Chemotaxonomy of Wild Diploid Triticum L. (Poaceae) Species in Iran
Mohammad Reza Rahiminejad
This study evaluates the taxonomic status and the variation
of wild diploid Triticum L. species using leaf flavonoid compounds
in Iran. Two-dimensional maps of these species were provided using polyamide
Thin Layer Chromatography. In order to show the taxonomic position of
these species, the cluster analysis based on Euclidean Distance Coefficient
and Ward Method and Factor analysis were applied (Principle Component
Analysis). The flavnoid compounds purification of each accession and species
was done. The results of cluster analysis showed that T. boeoticum
subsp. boeoticum Boiss. and T. boeoticum subsp. thaoudar
Reut. ex Boiss. are completely distinguished from each other. T. boeoticum
subsp. thaoudar shows more similarity to T. urartu Tum.
ex Gand., but T. boeoticum subsp. boeoticum displays affinity
to T. monococcum L. The results of this study showed variability
of flavnoid compounds at the diploid levels in Triticum species.
Also flavnoid profiles can be mentioned as well marker to show the taxonomic
position of these species.
Triticum L. genus (Poaceae; Triticeae) has four wild diploid species
(2n = 2x = 14; x = 7) growing in Iran (Bor, 1970, Rahiminejad and Kharazian,
2005). These species are as follow; T. boeoticum subsp. boeoticum
Boiss., T. boeoticum subsp. thaoudar Reut. ex Boiss.,
T. monococcum L. and T. urartu Tum. ex Gand. (Bor, 1968,
There have always been debates concerning the taxonomy of the genus
Triticum among the researchers. The complexes occurred among the species
were due to the high similarity and the hybridization between the Triticum
species which causes a high morphological similarity among these species
(Lange and Jochemsen, 1992). These evidences have obscured the morphological
limits of the species and caused taxonomic confusions in the number of
species and nomenclatural debates in the genus Aegilops (Morrison,
1993). Also, high levels of variations have been observed in this genus
(Waines and Barnhart, 1992).
Taxonomically, Dorofeev et al. (1979) reported that T. urartu
is in Section Urartu Dorof. and Filatenko, while T. monococcum
and T. boeoticum are grouped in Subgenus Boeoticum Migush.
and Dorof., Section Monococcon Dum. Furthermore, Van Slageren (1994) showed that Section Monococca Flaksb. includes T. monococcum and
Morphologically, T. urartu is very similar to T. boeoticum
(Yamagishi and Tanaka, 1978), while Kharazian et al. (2003) unfolded
that T. urartu is different from the two other diploid species.
Filatenko et al. (2001) reported that T. boeoticum, T.
monococcum and T. urartu are considered as separate species.
Controversially, Dhaliwal and Johnson (1976) showed T. monococcum
is similar to T. boeoticum. In addition, these species have
similar genome; AA, 2n = 2x = 14; x = 7 (Waines, 1995).
Chemotaxonomy is one of the important methods that display taxonomic
position of taxa (Crowford, 1990). It is now possible to study phenolic
profiles of high and low taxonomic levels, even of individual genotypes
(Mika et al., 2005). The use of the distribution patterns of natural
plant products-alkaloids, terpenes, phenolics, glucosinolites, terpenoids
and carbohydrates is well-established as a major tool for investigating
population structures, species, taxonomical problems and phyletic relationships
of genera. Taxonomically, the most important phenolics are the flavnoids,
which have a relatively common nucleus with great variety of types and
patterns of side-groups that characterize the individual compounds. There
is usually a considerable diversity of flavnoids in species (Nakipoklu,
2002). Also, extensive chemotaxonomic flavnoid research on the Poaceae
detected the presence of flavone C-glycosides and tricin 5-glucoside (Bouaziz
et al., 2001). Using thin layer chromatographic patterns, Dedio
et al. (1969) presented the original taxonomic relationships in
the genus Secale L. Thin-layer chromatography in these genus supported the
general taxonomic relationships based on the morphological and cytological
studies. The differences in chromatographic patterns were sufficient to
determine the position of the Section of Secale genus. In addition,
Bouaziz et al. (2001) showed the flavnoids of Hyparrhenia hirta
Stapf (Poaceae) which grows in Tunisia. The most important compounds
are 7-o-glucoside and vitexin. Also, Frey (1996) showed different phenolic
profiles among Trisetum Pers.
Moreover, since Iran is one of the secondary centers of genetic diversity
for diploid Triticum species, there is a need of using this genetic
resource effectively in wheat improvement in this country. So far, the
survey of flavnoid compounds in Triticum genus in Iran has not
been studied, therefore the aims of this study are as follow:
||Detect the flavnoid compounds among the accessions
||Determine the taxonomic position via flavnoids
||Identify variability in accessions of wild diploid Triticum
species in Iran
MATERIALS AND METHODS
Extraction, isolation and identification of flavnoids were based on the
protocol of Markham (1982). Using leaf flavnoid chemistry from 14 accessions
(Table 1) of wild Triticum species, the taxonomic
status and the phytochemical variations of these species were studied.
Two-dimensional maps (2DM) of these species were provided using methanol
extracts on polyamide (MN-Polyamid-DC6) Thin Layer Chromatography (TLC).
Moreover, Spots` detection in NP identifiers (Diphenylboric acid 2-Aminoethyle
Ester) was performed under UV-254 nm and the presence/absence of spots
was taken as a character state and it was applied in each accession of
these species. In addition, Rf values (migration distance of
the bands/distance of solvent front) in each accession were studied (Apaydin
and Bilgener, 2000; Gulen and Eris, 2004). In order to show the taxonomic
position of these species the Cluster analysis based on Euclidean Distance
Coefficient, Ward Method as well as Factor analysis (Principle Component
Analysis) via SPSS. V. 14 were applied; besides, the descriptive analysis
was studied to estimate the Coefficient Variation (CV). The flavnoid compounds
purification of each accession and species was carried out through a sephadex
(LH20) column and one-dimensional maps (1 DM) on polyamide TLC plates.
Identification of the purified compounds was performed based on their
UV spectra (200-500 nm) and shift reagent, such as AlCl3, AlCl3/HCl,
NaOAC, NaOAC/H3BO3 and Methanol (Markham, 1982).
||Locality of wild diploid Triticum species in
natural habitat of Iran
The two-dimensional flavnoid patterns of crude extracts of the taxa under
study displayed the variability for each accession. Total number of spots
were obtained in each species are presented as: (1) in T. boeoticum
subsp. boeoticum 6-7 spots, (2) in T. boeoticum
subsp. thaoudar 1-6 spots, (3) in T. monococcum 2-14 spots
and (4) in T. urartu 2-4 spots. Dark yellow spots were common in
T. urartu, T. boeoticum subsp. boeoticum and T.
monococcum, blue spots were in T. boeoticum subsp. boeoticum
and T. monococcum, orange spots were in T. boeoticum subsp.
thaoudar, boeoticum and T. monococcum but violet
spots were in T. urartu. Also, the presence and absence of flavnoid
spots were surveyed in each accession (Table 2, Fig.
1, 2). Based on the color of each profile, the kind
of flavnoid type is partially detected; T. urartu accessions have
3-OH flavonol, 5-OH flavonol or flavone, 4`-OH chalcone (Table
3, 4). In T. boeoticum subsp. boeoticum
accessions observed Hydroflavone, flavonol, 5-o-glycosides, 3-o-flavonol,
5-OH flavone, dihydroflavonol, biflavonyl, 2`-OH chalcone, 6`-OH chalcone.
T. boeoticum subsp. thaoudar have 2,4-OH chalcone. T.
monococcum have flavone, flavonol, 3-OH flavonol, 5-OH falvonol, 2`-OH
chalcone, 6`-OH chalcone.
Furthermore, Rf values of each accession were studied in aqueous
and organic solvent system. Both maximum and minimum Rf in
aqueous system were observed in T. boeoticum subsp. boeoticum
(1.76, 0.09, respectively), T. monococcum (1.72, 0.09, respectively).
Also, maximum and minimum Rf in organic system were in
T. monococcum (2.11, 0.14, respectively) (Table 5).
In aqueous solvent systems, most of the stains have Hydroxyl groups with
high polarity, but in organic solvent systems they have little polarity.
In order to determine the flavnoid compounds, the fractions of each accession
were surveyed. Absorption of UV spectrum via shift reagent such as AlCl3,
AlCl3/HCl, NaOAc and NaOAC/H3BO3 were
examined. Among T. urartu accessions, increased intensity (bathochromic
shift) of Band I displays Hydroxylation at 2` and 4-positions and variability
in B-ring o-Dihydroxylation. That is when Band II shows Oxygenation
at 6 or 8-position. In T. boeoticum subsp. boeoticum, Band
I shows Hydroxylation at 5 and 4-position and variability in B-ring o-Dihydroxylation
and A-ring o-Dihydroxylation while Band II shows Hydroxylation
at 7-position and Oxygenation at 6 or 8-position. In T. boeoticum
subsp. thaoudar increased intensity demonstrates Hydroxylation
at 7 and 2`-position and Oxygenation at 3`-position. Increased intensity
of Band I in T. monococcum accessions indicates the variability
of B-ring-o-Dihydroxylation and Hydroxylation at 4-position (Table
||Presence and absence of each spot in each accession
of Triticum accessions
|+: Presence of especial spot, -: Absence of especial
Similar flavnoid compounds between T. urartu and T. monococcum
are flavone, 4`-Hydroxychalcone, Pseudobaptisin, T. boeoticum
subsp. boeoticum and T. monococcum are flavone, 4`-Hydroxychalcone,
3`,4`,7` Trihydroxyflavone, between T. urartu and T. boeoticum
subsp. boeoticum are flavone, 4`-Hydroxychalcone, between T.
boeoticum subsp. thaoudar and boeoticum are 2-Hydroxychalcone,
4`-Hydroxychalcone, 2`-Hydroxy4`-Methoxychalcone, 2`,4- Dihydroxychalcone,
between T. monococcum and T. boeoticum subsp. thaoudar
are 2`-Hydroxychalcone, 2`-Hydroxy4`-Methoxychalcone, 4`-Hydroxychalcone
and between T. urartu and T. boeoticum subsp. thaoudar
is 4`-Hydroxychalcone. Therefore 4`-Hydroxychalcone one of the flavnoid
compounds which is common among these species (Table 3).
||Chromatogram of spots in Triticum species; b.b:
T. boeoticum subsp. boeoticum, b.th: T. boeoticum
||Chromatogram of spots in Triticum species; m:
T. monococcum, ur: T. urartu
||Flavnoid compounds among Triticum accessions
|+: Presence of especial flavnoid compound, -: Absence
of especial flavnoid compound
||Summary of the main flavnoid variation features among
diploid Triticum species
|+: Presence of flavnoid variation features, -: Absence
of flavnoid variation features
Cluster and Factor analysis among Triticum accessions
in aqueous and organic system; A and C in aqueous system, B and D
in organic system, b.b: T. boeoticum subsp. boeoticum,
b.th: T. boeoticum subsp. thaoudar, mn: T. monococcum
and ur: T. urartu
Relationships between these species based on the Rf data are
shown graphically in Fig. 1 and 2.
Maximum CV in aqueous solvent system is related to subsp. thaoudar
(75.3) and boeoticum (64.6) and in organic solvent system it is
in T. boeoticum subsp. thaoudar (64.2) and T. monococcum
(67.8), while minimum C.V. in both systems were in T. urartu (34.7
aqueous, 15.27 organic) (Table 6). The statistical analyses
such as Cluster analysis were investigated in both aqueous and organic
solvent systems. In aqueous system, two groups were comprised. (1)
T. monococcum, T. boeoticum subsp. boeoticum,
T. boeoticum subsp. thaoudar; T. boeoticum subsp.
thaoudar, T. urartu, T. monococcum;
T. urartu, T. boeoticum subsp. thaoudar, (2) T.
monococcum, T. boeoticum subsp. boeoticum. In most
of the clusters T. urartu and T. boeoticum subsp.
thaoudar grouped together, also T. monococcum and T.
boeoticum subsp. boeoticum were close together. Two subspecies
of T. boeoticum were discrete from each other and T.
monococcum clustered with two subspecies of boeoticum (Fig.
3). Using Factor analysis for aqueous system, it was observed that
two groups in two components were comprised; (1) T. urartu,
T. boeoticum subsp. thaoudar, T. monococcum (2) T.
monococcum, T. boeoticum subsp. boeoticum, T. boeoticum
subsp. thaoudar. In this analysis, two subspecies are exactly separated
(Fig. 3). The results of clustering in organic solvent
system proved three distinct groups, (1) T. urartu, T.
boeoticum subsp. thaoudar, T. monococcum, T. monococcum,
T. boeoticum subsp. boeoticum, T. urartu, T.
boeoticum subsp. thaoudar (2) T. monococcum,
T. boeoticum subsp. boeoticum. In this system, two subspecies
of T. boeoticum were distincted. Using Factor analysis for organic
data presented two comprised groups and therefore, affirmed the Factor
analysis performed for aqueous solvent system (Fig. 3).
Based on these results, aqueous solvent system is more appropriate for
determining taxonomic position than organic system.
Little information is available about the variation extension among the
wild progenitors of Triticum species (Hedge et al., 2000).
There are conflicting reports on the amount of diversity in diploid wheat
populations, specifically in chemotaxonomy. Based on the results of this
study, T. urartu accessions with B-ring o-Dihydroxylation,
Oxygenation at positions 6 and 8 and 4-Hydroxylation are common in
T. monococcum and T. boeoticum subsp. boeoticum
(Table 5), which is in accordance with Hammer et
al. (2000) using micro satellites. Van Slageren (1994) and Dubcovsky
and Dvorak (1995) reported that T. monococcum and T.
urartu were related in phylogenetic trees. Using flavnoid chemistry, it was
observed that T. monococcum and T. urartu were related,
noticeably T. urartu having 2`- Hydroxylation is similar
to T. boeoticum subsp. thaoudar (Table 5).
In this study two subspecies of T. boeoticum were distincted in
flavnoid variation, whereas these are exclusively similar in 7-Hydroxylation.
In cluster and Factor analysis T. urartu and T. boeoticum
subsp. thaoudar were approximately grouped together. Morphologically,
T. urartu is very similar to T. boeoticum (Yamagishi and
Tanaka, 1978). T. monococcum and T. boeoticum subsp. boeoticum
also grouped together and displayed affinity between them, which in accordance
with Dorofeev et al. (1979) and Jaaska (1993, 1997). In addition,
Dhaliwal and Johnson (1976) and Waines and Barnhart (1992) proved that
T. monococcum is similar to T. boeoticum. Furthermore,
using meiotic paring behavior and analysis of the polymorphisms of repeated
nucleotide sequences T. urartu was found to be as a donor of A
and B genome in polyploid wheat (Dhaliwal and Johnson, 1976, Dvorak et
al., 1993; Hammer et al., 2000). Controversially, Filatenko
et al. (2001) reported that T. boeoticum, T. monococcum
and T. urartu are considered as separate species. Also, Dorofeev
et al. (1979) and Kharazian et al. (2003) evolved that T.
urartu is different from the two other diploid species. Using isoenzyme
markers, Jaaska (1993) reported that T. boeoticum and T. urartu
were found to differ with regard to acid phosphatase, esterase and superoxide
dismutase, showing distinctly different gene pools.
||Rf values of Triticum accessions in
aquatic and organic phase
So far, the researches did not have any supporting evidence which could
clearly separate the wild diploid species (Morrison, 1993) and subspecies.
In this study two subspecies of T. boeoticum; thaoudar
and boeoticum were separated exactly (Fig. 3).
Using microsatellites, Hammer et al. (2000) reported that two subspecies
of T. boeoticum have different genomes. Based on flavnoid
compounds these four species seem to have genomic relationships because
of similarity in B-ring o-Dihydroxylation and 4-Hydroxylation,
flavone and 4`-Hydroxychalcone which can provide the gene flow among diploid
species. This similarity confuses the limits of diploid species. Noticeably,
T. monococcum accessions display oxidation patterns of flavones
which can be observed in Sciadopitysin, 3`,4`,7`-Trihydroxyflavone 7-o-rhamnoglucoside
and 3,3`,4`-Trihydroxyflavone. Both T. uraartu and T. monococcum
have Oxidation flavnoids patterns which are related to Pseudobaptisin
(Table 4). Therefore, these evidences show that T.
urartu can be mentioned as one of the diploid ancestors to other diploid
Triticum species. Nevertheless, these four wild diploid species
are different from each other (Dhaliwal, 1977).
Little information is available on the extent of genetic variation in
the wild Triticum species (Hedge et al., 2000, Moghaddam
et al., 2000). Hedge et al. (2000) reported that T. monococcum
and T. urartu accessions have polymorphic loci, but the low heterozygosity
is indirectly resulting in high hybridization among the accessions. The
flavnoid profiles among T. monococcum accessions present high variability
in their flavnoid patterns which confirm these results (Fig.
2). Noticeably, they reported that T. urartu has the highest
genetic diversity, which is not in accordance with present results (Table
6), T. urartu accessions have the least variability in their
profile (Fig. 2). Intraspecific alloenzymic variability
was found to be low in this species (Jaaska, 1993). Diversity and polymorphism
of flavnoid compounds in accessions of T. monococcum and
T. boeoticum subsp. thaoudar and boeoticum are more
than T. urartu (Table 5, 6
and Fig. 1, 2) which are in agreement
with Hedge et al. (2000) results, therefore maximum CV in aqueous
solvent system is related to subsp. thaoudar and boeoticum
and in organic solvent system it is in T. boeoticum subsp. thaoudar
and T. monococcum, while minimum CV in both systems were in T.
urartu (Table 6). Yaghoobi-Saray (1979) and Hedge
et al. (2000) reported higher levels of alloenzyme diversity in
T. monococcum and T. urartu. Genetic variability
which was approximately identical for diploid species may be brought about
by similar kinds of evolutionary forces for which all the species might
have had identical response. Phenolic profiles specifically demonstrate
genetic affinity and significant polymorphism among species (Mika et
al., 2005). These polymorphisms of flavnoid profiles have been showed
in Fig. 1 and 2.
||Variability of flavnoid compounds among Triticum
accessions in Iran
|*Standard Deviation, **Coefficient of Variation
The widespread occurrence of flavnoid compounds in Triticum genus
makes them useful markers for taxonomy and evolutionary relationships
(Mika et al., 2005). To use flavnoid compounds more widely as genetic
markers, these would have to be not only universal and abundant, but also
environmentally stable and convenient for identifying taxonomic position
(Fairbbothers et al., 1975). In addition, such studies should be
performed at the population level as well.
Authors are thankful to the University of Isfahan and Shahrekord to grant
the research project No. 28. This research was performed at Department
of Botany, Faculty of Sciences, University of Isfahan, Isfahan, Iran,
on October 2003.
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