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Chromosome Number and Karyotype Analysis of some Taxa of Vicia Genus (Fabaceae): Revision and Description.

Byline: Safia El-Bok Aziza Zoghlami-Khelil Takwa Ben-Brahim Ali Ouji Hamadi Hassen Oussama Lamine Cheima Jabri Rim Douggari and Mohamed El-Gazzah

Abstract

Karyotypes of 15 accessions belonging to five species of Vicia were determined. Chromosomes number varied between species and subspecies. V. cordata had chromosome count of 2n=10 V. angustifolia had 2n=12 V. narbonensis V. monantha ssp. calcarata and ssp. cinerea presented 2n=14. Both V. sativa ssp. amphicarpa accessions with aerial and underground pods showed 2n=14 and were first reported. Chromosome numbers of V. sativa ssp. sativa were verified and revised as 2n=10 12. Therefore our karyological data provided information about phylogenetic position of the analysed species. Vicia narbonensis had the most symmetrical karyotype and could be considered as the primitive among the studied species. Vicia monantha had metacentric chromosomes with high differences in relative size between the chromosomes of the complement showing an asymmetrical karyotype. V. sativa ssp. sativa was the only subspecies having a metacentric chromosome marker in its complement within the species. Both V. cordata and V.

angustifolia had all subtelocentric chromosomes showing an evolutionary tendency. Principal component analysis and Cluster analysis revealed that the studied species were grouped regarding to their evolutionary tendency. Our results support that the karyotypes analysis showed distinction between the taxa. 2014 Friends Science Publishers

Keywords: Karyotype; Asymmetry index; Chromosome evolution; Vicia ssp

Introduction

The genus Vicia (Leguminosae Vicieae) has long been a subject of active research because it contains several species of economic importance and many of which have been domesticated since the origin of agriculture (Gil and Cubero

1993). It comprises about 166 species chiefly located in Europe Asia and North America extending to the temperate regions of South America and Tropical Africa (Bisht et al.

1998) out of which 40 species are of considerable economic importance (Hanelt and Mettin 1989). The seed is traditionally used as an additive to voluminous feeds for

ruminants (Enneking 1995) while the forage is often grown for hay in mixtures with cereal tutor crops such as barley and oats (Van de Wouw et al. 2001). Though Vicia sativa ssp.amphicarpa which produces two pod types: aerial and underground has an ability to survive in marginal areas with low rainfall (about 250 mm year-1) and to produce nutritious herbage and pods which help address rehabilitation of degraded rangelands and increase feed production for small ruminants (Abd El Moneim and Elias 2003).

The basic chromosome numbers in the genus are n = 5 6 and (Maxted et al. 1991) and the great majority of the species have complements with 2n = 10 12 and 14 respectively (Hanelt and Mettin 1989; Murti et al. 2012). However plants with these three basic chromosome numbers can exist even within one species as is the case for Vicia sativa (Hollings and Stace 1974). Nevertheless some representatives of the section Cracca are polyploidy (Yamamoto 1973). Cytological studies have tended to focus on the two most important economic species: V. faba L. and V. sativa (Maxted et al. 1991; Maxted 1993). The study of rare and new species of Vicia has encouraged the simultaneous examination of their cytology. To obtain a better point of view regarding this subject we will consider the chromosomal information which gives us the possibility of classifying the species (Ayaz and Ertekin 2008).

Karyotype characteristics played a vital role in improvement and comprehension of the phylogenetic relationships between the related species (Lavia et al. 2009; Murti et al. 2012). In this study the chromosomes reveal the full range of cytogenetical possibilities for understanding the delimitation affinities and evolution of taxa (Moore 1978). Karyotype asymmetry is a good expression of the general morphology of plant karyotypes (Zarco 1986; Zuo and Yuan 2011). Changes in morphology of the chromosomes have been frequently related to evolution in higher plants. A symmetrical karyotype is characterized by the predominance of m and sm chromosomes of approximately the same size (Zuo and Yuan 2011). In Tunisia research works on taxonomy of Vicia species and subspecies are still rare and not developed enough. However the karyotypes of 15 species and subspecies of Vicia (most of them are

Tunisian) are analysed in this paper. The objectives are: (i) to chromosome counts of various sub species (ii) to revise the taxonomy of some Vicia species and subspecies by the use of cytological analysis and (iii) to evaluate the karyotype evolution and phylogenetic relationships of the studied species and subspecies.

Materials and Methods

In this study we examined 15 accessions from 5 species (Vicia narbonensis V. sativa V. monantha V. angustifolia and V. cordata). V. sativa is represented by two subspecies V. sativa sativa and V. sativa amphicarpa. V. monantha is represented by two subspecies i.e. V. monantha calcarata and V. monantha cinerea. The list of the studied material and its origin is given in Table 1. Somatic chromosomes were studied in root meristems of germinating seeds which were pre-treated with 0.1% colchicines at room temperature for 2 h then fixed in ethanol chloroform acetic acid (6:3:1) during 24 h at 4C and stored in 70% ethanol. Root tips were hydrolysed with 1N HCL at 60C during 15 mn and stained according to the Feulgen technique. After that root tips were washed briefly with distilled water.

Meristematic regions with 1 mm of length excised and squashed in a drop of 1% acetic orcein mixed with a drop of 45% acetic water (Jahier 1992). The slides were examined under an optical Microscope type Hund (H 600) and photomicrographs were taken with the same microscope fitted with a BenQ camera using an oil immersion objective (100 x). At least 10 metaphases were drawn for each accession (including 310 individuals) selecting the five best for measurements. The well spread chromosomes were photographed and arm length were measured on prints enlarged to a total magnification of 2000 (100 A- 20). Each chromosome was identified on the basis of its total chromosome length. The nomenclature used for the description of the chromosome morphology is that proposed by Levan et al. (1964) where the abbreviations m sm and st designate metacentric submetacentric and subtelocentric chromosomes respectively. Idiograms were drawn based on mean centromeric index and arranged in order of decreasing size.

Data Analysis

For the numerical characterization of the karyotypes the following parameters were calculated according to Yamamoto (1973); Seijo and Fernandez (2003); Sevimay et al. (2005) and Gaffazardeh Namazi et al. (2008): (i) total

chromosome length of the haploid complement (TCL); (ii) mean chromosome length (MCL); (iii) mean centromeric index (MCI); (iv) intrachromosomal asymmetry index (A1)=1-[(b/B)/n] (Zarco 1986); and (v) interchromosomal asymmetry index (A2)= s/x where b and B are the mean length of short and long arms of each pair of homologues respectively n is the number of homologues s is the standard deviation and x the mean chromosome length (Zarco 1986; Seijo and Fernandez 2003; Zuo and Yuan 2011). Karyotype asymmetry has been determined using the A1 and A2 indices (Zarco 1986) the categories of Stebbins (1971) the asymmetry index (AI) according to following equation: AI=CVCL CVCI/100; where CVCL is the relative variation in chromosome length and CVCI is the relative variation in centromeric index (Paszko 2006).

Means were compared by using one-way analysis of variance (ANOVA) after Bartlett's test homogeneity. Post hoc Duncan test was used following ANOVA and was performed to compare the chromosomes pair's in each accession. A significant difference was considered when P=0.05.

Clustering of the karyotype was performed using the average linkage method to examine karyotype similarity among species and subspecies. A data matrix of 15 OTus (operational taxonomic unit) x 10 variables was constructed. The TCL MCL MCI A1 and A2 indices number of m sm and st chromosomes as well as the position of satellites were considered. Correlation coefficients (r) between A1 and A2 indices and between TCL and MCI were also measured. Additionally a multivariate analysis (PCA) was performed based on data matrix of 15 OTUs times the 5 mentioned quantitative variables (Seijo and Fernandez 2003).

Results

General Karyotype Characteristics

Of 15 accessions analysed in this paper three karyotypes were found: 2n=10 is present in three accessions of V. cordata and one of V. sativa ssp. sativa 2n = 12 is present in two accessions of V. sativa ssp. sativa and one of V. angustifolia and 2n = 14 is present in four accessions of V. narbonensis var narbonensis in two accessions of V. sativa ssp. amphicarap and in two accessions of V. monantha ssp. calcarata and ssp. cinerea. For all analysed species and subspecies satellites were observed in one chromosomes pair except for two species Vicia sativa ssp. sativa (acc. C3) and V sativa ssp. sativa var. Mghila (acc. T2) which have two chromosomes pairs having satellites (Table 2).

The karyotype formula among the studied species and subspecies of Vicia is 10st for V. cordata (acc. Sr acc. 304 and acc. C1) followed in frequency by 14m for both of V. monanatha subspecies (acc. 157 and acc. 140) 14sm for Libanon and Syrian V. narbonensis accessions (acc. 13 and acc. 12) 10sm + 4st for Tunisian V. narbonensis accessions

Table 1: Species subspecies code accessions and origin of material analysed cytologically

Species/sub species/code accession###Origin

V. narbonensis acc. 856###Tunisia (Bizerte)

V. narbonensis acc. 488###Tunisia (INRAT))

V. narbonensis acc. 13###Lebanon (ICARDA)

V. narbonensis acc. 12###Syria (ICARDA)

V. sativa ssp. sativa acc. C3###Tunisia (Tunis)

V. sativa ssp. sativa acc. 12###Bangladesh (ICARDA)

V. sativa ssp. sativa var. Mghila acc. T2###Tunisia (Siliana)

V. sativa ssp. amphicarpa upground seeds acc. 139###Tunisia (Thala)

V.sativa ssp. amphicarpa underground seeds acc. 139###Tunisia (Thala)

V .angustifolia acc. C2###Tunisia (Tunis)

V. cordata acc. 304###Tunisia (Ouslatia/Kairouan)

V. cordata acc. Sr###Tunisia (Seriya/Sejnane)

V. cordata acc. C1###Tunisia (Tunis)

V. monantha ssp. calcarata acc. 157###Tunisia (Makthar)

V. monantha ssp. cinerea acc. 140###Tunisia (Thala)

Table 2: Names of species code accessions karyotype formula (KF) chromosomes number (2N) satellite position

(SAT) total length of the haploid complement (TCL) Mean chromosome length (MCL) range of chromosome length

(range) mean centromeric index (MCI) A1 and A2: the intrachromosome and interchromosome asymmetry index

respectively (Romero zarco 1986); Stebbins' types: intervals of Stebbins (1971) for karyotype asymmetry of some Vicia

species and subspecies

Species/subspecies/var###Code###KF###2N SAT TCL SE###MCL###Range###MCI SE###A1###A2###Stebbins'

###accession###Types

1-V. narbonensis###acc. 856###10sm+4st###14 7l###42.77 0.191 6.11b###407 -827 33.64 0.72 0.49 0.18 2A

2-V. narbonensis###acc. 488###10sm+4st###14 6l###45.57 0.14###6.51b###5-833###33.39 0.67 0.49 0.13 2A

3-V. narbonensis###acc. 13###14sm###14 7l###47.39 0. 24 6.77a###425-10###55.59 2.38 0.43 0.22 2A

4-V. narbonensis###acc. 12###14sm###14 6l###49.77 0.22###7.11a###475-126 56.22 1.64 0.44 0.19 1A

###efg

5-V. monantha ssp. cinerea###acc. 140###14m###14 2s###26.62 0.23###3.80###2-8###4369 077 0.21 0.35 1A

6-V. monantha ssp. calcarata###acc. 157###14m###14 2s###25.27 0.19###3.61fg 214 - 69 43.67 0.81 0.18 0.34 1A

7-V. sativa ssp. sativa###acc. C3###2m+2sm +6st 10 3l+5l 22.9 0.24###4.58d###286-783 27.53 2.36 0.58 0.27 3A

8-V. sativa ssp. sativa###acc. 12###2m +10st###12 4l###21.9 0.19###3.65fg 171-5###30.23 3.85 0.69 0.27 3B

9-V sativa ssp. sativa###var Mghila 2m +10st###12 3l+4s 19.92 0.17###3.32g###171-6###22.54 1.47 0.69 0.28 3B

10-V. sativa ssp. amphicarpa (upgs.) acc. 139 14sm###14 1l###29.33 0.19###4.19de 243-8###30.45 1.23 0.58 0.27 3A

11-V. sativa ssp. amphicarpa (undgs.) acc. 139 6sm + 8st###14 2l###23.73 0.13###3.38g###186-525 25.79 1.37 0.65 0.23 4A

###c

12-V. cordata###acc. C1###10st###10 5l###25.3 0.18###5.06###383-675 16.31 0.73 0.8###0.18 4A

13-V. cordata###acc. 304 10st###10 5l###17.77 0.09###3.55fg 286 -43###19.67 1.12 0.72 0.14 4A

14-V .angustifolia###acc. C2###12st###12 5l###23.64 0.13###3.94f###246 -538 21.21 0.55 0.73 0.19 4A

15-V. cordata###acc. Sr###10st###10 5l###19.6 0.12###3.92ef 275 -5###20.25 0.75 0.75 0.15 4A

(acc. 856 and acc. 488) 2m +10st for both V. sativa ssp. sativa (acc. 12 and var. Mghila) 14sm for upground V. sativa amphicarpa seeds (acc. 139 upgs.) 4st + 3sm for underground V. sativa amphicarpa seeds (acc. 139 undgs.) 2m+2sm+6st for V. sativa ssp. sativa (acc. C3) and finally 12st for V. angustifolia (acc. C2). Fig. 1 illustrates the mitotic metaphasis Fig. 2 the respective karyotypes and Fig. 3 the respective idiograms. In general karyotypes of the analysed species and subspecies have a predominance of subteloncentric chromosomes (st).

Vicia narbonensis

The total chromosome length was considerably long; it ranged from 42.77 to 49.8 m. There were two types of karyotypes: one of ten submetacentric and four subtelocentric chromosomes (10sm+4st) for Tunisian

accessions and one of 14 submetacentric chromosomes (14sm) for Libanon and Syrian accessions that have MCI around 50. All accessions posses' satellites (SAT chromosome) a secondary constriction connected to the long arm near centromeric region (Table 2; Fig. 2 3).

Vicia monantha

The somatic chromosome number was 14 and the chromosomes were median (14 m) for both subspecies which have MCI about 43.7 (Table 2). All the chromosomes have approximately the same length except the first pair which has the longest chromosomes (Fig. 3).

Vicia sativa

V. sativa ssp. sativa (2n=10 and 2n= 12): The somatic chromosome number was 2n=2x=10 for acc. C3 and 2n=2x=12 for both acc. 12 and var. Mghila (Fig. 1). For accessions with 2n=2x=12 chromosomes there is one marker metacentric chromosome pair and six subtelocentric chromosomes. For the acc. 12 the secondary constriction is connected to the long arm (chromosome 4) while for the var. Mghila there are two secondary constrictions one connected to the long arm (chromosome 3) and one connected to the short arm (chromosome 4). For the acc. C3 with 2n=2x=10 the karyotype is compound of one marker

V. cordata

The somatic chromosome number was 10 for all the accessions and only one type of karyotype formula was observed (10st). The total chromosome length was considered short; it ranged from 17.77 to 25.3 m. However the accession 304 has the shortest chromosome length (Table 2). The satellite (SAT chromosome) was connected to the long arm of the shortest chromosome of observed pairs.

V. angustifolia

The somatic chromosome number was 12 for the acc. C2.

The total chromosome length was relatively short (23.64 m). All the chromosomes were subtelocentric (Fig. 2 and Fig. 3) with a satellite connected to the long arm of the fifth chromosome pair.

Karyotype asymmetry

All the indices proposed to evaluate intra-chromosomal asymmetry of the species and subspecies of Vicia karyotypes (Table 3). Among these methods one qualitative classification and five different quantitative indices can be market out. According to Stebbins category species having

2A and 3A such as Vicia narbonensis acc. 856 acc 488 and acc. 13 as well as V. sativa ssp sativa acc. C3 and V. sativa ssp. amphicarpa acc. 139 upgds. have karyotypes moderately symmetrical. While V. narbonensis acc. 12 falls in Stebbins 1A category and has a moderately symmetrical karyotype. The most symmetrical karyotype was observed in both V. monantha ssp calcarata and ssp cinerea which fall in Stebbins 1A category while the most asymmetrical

Relationships between the total lengths of the haploid complement (TCL) and the mean centromeric index (MCI). Values of TCL and MCI are summarised in Table 2. 1.V. narbonensis var narbonensis (acc.856); 2. V. narbonensis var narbonensis (acc. 488). 3. V. narbonensis var narbonensis (acc. 13). 4. V. narbonensis var narbonensis (acc. 12). 5. V. monantha cinerea (acc 140). 6. V. monantha calcarata (acc.

157).7. V. sativa ssp. sativa (C3) 8. V sativa ssp. sativa (acc. 12). 9. V. sativa ssp. sativa (var. Mghila). 10.V. sativa ssp. amphicarpa (acc. 139 upgs.). 11. V. sativa ssp. amphicarpa (acc. 139 undgs.). 12. V. cordata (acc. C1). 13. V. cordata (acc. 304). 14. V. angustifolia (acc. C2). 15. V. cordata (acc. Sr)

Scatter diagram of the Romero Zarco (1986) asymmetry indices. Values of A1 and A2 are summarised in Table 2. 1. V. narbonensis var narbonensis (acc. 856); 2. V. narbonensis var narbonensis (acc. 488). 3. V. narbonensis var narbonensis (acc. 13). 4. V. narbonensis var narbonensis (acc.

12). 5. V. monantha cinerea (acc. 140). 6. V. monantha calcarata (acc. 157).7. V. sativa ssp. sativa (C3) 8. V sativa ssp. sativa (acc. 12). 9. V. sativa ssp. sativa (var. Mghila).

10.V. sativa ssp. amphicarpa (acc. 139 upgs.). 11. V. sativa ssp. amphicarpa (acc. 139 undgs.). 12. V. cordata (acc. C1). 13. V. cordata (acc. 304). 14. V. angustifolia (acc. C2). 15. V. cordata (acc. Sr)

karyotype was observed in acc. 12 and var. Mghila of V. sativa ssp sativa (3B) as well as the other accessions with Stebbins 4A category of asymmetry (Table 3).

ANOVA showed significant differences between the studied species and subspecies concerning MCL and MCI (Pless than 0.0001).The highest values of MCL are observed in V. narbonensis accessions while the lowest MLC values are obtained in V. sativa ssp. sativa (var. Mghila) and V. sativa ssp. amphicarpa (acc. undgs.). The remaining accessions have intermediate values (Table 2). The two accessions of V. monantha (acc. 140 and acc. 157) have a high MCI but a medium TCL. The remaining accessions possess the intermediate TCL and MCI (Fig. 4).

The scatter diagram of A1 and A2 asymmetry indices

Dendrogram showing the phonetic relationships among the studies species and sub species of Vicia constructed using the average linkage method. 1. V. narbonensis var narbonensis (acc.856); 2. V. narbonensis var narbonensis (acc. 488). 3. V. narbonensis var narbonensis (acc. 13). 4. V. narbonensis var narbonensis (acc. 12). 5. V. monantha cinerea (acc. 140). 6. V. monantha calcarata (acc. 157).7. V. sativa ssp. sativa (C3) 8. V sativa ssp. sativa (acc. 12). 9. V. sativa ssp. sativa (var. Mghila). 10.V. sativa ssp. amphicarpa (acc. 139 upgs.). 11. V. sativa ssp. amphicarpa (acc. 139 undgs.). 12. V. cordata (acc. C1). 13. V. cordata (acc. 304).

14. V. angustifolia (acc. C2). 15. V. cordata (acc. Sr) presents three groups of species: (1) the two accessions of V. monantha (acc. 140 and acc. 157) which have the highest inter-chromosomal asymmetry index (average of A2=0.345); (2): the four accessions of V. narbonensis which have intermediate values of A1 and A2; (3): the three accessions of V. cordata with the highest values of A1 and (4): the two accessions of V. sativa ssp. sativa (acc C3 acc. 12 and var. Mghila) the two accessions of V. sativa ssp. amphicarpa (acc. 139 upgs. and acc. 139 undgs.) and V. angustifolia (acc. C2) which have intermediate A2 but high values of A1.

Cluster analysis constructed on the basis of karyotype similarities (Fig. 6) using the average linkage method shows three major groups. The first group includes the four accessions of V. narbonensis the second is composed by the two accessions of V. monantha and the third gathers both accessions of V. sativa ssp. sativa and V. sativa ssp. amphicarpa the three accessions of V. cordata and V. angustifolia which is closed to V. cordata.

The principal component analysis (PCA) of the karyotypical traits shows that the first two components account for 93.91% of the total variation (Fig. 7). The first component (54.14%) is positively correlated to TCL MCL and MCI and negatively correlated to A1 while the second component (35.76%) is defined positively by A1 and negatively by A2. The arrangement of the species and subspecies obtained by this analysis is similar to that obtained by the cluster analysis.

Discussion

Our results showed a wide range of chromosome numbers and karyotype morphology in Vicia species. Chromosome count and their morphological

features have been frequently recorded for cytological characterization of germplasm (Sharma and Sharma 2013; Weiss-Schneeweiss and Schneeweiss 2013). Studies revealed that all analysed accessions of Vicia were diploid. The chromosome number of V. narbonensis accessions was 2n=14 (Cremonini et al.

1998; Kamel 1999; Venora et al. 2009). Furthermore we have found similar chromosome count in the two V. monantha accessions and the two V. sativa ssp. amphicarpa accessions. In V. sativa ssp. sativa the chromosome number was 2n=10 for acc. C3 and 2n=12 for respectively acc. 12 and var. Mghila. V. cordata had chromosome number of

2n=10 for all the accessions and finally V. angustifolia had chromosome count of 2n=12. These results were in agreement with Kamel (1999); Weber and Shifino-Wittman (1999); Yamamoto (1973) and Javier et al. (1998).

Karyotype formula and quantitative analysis were variable among the studied species and subspecies except those that correspond to the different accessions of V. monantha and V. cordata. The chromosomes observed in the study were mainly subtelocentric or submetacentric types as mentioned by Jalilian and Rahiminejad (2012). The presence of two different formulas in the four V. narbonensis accessions may be due to the differences in their geographic origins (Bakatoushi and Ashour 2009).

Primitive wild species had symmetrical chromosomes in their karyotypes (Stebbins 1971). Therefore the predominance of sm chromosomes in our V. narbonensis accessions of approximately similar size and the presence of only one pair with a secondary constriction revealed that these accessions might have retained some of their primitive wild traits as suggested by Zuo and Yuan (2011). In V. sativa ssp. sativa two different formulas were found in three Tunisian accessions with a metacentric chromosome marker in their chromosome complements. This chromosome was noticeably absent from most of the other Vicia sativa aggregate (Maxted et al. 1991; Kamel 1999; Weber and Shifino-Wittman 1999). Both acc. 12 and var. Mghila have 2m+10st while acc. C3 had 2m+2sm+6st. These differences in karyotype formula may be due to their different geographic origin and/or to the difference in their status. We also noticed the presence of two secondary constrictions in both the var. Mghila and the acc. C3. The predominance of st

chromosomes revealed that these accessions transformed from their primitive wild form. In V. sativa ssp. amphicarpa two different formulas were found within same accession. Aerial seed had 14 sm while underground seed had 6sm+8st. Ladizinsky and Temkin (1978) and Javier et al. (1998) also noticed similar change in the morphology of chromosome due pod disposition and environment could also impact on the morphology of chromosome. All the chromosomes in above aerial seed

were sub median. Thus karyotype of aerial seed was considered symmetrical while the underground seeds have a predominance of subtelocentric chromosomes showing an asymmetry tendency. In V. monantha and V. cordata there was only one formula for each 14m and 10st respectively. According to Paszko (2006) the former has a symmetrical karyotype while the latest one has a karyotype in which asymmetry was increased due to the centromere position. Karyotype formula of the acc. C2 of V. angustifolia was

12st which was close to those of V. cordata. Its karyotype was considered as asymmetrical due to predominance of st chromosomes (Ruffini Castiglione et al. 2012). Generally plant material showed two types of karyotype symmetrical and asymmetrical. The symmetrical karyotype was represented by V. narbonensis and V. monantha with a predominance of sm and m chromosomes while the symmetrical karyotype includes V. sativa V. cordata and

V. angustifolia with a predominance of st chromosomes.

On the basis of our results we revised the taxonomy of few accessions. The taxonomy of some species i.e. V. cordata and V. angustifolia was revised due to centromere position of the chromosomes these species were considered previously as V. sativa. The acc. C1 and the acc. Sr were classified as V. cordata due to the presence of st chromosomes in all the complement and the presence of a secondary constriction in the shortest chromosome. C2 was also classified in V. angustifolia on the basis of st chromosomes in all the complement (2n=12) with a satellite connected to the long arm of the fifth chromosome pair as reported by Weber and Shifino-Wittman (1999). Several studies showed that V. sativa was taxonomically complicated due to low morphological distinction between subspecies but little effort was done to describe relationships between each subspecies.

The species were grouped regarding to their evolutionary tendency. The comparison of the chromosomes and mechanisms of karytoype evolution helps to understand the process of diversification within various taxa (Sharma and Sharma 2013; Weiss-Schneeweiss and Schneeweiss 2013). V. narbonensis. species (2n=14) was considered the most primitive which further gave rise to

2n=12 and 10 through chromosomes rearrangement. The chromosome number of 2n=14 observed in V. monantha and V. sativa ssp. amphicarpa and the centromere position of median/submedian argues the primitive tendency of these two taxa. Nevertheless differences in relative size between the chromosomes of the complement showed asymmetrical karyotypes and suggest that this structure led to the diversification of these taxa.

V. sativa ssp. sativa V. angustifolia and V. cordata had lower chromosome number of 2n=10 and 2n=12 may have arisen through Robertsonian translocation between two chromosomes in the complement (Maxted et al. (1991). These species also showed low TCL values a predominance of st chromosomes and differences in relative size between the chromosomes of the complements suggesting asymmetry indices for the establishment of the evolutionary changes in Vicia genus. This group of species could be considered as the most evolved in our study. Based on Stebbins' system the studied Vicia species were placed in five classes: 1A 2A

3A 4A and 3B whose classes 1A and 2A were considered as primitives (Hejazi et al. 2010). During the speciation and divergence of Vicia genus cycles toward symmetry and asymmetry may have occurred as reported for different groups (Jones 1970; Stebbins 1971). The differences in the asymmetry of the karyotypes were great for which it may be assumed that diversity of the genus has been accompanied by very small changes in the structure of the chromosomes. This study can be a model for genetic improvement programs and diversity studies for others Vicia species.

Acknowledgments

Many people have made significant contributions to this work and they all deserve our acknowledgement and thanks for their contributions. First we wish to thank the Director of the Institute of Agronomic Research of Tunisia (INRAT) for his support. We also wish to thank all the members of the Laboratory of Animal and Forage Production for their thoughtful suggestions and help. The authors are also grateful to Dr Saeed Rauf for his help and his most pertinent advice for reading this manuscript.

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Author:El-Gazzah, Mohamed; El-Bok, Safia; Zoghlami-Khelil, Aziza; Ben-Brahim, Takwa; Ouji, Ali; Hassen, Ham
Publication:International Journal of Agriculture and Biology
Article Type:Report
Geographic Code:6TUNI
Date:Dec 31, 2014
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