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Meiotic studies in some species of tribe Cichorieae (Asteraceae) from Western Himalayas.

1. Introduction

The tribe Cichorieae (also known as Lactuceae) encompasses 95 genera and ca. 2500 species, primarily in temperate to subtropical zones of the Northern Hemisphere [1]. Members of the tribe are characterized by very important uniform characteristics, such as homogamous ligulate capitula and the presence of milky latex.

The presently investigated species are also known to have medicinal uses, such as Hieracium crocatum to cure gastric troubles, Lactuca dolichophylla to cure constipation, L. macrorhiza used as an ingredient to cure stomach ache, L. serriola to treat ailments of the urinary tract, and Taraxacum officinale used as diuretic and laxative [2-4].

Chromosome studies are valuable determinants in studying evolution. Many workers have studied the cytology of Indian Asteraceae, including members of tribe Cichorieae. Cytological studies on the family from Lahaul-Spiti area, an ecologically very fragile cold desert area of Western Himalayas, are almost lacking. As an attempt to fill this lacuna, the present investigations have been undertaken.

2. Materials and Methods

2.1. Plant Material. Exploratory surveys were made during the years from 2009 to 2013 in selected localities (Table 1) of Himachal Pradesh (Kullu and Lahaul-Spiti Districts). The cytologically worked-out plants were identified using regional floras and compared with the specimens deposited at the Herbarium of Botanical Survey of India, Northern Circle, Dehra Dun. The voucher specimens (Table 1) were deposited in the Herbarium, Department of Botany, Punjabi University, Patiala (PUN).

2.2. Meiotic Studies. For meiotic chromosome counts, unopened floral buds of suitable sizes were fixed in a freshly prepared Carnoy's fixative (mixture of alcohol, chloroform, and glacial acetic acid in a volume ratio 6: 3: 1) for 24 h. These were subsequently transferred to 70% alcohol and stored in refrigerator at 4[degrees]C until used for meiotic analysis. Meiocytes were prepared by squashing the developing anthers and stained with acetocarmine (1%). Chromosome number was determined at diakinesis/M-I/II/A-I/II from freshly prepared slides with light microscope Olympus. 500-600 pollen mother cells were analyzed for meiotic behaviour at different stages, metaphase-I/II (M-I/II), anaphase-I/II (A-I/II), and telophase-I/II (T-I/II).

2.3. Pollen Grain Analysis. Pollen fertility was estimated through stainability tests using glycerol-acetocarmine (1: 1) mixture and aniline blue (1%). Up to 450-800 pollen grains were examined for pollen fertility and size frequencies. Well-filled pollen grains with stained nuclei were taken as apparently fertile while shriveled and unstained pollens were counted as sterile.

2.4. Photomicrographs. Photomicrographs from the freshly prepared desirable slides having clear chromosome counts, dyads, triads, tetrads, and pollen grains were taken with a digital imaging system of Leica QWin.

3. Results

3.1. Chromosome Number

Hieracium crocatum Bunge ex. Ledeb. The present species revealed the diploid cytotype (2n = 10, Figure 1(a)), which is a first ever chromosome report for the species.

H. umbellatum L. Both the cytotypes, 2n = 10 (Figure 1(b)) and 2 n = 54 (Figure 1(c)), are the new records from the world, although the species is already known to have 2n = 18 [5] and 2n = 27 [6] from outside India. Polyploid cytotypes show some enlargement in vegetative and floral characters (Table 2).

Lactuca dissecta D. Don. The present report (2n = 16, Figure 1(d)) is in line with many previous reports from India [7-10] and abroad [11].

L. dolichophylla Kitam. The present chromosome report of 2n = 16 (Figure 1(e)) is in line with many previous reports from India [7, 8,10,12].

L. lessertiana (Wall. ex DC) C. B. Clarke. The present meiotic studies reveal a diploid cytotype (2n = 16, Figure 1(f)) which is a first ever chromosome report for the species.

L. macrorhiza (Royle) Hook. f. The present chromosome report of 2n = 16 (Figure 1(g)) is already confirmed by many workers [8,12,13] from India. There is no chromosome record from abroad for the species.

L. sativa L. Meiotic analysis of the species reveals the diploid cytotype (2n = 18, Figure 1(h)), which is in conformity with the previous works of Chatterjee and Sharma [14] and Gupta and Gill [15].

L. serriola L. The present chromosome report (2n = 18, Figure 1(i)) is confirmed by many workers from India [7,12, 15,16].

Prenanthes brunoniana C. B. Clarke. The present chromosome report of 2n = 16 (Figure 1(j)) is in conformity to only previous report from Garhwal, Uttarakhand, by Shetty [8].

Taraxacum officinale L. The present meiotic investigation revealed three cytotypes, 2n = 2x = 16 (Figure 1(k)), 2n = 3x = 24 (Figure 1(l)), and 2n = 4x = 32 (Figure 1(m)). Both diploid and tetraploid cytotypes are common and are reported by many workers [17]. Gupta et al. [13], besides triploid cytotype, also reported some other cytotypes, that is, 2n = 26, 27, 32, 38, and 40. Morphologically, the tetraploids do not show any robust and gigas effect due to polyploidy Table (2), but they certainly show a lot of variation in shape of leaves.

Tragopogon dubius Scop. Both the cytotypes, 2n = 14 (Figure 1(n)) and 2n = 28 (Figure 1(o)), are varied cytotypes at world level. The species is reported earlier with 2n = 24 by Koul and Gohil [18] and Mehra and Remanandan [12] from Kashmir Himalayas. From outside India, the species is known to have 2n = 12, 24, and 36 [5]. Morphologically, tetraploid cytotype does not show any gigas effect as compared to diploid (Table 2).

T. gracilis D. Don. The present chromosome report (2n = 14, Figure 1(p)) is a varied chromosome count for the species at world level. Earlier, Mehra and Remanandan [12] reported diploid cytotype with 2n = 12 from the Western Himalayas.

Youngia glauca Edgew. The present cytological investigation reveals the diploid cytotype (2n = 14, Figure 1(q)), which is a varied chromosome report. Earlier, there is a maiden cytological report of 2n = 16 [19] from Kinnaur valley.

Y. japonica (L.) DC. The present chromosome report (2n = 16, Figure 1(r)) confirms the earlier reports from different localities of India and abroad [20].

Y. tenuifolia (Wild.) Babcock and Stebbins. This chromosome report of 2n = 10 (Figure 1(s)) is for the first time reported from India. The same number is frequently reported from abroad [21, 22].

3.2. Meiotic Abnormalities. Meiotic abnormalities have been recorded in almost all the studied populations of different species in the form of cytomixis, chromatin stickiness, unoriented bivalent, bridges, laggards, or multipolarity at different stages of meiosis (Tables 3 and 4, Figures 2(a)-2(k)). From the data, triploid cytotype of Taraxacum officinale shows the highest percentage of chromatin transfer from prophase-I to telophase-II (Table 2, Figure 2(a)). Cytomixis usually led to the formation of pollen mother cells (PMCs) with different chromosome numbers and even empty PMCs in some cases (Figure 2(b)), as is evident in hexaploid cytotype of Hieracium umbellatum. Chromatin stickiness (partial or often complete clumping of bivalents) is found in 16.28 per cent of PMCs and unoriented bivalents were seen in 13.79 per cent of PMCs of tetraploid cytotype of T officinale (Table 3, Figure 2(c)). The present investigation also reveals abnormal meiosis in the form of chromosomal laggards (maximum in triploid cytotype of T. officinale) and bridges (maximum percentage in tetraploid cytotype of H. umbellatum) at anaphase-I/II and telophase-I/II (Figures 2(e)-2(f)). These meiotic abnormalities led to the abnormal microsporogenesis and the formation of heterogeneous sized pollen grains (Table 4, Figures 2(g)-2(k)) and also affect pollen viability (Table 1).

4. Discussion

4.1. Chromosome Number

4.1.1. Tribe Cichorieae. The ancestral basic numbers in the tribe are x = 4, 5, and 9, as suggested by Turner et al. [23]. But Stebbins et al. [24] proposed x = 9 as the base number for Cichorieae (Asteraceae in general), which is supported by Tomb et al. [25], with the other numbers (x = 3-9) derived through phylogenetic reduction through chromosomal aberrations, particularly translocation.

Hieracium L. About 360 species are cytologically known with 94 species being diploid and 152 triploid and 149 species are tetraploid and rarely possess aneuploidy (7 spp.). The genus forms agamic complex and is considered monobasic on x = 9. But, the present study reveals another cytotype (2n = 10), suggesting a new base number (x = 5) in the genus.

Lactuca L. A total of 150 species are known taxonomically, of which chromosome numbers for 87 species overall and 14 species from India are known. The chromosome number in the genus varies in the range of 2n = 10-48 and is polybasic on x = 5, 8, 9, and 17, of which x = 9 is the most dominant number.

Prenanthes L. Twenty-two species in the genus are known cytotaxonomically, including 1 from India. The most common base number is x = 8 represented with 19 species, including diploids (16 species) and tetraploids (3 species). However, the intraspecific polyploids are not available in the genus. Besides, x = 9 is also present in 3 species that are diploid. Hence, the genus is proposed to be dibasic on x = 8 and 9.

Taraxacum L. The genus is very complex, reinforcing the reason of having 347 cytologically (including 10 species from India) worked-out species. The chromosome numbers vary in the range of 2n = 8-64, the most common of which is 2n = 3x = 24 (230 spp.) on x = 8, followed by diploid (47 spp.) on the same base number. Genus is reported to have a series of base numbers on x = 4, 6, 8, 9, and 11, but only x = 8 is known to have well-developed polyploid races (2x-6x). Intraspecific polyploidy is also known to occur in x = 9 and 11 (1 species each).

Tragopogon L. 75 species in the genusare cytologically known, with chromosome number in the range of 2n = 12-36, almost all based on x = 6. The overall polyploidy in the genus is 26.6% (20 spp.), out of which 14 species show intraspecific polyploidy. The variable chromosome number of 2n = 14 is found in 4 species (including the present data) and 2n = 28 in only one species (from the present data).

Youngia L. A total of 35 species are taxonomically known, cytology is reported for only 14 species (including 5 from India), with 9 species showing polyploid cytotypes (3v, 4v, and 6v). The chromosome numbers reported so far are 2n = 10,15,16, 20, 24, 32, and 42, out of which 2n = 10 (43.7%) is the most common followed by 2n = 16 (31.2%). The genus is polybasic (x = 5, 7, and 8), of which x = 5 is most common.

4.2. Meiotic Abnormality. The phenomenon of inter-PMC migration of chromatin/chromosome between/among the contiguous meiocytes through cytomictic channels is termed as cytomixis (coined by Gates [26]). However, the phenomenon has been reported for the first time in gymnosperms by Arnoldy [27] and subsequently in angiosperms by Koernicke [28]. Since that time, cytomixis has been reported in a large number of plants [29]. Transfer of chromatin or chromosomes may take place through such inter-PMC cytomictic channels [30-32]. Some workers reported cytomixis to be more prevalent in polyploids than their diploid counterparts [33, 34]. Occasionally, either hypoploid meiocytes [35-37] or enucleated meiocytes or meiocytes with a hyperploid number of chromosomes have been reported [30, 37-39]. It is very much clear that the enucleated meiocytes die, but hypo- and hyperploid meiocytes could lead to the formation of gametes with variable chromosome number and size. Cytomixis is considered as a process of evolutionary significance because it results in change in gametic chromosome numbers [30,40]. Chromosome stickiness also results in the formation of fragmented chromatin. This chromatin stickiness, late or nondisjuncting bivalents, and chromosomal laggards seem to be responsible for chromosomal bridges [41]. All these meiotic abnormalities consequently 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 ploidal level through polyploidization (for review, see [42-45]).

As observed in the presently investigated data, the chromatin rearrangement due to meiotic abnormalities is considered the base of inter- or intraspecific diversity. Further, it provides a catalogue for studying different evolutionary trends such as breeding system or polyploidy and hybridization.

Conflict of Interests

The authors declare that there is no conflict of interests regarding the publication of this paper.

http://dx.doi.org/10.1155/2014/673456

Acknowledgments

The authors are grateful to the University Grants Commission of New Delhi (DRS SAP III) and I.P.L.S. (DBT) for providing financial assistance. The authors are highly thankful to the Head, Department of Botany, Punjabi University, Patiala, for providing necessary lab facilities during the work.

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Raghbir Chand Gupta, (1) Henna Goyal, (1) Vijay Singh, (1) and Rajesh Kumar Goel (2)

(1) Department of Botany, Punjabi University, Patiala 147002, India

(2) Department of Pharmaceutical Sciences and Drug Research, Punjabi University, Patiala 147002, India

Correspondence should be addressed to Vijay Singh; vijaykataria05@rediffmail.com

Received 28 August 2014; Accepted 16 October 2014; Published 13 November 2014

Academic Editor: Jennifer A. Tate

TABLE 1: The table showing details on taxon, geographical
coordination, accession number, average plant height,
flowering-fruiting period, chromosome number (2/j),
ploidal level (x), and pollen fertility of tribe
Cichorieae (Asteraceae) from Western Himalayas.

Taxon                Locality with           Accession
                     altitude (m) and         number
                     geographical             (PUP *)
                     coordinates

Hieracium crocatum   Trilokinath, Lahaul       52760
  Bunge ex Ledeb.      (H.P.), 2760
                       32[degrees]42'0"N,
                       76[degrees]41'0" E
H. umbellatum L.
    (P1)             Jispa, Spiti              52763
                       (H.P.), 3142
                       32[degrees]38'0"N,
                       77[degrees]10'0"E
    (P2)             Keylong, Lahaul           52774
                       (H.P.), 3080
                       32[degrees]34'48"N,
                       77[degrees]1'48"E
L. dissecta D.       Hadimba, Temple,          58562
  Don.                 Manali, 2438
                       32[degrees]14'32"N,
                       77[degrees]11'14"E
L. dolichophylla     Sissu, Lahaul (H.P.),     57504
  Kitamura             3130
                       32[degrees]29'0"N,
                       77[degrees]7'0"E
L. lessertiana       Batal, Spiti (H.P.),      52776
  C. B. Clarke         3890
                       32[degrees]21'28"N
                       77[degrees]37'10"E
L. macrorhiza        Chotadhara, Spiti         57510
  (Royle) Hook. f.     (H.P.) 3800
L. sativa L.         Hadimba, Temple,          58534
                       Manali, 2438
                       32[degrees]14'32"N,
                       77[degrees]11'14"E
L. serriola L.       Jispa, Spiti              58535
                       (H.P), 3200
                       32[degrees]38'0"N,
                       77[degrees]10'0"E
Prenanthes           Tandi, Keylong            52764
  brunoniana C. B.     (H.P), 2573
  Clarke               32[degrees]34'40"N,
                       77[degrees]1'36"E
Taraxacum
  officinale L.
    (P1)             Marhi, Manali             52687
                       (H.P), 3320
                       32[degrees]20'56"N,
                       77[degrees]13'4" E
    (P2)             Keylong, Lahaul           52693
                       (H.P), 3080
                       32[degrees]34'48"N,
                       77[degrees]1'48"E
    (P3)             Kibber, Spiti             57491
                       (H.P), 4205
                       32[degrees]19'54"N,
                       78[degrees]0'32"E
Tragopogon dubius
  Scop.
    (P1)             Keylong, Lahaul           57503
                       (H.P), 3080
                       32[degrees]34'48"N,
                       77[degrees]1'48"E
    (P2)             Lossar, Spiti             57505
                       (H.P), 4079
                       32[degrees]24'49"N,
                       77[degrees]49'11"E
T. gracilis D. Don   Koksar, Lahaul            52768
                       (H.P.), 3160
                       32[degrees]41'37"N,
                       77[degrees]23'54"E
Youngia glauca       Zingzingbar, Lahaul       52773
  Edgew.               (H.P), 4270
                       32[degrees]47'30"N,
                       77[degrees]19'28"E
Y. japonica (L.)     Hadimba, Temple,          57500
  DC.                  Manali, 2438
                       32[degrees]14'32"N,
                       77[degrees]11'14"E
Y. tenuifolia        Jispa, Spiti              52771
  (Wild.) Babe.        (H.P), 3200
                       32[degrees]38'0"N,
                       77[degrees]10'0"E

Taxon                  Average plant       Flowering-
                        height (cm)      fruiting period

Hieracium crocatum    42 [+ or -] 6.4    June-September
  Bunge ex Ledeb.

H. umbellatum L.
    (P1)              42 [+ or -] 2.3    June-September

    (P2)              46 [+ or -] 7.2    June-September

L. dissecta D.        16 [+ or -] 4.3    June-September
  Don.

L. dolichophylla      20 [+ or -]5.2     June-September
  Kitamura

L. lessertiana        32 [+ or -] 7.2    June-September
  C. B. Clarke

L. macrorhiza         25 [+ or -] 4.6    June-September
  (Royle) Hook. f.
L. sativa L.          55 [+ or -] 2.4    March-November

L. serriola L.        55 [+ or -] 2.5     April-October

Prenanthes           80 [+ or -] 13.1      June-August
  brunoniana C. B.
  Clarke

Taraxacum
  officinale L.
    (P1)             27.1 [+ or -] 3.2   March-November

    (P2)             8.3 [+ or -] 7.9

    (P3)             17.2 [+ or -] 1.6

Tragopogon dubius
  Scop.
    (P1)              50 [+ or -] 2.5    June-September

    (P2)              35 [+ or -] 7.3    June-September

T. gracilis D. Don   25.2 [+ or -] 4.9   June-September

Youngia glauca        73 [+ or -] 5.3      July-August
  Edgew.

Y. japonica (L.)      42 [+ or -] 4.7    June-September
  DC.

Y. tenuifolia         15 [+ or -] 4.2    June-September
  (Wild.) Babe.

Taxon                Chromosome    Ploidal
                       number       level
                        (2/j)        (x)

Hieracium crocatum   10 ([double     2x
  Bunge ex Ledeb.     dagger])

H. umbellatum L.
    (P1)             10 ([double     2.x
                      dagger])

    (P2)             54 ([double     6.x
                      dagger])

L. dissecta D.           16          2.x
  Don.

L. dolichophylla         16          2.x
  Kitamura

L. lessertiana       16 ([double     2.x
  C. B. Clarke        dagger])

L. macrorhiza            16          2.x
  (Royle) Hook. f.
L. sativa L.             18          2.x

L. serriola L.           18          2.x

Prenanthes               16          2.x
  brunoniana C. B.
  Clarke

Taraxacum
  officinale L.
    (P1)                 16          2.x

    (P2)                 32          4.x

    (P3)                 24          3x

Tragopogon dubius
  Scop.
    (P1)             14 ([double     2.x
                      dagger])

    (P2)             28 ([double     4.x
                      dagger])

T. gracilis D. Don   14 ([double     2.x
                      dagger])

Youngia glauca           14          2.x
  Edgew.

Y. japonica (L.)         16          2.x
  DC.

Y. tenuifolia            10          2.x
  (Wild.) Babe.      [(dagger])

Taxon                 Pollen
                     fertility
                        (%)

Hieracium crocatum     88.4
  Bunge ex Ledeb.

H. umbellatum L.
    (P1)               91.0

    (P2)               82.1

L. dissecta D.         88.1
  Don.

L. dolichophylla       79.2
  Kitamura

L. lessertiana         70.2
  C. B. Clarke


L. macrorhiza          71.3
  (Royle) Hook. f.
L. sativa L.           87.0

L. serriola L.         76.0

Prenanthes             90.1
  brunoniana C. B.
  Clarke

Taraxacum
  officinale L.
    (P1)               86.2

    (P2)               73.6

    (P3)               56.3

Tragopogon dubius
  Scop.
    (P1)               83.0

    (P2)               90.3

T. gracilis D. Don     79.2

Youngia glauca         83.0
  Edgew.

Y. japonica (L.)       78.2
  DC.

Y. tenuifolia          87.2
  (Wild.) Babe.

* Herbarium, Punjabi University, Patiala.
([double dagger]) First ever chromosomal report.
([dagger]) First cytotype report from India.

TABLE 2: Morphological comparison of different cytotypes
of tribe Cichorieae from Western Himalayas.

Taxon/voucher data      Chromosome         Leaf size
                          number      (mean [+ or -] SD)
                          (2/j)

Hieracium umbellatum
PI--52763                   10        4.0 [+ or -] 0.2 x
                                       1.0 [+ or -] 0.3
P2--52774                   54        7.0 [+ or -] 1.2 x
                                       1.5 [+ or -] 0.5
Taraxacum officinale
PI--52687                   16        20.0 [+ or -] 0.6 x
                                        1.9 [+ or -]0.9
P2--52693                   32        5.9 [+ or -] 1.2 x
                                        2.6 [+ or -]0.6
P3--57491                   24        6.9 [+ or -] 0.6 x
                                       1.9 [+ or -] 0.7
Tragopogon dubius
PI--57503                   14         15 [+ or -] 0.5 x
                                       0.6 [+ or -] 0.3
P2--57505                   28         15 [+ or -] 0.5 x
                                       0.9 [+ or -] 0.3

Taxon/voucher data      Shape of    Flower
                          leaf      colour

Hieracium umbellatum
PI--52763                Entire     Yellow

P2--52774               Serrate-    Yellow
                        toothed
Taraxacum officinale
PI--52687                           Yellow

P2--52693                           Yellow

P3--57491                           Purple

Tragopogon dubius
PI--57503                Linear     Yellow

P2--57505                Linear     Yellow
                       lanceolate

Taxon/voucher data      Average    Stomatal size ([micro]m)
                       number of      (mean [+ or -] SD)
                       capitula/
                         plant

Hieracium umbellatum
PI--52763                 17         23.21 [+ or -] 0.4 x
                                      31.54 [+ or -] 2.4
P2--52774                 22         29.80 [+ or -] 2.1 x
                                      34.26 [+ or -] 1.7
Taraxacum officinale
PI--52687                  3         17.6 [+ or -] 0.4 x
                                      14.5 [+ or -] 0.9
P2--52693                  4         18.75 [+ or -] 0.3 x
                                      20.1 [+ or -] 0.4
P3--57491                  6         18.30 [+ or -] 1.3 x
                                      18.98 [+ or -]2.8
Tragopogon dubius
PI--57503                 23         22.19 [+ or -] 0.5 x
                                      27.54 [+ or -] 0.7
P2--57505                 17         18.36 [+ or -] 1.2 x
                                      15.25 [+ or -] 0.8

Taxon/voucher data     Pollen size ([micro]m)   Stomatal
                         (mean [+ or -] SD)     index (%)

Hieracium umbellatum
PI--52763               16.75 [+ or -] 0.7 x      33.33
                         20.1 [+ or -] 1.2
P2--52774               15.0 [+ or -] 0.56 x      23.07
                         20.0 [+ or -] 0.9
Taraxacum officinale
PI--52687               18.75 [+ or -] 0.4 x      28.57
                         20.62 [+ or -] 0.8
P2--52693               18.98 [+ or -] 1.4 x      23.07
                         19.75 [+ or -] 1.6
P3--57491               18.75 [+ or -] 2.4 x      30.0
                         23.0 [+ or -] 2.5
Tragopogon dubius
PI--57503               23.0 [+ or -] 1.3 x       28.5
                         12.0 [+ or -] 0.8
P2--57505               22.75 [+ or -] 0.7 x      16.66
                         20.11 [+ or -] 1.4

* SD: standard deviation.

TABLE 3: Data on cytomixis and meiotic course in the
studied populations of tribe Cichorieae from Western
Himalayas.

Accession                    Cytomixis
number

            PMCs involved     Number      PMCs with
               (% age)       of PMCs     chromosomal
                             involved     stickiness
                               (%)        at M-I (%)

52760        0.8 (10/125)      1-2       3.10 (4/129)
52763        3.96 (8/126)      1-2       5.50 (6/109)
52774       11.71 (13/111)     2-4      11.57 (14/121)
52764        2.4 (3/125)       1-2       4.0 (5/125)
58562        5.4 (6/111)       1-2       3.17 (4/126)
57504            -/-            0        0.82 (1/121)
52776            -/-            0        1.66 (2/120)
57510        2.77 (4/144)      1-2       3.47 (5/144)
58534            -/-            0            -/-
58535         1.0 (1/97)       1-2      11.34 (11/97)
52687        4.35 (5/144)      2-3       5.50 (6/109)
52693        1.78 (2/112)      1-2       7.20 (9/125)
57491       29.91 (35/117)     2-6      16.28 (21/129)
57503        2.43 (3/123)      1-2       2.30 (3/130)
57505        1.72 (2/160)      1-2       1.37 (2/145)
52768            -/-            0            -/-
52773        0.88 (1/113)      1-2       3.53 (4/113)
57500            -/-            0            -/-
57771        1.61 (2/124)      1-2       1.62 (2/123)

Accession                    Meiotic course
number

              PMCs with         PMC with        PMCs with
              unoriented        bridges          laggards
              bivalents         (at A-I,         (at A-I,
              at M-I (%)      II/T-I, II)      II/T-I, II)
                                  (%)              (%)

52760        7.20 (9/125)    10.08 (12/114)   10.56 (13/123)
52763        2.30 (3/130)     4.31 (6/139)     1.55 (2/129)
52774       13.79 (16/118)   26.95 (38/141)   10.44 (14/134)
52764            -/-          4.0 (5/125)          -/-
58562        2.70 (3/111)         -/-              -/-
57504        1.80 (2/111)     4.50 (5/111)     0.82 (1/121)
52776            -/-              -/-              -/-
57510        2.0 (3/144)          -/-              -/-
58534            -/-              -/-              -/-
58535         6.1 (6/97)      7.21 (7/97)      2.06 (2/97)
52687       8.39 (11/131)     3.84 (8/130)    10.08 (12/114)
52693            -/-          3.10 (4/129)     3.84 (8/130)
57491       13.79 (16/116)   12.5 (17/136)    10.44 (14/134)
57503            -/-              -/-              -/-
57505        2.38 (3/126)     1.16 (2/125)     3.25 (4/123)
52768            -/-              -/-              -/-
52773        1.76 (2/113)     5.30 (6/113)     1.76 (2/113)
57500            -/-              -/-              -/-
57771            -/-              -/-              -/-

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

TABLE 4: Data on abnormal microsporogenesis on different
accession of tribe Cichorieae from Western Himalayas.

Taxon/accession   Monads         Dyads
numbers
                     WM (%)         WM (%)        WMN (%)

58533                --/--          --/--       0.97 (1/103)
52760             2.75 (3/109)   0.91 (1/109)   2.75 (3/109)
52763                --/--          --/--       1.80 (2/111)
52774             1.5 (2/130)    0.76 (1/130)   2.30 (3/130)
52764                --/--          --/--       1.6 (2/120)
58562                --/--          --/--          --/--
57504                --/--       0.95 (1/105)      --/--
52776                --/--          --/--       1.66 (2/121)
57510                --/--       0.80 (1/124)   1.61 (2/124)
58534                --/--       1.05 (1/95)       --/--
58535                --/--       1.72 (2/116)   4.31 (5/116)
52687                --/--       1.5 (2/130)    0.76 (1/130)
52593                --/--          --/--       0.8 (1/125)
57491             2.29 (2/87)    4.59 (4/87)    1.14 (1/87)
57503                --/--          --/--          --/--
57505                --/--          --/--       0.8 (1/125)
52768             1.66 (2/121)   4.13 (5/121)   0.82 (1/121)
52773                --/--       0.86 (1/115)   2.60 (3/115)
57500             1.66 (2/121)   0.82 (1/121)        0
52771                --/--          --/--       3.10 (4/129)

Taxon/accession   Triads
numbers
                     WM (%)        WMN (%)

58533                --/--       2.91 (3/103)
52760                --/--       2.75 (3/103)
52763             2.70 (3/111)   1.80 (2/111)
52774             3.07 (4/130)   2.3 (3/130)
52764                --/--          --/--
58562                --/--          --/--
57504             1.90 (2/105)   2.8 (3/105)
52776                --/--       4.13 (5/121)
57510                --/--       4.03 (5/124)
58534                --/--          --/--
58535             2.58 (3/116)   4.31 (5/116)
52687             0.76 (1/130)   2.30 (3/130)
52593             2.4 (3/125)    0.8 (1/125)
57491             2.29 (2/87)    3.44 (3/87)
57503                --/--          --/--
57505                --/--       1.6 (2/125)
52768             4.13 (5/121)   0.82 (1/121)
52773                --/--       2.60 (3/115)
57500             0.82 (1/121)   0.82 (1/121)
52771             0.75 (1/129)   6.97 (9/129)

Taxon/accession   Tetrads
numbers
                      WM (%)           WMN (%)

58533                 --/--        96.11 (99/103)
52760                 --/--        90.82 (99/109)
52763              3.60 (4/111)    90.09 (100/111)
52774              3.8 (5/130)     86.15 (112/130)
52764                 --/--        98.3 (118/120)
58562                 --/--         100 (123/123)
57504              1.90 (2/105)    92.38 (97/105)
52776                 --/--        95.04 (115/121)
57510              1.61 (2/124)    91.93 (114/124)
58534                 --/--        99.2 (139/140)
58535              5.17 (6/116)    81.89 (95/116)
52687              1.5 (2/130)     93.07 (121/130)
52593              1.6 (2/125)     94.4 (118/125)
57491             11.49 (10/87)     72.41 (63/87)
57503                 --/--         100 (121/121)
57505               0.8 (125)      96.8 (121/125)
52768              0.82 (1/121)    87.60 (106/121)
52773              1.73 (2/115)    92.17 (106/115)
57500             13.22 (16/121)   82.64 (100/121)
52771             8.52 (11/129)    80.60 (104/117)

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.
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Title Annotation:Research Article
Author:Gupta, Raghbir Chand; Goyal, Henna; Singh, Vijay; Goel, Rajesh Kumar
Publication:The Scientific World Journal
Article Type:Report
Date:Jan 1, 2014
Words:5515
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