Contribution to cytology of genus Salvia L. (Lamiaceae) in Iran

 
CARYOLOGIA                                                                                 Vol. 63, no. 4: 405-410, 2010

Contribution to cytology of genus Salvia L. (Lamiaceae) in Iran
Sheidai Masoud1,*, Behnaz Alijanpoo2 and Masoud Khayyami2
1
Faculty of Biological Sciences, Shahid Beheshti University, GC, Tehran, Iran
2
Biology Department, Urmia University, Urmia, Iran

Abstract — The genus Salvia L. (Lamiaceae) contains about 900 species distributed throughout the Old and
New world growing in temperate and subtropical areas, with about 70 species reported in Flora Iranica. Cyto-
logical studies of the Salvia have been mainly focused on chromosome number reports and karyotype analysis,
while study of the chromosomes behavior in meiosis is very limited. Meiotic studies were performed in ten Salvia
species of S. spinosa, S. reuterana, S. sclarea, S. ceratophylla, S. xanthocheiala, S. limbata, S. hypoleuca, S. staminea,
S. nemorosa and S. verticillata showing 2n = 14, 20, 22 and 32 chromosome numbers indicating the role played
by polyploidy and aneuploidy in Salvia species diversification. Among Salvia species, the highest value of rela-
tive total, terminal and intercalary chiasmata occurred in S. verticellata while the lowest value of relative total
and terminal chiasmata occurred in S. spinosa. Laggard chromosomes and chromosomes stickiness as well as
frequent tripolar and multipolar cell formation due to anaphase I and II failure were observed. Potential unre-
duced (2n pollen) pollen grains were formed due to meiotic irregularities. B-chromosomes of 0-2 were observed
in the species of S. sclarea, S. nemorosa and S. reuterana.

Key words: Chromosome pairing, Salvia, unreduced pollen grains.

                INTRODUCTION                                   ment of cancer. Many species of the Lamiaceae
                                                               are aromatic and are often used as herbs, spices,
    The genus Salvia L. (Lamiaceae) contains                   folk medicines and fragrances. In addition, Salvia
about 900 species distributed throughout the                   species are grown in parks and gardens as orna-
Old and New world growing in temperate and                     mental plants (ÖZDEMIR and SENEL 1999).
subtropical areas (STANDLEY and WILLIAMS 1973;                     There have been about 70 Salvia species re-
ÖZDEMIR and SENEL 1999). Western Asia and                      ported from Flora Iranica with 40% endemism
Mediterranean regions have been considered                     (RECHINGER et al. 1982). These species show im-
as the original centers of distribution for Salvia             portant center of diversity in Flora Iranica regions
(WU and LI 1982).                                              with extensive morphological variation. Some
    Salvia species are herbaceous, suffruticose or             species are very distinct while others show close
shrabby perennials, rarely biennial or annual,                 affinity with others and some species are in the
often strongly aromatic. These species are of                  sate of evolutionary flux (RECHINGER 1982). Inter-
horticultural, commercial and medicinal values                 specific hybridization is suspected to be operative
(BHATTACHARYA 1978). They contain monoter-                     in the genus leading to such a grate morphologi-
pene with antiseptic characteristics (ÖZDEMIR                  cal diversity (RECHINGER 1982).
and SENEL 1999) and the compounds obtained                         RECHINGER et al. (1982), in Flora Iranica state
from these species decrease DNA synthesis in the               that infra-generic taxa recognized in Salvia, are
cell, an important feature in diagnosis and treat-             unsatisfactory and produce a false impression
                                                               of the genus. The genus Salvia is a vast genus
                                                               with about 800 species distributed throughout
                                                               the world and it is necessary to have a clear idea
                                                               about the range of morphological variations in
                                                               the genus as a whole in order to decide about
    *Corresponding author: e-mail msheidai@yahoo.com           its classification. Therefore he suggests that
406                                                                                   MASOUD , ALIJANPOO   and KHAYYAMI

the only natural supra-specific categories to be                        MATERIALS AND METHODS
recognized are species-groups, which are often
small, not infrequently monotypic groups of                          Plant material - Plant materials collection was
clearly related species. He finally arranged the                 done in Central Alborze region during 2009 and
species in the four major informal groups based                  2010. Young flower buds could be obtained for in
largely on stamen structure within 3 of which,                   eleven populations of ten Salvia species growing
subdivisions based on habit for, leaf, calyx and                 wild in this region namely S. spinosa L., S. reuterana
corolla characters have been recognized.                         Boiss., S. sclarea (Mönch) Benth., S. ceratophylla L.,
    Various cytological studies performed on the                 S. xanthocheiala Boiss. ex. Benth., S. limbata C. A.
genus Salvia indicate surprisingly diverse chro-                 Mey., S. hypoleuca Benth., S. staminea Monthbr. &
mosome numbers in the genus. The genus seems                     Auch. ex. Benth., S. nemorosa L., and S. verticillata
to be polybasic, with different groups of species                L. The voucher specimens are deposited in Her-
in different parts having polyploid origins. The                 barium of Shahid Beheshti University (HSBU), de-
Mediterranean group seems to be characterized                    tails of which have been given in Table 2.
by x = 7 (AFZAL-RAFII 1976), those in Europe and                     Cytological studies - Meiotic studies were per-
Russia by x = 11 (PATUDIN et al. 1975), and those                formed on young flower buds collected from at
studied in California by x = 16 (EPLING et al.                   least 10 randomly selected plants from each spe-
1962). Studies made by PALOMINO et al. (1986),                   cies and population. Minimum 100 metaphase/
show that Salvia subgenus Calosphace is charac-                  diakinesis pollen mother cells (PMCs) and 500
terized by x = 11 and lower numbers.                             anaphase and telophase cells were analysed for
    Cytological studies of the Salvia have been                  data collection. Pollen satiability as a measure
mainly focused on chromosome number reports                      of fertility was determined by staining minimum
and karyotype analysis (for example ÖZDEMIR                      1000 pollen grains with 2 % acetocarmine: 50 %
and SENEL 1999; YILDIZ and GÜCEL 2006; PALO-                     glycerin (1:1) for about ½ hr. Round complete
MINO et al. 1986; AL-TURKY et al. 2000; FOLEY et                 pollens which were stained were taken as fer-
al. 2008; AFZAL-RAFII 1976; 1980; 1981; PATUDIN                  tile, while incomplete, shrunken pollens with no
et al. 1975; MIZIANTI et al. 1981; HAQUE 1981;                   stain were considered as infertile. .
BORGEN 1980; DÍAZ-LIFANTE et al. 1992; LEE                           Statistical analyses - χ2 test was performed
1967), while study of the chromosomes behavior                   to detect a significant difference in chiasma fre-
in meiosis is very scare (BAHATTACHARYA 1978;                    quency and chromosome pairing among the spe-
GHAFFARI and CHARIAT-PANAHI 1985; ESTILAI                        cies and populations studied. T-test analysis was
and HASHEMI 1990). We hope by studying mei-                      performed to study size difference between re-
otic peculiarities of Salvia species, a better un-               duced and unreduced pollen grains. At least 50
derstanding of species relationships would be                    larger pollen grains and 50 smaller pollen grains
obtained in future.                                              were randomly measured for t-test analysis.

TABLE 1 — Meiotic characteristics in Salvia species studied.

     Species        Locality     2n       TX         IX        TOX     RB      RD        PF     A1L        A2L    M1ST
S. nemorosa 1       Khojier      14        6.22     2.13        8.35   2.70    3.39     90.50    5.26      0.00    0.00
S. nemorosa 2       Gachsar      14        5.36     2.45        7.86   3.23    2.05     99.90    3.18      0.40    5.26
S. hypoleuca        Khojier      22       11.22     1.42       12.64   3.44    5.89     98.70   18.50      0.00    8.66
S. ceratophylla   Sorkhehesar    22        8.35     4.35       12.76   5.11    3.03     98.30    7.60      0.00   12.38
S. limbata          Khojier      22       10.85     2.84       13.70   4.56    4.69     99.80   15.87      7.14    0.00
S. verticellata     Gachsar      16       16.94     6.12       23.06   7.76    6.24     95.50    6.00      5.60    3.61
S. sclarea           Poloor      22        8.77     3.61       12.39   4.65    2.90     99.50    2.00      0.00    2.00
S. staminea          Abali       22        7.59     2.15        9.73   2.88    3.78     99.00    6.77      0.00    2.18
S. reuterana        Khojier      20       11.05     4.65       15.63   6.67    2.00     97.33   10.60      0.00    3.40
S. spinosa          Khojier      20        5.74     2.57        8.26   3.43    2.05     98.73   10.00      0.00    1.00
S. xanthochilla      Poloor      22        6.62     4.62       11.24   5.06    1.79     97.30    3.70      0.00    2.36
Abbreviations: TX = Terminal chiasmata, IX = Intercalary chiasmata, TOX = Total chiasmata, RB = Ring bivalents, RD = Rod
bivalents, PF = Pollen fertility percentage, A1L = Anaphase I laggards percentage, A2L = Anaphase II laggards percentage
and M1ST = Metaphase I stickiness percentage.
CONTRIBUTION TO CYTOLOGY OF GENUS SALVIA L .      ( LAMIACEAE )   IN IRAN                                           407

         RESULTS AND DISCUSSION                                al. (1981) and GHAFFARI and CHARIAT-PANAHI
                                                               (1985), S. hypoleuca, S. ceratophylla, S. limbata
    Ploidy level and chiasma frequency - Details               and S. sclarea and S. xanthocheila showed 2n = 22
of cytological characteristics are presented in Ta-            chromosome number, supporting the reports
bles 1-3, Fig. 1. The species studied had 2n = 14,             of AFZAL-RAFII, (1980; 1981), as well as ÖZ-
20, 22 and 32 chromosome numbers. S. nemoro-                   DEMIR and SENEL (1999), S. verticillata showed
sa showed 2n = 14 in both populations studied                  2n = 16 supporting the reports of AFZAL-RAFII,
supporting the earlier reports of MIZIANTI et                  (1980) and LÖVKVIST and HULTGÅRD (1999), S.

                                                                                                     10 µm

Fig. 1 — Representative meiotic cells in Salvia species studied. A = Meiocyte showing 2n = 22 in S. ceratophylla (arrow
indicated quarivalent). B = Meiocyte showing 2n = 20 in S. spinosa. C-E = Meiocytes showing 2n = 22 in S. limbata,
S. scalrea and S. hypoleuca respectively. F = A polyploid meiocyte in S. limbata. G and H = Meiocytes showing B-
chromosomes in S. reuterana and S. nemorosa (arrows) respectively. I and J= Micronuclei formation in S. hypoleuca
and S. reuterana respectively. K and L = Tripolar cells in S. scalrea and S. spinosa. M-O = Potential unreduced pollen
grains (larger pollen grains) in S. reuterana, S. xanthochilla and S. spinosa respectively. The bar corresponds to 10 µm.
408                                                                                MASOUD , ALIJANPOO   and KHAYYAMI

reuteriana showed 2n = 20, supporting the re-                and rod bivalents in metaphase of meiosis-I with
ports of AFZAL-RAFII, (1981), as well as GHAFFARI            almost high pollen fertility (>0.90%). In two mei-
and CHARIAT-PANAHI (1985). S. spinosa showed                 otic cells of S. ceratophylla, one ring quadrivalent
2n = 20 which seems to be new to science.                    was observed which due to low frequency were
    The occurrence of different basic chromo-                not included in final analysis. Such quadrivalents
some numbers in a single species has been re-                may be formed due to heterozygote transloca-
ported in the genus Salvia. For example in S.                tions among two pairs of chromosomes.
aegyptiaca L., 2n = 12, 26 and 28 have been re-                  Among Salvia species, the highest value of
ported by HAQUE (1981), BORGEN (1980), and                   relative total, terminal and intercalary chiasmata
DÍAZ-LIFANTE et al. (1992), in S. chanroenica ssp.           occurred in S. verticellata (2.12, 0.76 and 2.88
glomerifolia Chung, 2n = 14 and 16 have been re-             respectively) while the lowest value of the total
ported by LEE (1967), in S. nemorosa L. 2n = 14              and terminal chiasmata occurred in S. spinosa
and 16 were reported by MIZIANTI et al. (1981)               (0.88 and 0.57 respectively). The lowest value of
and GHAFFARI and CHARIAT-PANAHI (1985). All                  intercalary chiasmata occurred in Kojier popu-
these data indicate the role played by polyploidy            lation of S. nemorosa (0.30). χ2 test performed
and aneuploidy in Salvia species diversification.            showed no significant difference for chiasma
    The chromosome number so far reported in                 frequency and distribution among Salvia species
Salvia species, fall into different aneuploid series         indicating that during species diversification no
starting from x = 7 to 11. According to BAHAT-               significant change has occurred in the number of
TACHARYA (1978), base numbers 7 and 8 appear                 genes controlling chiasma formation. However
to be the primitive numbers from which second-               since variation in chiasma frequency and local-
ary base numbers became established and again                ization is genetically controlled (QUICKE 1993),
diversified in different directions, out of which            minor genetic changes leading to meiotic varia-
base number 11 is common in many species. FU-                tions is considered as a mean for generating new
JITA (1970), considered Salvia as the most primi-            forms of recombination influencing the variabil-
tive genus in the family Labiateae with the highest          ity within natural populations in an adaptive way
primary basic number 11 although Ajuga is often              (REES and DALE 1974).
considered as the most primitive genus. However,                 The correlation test showed no significant
according to BAHATTACHARYA (1978), due to cyto-              correlation between relative total, terminal and
logical numerical instability, the presence of highly        intercalary chiasmata as well as ring and rod
irregular cytological behavior and presence of ad-           bivalents with change in chromosome number
vanced karyotype in the Salvia species, they are re-         (ploidy level) of the species studied, indicating
cent and advanced members of a complex group.                the presence of a control over the mean value of
    Data with regard to chiasma frequency and                chiasmata/bivalent.
chromosome pairing has been given in Tables 1                    Meiotic abnormalities - Almost in all the spe-
and 2. The species studied mainly formed ring                cies studied, laggard chromosomes were ob-

TABLE 2 — Relative meiotic data and size of pollen grains in Salvia species studied.

     Species    Voucher No.      TXN          IXN        TOXN          RBN             RDN       NP          UP
S. nemorosa       8500980        0.89         0.30        1.19         0.39            0.48     27.18       41.91
S. nemorosa 2     8500990        0.77         0.35        1.12         0.46            0.29     27.18       41.92
S. hypoleuca      8500981        1.02         0.13        1.15         0.31            0.54     25.64       37.62
S. ceratophylla   8500982        0.76         0.40        1.16         0.46            0.28     30.92       52.63
S. limbata        8500983        0.99         0.26        1.25         0.41            0.43     27.52       34.53
S. verticellata   8500984        2.12         0.76        2.88         0.97            0.78     20.21       38.12
S. sclarea        8500985        0.80         0.33        1.13         0.42            0.26     30.91       42.59
S. staminea       8500986        0.69         0.20        0.88         0.26            0.34     37.87       45.88
S. reuterana      8500987        1.10         0.47        1.56         0.67            0.20     35.49       49.54
S. spinosa        8500988        0.57         0.26        0.83         0.34            0.21     28.48       38.10
S. xanthochilla   8500989        0.60         0.42        1.02         0.46            0.16     26.24       36.82
Abbreviations: TXN = Terminal chiasmata/bivalent, IXN =Intercalary chiasmata/bivalent, TOXN = Total chiasmata/biva-
lent, RBN =Ring bivalent/cell, RDN = Rod bivalent/cell, NP = Size of normal pollen grains (µm), UP = Size of unreduced
pollen grains. (µm).
CONTRIBUTION TO CYTOLOGY OF GENUS SALVIA L .   ( LAMIACEAE )   IN IRAN                                    409

served during anaphase I and II and (Table 2).             1-2% of pollen grains in the species studied.
The highest value of anaphase-I laggards (18.50)               The mean diameter of normal (reduced) pol-
occurred in S. hypoleuca followed by S. limbata            len grains ranged from 20.21 in S. verticellata to
(15.00), while S. sclarea showed the lowest value          37.87 µm in S. staminea, while the size of poten-
of the same (2.00). Laggard chromosomes would              tial unreduced (2n) pollen grains ranged from
lead to micronuclei formation observed in most             34.53 in S. limbata to 52.63 µm in S. ceratophylla.
of the species (Fig. 1, I and J).                          T-test analysis revealed a significant difference
    The chromosomes stickiness occurred from               (p 90% (Table 2), therefore it seems that meiotic            tion. To our knowledge this is the first report on
abnormalities may be partly the reason for some            the occurrence of unreduced pollen grain for-
degree of pollen sterility observed in Salvia spe-         mation in the genus Salvia.
cies. However it has been suggested that infertil-             B-chromosomes - B-chromosomes (Bs) of
ity in polyploids is not solely due to the produc-         0-2 were observed in the species of S. sclarea,
tion of aneuploid gametes formed by improper               S. nemorosa and S. reuterana (Fig. 1, D, G and
segregation of chromosomes during anaphase/                H). B-chromosomes were round in shape and
telophase stages, the genetic factors may also             did not pair with the A-chromosomes or among
bring about pollen sterility as evidenced in tet-          themselves. These chromosomes are also known
raploid strains of rye (HAZARIKA and REES 1967)            as accessory chromosomes reported in about
and Avena sativa cultivars (BAPTISTA-GIACOMELLI            1300 plants species and 500 animals species
et al. 2000).                                              (CAMACHO et al. 2000). They can reduce the
    Other meiotic irregularities observed were             growth and vigor of the plants when present
tripolar and multipolar cell formation due to              in high number; but in low number may ben-
anaphase I and II failure (Fig.1, K and L) and             efit the plant possessing them. One of the well
meiotic cells with double the chromosome num-              known effects of B-chromosomes is the change
ber, possibly due syncyte formation and absence            they bring about on chiasma frequency, thereby
of anaphase segregation (Fig. 1, F). Bahattacha-           affecting the genetic changes of the gametes and
rya (1978) also reported meiotic irregularities in         the next generation as reported in several grass
two varieties of S. splendens Ker-Gawl (2n = 22),          species including Festuca (JAUHAR and CRANE
leading to formation of tripolar cells and pollen          1990), Aegilops (SHEIDAI et al. 2002), Avena
grains with size variation and polyspory.                  (SHEIDAI et al. 2003). However we did not have
    Unreduced pollen grain formation - The occur-          many cells possessing B-chromosomes to anal-
rence of large pollen grains (possibly 2n pollen           yse statistically the effect of B-chromosomes on
grains) was observed along with smaller (normal)           chiasma frequency in Salvia species. According
pollen grains in all the species studied (Fig.1,           to our knowledge this is the first report on the
M-O). The large pollen grains comprised about              occurrence of B-chromosomes in Salvia.
410                                                                              MASOUD , ALIJANPOO   and KHAYYAMI

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