Karyological Study on Bellevalia and Muscari (Liliaceae) Species of Iran
Ali Asghar Maassoumi
An investigation of karyotype and chromosome numbers
was carried out in different populations of Bellevalia Lapeyr.
and Muscari Mill. species from Iran. In this research, six species
of Bellevalia and six species of Muscari were studied. Different
levels of ploidy were found in them. In Bellevalia with x = 4,
levels of ploidy were diploid, autopentaploid and hexaploid and in Muscari
with x = 9, it was diploid, autotriploid, tetraploid and autopentaploid.
In the present research, for first time, the karyotype of B. olivieri
was prepared and a population of B. longistyla with autopenploid
was observed. The variation of B chromosome number in Bellevalia was
Bellevalia Lapeyr. and Muscari Mill. are genera of bulbous
plants belonging to Liliaceae, subfamily Lilioideae and tribe Scilleae
(Engler, 1887). The both genera have a wide spread distribution. They
are present in the whole Mediterranean basin as far as Caucasus, temperate
Europe, North of Africa, South west of Asia (Losinskaya, 1935; Feinbrun,
1938-1940; Parsa, 1950; Bentzer, 1972; Garbari, 1973; Davis, 1984; Townsend
and Guest, 1985; Feinbrun, 1986; Assadi, 1988; Rechinger, 1990; Wendelbo,
Bellevalia and Muscari are comprising 18 and nine species
in Iran, respectively (Parsa, 1950; Wendelbo, 1967, 1980; Assadi, 1988;
Rechinger, 1990). Taxonomically Bellevalia closely is related to
Muscari and Hyacinthella. However cytologically, the larger
chromosome of Bellevalia make it easily recognizable from the two
other ones. The basic chromosome number is x = 4 in Bellevalia
(Johnson, 2003) and x = 9 in Muscari (Speta, 1998).
Previous karyological studies on Bellevalia and Muscari
showed in Table 1. The length of chromosome of Bellevalia
and Muscari species were variable between (6-10, 18-20) and (3-8)
micron, respectively (Feinbrun, 1938-1940; Bentzer, 1972).
The aim of this research was to study the karyotype of the mentioned
species and to test if the relationship based on morphological characters
are in accordance with cytological data. So we prepared karyogram of somatic
number of chromosome from B.fominii, B.tabriziana, B.
glauca, B.longistyla, B.olivieri, M. comosum,
M. caucasicum, M. tenuiflorum, M.longipes, M.
armeniacum var. szovitzianum and M. neglectum.
||Previous Karyological studies on studied Bellevalia
||Taxa, localities of Bellevalia and Muscari
MATERIALS AND METHODS
The materials were collected from the east, center and west of Iran in
February until June 2003, 2004 (Table 2). Voucher specimen
are deposited in herbarium of Tehran sciences and researches campus For
karyotype analysis, a pretreatment at room temperature for three hours
was usually applied before fixation of the root tips of six species of
Bellevalia and Muscari either in 0.002 M 8-Hydroxyquinoline.
After fixation in a cold mixture of ethanol and acetic acid (3:1), the
following procedure involved the maceration in 1 N HCl at 60 for 5-8 min,
washing in water, cutting off the meristems and squashing them in a drop
of 45% acetic acid (Krahulcova, 2003). Chromosomes were described according
to Levans terminology (Levan et al., 1964). Karyotypes were compared
using total form percentage (Forni-Martin et al., 1994) and calculated
the ratio of the longest to the shortest chromosome (Verma, 1980). Symmetry
karyotypes were determined using Stebbins two way system (Stebbins, 1971).
The somatic chromosome number and details of karyotype of Bellevalia
and Muscari studied species were shown in Table
3 and 4. B. fominii from sect. Nutans
(Fig. 1a, 2a) and B. tabrizina
from sect. Patens (Fig. 1b, 2b)
were diploid (2X = 2n = 8) with karyotype formula
||Somatic chromosome no., karyotype formula and symetoy
of Bellevalia and Muscari
|NO = Not Observed M = Median point, m = Median region,
sm = Submedian, st = Subterminal t = Terminal region
||Karyotypic details of the species studies of Bellevalua
|TL = Total chromatin length, S = Shortest chromosome,
L = Longest chromosome, L/S = Longest/Shortest, TF % = Total from
percentage, TV = Total Volume
(2m+1sm+1st) and (1m+2sm+1st), respectively. The rest of species belonged
to sect. Bellevalia. B. glauca was diploid (2X = 2n = 8) with karyotype
formula (3m+1sm) and the satellite was observed above the short arm of
first pair chromosome (Fig. 1c, 2c).
B. longistyla was autopentaploid (5X = 2n = 20+1B) with sm B-chromosome
and karyotype formula (1m+5sm+4st+1B) (Fig. 1d, 2d).
B. saviczii was hexaploid (6X = 2n = 24+1B) with sm B-chromosome
and karyotype formula (7m+2sm+3st+1B) (Fig. 1e, 2e).
B. olivieri was hexaploid (6X = 2n = 24+5B) with five sm, st and
t B-chromosome and karyotype formula (7m+1sm+3st+1t+ 5B) (Fig.
Also M. comosoum (1) was diploid (2X = 2n = 18) with (1M+2m+2sm+4st)
(Fig. 1g, 2g.), but in M. comosum
(2) was diploid (2X = 2n = 17) with karyotype formula (2m+4sm+3st) (Fig.
1h, 2h). M. caucasicum (1), (2) were
||Somatic chromosomes of B. fominii
||Somatic chromosomes of B. tabriziana
||Somatic chromosomes of B. glauca The arrow showing satellite
|| Somatic chromosomes of B. longistyla. The arrow showing
||Somatic chromosomes of B. saviczii. The arrow showing
||Somatic chromosomes of B. olivieri. The arrows showing
||Somatic chromosomes of M. comosum (1)
||Somatic chromosomes of M. comosum (2)
||Somatic chromosomes of M. caucasicum (1)
||Somatic chromosomes of M. caucasicum (2)
||Somatic chromosomes of M. tenuiflorum (1)
||Somatic chromosomes of M. tenuiflorum (2)
||Somatic chromosomes of M. longipes
||Somatic chromosomes of M. neglectum
|| Somatic chromosomes of M. armeniacum var. szovitzianum
||Karyotype of B. fominii
||Karyotype of B. tabriziana
||Karyotype of B. glauca
||Karyotype of B. longistyla. The arrow showing B chromosome
||Karyotype of B. saviczii. The arrow showing B chromosome
||Karyotype of B. olivieri
||Karyotype of M. comosum (1)
||Karyotype of M. comosum (2)
||Karyotype of M. caucasicum (1)
||Karyotype of M. caucasicum (2)
||Karyotype of M. tenuiflorum (1)
||Karyotype of M. tenuiflorum (2)
||Karyotype of M. longipes
||Karyotype of M. neglectum
||Karyotype of M. armeniacum var. szovitzianum
(2X, 3X = 2n = 18, 27) with karyotype formula (1M+3m+4sm+1st) and (7m+4sm+2st),
respectively (Fig. 1i, j, 2i,
j). M. tenuiflorum (1), (2) were diploid and
autotriploid (2X, 3X = 2n = 18, 27) with karyotype formula (5m+3sm+1st)
and (1m+5sm+6st+1t) and some secondary constrictions (Fig.
1k, l, 2k, l).
M. longipes was diploid (2x = 2n = 18) with karyotype formula (2m+4sm+3st)
(Fig. 1m, 2m). M. neglectum
was autopentaploid (5X = 2n = 45) with karyotype formula (7m+12sm+3st)
(Fig. 1n, 2n) and M. armeniacum
var. szovitzianum was tetraploid (4x = 2n = 36) with karyotype
formula (2M+10m+4sm+2st) (Fig. 1o, 2o).
The basic chromosome number of Bellevalia is X = 4. This genus
often are diploid with 2n = 8, but there is a polyploidy series of 2n
= 16, 24, 32. Aneuploidy occurs only at the octaploid level (Ozhatay and
Johnson, 1996). They have metacentric, acrocentric and telocentric B-chromosome
B. fominii was falling to class 2C. This species had the longest
chromosome among the studied species which didn`t have any chromosome
polymorphism. Also some weak constriction were above the long arms. In
Bothmers report, karyotype formulae is the same as the present research.
Bothmer reported one chromosome in pair No. 4 have a longer short arm
than usual which indicate a pericentric inversion (Bothmer and Wendelbo,
1981). B. tabriziana had the shortest and thickest chromosome among
the studied species. This species was falling to class 3C. Its chromosomes
were large M and short st while Persson and Wendelbo reported its chromosome
have been large M, sm and st (Persson and Wendelbo, 1979). The ploidy
level of B. tabriziana in this research paper was similar to Perssons
report. B. glauca was falling to class 1C. The satellite was observed
above the short arm of first pair chromosome. The chromosome polymorphism
was occurred on first pair chromosome. The long arms of first pair were
not equal. It is related to pericentric inversion. This species had the
biggest and the thickest chromosomes among the studied sect. Bellevalia.
B. longistyla was falling to class 2C. The autopentaploid population
for this species is recorded for the first time. No obvious chromosome
polymorphism was observed. B. saviczii with one B chromosome
was falling to class 2C. Its chromosomes were smaller and thinner
than the other studied species of sect. Bellevalia in this research.
In present paper, allocyclic event and secondary constriction in long
arm were observed. Only one sm B-chromosome also, was found while in Bothmers
report its number was 1, 3, 7. Getner also reported 1-8 B-chromosome which
the most and the least of them were m and M, t (Getner, 2005). The
chromosome polymorphism was occurred in pairs No. 2, 3, 4. Getner also
confirmed pericentric inversion. He believed the acrocentric was changed
to metacentric. In its report aneuploidy, pentasomic (2n-1 = 23), heptasomic
(2n+1 = 25) and octasomic (2n+2 = 23) were found in B. saviczii which
collected from Shiraz (South of Iran) (Getner, 2005). Zakharieva reported
pentaploid population of B. saviczii (Zakhariyeva and Makushenko,
1969). B. olivieri was falling to class 2C. The detail of karyotype
is presented for first time. It has five B-chromosomes which were variable
as st, sm`, m. Asymmetric karyotype was bimodal. Pericentric inversion
and to minor extent translocations seems to be the main background for
the chromosome polymorphism (Bothmer and Wendelbo, 1981) while Feinbrun
believed allopolyploidy occurred in Bellevalia (Feinbrun, 1938-1940).
In regarding to Muscari, M. comosum (1), (2) were diploid
and falling to class 3C but first population showed decreasing aneuploidy.
Also one of the small chromosomes didn`t have homologous. One of the first
pair chromosome was longer than the other long st chromosome. Asymmetric
karyotype was trimodal. In Bentzers report, M. comosum was 2n
= 18, 27 (Bentzer, 1972). Both of the population of M. caucasicum
were diploid and autotriploid and falling to class 2C. Asymmetric karyotype
was trimodal. Both of the population of M. tenuiflorum
were diploid and autotriploid that falling to class 2C and 3C. Some constrictions
were recognized. They had trimodal asymmetric karyotype. In this specimen
allocyclic event existed.
||M. longipes was diploid and falling to class
||M. neglectum and M. armeniacum var. szovitzianum
were autopentaploid and tetraploid that falling to class 2C
Most of reports of polyploidy level in Bellevalia showed diploid
and tetraploid population. Few reports exists about triploid (Musano and
Maggini, 1976) and octaploid populations (Zakhariyeva and Makushenko,
1969; Bothmer and Wendelbo, 1981; Pogosjan and Torosyan, 1983). Variation
of ploidy and similarity of morphological characters have been found among
the species. In terms of morphological characters, Morphologically, Bellevalia
sect. Bellevalia is closer to Muscari subgen. Leopoldia.
Also subgen. Botryanthus is more advanced than subgen. Leopoldia.
The morphology of chromosomes confirmed taxonomic position. Most of studied
Bellevalia species were placed in class 2C of Stebbins system
except B. glauca and B. tabriziana.. It is representing
more primitive karyotypes.
The karyotypes of both genus are markedly asymmetrical because polymorphism
as confirmed by Garbari (1969, 1973), Bentzer and Ellmer (1975), Bentzer
and Landstorm (1975), Dalgic (1991), Corsi et al. (1996) and Bareka
and Kamari (2001).
Also, data regarding the total chromatin length and size of the longest
and shortest chromosomes were shown in Table 4. In
Bellevalia studied species, B. saviczii had the highest amount
of total chromatin length (236.12 μM) and B. tabrizana had
the least (32.24 μM). Also in Muscari, M. tenuiflorum
(1) had the highest amount of total chromatin length (189.44) and
M. longipes had the least (54.29). Also, M. comosum (2)
with 2n = 18 had 72.19 μM total chromatin length while M. caucasicum
(2) with 2n = 27 had 66.24 μM.
M. tenuiflorum (1) with 2n = 18 had 189.44 total chromatin length
and M. tenuiflorum (2) with 2n = 27 had 138.19. Here M. tenuiflorum
(1) with 2n = 18 had 189.44 total chromatin length, M. neglectum
with 2n = 36 had 180.26 and M. armeniacum var. szovitzianum
with 2n = 45 had 178.86. All of the above cases, are indicating that polyploidy
has been accompanied with chromatin loss. Mean while Stebbins class about
karyotype was shown in Table 3. Most of studied Bellevalia
species were placed in class 2c of Stebbins system except B. glauca
and B. tabriziana. It is representing more primitive karyotypes.
Most of studied Muscari species were placed in class 2c. Except
the both of M. comosum and M. tenuiflorum (2) populations
which were placed in 3C because they have trimodal asymmetric karyotype
with 3 sizes of chromosomes.
The results were obtained from the present study can be summarized as
||Polyploidy variation in Bellevalia was more than
Ploidy increasing didn`t have corresponding to total chromatin
||Chromosome morphology of Muscari subgen. Leopoldia
is larger and similar to Bellevalia. This subgenus is more
primitive than subgen. Botryanthus and closer to Bellevallia
||In B. longistyla and B. olivieri autopentaploidy and
hexaploidy were observed.
||The results of karyological study proved taxonomical position of
studied species of sect. Bellevalia.
||Presence of M chromosome in M. armeniacum var. szovitzianum
and M. caucasicum (1) probably due to pericentric inversion.
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