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Cytomorphological Study of Trigonella Disperma (Fabaceae) in Iran

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Cytomorphological Study of Trigonella Disperma (Fabaceae) in Iran © 2011 The Japan Mendel Society Cytologia 76(3): 279–294 Cytomorphological Study of Trigonella disperma (Fabaceae) in Iran Massoud Ranjbar*, Roya Karamian and Zahra Hajmoradi Department of Biology, Herbarium Division, Bu-Ali Sina University, P.O. Box 65175/4161, Hamedan, Iran Received May 5, 2010; accepted April 28, 2011 Summary Chromosome number, meiotic behaviour and morphological characters related to habit and pollen grains were studied in 8 populations of Trigonella disperma of Trigonella sect. Ellipticae native to Iran. All populations are diploid and possess a chromosome number of 2n=2x=16, which is consistent with the proposed base number of x=8. This taxon displayed regular bivalent pairing and chromosome segregation at meiosis. However, some meiotic abnormalities observed included varied degrees of sticky chromosomes with laggards and bridges in metaphase I or II, asynchronous nuclei in metaphase II, desynapsis in metaphase I, and cytomixis. This paper reports the first known study of the meiotic chromosome number and behaviour of T. disperma. We evaluated and determined the population limits within T. disperma, employing multivariate statistics. Results from meiotic behav- iour supports the phenetic grouping. We found a striking association between morphological pat- terns of the pollen grains, meiotic behaviour and habit morphology. Our results showed that Iranian populations of T. disperma represent the same chromosome, suggesting that the pollen size in differ- ent populations can be served to obtain useful information. The environment of origin seems to have an effect on the chromosomes within different populations. Key words Fabaceae, Meiosis, Morphology, Pollen, Trigonella disperma. The tribe Trifolieae in the family Fabaceae consists of 6 genera: Medicago L., Melilotus Mill., Ononis L., Parcochetus Buch.-Ham. ex D. Don, Trifolium L. and Trigonella L. (Heyn 1981, Lock and Simpson 1991, Mabberley 1997). This tribe as proposed by Berchtold and Presl (1820) is characterised by having trifoliate leaves and stipules that are adnate to the petioles. In the early 20th century, Schulz (1901) used seed characters to form a tribe called Trifolieae, uniting the genera, Parochetus, Ononis and Trifolium. Hutchinson (1964) separated the genus Ononis into the monogeneric tribe Ononideae and he expanded the tribe Trifolieae to include the genera Factorovskya, Medicago, Melilotus, Parochetus, Trifolium and Trigonella. This treat- ment of the genera has been adopted by subsequent authors (Davis 1970, Townsend and Guest 1974, Meikle 1977). Small et al. (1981, 1990) separated the genus Ononis and subdivided the other 6 genera into 2 subtribes: subtribe Trifoliinae consisting only of the genus Trifolium, and subtribe Trigonellinae including the rest of the genera. Progression of modern tools in taxonomy led to a flux of taxonomic opinion of these genera and tribes. Comprising of ca. 135 species, the genus Trigonella is one of the largest genera in the tribe Trifolieae and widely distributed in the dryer regions around the eastern Mediterranean, western Asia, southern Europe, northern and southern Africa, with only 1 species found in southern Australia (Townsend and Guest 1974, Polhill 1981, Kawashty 1998). Trigonella consists of perennial or annual herbs with pinnately trifoliate leaves, often exhaling an odour and are, like other grain legumes, important for food and medicine (Chopra et al. 1956, * Corresponding author, e-mail: [email protected] 280 M. Ranjbar et al. Cytologia 76(3) Girardon et al. 1989, Balodi and Rao 1991, Bhatti et al. 1996, Dangi et al. 2004). In Flora Iranica (Rechinger 1984) the genus is represented by 58 annual and perennial species in 12 sections. Little is known about the nature of genetic variability in diploid species of Trigonella and the taxonomic relationships between different taxa (Singh and Roy 1970). An important part of peren- nial members (nearly 7) of the genus endemic to Iran belong to T. sect. Ellipticae. This section is characterised by having glabrous leaves, stems and pods, and yellow or occasionally violet flowers. The taxonomy of the T. sect. Ellipticae continues to be subject of much confusion, mainly because of different approaches to species delimitation, resulting in various numbers of recognised spe- cies (Boissier 1872, Hedge 1970). Hence, investigations in different aspects such as morphology, chromosome number, meiotic behaviour and pollen morphology may be useful to solve taxonomic problems within this section. Multivariate statistical methods provide an objective analysis of patterns of morphological variation within a taxon and are a suitable tool for systematists in deciding on taxon delimita- tion. Some recent studies employing this method are e.g. Baum and Bailey (1992), Tyteca and Dufrene (1994), Matos (1995), Eakes and Lammers (1996), Eriksen (1997), Aldasoro et al. (1998), Sepp and Paal (1998), Shaw (1998), Verboom and Linder (1998), Brunell and Whitkus (1999), Eddie and Ingrouille (1999), Wilkin (1999), and Cupido (2003). Cytogenetic investigations were conducted on Trigonella by Singh and Roy (1970), Singh and Singh (1976), Agarwal and Gupta (1983), Ahmad et al. (1999) and Dundas et al. (2006). The mitotic chromosome number of an accession from Nisyros (Greece) of Trigonella balansae Boiss. & Reuter, an annual pasture legume of Eurasian origin, was first reported by Kamari and Papatsou (1973). Studies on the impact of karyotypic data on the interspecific and phylogenetic relationships and also on meiotic behaviour in the genus are still limited. Study on the pollen grains of legumi- nous plants (Clarke and Kupicha 1976, Ferguson and Skvarla 1981, Ferguson 1990, Ferguson and Stirton 1993, Diez and Ferguson 1994, Hughes 1997, among others), dealt mainly with the descrip- tion of the pollen grains of certain genera or tribes. This article follows previous studies conducted on fodder legumes in Iran (Ranjbar et al. 2004. 2006, 2009a, 2009b, 2010a, 2011a, 2011b, 2011c) and aims at increasing the knowledge about pat- terns of morphological variation, chromosome numbers, meiotic behaviour and pollen morphology in different populations of Trigonella disperma Bornm. (1910: 326) of section Ellipticae in Iran. It is an endemic and also a widespread perennial species of the genus in the western Iran. As a mem- ber of Leguminosae, it improves the nitrogen content of the soil by fixing the atmospheric nitrogen. The species is an aromatic plant; often emitting an odour consists of hydrocarbons and oxygenated sesquiterpenes as the most abundant components (Ranjbar et al. 2009b). In our previous study, ana- tomical characters related to epidermis cells of the leaves and peduncles were used to assess intra- specific relationships within T. disperma and their taxonomic significance was discussed (Ranjbar et al. 2010a). The present study establishes relationships between chromosomal criteria, pollen morphology and taxonomic delimitation. Further investigations conducted on native populations of T. disperma coming from other parts of Iran would help us to verify the emitted hypothesis for better understanding of the mechanism of evolution in these populations. Materials and methods Morphology Plants were collected from different regions within the natural geographical distribution of T. disperma during several excursions in Iran (Table 1). The collected materials were in the vegeta- tive or fruiting phase and deposited at BASU, Hamedan, Iran. Several herbarium sheets have been examined for each population from the following herbaria: FUMH, PR, TARI, W, WU, Herbarium of Isfahan University, Herbarium Research Centre of Natural Resource and Animal Affairs of 2011 Cytomorphological Study of Trigonella disperma (Fabaceae) in Iran 281 Table 1. Collection data of 8 populations of T. disperma Taxa Locality Altitude (m) Voucher specimen Abbreviation T. disperma Kurdestan: Serishabad 1750 m 14482 (BASU) DIS 82 T. disperma Kurdestan: The first village after Serishabad 1750 m 14483 (BASU) DIS 83 T. disperma Hamedan: Mahnian to Avaj 2062 m 14484 (BASU) DIS 84 T. disperma Kurdestan: Hamekasi 2035 m 14485 (BASU) DIS 85 T. disperma Hamedan: Avaj to Abgarm, 15 km to Abgarm 1732 m 14486 (BASU) DIS 86 T. disperma Hamedan: Kaboudrahang, 5 km after Kantapeh village 2200 m 14479 (BASU) DIS 79 T. disperma Zanjan: Abhar to Gheydar, before Kinevers village 1700 m 14480 (BASU) DIS 80 T. disperma Hamedan: 16 km before Avaj 1900 m 14481 (BASU) DIS 81 Table 2. Mean values of morphological characters and character state matrixes of 8 populations of T. disperma (abbreviations are as listed in Table 1) DIS DIS DIS DIS DIS DIS 7 DIS DIS Morphological characters 82 83 84 85 86 9 80 81 Stem length (cm) 34 31.5 36 24.5 34.5 30 27 30 Internodes length (cm) 1.75 2.6 2 2 1.75 3.25 2.5 2.4 Leaf length (mm) 13.5 9 12 8 8 9 13 11.5 Leaflet length (mm) 6.5 8.5 7.5 6.5 7 8.5 9.5 8.5 Leaflet width (mm) 4.5 5.5 6 5 5 5.75 6 15 Petiole length (mm) 8.5 6.5 10 6.5 5.5 7 10.5 10 Stipule length (mm) 3.5 4 4 3.5 2.5 5 4.5 4 Stipule width (mm) 0.65 0.75 0.75 0.75 1.25 0.75 0.75 0.75 Peduncle length (cm) 3.25 4 2.1 3.25 1.5 2.9 2.75 2.85 Calyx tube length (mm) 3 3.5 3 3 2.5 3 3 3 Calyx tooth length (mm) 1 1.5 2 1 1.5 1 1 2 Calyx length (mm) 4 5 5 4 4 4 4 5 Standard length (mm) 7 9 10 9 8 10 8 10 Standard width (mm) 6 7 7 8 6 9 6 7 Wing length (mm) 8 9 10 9 8 10 8 9 Wing width (mm) 2.25 3 3 2.5 2 2.75 2 2.5 Wing claw length (mm) 3 3 3 3 3 2.5 3 3 Keel length (mm) 7 8 9 8 7 9 8 9 Keel width (mm) 3 3 3 3 3 3.5 3 3 Number of flowers 10 9 10 9 9 8 6 8 Pedicel length (mm) 2.5 3 4 2.5 3.5 3 2.5 2.5 Ovary stipa 1 1.5 1 0.75 1 1.25 1 1 Hair density on leaflet 1 1 1 1 1 0 1 1 (loose=1, sparse=0) Hair density on calyx 0 1 0 1 1 0 1 1 (sparse-loose=1, sparse-glabrous=0) Hair density on stipule 0 1 1 1 1 0 1 1 (loose=1, sparse=0) Hair density on bract 2 2 0 1 1 2 2 2 (loose=1, glabrous=0, sparse-glabre=0) Standard colour 1 1 1 0 1 2 0 1 (yellow=0, mild yellow=1, dark yellow=2) Standard shape 0 0 0 0 0 1 0 2 (orbit fan-shaped=0, obovatum=1, emarginate=2) Tabriz, Mashhad, Isfahan, Shiraz, Kerman and Zahedan.
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