6. botany-ijbr-chromosome numbers and abnormal

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www.tjprc.org [email protected]

CHROMOSOME NUMBERS AND ABNORMAL MALE MEIOSIS IN SIX SPECIES OF

CAMPANULACEAE FROM NORTHWESTERN HIMALAYAS

RAGHBIR CHAND GUPTA, REYAZ AHMAD MALIK, VIJAY SINGH & SANTOSH KUMARI

Department of Botany, Punjabi University, Patiala, Punjab, India

ABSTRACT

The present paper reveals meiotic analysis on 12 populations comprising 6 species belonging to 3 genera of the

family Campanulaceae from various localities of Northwest Himalayas. The chromosome number has been reported for the

first time in Campanula aristata (2n=32). Besides, there arefirst ever reports of tetraploid cytotypes in

Codonopsisrotundifolia (2n=32) and Cyananthus lobatus (2n=28). 0-2B chromosomes have been reported for the first time

in Campanulacashmeriana . Most of the populations show laggards, chromosome stickiness and cytomixis from early

prophase to telophase-II, leading to the formation of aneuploid cells or meiocytes with double the normal chromosome

number. Such meiotic abnormalitiesproduce unreduced pollen grains and the reduce pollen viability.

KEYWORDS: Campanulaceae, Chromosome Number, Meiotic Abnormalities, Northwest Himalayas

INTRODUCTION

Campanulaceae commonly known as bellflower or herbal family is almost cosmopolitan but concentrated in the

Northern Hemisphere. Campanulaceae Sensustricto contains about 600 species in 35 genera (Kovanda, 1978) to 950 species

(1045 as per Lammers, 2007) in 55 genera (Takhtajan, 1987) and Campanulaceae Sensu lato it contains about 90 generaand 2200 species (Judd et al., 2002). In India, it is represented by 8 genera and 48 species. Three genera and 12 species

occurs in Northwest Himalayas. Most of the species are herbaceous followed by shrubs and rare number of small trees.

Plants of this family often have an arid milky juice (Bailey, 1949). The family is an important source of variety of

polyacetylenic compounds which have proved to in-vitro toxicity and phytotoxicity against bacteria, fungi, and human

fibroblast and erythrocyte (for reviews: Badanyan et al.,1973; Bantley et al., 1973; Warren et al., 1980; Wat et al., 1980).

Campanula L.: It is the largest genus in the family Campanulaceae with about 300 species, out of which 150

species are distributed inthe Mediterranean region (Cronquist, 1988; Heywood, 1998). Besides, well known for ornamental

value, some species are known for medicinal importance like C. rotundifolia in heart and lung problems (Moerman, 1998).

Codonopsis Wall.: The genus is represented by 55 species in the world and mainly distributed in East Asia,

Himalaya and Malaysia (Morris and Lammers, 1997). Few species like, C. pilosula , contains an alkaloid Codonopsine, is

very nutritious and often referred to as poor man's ginseng (Wang et al., 1996).

Cyananthus L.: It is a genus that consists of ca. 20 species of annual or mostly perennial herbs from high

mountains of Central and East Asia (Zhou et al., 2013). Most of the species are laxative and used in the treatment of

various serous disorders (as dry up serous fluid) and constipation (Manandhar, 2002).

The family is well known for its medicinal and ornamental values as comprising beautiful blue flowers like,

Campanula , Symphyandra , Edraianthus , Cyananthus , etc. which attracts the researcher for the study of evolution,

International Journal of Botanyand Research (IJBR)ISSN(P): 2277-4815; ISSN(E): 2319-4456Vol. 4, Issue 3, Jun 2014, 39-50 TJPRC Pvt. Ltd.


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40 Raghbir Chand Gupta, Reyaz Ahmad Malik, Vijay Singh & Santosh Kumari

Impact Factor (JCC): 1.6913 Index Copernicus Value (ICV): 3.0

taxonomy, pollination, anatomy and biogeography. The present work has, therefore been undertaken to know the

cytological behavior of different species belonging to family Campanulaceae from the Northwestern Himalayas.

MATERIALS AND METHODS

Meiotic surveys have been made during the years from 2009-2013 to selected localities (Table 1) of Kashmir and

Himachal Pradesh (Kullu and Sirmaur Districts). Young flower buds of plants growing in their natural habitat were

collected and fixed in Carnoys fixative (Alcohol: Chloroform: Acetic acid in 6: 3: 1 ratio). After 24 hrs, the buds were

transferred in rectified alcohol for longer storage purposes. Meiotic studies were made by squashing in standard

acetocarmine. Photographs of pollen mother cells (PMCs) were made with freshly prepared slides by using a Nikon 80i

Eclipse Microscope. Pollen fertility was estimated by their stainability in 1% glyceroacetocarmine (1:1). Filled and stained

pollen grains considered as fertile whereas shrunken and unstained pollen grains are sterile. The specimens were submitted

to the Herbarium, Department of Botany, Punjabi University, Patiala. The data regarding the number of cytologically

worked out species, number of cytotypes, and frequency of polyploids of a particular genus has been compiled onworldwide and India basis from various Chromosomal Atlases and Indexes to Plant Chromosome Numbers by Darlington

and Wylie (1955), Fedorov (1974), Moore (1973, 1974, 1977), Goldblatt (1981, 1984, 1985, 1988), Goldblatt and Johnson

(1990, 1991, 1994, 1996, 1998, 2000, 2003, 2006), Kumar and Subramaniam (1986), and Khatoon and Ali (1993), various

journals, Internet, as well as presently studied species.

RESULTS

Presently, 12 populations of 6 species belonging to 3 genera of Campanulaceae have been worked out from

different localities of Kashmir and Himachal Pradesh at an altitude ranging 2400-4000 m. The data pertaining to taxon,

voucher data, average plant height, chromosome number, ploidy level, pollen fertility, pollen size and localities with

geographical co-ordination has been discussed in Table 1.

Chromosome Numbers

Campanula aristata Wall. in Roxb: The present chromosome report of 2n=32 (Figure 1a, b), is first ever

cytological report for the species.

C. cashmeriana Royle: In the present investigations, B-chromosomes have been reported for the first time in the

tetraploid cytotype (2n=28+0-2B, Figure 1c, d). The only previous cytological report (2n=28) for the species from India is

that of Jee et al .(1983, 1989), besides by some workers from outside India.

C. colorata Wall: Cytologically, both the populations are diploid (2n=28, Figure 1e), which is in conformity to

the previous reports by Sarkar et al. (1976) from India. Besides, another cytotype (2n=24) has been reported from India by

Kishore (1951).

Codonopsis ovata Benth: All the presently investigated accessions are diploid (2n=16, Figure 1f), and is in

conformity to the single report of Jee et al. (1989) from India.

C. rotundifolia Benth: The present chromosome number of 2n=32 (Figure 1g) is first tetraploid chromosome

count for the species, which is in line with the earlier report by authors (Malik and Gupta, 2013). Previously, the species is

known to exist at diploid level (2n=16) by Zhukova (1967) and octaploid level (2n=68) by Druskovic and Lokova (1995)


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Chromosome Numbers and Abnormal Male Meiosis in Six Species of Campanulaceae from Northwestern Himalayas 41


Cyananthus lobatus Wall. ex Benth: The present chromosome report of 2n=28 (Figure 1h) is first tetraploid

chromosome count for the species.

Meiotic Abnormalities

Meiotic abnormalities have been recorded in almost all the studied populations of different species in the form of

cytomixis, chromatin stickiness, unoriented bivalents, bridges, laggards or multipolarity at different stages of meiosis

(Table 2, Figure 2ak). The highest percentage of chromatin transfer from prophase-I to telophase-II has been recorded in

Codonopsis rotundifolia (Table 2, Figure 2a). Cytomixis usually led to the formation of pollen mother cells (PMCs) with

different chromosome numbers, and even empty PMCs in some cases (Figure 2b), as being evident in Campanula

cashmeriana . All these meiotically abnormal populations quite frequently showed the presence of chromosomal laggards

and bridges atanaphase and telophase (Figure 2c, d). Chromatin stickiness (partial or often complete clumping of bivalents,

Figure 2e) and unoriented bivalents were seen from prophase I to metaphase I and heighest frequency is found in

Campanula cashmeriana (24.64%) and Codonopsis rotundifolia (16.28%), respectively (Table 2). These meioticabnormalities led to abnormal microsporogenesis and the formation of heterogenous sized pollen grains

(Table 3, Figure 2f-k), and also affects pollen viablitity (Table 1).

DISCUSSIONS

Chromosome Numbers

Campanula : A total of 318 species are chromosomally known in the genus with the chromosome ranging from

2n=14102. The most common base number is x=17 as there are 193 species showing 2n=34 (diploid) and 26 species

showing 2n=68 (tetraploid). There is a lot of chromosome number variation in Campanula . The other chromosome

numbers available are 2n=14 (4 species), 2n=28 (13 species), and 2n=56 (5 species), all based on x=7. Base number x=8 is

also well represented since there are 13 species with 2n=16 and 20 species with 2n=32. Besides, 2n=20 is met with 18

species, 2n=30 in 9 species which support x=10 and /or x=15. Further, there is a significant number of species based on

x=12 showing 2n=24 (8 species) and 2n=48 (5 species). However, the intraspecific polyploidy is shown by 16 species on

x=17 only. The base number x=14 is considered as the secondary base number derived through polyploidization from x=7.

The genus is considered polybasic with a domination base number of x=17. From India, chromosome reports are available

for 3 species, of which two share 2n=28 and 2n=24, one with 2n=32 (From present data).

Codonopsis: The chromosome number is known for 9 species, all showing 2n=16, hence the genus is monobasic.

From India, now 2 species are worked out cytologically.

Cyananthus : Out of 23 taxonomically known species, only 3 are known cytologically. All the three species exist

on a single base number x=7, as the chromosomal range from 2n=14-28 (from present data).


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Table 1: The Table Showing Data on Voucher Number, Average Plant Height, Flowering - Fruiting, ChromosomeNumber (N), Ploidy Level (X), Pollen Fertility, Pollen Size, and Latitude and Altitudes Various Accession of Family

Campanulaceae from Northwest Himalayas

S.No.

TaxonData

(PUN)*

VoucherHeight

(cm)

PlantFruitin

g(Mean+SD)

FloweringNumber

(n)

Chromosome

Level (x)Fertility(

%)

Ploidy

Pollen Size(m)

Pollen(Mean+SD

)

Locality

with GeographicalCoordination,District

Altitude (m) & PlantHabitat

1Campanulaaristata

58812 333.2 June-August

8 2x 82 19.251.918.902.1

Marhi, 3235N7722E

Kullu, 3360, on rockyslopes

58813 352.9June-

August8 2x 82

19.890.919.102.8

Marhi, 3235N7722E

Kullu, 3360, on rockyslopes

2 C.cashmer iana 55183 301.5June-

August 14+1B* 2x 59

12.221.611.202.1-22.892.122.471.3

Aharbal, 3338 N

7446EKulgam, 2400, on

slopes near shadyplaces

3C.

colorata

55184 522.1May-

August 14 2x 7523.871.221.521.1

Aharbal, 3338 N7446 E

Kulgam, 2400, onshady grassy slopes

58640 452.4 May-August 14 2x 75

13.560.614.210.8

22.840.9420.230.5

Tisri, 30 51' N 77 33'E,

Sirmaur, 3100, shadygrassy slopes

4Codonopsi

s ovata

57002 421.4 June-August 08 2x 8257.660.450.591.2

Thajwas, 34 17' N 7517' E

Rocky shady slpoes

56930 400.5 June-August

08 2x 76 54.890.454.350.9

Amarnath Road, 340034 17' N 75 17' E,

56993 411.2 June-August 08 2x 7555.880.750.711.5

Apahrwat, 34 00' N75 17' E

Gulmarg, 4000Rocky shady slpoes

5C.

rotundifolia

57003 1700.3

June-Septemb

er16 4x 78

37.021.335.02 1.844.050.742.810.9

Aharbal, 3338 N7443 E

Kulgam, 2400, ongrassy slopes

570041700.

3

June-

September 16

4x 82

38.590.837.500.9

43.262.141.922.4

Gurez, 3338 N

7446Eon rocky shady palces

6 Cyananthus lobatus

58810 148.2June-

September

14 4x 80 39.122.138.341.9

Rohtang, 32 22 N,77 14 E

Kullu, 3978, on grassyslopes

58811 302.3June-

September

14 4x 8037.652.538.272.2

Marhi, 3235N7722E

Kullu, 3360, on grassyslopes

* Herbarium, Punjabi University, Patiala, First ever chromosomal report, First ever cytotype report, * First report forB-chromosomes.


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Table 2: Data On Cytomixis and Meiotic Course in the Studied Populations of Family Campanulaceae from NorthwestHimalayas

AccesionNumber Involved

Cytomixis Meiotic course

PMCsInvolved

No. of

PMCsStickiness at M-I

PMCs withChromosomal

PMCs withUnorientedBivalents atM-I

PMC

withBridges(at A-I,II/T-I,II)

PMCs

withLaggards(at A-I,II/T-I,II)

Campanulaaristata

58812 2.43(3/123) 1-2 9.60 (12/125) 1.16 (2/125)0.8

(1/125)8.0

(10/125)

58813 5.42 (7/129) 1-2 9.30 (12/129) 3.10 (4/129) 3.84(8/130)

10.56(13/123)

Campanula cashmeriana

551834.35%(5/144)

2-3 24.64% (35/142)8.39%(11/131)

2.20(3/136)

15.12%(18/119)

C. colorata

55184 3.96 (8/126) 1-2 5.50% (6/109) 2.30 (3/130)4.31(6/139)

1.55(2/129)

5864025.62(31/121) 2-6 14.50% (19/131) 2.38 (3/126)

12.5(17/136)

10.44(14/134)

Codonopsis ovata

55474 1.96 (2/102) 1-2 11.53% (15/130) 4.65 (6/129) 1.92%(2/104) 1.80(2/11

1)

58302 1.72 (2/160) 1-2 19.56% (27/138) 3.84 (5/130) 11.71%(15/128)8.03

(9/112)

58303 2.50 (3/120) 1-2 20.15% (26/129) 5.46%(7/128)13.79%(16/116)

9.73(11/113)

C. rotundifolia

57003 31.4%(38/121) 2-6 2.85% (3/105)14.1%(19/135)

27.34%(35/128)

57004 29.91%(35/117)

2-6 9.32% (11/118) 16.28%(21/129)

26.95%(38/141)

Cyananthus lobatus

58810 2.38(3/126) 1-2 1.78 (2/112)3.25(4/123)

58811 2.25 (3/133) 1-2 11.71 (13/111) 11.57(14/121)7.20

(9/125)10.08

(12/119)

Figures in parenthesis denote observed number of abnormal PMCs in the numerator and total number of PMCs

observed in denominator, WMN without micronuclei, WM with micronuclei

Table: 3. Data on Abnormal Microsporogenesis on Different Accession of Camapnulaceae from Northwest Himalayas

Taxon/AccessionNumbers

MonadsWM(%)

Dyads Triads Tetrads

WM(%)

WMN(%) WM (%)

WMN(%) WM (%)

WMN (%)

Campanulaaristata

58812 1.49(2/134)0.74

(1/134)4.45

(6/134) 02.23

(3/134)3.73

(5/134)87.31

(117/134)

58813 1.51(2/132) 0.75(1/132) 6.81(9/132) 0.75(1/132) 3.78(5/132) 0 87.12(115/132)


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Campanula cashmeriana

55183 1.66(2/121)0.82

(1/121) 00.82

(1/121)0.82

(1/121)13.22

(16/121)82.64

(100/121)C. colorata

55184 0 0 1.66(2/121) 04.13

(5/121) 095.04

(115/121)

58640 0.94(1/106) 00.94

(1/106)0.94

(1/106)5.66

(6/106)3.77

(4/106)87.73

(93/106)Codonopsis ovata

55474 0.85(1/117)5.12

(6/117)1.70

(2/117)9.40

(11/117) 00.85

(1/117)82.05

(96/117)

58302 0 7.2(9/125)1.60

(2/125)4.0

(5/125)1.60

(2/125)5.60

(7/125)80.0

(100/125)

58303 0.74(1/134)0.74

(1/134) 04.45

(6/134) 0 090.59

(106/117)C. rotundifolia

57003 1.44(1/69) 1.44(1/69) 2.89(2/69) 1.44(1/69) 4.34(3/69) 10.14(7/69) 78.26(54/69)

57004 2.29(2/87)4.59

(4/87)1.14

(1/87)2.29

(2/87)3.44

(3/87)11.49

(10/87)72.41

(63/87)Cyananthus lobatus

58810 4.36(5/114)1.75

(2/114)3.50

(4/114)5.26

(6/114)4.38

(5/114)1.75

(2/114)78.9

(90/114)

58811 1.66(2/121)4.13

(5/121)0.82

(1/121)4.13

(5/121)0.82

(1/121)0.82

(1/121)87.60

(106/121)

Figures in parenthesis denote observed number of abnormal PMCs in the numerator and total number of PMCs

observed in denominator, WMN - without micronuclei, WM with micronuclei.

The most common chromosome number in the Campanulaceae is n=17 and this appears to have evolved

independently several times in relatively unrelated genera (e.g. Campanula, Nesosodon, Ostrowskia and Canaria ).

The family is supposed to be monobasic on x=8 (For reviews, see: Bocher 1964; Contandriopoulos 1984), but Raven

(1975) and Zhang et al. (2011) suggested x=7 as the ancestral number. The ancestral base number of x=7 is supported by

counts for Cyananthus (Hong and Ma, 1991) which is considered as most primitive genus within the family

(For reviews, see: Hutchinson, 1969; Takhtajan, 1969; Carolin, 1977; Cronquist, 1988) based on its superior ovary.

Chromosome breakage may result into x=8 and x=9, such as in Codonopsis . The other possibility to form x=18 is

autopolyploidization of x=9 and that form paleopolyploids and then gradually modern secondary diploids or polyploids

(Zhang et al., 2011).

Meiotic Abnormalities

The present study of different species at population-level depicts the occurrence of different frequency of meiotic

abnormalities in the form of cytomixis, unoriented bivalents, chromatin stickiness, chromatin bridges and laggards, or

multipolarity, which certainly reflects intraspecific genetic diversity (for reviews, see: Baptista-Giacomelli et al., 2000;

Sheidai et al., 2003). The chromatin migration (one or a few chromosomes) into the neighbouring meiocytes leads to the

formation of hyper-/hypoploid cells, syncyte formation, and even double the chromosome number, eventually producing

unreduced pollen grains (for reviews, see: Falistocco et al., 1995; Sheidai and Fadaei, 2005; Ghaffari, 2006; Fadaei

et al., 2010; Jeelani et al., 2011; Kumar et al., 2013). Chromatin transfer and stickiness might be the projection ofgenetic/environmental factors or genomicenvironmental interaction (for reviews, see: Nirmala and Rao, 1996;


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Baptista-Giacomelli et al., 2000; Bellucci et al., 2003; Fadaei et al., 2010; Kumar and Singhal, 2011).

Chromosome stickiness also results in the formation of fragmented chromatin. Besides, non-synchronous behavior of

bivalents may lead to unoriented bivalents, as they fail to arrange on spindle apparatus. This chromatin stickiness, late or

non-disjuncting bivalents, and chromosomal laggards seems to be responsible for chromosomal bridges(Kumar et al., 2013). All these meiotic abnormalities assert an effect on microsporogenesis, leading to the formation of

monads, dyads, triads, or polyads with or without micronuclei, which ultimately produce heterogeneous sized

(large and small) fertile pollen grains and reduced pollen fertility. The size difference may be due to the formation of

unreduced gametes (2n), which may produce plants with higher ploidy level through polyploidization

(for reviews, see: Villeux, 1985; Thomson, 1995; Sheidai et al., 2008; Fadaei et al., 2010; Jeelani et al., 2011).

CONCLUSIONS

The chromatin rearrangement due to meiotic abnormalities (as observed in presently investigated data) are

considered the base of inter/ or intraspecific diversity. Further, it provides a catalogue for studying different evolutionarytrends such as breeding system or polyploidy and hybridization. Thus, these meiotic variability persuades the further

cytological exploration on different species of Campanulaceae from various localities of Northwestern Himalayas at

population basis..

ACKNOWLEDGEMENTS

The authors are grateful to the I.P.L.S. (DBT) and University Grants Commission, New Delhi (DRS SAP III) for

providing financial assistance. We are highly thankful to the Head, Department of Botany, Punjabi University, Patiala for

providing necessary lab facilities during the work.

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Figure Legends

Figure 1 (a-h): Chromosome Numbers in Six Species of Campanulaceae from Northwestern Himalayas. (a-b)Campanula Aristata, PMC at Metaphase-I Showing 16 II, PMC at Anaphase-I Showing 16:16 Distribution ofChromosome (c-d) C. Cashmeriana, PMC at Metaphase-I Showing 14 II , PMC at Anaphase Showing 14:14

Chromosomal Distribution and I B Chromosome (arrowed) (e) C. Colorata, PMC at Anaphase-I showing 14:14Distribution of Chromosome (f) Codonopsis Ovata, PMC at Metaphase-I with 8 II, (g) C. Rotundifolia, PMC at


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Anaphase-I Showing 16:16 Distribution of Chromosome (h) Cyananthus Lobatus, PMC at Diakinesis Showing 14 II.(II - Bivalents, Scale 10 m)

Figure 2 (a-k): Meiotic Abnormalities in Studied Populations of Campanulaceae. (a) Different Groups of PMCsInvolved in Cytomixis. (b) PMC Showing Extra Chromatin Mass due to Cytomixis (Arrowed) (c-d) Both PMCsshowing Laggards (arrowed) (e) PMC at Metaphase-I Showing Chromatin Stickiness (f) Monads (g) Dyads with

Micronucleous (h) Triad (i) Tetrad with Micronucleous (j-k) Heterogenous Sized Fertile and Sterile Pollen Grains.(Scale 10 m)


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6. botany-ijbr-chromosome numbers and abnormal

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