Cytological studies of Brassicaceae Burn. (Cruciferae Juss.) from western Himalayas
Цитологические исследования проведены на 12 видах семейства Brassicaceae Burn. из популяций различных географических зон Западных Гималаев. Определены хромосомные числа для Barbaraea intermedia (n = 16), Cardamine loxostemonoides (n = 8), Nasturtium officinale (n = 8), Sisymbrium orientale (n = 14)...
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Інститут клітинної біології та генетичної інженерії НАН України
2013
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| Цитувати: | Cytological studies of Brassicaceae Burn. (Cruciferae Juss.) from western Himalayas / S.M. Jeelani, S. Rani, S. Kumar, S. Kumari, R.C. Gupta // Цитология и генетика. — 2013. — Т. 47, № 1. — С. 26-36. — Бібліогр.: 29 назв. — англ. |
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nasplib_isofts_kiev_ua-123456789-1265182025-02-09T14:00:45Z Cytological studies of Brassicaceae Burn. (Cruciferae Juss.) from western Himalayas Цитологическое изучение семейства Brassicaceae Burn. (Cruciferae Juss.) из западных Гималаев Цитологічне вивчення сімейства Brassicaceae Burn. (Cruciferae Juss.) із західних Гімалаїв Jeelani, S.M. Rani, S. Kumar, S. Kumari, S. Gupta, R.C. Оригинальные работы Цитологические исследования проведены на 12 видах семейства Brassicaceae Burn. из популяций различных географических зон Западных Гималаев. Определены хромосомные числа для Barbaraea intermedia (n = 16), Cardamine loxostemonoides (n = 8), Nasturtium officinale (n = 8), Sisymbrium orientale (n = 14) и добавлены к известным сведениям об этих видах. Хромосомные числа семи видов, Barbaraea intermedia (n = 8), B. vulgaris (n = 8), Capsella bursa-pastoris (n = 8), Descuriania sophia (n = 10), Rorippa islandica (n = 8), Sisymbrium strictum (n = 7) и Thlaspi alpestre (n = 7), были определены впервые в Индии. Течение мейоза в популяциях семи видов (Barbaraea intermedia, Capsella bursa-pastoris, Coronopus didymus, Descuriania sophia, Nasturtium officinale, Sisymbrium orientale и S. strictum) изменяется от нормального до аномального, в то время как в популяциях Barbaraea vulgaris и Sisymbrium irio наблюдается аномальный ход мейоза. Мейотические аномалии проявляются в виде цитомиксиса, слипшихся хромосом, дезориентированных бивалентов, межбивалентных соединений, формировании отставших хромосом и мостов, что в целом приводит к нарушениям микроспорогенеза. Гетерогенные по размеру фертильные пыльцевые зерна и сниженный репродукционный потенциал наблюдались во всех мейотически аномальных популяціях. В то же время ход мейоза во всех популяциях Cardamine loxostemonoides, Rorippa islandica и Thalspi alpestre был нормальным и сопровождался высокой фертильностью пыльцы. Цитологічні дослідження проведені на 12 видах родини Brassicaceae Burn. з популяцій різних географічних зон Західних Гімалаїв. Визначені хромосомні числа для Barbaraea intermedia (n = 16), Cardamine loxostemonoides (n = 8), Nasturtium officinale (n = 8), Sisymbrium orientale (n = 14) і додані до відомостей про ці види. Хромосомні числа семи видів, Barbaraea intermedia (n = 8), B. vulgaris (n = 8), Capsella bursa-pastoris (n = 8), Descuriania sophia (n = 10), Rorippa islandica (n = 8), Sisymbrium strictum (n= 7) та Thlaspi alpestre (n = 7), були визначені вперше в Індії. Проходження мейозу в популяціях семи видів (Barbaraea intermedia, Capsella bursa-pastoris, Coronopus didymus, Descuriania sophia, Nasturtium officinale, Sisymbrium orientale і S. strictum) змінюється від нормального до аномального, тоді як в популяціях Barbaraea vulgaris і Sisymbrium irio спостерігається аномальний хід мейозу. Мейотичні аномалії проявляються у вигляді цитоміксису, злиплих хромосом, дезорієнтованих бівалентів, міжбівалентних сполук, формуванні відсталих хромосом і мостів, що в цілому призводить до порушень мікроспорогенезу. Гетерогенні за розміром фертильні пилкові зерна та знижений репродукційний потенціал спостерігалися в усіх мейотично аномальних популяціях. В той же час хід мейозу в усіх популяціях Cardamine loxostemonoides, Rorippa islandica і Thalspi alpestre був нормальним і супроводжувався високою фертильністю пилку. Cytological studies have been carried out on 12 species of Brassicaceae Burn. on population basis from different geographical areas of Kashmir and Himachal Pradesh in the Western Himalayas. Variable chromosome reports for Barbaraea intermedia (n = 16), Cardamine loxostemonoides (n = 8), Nasturtium officinale (n = 8), Sisymbrium orientale (n = 14) on world-wide basis have been added to the previous reports of these species. The chromosome numbers in seven species as Barbaraea intermedia (n = 8), B. vulgaris (n = 8), Capsella bursa-pastoris (n = 8), Descuriania sophia (n = 10), Rorippa islandica (n = 8), Sisymbrium strictum (n = 7) and Thlaspi alpestre (n = 7) have been worked out for the first time from India. The meiotic course in the populations of seven species such as Barbaraea intermedia, Capsella bursa-pastoris, Coronopus didymus, Descuriania sophia, Nasturtium officinale, Sisymbrium orientale and S. strictum varies from normal to abnormal while all the populations of two species Barbaraea vulgaris and Sisymbrium irio show abnormal meiotic course. Meiotic abnormalities are in the form of cytomixis, chromosomal stickiness, unoriented bivalents, inter-bivalent connections, formation of laggards and bridges, all resulting into abnormal microsporogenesis. Heterogenous sized fertile pollen grains and reduced reproductive potentialities have invariably been observed in all the meiotically abnormal populations. However, the meiotic course in all the populations of Cardamine loxostemonoides, Rorippa islandica and Thalspi alpestre is found to be normal with high pollen fertility. The authors are grateful to the University Grants Commission, New Delhi under the DRS SAP II and ASIST programmes and Rajiv Gandhi National Fellowship Scheme for providing financial assistance. We are highly thankful to the Director Botanical Survey of India, DehraDun for their help in the identification of the plant species. 2013 Article Cytological studies of Brassicaceae Burn. (Cruciferae Juss.) from western Himalayas / S.M. Jeelani, S. Rani, S. Kumar, S. Kumari, R.C. Gupta // Цитология и генетика. — 2013. — Т. 47, № 1. — С. 26-36. — Бібліогр.: 29 назв. — англ. 0564-3783 DOI: 10.3103/S0095452713010076 https://nasplib.isofts.kiev.ua/handle/123456789/126518 en Цитология и генетика application/pdf Інститут клітинної біології та генетичної інженерії НАН України |
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Digital Library of Periodicals of National Academy of Sciences of Ukraine |
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English |
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Оригинальные работы Оригинальные работы |
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Оригинальные работы Оригинальные работы Jeelani, S.M. Rani, S. Kumar, S. Kumari, S. Gupta, R.C. Cytological studies of Brassicaceae Burn. (Cruciferae Juss.) from western Himalayas Цитология и генетика |
| description |
Цитологические исследования проведены на 12 видах семейства Brassicaceae Burn. из популяций различных географических зон Западных Гималаев. Определены хромосомные числа для Barbaraea intermedia (n = 16), Cardamine loxostemonoides (n = 8), Nasturtium officinale (n = 8), Sisymbrium orientale (n = 14) и добавлены к известным сведениям об этих видах. Хромосомные числа семи видов, Barbaraea intermedia (n = 8), B. vulgaris (n = 8), Capsella bursa-pastoris (n = 8), Descuriania sophia (n = 10), Rorippa islandica (n = 8), Sisymbrium strictum (n = 7) и Thlaspi alpestre (n = 7), были определены впервые в Индии. Течение мейоза в популяциях семи видов (Barbaraea intermedia, Capsella bursa-pastoris, Coronopus didymus, Descuriania sophia, Nasturtium officinale, Sisymbrium orientale и S. strictum) изменяется от нормального до аномального, в то время как в популяциях Barbaraea vulgaris и Sisymbrium irio наблюдается аномальный ход мейоза. Мейотические аномалии проявляются в виде цитомиксиса, слипшихся хромосом, дезориентированных бивалентов, межбивалентных соединений, формировании отставших хромосом и мостов, что в целом приводит к нарушениям микроспорогенеза. Гетерогенные по размеру фертильные пыльцевые зерна и сниженный репродукционный потенциал наблюдались во всех мейотически аномальных популяціях. В то же время ход мейоза во всех популяциях Cardamine loxostemonoides, Rorippa islandica и Thalspi alpestre был нормальным и сопровождался высокой фертильностью пыльцы. |
| format |
Article |
| author |
Jeelani, S.M. Rani, S. Kumar, S. Kumari, S. Gupta, R.C. |
| author_facet |
Jeelani, S.M. Rani, S. Kumar, S. Kumari, S. Gupta, R.C. |
| author_sort |
Jeelani, S.M. |
| title |
Cytological studies of Brassicaceae Burn. (Cruciferae Juss.) from western Himalayas |
| title_short |
Cytological studies of Brassicaceae Burn. (Cruciferae Juss.) from western Himalayas |
| title_full |
Cytological studies of Brassicaceae Burn. (Cruciferae Juss.) from western Himalayas |
| title_fullStr |
Cytological studies of Brassicaceae Burn. (Cruciferae Juss.) from western Himalayas |
| title_full_unstemmed |
Cytological studies of Brassicaceae Burn. (Cruciferae Juss.) from western Himalayas |
| title_sort |
cytological studies of brassicaceae burn. (cruciferae juss.) from western himalayas |
| publisher |
Інститут клітинної біології та генетичної інженерії НАН України |
| publishDate |
2013 |
| topic_facet |
Оригинальные работы |
| url |
https://nasplib.isofts.kiev.ua/handle/123456789/126518 |
| citation_txt |
Cytological studies of Brassicaceae Burn. (Cruciferae Juss.) from western Himalayas / S.M. Jeelani, S. Rani, S. Kumar, S. Kumari, R.C. Gupta // Цитология и генетика. — 2013. — Т. 47, № 1. — С. 26-36. — Бібліогр.: 29 назв. — англ. |
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Цитология и генетика |
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26 ISSN 0564–3783. Öèòîëîãèÿ è ãåíåòèêà. 2013. ¹ 1
S.M. JEELANI, S. RANI,
S. KUMAR, S. KUMARI,
R.C. GUPTA
Department of Botany, Punjabi University, Patiala, 147 002 India
E-mail: sandrabi555@gmail.com
CYTOLOGICAL STUDIES OF
BRASSICACEAE BURN. (CRUCIFERAE
JUSS.) FROM WESTERN HIMALAYAS
Cytological studies have been carried out on 12 species
of Brassicaceae Burn. on population basis from different
geographical areas of Kashmir and Himachal Pradesh in
the Western Himalayas. Variable chromosome reports for
Barbaraea intermedia (n = 16), Cardamine loxostemonoides
(n = 8), Nasturtium officinale (n = 8), Sisymbrium orientale
(n = 14) on world-wide basis have been added to the
previous reports of these species. The chromosome numbers
in seven species as Barbaraea intermedia (n = 8), B. vulgaris
(n = 8), Capsella bursa-pastoris (n = 8), Descuriania
sophia (n = 10), Rorippa islandica (n = 8), Sisymbrium
strictum (n = 7) and Thlaspi alpestre (n = 7) have been
worked out for the first time from India. The meiotic course
in the populations of seven species such as Barbaraea in-
termedia, Capsella bursa-pastoris, Coronopus didymus,
Descuriania sophia, Nasturtium officinale, Sisymbrium
orientale and S. strictum varies from normal to abnormal
while all the populations of two species Barbaraea vulgaris
and Sisymbrium irio show abnormal meiotic course.
Meiotic abnormalities are in the form of cytomixis, chro-
mosomal stickiness, unoriented bivalents, inter-bivalent
connections, formation of laggards and bridges, all resulting
into abnormal microsporogenesis. Heterogenous sized fertile
pollen grains and reduced reproductive potentialities have
invariably been observed in all the meiotically abnormal
populations. However, the meiotic course in all the popu-
lations of Cardamine loxostemonoides, Rorippa islandica
and Thalspi alpestre is found to be normal with high pollen
fertility.
Introduction. The family Brassicaceae includes
332 genera and 3,709 species [1] widely distributed
throughout the world but abundant in north
temperate zone with Mediterranean as major
centre. In Indian subcontinent, 61 genera and 168
species [2] are taxonomically known which are
mostly found in temperate and alpine Himalayas.
Starting with the first comprehensive cytological
studies [3–5], at present more than 9.000 chro-
mosomal counts are known for 232 (68.6 %)
genera and 1.558 (42.0 %) species of the family
all over the world [1]. The chromosome numbers
in the family vary from n = 4 in Stenoptalum
nutans from Australia to n = 128 in Cardamine
concatenate and C. diphylla from eastern North
America [1]. From the Indian subcontinent, the
chromosomal counts are known for nearly half
the number of species of the family i.e. 90 species
belonging to 41 genera. It is noted that cytological
evaluation of Brassicaceae has got poor attention
from Western Himalayas, in general, and for the
present studied areas in particular. As the family
is well documented with natural allopolyploidy,
dysploid aneuploid series and mixoploidy coupled
with economic importance, making it very good
collection of plant material to be studied further
from cytological perspective. To make additions
to the chromosomal database and to study the
genetic diversity at intra- and interspecific levels,
concerning lesser known angiospermic families,
from newer here as areas the present meiotic
studies on the aforesaid taxa of Brassicaceae have
been carried out.
Materials and methods. For meiotic studies,
flower buds were collected from different lo-
calities of selected areas of Western Himalayas
(Table 1). Smears of appropriate sized flower
buds were made after fixing in Carnoy’s fixative,
using standard acetocarmine technique. Pollen
fertility was estimated by mounting mature pollen
grains in glycero-acetocarmine (1:1) mixture or
1 % aniline blue. Well-filled pollen grains with
stained nuclei were taken as apparently fertile,
while shrivelled and unstained pollen grains were
counted as sterile. Photomicrographs of pollen
mother cells and pollen grains were made from
freshly prepared slides using Nikon 80i eclipse
Digital Imaging System. Voucher specimens are
deposited in the Herba rium, Department of
Botany, Punjabi University, Patiala (PUN).
Results. Detailed meiotic studies have been
carried out on 29 populations belonging to 12 © S.M. JEELANI, S. RANI, S. KUMAR, S. KUMARI,
R.C. GUPTA, 2013
27
Cytological studies of Brassicaceae Burn. (Cruciferae Juss.)
ISSN 0564–3783. Öèòîëîãèÿ è ãåíåòèêà. 2013. ¹ 1
Table 1
Data showing location, altitude, accession number, present chromosome number, ploidy level, meiotic course
and pollen fertility in different taxa of family Brassicaceae Burn. from Western Himalayas
Taxa
Locality/Altitude (m)/
Accession Number
Present
Chromosome
Number (n)/
Figure Number
Ploidy
level
Meiotic
course**/
Pollen
fertility,
(%)
Barbaraea intermedia Boreau
B. vulgaris R.Br.
Cardimine loxostemonoides
O.E.
Capsella bursa-pastoris (L.)
Medik
Coronopus didymus (L.)
Smith
Descuriainia sophia Webb ex
Prantl.
Nasturtium officinale R.Br.
Rorippa islandica (Oeder)
Berbas
Sisymbrium irio L.
S. orientale L.
Ferozpur Nallah Distt. Baramulla
(Kashmir)/2,400 /54270
Triund, Distt. Kangra N/(H.P*.)/3,000/52744
Chhota-Bhangal, Distt. Kangra
(H.P.)/2,300/52796
Ferozpur Nallah, Distt.Baramullah
(Kashmir)/2,200/52473
Ratnipora, Distt. Pulwama (Kashmir)/
1,750/52472
Lower Munda, Distt. Anantnag (Kashmir)/
2,300/54193
Ratnipora, Distt. Pulwama
(Kashmir)/1,750/54192
Chhota-Bhangal, Distt. Kangra
(H.P.)/2,300/52722
Salai, Distt. Kangra
(H.P.)/1,600/52733
Shillai, Distt. Sirmaur
(H.P.)/1,800/52653
Triund, Distt. Kangra (H.P.)/3,000/52703
Dharmkot, Distt. Kangra (H.P.)/2,800/
52677
Yusmarg, Distt. Budgam
(Kashmir)/2,600/54202
Pehalgam, Distt. Anantnag (Kashmir)/
2,300/54201
Ferozpur Nallah, Distt. Baramullah
(Kashmir)/2,100/54205
Ranhear, Distt. Kangra (H.P.)/800/52736
Nahan, Distt. Sirmaur (H.P.)/900/55242
Keller, Distt. Shopian (Kashmir)/2,000/
54827
Aharbal, Distt. Kulgam (Kashmir)/2,100/
54271
Saketi, Distt. Sirmaur (H.P.)/800/55241
Renukaji, Distt. Sirmaur (H.P.)/650/54234
Keller, Distt. Shopian (Kashmir)/2,200/
54235
Chhota-Bhangal, Distt. Kangra (H.P.)/
2,300/52741
Triund, Distt. Kangra (H.P.)/3,000/52740
Churdhar, Distt. Sirmaur (H.P.)/3,630/
54206
8 (Fig. 1, a)
16 (Fig. 1, b)
8 (Fig. 1, c)
8 (Fig. 1, d)
8
16 (Fig. 1, e)
16
8 (Fig. 1, f)
16
16
16 (Fig.1, g)
16
10 (Fig. 1, h)
10
8
8 (Fig. 1, i)
8
8 (Fig. 1, j)
14 (Fig. 1, k)
7 (Fig. 1, l)
14
14 (Fig 1, m)
8
8
8
2x
4x
2x
2x
2x
4x
4x
2x
4x
4x
4x
4x
2x
2x
2x
2x
2x
2x
4x
2x
4x
4x
2x
2x
2x
A/64. 42
N/96.50
A/60.78
N/97.55
N/83.87
A/69.01
A/62.42
A/60.00
N/97.98
N/98.65
A/65.78
N/94.54
A/63.45
N/88.78
N/92.65
A/65.50
A/61.45
N/92.00
A/63.47
A/60.87
A/66.94
A/65.76
N/96.45
A/62.00
A/60.50
28 ISSN 0564–3783. Öèòîëîãèÿ è ãåíåòèêà. 2013. ¹ 1
S.M. Jeelani, S. Rani, S. Kumar et al.
Taxa
Locality/Altitude (m)/
Accession Number
Present
Chromosome
Number
(n)/Figure
Number
Ploidy
level
Meiotic
course**/
Pollen
fertility,
(%)
S. strictum L.
Thlaspi alpestre L.
Batnoor Tral, Distt. Pulwama (Kashmir)/2,300/
52475
Chhota-Bangal, Distt. Kangra (H.P.)/2,300/
52704
Nauradhar, Distt. Sirmaur (H.P.)/1,800/54209
Baragran Distt. Kangra (H.P.)/3,000/52742
7 (Fig 1, n)
7
7
7 (Fig. 1, o)
2x
2x
2x
2x
A/69.78
A/65.75
N/96.98
N/93.50
Continuation of table 1
* H.P. = Himachal Pradesh, ** N = Normal, ** A = Abnormal.
Taxa/Accessions
Cytomixis Meiotic
% age of PMCs involved
Meiosis-I/Meiosis-II
Number of PMCs
involved
Chromosomal stickiness
at M-I (%)
Barbaraea intermedia
54270
52796
B. vulgaris
52796
Capsella bursa-pastoris
54193
54192
52722
Coronopus didymus
52703
Descuriainia sophia
54202
Nasturtium officinale
55736
55242
Sisymbrium irio
54271
55241
54234
S. orientale
54489
52740
S. strictum
52475
52704
13.86(14/101)/5.71(6/105)
/
9.73(11/113)/8.04(7.87)
20.83(25/120)/15.38(20/130)
7.14(7/98)/4.08(4/98)
/
5.21(6/115)/6.83(8/117)
5.21(6/115)/6.42(7/109)
/
7.33(8/109)/5.08(6/118)
/
5.94(6/101)
/
6.95(8/115)/5.00(6/120)
/
11.53(15/130)/8.00(10/125)
12.87(17/132)/12.59(16/127)
2 3
2 3
2 6
2 3
2 3
2 3
2 3
2 3
2 3
2 3
2 3
6.06(6/99)
1.98(2/101)
13.91(16/115)
5.55(4/72)
3.38(4/118)
3.73(4/107)
1.81(2/110)
4.65(6/129)
6.15(8/130)
2.04(2/98)
3.05(4/131)
5.45(6/110)
Data on cytomixis, meiotic course and pollen size in different
Figures in parenthesis denote observed number of abnormal PMCs in the numerator and total number of PMCs
29
Cytological studies of Brassicaceae Burn. (Cruciferae Juss.)
ISSN 0564–3783. Öèòîëîãèÿ è ãåíåòèêà. 2013. ¹ 1
species of 9 genera of the family Brassicaceae
from different localities with altitudinal range of
800–3,000 m from two districts of Himachal
Pradesh and 2,000–2,600 m from six districts of
Kashmir. The data regarding locality with alti-
tude, accession number, present chromosome
numbers with figure numbers, ploidy level,
nature of meiotic course and pollen fertility of
the presently worked out populations has been
presented in Table 1.
Two populations of Barbaraea intermedia,
one each from Kashmir and Himachal Pra-
desh shows intraspecific variability in the form
of 2x (n = 8) and 4x (n = 16) cytotypes, res-
pectively. Single population of B. vulgaris col-
lected from Himachal Pradesh and two po-
pulations of Cardamine loxostemonoides col-
lected from Kashmir represent 2x cytotypes,
all exhibiting n = 8. Capsella bursa-pastoris is
another species depicting intraspecific variabi-
lity as one population from Himachal Pradesh
shows 2x cytotype (n = 8) whereas two
populations from Kashmir and two populations
from Himachal Pradesh show 4x cytotype (n =
= 16) pointing wide spread occurrence of tet-
raploids of this species in the Western Himalayas.
Both the populations of Coronopus didymus,
studied from Himachal Pradesh have the tet-
Table 2
species of Brassicaceae Burn. from Western Himalayas
course showing PMCs with
Size ( m) of fertile pollen grains
(Range)Unoriented bivalents
at M-I (%)
Bridges at Meiosis-I/Meiosis-II
(%)
Laggards at Meiosis-I/ Meiosis-II
(%)
6.45(6/93)
3.48(4/115)
12.50(15/120)
5.55(4/72)
2.63(3/114)
1.85(2/108)
1.76(2/113)
6.40(8/125)
5.08(6/118)
4.04(4/99)
4.65(6/129)
3.14(4/127)
7.23(11/152)/8.92(10/112)
14.28(15/105)/9.52(10/105)
8.84(10/113)/7.50(9/120)
13.04(15/115)/10.00(12/120)
9.67(6/62)/12.90(8/62)
10.00(5/50)/8.00(4/50)
4.80(6/125)/4.13(5/121)
6.25(7/112)/1.53(2/130)
4,16(4/96)/5/21(6/115)
4.87(6/123)/
6.95(8/115)/5.00(6/120)
6.36(7/110)/6.08(7/115)
6.83(8/117)/
6.89(8/116)/3.36(4/119)
7.47(8/107)/5.12(6/117)
5.98(7/117)/5.21(6/115)
4.95(6/121)/4.20(5/119)
8.86(7/79)/11.22(11/98)
11.76(12/102)/8.84(10/113)
9.23(12/130)/
14.61(19/130)/15.00(18/120)
11.29(7/62)/8.06(5/62)
11.11(6/54)/9.25(5/54)
6.25(8/128)/5.88(7/119)
4.87(6/123)/1.73(2/115)
5.04(6/119)/3.60(4/111)
6.95(8/115)/
7.20(9/125)/5/60(7/125)
8.00(10/125)/9/91(12/121)
5.21(6/115)/6/66(8/120)
6.66(8/120)/7.69(9/117)
13.60(17/125)/12.30(16/130)
9.30(12/129)/7.63(10/131)
5.00(6/120)/5.78(7/111)
15.08×13.00 15.16×13.08
19.96×18.47 16.45×16.20
18.18×17.05 17.24×17.10
14.17×13.35 16.25×15.45
18.83×17.24 16.95×16.05
15.78×14.00 15.17×13.14
16.78×14.00 15.26×13.87
18.78×16.00 15.05×17.90
16.75×15.90 14.56×13.26
observed in denominator.
30 ISSN 0564–3783. Öèòîëîãèÿ è ãåíåòèêà. 2013. ¹ 1
S.M. Jeelani, S. Rani, S. Kumar et al.
Fig. 1. Barbaraea intermedia (a) – PMC at A-I (n = 8); B. intermedia (b) – PMC at A-I (n = 16); B. vulgaris (c) –
PMC at A-I (n = 8); Cardimine loxostemonoides (d) – PMC at M-I (n = 8); Capsella bursa-pastoris (e) – PMC
at M-I (n = 16); Capsella bursa-pastoris (f) – PMC at M-I (n = 8); Coronopus didymus (g) – PMC at M-II
(n = 16); Descuriania sophia (h) – PMC at A-I (n = 10); Nasturtium officinale (i) – PMC at A-I (n = 8); Ror-
ippa islandica (j) – PMC at A-I (n = 8); Sisymbrium irio (k) – PMC at M-II (n = 14); S. irio (l) – PMC at A-I
(n = 7); S. orientale (m) – PMC at Diakinesis (n = 14); S. strictum (n) – PMC at M-I (n = 7), Thlaspi alpes-
tre (o) – PMC at M-I (n = 7). Bar = 10 m
31ISSN 0564–3783. Öèòîëîãèÿ è ãåíåòèêà. 2013. ¹ 1
Cytological studies of Brassicaceae Burn. (Cruciferae Juss.)
Fig. 2. Transfer of chromatin (arrowed) in pollen
mother cells (PMCs) at different stages of meiosis with
one empty (double headed arrow) PMC (a, b); PMC
showing chromatin stickiness at metaphase I (c); PMC
showing unoriented bivalent at M-I (d); PMC showing inter-bivalent connections at M-I (e); PMC with multiple
bridges (arrowed) at anaphase I (f); PMC showing bridge (arrowed) at telophase I (g); PMC with laggards (ar-
rowed) at anaphase I (h); PMCs showing laggards (arrowed) at telophase I (i–k); PMC with six poles at telophase
II (l); Diad (m); Triad (n); Tetrad with micronuclei (arrowed) (o, p); Fertile pollen grains differing in size (q);
Fertile and sterile pollen grains (r). Bar = 10 m
32 ISSN 0564–3783. Öèòîëîãèÿ è ãåíåòèêà. 2013. ¹ 1
S.M. Jeelani, S. Rani, S. Kumar et al.
raploid cytotypes (n = 16). Two populations of
Descuriainia sophia have been collected from
Kashmir exhibiting meiotic chromosome number
n = 10 and diploid status based on x = 10. In
case of Nasturtium officinale, one population from
Kashmir and two populations from different parts
of Himachal Pradesh, all depict 2x cytotypes
with n = 8. Likewise Rorippa islandica based on
single population studied from Kashmir is also 2x
(n = 8). For Sisymbrium irio one population form
district Kangra of Himachal Pradesh is reported
to be 2x (n = 7) whereas two other populations,
one from Kashmir and other from district
Sirmaur of Himachal Pradesh on the contrary
are noted to be 4x (n = 14). S. orientale makes
interesting case where three populations studied
from Himachal Pradesh represent only 2x (n =
8), but a single population studied from Kashmir
is found to be 4x based on x = 7. In case of
S. strictum, one population from Kashmir and
two from Himachal Pradesh as well as for Thalspi
alpestre, a single population from Himachal
Pradesh, all unequivocally represent 2x cytotypes
strictly with n = 7.
The meiotic abnormalities have been recorded
in some populations of Barbaraea intermedia,
Capsella bursa-pastoris, Coronopus didymus, Des-
curiania sophia, Nasturtium officinale, Sisymbrium
orientale and S. strictum and in all the popula-
tions of Barbaraea vulgaris and Sisymbrium irio.
In such populations, the meiotic abnormalities
in the form of cytomixis, chromatin stickiness,
unoriented bivalents, inter-bivalent connections,
bridges, laggards or multipolar spindle have
been observed at different stages of meiosis
(Table 2; Fig. 2, g–k). Regarding cytomixis,
the chromatin transfer from early prophase
to pollen grain formation has been observed
in most of these populations, with the highest
percentage recorded in one population of
Capsella bursa-pastoris involving 2–6 PMCs
(Table 2). Cytomixis in these species results
into the production of hypo- and hyperploid
PMCs and even empty PMCs (Fig. 2, b).
Chromatin stickiness involving few bivalents
or whole complement is seen from prophase-I
to metaphase-I (Fig. 2, c). The unoriented
bivalents at metaphase-I (Fig. 2, d) have also
been observed with highest frequency noted in
one population of Capsella bursa-pastoris (Table
2). Interbivalent connections have been observed
at M-I in one of the populations of Capsella
bursa-pastoris from Chotta Bhangal (Fig. 2, e).
These meiotically abnormal populations quite
frequently show the presence of chromosomal
laggards and bridges at anaphases and telophases
(Fig. 2, f–k). Instead of regular spindle
formation, abnormal split spindles and three
poles per PMC at T-I and even six poles at T-II
(Fig. 2, l) have been seen in significant number
of PMCs in Capsella bursa-pastoris, Des-
curiania sophia, Nasturtium officinale and Sisym-
Data on abnormal
Taxa/Accessions Monads
WMN WM
Barbaraea intermedia
54270
52796
Barbaraea vulgaris
52796
Capsella bursa-pastoris
54193
54192
52722
Coronopus didymus
52703
Descuriainia Sophia
54202
Nasturtium officinale
55136
55242
Sisymbrium irio
54271
54241
54234
S. orientale
54489
52740
S. strictum
52475
52704
2.06(2/97)
5.60(6/107)
2.00(2/100)
1.98(2/101)
4.76(6/126)
4.31(6/139)
1.96 (2/102)
1.04(1/96)
1.90(2/105)
0.86(1/115)
1.96(2/102)
0.98(1/102)
0.97(1/103)
1.83(2/109)
2.88(3/104)
1.03(1/97)
2.00(2/100)
0.99(1/101)
2.38(3/126)
0.71(1/139)
0.98(1/102)
1.04(1/96)
0.95(1/105)
1.73(2/115)
0.98(1/102)
0.91(1/109)
1.92(2/104)
Indication. WMN = without micronuclei; WM = with
ber of PMCs observed in denominator.
33ISSN 0564–3783. Öèòîëîãèÿ è ãåíåòèêà. 2013. ¹ 1
Cytological studies of Brassicaceae Burn. (Cruciferae Juss.)
brium orientale. This results into abnormal micro-
sporogenesis leading to the formation of monads,
diads, triads or polyads, along with micronuclei
(Fig. 2, m–p; Table 3) and leads to the formation
of heterogeneous sized fertile pollen grains with
reduced pollen fertility (Fig. 2, q, r). The pollen
fertility in the populations with normal meiosis
has been recorded to be nearly cent per cent.
Discussion. As evident from the great amount
of variation reported in chromosome numbers of
Brassicaceae in literature on world-wide basis, it
is clear that family Brassicaceae represents range
of genera depicting mono-, di- and polybasic
nature. Some authors have suggested the base
numbers to be considered from x = 4–13,
considering even some higher numbers as 32, 42,
etc. to be taken as base numbers, the later most
probably represent single and isolated gametic
number report for any species of the genus [1].
On the other hand some authors have reported
only x = 5–8 as primary base numbers [6]. Both
the statements reveal unanimously that x = 8
is the most popular base number. Analysis of
overall chromosome number data proves that x
Microsporogenesis, %
Diads Triads Tetrads
WMN WM WMN WM WMN WM
7.21(7/97)
1.86(2/107)
4.00(4/100)
1.98(2/101)
6.34(8/126)
1.43(2/139)
1.96(2/102)
1.04(1/96)
1.94(2/103)
2.85(3/105)
1.73(2/115)
3.19(3/94)
2.94(3/102)
0.97(1/103)
1.83(2/109)
0.96(1/104)
2.06(2/97)
1.86(2/107)
2.00(2/100)
0.99(1/101)
3.17(4/126)
5.03(7/139)
2.94(3/102)
1.04(1/96)
0.95(1/105)
2.60(3/115)
1.06(1/94)
1.96(2/102)
0.98(1/102)
1.83(2/109)
1.92(2/104)
4.12(4/97)
6.54(7/107)
5.00(5/100)
0.99(1/101)
5.55(7/126)
11.51(16/139)
1.96(2/102)
2.08(2/96)
3.88(4/103)
1.90(2/105)
1.06(1/94)
1.96(2/102)
3.92(4/102
1.94(2/103)
3.66(4/109)
0.96(1/104)
2.06(2/97)
0.93(1/107)
2.00(2/100)
3.96(4/101)
3.17(4/126)
6.47(9/139)
0.98(1/102)
2.08(2/96)
1.90(2/105)
1.73(2/115)
2.12(2/94)
0.98(1/102)
0.97(1/103)
2.75(3/109)
1.92(2/104)
69.07(67/97)
65.42(70/107)
62.00(62/100)
67.32(68/101)
57.14(72/126)
52.51(73/139)
67.64(69/102)
69.79(67/96)
75.72(78/103)
68.57(72/105)
67.82(78/115)
71.27(67/94)
67.64(69/102)
70.58(72/102)
72.81(75/103)
64.22(70/109)
65.38(68/104)
12.37(12/97)
17.75(19/107)
21.00(21.00/100)
21.78(22/101)
17.46(22/126)
17.98(25/139)
21.56(22/102)
21.87(21/96)
18.44(19/103)
20.95(22/105)
21.73(25/115)
21.27(20/94)
21.56(22/102)
23.52 (24/102)
22.33(23/103)
22.93(25/109)
25.00(26/104)
Table 3
microsporogenesis in different species of Brassicaceae Burn. from different areas of Western Himalayas
micronuclei. Figures in parenthesis denote observed number of abnormal PMCs in the numerator and total num-
34 ISSN 0564–3783. Öèòîëîãèÿ è ãåíåòèêà. 2013. ¹ 1
S.M. Jeelani, S. Rani, S. Kumar et al.
= 8 is represented nearly by 20 % of taxa in
the family followed by x = 7 being present in
19.13 % taxa [1]. Another frequent base number
x = 10 is found in 10.24 % taxa and seems to
be secondarily derived base number. The same
holds true for Indian representatives in general
and presently investigated taxa from Western
Himalayas in particular.
The meiotic chromosome numbers for Barba-
raea intermedia (n = 8), B. vulgaris (n = 8), Des-
curiania sophia (n = 10), Rorippa islandica (n = 8),
Sisymbrium strictum (n = 7) and Thlaspi alpestre
(n = 7) have been worked out for first time from
India confirming the same chromosome number
for these species from outside India. But at the
same time other interaspecific cytotypes are also
known in literature on world-wide basis for some
of these species to be taken into account such
as Barbaraea vulgaris 2n = 14 [7], Descuriania
sophia 2n = 14, 28, 56 [5, 8, 9], Rorippa islandica
2n = 32 [10]. Out of the four species where new
and varied cytotypes have been reported for the
first time, Barbaraea intermedia (n = 16) and
Sisymbrium orientale (n = 14) make an addition
of tetraploid cytotypes on world-wide basis. In
case of Cardamine loxostiemonoides present di-
ploid report (n = 8) against tetraploid cytotype
2n = 32 [11] and for Nasturtium officinale also
present diploid report (n = 8) against another
diploid cytotype with 2n = 14 [12], tetraploid
(2n = 32) and octaploid (2n = 64) cytotypes
[5] are recorded on world-wide basis. The
diploid cytotype (n = 8) in Capsella bursa-
pastoris is reported for the first time from
India and confirms the previous reports of
2n = 32 [13–15] from outside India. Sisymbrium
irio is another species with presently worked
out diploid and tetraploid cytotypes like those
of Barbaraea intermedia and Capsella bursa-
pastoris. Although Sisymbrium irio is one species
with high amount of intraspecific variability
existing in the form of 2n = 14, 21, 28, 42,
56 numbers showing diploid to octaploid
cytotypes collected from the same region of
India [16] but in spite of wide screening at
present only diploid and tetraploid cytotypes
could be found. In Coronopus didymus present
tetraploid chromosome number count of n =
16 is in line with the earlier report of 2n =
32 from India [17] and from outside India [4,
5, 18]. From this entire account one thing
emerges out that there exists a lot of variation
of chromosome numbers in the species of the
family Brassicaceae. Allopolyploidy coupled
with hybridization resulting from different
base number seems be major evolutionary fac-
tor for generating higher chromosome numbe-
red taxa as postulated earlier [19]. The
phenomenon of mixoploidy has also been
described for certain members of Brassicaceae
[20], and explained that mostly gamete
formation in such plants is not affected and show
behaviour of normal diploids, but sometimes
as an aberration some of the aneuploid and
polyploidy cells may enter the germline to give
rise to dysploid and polyploidy plants especially
in apomicts and spontaneous hybrids.
The meiotic abnormalities are considered
to be resulting from genetic [21–23] and
environmental factors [24] as well as geno-
mic-environmental interaction [25]. Chroma-
tin transfer, production of hypo- and hyper-
ploid PMCs and meiotic irregularities in ana-
phase segregation of chromosomes may be
the possible mechanisms for the formation
of large sized pollen grains and low pollen
fertility as has been earlier reported in se-
veral angiospermic plants [26–29]. All these
factors seem to be equally applicable in
the presently investigated populations with
meiotic irregularities in the form of cytomixis,
chromosomal stickiness, unoriented bivalents,
interbivalent connections, laggards and bridges.
The occurrence of sufficient variation of chro-
mosome numbers as well as change of meiotic
behavior at intra- and interspecific level of
these 12 species calls for further need of the
extensive cytological exploration at population
basis of members of Brassicaceae from different
phytogeographical areas of India to complete
the data bases for Indian Brassicaceae.
The authors are grateful to the University
Grants Commission, New Delhi under the DRS
SAP II and ASIST programmes and Rajiv Gandhi
National Fellowship Scheme for providing fi-
nancial assistance. We are highly thankful to the
Director Botanical Survey of India, DehraDun for
their help in the identification of the plant species.
35ISSN 0564–3783. Öèòîëîãèÿ è ãåíåòèêà. 2013. ¹ 1
Cytological studies of Brassicaceae Burn. (Cruciferae Juss.)
S.M. Jeelani, S. Rani, S. Kumar, S. Kumari,
R.C. Gupta
ÖÈÒÎËÎÃÈ×ÅÑÊÎÅ ÈÇÓ×ÅÍÈÅ
ÑÅÌÅÉÑÒÂÀ BRASSICACEAE BURN.
(CRUCIFERAE JUSS.) ÈÇ ÇÀÏÀÄÍÛÕ ÃÈÌÀËÀÅÂ
Öèòîëîãè÷åñêèå èññëåäîâàíèÿ ïðîâåäåíû íà
12 âèäàõ ñåìåéñòâà Brassicaceae Burn. èç ïîïóëÿ-
öèé ðàçëè÷íûõ ãåîãðàôè÷åñêèõ çîí Çàïàäíûõ Ãè-
ìàëàåâ. Îïðåäåëåíû õðîìîñîìíûå ÷èñëà äëÿ Barb-
araea intermedia (n = 16), Cardamine loxostemonoides
(n = 8), Nasturtium officinale (n = 8), Sisymbrium
orientale (n = 14) è äîáàâëåíû ê èçâåñòíûì ñâå-
äåíèÿì îá ýòèõ âèäàõ. Õðîìîñîìíûå ÷èñëà ñåìè
âèäîâ, Barbaraea intermedia (n = 8), B. vulgaris (n =
= 8), Capsella bursa-pastoris (n = 8), Descuriania so-
phia (n = 10), Rorippa islandica (n = 8), Sisymbri-
um strictum (n = 7) è Thlaspi alpestre (n = 7), áûëè
îïðåäåëåíû âïåðâûå â Èíäèè. Òå÷åíèå ìåéîçà â
ïîïóëÿöèÿõ ñåìè âèäîâ (Barbaraea intermedia, Cap-
sella bursa-pastoris, Coronopus didymus, Descuriania
sophia, Nasturtium officinale, Sisymbrium orientale è
S. strictum) èçìåíÿåòñÿ îò íîðìàëüíîãî äî àíî-
ìàëüíîãî, â òî âðåìÿ êàê â ïîïóëÿöèÿõ Barbaraea
vulgaris è Sisymbrium irio íàáëþäàåòñÿ àíîìàëüíûé
õîä ìåéîçà. Ìåéîòè÷åñêèå àíîìàëèè ïðîÿâëÿþòñÿ
â âèäå öèòîìèêñèñà, ñëèïøèõñÿ õðîìîñîì, äåçî-
ðèåíòèðîâàííûõ áèâàëåíòîâ, ìåæáèâàëåíòíûõ ñîå-
äèíåíèé, ôîðìèðîâàíèè îòñòàâøèõ õðîìîñîì è
ìîñòîâ, ÷òî â öåëîì ïðèâîäèò ê íàðóøåíèÿì
ìèêðîñïîðîãåíåçà. Ãåòåðîãåííûå ïî ðàçìåðó ôåð-
òèëüíûå ïûëüöåâûå çåðíà è ñíèæåííûé ðåïðî-
äóêöèîííûé ïîòåíöèàë íàáëþäàëèñü âî âñåõ ìåé-
îòè÷åñêè àíîìàëüíûõ ïîïóëÿöèÿõ.  òî æå âðåìÿ
õîä ìåéîçà âî âñåõ ïîïóëÿöèÿõ Cardamine loxos-
temonoides, Rorippa islandica è Thalspi alpestre áûë
íîðìàëüíûì è ñîïðîâîæäàëñÿ âûñîêîé ôåðòèëü-
íîñòüþ ïûëüöû.
S.M. Jeelani, S. Rani, S. Kumar, S. Kumari,
R.C. Gupta
ÖÈÒÎËÎò×ÍÅ ÂÈÂ×ÅÍÍß Ñ²ÌÅÉÑÒÂÀ
BRASSICACEAE BURN. (CRUCIFERAE JUSS.) ²Ç
ÇÀÕ²ÄÍÈÕ Ã²ÌÀËÀ¯Â
Öèòîëîã³÷í³ äîñë³äæåííÿ ïðîâåäåí³ íà 12 âè-
äàõ ðîäèíè Brassicaceae Burn. ç ïîïóëÿö³é ð³ç-
íèõ ãåîãðàô³÷íèõ çîí Çàõ³äíèõ óìàëà¿â. Âèçíà÷åí³
õðîìîñîìí³ ÷èñëà äëÿ Barbaraea intermedia (n = 16),
Cardamine loxostemonoides (n = 8), Nasturtium offici-
nale (n = 8), Sisymbrium orientale (n = 14) ³ äîäàí³
äî â³äîìîñòåé ïðî ö³ âèäè. Õðîìîñîìí³ ÷èñëà ñåìè
âèä³â, Barbaraea intermedia (n = 8), B. vulgaris (n = 8),
Capsella bursa-pastoris (n = 8), Descuriania sophia (n =
= 10), Rorippa islandica (n = 8), Sisymbrium strictum (n=
= 7) òà Thlaspi alpestre (n = 7), áóëè âèçíà÷åí³ âïåðøå
â ²í䳿. Ïðîõîäæåííÿ ìåéîçó â ïîïóëÿö³ÿõ ñåìè
âèä³â (Barbaraea intermedia, Capsella bursa-pastoris,
Coronopus didymus, Descuriania sophia, Nasturtium
officinale, Sisymbrium orientale ³ S. strictum) çì³íþº-
òüñÿ â³ä íîðìàëüíîãî äî àíîìàëüíîãî, òîä³ ÿê â
ïîïóëÿö³ÿõ Barbaraea vulgaris ³ Sisymbrium irio ñïîñ-
òåð³ãàºòüñÿ àíîìàëüíèé õ³ä ìåéîçó. Ìåéîòè÷í³
àíîìà볿 ïðîÿâëÿþòüñÿ ó âèãëÿä³ öèòîì³êñèñó, çëèï-
ëèõ õðîìîñîì, äåçîð³ºíòîâàíèõ á³âàëåíò³â, ì³æ-
á³âàëåíòíèõ ñïîëóê, ôîðìóâàíí³ â³äñòàëèõ õðîìî-
ñîì ³ ìîñò³â, ùî â ö³ëîìó ïðèçâîäèòü äî ïî-
ðóøåíü ì³êðîñïîðîãåíåçó. Ãåòåðîãåíí³ çà ðîçì³-
ðîì ôåðòèëüí³ ïèëêîâ³ çåðíà òà çíèæåíèé ðåïðî-
äóêö³éíèé ïîòåíö³àë ñïîñòåð³ãàëèñÿ â óñ³õ ìåéî-
òè÷íî àíîìàëüíèõ ïîïóëÿö³ÿõ.  òîé æå ÷àñ õ³ä
ìåéîçó â óñ³õ ïîïóëÿö³ÿõ Cardamine loxostemonoides,
Rorippa islandica ³ Thalspi alpestre áóâ íîðìàëüíèì
³ ñóïðîâîäæóâàâñÿ âèñîêîþ ôåðòèëüí³ñòþ ïèëêó.
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Received 26.10.11I
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/ENU (Use these settings to create Adobe PDF documents best suited for high-quality prepress printing. Created PDF documents can be opened with Acrobat and Adobe Reader 5.0 and later.)
>>
/Namespace [
(Adobe)
(Common)
(1.0)
]
/OtherNamespaces [
<<
/AsReaderSpreads false
/CropImagesToFrames true
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(InDesign)
(4.0)
]
/OmitPlacedBitmaps false
/OmitPlacedEPS false
/OmitPlacedPDF false
/SimulateOverprint /Legacy
>>
<<
/AddBleedMarks false
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/ConvertColors /ConvertToCMYK
/DestinationProfileName ()
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/Downsample16BitImages true
/FlattenerPreset <<
/PresetSelector /MediumResolution
>>
/FormElements false
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/MultimediaHandling /UseObjectSettings
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/PDFXOutputIntentProfileSelector /DocumentCMYK
/PreserveEditing true
/UntaggedCMYKHandling /LeaveUntagged
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/UseDocumentBleed false
>>
]
>> setdistillerparams
<<
/HWResolution [2400 2400]
/PageSize [612.000 792.000]
>> setpagedevice
|