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