been reported for the first time. The ANOVA test per- formed among Melica species and populations studied, revealed a significant difference in the amount of total, terminal and intercalary chiasmata as well as amount of rod bivalents (pϽ0.01), in- dicating partly their genetic differences. Two populations of Milium vernale showed the presence of 2nϭ2xϭ14 and 2nϭ2xϭ18, while two populations of M. schmidtianum possessed 2nϭ2xϭ14 chro- mosome number, which is new for M. chmidtianum. Key words Cytogenetic, Melica, Milium, Iran. The genus Melica L. is the largest genus of the family Meliceae (Gramineae, Pooideae), com- prised of about 80 species of perennial rhizomatous grasses, distributed all over the world except tropical regions and Australia (Clayton and Renovize 1986), mainly grown in North temperate, Southern Africa and South America and are mesophytic to xerophytic; shade species and species of open habitats. The Melica species are bisexual, with bisexual spikelets; with hermaphrodite florets; exposed- cleistogamous, or chasmogamous. The inflorescence is few spikeleted to many spikeleted; a single raceme, or paniculate; open, or contracted; with capillary branchlets, or without capillary branch- lets. The genus Melica includes phylogenetically old forms in East Asia and many forms (in a con- tinuous evolution) in the (semi) arid regions of West Eurasia, North and South America. The genus chromosome base number is xϭ9 and diploid and tetraploid levels have been reported. The somatic chromosome number of 2nϭ14 has been rarely reported in this genus (Dallwitz and Watson 1992). Due to the presence of a large number of species in the genus Melica, controversy exists about taxonomic treatment of the genus and different authors have considered different number of subgenera and sections for this genus (for example see: Stebbins 1982, 1986, Tzvelev 1976, Torres 1980). Polyploidy and inter-specific hybridization is considered to be of high importance in the evolu- tion of Gramineae (Stebbins 1982, 1986), however the genus Melica is an exception by possessing a very homogenous group of mainly diploid species with the rare occurrence of change in the basic chromosome number and a very low level of inter-specific hybridization (Teresa and Bibsy 2000). Many species of the genus Melica are of limited distribution and are of less fodder value due to the presence of prussic acid which is harmful to the cattle. However two species of M. cupani and M. ciliata are known to be of forage value. Controversy also exists about the number of Melica species growing in Iran, as Parsa (1950) * Corresponding author, e-mail: [email protected], [email protected] 186 M. Sheidai and K. Moghaddam Cytologia 74(2) reported 5 species from Iran while Bor (1970) in Flora Iranica reported 8 species from the country including 2 sub-species for M. persica, 3 sub-species for M. jacquemontii and 2 varieties for M. cil- iata. Therefore according to Bor (1970), in total 12 species, sub-species and varieties of Melica grow in Iran. Recently, Assadi (1996) has reported M. picta from Iran. The genus Milium L. (Aveneae, Poaceae) is comprised of 3–7 annual or perennial stolonifer- ous, or caespitose species, mainly grown in the North temperate regions of the old world and also the east part of North America. They are commonly adventive, mesophytic and xerophytic; shade species and species of open habitats. The species are bisexual, having bisexual spikelets with her- maphrodite florets. The inflorescence is paniculate; open; with capillary branchlets; espatheate; not comprising ‘partial inflorescences’ and foliar organs. The chromosome base number of the genus Milium is xϭ4, 5, 7, and 9 and diploid, tetraploid and hexaploid levels have been reported (Clayton and Renovize 1986, Tzvelev 1976, Simon and Tomas 1991, Dallwitz and Watson 1992). Parsa (1950) reported 1 species of M. vernale with 2 varieties from Iran while Bor (1970) in Flora Iranica reported 4 species from the country. The present report is a part of biosystematic study of two genera of Melica and Milium concerned with cytogegetic analysis of 15 populations of 7 species and subspecies of the genus Melica as well as 4 populations of 2 species of the genus Milium growing wild in Iran for the first time. According to our knowledge, almost all the meiotic reports of the present study regarding the chiasma frequency and distribution as well as chromosome pairing are new to science. The chromo- some number of Melica picta and M. persica subsp. persica and a new ploidy level (2nϭ2xϭ14) in Milium schmidtianum have been reported for the first time. Materials and methods Plant material Meiotic studies were performed in 15 populations of 7 species and subspecies of the genus Melica as well as 4 populations of 2 species of the genus Milium growing wild in Iran. The Melica species studied are: 1–M. persica Kunth subsp. persica (2 populations), 2–M. persica Kunth subsp. inaequiglumis (Boiss.) Bor (2 populations), 3–M. jacquemontii Decne subsp. jacquemontii (1 popu- lation), 4–M. jacquemontii Decne subsp. canescens (Regel) Bor (1 population), 5–M. jacquemontii Decne subsp. hohenackeri (Boiss.) Bor (2 populations), 6–M. ciliata L. var. ciliata (2 populations), and 7–M. picta C. Koch. (1 population). The Milium species studied are: 8–M. vernale L. (2 popu- lations), and 9–M. schmidtianum C. Koch. (2 populations). The voucher specimens are deposited in the Herbarium of Shahid Beheshti University (HSBU) and TARI. Cytological preparation and meiotic analysis Young flower buds were collected from 10 randomly selected plants of each species/population and fixed in glacial acetic acid : ethanol (1 : 3) for 24 h. Flower buds were washed and preserved in 70% ethanol at 4°C until used (Sheidai et al. 2002). Cytological preparations used the squash tech- nique and 2% aceto-orcein as the stain. Fifty to 100 pollen mother cells (PMCs) were analysed for chiasma frequency and distribution at diakinesis metaphase stage and 500 PMCs were analysed for chromosome segregation during the anaphase and telophase stages. The Pearson correlation test was performed for meiotic characteristics among populations studied. The analysis of variance (ANOVA) followed by the Least Significant Test was performed among populations and species studied to detect significant difference in the relative chiasma fre- quency and distribution as well as chromosomes association (Sheidai et al. 2006). In order to group the species studied based on similarity in their meiotic characteristics, differ- ent clustering methods of UPGMA (unweighted paired group with arithmetic average) and WARD 2009 Cytology of Melica and Milium 187 Fig. 1. Representative meiotic cells in Melica and Milium species and populations studied. AϭMeiotic cell showing nϭ9 in Vardavard population of M. persica subsp. persica. BϭMeiotic cell showing nϭ9 and one B-chromosome (arrow) in Mazandaran population of M. ciliata var. ciliate. CϭMeiotic cell showing nϭ9 in Gilan population of Milium vernale. DϭMeiotic cell showing nϭ9 in Tehran popu- lation of M. persica subsp. persica. EϭMeiotic cell showing nϭ9 in M. jacquemontii subsp. jacque- montii. FϭMeiotic cell showing nϭ7 in Mazandaran population of Milium vernale. GϭMeiotic cell showing nϭ9 in Gajereh population of M. jacquemontii subsp. hohenackeri. HϭMeiotic cell show- ing nϭ7 in Ardebil population of M. schmidtianum. IϭMeiotic cell showing nϭ9 in M. picta. JϭMeiotic cells showing a single quadrivalent (arrow) in Arasbaran population of M. schmidtianum. (minimum spherical cluster method) as well as ordination based on principal coordinate analysis (PCO) were performed (Sheidai and Noormohammadi 2005). In order to avoid the chaining effect of clustering methods, the Neighbor joining method was also performed on cytogenetic data. The Euclidean and taxonomic distances were used as dissimilarity coefficient in cluster analy- sis of cytological data (Sheidai et al. 2006). SPSS Ver. 9 (1998) and NTSYS Ver. 2.2 (1998) were used for numerical analyses. 188 Table 1. Meiotic characteristics in Melica and Milium species studied. Sp n TX IX TOX RB RD I IV TXN IXN TOXN RBN RDN IN IVN per 1 9 15.33 0.27 15.63 6.87 2.13 0.00 0.00 1.70 0.03 1.74 0.76 0.24 0.00 0.000 per 2 9 14.61 0.56 15.17 6.94 1.86 0.08 0.00 1.62 0.06 1.69 0.77 0.21 0.01 0.000 ina 1 9 15.87 0.37 16.23 7.60 1.40 0.00 0.00 1.76 0.04 1.80 0.84 0.16 0.00 0.000 ina 2 9 14.23 1.26 15.49 7.00 1.17 0.06 0.00 1.58 0.14 1.72 0.78 0.13 0.01 0.000 jac 9 11.29 0.32 11.62 5.88 3.11 0.02 0.00 1.25 0.04 1.29 0.65 0.35 0.00 0.000 M. SheidaiandK.Moghaddam can 9 17.23 0.15 17.38 8.85 0.15 0.00 0.00 1.91 0.02 1.93 0.98 0.02 0.00 0.000 hoh 1 9 16.39 0.52 16.88 8.47 0.50 0.06 0.00 1.82 0.06 1.88 0.94 0.06 0.01 0.000 hoh 2 9
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