Cytological features of green algae

03/15/19 Cytological features of green algae 1
CYTOLOGICAL FEATURES OF GREEN
ALGAE
By
Prof (Dr.) Ichha Purak
Department of Botany
Ranchi Women’s College,Ranchi

 Nuclear Structure of Green Algae
 Cell Cycle and process of Cell division
 Mitosis and Meiosis
 Chromosome Types recorded in green algae
 Karyotypes : Ideograms
 Chromosome numbers : Basic chromosome number
Polyploidy and Aneuploidy
Resistance or Susceptibility towards chemicals
 References

NUCLEAR STRUCTURE OF GREEN ALGAE
Cells in green algae are eukaryotic in organization
Uninucleate : Chlamydomonas,Oedogonium,Spirogyra, Zygnema,Chara
Multinucleate : Cladophora, Rhizoclonium,Bryopsis,Caulerpa
Nucleus is surrounded by two layered nuclear membrane with nuclear
pores linked with Endoplasmic Reticulum
Interphase nucleus contains uncoiled and expanded chromatin ,one or
more nucleoli and a few deeply stainable lumps (chromocentres)
The nucleoli and chromatin material remain suspended in the granular
matrix of the nucleus.

CELL CYCLE AND PROCESS OF CELL
DIVISIONNuclear division is mitotic in vegetative cells and meiotic in
reproductive cells. Both mitosis and meiosis are like higher plants.
Cell division involves division of nucleus ( Karyokinesis ) followed by
Cytokinesis
The nuclear membrane and nucleoli degenerate at the end of prophase
and are reorganized after the telophase
During Interphase (Resting phase ) : time gap between two divisions, the
nucleus as well as cells prepare them for next division and comprise G1,
S and G2 substages. During these stages, proteins, enzymes and other
necessary molecules are synthesized and replication of DNA takes place.
Mitosis involves Prophase, Metaphase, Anaphase and Telophase as that of
higher plants.

The process of mitosis in green algae follows the pattern as that of higher
plants.The interphase nucleus enlarges considerably (2-3 times) with the
onset of prophase.
The granular structure in prophase nucleus later results discontinuously
stained long chromatin threads as a network.
Late prophase is marked by disappearance of nuclear membrane and
nucleolous.
The chromatin threads start condensation to form rod like chromosomes.
At metaphase chromosomes are arranged on the equatorial plate.
Chromosomes at this stage are thick and rod shaped with smooth outline .
Chromosomes get splitted in to two chromatids attached only at
centromere.
PROCESS OF MITOSIS

Two chromatids of a chromosome start moving apart towards opposite
poles indicating onset of anaphase
The position of centromere can be traced comfortably in orders like
Chaetophorales, Oedogoniales and Charales
The chromatids assume different shapes ( V,J and I ) depending upon the position of
centromere ( Median, submedian, subterminal and terminal )
At telophase two sets of chromatids, undergo decondenstion and nuclear
membrane and nucleolous reappear organizing two identical daughter
nuclei
Cell plate is laid down in between the two daughter nuclei . These
daughter cells undergo a period of rest before undergoing next division.

CELL PLATE PHYCOPLAST AND PHRAGMOPLAST
Two modes of nuclear and cell division are reported in green algae
i) Phycoplast- The mitosis is intranuclear,nuclear membrane not
completely degraded. Additional Microtubules are disposed
transversely to long axis of spindle at the equator at the telophase ,it
is termed as phycoplast.In this case two daughter nuclei tend to lie
closer to each other. Example Volvocales, Chlorococcales,
Microsporales,Ulvales, Chaetophorales and Oedogoniales
ii) .
ii) Phragmoplast -In other case spindle and nuclear envelope are
open at metaphase (completely degraded). At telophase the
microtubules of intranuclear spindle are persistent and result in
formation of phragmoplast, these fibres ( vertical microtubules )
increase in number to form a barrel shaped region as a result of
which the two daughter nuclei are distantly placed example
Coleochaetales, Conjugales(Zygnemales), Klebsormidales
and Charales
03/15/19 7Cytological features of green algae

Chara corallina
Interphas
e
Prophase
Anaphas
e
Telophase

Chara corallina
Metaphase

PLATE SHOWING PROPHASE ,METAPHASE ,ANAPHASE & TELOPHASE
ANAPHASE
PROPHASE
TELOPHASE
Metaphase
Chara fibrosa

4 CELLS OF ANTHERIDIAL FILAMENT SHOWING METAPHASE
METAPHASE
Chara fibrosa ( n=56 )

03/15/19 12
Nuclear cytology of Green Algae since 1940
Significant contributions on chromosomes and karyotyping from India and
abroad:
Godward (1940- 1950 onwards)
Guerlesquin (1963)
Sunderlingam (1946)
Sinha (1958 onwards)
Sarma (1961 onwards)
Noor (1967), Ramjee (1970)
Choudhary (1972) ,Tripathi (1974)
Verma (1969)
Chatterjee (1971) Ray (1976)
Bhatnagar (1978 onwards)
Patel (1979) Jawale (1980) Subramanian (1981)
Mukherjee (1976) Subramanyam and Choudary (1991)
Purak(1986), Bharati (1990)
Pundhir and Vidyavati (1993)
In India significant work has been done in field of Algal Cytology from BHU, Calcutta
University, Ranchi University, Bihar University, Gujrat; Madras; Bangalore &
Bareilly.
Cytological features of green algae

CHROMOSOME TYPES RECORDED IN GREEN
ALGAE
Apparently long chromosomes
Extremely small chromosomes
Polycentric or diffused centric chromosomes
Nucleolar organizing chromosomes
Lampbrush chromosomes

About 250 taxa of green algae have been analysed Cytologically with the help of light
microscopic study. On the basis of extensive studies, following chromosome types
have been recognized in green algae (Sarma, 1982, 1984)
Apparently long chromosomes In Oedogoniales, Cladophorales and
Charales. Chromosomes range upto 12 µm and may have single distinct localized
centromere (Singh and Choudhary,1990). V. J and I shaped configurations of anaphase
chromosomes indicate metacentric, sub metacentric and telocentric nature of
chromosomes.
Extremely small chromosomes (0.2µm-1.0µm ) in orders Volvocales,
Chlorococcales, Ulotrichales, Chaetophorales and some species of filamentous
Conjugales and desmids without any indication of the position of the centromere.
Later on ultrastructural studies provided clues of localized centromere even in very
minute chromosomes and are able to move towards opposite poles during anaphase
through well organized spindle.

Polycentric or diffused centric chromosomes, centromere not
localized, but is diffused throughout length e. g. several taxa of Spirogyra and
desmids
In case of polycentric chromosomes of Conjugales, sister chromatids at metaphase
and anaphase stages lie parallel to each other indicating their poly or diffused centric
nature ( Godward, 1954,Sarma and Srivastava,1993)
Nucleolar Organizing chromosomes (N O chromosomes) have been
recorded in several algae . Godward (1956) has critically analysed the structure,
behaviour and role of secondary constriction in chromosomes of Conjugales and
mentioned about the presence of terminal satellite separated from main body of
chromosome by sub terminal constriction .
Further Electron microscopic studies by Godward (1988) have shown that during
development of nucleolous , the organizer appear as fibrillar material (Nucleolemma)
and interpreted this as chromatin expanded for transcription.

Lampbrush chromosomes have been recorded in the primary nucleus of
green alga Acetabularia of Siphonales (Spring et al 1974) and Koop et al (1979). De
and Berger (1990) studied detailed morphology, meiotic pairing using fluorescence
microscopy and reported certain chromosomes having distinct loops
Electron microscopy of spread
preparation of isolated nuclear
components of Acetabularia revealed the
small lampbrush chromosomes (4-30 µm
long) having typical loops (upto 20 µm
long) comparable to those of animal
lampbrush chromosomes . Association of
some of these chromosomes with
nucleolar structure and with nuclear
envelope were also recognized.
Lampbrush-Type Chromosomes
In the Primary Nucleus of the Green Alga
Acetabularia mediterranea

KARYOTYPES IN GREEN ALGAE
Symmetric and primitive- Karyotypes with low chromosome number,
almost of same size, large size of chromosome, with median and
submedian centromere : examples orders like Cladophorales,
Oedogoniales and Charales
Asymmetric and advanced -high number of chromosomes , small size
of chromosomes, with appreciable difference in size of smallest and
longest chromosome ,terminal and sub-terminal position of centromere :
examples orders like Ulotrichales, Siphonales and Chlorococcales
Analysis of karyotype symmetry can be used as a tool to trace
intervarietal,interspecific or intergeneric karyotypic evolution.

Karyological studies play an important role in understanding the
evolutionary pattern as opined by Stebbins (1971) and help in solving
systematic position of certain controversial genera
a) Sphaeroplea and Microspora have been shown to be more clearly
related to Ulotrichales than to Cladophorales or Siphonales (Sarma,1962)
on the basis of its karyology
b) Consideration of Schizomeris as an ecophene of Stigeoclonium
( Campbell and Sarafis,1972) has been opposed by Sarma and Choudhary
(1975) on cytological grounds and according to them Schizomeris
deserves a generic status.
Karyotypes have been classified into 12 categories (1A- 4C) as per
Stebbins(1958-1971) taking into account both the degree of difference
between the largest(LC) and Smallest(SC) chromosome of complement
and position of centromere on chromosome.03/15/19 Cytological features of green algae 18

A- Photomicrograph of metaphase plate(n=56) B- Outline drawing of metaphase
A B
PLATE-Karyological
features of Chara fibrosa
var.fibrosa
f. tylacantha(Nordst.) R D W
(Purak and Noor ,2018)
C- Karyogram showing 56 chromosomes
KARYOTYPIC FORMULA
( 8M 6m) B + ( 10 M 22m 4sm 4 t) C + (2 t) D
C

CHROMOSOME NUMBER RECORDED IN GREEN
ALGAE
SN Order Chromosome numbers recorded (n)
1 Volvocales 4, 7,8, 16, 36, 38±4
2 Chlorococcales 4, 10,14,18, 19, 32,80
3 Ulotrichales 3, 4, 9, 10,13,16, 48
4 Cladophorales 12, 24, 36, 48, 72, 96
5 Chaetophorales 3, 4, 5, 8, 10, 13, 26, 56
6 Conjugales 2, 24, 92, 592
7 Oedogoniales 9,11, 13, 15, 16,17, 19, 21, 22, 25, 32, 37
41,46,49
8 Siphonales 4,6,8, 10,12, 16, 20
9 Charales Chara 7, 14, 21, 28, 35, 42, 49, 56, 63, 70
Nitella 6, 9, 12, 15, 18, 21, 24, 27, 30, 36, 42, 48
Tolypella 10, 11, 15, 20, 50
Lamprothamnium 14, 42
Lycnothamnus 14, 28, 35
Table : Chromosome numbers in different orders of Chlorophyceae

In green algae chromosome number ranges from n=2 in some species of Spirogyra to
n=592 in a desmid Netrium digitus (King,1960). Both belonging to order conjugales
The lowest chromosome number in Volvocales is n=4( Chlamydomonas monoiae)
and highest is n=38±4 in Chlamydomonas eugametus.The chromosomes are
mostly minute dot like. Aneuploidy is prevalent in the group (Sarma,1982 )
Chromosome number in Chlorococcales varies from n=4 ( Scenedesmus platydiscus)
to n=80 (Eraemosphaera viridis )
Ulotrichales show a range of n=3 (Prasiola japonica ) to n=48 (Hormidium
crenulatum .The chromosomes are extremely small ,in Ulothrix zonata , Ulva
lactuca and Enteromorpha. Sarma(1962) reported n=16 for Sphaeroplea
annulina. He also mentioned that Sphaeroplea should be included under
Ulotrichales on cytological grounds
Sinha (1967) reported polyploid counts 48 ,72 and 96 in four fresh water
forms of Cladophora glomerata ,where as its marine form exhibits diploid
count of 24 chromosomes.03/15/19 Cytological features of green algae 21

Courtesy: Sarma YSRK (1962) Nuclear cytology of Sphaeroplea annulina (Roth)Ag. Cytologia 27 :71-78

Courtesy: Sinha JP (1963) Cytotaxonomical Studies on Cladophora flexuosa (Griff.) Harv., a Marine sp.
Cytologia 28 : 1-11
03/15/19 23
.12 bivalents at meiotic metaphase

Courtesy: Sinha JP (1967) Cytotaxonomical Studies on Cladophora glomerata Four Freshwater Forms
Cytologia 32: 507-518
03/15/19 24
96
72
48

Chromosome count 2n=24 in Cladophora glomerata
(Courtesy: Noor, 1965, 1967)
12 bivalents at meiotic metaphase of Cladophora
glomerata
Rhizoclonium implexum n=12 a- Metaphase, b- outline drawing of metaphase plate c- late
anaphase
a b c
a
b c d
03/15/19 25
a b c

Hydrodictyon reticulatum
(L) langerheim n=18. a
Metaphase b- outline
drawing
a b c
Draparnaldiopsis indica (bhardwaj) n=8. a- early prophase b- metaphase
c- outline drawing
Courtesy: Noor (1967)
a b

Species No of nuclei
/cell
Diameter
of nuclei
( µm)
No of
nucleolus/nucl
ei
R
dimorphum
2-4 6-7 2
R stagnale 2-16 4-4.5 2
R crispum 4-12 4.6-5.5 2
03/15/19 27
Courtesy: Verma, B N (1980)
Mitotic metaphases (24)
Meiosis: R. dimorphum a- synchronous division
b- diakinesis 12 bivalents
a b

Courtesy: Verma B N (1979)
P kewensis n = 24
P polymorpha n = 36

N=4 is lowest chromosome count for Chaetophorales (Draparnaldiopsis indica)
and Fritschiella tuberosa whereas n=56 forms the highest count (Trentepohlia
treubiana and Trentepohlia unarata) .Michetti et al (2010 ) have reported 3,5 and 8
chromosomes in four species of Stigeoclonium from Argentina
The different chromosome counts for different genera of conjugales show that
euploidy has played significant role in speciation of Spirogyra but on the whole
aneuploidy plays a role in species diversification. Most common chromosome
number in Spirogyra is n=24
Sinha J P (1962) has cytologically analysed 2 spp of Oedogonium . He reported
chromosome Number 19 in O. cardiacum and 41 in other species of Oedogonium
. Noor (1965) have recorded 13 chromosomes in Oedogonium annulus.
Singh and Choudhary (1990) made cytological analysis of 9 macrandrous taxa of
Oedogonium recording chromosome numbers n= 13, 17,19,21,22,25, 32 and 37.

Courtesy: Sinha JP (1962) Cytological studies on
Oedogonium cardiacum Wittrock and another
Oedogonium Sp. Cytologia 28:194-200
03/15/19 30
19 chromosomes
41 chromosomes
Metaphase plate
showing 13
chromosomes in
Oedogonium annulus
Noor(1967)

The lowest chromosome number of n=6 is exhibited by several taxa of Nitella while
highest number n=70 is recorded in Chara zeylanica . Almost all Charophycean
genera like Chara, Nitella, Tolypella, Lychnothamnus and L.amprothamnium show
varied euploid series.
Chromosome number in Charales
On the basis of cytological evidences the view of Chaetophoralean
origin of charophytes put forward by Desikachary and Sunderlingam
(1960) has been discarded by Sarma Khan and Ramjee (1970)
More than 150 taxa belonging to Indian Charophytes have been subjected to
cytological studies.(Khan and Sarma 1980)

Sundaralingam 1946
 Khan and Sarma 1964
 Sarma 1968
 Sinha and Noor 1967,1968,1969,1971
Noor 1968
Sinha and Ahmad 1971,1973
 Sinha and Verma 1971
 Sarma,Khan and Ramjee 1970
Chatterjee 1971,1975,1976,1979
Chatterjee and Ray 1976
 Channaveeraiah and Bharati 1974
 Ramjee and Bhatnagar 1976
 Bhatnagar 1976
 Patel and Jawale 1978
 Purak 1986
 Noor and purak 1989
 Purak and Sinha 1998
 Vidyavati and Pundhir 1993
CHAROPHYTOLOGIST FROM DIFFERENT PARTS OF INDIA

Table No 1 Basic chromosome numbers of genera
of Charales
S.N. Name of Genus Basic Chromosome Number
1 Chara 7
2 Nitella 3
3 Tolypella 5
4 Lychnothamnus 7
5 Lamprothanium 7

Different chromosome counts (n) recorded in
varied Indian taxa of Charales
Genus Chromosome numbers (n )
Chara 7,8,14,21,26, 27,28,35,42,48,49,56,63,70
Nitella 6, 9,12,15,18,19,21,24,27,29,30,36,42,48
Tolypella 10,11,15,20,50
Lychnothamnus 14,28,35,60
Lamprothamnium 14,42

Some aneuploid counts have also been recorded in a few
Aneuploid chromosome counts recorded in
few charophyte taxa
Aneuploid
count
Taxa Reference
8 Chara fibrosa f. hydropitys
Nitella furcata f. dictyosperma(H &J Gr)
N itella gracilis annandale (Pal) RDW
Noor and Mukherjee 1975,1976
Subramanyan, Choudary, and Sarma
1991
-do-
11 Tolypella nidifica var. glomerata (Desv.)
RDW
T. nidifica f. stenhammariana (walm in
lilj)RDW
Tolypella intricata f.prolifera (Ziz ex A.Br.)
Bhatnagar and Ramjee 1976
Labh and Verma 1984
Sarma & Ramjee,1969
19 N. furcata f.papuana (Zanev) RDW Noor and Mukherjee 1975
26 Chara globularis var. kokeilli (A.Br.) RDW Sarma and Bala 1983
27 Chara setosa Klein ex.Willd Prasad and Verma 1985
29 Nitella acu minata var. acuminata f.
belangeri (A.Br.) RDW
Bhatnagar and Johri1988
48 Chara zeylanica f. elegans Bhatnagar and Johri 1988
60 Lychnothamnus . barbatus f. barbatus
(Meyen) Leonh
Subramanian and Ganesan1983

Prophase Chara
zeylanica
Metaphase
C.hydropitys
Anaphase Nitella
batrachosperma
Telophase N. batrachosperma
Different stages of mitosis of Charophytes (Noor,1967)
03/15/19
36Cytological features of green algae

Nitella acuminata (n=18) N. dualis (n=18) N. gracilis (N=18)
Chara braunii (n=14) C. globularis (n=28) C. corallina (n=42)
Some metaphase plates : Charophytes (Noor, 1967)
03/15/19
37Cytological features of green algae

Noor M N and Mukherjee ,S (1975) . On the
Aneuploid chromosome number (8) in
Chara hydropitys Reich from India.
14
8
Noor M N and Mukherjee s
(1977 )Some new records
of chromosome number in
Indian Charophyta
Chara braunii f. kurjii n=7& 14
Chara zeylanica n=49 *
Chara vulgaris n=35 *
Nitella pseudoflabellata n=21*
Nitella gracilis n=36
49
21
36

Chara zeylanica var. zeylanica f. elegans (n = 48)
03/15/19 39
Courtsey : Bhatnagar S K and Johri M (1986)
Some aneuploids in Indian Charophyta .Acta
Bot. Neeth. 35 (4): 377-381
Nitella acuminata var. acuminata f.belangari (n = 29).
Prasad P K and Verma B N ( 1985 ): Aneuploid count
for Chara setosa Klein ex. Willd. Cytologia 50:241-245
n=27 (Aneuploid number )
Chara setosa

B. V. S. Subrahmanyam and Y. B. K.
Chowdary (1992)
Karyological studies of Some Indian
Charophyta.Cytologia 57: 409-415
Chara vulgaris n=35
Chara zeylanica n=42
Nitella heterodactyla n=18

L Labh and B N verma (1985)
New counts for Chara fibrosa
n=14,28,42 and 63
Have been recorded
Different varieties of
Chara fibrosa

Plate 19 Nitella furcata f. microcarpa n=36
Plate 20 Nitella furcata f. oligospira n= 18
Plate 21 Nitella furcata f. oligospira n=24
Plate 22 N. pseudoflabellata f. pseudoflabellata n=36
Plate 23 Nitella gracilis f. confervacea n= 36
Some slides from Ph. D. Thesis : Ichha Purak (1986)
Under Supervision of
Professor M N Noor
03/15/19 42
CYTOTAXONOMIC AND MUTAGENIC STUDIES OF SOME MEMBERS OF
CHAROPHYCEAE AND OTHER GREEN ALGAE

Plate 24 Nitella hyalina f.hyalina n=18
Plate 25 Nitella hyalina f. formosa n=24
Plate 26 Nitella hyalina f. maxima n=18
Plate 27 Nitella translucens f. axillaris n=42
Plate 28 Nitella tricellularis var. tricellularis n=36
03/15/19 43
Purak,1986

Plate 29
Plate 30
Plate 31
Plate 32
Plate 33
Plate 34
Plate 29 Chara braunii f. braunii n=14
Plate 30 C brauni f. coromandelina n=14
Plate 31 C braunii f. novimexicana n=14
Plate 32 Chara braunii f. kurzii n=14
Plate 33 C .braunii f. oahuensis n=28
Plate 34 C braunii f. perrottetii n=14
03/15/19 44
Purak,1986Cytological features of green algae

Plate 35
Plate 36 Plate 37
Purak,1986
Plate 35 Chara corallina f. corallina (n=42)
Plate 36 Chara corallina f. kyusyensis (n=42)
Plate 37 Chara fibrosa f. tylacantha (n=56)

Chakrabarty R and Ray,S (2016) reported n=18 and n=21 for two
populations of Nitella hyalina (DC) Ag collected from district Burdwan and
Birbhum of West Bengal. These two populations do not show any variation
in morphological features but variation in chromosome number and
karyotypes.
Chakrabarty, R and Ray,S (2016) Chromosomal variations and cytotaxonomical
considerations in two populations of Nitella hyalina (Charophyceae, Characeae) from
West Bengal, India. Phykos 46 (2): 14-19
Figure- metaphase plates of Nitella hyalina a- Population 1 n=18 and b- Population 2 = 21

RESISTANCE OR SUSCEPTIBILITY TOWARDS CHEMICALS
High doses of chemicals induce abnormalities as chromosome breaks,
clumped metaphases,anaphasic bridges ,laggards etc.
03/15/19 47
Chemicals used : Colchicine,Triacontal, antibiotics,Herbicides,Pesticides
Experimental materials : Rhizoclonium,Oedogonium,Chara,Nitella,
Tolypella,Spirogyra
Significant contributors :
Noor (1969)
Sarma and Tripathi (1973,76)
Sarma and Abhyavardhani (1980)
Singh, Verma and Bhatnagar (1980)
Noor and Purak (1989,1993)
Pal and Chatterjee (1987,1989)
Bhatnagar and Johri 1987
Some of the chemicals in lower concentration are stimulatory to mitotic
division,whereas higher concentrations are inhibitory to division.

Sarma Y S R K and Tripathi, S. N (1976) Effects of Chemicals on Some Members of Indian
Charophyta I Caryologia 29(3):247-262 ·
Chemicals used: Colchicine,MH, Caffeine,
Theobromine

Sarma Y S R K and Tripathi, S. N (1976)
Effects of chemicals on some members of
Indian Charophyta. II Caryologia 263-276
Chemicals used : GA,
2,4-D,Coumarin,Acenaphthene

Bhatnagar S K ,Verma,A and Singh V K (1989) Mutagenicity of Triacontanol in Tolypella prolifera
(Div. Charophyta) Cytologia 54: 183-189
Recommended application of
Triacontanol in place of polyploid
inducing colchicine.

Noor M N and Purak I (1989) Effects of two antibiotics on nuclear division of Chara braunii
Gm, (Characeae) Cryptogamie Algol 10(2) : 143-152
03/15/19C 51Cytological features of green algae

Purak I and Noor M N (1991) Effect of Chloramphenicol on growth pattern and cytological
behaviour of Chara corallina .Phykos 30(1&2): 123-128
Figs. 1-7 Effect of chloramphenicol on Chara
corallina
1. Normal coumt n=42
2. Rotation of Anaphase plates
3. Anaphasic bridge
4. Clumped metaphases
5. Laggards
6. Rings at metaphase
7. Erosion of chromosomes

Bharati,S G (1990) Morphological and Cytological studies on two taxa of Chara
fibrosa Ag.Ex. Bruz. EM RDW Indian Bot.soc. 69: 315-318

Bharati, S G and Chennaveeraiah, M S (1983) Lychnothamnus barbatus var. gigantii
var. Nov. Cytologia 48: 843-846.
Bhatnagar, S K (1981) Cytotaxonomy of Charophytes of Rohilkhand division and
different effect of different chemicals on selected taxa Ph.D thesis, Rohilkhand
University, Bareily.
Bhatnagar, S K (1983) The concept of basic chromosome numbers in Charophyta. A
review. Cryptogamie, Algologie 6: 111-116

Bhatnagar, S K and Johri,M (1985) Mutagenic efficacies of tri-aconatol in Chara
braunii Gm.(Charopohyta) with reference to its application in chromosome analysis.
Crypto.Algol 6(4): 273-280

Bhatnagar S K and Johri M (1986) The genus Tolypella from Indian subcontinent: A
cytotaxonomic approach.Cytologia 51: 413-419
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