karyomorphological studies in two species of allium l

Karyomorphological studies in two species of Allium L.

Keywords: Allium wallichii Kunth., Allium hookeri Thw., karyomorphology, chromosome classification, karyotype formula, metacentric.

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Journal of Research in

Plant Sciences An International Scientific

Research Journal

Authors:

Harinita Toijam,

Borah SP, Bhaben Tanti

and Borthakur SK.*

Institution:

Department of Botany,

Gauhati University,

Guwahati-781014,

Assam, India

Corresponding author:

Borthakur SK.

Email:

Web Address: http://plantsciences.info/ documents/PS0060.pdf.

Dates: Received: 31 May 2013 Accepted: 03 July 2013 Published: 31 July 2013

Article Citation: Harinita Toijam, Borah SP, Bhaben Tanti and Borthakur SK. Karyomorphological studies in two species of Allium L. Journal of Research in Plant Sciences (2013) 2(2): 213-221

An International Scientific Research Journal

Original Research

Jou

rn

al of R

esearch

in

Plan

t Scien

ces

Journal of Research in Plant Sciences

213-221 | JRPS | 2013 | Vol 2 | No 2

www.plantsciences.info

ABSTRACT: A detail karyomorphological study was undertaken in Allium wallichii Kunth. and Allium hookeri Thw. The study included determination of somatic chromosome number, total chromosome length, volume, arm ratio and centromeric position. The karyotypic study of Allium wallichii Kunth. and Allium hookeri Thw. showed that, both of them have somatic chromosome number 2n=16 and 2n=22 respectively. Chromosome classification was done on the basis of the position of the centromere. In case of A. wallichii, the total chromosome length was found to be 150.94 µm while the relative chromosome length varied in between 4.22 µm - 8.19 µm. The arm ratio was varied from 1.00 - 3.06 .In A. wallichii, out of 16 somatic chromosomes, 9 were of metacentric, 6 were of sub-metacentric and 1 was of sub-telocentric type. For A.hookeri, the total chromosome length was found to be 164.05 µm and the relative chromosome length was varied in between 2.59 µm- 6.95 µm; arm ratio also varied from 1.00 - 4.36. Out of 22 somatic chromosomes, 2 chromosomes were of metacentric, 13 were of sub-metacentric and 7 were of sub-telocentric type. No satellite chromosomes were found in any one of the two species. The karyotype formulae deduced for the two species of Allium may be represented as: A. wallichii Kunth.: m9 +sm6 +st1 +t0 = 2n = 16 and A. hookeri Thw.: m2 +sm13 +st7 +t0 = 2n = 22.

INTRODUCTION

Allium wallichii Kunth. and Allium hookeri

Thw. belong to the family Liliaceae. Recent estimates

accept about 750 species in the genus Allium (Stearn,

1992), and 650 more synonymous species names exist

(Gregory et al., 1998). Alliums are perennial bulbous

plants that produce chemical compounds known as

cystein sulfoxide. And these sulfur containing

compounds give them a characteristics onion or garlic

taste and smell and many of them are used as food

plants. Many species of Allium are bulbous as well as

there are non bulbous species and variety also and many

of them have ornamental values. The young and tender

leaves of A. wallichii are used as vegetables and the dried

ones are used as spices (Manadhar, 1980; Dobremez,

1982). The bulbs are also boiled, fried with ghee and

used for the treatment of cholera and diarrhea (Coburn,

1984). A. hookeri also have some medicinal values. They

are used for treating cold and cough, for healing burn

injuries and wounds (Kala, 2005) and lowering the high

body temperature (Singh et al., 2003). The fresh leaf and

root extract is taken for anthelmintic and also used for

messaging (Sangtam et al., 2012).The bulbs and leaves

of A. cepa, A. sativum, A. cepa var. aggregatum,

A. fistulosum and A. chinense are used as vegetables and

as component of soup. Not only the members of this

family include many vegetables but also some of them

have great medicinal value, e.g. Allium sativum,

Allium hookeri, etc. The status of chromosome research

has undergone fundamental changes in recent years from

an emphasis on qualitative description of structure to an

interdisciplinary phase involving quantifications of

chemical components, identification of segments, even at

the molecular level, as well as in situ cytophotometry,

fluorometry and structural cytology. And also the

application of these techniques led to a more in depth

understanding of the structure, number and behavior of

chromosomes and such a useful study of chromosomes

serve as an aid for dealing with the systematic

relationships, phylogeny and evolution of related plant

groups and also helps to find out the possible modes of

karyotypic variation within and between species

(Mathew and Mathew, 1982 and Aswathanarayana,

2003). In fact, it cannot be denied that all these basic

aspects lay the foundation of the entire discipline of the

present day molecular genetics. So, the various aspects

of research, application of tissue culture, investigation on

molecular genetics and genetical engineering are based

on the basic chromosome research for better utilization

of plants towards commercial cultivation.

Chromosomal differences reflect the general

differences in genic content of the individuals. Thus,

chromosome morphology may be used to establish the

distinctiveness of individual plant species. Improving the

quality of this crop plant to meet the different needs

would involve hybridization programme. So in order to

improve these plants we need to know the thorough

knowledge of chromosome structure and number which

will facilitate the development of an appropriate

hybridization programme and thereby we can manipulate

both the chromosome structure and number for the

genetic improvement of such an important crop plant.

Despite the importance of these two species of

Allium in terms of their culinary as well as medicinal

values, they are poorly explored scientifically with

regard to survey, characterization and conservation.

Some cytological as well as taxonomic investigation of

certain species of Allium has been made but they are so

far quite inadequate. No systematic karyomorphological

classifications of chromosomes have been available for

them, except a few earlier reports (by Ved Brat, 1965).

Therefore, the aim of this study is to provide more

information on the mitotic chromosome in

Allium wallichii Kunth. and Allium hookeri Thw. and to

carry out the karyotypic analysis.

214 Journal of Research in Plant Sciences (2013) 2(2): 213-221

Toijam et al.,2013

MATERIALS AND METHODS

The species of plant Allium: Allium wallichii

Kunth. and Allium hookeri Thw. were collected from

Darjeeling and Manipur (India) respectively.

Methods for cytological study

For detail karyotypic studies, karyotypes were

prepared from the somatic chromosomes. For cytological

studies, root tips were collected from the plants between

7.30a.m to 8.30a.m. and washed them thoroughly with

distilled water and then they were pretreated with para-

dichlorobenzene for 3 hours at 4° C. Then, fixation was

done in Carnoy’s fluid (1:3 glacial acetic acid and

ethanol) for 24 hours at room temperature. After fixation

the root tips were washed with 70% ethanol and finally

stored in 70% ethanol.

Hydrolysis and staining of chromosomes

For preparation of slides, root tips were first

hydrolyzed in 0.1 N HCl at 60° C for 10 -12 mins and

then washed with double distilled water and then they

were kept in 45% acetic acid for 5 – 7 mins and then

transferred to 2% aceto orcein solution, warmed over a

flame for 15 mins and kept for 2 -3 hours at room

temperature (Sharma and Sharma, 1980).

Squash preparation

For preparation of squash the meristematic parts

of root tips were cut out and transferred to a small drop

of 45% acetic acid on a grease free clean slide; then

covered with a cover slip and squashed continuously

without heating and finally sealed with paraffin. Then

temporary slides were prepared and observed under a

compound microscope at a magnification of 10X, 45X

and 100X (using oil immersion). Well scattered

metaphase stages were selected for karyomorphological

analysis of the chromosomes by using camera lucida

apparatus. Drawings of the chromosomes were also

made. The following parameters were considered for the

karyomorphological analysis:-

(i) Total chromosome length (ii) Arm ratio =

Length of long arm/Length of short arm (iii) Volume of

the chromosome (πr2h) (iv) Relative length of the

chromosome = (Length of the individual chromosome/

Total chromatin length of the diploid set) X 100. (v) On

the basis of length, chromosomes will be categorized as:

Type A, Type B, Type C, Type D, Type E and Type F

(vi) F% = (Length of the short arm / Total length of

individual chromosome) X 100. (vii) Total form percent

or TF % = (Total sum of short arm length / Total sum of

chromosome length) X 100. (viii) On the basis of the

centromeric position, the chromosomes may be classified

into metacentric, sub - metacentric, sub - telocentric, and

telocentric (Levan et al., 1964; Tanti et al., 2009; Tanti

et al., 2012).

The chromosomes were grouped into different

categories depending upon their length. These were Type

A= 11.00µm and above, Type B = 10.00µm – 10.99µm,

Type C = 9.00µm – 9.99µm, Type D = 8.00µm –

8.99µm, Type E = 7.00µm – 7.99µm, Type F = 6.99µm

and below.

Finally the chromosomes were classified on the

basis of their centromeric position by following Levan

et al., (1964).

RESULTS

The detail karyomorphological analysis of the

two species of Allium L. is given below:

Allium wallichii Kunth.:

The chromosome no. of Allium wallichii Kunth

was found to be 2n=16 in the somatic cells. The

chromosome length varied from 6.38µm to 12.36µm

while their volumes ranged from 5.00µm3 to 14.96µm3.

The relative length of the chromosomes varied from

4.22µm to 8.19µm. On the basis of the length, the

chromosomes were classified into Type A, Type B,

Type C and Type D (Table 1). The total genomic

chromosome length was found to be 150.94 µm. The

different types of chromosomes categorized on the basis

of the length are represented as:

Journal of Research in Plant Sciences (2013) 2(2): 213-221 215

Toijam et al.,2013

A6 +B2 +C0 +D2 +E3 +F3 = 2n= 16


Toijam et al.,2013

Ch

r.

Typ

e

Ch

r.

No.

Ch

rom

oso

me

Len

gth

T

ota

l

Len

gth

(l +

s) µ

m

Rel

ati

ve

Ch

r.

Len

gth

(µ

m)

Arm

Rati

o (

l/s)

C

hro

moso

me

Cen

trom

eric

Ind

ex (

F%

) P

osi

tion

of

Cen

tro-m

ere

Nom

encl

atu

re

of

Ch

rom

oso

me

Lon

g A

rm

(l)

µm

S

hort

Arm

(s)

µm

R

ad

ius

(r)

µm

V

ol.

(π

r2h

)

µm

3

A

1

8.1

6

4.2

0

12.3

6

8.1

9

1.9

4

0.6

0

13.9

7

33

.9

8

sm

Su

bm

etac

entr

ic

A

2

8.1

6

4.2

0

12.3

6

8.1

9

1.9

4

0.6

0

13.9

7

33

.98

sm

Su

bm

etac

entr

ic

A

3

8.0

0

4.0

1

12.0

1

7.9

6

1.9

9

0.6

3

14.9

6

33

.38

sm

Su

bm

etac

entr

ic

A

4

8.0

0

4.0

1

12.0

1

7.9

6

1.9

9

0.6

3

14.9

6

38

.38

sm

Su

bm

etac

entr

ic

A

5

7.6

1

3.4

4

11.0

5

7.3

2

2.2

1

0.6

0

12.4

9

31

.13

sm

Su

bm

etac

entr

ic

A

6

7.5

8

3.4

2

11.0

0

7.2

8

2.2

1

0.5

5

10.4

4

31

.09

sm

Su

bm

etac

entr

ic

B

7

6.4

4

3.8

1

10.2

5

6.7

9

1.6

9

0.5

5

9.7

3

37

.17

m

Met

acen

tric

B

8

6.4

0

3.8

0

10.2

0

6.7

5

1.6

8

0.5

5

8.9

9

37

.25

m

Met

acen

tric

D

9

5.4

4

3.4

0

8.8

4

5.8

6

1.6

0

0.5

3

7.5

0

38

.46

m

Met

acen

tric

D

10

5.4

4

3.4

0

8.8

4

5.8

6

1.6

0

0.5

2

7.5

0

38

.46

m

Met

acen

tric

E

11

6.0

0

1.9

6

7.9

6

5.2

7

3.0

6

0.5

2

6.7

5

24

.62

st

Su

bte

loce

ntr

ic

E

12

4.7

1

3.1

2

7.8

3

5.1

9

1.5

1

0.5

2

6.6

5

39

.84

m

Met

acen

tric

E

13

3.5

1

3.5

0

7.0

1

4.6

4

1.0

0

0.5

2

5.5

0

49

.92

m

Met

acen

tric

F

14

3.5

0

2.9

6

6.4

6

4.2

7

1.1

8

0.5

0

5.0

7

45

.82

m

Met

acen

tric

F

15

3.4

8

2.9

0

6.3

8

4.2

2

1.2

0

0.5

0

5.0

0

45

.45

m

Met

acen

tric

F

16

3.4

8

2.9

0

6.3

8

4.2

2

1.2

0

0.5

0

5.0

0

45

.45

m

Met

acen

tric

Tab

le 1

. A

lliu

m w

all

ich

ii K

un

th.

These 16 somatic chromosomes comprised of 9

metacentric chromosomes, 6 submetacentric

chromosomes and 1 subtelocentric chromosome. The

karyotypic formula for Allium wallichii Kunth. is

represented as:

The microphotograph of the chromosomes are

represented in Fig. 1. The idiotype and camera lucida

diagram are shown in Fig. 3 and 5A respectively.

Allium hookeri Thw.:

The chromosome no. of Allium hookeri Thw.

was found to be 2n=22 in the somatic cells. The

chromosome length varied from 4.25µm to 11.41µm

while their volumes ranged from 3.34µm3 to 12.89µm3.

The relative length of the chromosomes varied from

2.59µm to 6.95µm. On the basis of the length, the

chromosomes were classified into Type A, Type B, Type

C and Type D (Table No.2). The total genomic

chromosome length was found to be 164.05 µm. The

different types of chromosomes categorized on the basis

of the length are represented as:

These 22 somatic chromosomes comprised of 2

metacentric chromosomes, 13 submetacentric

chromosomes and 7 subtelocentric chromosomes. The

karyotypic formula is represented as:

The microphotograph of the chromosomes is

represented in Fig. 2. The idiotype and camera lucida

diagram are shown in Fig. 4 and 5B respectively.

DISCUSSION

Karyomorphology and chromosome number of a

variety or species are useful in its identification. Each

plant species is characterized by its karyotype. The

importance of karyotype analysis in different plant

species is well known. Karyomorphology and

chromosome number of a variety or species are useful in

its identification and also in establishing the relationships

among related species .The study of variation in

chromosome morphology indicates the progress of

microevolution and to some phenomenon of speciation.

Chromosomal differences are the sources of

genetic variation. Karyomorphology and chromosome

number of a variety or species are useful in its


Toijam et al.,2013

M9 +sm6 +st1 +t0 = 2n = 16.

A2 +B2 +C0 +D4 +E4 +F10 = 2n= 22

m2 +sm13 +st7 +t0 = 2n = 22.

Figure 1. Microphotograph of chromosomes of

Allium wallichii Kunth. 2n=16

Figure 2. Microphotograph of chromosomes of

Allium hookeri Thw. 2n=22

Figure 3. Idiotype of Allium wallichii Kunth. (2n=16)


Toijam et al.,2013

Ch

r.

Typ

e

Ch

r.

No.

Ch

rom

oso

me

Len

gth

T

ota

l

Len

gth

(l+

s) µ

m

Rel

ati

ve

Ch

r. L

ength

(µm

)

Arm

Rati

o (

l/s)

Ch

rom

oso

me

Cen

trom

eric

Ind

ex (

F%

)

Posi

tion

of

cen

tro-m

ere

Nom

encl

atu

re o

f

Ch

rom

oso

me

Lon

g A

rm

(l)

µm

Sh

ort

Arm

(s)

µm

Rad

ius

(r)

µm

Vol.

(π

r2h

)

µm

3

A

1

9.2

6

2.1

5

11.4

1

6.9

5

4.3

0

0.6

0

12.8

9

18

.84

st

Su

bte

loce

ntr

ic

A

2

9.2

6

2.1

5

11.4

1

6.9

5

4.3

0

0.6

0

12.8

9

18

.84

st

Su

bte

loce

ntr

ic

B

3

8.5

1

2.2

4

10

.75

6.5

5

3.7

9

0.6

0

12.1

5

20

.83

st

Su

bte

loce

ntr

ic

B

4

8.4

0

2.1

4

10

.54

6.4

2

3.9

2

0.6

0

11.9

1

20

.30

st

Su

bte

loce

ntr

ic

D

5

6.4

5

2.5

1

8.9

6

5.4

6

2.5

6

0.5

5

8.5

1

28

.01

sm

Su

bm

etac

entr

ic

D

6

6.4

2

2.5

1

8.9

3

5.4

4

2.5

5

0.5

5

8.4

8

28

.10

sm

Su

bm

etac

entr

ic

D

7

6.0

2

2.1

4

8.1

6

4.9

7

2.8

1

0.5

5

7.7

5

26

.22

sm

Su

bm

etac

entr

ic

D

8

6.0

2

2.1

4

8.1

6

4.9

7

2.8

1

0.5

5

7.7

5

26

.22

sm

Su

bm

etac

entr

ic

E

9

4.0

0

3.9

2

7.9

2

4.8

2

1.0

2

0.5

5

7.5

2

49

.49

m

Met

acen

tric

E

10

3.9

2

3.9

0

7.8

2

4.7

6

1.0

0

0.5

5

7.4

2

50

.00

M

Met

acen

tric

E

11

5.1

0

2.1

4

7.2

4

4.4

1

2.3

8

0.5

5

6.8

7

29

.55

sm

Su

bm

etac

entr

ic

E

12

5.0

2

2.1

4

7.1

6

4.3

6

2.3

4

0.5

0

5.6

2

29

.88

sm

Su

bm

etac

entr

ic

F

13

4.9

8

1.4

1

6.3

9

3.8

9

4.3

6

0.5

0

5.0

1

22

.06

st

Su

bte

loce

ntr

ic

F

14

4.3

8

1.7

7

6.1

5

3.7

4

2.4

7

0.5

5

5.8

4

28

.78

sm

Su

bm

etac

entr

ic

F

15

3.9

8

2.1

0

6.0

8

3.7

0

1.8

9

0.5

2

5.1

6

34

.53

sm

Su

bm

etac

entr

ic

F

16

4.3

0

1.7

8

6.0

8

3.7

0

2.4

1

0.5

2

5.1

6

29

.27

sm

Su

bm

etac

entr

ic

F

17

3.9

1

2.1

4

6.0

5

3.6

8

1.8

2

0.5

2

5.1

3

35

.37

sm

Su

bm

etac

entr

ic

F

18

3.9

1

2.1

4

6.0

5

3.6

8

1.8

2

0.5

2

5.1

3

35

.37

sm

Su

bm

etac

entr

ic

F

19

3.5

5

1.4

2

4.9

7

3.0

3

2.5

0

0.5

0

3.9

0

28

.57

sm

Su

bm

etac

entr

ic

F

20

3.5

5

1.4

2

4.9

7

3.0

3

2.5

0

0.5

0

3.9

0

28

.57

sm

Su

bm

etac

entr

ic

F

21

3.5

5

1.0

5

4.6

0

2.8

0

3.3

8

0.5

0

3.6

1

22

.82

st

Su

bte

loce

ntr

ic

F

22

3.2

0

1.0

5

4.2

5

2.5

9

3.0

4

0.5

0

3.3

4

24

.70

st

Su

bte

loce

ntr

ic

Tab

le 2

. A

lliu

m h

oo

keri

Th

w.

identification and also in establishing the relationships

among related species (Lavania and Srivastava, 1999;

Liu et al., 2000; Frame, 2001; Sharma and Sen, 2002;

Salimuddin and Ramesh, 2005; Ying et al., 2006). In the

present investigation, karyotype analysis of A. wallichii

Kunth. showed that the species contain diploid

chromosome number, 2n=16 and A. hookeri Thw.

contain chromosome number, 2n=22. The

karyomorphological studies in both the species of

Allium L. revealed a number of features. The cytological

study reveals that the length of the chromosomes and

gross appearance of the karyotype show a general

resemblance in both the species investigated but, upon

closer examination the karyotypes however showed

variations in the type of chromosomes, absolute

chromosome size, variation in the position of centromere

and variation in relative chromosome size. According to

Ved Brat (1965), most Allium species possess

metacentric to submetacentric types of chromosomes,

but some subtelocentric and telocentric chromosomes are

also present in few species. With respect to chromosome

morphology the karyotype of A. wallichii have a majority

of metacentric and submetacentric types of chromosomes

with the former being more frequent. This shows the

symmetry in morphology of the chromosomes which is a

reflection of relatively primitive karyotypes of the

members of this genus (Stebbins, 1971). In case of

A. hookeri, the chromosomes mostly consist of

submetacentric and subtelocentric chromosomes. It

consists of 13 submetacentric, 7 subtelocentric and only

2 metacentric chromosomes. So it may be described as

asymmetrical, since the karyotype consisting of more

numbers of submetacentric and subtelocentric

chromosomes may be termed as asymmetrical (Stebbins,

1971). The evolution of karyotype is a continuous

process as Jones (1978) has pointed out that the

karyotype alterations during course of evolution are

brought about by various structural alterations of

chromosomes. This implies that the more number of

submetacentric and subtelocentric chromosomes in

A. hookeri Thw. has shown that the genome of this

species has undergone some amount of structural

modifications. This increase in the number of

submetacentric and subtelocentric chromosomes of this

species may be attributed to the phylogenetic changes

occurring in the length of the chromosomes that causes

shortening in the size of the arms leading to the shifting

of the centromeric position as well as reduction in the

absolute length of the chromosomes (Delaunay, 1926).

No secondary constriction could be detected in

any one of the chromosomes of the two species under

investigation.


Toijam et al.,2013

Figure 4: Idiotype of Allium hookeri Thw. (2n=22). Bar indicates 10µm

Figure 5: Camera lucida diagram of Allium

wallichii Kunth. and Allium hookeri Thw.

CONCLUSION

It is an established fact that karyomorphological

analysis forms a prerequisite for the genetic

improvement of any plant species. Therefore, the

detailed cytological investigations undertaken in these

two species A. wallichii. and A. hookeri will contribute

significantly towards their genetical improvement.

Realizing their importance and utility, ex-situ

conservation measures may be initiated for protection,

preservation and regeneration of such a remarkable plant

species. It may further help in research such as breeding

programme, gene exploitation as well as germplasm

conservation.

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