PART I
CYTOLOGICAL STUDIES IN
ASTERACEAE
INTRODUCTION
The Asteraceae family is one of the most numerous within the Phanerogames.
This dicotyledonous family is widely distributed and constitute about 10% of
the entire population of flowering plants. The members of this family show a
remarkable diversity in hattit. Asteraceae has been thought to be at or very
near the peak of dicot evolution. This family is readily distinguishable from all
other families by the flowers aggregated together in head or capitulam. The
number of florets in a head varies enormously from several thousands as in
huge heads of some sunflowers, to a single flower, as in Echinops. where, the
single flowered head is hc~wever, generally associated in secondary heads.
Florets of a head is bisexual or unisexual (monecious or dioecious) or the
outer (ray florets ) female or asexual. Many members of the family are of
economic or medicinal value from the presence of ethereal and fatty oils,
resins and bitter princirles. Many indegenious plants are used as
grandmother's remedy for common colds, chills and fever.
Dumort (1822) was the first to propose the name Asteraceae and later the
International code of Botan cal Nomenclature approved that it can be used as
an alternative to Composita~:. According to Turner et al. (1979) the dispersion
centre of the family was probably South America. The plants are most
abundant in the tropical and temperate land but are also found in artic and
alpine regions. In India the plants occur in all possible climates and places but
are less common in areas under rain forests. They grow both in hills and plains
and play an important role ir vegetation.
The family is sub divided into two subfamilies viz, Tubuliflorae and
Liguliflorae, under the sub family Tubuliflorae 12 tribes are included viz.
Vernonieae (Vemoniaceae* ), Eupatorieae (Eupatoriaceae*), Astereae
(Asteroideae*), lnuleae (Inuloideae*), Heliantheae (Helianthoideae*),
Helinieae (Helenioideae*), Anthemideae (Anthemideae*), Senecioneae
(Senecionideae*), Calendulae (Calendulaceae*) Arctotideae (Arctotideae*).
Cynareae (Cynaroideae*) and Mustisiea (Mutisiaceae*) and one tribe
Cichorieae (Cichoriaceae*) under the sub family Liguliflorae (Bentham and
Hooker, 1883). The work fi)llowed here is on the basis of Bentham and
Hooker system of classification. Bentham and Hooker have placed the family
Asteraceae in their second order- the Asterales of Class Dicotyledones,
Division Gamopetalae and Series Inferae.
There is considerable difference of opinion regarding the number of
constituent taxa within the family. Lawrence (1956) recognized the Asteraceae
spread over 950 genera and 20000 species. Willis (1966) considered only 900
genera and 13000 species, where as according to Cronquist (1968) the number
of species included in this lamily is 19000. Jones and Luchsinger (1987)
represented by about over 20000 species, which are world wide in distribution.
140 of its genera and over 700 species have been reported from India. Fifty
two percent of the species art: endemic. In the world wide genus Senecio. the
largest in the family, with abcut 2500 species. Vernonieae is one of the 13
* name as in Flora of British 'ndia
pantropical tribe comprising more than 1500 species (Johri and Singh, 1997).
The Eupatorieae are a very diversified tribe and consists of more than 2300
species and 180 genera (Watanabe et al. 1995). The tribe lnuleae comprises
38 genera and around 480 species distributed mainly in Eurasia (Panchami
and Vijayavalli, 1998).
A majority of the plants are herbs - annual or perennial, some are shrubs, a
few are herbaceous or woody climbers, rarely trees. Majority are mesophytes.
Apart from mesophytes, sc~me occur as xerophytes, aquatic or marsh plants
and epiphytes. True water Aants are rare. Many taxas possess a milky juice
while in the other members the juice is watery, resinous and bitter. The tribe
Cichorieae is charecterised by a system of laticiferous vessels which
accompany the phloem tiaue; they are formed by the disappearance of the
transverse walls from longitudinal rows of cells and are also freely joined by
cross union. In the other lribes oil containing passages occur in the cortex
outside the vascular bundles running from the root through the stem and
generally continued in the leaf. The carbohydrate reserve material inuline is
dissolved in the cell sap of the roots and tubers of many members of the
family.
The remarkable success ol'the Asteraceae as evidenced by its great numerical
preponderance in genera and species over other families and the abundance of
many of its species, and its world wide distribution, it at any rate in part due to
the admirable adaptations of its flowers for cross pollination by a great variety
of insects (Rendle, 1938). Plentiful supply of easily accessible nectar. its
protection from rain, the close association of the flowers and the possession of
pollen mechanism, ensures c r x s pollination in the event of insect- visits. In
additon to these characters other factors are numerous small fruits produced
from capitulam and each of these fruits has direct aids to distribution by the
wind and water. They are chaffy character of small fruits , as in many
Anthemidea, or a covering of loose wooly hairs on the fruit itself or on the
bract; or more generally in the parachute development of the pappus.
The family is of great eccnomic importance. Several types of plants are
included in the family. Many plants possess medicinal properties and many are
grown as ornamentals. Coml~lex antibiotic preparations are also obtained from
some members of Asteraceile (Smimov et a]. 1995). Insecticides, oils. dyes,
and edible products are preoared from some members. The pharmacological,
medicinal, economical and other industrial uses of various taxa used in the
present study have been disc:ussed in the second part ofthe dissertation.
Genetic and cytological studies have been extensively carried out for more
than hundred years. Most >f the cytogenetic descriptions in the early stages
consists of informati01 only on the chromosome number without
morphological karyotype assessment of the group. Recently majority of
chromosomal and cytogenetical studies have been based on visible
characteristic of the chromosome. Karyotype analysis, a well established
method, is based on the morphological characteristic of chromosome and
widely used in cytogenetical analysis. The remarkable constancy of
chmmosome morphology ansj number within genera has been invaluable in
the study of plant systamatiss. In addition, many of the groups that have
distinctive chromosome nunbers are among the most readily defined
taxonamicaly. Therefore, it is attempted to find clue regarding phylogenetic
relationships through out the family by assessing chromosomal variations.
Polyploidy is very defused among plant species (Stebbins, 197 1 ; Grant, 1981 ;
Gill and Gill, 1994) and acsording to Lewis (1980) polyploidy and diploidy
have been important mechanism in the evolution of many plant groups.
Chromosome counting of same species may show different chromosome
numbers, from the distinct cytotypes of the species. Generally the
polyploidiied organisms have a great genetic plasticity due to a greater genetic
variability present in the genome (Mariano and Morales, 1999). However,
cytogenetic studies, such as determination of chromosome number and
morphology, along with bending patterns have been used increasingly in the
taxonomic determination of many species, where phenotypic or
morphoanatomic traits an: in sufficient to discriminate between species
Pioneering attempts on the cytology of Asteraceae are those by Tahara ( 1 9 15)
and Geisler (1931). Suffisient information is available on the members of
Asteraceae by virtue of the major cytological surveys conducted by Turner
et.al (1961,1964,1965) arid Mehra et al. (1965). Powell and PoweIl(1978)
recorded chromosome nurnbers of 100 species and 54 genera of Asteraceae.
Lawrence (1980); Adame and Falavera (1980); Luque et al. (1984); Jalals and
Pellinen (1985); Rabakonandrianiha and Carr (1987); Stahevitch and Wojtas
(1988); Husaini and Iwo (1990) Atlagic et a1.(1992); Endo and lnada (1992);
Tavassoli and Derakhshndesh (1993) Ayodele (1994); Strother and Panero
(1994); and Razaq et al. (1994); have also been made attempts in various
species of the family. However, almost all these cytological studies have
concentrated mainly on the determination of chromosome number and
provided scarcely any data on chromosome morphology and structure in
detail, nevertheless, this work laid the foundation for later researches.
Detailed karyotype analysi:; of Asteraceae members were carried out by
Khamdamov and Noskova I 1986); Jayaramu and Chanerji (1986); Ruas and
Ruas (1987); Chui et al. (1989); Watanabe et al. (1990); Qiao et al. (1990);
Herickhoff et al. (1 994); Branas et al(1994); Dagne (1995) Xiong et al. (1995)
and Arturo et al. (1996). Tlie study ofessential oil composition and karyotype
analysis are very meagre. Maffi et al. (1993) reported Essential oils.
chromosome numbers and Karyotypes from Achillea species. However, all
these karyomorphological studies mostly depended on the conventional
method.
Cytology is believed as a dependable tool for solving taxonomic problems and
for elucidating systematic relationships, phylogeny, and evolution of related
plant groups. The irformation like chromosome number, structure,
morphology and behaviol during mitotic and meiotic division have been of
considerable value in understanding inter relationships and delimitation of
taxa. (Yoshikane and Naohiro 1991). Therefore, these factors are used as
classificatory criteria in the same manner as the morphological characters:
since the chromosomes have direct relation to the genetic system of which
they are. an integral part ( Den Harlog et al. 1979).
In Asteraceae, chromosome numbers have provided to be of great value in
the determination of tribes (Naik, 1992). Raven et al. (1960) Raven and Kyos
(1%1) , Hair (1962), Omdui'f et al. (1963, 1967) Payme et al. (1964) and
Solbrig et al. (1964) have indicated that cytologically the tribe Helinieae is a
diverse group of various aftinities. The use of chromosomal characters in the
delimitation of genera can be seen in some Asteraceae members (Babcock,
1947).
Katyotype analysis have been useful in classifLing phylogenetic and
evolutionary relationship between some related species and species groups,
where differences in karyotyopes between taxa are not distinct, however, it has
been difficult to evaluate heir differences using conventional methods, and
interpretation of the differences have often been critized for lack of statistical
analysis (Watanabe et al. 1990). Several indices measuring karyotypic
differences are available i f homologies of chromosome and chromosome arms
are accurately ascertained (Duncan and Smith, 1978). It is hardly possible to
obtain definite evidence con chromosome homologies in plants, because G-
banding techniques are immature. Hence more objective method for assessing
karyolypic differences is needed. To answer this deficiency, in the present
study, it is used a numerical method for describing karyotypic difference or
similarities
In some Asteraceae members karyotype analysis is requiring the identification
of homologues are unreli:lble, because not all chromosomes can be
distinguished by their Length and cenhomere position, and no useful additional
cytological markers are avitilable (Koopman et al. 1996). Therefore the
karyotypes are established using numerical parameters describing the
chromosome length, area, perimeter, visual apparent three dimensional
volume, uniformity coeff11:ienf variation coefficient, disparity index of
chromosomes, total forma penentage (mean cenhomeric index value or TF%)
and number of discernible satellites. In some groups karyotypic differences
between species are largely quantitative and have been difficult to be assessed
by conventional quantitative methods.
Chromosome identification and mappings are indispensable in cytological and
genome analysis. There are limitation for the conventional measuring and
characterization of chromosome complement by visual evaluation, especially
for very small chromosomes. The ordinary karyotype analysis has provided
only limited success from the view point of chromosome identification, not
only in the plants with small chromosomes, but in many other plant species as
well (Fukui and Mukai, 15188). It had also been diff~cult to identify the small
chromosomes because of the similarity on the morphology at the mitotic
metaphase stage. In addition to that stainability of chromosome is not always
good. Only N and C banding methods are presently available in many cases,
these techniques, however, cannot always be applied. Therefore image
analysis of the chromoson~e by the chromosome image analysis system
(Fukui, 1985, 1986 a) wa:, employed in order to obtain data, which are
quantitatively accurate. Seniiautomatic karyotyping including numerical data
acquisition, pairing and arrangement of chromosome by digital manipulation
of the image using computer devices results in a detailed construct of
descriptive data (Fukui, 1988).
In the present investigation, karyomorphometrical analysis was conducted on
forty seven species of thirty four genera with the aid of improved techniques
(Sharma and Sharma, 198C; Fukui and Kamisugi, 1995). This study was
designed and aimed to bring out the Karyotype characteristic of different
species in the family Asteraceae and to check for marked symmetry or
asymmetry in the chromoiomal complements. The basic chromosome
numbers were employed in formulating phylogenenetic speculations and to
find out the direction of evolution in the family. Detailed
karyomorphometrical studies pertaining to chromosome length, area,
perimeter, volume, disparity index, variation coefficient and total forma
percentage are high- lighted n the present investigation in order to throw light
on the phylogenetic relation:,hip, the systematic position and affinities of the
different genera and different species of the family Asteraceae. This study is
also report original counts of chromosome numbers in Asteraceae and to
compare them with member; reported previously and to statistically test the
correlation between changes in chromosome number and karyotype. Further
an attempt is also made based on the previous and present cytological data to
decipher the inter relation:;hips among the various taxa to obtain a clear
understanding of evolutionay process at work in the family Asteraceae.
MATERIALS
Materials for the present stucy were collected from different localities, wild as
well as cultivated areas of South India. Forty seven species examined belong
to thirty four genera representing 9 tribes. Table - 1 show the collection sites
of the forty seven taxa used in the study. Voucher specimens are deposited in
the herbarium of Sacred Heart College, Thevara, Kochi, Kerala, South India.
1. Eleplantopus scabei- Linn.(Fig. 1 a)
The plant is a stiff sub - scapigerous herb with ohovate, oblong basal
leaves upto 16 cm long, narrow sessile cauline ones. Stiff heads with
purple flowers. Heads homogarnous of 2 - 5 flowers collected in a
head like clusters supported by 3 broadly ovate - cordate leaf bracts.
2. PhyUocephalum rutzgucharii (Gamble) Narayana.(Fig.2a)
( Centratherum ranpachurii Gamble)
Plant is an annul herb; stem loosely moniliform hairy, leaves alternate
4 - 9 cm x 1.5 - 3 c.m, ovate or elliptic - lanceolate. Heads solitary or
2 - 3 at tip of long slender peduncle. Homogamous.
3. Vernonia cinerea Less (Fig.3a)
This species is one of the commonest in Indian weeds. Stem slender,
15 - 17 cm high, g.ooved and ribbed. Leaves 2.5 - 5.0 cm; variable in
shape, broadly elli!~tic or lanceolate,
1 1
us. Flowers vinkish
and purple, in minute heads, in rounded or flat topped corymbs. Heads
homogamous.
4. Adenostemma lavenia (Linn) Kuntze. (Fig.4a)
The plant is an erect herb, stem, leaf viens and petals glandular
pubescent. Leaves 5 - 15 x 3 - 8 cm broadly ovate to elliptic -
lanceolate, Heads inteminal lax panicles, homogamous .
5 . A g e ~ t u m conyzoides L ~ M (Fig.5a)
The plant is an erect herb, annual, 30 - 60 cm, hispidly hairy, leaves
petioled, ovate crenate, heads small in dense terminal corymbs,
Homogamous flowers blue or white.
6. Ageratum haustonianwn Miller. (Fig.6a)
Herb to 1 m, annual, hairy. Leaves serrate, acuminate, corymb dense,
capitula blue. Flow(xs deep blue. (often higher altitude specimens of
A.conyzoides, with progressively larger and deeper blue capitula, are
confused with this sl~ecies).
7. Chromolaena odor~zta (Linn) R.Kig & H. Robinson. (Fig.7a)
(Errpotorium odoratum. Linn.)
Aromatic, erect, vis8:id -pubescent sub shrub to 3 m. Leaves opposite,
simple obovate, to deltoid ovate, acute, crenate, serrate, sub palmately
3 nerved. Capitula corymbose, stalked. corolla white to purple.
Flowers homogamc~us.
8. Eupoforium trblinenve Vahl. (Fig.8a)
(E.ayapana Vent.)
An aromatic under shrub, 0.9 - 1.2 m high with trailing stem, rooting
at the nodes. Subsessile lanceolate leaves and lax corymbs of bluish
flower heads. Flower!; homogamous.
9. Mikania cordata (Bu1m.F) Robins. (Fig.9a)
( M. scandens, Willd.:~
The plant is a climbing shrub, leaves long petioled, ovate, acute or
acuminate base rounded cordate or truncate crenate or angled. Some
times villous beneath. Heads 4 flowered corymbose terminating lateral
branches, homogamous.
10. Conyza bonariensis (Idinn) Cronq.(Fig.lOa)
Sericeous herb. Leaves linear - lanceolate, hirsute above and below,
margin entire to sparingly serrate, sub sessile, panicles terminal lax
racemose, capitula cre Im.
1 1 . Conyza canadensis (Linn) Cmnq.(Fig. l la)
It is an annual plant with an erect branched stem, densely covered with
narrowly lance-shapea leaves, and bearing many flower heads in dense
clusters. Each flower heads has many central tubular disk florets and
several outer rows of ray florets. Both are yellow and white.
.? a- : * - 12. Dichrocephala chrysanthemifolia DC.(Fig.l2a)
The plant is an annual herb. Leaves alternate, sessile, auricled at base,
obovate, heads very small, panicled, heterogamous globose, not rayed.
13. Erigeron mucronatus DC.(Fig. 1 3a)
The plant is an annual herb. Leaves narrow linear. Head
heterogamous and rayed. Corollas of ligulate flowers narrow, white,
pink or purple.
14. Blmea lacera DC. (Fig.l4a)
Strongly scented her11 to 75 cm, glandular pubescent, interspersed with
eglandular hairs. Leaves elliptic oblanceolate, 2.5 - 6 x 1.3 - 3.5 cm
capitula 5 - 7, shortly stalked in dense corymbose, spiciform, panicles
terminating the branchlets. Corolla yellow in disc florets.
15. Blumea mollis (D.Don) Merr.(Fig.l 5a)
Villous silky hairy, !items erect sub simple very leafy, leaves petioled,
obovate, irregularly toothed. Head 0.6 cm collected into terminal
spiciform dense cymcs. Heads heterogamous. Corolla purple.
16. Blumea oxyodonla 3C.(Fig. 16a)
Low herb with several ascending branches. Leaves radical and cauline,
thinly scabrid above, softly pubescent below, capitula paniculate,
shortly stalked,bisex~lal florets, corolla yellow.
17. SphaeMntksrs indicus Idinn. (Fig. l7a)
The plant is an annual herb with spreading branch, leaves alternate,
toothed, decurrent on the stem. Heads small heterogamous not rayed.
Pink or purple flowe~s, collected together in close terminal globose
clusters.
18. Vicoa indira DC.(Fig.l8a)
Plant is a herb. Leav:s alternate, sessile, oblong, lanceolate. auricled at
base. Heads heterogmous, rayed, solitary, terminal or leaf opposed.
Yellow flowers.
19. Acanthospermum hispidum DC.(Fig. l9a)
Erect branched, h~spid-hairy plants. Leaves obovate, spathulate
palmately veined and alternate. Heads solitary sessile, yellow.
Involucral bracts (outer) ciliate. Achenes with many, hooked, lateral
spinules and two s might apical spines.
20. Bidenspilosa Linr .(Fig.20a)
The plant is a ver/ variable erect herb, leaves 3 fid - 3 foliate. Heads
on long stout peduncles very variable in length, with white rays.
Heads heterogamous
21. Cosmos b@innalus Cav. cv. Orange.(Fig.2la)
Tall herbs, to 80 cm high, leaves 2 - 4 pinnatisect. lobes entire,
glabrous, 7 - 8 1:m long. Peduncle long ligules orange red to yellow,
sometimes white; disc florets orange in colour. Grown in gardens,
occuning as escape ancl naturalizing in plains and upper ghats.
22. Cosmos bipinnaius Cav. cv. Yellow.(Fig.22a)
Erect herbs, leaves lorig, opposite, bipinnatisect, petioles upto 2.5 cm
long sheating at base, tleads terminal, Disc florets yellow.
23. Cosmos caudalur Kunth.(Fig.23a)
Erect annual herb 0.5 - 1.5 m tall. Leaves bipinnatisect or some what
tripinnatisect, pinnuk:~ opposite, capitulam solitary, axillary or
terminal, long stalked, heterogamous,wrolla purple.
24. Ecliptaprosiraia (Linl~) Linn.Vig.24a)
The plant is an annu:~l herb with small flowers. Leaves lanceolate,
oblong and strigose. Branches erect or prostrate. Leaves opposite.
Heads small, heterogarnous with white ray florets.
25. Golinsogaparvifora Cav. (Fig.25a)
The plant is a weak, erect glabrous herb, annual branched stem and
simple opposite leave:; bearing stalked clusters of flower heads in their
axils. The small flow.:r heads are formed mainly of yellow disc florets
with a few white ray florets.
a 1 9a Acuni/~o.iperrnum hupufvrn 20a Brdem pilaur, 2 l a. ( ' a m h ~ p i m ~ m cv. mange, 22a ( b s m bspinmw m. yellow, 23a C o s m unahtw, 24a lidlipru proIytrata. 2 k. ( ialiit.qa pnvflwq 26a Melampoth urn puludann. ?7a Purihenium hyteruphore.~. =
26. Melampodiumpaludosnr B.H.& K. (Fig.26a)
This is an annual herb with profuse dichotomously branched stem.
The leaves are opposite: and ovate. Flower heads are star-like, lemon
yellow in colour with a raised orange to brownish central disc.
27. Parfhenium hysteropht>rus Linn.(Fig,27a)
The plant is a herb. 1.0 rn in height, stem long; tridinally grooved,
leaves irregularly dissrzted, head is corymbose head; homogarnous
white flowers.
28. Sigesbeckia oriental& Linn.(Fig.28a)
A large annual herb ~ i t h yellow flowers and large ovate-triangular
deeply cut leaves, the lower heads glandular and very sticky, adhering
to the clothing. Leav:s opposite, toothed, shortly petioled. Heads
heterogamous, rayed.
29. Spilanthes caba Dc.(I:ig.29a)
The plant is a herb, occurring through out the greater parts of lndia.
Stem erect or decumbent at base, leaves opposite, ovate-lanceolate,
dentate or almost entire, florets yellow in solitary or sub solitary long
peduncled conical heads. Flowers homogarnous.
30. Spilanthes ciliata H.6.K (Fig.30a)
(S. acmella Murr.)
Diffuse herbs rooting at lower nodes. Stem terete. Leaves to 7 x 4 cm
ovate, base rounded cr sub cordate margins serrate, apex acute, petiole
1 - 2.5 cm long. Heads rayed axillary, usually solitary, rarely 2 -3 in
each axil, turning cortical, yellow, peduncle 3 -8 cm long. lnvolucral
bracts 2 seriate shorter than ray florets.
3 1. Spilunthes radicans Jacq.(Fig.3 I a)
Erect herb, stem minutely pubescent. Leves to 7 x 4 cm ovate, base
obtuse, margins faimly serrate or entire, apex acute, petiole 1 -2 cm
long. Heads axillary, solitary 5 - 8 mm. across discoid, white,
peduncle 4 - 7 cm lor~g.
32. Spilanthes uliginosa Sw.(Fig.32a)
This plant is a very common creeping herbaceous weed. The flower
heads are conical and solitary on peduncles, yellow in colour,
heterogamous ray florets few.
33. Synedrella nodifrora, Gaertn.(Fig.33a)
The plant is an erect dichotomously branched herb, stem and branches
terette, glabrous, leaves ovate lanceolate, shortly petioled serrate,
scaberulous, 3 nerved. Heads sessile, axillary and terminal. Heads
small heterogamous, flowers yellow.
34. Tithonia diversifolia ,\.Gray.(Fig.34a)
The plant is a large shrub. Leaves petiolate and dissected (palmately
compound). Heads large, heterogamous and rayed, yellow.
35. Tridaxprocumbens Linn (Fig.35a)
The plant is a weak ztraggling herb 30 - 60 cm long with few leaves
2.5 - 5 cm long and very long slender solitary peduncles 25 cm long
and more. Head 2' cm diameter. Heads very long peduncled
heterogamous ,rayed, ray flowers white.
36. Wedelia chinensis (Osbeck) Merrill (Fig.36a)
(W. Calendulacea Le:;s.)
It is a climbing shn~b. Leaves linear-oblong or oblanceolate, sub
sessile, entire, roughly scabrous, heads solitary on slender axillary
peduncles 5 - 12.5 1:m long. Head heterogamous with yellow ray
flowers.
37. Wedelia frilobata (Lirn) Hitch.(Fig.37a)
It is a climbing s h r ~ b scabrid pubescent or hirsute herbs or under
shrubs, leaves opposite, trilobed often triple nerved. Heads
heterogamous rayed, axillary or terminal, radiate, yellow flowers.
38. Zinnia elegans Jacq.(Fig.38a)
Erect herbs. Leaves :~.5 - 10 x 1.5 - 5 cm, opposite, ovate, elliptic
ovate or ovate - oblorg, scahrid obtuse or subcute, 5 nerved from the
base. Heads 5 - 8 cm cross, pink, terminal, solitary.
39. Tagetes ereda Linn cv. orange.(Fig.39a)
A stout branching herb, 60 cm tall, leaves strong scented, pinnately
dissected segments 1 - 5 cm long oblong or lanceolate, serrate, flower
heads solitary, orange in colour,rays many long clawed.
40. Tagetes erecta Linn cv pale yellow (Fig.40a)
Stout herb, tall, native to Maxico extensively cultivated in gardens all
over India. Flower heads solitary .Flower heads yellow in colour.
41 Tagetes ereda Linn I:V Yellow. (Fig.4la)
Leaves strong scented, pinnalely dissected. Flower heads solitary
yellow in colour.
42. Tagetespatula Linn. (Fig.42a)
Sub shrub to lm. Stem dark red in colour .leaves aromatic, alternate,
pinnate, capitulam solitary, terminal or axillary, heterogamous, corolla
orange red in colour.
* - 37a Wedeliu rriiobuiu, 38a. Z~nniu elegans. 39a Tageles erecia cv.orunge, @a. 7agefe.s ereom cv. polf yellow. 41a. lagetes erect0 c v . ~ Y e l l ~ ~ , 42a. , I , ugeies piiltdu, 43a. ('hrysanthernum pur~henium, 44a. ('&.r.r~~'e~haIurn
crepIiliide,s. 4 5a. Enrilia sonch$ioIiu.
43. Chrysanthemum parthenium (Linn) Benth.(Fig.43a)
This medium high, densely branched and leafy plant attracts attention
by its distinctive aroma. The stems are erect, bearing many soft much
dissected leaves and t-rminated by clusters of flower heads. These
are small with yellow tubular florets in the center and white ray florets
around the outside.
44. Crossocephalum cwpidioides (Benth) S. Moore.(Fig.44a)
Stout herb to 1.5 rn, branchlets striate, brittle, leaves ovate to
oblanceolate (lower part pinnatipartite) base attenuate to decurrent,
margin coarsely dentate, petiole to 8 cm, capitula, 1.5 crn across.
Florets brick red.
45. E& sonchifolia (],inn) Dc.(Fig.45a)
The plant is a small herb, stems and leaves soft, fistular, glauceous,
glabrous or nearly so, the leaves lyrate pinnatifed with large terminal
lobe, upto 10 cm lsmg the basal leaves petioled cauline, accurately
auricled corolla lobes very short. Head small. Homogenous.
46. Notoniagrandiflru Dc.(Fig.46a)
Plant is a fleshy shrub reaching 1.5 m in height, with pale yellow
flowers, turning green. Leaves obovate or oblanceolate or sub
orbicular, obtuse, vwiable in size but some times reaching 17 cm long
and 8 cm broad, quite entire glacious green heads 2.0 cm - 3.0 cm
long. Head large, homogamous not rayed all bisexual in long
peduncled corymbs.
47. Sonchus oleraceus Lilm.(FigA7a)
It is an annual plant with an upright hollow stem. Leaves radical in
young plant, in mature plants they are cauline, exstipulate, sessile and
auriculate. Inflorescer~ce bearing several umbellate heads which are
homogamous, flowm are yellow in colour.
Table 1: List of members investigated
Serial Name o f taxa Locality of collection Altitude in No. meter (approx)
1 Elephantopus scaber Neriyamangalam 65
2 PhyUocephalum rangacharii Malliyankara Sea level
3 Vernonia cinerea Kochi Sea level
4 Adenostemma lavenia Malliyankara
5 Ageratum conyzoides Kochi
Ageratum haustonianum Munnar
Chrodaena odoraia
Eupatorium triplinewe
Mirkonia cordata
Conyza bonariensis
Conyza canadensis
Dichrocephala ch~ysanfht~m~olia
Erigeron mucronatus
Blumea lacera
Blumea mollis
Blumea oxyodonta
Sphaeranthus indicus
Vicoa indica
Acanihospermum hispidum
Bidens pilosa
Kochi
Aluva
Kochi
Kochi
Neriyamangalam
Munnar
Munnar
Kochi
Kochi
Kochi
Thrissur
Thrissur
Vandanam
Munnar
2 1 Cosmos br$innatus cv. orange Kochi
22 Cosmos bipinnatus cv. yellow Kochi
Sea level
Sea level
I050
Sea level
10
Sea level
Sea level
65
I050
1050
Sea level
Sea level
Sea level
80
80
Sea level
1050
Sea level
Sea level
Cosmos caudatus
Eclipta prostrata
Galinsoga pawiflora
Melampodium paludosm
Parthenium hysterophorus
Sigesbeckia orientalis
Spilanthes calva
Spilanthes ciliata
Spilanthes mdicans
Spilanthes uliginosa
Synedrella nodiflora
Tithonia diversifolia
Tridax procumbens
Wedelia chinensis
Wedeli0 trilobata
Zinnia elegans
Tagetes erecta cv. orantre
Tagetes erecta cv. pale e ell ow
Tagetes erecta cv. yell0 u
Tagetes patula
Chrysanthemum parthenium
Crassocephalum crepidioides
Emdia sonchifolia
Notonia grandifora
Sonchus oleraceus
Kochi
Kochi
Munnar
Kochi
Alapuzha
Munnar
Munnar
Kochi
Neriyamangalam
Malliyankara
Kochi
Munnar
Kochi
Angamali
Kochi
Munnar
Kochi
Kochi
Kochi
Munnar
Munnar
Kochi
Kochi
Kochi
Munnar
Sea level
Sea level
I050
Sea level
Sea level
1050
1050
Sea level
65
Sea level
Sea level
1050
Sea level
Sea level
Sea level
1050
Sea level
Sea level
Sea level
1050
1050
Sea level
Sea level
Sea level
1050
METHODS
a) Mitotic squash experiments
The cytology of Forty s e v e ~ species belonging to thirty four genera from
South India was investigated with the help of improved cytotechniques
(Sharma and Sharma 1980). Squash experiments were carried out on root tip
meristem at mitotic metaphase stage. The root tips are collected from the
plants of various species planted in the experimental botanical garden at the
periods showing peak mitotic frequency, i.e, 9.00 a.m. to 1 1 a.m. The root tips
were pretreated with saturatecl solution of para dichloro benzene with traces of
aesculin for 3 hours. It is fcund to be most suitable for many members of
Asteraceae. Eventhough 0.0(14 M 8 - hydroxy quinoline at 18 to 20°c for 2
hours is also used for some genera like Notonia . The root tips immersed in
cytostatic chemicals are initially chilled at 0-5% for 4 - 7 minutes and then
kept at 12 - 20% for I - 3 hours for obtaining best results.
The pretreated root tips are then washed thoroughly with distilled water and
fixed in 1 : 3 acetic acid - ethyl alcohol mixture (Carnoy, 1886) overnight,
followed by 3 - 7 minutes reatment in 45% acetic acid. Root tips were
hydrolysed in IN Hcl at 60% for 5 - 7 minutes and squash preparations were
made in 1% acetoorcein (Shanna and Sharma, 1980). The apical 0.5 - 1.0 mm
root tips were cut and placed on slide, squashed gently with a needle in 45%
acetic acid and covered with cover glass. The preparation was temporarily
sealed.
b) Meiotic smear experiments.
Pollen mother cell (PMC) analysis were carried out on those members which
bloomed frequently in the experimental garden. Young flower buds of 10 - 12
days old were picked b e t w ~ ~ n 9 a.m. and I 1 a.m. each day and fixed in
Carnoy's fluid (6:3:1 Ethanol, Acetic acid, and chloroform) for 12 hours. The
next day anthen were washed three times in distilled water and stored in 70%
Ethyl alcohol. Squash preparations were made in 1% aceto carmine (Shanna
and Shanna 1980).
c) Detailed karyomorpbologieal studies by image analysis system.
Pbotogrnphs and image processing
Well spread metaphase plates were photographed using 125 ASA 35 mm
Orwo film in a Lei& photc~graphic attachment and suitably enlarged. A 200
dpi scanner scans each original photograph. The software Adobe Photoshop
was used for digitalizing and reproduction. The contrast of each image has
been increased by raising the resolution upto a satisfactory level. Acquisition
of quantitative data from large number of plant chromosomes and also semi
automatic karyotyping can be easily carried out by using image analysis
system. The generated mages were checked by the visual inspection
comparing with the original photo micrographs. After storage of original
digital images of the metaphase spreads in floppy discs, these images were
analyzed by using compllter devices. The image was recovered from the
floppy disc and the original digital image for the analysis was then generated
automatically. A binary imabe that was essential for the object identification
by the computer was generated by interactive setting of the lower and upper
thresholds of the gray levels These thresholds were defined properly so that
the gray values of the pixels that consisted of chromosome images were
included. Binarization was iutomatically carried out by changing the pixels
values stood between the two thresholds to white and all other to black. The
large background dust particles and spots whose gray values also fell between
the two thresholds were elin~inate by adjusting the gray values of the pixels
outside the chromosomal region by interactive setting of lower and upper
thresholds of the gray levels. This will result in a binary chromosomal image
with a clean background.
Quantitative karyotyping of the chromosomes.
Measurement of each chroinosorne from enhanced image were made by
AutoCAD (Version2000) loaded on a personal computer. This image was
automatically coloured differently by the computer generated colouration
based on the actual gray values of the pixels. Pseudocolouration considerably
improved the density distribution of the objects and the recognition by
humans. This will help to detect the primary and secondary constriction of the
chromosomes. These constrictions were namely marked by the overlay
straight lines on the pseudocoloured images. After this midrib lines were
drawn interactively on eact chromosome. Extraction of the midrib lines,
breakage at the position of the constriction and identification by different
Fig. 1
Fig. 1-9: Different steps of karyotype analysis by the image analysis systems. 1 - metaphe plate, 2 - binarization, 3 - background cleaning, 4 - metaphase plate with clear background, 5 - selsction of a chromosome for measurement, 6 - enhanced image with marking in the constriction, 7 - drawing marginal line, 8 - differentiation by applying various colours and 9 - midrib lines.
colours were subsequently carried out automatically. The outer margin of each
chromosome image also marked t!y drawing surrounding line. Numerical data
such as the arm length, area, perimeter and an apparent visual three
dimensional volume of each chromosomes were obtained in pixel units. In all
the karyotypes, ratio of the short arm to the total length of the chromosome in
percentage, Forma percentage or centromeric index (F%) is determined after
Krikorian et a1 (1983). On the bmis of F% the nature of primary constriction
in the chromosomes are classifietl as follows:
Nature of primary constriction
Mc:dian
Ntarly median
N~:arly sub median
Sub median
Nearly sub median
Nearly sub terminal
Sub terminal
Irlearly sub terminal
E.xtremely sub terminal
Terminal
The values obtained for the ch~.omosome morphology were:
Total arm length of each chromosome (long arm length + short arm length)
and relationship of the arm (R3)
RB is calculated by the following formula
RB = long arm length 1 short arm length
Arm ratio (AR) has been widely utilized for the classification of chromosome
types (Leven et a1.1964) has been considered empirically to be a more stable
parameter of the chromosomal morphology. The AR was defined as the ratio
of the length of the short arm to that of the long arm (SiL) for each
chromosome.
The following indices were also calculated for each chromosome is
determined after Watanabe et d. (1 990)
2ai Relative long arm length (RLI,) = 2n
Z (aj+ bj) j=l
2bi Relative short arm length (RS L) = -57-
1 (aj+ bj) j=l
Relative chmosome length (RL) = RLL +RSL
Arm difference ratio (AD) = (ai - bi)/ (ai + bi)
where ai = long arm length o Fchromosome i, bi = short arm length of chromosome i, and
2n
C (aj+ bj) =total diploid chromosome length j=1
The disparity index (DI) of chromosome in a karyofype is calculated after Mohanty et a1.(1991) by the f3rmula:
longest chromosome - shortest chromosome DI = x 100
longest chromosome + shortest chromosome
The variation coefficient among chromosome complements (VC) is determined after Verma (1980) as follows:
standard deviation vc = - x 100
mean lengths of chromosome
The total forma percentage or the mean centromeric index value (TF%) is calculated in each taxa after Husiwara (1962), by the formula.
Total sum of short arm length TF% = x 100
Total sum of chromosome length
The uniformity coefficient ([erimeterlarea) (Ojeda and Torres, 1996) were
also calculated. All the numerical data are prepared after comparing at least
five well spread metaphase plates. The various calculations were done by the
computer package Microsoft Excel
Quantitative idiograms of chromosomes
Based on the data relating to the length, the idiograms were presented
combined with the results of quantitative image analysis of chromosomes.
The chromosomes were arranged semi automatically according to the length,
arm ratio, uniformity coefficient, three-dimensional volume and idiograms
generated with the aid of computer software Adobe Photoshop.
OBSERVATIONS
In the present investigation 47 taxa of 34 genera were analysed. The normal
somatic chromosome number ranges from2n = 10 to 2n = 78. However,
numerical variations are previilent in some taxa. The ploidy level exhibited by
different taxa ranges from diploidy to hexaploidy. The chromosome pairs
with secondary constriction are found to range from one to four. The
karyotypes are characterised by comparatively small chromosomes ranging
from 0.41 pm to 2.54 pm in length. The chromosomes in each karyotype
decreased in size progressiv~:ly and they had nearly median to nearly sub
median primary constriction.
The general description of the common chromosome types is given below
followed by the karyotype deszription of each of the members investigated.
Type A: Comparatively long chromosomes with two constrictions, one median
to nearly median and the other nearly sub median.
Type B: Relatively long chron~osomes p0.8) with nearly median to nearly sub
median primary constriction.
Type C: Small chromosomes (<0.8) with nearly median to nearly sub median
ptimary constriction.
Figs. I b, l e - Elephanropus scuber : I b - mitotic rnetaphase (2n=22), le - meiotic ~netaphase (n=I I ) ; 2b.2e - Phyliocephalum rangacharii : 2b - mitotic metaphase ( 2 ~ 1 8 ) . 2e - meiotic metaphase (n=9); 3b,3e - Vernonia cinerea : 3b - mitotic rnetaphase (2n=18), 3e - meiotic metaphase (n=9); 4b - Adenostemma luvenia mitotic metaphase (2n=20); 5 b . 5 ~ - Ageralum conyzoides : 5b - mitotic metaphase (2n=40). 5c - somatic variant ( 2 ~ 3 0 ) ; 6b - Ageruturn hausronianurn mitotic metaphase (2n=40); 7b Chromolaena odorara mitotic metaphase ( 2 ~ 6 0 ) ; 8b - Eupatorium lriplinerve mitotic metaphase (2n=50); 9b,9c,9e - Mikania cordara: 9b - mitotic rnetaphase (2n=36), 9c - somatic variant (2n=34); 9e - meiotic metaphase (n=18); lob - Conyza bonariensis mitotic metaphase ( 2 n ~ 5 4 ) . Bar represents 5pm each
Elephantopus scaber
Nomal somatic chromosome number
Karyotype formula
Number of chromosome pairs with secondary constriction
Range of chromosome length in pm
Total chromosome length in prn
Average chromosome length in pm
T.F. value (%)
Variation coefficient (VC)
Disparity index (Dl)
Total volume of chromosomes in umf
2n = 22 (Fig. I b) A2 86 C14 1
1.1 1-0.63 17.88 0.81 45.74 18.75 27.58 0.72
Table 2: Detailed karyotype analysis of Nephanlopus scaber
Total Short arm
Type length length F in pm in um
Nearly sub median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Normal somatic chromosome number
Karyotype formula
Number of chromosome pain with semndarr constridon
Range of chromosome length in pm
Total chromosome length in urn
Average chromosome length in pm
T.F. value (%)
Variation coefficient (VC) Disparity index (D.1)
Total volume of chromosomes in pm'
Table 3 : Detailed karyolype analysis of Ph!fllocephalum rangachsri
Short Total arm Volume p,A
A? 4D RE F % inpm, Nature of primary
Type length length RL ratto constriction in pm in pm
A 1.14 0.37 0.11 0.84 0.09 1.19 32.84 0.0210 2.64 Nearlysubmedian 0.32
A 1.00 0.37 0.10 0.99 0.01 1.01 37.17 0.0329 2.99 Nearlysubmedian 0.25
B 0.82 0.38 0.11 0.87 0.07 1.15 46.41 0.0229 3.54 Nearly median
B 0.81 0.35 0.11 0.78 0.13 1.29 43.73 0.0199 3.01 Nearly median
C 0.77 0.36 0.10 0.88 0.06 1.14 46.80 0.0195 3.02 Nearly median
C 0.76 0.34 0.10 0.81 0.11 1.24 44.85 0.0106 3.87 Nearly median
c 0.73 0.35 0.10 0.90 0.05 1.12 47.28 0.0162 2.82 Nearly median
C 0.72 0.35 0.10 0.98 0.01 1.02 49.46 0.0199 2.94 Nearly median
c 0.67 0.31 0.09 0.88 0.06 1.14 46.78 0.0130 3.74 Nearly median
Vernonla cinerea
Normal somatic chromosome number
Karyolype formula
Number of chromosome pairs with seo~ndary constriction
Range of chromosome length in pm
Total chromosome length in pm
Averege chrommme lmgth in pm
T.F. value (%)
Variation coefficient OJC) Disparity index (D.1)
~ o t a l volume of chromosomes in pm'
Table 4 : Detailed karyolype analysis of Vemonia cinerea
Short Total a, Volume P,A Nature of primary
TYW length lensul Arm AD RB F % in pm, RL ratlo consttiction
in pm . ~n pm
A 1.29 0.47 0.11 034 0.03 1.07 36.09 0.0505 2.66 Nearlysubmedian 0.33
Nearly sub median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Normal somatic chromosome number
Karyotype formula
Number of chromosome pairs with secondact
Range of chromosome length in pm
Total chromosome length in vm
Average chromosome length in Urn
T.F. value (%)
Variation coefficient (VC)
Disparrty index (D.1)
Total volume of chromosomes in um'
2n = 20 (Fig.4b)
A2 818
constriction 1
1.53 - 0.92
23.02
1.15
44.39
35.1
24.89
1.4
Table 5: Detailed karyotype analysis of Adem,siemma lavenia
Shod Total am
Type length Volume RL Am AD RE F% In pm, PIA Nature of primary length ratlo constriction
In pm ln urn
A 1.53 0.59 0.10 1.00 0.30 l .W 38.67 0.0681 3.02 Nearly median 0.35
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Neatly median
Nearly median
Normal somatic chromosome number
Kalyotype formula
Number of chromosome pairs with sewndarl constriction
Range of chromosome length in pm
Total chromosome length in pm
Average chromosome length in pm
T.F. value (%)
Variation coefficient (VC)
Disparity index (Dl)
Total volume of chromosomes in pm'
Table 6 : Detailed karyotype analysis of AgJemtum wnyzoides
Short Total arm Arm Volume
4D RB F% inpm3 P/A Nature of primary Type length length in pm RL ratio constridion
in pm
Nearly submedian
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Ageratum haustonfanurn
Normal somatlc chromosome number
Karyotype formula
Number of chromosome pairs with secondary constriction
Range of chromosome length in pm
Total chromosome length in pm
Average chromosome length in pm
T.F. value (%)
Variation coefficient (VC)
Disparity index (D.1)
Total volume of chromosomes in pm'
Table 7 : Detailed karyoptype analysis of Ageratum haustonianum
Short Total arm
Type length length in pm
in um
Arm ratio
Volume F96 in ~ m '
Nature of primary constriction
Nearly sub median
Nearly sub median
Nearly median
Nearly median
Nearly median
Nearly median
Median
Nearly median
Median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Chnamolaena odorab
Normal somatic chromosome number
Karyotype formula
Number of chromosome pairs with secondary ans strict ion
Range of chromosome length in IJm
Total chromosome length in pm
Average chromosome length in pm
T.F. value (%)
Variation coefficient (VC)
Disparity index (Dl)
Total volume of chromosomes in urn'
Table 8: Detailed karyotype analysis of Chmmolaena d r a b
Short Total am Volume Arm /rD RE F% in pm, PIA
Nature of prirnaly Type length length RL rabo conshidion
in pm in pm
Neatly sub median
Nearly sub median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Median
Median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Neatly median
Nearly median
Nearly median
Nearly sub median
Nearly median
Neatly median
Neatly median
Nearly rnedtan
Nearly median
Normal somatic chromosome number
Karyotype formula
Number of chromosome pairs with secondary constriction
Range of chromosome length in pm
Total chromosome length in pm
Average chromosome length in vm
T.F. value (%)
Variation coethclent (VC)
Disparity index (D.1)
Total volume of chromosomes in pm'
Table 9 : Detailed karyolype analysis of Eupaiorium triplinerw,
Total Type length
in pm
arm Arm -- . Volume Nature of orimam length RL rat In pm
0.58 0.04 0.93 0.03 1.07 34.97 0.1642 3.51 Nearly sub median 0.46
0.55 0.04 0.95 0.(12 1.05 34.38 0.2- 2.95 Nearlysubmedian 0.48
0.46 0.04 0.67 0.;0 1.49 29.22 0.0823 3.46 Nearly sub median 0 43
0.54 0.04 0.82 0.10 1.22 45.w 0.0531 3.25 Nearly median
0.58 0.04 0.97 0.~2 1.03 49.20 0.0737 3.98 Nearly median
0.57 0.04 0.95 0.c3 1.05 48.72 0.1647 3.41 Nearly median
0.51 0.04 0.83 0.C9 1.21 45.27 0.1425 3.30 Neally median
0.47 0.04 0.71 0.17 1.41 41.55 0.0931 3.24 Nearly median
0.55 0.04 0.97 0.C1 1.03 49.35 0.0809 3.32 Nearly median
0.52 0.04 0.88 0.C6 1.13 46.91 0.0556 3.73 Nearly median
0.54 0.04 0.99 OC 1 1.01 49.65 0.0808 3.59 Nearly median
0.54 0.04 0.98 0.C1 1.02 49.53 0.0470 3.75 Nearly median
0.53 0.04 0.97 0.01 1.03 49.33 0.0931 3.53 Nearly median
0.41 0.04 0.66 0.20 1.51 39.85 0.0446 3.33 Nearly median
0.45 0.04 0.80 0.11 1.25 44.54 0.0686 3.96 Nearly median
0.49 0.04 0.98 0.01 1.02 49.46 0.0578 3.68 Nearly median
0.49 0.04 0.98 0.01 1.02 49.42 0.0868 3.79 Nearly median
0.48 0.04 0.95 0.03 1.06 48.64 0.0288 3.91 Nearly median
0.48 0.04 0.99 0.01 1.01 49.71 0.0389 3.47 Nearly median
B 0.95 0.45 0.03 0.89 0.16 1.12 47.07 0.0424 4.10 Nearly median
B 094 0.45 0.03 0.94 0.03 1.06 48.46 0.0402 3.63 Nearly median
B 0.90 0.45 0.03 0.97 0.01 1.03 49.31 0.1190 3.20 Nearly median
B 0.90 0.44 0.03 0.97 0.02 1.03 49.21 0.0766 3.52 Nearly median
B 0.89 0.44 0.03 0.98 0.11 1.02 49.42 0.0625 3.24 Nearly median
B 0.66 0.42 0.03 0.93 0.13 1.07 46.28 0.0449 3.93 Nearly median
Normal somatic chromosome number
Karyotype formula
Number of chromosome pairs with secondary constriciion
Range of chromosome length in pm
Total chromosome length in pm
Average chromosome length in pm
T.F. value (%)
Variation coefficient (VC)
Disparity index (Dl)
Total volume of chromosomes in pm3
Table 10 : Detailed karyotype analysis of Mnrania &at3
Total Shoe arm Volume p/A Nature of primary
Type length length RL A" ratlo AD RE F% inpm, constriction in pm in pm
Volume p/A Nature of primary A" AD RE F% inpm, ratlo constriction
Nearly sub median
Nearly sub median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Normal somatic chromosome
Karyotype formula
number
Number of chromosome paws with secondaw mnstritimn
Range of chromosome length in pm
Toial chromosome length in Vm
Average chromosome length in pm
T.F. value (%)
Variation d c i e n t (VC)
Disparity index (D.1)
Total volume of chromosomes in urn'
Table 11 : Detailed karyolype analysis of Cor~yza bonariensis
Nearly median
Narrly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly M a n
Median
Nearly median
Nearly median
Nearly median
Nearly median
early mediin
Nearly median
6 1.08 0.52 0.03 0.94 0.03 1.07 48.38 0.0298 3.45 Nearly mediin
6 1.07 0.51 0.03 0.92 0.04 1.08 48.02 0.0333 3.03 Nearly median
6 1.06 0.48 0.03 0.83 0.09 1.21 45.32 0.0986 2.82 Nearly median
6 105 0.50 0.03 0.90 0.05 1.11 47.44 0.0237 3.51 Nearly median
8 1.04 0.46 0.03 0.79 0.11 1.26 44.28 0.0361 3.70 Nearly median
6 0.96 0.47 0.03 0.97 0.01 1.03 49.29 0.1114 3.03 Nearly median
I l b 12b 13b - 14b -
bigs. I 1 b, l le - CO~IJ.;(J canadensis: 1 l b - mitotic metaphasc (2n = 18). I le - mciotic mctaphase (2n = 9); I2b - Dichrocephula chrysanrhetnifolia mitotic metaphase (2n = 18); 13b - Erigeron mucronatus mitotic metaphase (2n = 18); 14b, l4e -Blumeu iacera; 14b - mitotic metaphase (2n = 36), 14e - meiotic metaphase (n = 18); 15b, I Sc, 15e - Blumea mollis: 15b - mitotic metaphase (2n - 18); 15c . somatic variant (2n = I6),15e - meiotic metaphase (n = 9); 16b - Blurnea oxyodonra milotic meraphse (2n : 28); 17b,17c,I 7c1 - Sphaeranrhus indicus: 17b - mitotic metaphase (2n = 201, 17c - somatic variant (2n = 1 X), 17c1 - somatic variant (2n = 16); 18b,l8e - Vicoa indira: 18b - mitotic metaphase (2n - 18), 18e - meiotic rnetaphase (n = 9); 19b - Auanthospermum hispidum mitotic mctaphase (2n = 22).
Bar represents 5prn each
Normal somatic chromosome number
Karyolype formula
Number of chromosome pairs with secondary constriction
Range of chromosome length in Vm
~ o t a l chm- length in pm
Average c h m m m length in pm
T.F. value (%)
Variation mefiicient (VC)
Disparity index (D.1)
Total volume of chromosomes in pm'
Table 12 : Detailed karyotype analysis of Cmy2a canedensis
Short arm Type length
t e r n RI
inpm in,,",
VOlUrn P,A A? AD RB F% in(rm$ ~a tum of primary
- rabo consbidion
A 1.52 0.43 0.12 0.64 0.22 1.56 28.18 0.1210 2.80 Nearlysubmedian 0.43
€3 1.05 0.45 0.12 0.73 0.15 1.36 42.31 0.0346 2.85 Neariy median
B 1.04 0.50 0.12 0.95 0.03 1.06 48.64 0.0487 3.13 Nearly median
B 0.98 0.48 0.11 0.97 0.02 1.03 49.22 0.0713 2.85 Nearly median
B 0.95 0.42 0.11 0.80 0.11 1.24 61.58 0.0462 2.93 Nearly median
B 0.92 0.46 0.10 0.98 0.01 1.02 49.50 0.0719 2.99 Nearly median
B 0.68 0.43 0.10 0.98 0.02 1.04 49.00 0.0522 2.95 Nearly median
B 0.84 0.39 0.09 0.86 0.08 1.16 46.21 0.0469 3.23 Nearly median
C 0.77 0.35 0.09 0.81 0.10 1.23 44.87 0.0581 3.42 Nearly median
Normal somatic chromosome number
Karyotype formula
Number of chromosome pairs with secondary constriction :
Range of chmmwome length in pm
Total chromosome length in pm
Average chromosome length in pm
T.F. value (YO) Variation coefficient (VC)
Disparity index (0.1)
Total volume of chromosomes in pm'
Table 13 : Detailed karyotype analysis of Dichmphala chrysanthemifola
Short Total arm Arm Volume p,A Nature of primary
Type length length RL ratio AD RB F% in pmS in pm constridion
in pm
Nearly sub median
Nearly median
Nearly median
Nearly median
Nearly median
Neariy median
Neariy median
Nearly median
Nearly median
Normal somatic chmmosome numbel
Kafyotype formula
Number of chromosome pairs with sewndary wnstriction
Range of chromosome length in prn
Total chromosome length in pm
Average chromosome length in pm
T.F. value (%)
Variation coefficient (VC)
Disparity index (0.1)
Total volume of chromosomes in urn'
Table 14 : Detailed karyotype analysis of Efigemn mucmnatus
Short Total arm Volume P,A Nature of primary
Type length length RL :z AD RB F% constriction inpm . In pm
A 1.46 0.52 0.13 0.82 0.10 1.21 35.78 0.0886 3.63 Nearlysubmedian 0.31
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearty median
Nonnal somatic chmmosome number
Karyolypa formula
Number of chromosome pairs with sewndary conitridion :
Range of chromosome length in pm
Total chromoMnmt length in pm
Average chromoawmt length in pm
T.F. value (%)
Variation coeffident (VC)
Disparity index (D.1)
Total volume of chromosomes in pm'
2n = 36 (Fig. 14b)
A4 632
2
2.07 - 0.88
43.88
1.22
42.9
25.86
40.33
1.82
Table 15 : h i l e d karyotyps analysis of B I u m lacera
Short Total arm
T Y P ~ ienpth RL AD RB F% P,A Nature of primary
in pm Fngm in pma w n s m o n In pm
A 2.07 0.88 0.07 0.80 0.11 1.25 31.85 0.1069 2.66 Nearly sub median 0.58
A 1.91 0.59 0.07 0.63 0.23 1.59 31.19 0.0410 2.89 Nearlywbmedian 0.37
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Normal somatic chromosome number
Karyotype formula
Number of chromosome pairs with secondary constriction : Range of chromosome length in pm
Total chromosome length in pm
Average chromosome length in pm
T.F. value (%)
Variation M c i e n t (VC)
Disparily index (0.1)
Total volume of chromosomes in Mm'
Table 16 : Detailed kalyotype analysis of Bfumea mollis
Short Total arm Arm Volume p,A Nature of primary Type 1- length RL ratio .\D RB FSb in pm, consbidion
in lrm in pm
Nearly medin
Nearly median
Nearly median
Nearly median
Nearly median
Neady median
Nearly median
Nearly median
Nearly median
Normal somatic chromosome number
Karyotype formula
Number of chromosome paws with secondary constriction
Range of chromosome length in pm
Total chromosome length in pm
Average chromosome length in pm
T.F. value (%)
Variation coefficient O/C)
Disparity index (D.1)
Total volume of chromosomes in pm'
Table 17 : Detailed karyolype analysis of Blumea oxyodonla
~~~
Short Total arm Typm length Volume
Arm 4 RB F% in pm3 P/A Nature of primary lensth inpm . RL ratio constriction m pm
A 2.17 0.79 0.13 0.70 (. I8 1.43 36.25 0.0533 1.84 Nearlysubmedien 0.26
Nearly sub median
Nearly sub median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Normal somatic chromosome number
Karyotype formula
Number d chromosome pairs with samndary mnstriction
Range of chromosome length in pm
Total chromosome length in pm
Average chromosome lenglh in pm
T.F. value (%)
Variation d l d e n t PC)
Disparity index (0.1)
Total volume of chromosomes in vm'
Table 18 : Detailed kary- analysis of !:phaemnfhus indicus
2n = 20 (Fig. 17b)
A4 616
2
1.90 - 0.91 26.16
1.31
42.66
21.04
35.23
1.76
Short Total arm Volume p,A Nature of primary
T W length length RL :E AD RE F% inpmS consiridon in vm in pm
A 1.90 0.69 0.11 0.98 0.01 1.02 36.23 0.1466 2.36 Nearlysubmedian
B 1.42 0.56 0.11 0.65 0.21 1.55 39.25 0.0965 2.54 Nearly median
B 1.37 0.63 0.10 0.85 0.08 1.18 45.98 0.0701 3.00 Nearly median
B 1.30 0.63 0.10 0.93 0.04 1.07 48.23 0.0846 2.93 Nearly median
B 1.25 0.55 0.10 0.78 0.12 1.28 43.77 0.1106 2.63 Needy median
B 1.16 0.51 0.09 0.79 0.12 1.27 44.04 0.1111 3.05 early median
8 1.06 0.48 0.08 0.83 0.09 1.21 45.31 0.0980 2.14 Nearly median
B 1.08 0.52 0.08 0.93 0.04 1.08 48.16 0.0567 2.36 Nearly median
B 0.91 0.45 0.07 0.98 0.01 1.02 49.60 0.0483 2.60 Nearly median
Normal somatic chromosome number
Karyotype formula
Number of chromosome pairs with secondary constriction
Range of chromosome length in pm
Total chrmosmne length in pm
Averaga chromosome length in pm
T.F. value (%)
Variation we f fwn t (VC)
Disparity index (D.1)
Total volume of chromosomes in urn'
Table 19 : Detailed karyotype analysis of b i m e indica
Short Total arm volume p,A
A? AD RB F% inpm, Nature of primary Type length length RL ratlo wlgbiClion in um in
Nearly sub median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Normal somatic chromosome number
Karyolype formula
Number of chmmosome pairs with sewndory constriction
Range of chromosome length in pm
Total chromosome length in urn
Average chromosome length in pm
T.F. value (%)
Variation d ~ c i e n t WC)
Disparity index (D.1)
Total volume of chromosomes in pm'
Table 20 : Detailed karyotype analysis of ,\canUlospermum hispidurn
Short Total arm Volume p,A Nature of primary Type length length RL AD RB F% in pm. combiCtion
in pn In pm
A 1.71 0.56 0.10 0.82 0.10 1.21 32.69 0.0872 2.78 Nearlysubmedian 0.47
A 1.66 0.52 0.10 0.74 0.15 1.35 31.09 0.0923 3.32 Nearlysubmedian 0.45
A 1.63 0.53 0.10 0.75 0.14 1.33 32.44 0.0039 4.61 Nearlysubmedian 0.40
B 1.25 0.59 0.10 0.88 0.06 1.13 46.94 0.0912 2.46 Nearly median
B 1.09 0.52 0.09 0.92 0.04 1.09 47.78 0.0795 2.70 Nearly median
B 0.94 0.43 0.08 0.86 0.08 1.16 46.20 0.0412 3.75 Nearly median
B 0.92 0.46 0.07 0.99 0.01 1.01 49.71 0.0395 2.73 Needy median
B 0.88 0.40 0.07 0.85 0.08 1.18 45.85 0.0387 3.49 Neady median
B 0.85 0.39 0.07 0.83 0.09 1.20 45.39 0.0445 2.90 Nearly median
B 0.81 0.34 0.07 0.71 0.17 1.40 41.58 0.0202 3.78 Nearly median
C 0.72 0.33 0.06 0.85 0.08 1.18 45.90 0.0165 3.78 Nearly median
Figs.20b - Bidens pilosa mitotic metaphase (2n = 72); 2 I b,2 1c,2 l e,2ie1- Cosmos bipinnatus 0. orange: 21b - mitotic metaphase (2n = 241, 2 1c - somatic variant (2n = 22), 21 e - meiotic metaphase (n = 1 I), 2 1 el - meiotic metaphase (n = 12); 22b,22e - Cosmos bipinnah cv. yellow: 22b - mitotic metaphase (2n = 24), 22e - meiotic metaphase (n = 12); 23b - Cosmos caudatw mitotic rnetaphase (2n = 24); 24b,24c,24e - Eclipta prostrata: 24b - mitotic metaphase (2n = 22), 24c - somatic variant (2n = 18), 24e - meiotic metaphase (n = 1 I); 25b - Galinsoga pamiflora mitotic rnetaphase (2n = 32); 26b - Melampodium paiudosm mitotic metaphase (2n = 24); 27b - Parthenium hysrerophorus mitotic metaphse (2n = 36), 28b - Sigesbeckia orientalis mitotic metaphase (2n = 30); 29b - Spilanthes calva mitotic metaphase (2n = 78).
Bar represents 5 p - i each
Normal somatic chromosome number
Karyotype formula
Number of chromosome pairs with secondary constriction
Range of chromosome length in pm
Total chromosome length in pm
Average chromosome length in vm
T.F. value (%)
Variation coefficient (VC)
Disparity index (D.1) Total volume of drromosomes in pm'
Table 21 : Detailed kaiyolyp analysis of Bidens piloss
Short Total TYW length length RL krb", AD RB F% PIA Nature of primary
inpm . constridion m pm
A 1.30 0.47 0.03 0.78 0:12 1.28 38.15 0.0671 2.80 Nearly sub median 0.23
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Cosmos bipinnalvr cv. orange
Normal somatic chromosome number
Karyotype formula
Number of chromosome pain with secondary constriction
Range of chromosome length in pm
Total chromwome length in pm
Average chromosome length in pm
T.F. value (%)
Variation coefficient (VC)S
Disparity index (D.1)
Total volume of chromosomes in pm3
Table 22 : Detailed kalyotype analysis of Co::mos bipinnafus ccv. m g e
Short Total arm Volume P,A A? AD RB F% inpms Nature of primary
Type length length RL ratto constriction in pm in pm
A 1.63 0.44 0.09 0.58 026 1.72 27.24 0.1191 3.13 Nearly sub median 0.42
B 1.54 0.75 0.12 0.96 002 1.04 48.69 0.0881 3.09 Nearly median
B 1.20 0.48 0.09 0.67 0.20 1.49 40.08 0.0514 3.16 Nearly median
B 1.17 0.52 0.09 0.80 C . l l 1.25 44.47 0.1203 3.11 Nearly median
B 1.14 0.51 0.09 0.81 C . l l 1.24 44.73 0.0563 3.28 Nearly median
B 1.03 0.48 0.08 0.87 C.07 1.15 46.50 0.0237 4.28 Nearly median
B 0.99 0.48 0.08 0.95 C.03 1.06 48.65 0.0516 3.71 Nearly median
B 0.96 0.39 0.07 0.70 I 1.43 41.15 0.1027 3.72 Nearly median
B 0.95 0.49 0.07 1.06 -0.03 0.94 51.46 0.0255 3.67 Median
B 0.89 0.41 0.07 0.85 (1.08 1.18 45.87 0.0390 3.54 Nearly median
B 0.81 0.40 0.06 1.00 (1.00 1.M) 50.01 0.0615 3.72 Median
C 0.76 0.40 0.06 1.11 -tl.05 0.90 52.69 0.0371 3.87 Median
Cosmos blpinnatus cv. ydlow
Normal somatic chromosome number
Karyotype formula
Number of chromosome pairs with secondaly constriction
Range of chrornoaome length in pm
Total chromosome length in pm
Average chromcsome length in um
T.F. value (%)
Variation M c i e n t (VC)
Disparity index (D.1)
Total volume of chromosomes in pma
Table 23 : Detailed karyotype analysis of C a s m bipinnatus cv. yellow
Short Total arm volume P,A RL :z ,+D RB F% inurn. Nature of primary
Type length in pm
constriction in pm
A 1.75 0.60 0.08 0.95 0.02 1.05 34.11 0.0656 2.67 Nearlysubmedian 0.53
A 1.68 0.56 0.06 0.96 C.02 1.04 33.02 0.0804 2.83 Nearly submedian 0.55
A 1.66 0.63 0.08 0.98 C.01 1.02 37.74 0.1091 2.39 Neatly median 0.39
A 1.54 0.53 0.07 0.91 C.05 1.10 34.68 0.0504 2.84 Nearlysubmedian 0.42
B 1.36 0.67 0.09 0.94 C.03 1.06 48.56 0.3049 3.13 Nearly median
B 1.21 0.59 0.08 0.97 C.02 1.03 49.19 0.1054 2.41 Nearly median
B 1.12 0.52 0.07 0.66 (,.07 1.16 46.26 0.1563 2.80 Nearly median
B 0.93 0.46 0.06 0.97 ('.01 1.03 49.33 0.0690 2.60 Nearly median
B 0.62 0.36 0.05 0.77 (1.13 1.30 43.48 0.0436 2.62 Neariy median
C 0.79 0.39 0.05 0.99 (1.00 101 49.84 0.1299 2.56 Nearly median
c 0.72 0.34 0.05 0.91 (1.05 1.10 47.70 0.0192 3.02 Nearly median
C 0.70 0.32 0.05 0.67 (1.07 1.15 46.56 0.0237 3.26 Nearly median
Casmos caudatus
Normal somatic chromosome number
Karyotyp formula
Number of chromosome pairs with secondary constriction
Range of chromosome length in pm
Total chromosome length in pm
Average chromosome length in pm
T.F. value (%) Variation coefidant (VC)
Disparity index (D.1)
Total volume of chromosomes in pm'
Table 24 : Detailed karyotype analysis of Cosmos caudatus
Short Total am RL tz 4D RB F% PIA
Nature of primary T Y P ~ length l.ngm in pms constribion
in vm in pm
A 1.49 0.56 0.11 0.79 0.12 1.26 37.36 0.0011 2.83 Neariysub median 0.23
B 1.19 0.50 0.10 0.73 1.16 1.38 42.08 0.0362 2.84 Nearly median
B 1.07 0.51 0.09 0.89 1.06 1.12 47.16 0.0462 3.21 Nearly median
B 1.06 0.47 0.09 0.81 3.10 1.23 44.81 0.0326 2.64 Nearly median
B 1.00 0.44 0.08 0.77 3.13 1.31 43.38 0.0193 3.50 Nearly median
B 0.97 0.44 0.08 0.82 010 1.22 45.09 0.0197 2.89 Nearly median
B 0.95 0.44 0.08 0.86 0.07 1.16 46.31 0.0519 317 Nearly median
B 0.90 0.33 0.08 0.57 0.27 1.75 36.31 0.0265 3.59 Nearlysubmedian
B 0.88 0.42 0.07 0.93 0.03 1.07 48.29 0.0191 3.38 Nearly median
B 0.84 0.35 0.07 0.72 0.16 1.39 41.77 0.0323 4.22 Nearly median
C 0.79 0.38 0.07 0.92 0.04 1.08 48.03 0.0261 2.93 Nearly median
C 0.76 0.36 0.06 0.90 0.05 1.11 47.42 0.0274 3.64 Neafly median
Normal somatic chromosome number
Karyolype formula
Number of ch ro rnom pairs with secondary constriction :
Range of chromosome length in pm
Total chromosome length in pm
Average chromosome length in prn
T.F. value (%)
Variation UwIKIcient (VC)
Disparity index (D.1)
Total volume of chromoswnes in pm'
2n = 22 (Fig. 24b)
A6 84 C12
3
1.63 - 0.61
20.11
0.91
42.11
35.91
45.53
0.60
Table 25 : Detailed karyotype analysis of Edipta prostrata
Short Total arm Volume p,A Type length RL :z AD RB F% inurn, Nature of primary
in pm constriction in pm
A 1.63 0.57 0.13 0.75 0.14 1.33 35.20 0.0385 3.15 Nearly submedian 0.29
A 1.31 0.42 0.09 0.84 0.09 1.20 32.35 0.0365 3.47 Nearlysubmedian 0.38
A 1.23 0.41 0.09 0.84 0.09 1.19 32.91 0.0165 3.10 Nearlysubmedian 0.34
B 1.03 0.48 0.10 0.69 0.06 1.13 46.99 0.0637 3.05 Nearly median
B 0.87 0.42 0.09 0.94 0.03 1.06 48.54 0.0318 3.48 Nearly median
C 0.78 0.37 0.08 0.92 0.04 1.09 47.78 0.0212 3.40 Nearly median
C 0.66 0.33 0.07 0.95 0.03 1.06 48.59 0.0139 3.69 Nearly median
C 0.67 0.33 0.07 0.95 0.03 1.05 48.70 0.0306 3.73 Nearly median
C 0.63 0.30 0.06 0.91 0.05 1.10 47.64 0.0212 3.65 Nearly median
C 0.62 0.31 0.06 0.99 0.01 1.01 49.65 0.0109 3.87 Nearly median
C 0.61 0.29 0.06 0.92 0.04 1.09 47.86 0.0153 3.72 Nearly median
Normal somatic chromosome number
Kayotype formula
Number of chmmosome pairs with secondan, constriction
Range of chromosome length in vm
Total chromosome langth in vm
Average chromosome length in vm
T.F. value (%)
Variation coefficient (VC)
Disparity index (D.1)
Total volume of chromosomes in vm'
2n = 32 (Fig. 25b)
A4 016 C12
2
1.34 - 0.68
28.62
0.95
44.16
19.95
32.67
1.26
Table 26 : Detalied karyotype analysis of Gali~~soga pamr7m
Short Total am Type length length Volume p,A Nature of primary RL All RB FX in vm,
in lrm in vm constriction
A 1.27 0.38 0.06 0.73 0.11; 1.37 30.22 0.0861 2.78 Nearlysubmedkn 0.36
B 1.07 0.52 0.08 0.95 0.0:l 1.06 48.64 0.0707 3.23 Nearly median
B 0.95 0.46 0.07 0.97 0.0; 1.04 49.13 0.0332 3.04 Nearly median
B 0.94 0.43 0.07 0.84 0.05 1 19 45.71 0.0454 2.98 Nearly median
B 0.90 0.44 0.06 0.94 0.03 1.08 48.57 0.0267 3.42 Nearly median
B 0.87 0.37 0.06 0.74 0.15 1.36 42.37 0.0463 3.32 Nearly median
B 0.63 0.37 0.06 0.79 0.12 1.27 44.15 0.0240 2.91 Nearly median
B 0.82 0.38 0.06 0.88 0.06 1.14 46.78 0.0775 2.83 Nearly median
B 0.80 0.40 0.06 0.97 0.02 1.03 49.23 0.0156 3.21 Nearly median
C 0.79 0.39 0.06 0.96 0.02 1.04 46.92 0.0226 3.20 Nearly median
C 0.79 0.35 0.06 0.78 0.12 1.28 43.89 0.0314 3.28 Nearly median
C 0.76 0.39 0.05 0.96 0.01 1.02 49.40 0.0209 3.22 Nearly median
C 0.76 0.37 0.05 0.98 0.01 1.03 49.38 0.0308 2.77 Nearly median
C 0.72 0.32 0.05 0.80 0.11 1.24 44.58 0.019i 3.18 Nearly median
C 0.68 0.31 0.05 0.85 0.08 1.17 46.08 0.0141 3.44 Nearly median
Normal somatic chmmcwwne number
Keryotype formula
Number of chromosome pairs with secondary constriction :
Range of chromosome length in !nn
Total chromosome length in pm
Average chromosome length in pm
T.F. value (%)
Variation coefficient (VC)
Disparity index (D.1)
Total volume of ch rommes in pm'
Table 27 : Detailed karyotype analysis of Melitmpadium paludosm
Shalt Total Volume P,A A? AD RE F% inma
Nature of primary Tw length length RL ratlo mnstriction
in vm in pm
Neatly median
Nearly medin
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Normal somatic chromosome number
Karyotype formula
Number of chromosome pairs with secondary constriction
Range of chromosome length in pm
Total chromosome length in prn
Average chromosome length in prn
T.F. value (%)
Variation coeffiuent (VC)
Disparity index (Dl)
Total volume of chromosomes in pm'
Table 28 : Detailed karyotyp analysis of Panhenium hystemphms
2n = 36 (Fig. 27b)
A2 820 C14
lorn' arm Volume plA Nature of primary Type lenga RL tr AD RE F% constridion
in pm in ,,m
Short Total arm Volume plA Nature of primary
Type lenga RL tr AD RE F% constridion in pm in pm
A 1.41 0.45 0.07 5.24 0.12 1.26 32.04 0.0363 3.14 Nearlysubmedian 0.39
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Normal somatic chromosome number
Kacyoiype formula
Number of chromosome pairs wiih secondaN constriction
Range of chromosome length in pm
Total chromosome length in pm
Average chromosome length in pm
T.F. value (%)
Variation wefficient (VC)
Disparity index (D.1)
Total volume of chromosomes in prn'
2n = 30 (Fig. 28b)
A2 84 C24
1
1.24 - 0.61
23
0.76
46.9
19.27
34.05
0.74
Table 29 : Detailed karyotype analysis of Si{~esbeckia orientalis
Short Total an Arm 4D RE F% PIA Nature of primary
Type lenm length RL ratto wnstriction in pm in irm
Nearly sub medain
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Median
Nearly median
Nearly median
Nearly median
Nonal somatic chromosome
Karyotype formula
number
Number of chromosome pairs with secondary constriction
Range of chromosome kngth in pm
Total chromosome knglh in pm
Average chromosome length in pm
T.F. value (%)
Variation coefftcient CJC)
Disparity index (13.1)
Total volume of chromosomes in urna
Table 30 : Detailed karyotype analysis of Spilanlhes calva
Short Total an
Volume P,A Nature of primary Type length length RL ZO AD RB F% inpm,
in pm constriction in pm
0.14
0.06
0: 3
0:4
0.(11
0.03
OC'O
0.C 1
0.02
0.00
0.1 1
0.02
0.0'3
0.1 1
0.01
0.01
0.20
o.l:! 0.01
0.0''
0.1'1
Nearly sub median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly medin
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Neady median
Nearly median
Nearly median
Nearly mediin
Nearly median
Nearly median
Nearly median
Nearly median
bigs. 3Ub - Spilanrhes ciliaru i-nitotic metaphase (2n = 52); 3 1 b - Spilnnrhes radicans mitotic metaphase (2n = 78); 32b - Spilanrhes uliginosa mitotic metaphase (2n = 52); 33b - Synedrella nodgora mitotic metaphase (2n - 40); 34b - Tithonia diversifolia mitotic metaphase (2n = 34); 35b,35e - Tridax procumbens: 35b - mitotic metaphase (2n = 36), 35e - meiotic tnetaphase (n = 18); 36b - Wedelia chinensis mitotic metaphase (2n =50); 37b - Wedelia irilobata mitotic metaphase (2n = 50); 38b - Zinnia ele'qans mitotic rnetaphase (2n = 24); 39b ,39c - Tagetes erecta cv. orange: 39b - mitotic metaphase (2n - 48), 39c - somatic variant (2n = 24).
Rar represents 5pm each
Normal somatic chromosome number
Kafyotype formula
Number of chromosome pairs with secondarr constriction
Range of chromosome length in vm
Total chromosome length in vm
Average chromosome length in pm
T.F. value (%)
Variation coefficient (VC)
Disparity index (D.1)
Total volume of chromosomes in vm'
Table 31 : Detailed karyotype analysis of Spilanthes cifiata
Short Total arm Volume P,A Nalure of primary
T Y P ~ length length RL A" ratlo AD RB F% inpm, constriction in pm in vm
A 1.12 0.32 0.04 0.64 0.22 1.57 28.21 0.0347 2.57 Nearly submedian 0.31
A 0.96 0.30 0.04 0.70 0.16 1.43 30.82 0.0231 3.24 Nearly sub median 0.24
A 0.95 0.32 0.03 0.91 0.05 1.10 34.09 0.0204 3.43 Nearly sub median 0.27
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Median
Nearly median
Nearly median
Nearly sub median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Neady median
Nearly median
C 0.65 0.30 0.03 0.87 0.07 1.15 46.42 0.0147 3.69 Nearly median
c 0.63 0.28 0.03 0.83 0.09 1.21 45.28 0.0207 3.40 Nearly median
C 0.59 0.28 0.03 0.92 0.04 1.08 48.01 0.0102 3.77 Nearly mediin
C 0.58 0.29 0.03 0.97 0.02 1.04 49.12 0.0159 3.04 Nearly median
C 0.56 0.28 0.03 0.97 0.01 1.03 49.31 0.0096 3.33 Nearly median
C 0.53 0.25 0.03 0.85 0.08 1.18 45.90 0.0203 3.73 Nearly median
C 0.50 0.24 0.03 0.88 0.06 1.14 46.79 0.0104 3.41 Nearly median
Spilanihes radicans
Normal somatic chromosome number
Karyotype formula
Number of chromosome pairs with secondary conshiction
Range of chromosome length in pm
Total chromosome length in pm
Average chromosome length in pm
T.F. value (%)
Variation coefficient (VC)
Disparity index (D.1)
Total volume of chromosomes in pm'
Table 32 : Detailed kalyotype analysis of Sprlanthes radicans
Short Total an Type length RL :z /iD RE F% Volume P/A Nature of primary
in pm length in pm' constriction in pm
A 1.62 0.61 0.04 0.97 001 1.03 37.56 0.0323 3.26 Nearly median 0.39
A 1.54 0.50 0.03 0.86 0 07 1.16 32.69 0.0396 3.20 Nearly sub median 0.45
A 1.46 0.46 0.03 0.77 013 1.30 31.82 0.0242 3.06 Nearlysubmedian 0.39
B 1.34 0.58 0.04 0.75 014 1.33 42.95 0.0288 3.66 Nearly median
B 1.05 0.44 0.03 0.71 0 17 1.41 41.48 0.2766 3.56 Nearly median
B 1.03 0.44 0.03 0.76 014 1.32 43.05 0.0277 3.18 Nearly median
B 0.92 0.41 0.03 0.81 0 10 1.23 44.79 0.0305 2.82 Nearly median
B 0.84 0.37 0.03 0.78 012 1.28 43.89 0.0248 3.14 Nearly median
B 0.84 0.41 0.03 0.98 001 1.02 49.43 0.0366 3.41 Nearly median
B 0.83 0.37 0.03 0.79 012 1.27 44.09 0.0196 3.45 Nearly median
B 0.81 0.29 0.03 0.56 0 28 1.79 35.86 0.0245 3.71 Nearly sub median
C 0.80 0.37 0.03 0.85 008 1.18 45.94 0.0175 3.40 Nearly median
C 0.79 0.33 0.03 0.72 016 1.39 41.87 0.0207 3.34 Nearly median
C 0.77 0.39 0.02 1.01 -0.01 0.99 50.31 0.0109 3.32 Nearly median
c 0.75 0.36 0.02 0.92 004 1.09 47.88 0.0156 3.42 Nearly median
C 0.74 0.36 0.02 0.95 002 1.05 48.75 0.0251 3.40 Nearly median
C 0.74 0.35 0.02 0.91 005 110 47.55 0.0381 3.53 Nearly median
c 0.73 0.36 0.02 0.98 001 1.02 49.51 0.0085 3.54 Nearly median
C 0.72 0.32 0.02 0.81 0 11 1.24 44.74 0.0077 3.94 Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly medtan
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly sub median
Nearly sub median
Normal somatlc chromosome number
Karyotype formula
Number of chromosome pairs with secondar) constriction
Range of chromosome iength in pm
Total chromosome length in pm
Average chromosome length in pm
T.F. value (%)
Variation weffment (VC)
Disparity index (D.1)
Total volume of chromosomes in um3
Table 33: Detailed karyotype analysis of Spilanfhes uliginosa
Total Type length
In pm
arm length
Arm AD RE RL ratio
2n = 52 (Fg. 32b)
A4 834 C14
2
1.25-061
47.75
0.918
45.92
17.73
34.4
2.13
Volume p,A Nature of primary F% in pm= constriction
Nearly sub median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearty median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
C 0.79 0.37 0.03 0.91 O.O!j 1.10 47.51 0.0372 2.79 Nearly median
C 0.78 0.38 0.03 0.94 0.03 1.06 48.51 0.0384 2.77 Nearly median
C 0.77 0.34 0.03 0.81 0.11 1.24 44.63 0.0101 3.25 Nearly median
C 0.76 0.36 0.03 0.92 0.04 1.09 47.86 0.0231 2.80 Nearly median
C 0.70 0.30 0.03 0.76 0.14 1.32 43.12 0.0394 2.88 Nearly median
C 0.61 0.28 0.03 0.85 0.08 1.18 45.90 0.0531 3.06 Nearly median
Normal somatic chromosome number
Karyotype formula
Number of chromosome pain with sewndaty mnstriction
Range of chromosome length in pm
Total chromosome length in vm
Average chromcsome length in pm
T.F. value (%)
Variation coefficient (VC)
Disparity index (D.1)
Total volume of chromosomes in uma
Table 34 : Detailed karyotype analysis of Synedrella nodifbra
2n = 40 (Fig. 33b)
A4 616 C20
2
1.67 - 0.57
35.32
0.883
42.24
34.58
40.86
1.65
Shoa Total arm Volume p,A Nature of primary Type length RL tE RB F% in pm.
inpm . wnstriction In vm
Nearly sub median
Nearly sub median
Nearly sub median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly sub median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly medin
marly median
Nearly median
Nearly median
Nearly median
Nearly median
Normal somatic chromosome number
Karyotype formula
Number of chromosome pairs with secondary c3nstriction
Range of chmmosome length in vm
Total chromosome length in pm
Average chromosome length in Vm
T.F. value (%)
Variation meffident (VC)
Dispariky index (D.1)
Total volume of chromosomes in pm'
Table 35 : Detailed karyotype analysis of TiUm~ria d i~~mYdia
Short Total am A m AD F% Volume P,A Nature of primary Type length length RL ratio in pma
in vm in vm constriction
A 1.79 0.55 0.07 0.65 0.2:! 1.55 30.74 0.1271 2.66 Nearlysubmedian 0.39
A 1.69 0.61 0.07 0.81 0.111 1.23 36.01 0.0359 2.89 Nearlysubmedian 0.33
B 1.66 0.76 0.08 0.90 O.O!i 1.12 47.27 0.0680 2.71 Nearly median
B 1.40 0.67 0.07 0.91 O.O!i 1.09 47.74 0.0444 3.07 Nearly median
B 1.31 0.50 0.07 0.62 0.2:1 1.61 38.30 0.0434 3.26 Nearly median
B 1.16 0.50 0.06 0.76 O.lr. 1.32 43.18 0.1491 2.67 Nearly median
B 1.12 0.53 0.06 0.90 O.O!, 111 47.29 0.0506 2.77 Nearly median
B 1.08 0.54 0.06 0.99 0.0(1 1.01 49.87 0.0373 3.70 Nearly median
B 1.05 0.50 0.05 0.93 0 108 48.07 0.0143 2.99 Nearly median
B 1.02 0.45 0.05 0.79 0.1; 1.26 44.21 0.0362 2.91 Nearly median
B 0.99 0.44 0.05 0.80 0.1' 1.25 44.46 0.0676 2.64 Nearly median
B 0.98 0.44 0.05 0.83 O.l i1 1.21 45.24 0.0195 3.43 Nearly median
B 0.96 0.47 0.05 0.97 0.0;' 1.03 49.22 0.0127 3.53 Nearly median
B 0.94 0.43 0.05 0.84 0.OIi 1.20 45.54 0.0468 2.67 Nearly median
B 0.87 0.40 0.04 0.67 O.Oi 1.14 46.62 0.0271 3.23 Nearly median
C 0.80 0.38 O M 0.93 0 1.08 48.09 0.0170 3.61 Nearly median
C 0.73 0.35 0.04 0.92 0 . 1.09 47.86 0.0072 2.36 Nearly median
Normal somatic chromosome number
Karyotype formula
Number of chromosome pain with secondary a)nstrlction
Range of chromosome length in pm
Total chromosome length in pm
Average chromosome length in pm
T.F. value (%)
Variation coe f f t n t (VC)
Disparity index (D.1)
Total volume of chromosomes in um'
Table 36 : Detailed karyotype analysis of Trida): procumbens
2n = 36 (Fig. 35b)
A6 830
3
2.23 - 1.25
58.14
1.615
40.9
15.38
28.16
4.5402
Short Total arm Volume p,A RL AD RB F% inpm, Nature of primary Type length length
in pm mnsMdion In pm
A 2.23 0.71 0.12 0.64 0.2:! 1.55 31.61 0.1344 2.64 Nearlysubmedian 0.43
A 1.96 0.63 0.09 0.88 O.Mi 1.14 31.96 0.0587 2.25 Nearlysubmedian 0.62
A 1.94 0.62 0.09 0.88 0.0:' 1.14 32.20 0.1748 3.02 Nearly sub mediin 0.60
B 1.79 0.75 0.12 0.73 0.lli 1.38 42.05 0.0375 2.65 Nearly median
B 1.76 0.69 0.12 0.66 0.2' 1.53 39.59 0.0692 2.47 Nearly median
B 1.70 ' 0.84 0.12 0.97 0.0'1 1.03 49.35 0.2553 2.41 Nearly median
B 1.67 0.75 0.11 0.82 0.10 1.21 45.17 0.0737 2.62 Nearly median
B 1.64 0.70 0.11 0.74 I ! 1.36 42.43 0.1250 2.96 Nearly median
B 1.62 0.75 0.11 0.86 0.0" 1.16 46.28 0.1046 2.99 Nearly median
B 1.54 0.67 0.11 0.78 I ! 1.28 43.86 0.0923 2.73 Nearly median
B 1.50 0.68 0.10 0.84 0.0!1 1.19 45.69 0.1111 2.67 Nearly median
B 1.49 0.56 0.10 0.60 O.Z!i 1.68 37.36 0.0023 2.42 Nearlysubmedian
B 1.45 0.68 0.10 0.88 0.0" 1.14 46.74 0.3176 3.43 Nearly median
B 1.44 0.52 0.10 0.56 0.2;3 1.79 35.81 0.0677 2.97 Nearlysubmedian
B 1.39 0.56 0.10 0.68 0.13 1.46 40.60 0.2996 3.54 Nearly median
B 1.38 0.59 0.09 0.74 0.15 1.36 42.42 0.1699 2.90 Nearly median
B 1.31 0.58 0.09 0.79 0.lZ 1.26 44.15 0.1015 3.57 Nearly median
B 1.25 0.61 0.09 0.94 0.03 1.06 48.49 0.0752 3.03 Nearly median
Normal somatlc chromosome number
Karyotype formula
Number of chromosome pairs with sewndar) constriction : Range of chromosome length in pm
Total chromosome length in pm
Average chromosome length in pm
T.F. value (%)
Variation coefficient (VC)
Disparity index (Dl)
Total volume of chromosomes in pm'
2n = 50 (Fig. 36b)
A4 68 C38
2
1.01 - 0.48
35.81
0.716
44.84
18.68
35.57
1.41
Table 37 Deta~led karyoh/pe analysis of Wedt,lia chinensis
Short Total arm
Arm Type length length Volume P,A Nature of primary
RL AD RB F% in pm'
in pm in pm constriction
A 1 01 0 33 0.04 0.85 0.06 118 32.22 0.0680 3.45 Nearly submedian 0 30
A 0.97 0.32 0.04 0.92 0.04 1.09 33.46 0.0636 3.65 Nearlysubmedian 0.29
Nearly median
Median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Median
Median
Nearly median
Median
Nearly mediin
Nearly sub median
Nearly median
Median
Nearly median
C 0.60 0.22 0.03 0.58 0 26 1.72 36.79 0.0468 3.67 Nearly sub median
C 0.59 0.31 0.03 1.12 -13.06 0.89 52.88 0.0115 4.08 Median
C 0.57 0.27 0.03 0.90 005 1.11 47.45 0.0090 4.34 Nearly median
C 0.53 0.27 0.03 0.99 000 1.01 49.86 0.0062 4.35 Nearly median
C 0.48 0.21 0.03 0.76 0.14 1.32 43.10 0.0136 3.69 Nearly median
Normal somatic chromosome number
Karyotype formula
Number of chromosome pairs with secondary constriction
Range of chromosome length in pm
Total chromosome length in pm
Average chromosome length in pm
T.F value (76)
Variation coefficient (VC)
Disparity index (Dl)
Total volume of chromosomes in pm'
2n = 50 (Fig. 37b)
A4 B2 C44
2
1.15-0.41
30.78
0.61
43.14
25.38
47.43
0.89
Table 38 Deta~led karyotype analysls Wede ra tnlobata
Short Total arm
Volume p,A A? PD RE F% in pm, Nature of primary
Type length length In pm
RL ratlo constriction In pm
Nearly sub median
Nearly sub median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly sub median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
C 0.51 0.19 0.03 0.59 0.:'6 1.69 37.17 0.0126 4.56 Nearly sub median
C 0.49 0.20 0.03 0.67 O.:M 1.50 40.07 0.0160 4.50 Nearly median
c 0.49 0.22 0.03 0.81 0. 11 1.24 44.71 0.0160 4.03 Nearly median
C 0.47 0.21 0.03 0.79 012 1.27 44.00 0.0131 4.44 Nearly median
C 0.41 0.17 0.03 0.68 0.19 1.48 40.40 0.0116 4.45 Nearly median
Normal somatic chromosome numbel
Kafyotype formula
Number of chromosome pairs with seconda~y constriction
Range of chromosome length in vm
Total chromosome length in pm
Average chromosome length in pm
T.F. value (%)
Variation memuen1 (VC)
Disparity index (D.1)
Total volume of chromosomes in pm'
Table 39 : Detailed kafyotype analysis of Zinnia elegans
Short Total arm Arm Volume p,A Nature of primafy
Type length length RL ratio 4D RB F% in pm, inpm . constriction
In pm
Nearly sub median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Normal somatic chromosome number
Karyotype formula Number of chromosome pairs with secondary iXJnStricti0n :
Range of chrommme length in vm
Total chromosome length in vm
Average chromosome length in vm
T.F. value (%)
Variation coefficient (VC)
Disparity index (D.1) Total volume of chmmosomes in pm'
Table 40 : Detailed karyotype analysis of Tagetas emcta cv orange
Short Total arm ~ r m Volume plA Nalure of primary
Type length length RL ratio 4D RB F% mnstridion in vm in vm
Nearly median
Nearly sub median
Nearly median
Nearly sub median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Median
Nearly median
Nearly median
Median
Nearly median
Nearly median
Nearly median
Nearly sub median
Nearly median
Nearly median
C 0.76 0.30 0.03 0.66 C.21 1.53 39.60 0.0072 4.43 Nearly median
C 0.75 0.36 0.03 1.03 4.02 0.97 50.62 0.0099 4.29 Median
C 0.67 0.30 0.03 0.82 010 1.22 45.02 0.0101 4.76 Nearly median
C 0.60 0.21 0.03 0.52 0 31 1.92 34.26 0.0078 4.78 Nearlysub median
Figs. 40b - Tagures erecla cv pale yellow mitotic mclapha5e (211 = 48); 4lb,J lc.4 lcl - Tageles erecra cv. yellow: 4 1 b - mitotic m e t a p h a e (2n = 24), 4 1 c - somatic variant (2n = 22), 41c1 - somatic variant (2n = 18); 42b - Tagetes paflda mitotic metaphase (2n = 24); 43b - Chrysanthemun~ parrheniurn mitotic metaphasc (2n = 18); 44b - Crassocephalum crepidiodes mitotic metaphase (2n = 40); 45b.45e - Emilia sonchijblia: 45b - mitotic metaphase (2n -- lo), 45e - meiotic metaphase (n = 51, 46b,46e ~Vofonia grandflora : 46b - mitotic metaphase (2n = 20), 46e - meiotic metaphase (n = 10); 47b - Sonchm oleraceus mitotic melaphase (2n = 1 8)
Rar represents 5pm each
Normal somatic chromosome number
Karyolype formula
Number of chromosome pairs with secondary constriction :
Range of chromosome length in pm
~ o t a l chromosome length in pm
Average chromosome length in pm
T.F. value (Ye) Variation coefiicient (VC)
Disparity index (D.1)
Total volume of chromosomes in pm'
2n = 48 (Fig. 40b)
A4 636 C8
2
1.72- 0.72
48.27
1.01
43.27
22.82
40.74
3.62
Table 41 : Detailed karyotype analysis of Tiwtes emfa cvpc~le yellow
Short Total arm Volume p,A Nature of primary
Type length length
RL AD RB F?6 in pm, constridbn in in prn
~edr ly sub median
Nearly sub median
Nearty sub median
N~arly median
NMrly median
NIarly median
Nsdrly median
N6drly median
Nearly sub m i a n
Nearly median
Nearly median
Nearly median
Nearly medlan
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
C 0.80 0.37 0.03 0.85 0.08 1.17 46.M) 0.0637 2.79 Nearly median
C 0.76 0.37 0.03 0.95 0.13 1.05 48.75 0.0431 2.77 Nearly median
C 0.74 0.33 0.03 0.78 0.12 1.27 43.97 0.0316 3.03 Nearly median
C 0.72 0.32 0.03 0.79 012 1.27 44.06 0.0597 3.23 Nearly median
Tagetes ereeta cv yellow
Normal somatic chromosome number
Karyotype formula
Number of chromosome pairs with secondary constriction
Range of chromosome length in pm
Total chromosome length in Mm
Average chromosome length in pm
T.F. value (%)
Variation coefficient (VC)
Disparity index @.I)
Total volume of chromosomes in vm'
2n = 24 (Fig. 41b)
A2 08 C14
1
1.20 - 0.67 19.51
0.81
44.38
18.1
28.34
0.4
Table 42 : Detailed karyotype analysis of Tagetes emta cv yelbw
Short Total arm Volume p,A Nature of primary TYW length length RL :G AD RB F% invm, wnsbiction
in vm in
A 1.20 0.39 0.09 0.88 0.16 1.14 32.90 0.0218 2.91 Nearlysubmedian 0.36
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Normal somatic chromosome number
Karyotyp formula
Number of chromosome pairs with secondary constriction :
Range of chromosome length in pm
Total chromosome length in pm
Average chromosome length in pm
T.F. value (Oh)
Variation mefficient (VC)
Disparity index (D.1)
Total volume of chromosomes in urn'
2n = 24 (Fig. 42b)
A2 812 C10
1
1.36 - 0.66
21.01
0.87
45.12
23.92
34.65
0.43
Table 43 : Detailed karyotype analysis of Taqetes pafula
Short Total arm Volume .1D RB F% P,A Nature of primary
Type length inpm . RL ratlo constriction
In urn
A 1.36 0.46 0.10 0.72 (1.16 1.38 33.71 0.0013 2.87 Nearlysubmedian 0.27
B 1.16 0.55 0.11 0.91 0.05 1.10 47.59 0.0663 2.81 Nearly median
B 1.10 0.55 0.10 1.00 0.00 1.00 49.90 0.0290 2.76 Nearly median
B 0.86 0.42 0.08 0.95 11.03 1.06 48.66 0.0153 3.35 Nearly median
B 0.83 0.39 0.08 0.87 0.07 1.15 46.58 0.0188 3.20 Nearly median
B 0.81 0.38 0.08 0.88 0.06 1.14 46.81 0.0170 3.03 Nearly median
B 0.80 0.32 0.08 0.68 0.19 1.48 40.33 0.0089 3.81 Nearly median
C 0.77 0.39 0.07 1.00 1.00 1.00 49.93 0.0195 4.00 Nearly median
C 0.74 0.34 0.07 0.85 1.08 1.18 45.80 0.0140 3.60 Nearly median
C 0.71 0.31 0.07 0.77 1.13 1.30 43.49 0.0092 3.55 Nearly median
C 0.69 0.33 0.07 0.95 1.02 1.05 48.80 0.0101 4.19 Needy median
C 0.66 0.30 0.06 0.83 2.09 1.20 45.45 0.0090 3.83 Nearly median
Normal somatic chromosome number
Karyotype formula
Number of chromosome pairs with samndary omstridion :
Range of chromosome length in pm
Total chromosome length in pm
Average chromosome length in pm
T.F. value (%)
Variation coefficient (VC)
Disparity index (D.1)
Total volume of chromosomes in pm'
2n = 18 (Fig. 43b)
A2 814 C2
1
Table 44 : Detailed karyotype analysis of Chrysanthemum parthenium
Short Total arm ~ r m Volume p,A AD RB F% in pm, Nature of primaty
Type length length in pm RL ratio constriction
in pm
Nearly sub median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Naarty median
Nearly median
Normal somatic chromosome number
Karyotyp formula
Number of chromosome pairs with semndary c,>nstriction
Range of chromosome length in pm
Total chromosome length in pm
Average chromosome length in pm
T.F. value (%)
Variation coefficient (VC) Disparity index (D.1)
Total volume of chromosomes in pm'
Table 45 : Detailed karyotyp analysis of Crassooephalum mpidioides
Short Total an RL AD RB F% ' PIA Nature of primary Type length length
in pm in mnstriction
A 1.92 0.55 0.05 0.65 0.21 1.53 28.82 0.1430 2.58 Nearly sub median 0.52
A 1.80 0.55 0.04 0.85 0.06 1.18 30.42 0.1090 2.76 Nearlysubmedian 0.61
Nearly sub median
Nearly median
Nearly median
Median
Nearly median
Nearly median
Median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Nearly median
Median
Nearly median
Emllla sonchHorIa
Normal somatic chromosome number
Karyotype formula
Number of chmmosome pairs with m n d a r y mstriction
Range of chromosome length in pm
Total chromosome length in pm
Average chmmosome length in pm
T.F. value (Om)
Variation coefficient (VC)
Disparity index (D.1)
Total volume of chromosomes in pma
2n = 10 (Fig. 45b)
A2 68 1
2.54 - 1.76 20.25
2.03
43.9
13.49
18.13
4.57
Table 46 : Detailed karyolype analysis of Emila sonchifdia
Short Total arm ~ r m Volume P,A A[) RB F% in Nature of primary
Type length length RL ratio constridion in vm in pm
A 2.54 0.93 0.19 0.95 0.C3 1.06 36.46 0.8783 2.12 Nearlysubmedian 0.63
B 2.02 0.75 0.20 0.59 0.26 1.71 36.96 0.4890 2.32 Nearlysubmedian
B 1.97 0.91 0.19 0.85 0.08 1.18 45.93 0.2998 2.05 Nearly median
B 1.83 0.95 0.18 1.09 -O.(kl 0.92 52.17 0.2958 2.42 Nearly median
B 1.76 091 0.17 1.07 -0.M 0.93 51.77 0.3220 2.39 Nearly median
Normal somatic chromosome number
Karyolype formula
Number Of chromosome pairs with secondary constrict~on
Range of chromosome length in pm
Total chrcinwonre length in pm
Average chromosome length in pm
T.F. value (%)
Variation coefficient (VC)
Disparity index (D.1)
Total volume of chromosomes in pm'
2n = 20 (Fig. 46b)
A2 B18
1
2.18- 1.26
32.55
1.62
45.46
20.2
25.71
4.5
Table 47 : Detailed karyotype analysis of Notonia grandiflorn
Short Total arm Type length RL tz A l l RB FK PIA Nature of primary
in pm wnswcuon in pm
A 2.16 0.74 0.11 1.37 4 . "6 0.73 46.75 0.1822 2.55 Nearly median 0.42
B 2.17 0.98 0.13 0.83 0.10 2.21 45.23 0.2573 2.81 Nearly median
B 1.77 0.58 0.11 0.48 0.35 2.07 32.61 0.3549 2.12 Nearlysubmedian
B 1.80 0.88 0.11 0.96 0.02 1.04 48.93 0.4926 2.67 Nearly median
B 1.62 0.76 0.10 0.88 0.05 1.14 46.80 0.1667 2.54 Nearly median
B 1.51 0.79 0.09 1.11 -0.05 0.90 52.61 0.0612 2.56 Median
B 1.36 0.66 0.08 0.94 0.0:) 1.06 46.52 0.2240 2.51 Nearly median
B 1.34 0.63 0.08 0.89 O.@i 1.12 47.17 0.1261 2.73 Nearly median
B 1.27 0.50 0.08 0.65 0.2:! 1.55 39.24 0.1009 2.95 Nearly median
B 1.26 0.61 0.08 0.93 0.04 1.07 48.21 0.2841 2.62 Nearly median
Sonchus olencws
Normal somatic chromosome number
Karyotype formula
Number of chromosome pairs with secondary constriction
Range of chromosome length in prn
Total chromosome length in urn
Average chromosome length in prn
T.F. value (%)
Variation coefficient (VC)
Disparity index (D.1)
Total volume of chromosomes in urnS
Table 48 : Detailed karyolype analysis of Sonchus oleraceus
2n = 18 (Fig. 47b)
A2 814 C2
Short Total arm Volume
RL :E Al l RB F% in P/A Nature of primary ~ y p e length ,engch
in vrn constriction
In urn
A 1.36 0.50 0.13 0.89 0.[16 1.12 36.68 0.0150 2.84 Nearlysubmedian 0.30
B 1.05 0.52 0.12 0.98 0.C1 1.02 49.51 0.0225 3.37 Nearly median
B 0.96 0.42 0.11 0.77 0.13 1.30 43.56 0.0222 2.76 Nearly median
B 0.91 0.39 0.11 0.74 0.15 1.35 42.51 0.0151 3.50 Nearly median
B 0.69 0.43 0.11 0.93 0.03 1.07 48.28 0.0185 3.38 Nearly median
B 0.85 0.42 0.10 0.98 0.01 1.02 49.60 0.0173 3.73 Nearly median
B 0.83 0.41 0.10 0.96 O.O.! 1.04 49.08 0.0155 3.31 Nearly median
B 0.81 0.35 0.10 0.77 0 . 1 1.29 43.61 0.0188 3.72 Nearly median
C 0.79 0.36 0.09 0.83 0.0!1 1.20 45.35 0.0216 3.60 Nearly median
A B B B C C C C C C C
A A B B B B B B C
A B B B B B B B B B E B B B B B B C C C
Fig. Id to 5d -Comparative idiogram chart of various taxa studied
I d - Elephantopus scaber (2n = 22) 2d - PhV,Iocephalum rangacharii (2n = 18) 3d - Vernonia cinema (2n = 18) 4d - Adenostemma lavenia (2n = 20) 5d - Ageratum mnyzoides r2n = 40)
A A A B B B B B B B B B B B B B B B B B B B B B B
A A B B C C C C C C C C C C C C C C
10d I
A B B B B B B B B B B B l > B B B B B B B B B B B B B B
Fig. 6d to 10d - Comparative idiogram chart of various taxa studied
6d - Agemtum haustonianum (2n = 40) 74 - Chromolaena odomta (Zn = 60) ad - Eupatorium triplinewe (2n = 50) 9d - Mik~nia cordata (2n = 36) 10d - Conyza bonariensis (2n = 54)
l l d I
A B B B B B B B C
A B B B B B B B C
A B B B B B B B B
Fig.1 ld to 15d - Comparative idiograrn chart of various taxa studied
I l d - Conyza canadensis (2n = 18) 126 - Dichrocephala chrysanthemifolia (2n = 18) 136 - Erigeron mut:ronatus (2n = 18) 14d - Blumea lacem (2n = 36) 156 - Blumea mollis (2n = 18)
I
A B B B B B B B B B B C C C
A A B B B B B B B B B B B B B B B B B B B B B B B C C C C C C C C C C C
Fig. 16d to 20d - Comparative idiogram chart of various taxa studied
16d - Blumea oxyodonta (2n = 28) 17d - Sphaeranthus indicus (217 = 20) 18d - Vicoa indica (2n = 13) 19d - Acanthospermum hispidum (Zn = 22) 20d - Bidens pilosa (2n = 52)
A A B B B B B B B B C C C C C C
Fig. 21d to 25d -Comparative idiogram chart of various taxa studied
21d - Cosmos bipinnatus cv orange (2n = 24) 22d - Cosmos bipinnatus cv yellow (2n = 24) 236 - Cosmos caudatus (2r = 24) 24d - Edipta prostrata (2n = 22) 25d - Galinsoga parviflom (2n = 32)
A B C C C C C C C C C C
A B B B B B B B B B B C C C C C C C
A B B C C C C C C C C C C C C
A A A B B B B B B C C C C C C C C C C C C C C C C C
Fig. 26d to 306 - Comparative idiogram chart of various taxa studied
26d - Melampodium paludom (2n = 24) 276 - Parfhenium hystemphorc~s (2n 36) 286 - Sigesbeckia orientalis (2n = 30) 29d - Spilanthes calva (2n = 78) 306 - Spilanthes ciliata (217 = 5.2)
A A A B B B B B B B B C C C C C C C C C C C C C C C C C C C C C C C C C C C C
A A B B B B B B B B B B B B B B B B B C C C C C C C
A A B B B B B B B B C C C C C C C C C C
Fig. 31d to 356 - Comparative idiogram chart of various taxa studied
31d - Spilanthes rndicans (;!n = 78) 32d - Spilanthes ul@inosa (2n = 52) 336 - Synedrella nodiflorn (:!n 40) 1 346 - Tithonia diversifolia (2n = 34) 35d - Tridax procumbens (2n 36) scale 1 pm
A A B C C C C C C C C C C C C C C C C C C C C C C
A B B B B B B B B B B C
Fig. 36d to 406 - Comparative idiogram chart of various taxa studied
36d - Wedelia chinensis (2n = 50) 37d - Wedelia trilobata (2n = 50) 384 - Zinnia elegans (2n = 24) 396 - Tagetes erecta cv orange (2n = 48) 40d - Tagetes erecta cv pale yellow (2n = 48)
A B B B B C C C C C C C
A B B B B B B C C C C C
A B B B B B B B C
A A A B B B B B B B B B B B B B B B B C
Fig. 41d to 44d - Comparative idiogram chart of various taxa studied
41 d - Tagetes e m t a cv yellouc (2n = 24) 42d - Tagetes patula (2n = 24 4
436 - Chrysanthemum parthenium (2n = 18) 44d - Crassocephalum crepidioides (217 = 40)
A B B B B B B B B B
i 1 m . . I . . 1 1 1 1 1 3 . 1 . A B B B B B B B C
Fig. 45d to 47d - Comparative idiogram chart of various taxa studied
45d - Emilia sonchifolia (2n = 10) 46d - Notonia grandiflora (2n = 20) 47d - Sonchus oleraceus (211 = 18)
scale I prn
Table : 49.
Summary of the Karyomorphometerial analysis on the forty seven taxa investigated
No Name of the taxa 2n PL KF CP RCL ACL TCL CLH DI VC TF Volume
Elephantopus scaber 22 2x
nl-..lJ---..h"r,,w ""w*",-h"rii A ,,I..V.,lZ,.-...... . -... o- - 18 2x
Vemnia cinerea 18 2x
Adenostemma lavenia 20 4x
Ageratum conyoides 40 4x
Ageratum haustonianum 40 4x
Chromolaem odorata 60 6x
Eupatorium triplinewe 50 5x
Mikania cordata 36 2x
Conyza bonariensis
Conyza canadensis
Dichrocephala chrysanthemifolia
Erigeron mucronatus
Blumea lacera
Blumea mollis
QI.,,"" ",,"do,tn " . -., . - - - . ., - - -
Sphaeranthus indicus
Vicoa indica
Acanthospennum hispidum
Bidem pilosa
Cosmos hipinnatus cv orange
Cosmos hipinnatus cv yellow
Cosmos caudatus
Eclipta prostrata
Galinsoga parvijlora
Melampodium paludosm
Parthenium hysterophorus
Sigesbeckia orientalis
Spilanthes calva
Spilanthes ciliata
Spilanthes radicans
Spilanrhes uliginosa
SynedrelIa nodzjlora
Tithonia divers ifolia
Tridar procumbens
Wedelia chinensis
Wedelia trilobata
38 Zinnia elegans 24 2x A292OC2 1 2.00-0.79 1.07 25.73 12.86 43.36 29.55 45.66 1.59
39 Tagetes erecta cv. orange 48 4x A2B36C10 1 1.44-0.60 0.92 44.34 11.08 41.i7 18.89 44.06 1.14
40 Tagetes erecta cvpale yellow 48 4x A4B36 C8 2 1.71-0.72 1.01 48.27 12.06 40.74 22.82 43.27 3.62
41 Tagetes erecta cv. yellow 24 2x A2B8C14 1 1.20-0.67 0.81 19.51 9.76 28.34 18.10 44.38 0.40
42 Tagetes patula 24 2x A2B12C10 1 1.36-0.66 0.87 21.01 10.50 34.65 23.92 45.12 0.43
43 Chrysanthemum parthenium 18 2x A2B14C2 1 1.56-0.64 1.12 20.20 10.10 41.81 23.62 45.89 0.62
44 Crassocephalurn crepidioides 40 4x A6B32 C2 3 1.92-0.78 1.42 57.11 14.27 42.22 21.67 42.07 4.52
46 Notonia grandiflorn 20 4x A2B18 1 2.20- 1.30 1.62 32.55 16.30 25.71 20.20 45.46 4.50
47 Sonchus oleraceus 18 2x A2914C2 1 1.36-0.79 0.93 16.88 8.44 26.51 17.92 44.90 0.33
PL - Ploidy level, KF - Karyotype formula, CP - Chromosome pairs with secondary constriction, RCL - Range of chromosome length in pn, ACL - Average chromosome length in p, TCL - Total chromatin length in p, CLH - Chromatin length of haploid complement in p, DI - Disparity index, VC -Variation coefficient, TF -Total forma percentage, Volume - Total volume of chromosome.
Chromosome number E l % of Plants
Fig.48 Range of chromosme numbers recoganized in the present study
Base Number El% of Plant
Fig.49 Range of Base Numbers recoganised in the present study
Fig.50 Range of ploidy level recoganized in the present study
Elephantopus scaber Phyllocephalurn rangacharii Vemonia cineree
Fig.51: Comparison of the major karyotypic parameters in various taxa of the tribe Vernonieae
Fig.52 : Comparison of the major karyotypic parameters in various taxa of the tribe Eupatorieae
Conyzs bonaflensis Conyza canadensis Dichrocephala Erigemn chrysanthemifofia mucronetus
Fig.53 : Comparison of the major karyotypic parameters on various taxa of the tribe Astereae
Blurnee lecem BIumea mollis Blumea Sphaemnthus Vima indica oxyodont8 indicus
Fig.54 : Comparison of the major karyotypic parameters on various taxa of the tribe lnuleae
QTCL
TF%
Acanfhospennum Bidens pilosa Eclipfe prostrata Galinsoga parviffora hispidurn
Fig.55 : Comparison of the major karyotypic parameters on various taxa of the tribe Heliantheae
Cosmos bipinnatus cv Cosmos bipinnalus cv yellow Cosmos caudatus orange
Fig.56 : Comparison of the major karyotypic parameters in different cyfotypes of the genes Cosmos
Melampodium Parthenium Sigesbeckia Synedrella nodflora paludosm hystemphorus orientalis
Fig.57 : Comparison of the major karyotypic parameters on various taxa of the tr~be Heliantheae
Spilanthes calve Spilanthes ciliafa Spilanthes radimns Spilanthes uliginosa
Fig.58 : Comparison of the major karyotypic parameters in different cytotypes of the genus Spilanthes
Tifhonia Tridex Wedelta Wedelia Zinnia elegans diversifolia procumbens chinensis trilobata
Fig.59 : Comparison of the major karyotypic parameters on various taxa of the tribe Heliantheae
Tagefes erecta cv Tagetes erecte cv Tegetes erect@ ev Tegetes pefula orange pale yellow yellow
Fig.60 : Comparison of the major karyotypic parameters in different cytotypes of the genus Tagetes
Chryssnthemum Crassocephelum Emilia sonchifolia Nofonia Sonchus partbenium crepidioides grandflora oleraceus
avc I UTF%
Fig.61 : Comparison of the major karyotypic parameters on various taxa studied
DISCUSSION
a) Chromosome numbers in the genera investigated
Chromosome number determination from fifty seven populations attributed to
forty seven species from thirty four genera of Asteraceae were made.
Cytologically the family is variable. The chromosome spectrum varies from
2n = 10 to 2n = 78. With majority of species concentrated in the number 2n =
18, followed by 2n = 24, 2n = 3t3 and 2n = 40. The presence of identical
numbers in unrelated genera is a vary noteworthy feature in this family. The
presence of such widely differen series of chromosome numbers in the
species of even the same genus and in genera placed under different tribes
indicated that the different chromosame numbers can be derived one from the
other.
The counts confirm the chromosom: numbers of all previous reports in some
species or are consistent with one of variable numbers reported by several
authors in other species. Of the 'ifty seven plants in which chromosome
number determinations were made hventy six (55.31%) are diploids and
twenty one (44.68%) are polyploids. High chromosome numbers such a . n =
15, 16,17 and 25, have been considered to be the result of polyploid increase
followed by dysploid loss (King and Kobnison, 1987).
Sub family: Tubuliflorae
Tribe: Vernonieae
In this tribe three taxa belonging to three genera have been examined. The
chromosome number of phyllc~cephalum rangacharii is 2n = 18,
Elephantopus scaber is 2n = 22 and Vernonia cinera is 2n = 18. According to
the previous reports (Shetty, 1967; Narayana, 1979; Ayyangar and
Sampathkumar, 1969a; Sharma and Sarkar, 1967-68; and Bir and Sidhu,
1980); the somatic chromosome numbers evaluated in these species are 2n =
18 (Phyllocephalum rangacharir) 2n = 18, 22, 44 (Elephantopus scaber) and
2n = 18, 36 (Vernonia cinerea).Tt~is seems to be a confirmation of an earlier
report made on these taxas.
Tribe : Eupatorieae:
Five genera have been examined. The chromosome count made on the lone
member investigated in the genus Adenostemma is 2n = 20 (Adenostemma
lavenia). This somatic chromosome number agrees with the previous reports
(Powell and Cuatrecasas, 1970; Mi yagi, 1971). The second genus Ageratum is
seems to be plastic. In the present study the genus Ageratum represented by
two species. The chromosome number ranges from 2n = 20 (A. conyzoides) to
2n = 40 (A. haustonianum). Somatic variations are also found in Ageratum
conyzoides (2n = 30). This chromosome count has not been reported so far in
this species. However, it is a new count. Reported somatic chromosome
numbers of this genus are 2n = 20, 38 and 40. (Bir and Sidhu, 1980; Olsen,
1980; Mehara and Remanandan, 1975; Sharma and Varma, 1960; Jansen and
Stuessy, 1980). This variation fourid for the chromosome number results from
the establishment of different tytotypes giving a numeric chromosome
polymorphism for the species. For species Chromolaena odorafa, the somatic
chromosome number is found to txe 2n = 60. Eupatorium triplinewe root tip
cells showed 2n = 50 and both these populations, agree with the previous
reports (Olorode, 1974a; Khonglam and Singh, 1980). The counts made on
this genus Mikania reveal the codominance of two chromosome numbers 2n =
34 and 2n =36. Data in the literature also suggest a numerical chromosome
variation for the Mikania cordata species as 2n = 30, 34, 36 and 38. (Fedarov,
1969; Rabakonandrianina, 1987). Cytogenetic studies have shown that among
population of the same species there may be numerical variation, as o b s e ~ e d
by Maffie (1996) in Mikaria micrantha pollulation, where a numberical
variation of 2n = 36 to 2n = 72 war recorded.
Tribe: Astereae.
Four taxas belonging to 3 genera have been examined. The genus Conyza is
represented by two species. One with 2n = 54 (C. bonariensis) and the other
with 2n = 18 (C.canademis) in the present study. A review of literature
reveals that the chromosome numner in Conym ranges from 2n = 18 to 2n =
54, with a high frequency for the number 2n = 18 (Fedarov, 1969).
In Dichrocephala chrysanthemifolia the somatic chromosome number found
with 2n = 18 agrees with the prc:vious report (Mathew and Mathew, 1978).
Only one species of Erigergn studied was found to be with 2n = 18
(E.mucronnhrs). The previou!; reports show 2n = 18, 32, 34 and 36 (Carano,
1921; Mehra and Remanandan, 1974). Thus this species seems to be highly
plastic in its chromosome number.
Tribe.: Inuleae.
Only three genera have been examined. The genus Blumea the chromosome
number 2n = 18 and 2n = 28 in B.mollts and B.oxyodonta respectively and
2n = 36 in one cytotype (B.l~cera) investigagted. The result of the cytogenetic
analysis of the B.mollis populations also show a chromosome variation with
2n = 16 chromosomes. Chrc~mosome counting of the same species may show
different chromosome numbers, from the distinct cytotypes of the species. The
chmmosome numbers obsemed in members of Inuleae range from 2n = 14
to 28 (n = 7, 8, 9, 10, 11, 12 and 14) (Panchami and Vijayavalli, 1998). The
results of the cytogenetic analysis of the species Sphaeranthus indicus show a
chromosome variation will1 2n = 16, 2n = 18 and 2n = 20 and the most
frequent number was 2n = 20. This variation found for chromosome number
results from the establishment of different cytotypes, giving a numeric
chromosome polymorphisn~ for the species. In one member investigated in
the genus Vicoa, Vicoa ind;ca possesses 2n = 18. Previous studies (Mehra and
Sidhu, 1960) support the dominance of this number in Vicoa indica.
Tribe: Heliantheae
The chromosome numbers of hwenty two plants were observed; which belong
to 15 genara and 18 species. Chromosome numbers observed in the present
study were 2n = 22 in Acantho,permum hispidum; 2n = 72 in Bidens pilosa ;
2n = 24 in Cosmos bipinnatus and C. caudatus; 2n = 22 in Eclipra prostrata;
2n = 32 in Galinsogapmifror~~; 2n = 24 in Melampodium paludosm; 2n = 36
in Parthenium hysterophores; 2n = 30 in Sigesbeckia orientalis; 2n = 78 in
Spilanthes calva and S. radicans; 2n = 52 in S. ciliata and S. uliginosa; 2n =
40 in Synedrella nodifora; 2n = 34 in Tithonia diversifolia; 2n = 36 in Tridm
procumbens; 2n = 50 in Wedelia chinensis and in Wedelia trilobata and 2n =
24 in Zinnia etegans.
Data in the literature also sug:gest a numerical chromosome variation for the
Bidens pilosa species from 2n = 24 to 2n = 96 (Mariano and Morales, 1999).
The previous reports show that the number 2n = 72 is the most frequent, with
about 70% of the population coinciding with this results. Generally the
polyploidid organism have a greater genetic plasticity, due to a greater
genetic variability present in the genome.
On the contrary, Cosmos b i p i n m s is characterized by intraspecific
variations. The chromosome 2n = 24 occur in two cultivars (cv orange and cv
yellow) and C.bipinnates c\ oranage also show variation 2n = 22. The
occurrence of different cytoiypes in the same species is not very stable and
very alter either by the duplication or loss by individual chromosome (Morton,
1962). The dominance of the chromosome numbers 2n = 52 and 2n = 78 in
the genus Spilanfhes is very well established in literature. (Mathew and
Mathew, 1978; Mehra and R~:manandan, 1969).
hesent cytogenetic analysis of the Eclipta prosfmta shows a chromosome
variation with 2n = 18 and 2n = 22 observed in many plants analysed. This
somatic chromosome numbers agree with the previous report (Mohan et al.
1962). The chromosome number counted in the present analysis (2n = 22,24,
30, 32, 34, 36, 40, 50, 52 and 78) confirmed in all the populations studied in
this tribe, agrees with the previous reports.
Tribe: Helinieae
Seven populations belonging to the lone genus Tagetes have been examined.
This genus is characterized by intraspecific variation. The result of the present
cytogenetic analyses of the s x Tagetes erecta populations show chromosome
variation with 2n= 18,2n = 2 1,2n = 24 and 2n = 48 chromosome and the most
frequent number was 2n = 24 observed in three populations analysed. Only
one plant of Tagetespatula species studied was found to be with 2n = 24. As
regards the chromosome number this genus seems to be heterogenous with the
frequent occurrence of the somatic numbers 2n = 18, 24 and 48. A similar
trend has also been observed in literature.(Mathew and Mathew, 1980;
Shanna and Sarkar, 1967-1968; Federov, 1969). Intraspecific variation is thus
very well pronounced in this taxa. Variation in chromosome number may be
due to nondisjunction, somatic reduction or even by partial endomitosis.
Nondisjunction, in the somatic tissue, involves unequal anaphasic separation,
which results in unequal distribution of chromosomes in the daughter nuclei.
Tribe: Anthemideae
One plant has been examined. For the taxa Chrysanthemum parthenium, the
somatic chromosome number is found to be 2n = 18. The absence of any
somatic variation numbers suggests the stability of the genomes.
Tribe: Senecioneae
Three genera have been examined. Crassocephalum crepidioides root tip cells
showed 2n = 40. In the present study it is revealed that Emilia sonchifolia and
Notonia grandgora possess 2n = 10 and 2n = 20 respectively, agrees with the
previous reports (Gill et al., 1980; Narayan and Shukur, 1968; Federov, 1969).
Sub family: Liguliflorae
Tribe: Cichorieae
Sonchus oleraceus is the lone plant examined in this tribe. This taxa showed
the chromosome number 2n = 18. This is in conformity with the reports made
on the same taxa. However, 2n = 16, 18, 32 and 36 were also reported
(Federov, 1969) and frequent number is 2n =IS. Hence this species seems to
be highly plastic as far as the chromosome number is concerned.
b) Basic chromosome number
Basic chromosome number forms one of the widely used characters in
formulating phylogenetic speculations. It is generally regarded as a
dependable and stable marker of the direction of evolution and widely being
used in formulating phylogenetic speculations (Jones, 1970, 1974, 1978). It is
known that many basic chromosome numbers are involved in the origin of the
polyploid series in Angiosperms (Grant, 1982).
Based on the previous reports and the present investigation of somatic
chromosome number, an attempt is being made to discuss the possible
direction of basic chromc~some number evolution in this family. The
investigations are conducted on forty seven taxa belonging to thirty four
genera of nine tribes. The basic chromosome numbers vary widely in this
family and thus it appears that the family is a highly evolved one. Both
primary and secondary base numbers are involved in the evolution of the
various taxa studied. The primary base numbers (XI) range from 5, 7 and 9
and secondary base numbers (X2) range from 10, 11, 12, 13, 15, 16, 17, 18
and 20. The basic number X = 9 was found in majority oFthe members (30%)
followed by the numbers X. = 10, X = 12 and X = 11 with percentagel9,17 and
13 respectively. It is likely that protopolyploidy plays an active role in the
evolution of these basic numbers (Fujita, 1970). The ancestral basic
chromosome number of /isteraceae appears to be X = 9. The frequency of
polyploidy was 23% (Tomb et al. 1978).
Grant, (1981) proposes the original primary base numbers of angiosperms to
range From X = 7 to 9 and they seem to be ancestral in the phylogenetic sense.
From these primary base numbers many polyploid series develop both by
autopolyploidy and amphipol!lploidy. The latter involves both hyper and
hypoploidy (Fernandes and l,eitao, 1984). Thus it can be confirmed that
polyploidy had influenced and played a role in the evolution of base numbers.
Sub family: Tubuliflorae:
Tribe: Vernonieae.
The common base numbers of this tribe found in the present investigation are
X = 9 and X= 11. Phyllocephalum and Vernonia possess X = 9 and
Elephantopus X = 11. Literature evidences reveal the dominance of two
common base numbers X = 9 and X = 10 in Vernonia (Ayodele, 1994).
Tribe: Eupatorieae
In this tribe the common base: numbers found are 5, 10 and 18. The basic
chromosome number found in the lone member investigated in the genus
Adenostemma was primary in llature with X = 5 chromosomes. Other 3 genera
Ageratum, Chromolaena and Eupatorium are monobasic with the secondary
number X = 10, found in all the four investigated members. The existence of
these basic numbers was confirmed by Watanabe, (1990). Bremer et a1.(1994)
postulated that an initial reducrion in base number in the tribe Eupatorieae was
followed by an increase in the basic chromosome number.
A broad range of basic chronlosome numbers from X = 4 to 25, has been
reported in this tribe. 73% of the reported genera, and 53% of the reported
species have chromosome numbers based on X = 10. Thus the predominant
chromosome number among species, genera and sub tribes X =I0 was
doubtlessly regarded as the uitimate base number in this tribe by most of the
previous workers (Watanabc:, 1995). The basic chromosome number of
M i h i a cordota in the preser~t investigation is X = 18. Two base chromosome
numbers of I7 and 18 are also confirmed in Mikmria cordata by many
workers. There are many reports of 2n = 34, suggesting the occurrence of X
= 17 (Rabakonandrianina and Carr, 1987). On the other hand, some workers
(Grant, 1953; Turner and King,1964) have suggested that species with the
ancestral base number of X = 5 and 4 might have given rise through
successive alloploidy to those taxa with base number of 9, 10 and 17.
Tribe: Astereae:
The basic chromosome numbers found in the investigated members of this tribe is
X = 9. The present study includes 3 genera. The genus Conyza is mono basic
with the X = 9, found in two investigated members (C. bonmiensis, and C.
canadensis). Dichrocephala and Erigeron, both are diploids with
chromosome number 2n = 18. Thus the basic number is X = 9. Robinson et
al., (1981) has proposed X = 10 as the base number for the entire Astereae.
Tribe: Inuleae:
The basic lines in the tribe are >: = 7, 8,9, 10, 11 and 12. It is suggested that X
= 5 could be the ancestral basic number in the tribe. In the genus Blumea 4
basic numbers exist such as X .= 8,9, 10 and 1 1. It is suggested that X = 10 in
BIwnea could be the earlier evolved condition from which the lower
constitution found among thc: South Indian taxa which has evolved by
progressive reduction through the formation of B-chromosomes followed by
their elimination ( 8 t 9 t 10 +11) (Panchami and Vijayavalli, 1998). The
occurrence of 2 numbers (7 and 9) in 3 different species of Blumea,
investigated in the present study appears to c o n f m this suggestion. Similar
suggestions have been made by a few earlier workers (Mathew and Mathew
1987). But Mehra and Remanandan, (1975) suggested that Blumea is based
on X = 11. The basic numbers in other genera investigated in the present
study are X = 10 (Sphaeranthus) and X = 9 (Vicoa). In contrast Turner
(1970), when discussing possible base numbers in this tribe speculated that X
= 4 and 5 may be ancestral for the tribe with the former giving rise to X = 8
and X = 12 and aneuploid number of X = 7 and X = 13, respectively. Under
this scheme the base of X = 5 given rise to genera with n = 10 and
subsequently aneuploid derivatives with n = 9 and 11. This hypothesis was
suggested in past because of the apparent absence of taxa based on X = 6.
Tribe: Heliantheae:
Chromosome numbers also vary considerably in the tribe Heliantheae, ranging
from X = 4 to X = 19. However, X = 10 is rare, while X = 8 and 9 are
common and higher base numbers of X = 17, 18 and 19 are also prevalent
(Stuessy, 1977). h e basic chromosome numbers found in the investigated
members of this tribe include X = 9, 1 1, 12, 13, 17 and 20. Both ascending
and descending dysploidy, autopolyploidy and amphipolyploidy were found to
be responsible for the evolutic)n of various taxa coming under this tribe. Most
of the cytologically known genera are monobasic while a few other with
dibasic and polybasic conditions are also present. In the genus
Acanthospemum, which is 3 monobasic genus with X = 11. Most of the
previous workers are also of the opinion that X = 1 1 is the original number. In
the present investigation suggest X = 12 for the Bidens pilosa species. This
result c o n f m the result of Bamso (1991) of X = 12 for the Bidens genus.
Data in the literature also suggest a numerical chromosome variation for the
Bidenspilosa species from 2n = 24 to 2n = 96 (Mariano and Morales, 1999).
Cosmos the genus is monotmsic with the secondary number X = 12, found in
all the three investigated n~ernbers. Literature survey confirms the existence
of X = 12 in Cosmos (Mathew, 1978). Two different chromosome numbers
have been reported for this species by earlier workers such as n = 12 and n =
1 1 which indicate that this species exist at least in two cytotypes .
The genus Eclipfa exhibits the secondary basic number X = 11. Previous
workers also confirm the existence of X2 = 1 1 in Eclipta, hence chance for the
occurrence of amphipolyploidy from the primary base numbers XI = 5 and
XI = 6 as well as ascending or descending dysploidy from the respective
secondary base numbers are equal during the process of evolution.
Galinsoga parvifora in this present investigation shows n = 16. In the
absence of authentic basic chromosome number reports on this species of the
genus, the question regarding the basic number remains a matter of
speculation. The basic numter of this species may be X = 16. The base
number found in Melampodium paludosm is X = 12 which is secondary in
nature and well supported by an earlier study (Mathew, 1978). The basic
chromosome numbers in other genera are X = 9 (Parthenium and Tridar) X =
IS (Sigeskckia) X = 17 (Tithoiiia) X = 20 (SynedreNa).
The genus Spilanthes is monobasic with the secondary number X2 = 13;
found in all the four investigated members. The existence of these basic
numbers were confirmed by Mathew (1978). This secondary basic number X
= 13 might be a case evolved through polyploidy (Gill, 1970) involving the
primary base numbers X = 6 and 7. The present cytotypes with n = 39 and 2n
= 78, obviously constitute a polyploid series on the n = 26 type reported by
Mehra and Ramanandan, (1974) The reported occurrence of types with n = 7
as well as n = 12 indicates a pmsible polyploidy origin of the n = 26 type
through n = 13 which in turn originated by hybridisation of two lower
chromosomal types such as n = 7 and n = 6.
Two species of Wedelia examined in the present study reveals X = 10 is the
basic number of this genera. Brised on the available reports of basic numbers
in the genus Wedelia suggested a basic number of X = 10. But based on the
occurrence of n = 15 and 25 in Indian species Mehra and Ramanandan, (1974)
on the other hand, have suggested n = 5 as the possible basic number of the
genus. If X = 10 is taken as the basic number of the genus, the present two
species should be pentaploid. The basic chromosome number found in the
lone member investigated, in the genus Zinnia (Zelegans) was secondary in
nature with X = 12 chromosomc s
Tribe: Helinieae:
The tribe Helinieae possess chromosome numbers ranging from X = 3 to X
= 20 (Ito et al., 2000) All the four investigated members of the genus Tagetes
are multiplies of the secondary basic chromosome number X = 12. Evidences
from the literature reveal the predominance of this base number in the genus.
This secondary base number can be found only through autopolyploidy
(Stebbins, 1966) from the primiuy basic number, X=6.
Tribe: Anthemideae:
Chrysanthemum parthenium (2 n= 1 S), the lone member cytological1y screened
in this tribe reveals the presence of the primary basic chromosome number X
= 9, which has been confirmed by earlier reports.
Tribe: Senecioneae:
The base number for the tribe is X = 10 which is found in various multiples
(Robinson et al., 1997). From the present evolution both primary (X = 5) as
well as secondary (X = 10) base numbers seem to be prevalent in this tribe.
The secondary basic number X = 10 has been well established in
Crassocephalum (2n = 40) and Notonia (2n = 20). In Emilia sonchfofia the
somatic number found from this present study is 2n = 10. Thus the basic
number is X = 5. The higher basic number (X = 10) is considered to be
derived from lower number X = 5 by autopolyploidy (Stebbins, 1966).
According to Robinson (1997) occurrences of n = 5 in Emilia, n = I0 in some
Senecio, and n = 14 to 16 in members are considered as reduction.
Tribe: Cichorieae:
The basic chromosome number found in the lone member investigated,
Sonchus oleraceus was primary in nature with X = 9 chromosome. A new
base number (X = 10) in this genera is reported by Razaq et al. (1 994 ).
c) Polyploidy
Polyploidy is a very wide spread cytogenetic phenomenon found in over 30%
of dicotyledons and 50% w' monocotyledons (Love and Love, 1975). It is a
mechanism which involves multiplication of the whole chromosome
complement and there by an increase of gene number and variety, producing
radically different well adopte,d genomes (Stebbins, 1950). Polyploids are
considered to be more hardy and adaptable to extreme climatic conditions
(Hagemp, 193 1; Tischler, 1935; Love and Love, 19424 1943).
Polyploidy is one of the best genetical and evolutionary progresses, which has
greatly contributed to speciation and evolution of higher plants (Gottschalk
1985). This is mainly due to the ability of polyploids to increase the chances
of fertilization by breaking reproductive bamers, which permit natural
selection and establishment of species even under adverse environmental
conditions (Winge, 1917; Stebbins, 1971, 1974). There can be no doubt that
polyploid species are highly successful. A comparison of the geographic
distribution of polyploids ant1 diploids in plants show a greater adaptability of
polyploids (Love and Love, 1943). However, the evolutionary potentialities
of a diploid are likely to be g=ater in the long run (White, 1937).
The family A s t e m a e is characterised by relatively high frequency of
polyploidy. The present study reveals that herbaceous elements predominate
as compared with the shmb~y elements of the family. Growth habit is one of
the factors, which influence the frequency of polyploids in angiosperms
(Baquar, 1976). In Angiosperm, the highest percentage of polyploidy was
found in perennial herbs while annuals and woody plants have lower
percentage of ploidy (Stebbins, 1938; Fagerlind, 1944). Among the
polyploids, the majority are tetraploids. The various genera which show a
predominance of polyploicly include Adenostemma, Ageratum, Chromolaenu,
Eupatorium, Conyza, Blumea. Bidens, Parthenium, Spilanthes, Tridax,
Wedelia, Tagetes, Crassocephalum and Notonia.
Sub family: Tubuliflorae.
Tribe: Vernonieae.
The tribe shows a lower percentage of polyploids. Among the three members
studied, all are diploids with primary and secondary basic number X = 9 and
X = 11.
Tribe: Eupatorieae.
Different degrees of polyploidj has been observed in this tribe. The role of
polyploidy in the mechanism ol'speciation is obvious in the tribe Eupatoreae.
The cytological data obtained from the 6 members of this tribe indicated that 5
members are polyploids. MlRania cordata (2n = 36), the lone diploid
representative, exhibits a aneuploid variant number 2n = 34. Ageratum
conyzoides and Ageratum hmcstonianum (2n = 4X = 40) Chromolaena
odorata (2n = 6X = 60) and Eupatorium triplinewe (2n = 5X = 50) are
polyploids based on X = 10. There is no sign of a polypoid state at 2n = 20. In
addition, new polyploid series (3X, 4X, 5X and 6X) based on X = 10 have
developed extensively in the genera of Eupatorium and Chromolaena (Ito et
al. 2000). In the course of eletkrophoresis studies, found that diploid species
of Eupatorium with a chromosome number of n = 10 had extensive gene
duplications (Yahara et al. 1989). This finding casts further doubt on the
hypothesis that X = 10 was the ultimate base number of Eupatorium as well as
Eupatorieae, because diplolid vascular plants have a minimum highly
observed number of isozymes (Ciottlieb, 1981). A.conyzodies (2n = 40) exhibit
somatic variant (2n = 30), it is ;I triploid compliment of the basic set X = 10
chromosomes. Another member investigated in this tribe Adenostemma (2n =
4X = 20) is a tetraploid with the basic chromosome number X = 5.
Tribe: Astereae:
Out of four taxa investigated, one exhibit polyploidy. Two species studied
under the genus Conym, one is hexaploid (C. bonariensis) and other is diploid
(C.canadensis) These two sets ;ire evolved from the base figure X = 9. Other
diploid species observe in this tribe are Dichrocephala chiysanthemifolia (2n
= 18) and Erigeron mucronatus (2n = 18). All the investigated members of
this tribe are characterised by the absence of variants. The stability of the
chromosome number (2n = 18) indicated that distinctly well differentiated
genomes are involved in the origin of these diploids.
Tribe: Inuleae.
Out of the three taxa investigated in the genus Blumea, B. lacera and B.
Oxyodoto are tebaploids with 2n = 36, and 2n = 28 respectively. The diploid
BIumea mollis possess a somatic number 2n = 18. Here the cause for
ployploidy in the two species (B. lacera and B. oxyodonta) may be due to the
parallel evolution of two basic figures X = 9 & X = 7 and formation of
multiplies which develop into two series of polyploids during the course of
evolution. The diploid memter B. mollis (2n = 18) exhibits a t e t m m i c
variant number 2n = 16. This might have been probably caused by
multipolarity that leads to unbalanced chromosome compliment.
Sphaeranthus indicus is diploid with a basic set of X = 10 also exhibit
aneuploid somatic variants 2n = 18 and 2n = 16. Aneuploidy is very common
in this tribe at both generic and specific levels. It is caused by the gain or loss
of one or more chromosomes from the haploid set. The lone species of Vicoa
examined in this study, Vicoa indica is a diploid cytotype (2n = 18) with the
base number X = 9.
Tribe: Heliantheae:
In contrast to the other tribes, it shows a lower percentage of polyploids
among the members studied. Robinson et al., (1981) proposed that ancient
polyploidy played an important role in the early history of the Heliantheae,
based on the distribution of chromosome number. Acanthospermum hispidum,
the lone member of the genus Acanthospermum investigated in the present
study is diploid in nature (:ln = 22) with the basic figure of X = 11. The
result confms the result of Barroso, (1991) of X = 12 for the Bidens genus
and this is the most common member. Cytotypes with different ploidy levels
were found in previous reports, derived from X = 12, thus showing that
polyploidy were the main evolutionary process acting in B. pilosa (Mariano
and Morales, 1999).
In Cosmos the chromosome number counted, 2n = 24 which confirmed in both
populations (C. bipinnatus anti C. caudatus), agrees with the previous reports.
It belongs to a diploid level with basic number X = 12, which is the most
common in the genus, C. bipi.mafus CV. orange (2n = 24) exhibit aneuplopid
somatic variant (2n = 22). Ar~euploidy is common in this genus. It is caused
by the gain or loss of one or rnore chromosomes from the haploid set (White,
1937). The genus Eclipfa with one studied species shows, 2n = 22 based on X
= 1 1. This agrees with the pn:vious reports. Somatic variant (2n = 18) is also
found in this species which are multiples of the lowest one. The incidence of
polyploidy is very much striking in the genus Spilanthes. All the members
studied showed tetraploid or hexaploid condition, with the basic number X =
13. A perusal of the literature shows that so far all the investigated Spilanthes
species are at polyploid level. The present cytotypes with n = 39 and n =26
obviously constitute a ployploid series reported by Mehra and Remanandan,
(1974). The reported occurrence of types with n = 7 as well as n = 12
indicates a possible polyploid origin of the n = 26 type through n = 13 which
in turn originated by hybridisation of two lower chromosomal types such as
n=7and n = 6 .
The other members investigated in this tribe Galinsoga parvijora (2n = 2X =
32), Melampodium paludosm 1:2n = 2X = 24). Siegesbeckia orientalis (2n = 2n
= 30), SynedreIla nod~$ora (2n = 2X = 40), Tithonia diversifolia (2n = 2X =
34), and Zinnia Iegons (2n = :!X = 24) are diploid with basic number X = 16,
12, 15, 20, 17 and 12 respectively. The high basic numbers considered as a
secondary one. Such a high basic numbers might have originated either
through polyploidy followed by aneuploidy or dibasic polyploidy involving
the primary base number X = 7,8 and 9.
Other two taxa investigated in h i s tribe, Parrhenium hysterophores and Tridax
procumbetas, reveal the existence of tetraploidy from the basic figure X = 9.
An increase in the number of ~:hromosomes through autopolyploidy provides
increased possibilities for new gene combinations, which are of considerable
importance in evolution. The genus Wedelia, which was found to exhibit
pentaploidy in two species investigated. Moreover the vegetative mean of
reproduction, which is prevalent in the investigated taxa seem to bear a
correlation with the high degrees of polyploidy. Polyploidization might have
lead to the establishment of new gene combinations that have triggered off
new developmental changes leading to a later shift towards the asexual mode
of reproduction. (Stebbins, 1980; Gustafsson, 1947b). Thus the polyploidy
and aneuploidy was found to play an important role in the evolution of
Wedelia.
Tribe: Helinieae:
The various cytotypes of the genus Tagetes exhibits polyploidy and varying
degrees of somatic variation. The delicately balanced system of gene
interaction in tetraploids is disturbed by the doubling of chromosomes,
resulting in the formation of somatic variants (Kuckuck and Levan, 1951).
Out of four taxa investigated in this genus, Tageres erecta cv. orange and
rerecta cv. pale yellow are :etraploids with 2n = 48. The diploids Tagetes
erecta cv. yellow and Tagete,~ patula possess somatic number 2n = 24 each.
Each though both the polyplc4d and diploid taxa of Tagetes are multiplies of
the basic figure X = 12, the liigher levels of ploidy attained by T. erecta cv.
orange and T. erecta cv. pale yellow probably point towards their greater
adaptability. The aneuploid and hypoploid variations found in their polyploid
taxa reveal the on going genetic and evolutionary processes which may help to
break the reproductive barriers enabling natural selection and thereby
speciation (Stebbins, 1974).
Tribe: Anthemideae.
The single taxa investigated in the tribe Chrysanthenum parthenium is
characterised by its diploid stales from the base figure X = 9.
Tribe: Senecioneae.
A brief review is offered by Robinson et al. (1997) of major chromosome
number variations in the senecioneae based on recent delimitation of the tribe.
According to him the base number for the hibe X = 10, which is found in
various multiplies. Some mc:mbers with a base of X = 9, considered an
aneuploid reduction from X = 10. Number based on X = 30 and aneuploid
reduction are found in many Australian Senecio. Three members are studied
in the present investigation, two are diploid, and one is a tetraploid.
Crassocephalum crepdioides (:!n = 4X = 40) and Notonia (2n = 2X = 20) have
the basic chromosome number X = 10; and Emilia sonchifolia (2n = 2 X = 10)
is X = 5. Occurrence of n = 5 in Emilia is considered as reduction.
Tribe: Cichorieae.
Sonchus oleraceus the lone t;vca investigated is diploid and with X = 9 as the
basic chromosome number
d) Karyomorphometrical analysis
In Angiosperms the specie; of several families both dicotyledonous and
monocotyledonous are found to exhibit a direct relationship between their
phylogeny and the chromosome constitution. The chromosomes being the
carriers of heredity, both 3tructural and numerical changes in them can
influence the genetic evolutionary process at work than do any other type of
changes. Detailed information regarding the chromosome architecture in
higher plants can thus serve as a useful tool to understand their systematic
relationships and for tracing the trend and direction of their evolution (Love
and Love, 1975).
Some of the major karyotyp characteristics of considerable evolutionary and
taxonomic significance are ( I ) differences in the absolute chromosome size (2)
differences in the position $of centTomere (3) differences in total chromatin
length (4) differences in karyotype formula and (5) number as well as position
of satellites.
Since, most of the members of Asteraceae possess small chromosomes the
detailed karyomorphological study by conventional method is not easy. So far
only a few plant species wen. reported for their karyomorphology. An image
analysis system provides a better opportunity for studying the various
chromosomal parameters. 'Illis method has only been used recently in plant
cytology and chromosome studies (Fukui, 1985, 1986). It appears to be a very
powerful tool which to generate the quantitative chromosome data and
idiograms, which cannot be obtained by conventional methods. This system
has achieved a drastic reduction of the researchers' time and efforts while
maintaining a high standard of information in the quantification and
karyotyping of chromosomes. It has become possible to obtain various kinds
of quantitative data on the chromosome images. The length of the
chromosomes and the ratio between the short and long arm values has
represented almost all thr: numerical information available on the
chromosomes for the past decades. It has become however possible to obtain
not only one dimensional daia of the chromosomes such as length, but also
two and three dimensional data of area and volume within a limited time by
using image analysis system. Also it is possible to make an accurate pairing
by PerimeterIArea (PIA) ratio mainly in those cases where chromosome sizes
are really small. Relative chromosome Length (RL) and Arm Difference ratio
(AD) are also considered to find out homologous chromosomes for accurate
pairing. Chromosome pairs whose ADS and RLs are not significantly different
(P<O.OS) were judged as morl~hologically identical (Watanabe et al. 1990). In
addition to this imaging techniques provide pseudo colouration, which is a
very useful tool for identifying the primary and secondary constriction of
small chromosomes very c1t:arly. Idiograms can be prepared semi
automatically and by using computer devices. By the traditional methods of
analysis this procedure would be more difficult due to the small size of
chromosomes. At the same time biases in the analytical processes originating
from the differences in the researchers skill and experiences can be minimised
by employing this method. Hence this method would provide useful
information for chromosome analysis and may be helpkl as an essential tool
in chromosome research (Fukui, 1988; Fukui and Kakeda, 1990; Iijima, 1991).
The general feature noted i r ~ the family Asteraceae is the wide range of
chromosomes with small sized chromosomes in most of the species.
However, the chromosome c:omplements in the various members differ in
minute karyotypical details (vide Table 49). With regard to the gross
morphology, chromosomes a-e nearly sub median to nearly median in nature.
The chromosome range from1 2.54 to 0.41 pm in length. The chromosomes
with secondary constriction ranges from 2 to 8 in number. The average
chromosome length (ACL) baries from 0.74 to 2.3 pm. The total chromatin
length (TCL) shows a very wide variation with 14.82 being the minimum and
90.32 being the maximum value. The disparity index values ranges between
18.13 - 48.63, the coefficier~t of variation ranges from 13.49 - 35.97 and total
chromatin index (TF%) ranges from 40.3 - 50.74.
These variations found in the karyotypic parameters suggest that Asteraceae
members are characterized by symmetrical to slightly asymmetric karyotypes.
An increase in the range of chromosome length as well as the increase of sub
metacentrics at the expense of metacentrics is accompanied by an increase in
the coefficient of variation li:ading to a symrnelry (Stebbins, 1958). Thus
karyomorphological studies are of considerable importance in order to throw
light on the phylogenetic '-elationship among taxa of flowering plants
(Iwatsubo and Naruhashi,1991).
Sub family: Tuhuliflorae
Tribe: Vemonieae
Three genera represented in the present study are diploids. As regards the
chromosome with secondary constrictions Phyllocephalum rangacharii and
Vemonia cinerea possesses four chromosomes while Elephantopus scaber has
only two. The karyotype o!' these taxa is somewhat symmetrical and thus
shows primitiveness. Comparatively lower variation wefticient and higher
TF% is also primitive featun:. The range of chromosome length is lesser in
Elephantopus scaber as compared to other two members. Normally a low
disparity index value correspc~nds to the homogeneity of chromosomes in most
of the higher as well as lower plants (Mohanty et al.1991).
Tribe: Eupatorieae.
Five genera studied for their detailed karyomorphology. Chromosomes are
moderately small with symn~etrical karyotype except Chromolaena odorata.
In three members (Adenosternma, Chromolaena and Eupatorium) the absence
of 'C' type chromosome is a speciality. The differences observed in the
karyotype formula (KF), averase chromosome length and chromatin length of
haploid complement (CLH) among different taxa might have been probably
due to minute structural alterations of chromosomes. Speciation depends more
on chromosomal rearrangements and mutation of individual genes, than on
changes in the total amount of genetic content (Stebbins,l959). The high
value of average chromosome length shown by Chromolaena probably show
its primitiveness, where as the lower value for this parameter found in Mikania
denotes its evolved nature. A decrease in chromatin length is one of the
factors responsible for evolution of higher plants (Babcock and Cameron,
1934). The comparatively high disparity index (Dl) value found in Ageratum
followed by Mikania cordata corresponds to the heterogeneous assemblage of
chromosomes in these taxa.
Tribe: Astereae.
Karyomorphology of four taxa have been studied. The karyotype in these taxa
studied is found to be more or less homogenous and symmetric. The
chromosomes are found to t)e medium in size. All the members except one
species of Conyza (C. bon0riensi.s ) are diploid. Number of chromosomes
with secondary constriction is two in all the taxa studied. Karyotype formula
is symmetrical in all the cases except C.bonariensis, with the absence of 'C'
type chromosomes. All the other parameters (Dl, VC and TF%) seems to be
more or less symmetrical i n all these investigated members. A symmetric
karyotype is considered to t ~ e a primitive one (Stebbins, 1959). Even though
the karyotypic features of these members show general uniformity, in finer
details they appear to show recognizable difference with regard to the
distribution of secondary constrictions, centromere position of a few
individual chromosomes and there was no appreciable intra karyotype size
differences. It is remarkable that only minute structural alterations exhibited.
In the light of chromosome data and karyomorphological information of four
members reported here, it appears that both numerical and structural alteration
in chromosomes have not played any major role in speciation and evolution of
this tribe.
Tribe: Inuleae.
Karyomorphology of five taxa have been studied. The different species of
Blumea are exceedingly variable both in chromosome numbers and in
morphometric characters of their karyotypes. This variability of the genus
evidently reflects an important side of its evolution. The number of
chromosomes with secondary constrictions varies in different species. In
diploid taxa they are one pair ir~ number. The tetraploid species possess one to
two pairs of satellite chromos~~mes. The three species are characterized by
their homogeneous symmetrical karyotypes with lesser range of chromatin
length, a primitive condition. Among the karyotype of five taxa Vicoa indica
exhibits a short sized chromosomes, comparatively low values of average
chromosome length, disparity index as well as variation coefficient. From the
table 49 it is clear that the higher values for average chromosome length,
chromatin length of haploid complement, along with a low variation
coefftcient reveal primitivene:;~ of Sphaeranthus indicus over Vicoa indica
and Blumea.
Tribe: Heliantheae
Twenty taxa were studied karyomorphologically. The data indicate that, all
the tax8 have more or less symmetrical karyotype, which is considered to be
primitive. On a close examination of karyotype of these members it reveals
that karyotype asymmetry is progressively greater among the higher
polyploids. Similarly karyoqpic size difference of chromosome which is
brought about by differential deletion of segments of individual chromosomes
as well as through occurrence and establishment of unequal transaction
between non homologous chromosome is also seem to be greater among the
polyploids. The evidence thl~s indicate that the above factors which lead to
karyotype speciation also haw played some role, along with polyploidy, in the
evolution of genera.
For species in Spilanthes witn X = 13, the size ranged from 0.49 - 1.67 pm.
For species in Cosmos with >: = 12, the size ranged from 0.7 - 1.49 pm. For
species of Wedelia with X = 10, the size ranged from 0.41 - 1.15 pm.
Spilanthes and Wedelia species are polyploids compared to the diploid species
of Cosmos. Thus the significant chromosome difference between genera is
also confirmed. In Wedelidl and Spilanthes the lesser values for all the
parameters of karyomorpholclgy show its rather evolved nature. Such type of
variation is important from the evolutionary view point. Long chromosomes
with symmetrical karyotyp~: are further evidence of primitiveness
(Sharma,1984). In Tridaxproc-umbens even though the karyotype shows some
trend towards asymmetry, the higher values of average chromosome length,
chromatin length of basic corriplement reveal its trend towards primitiveness.
The chromosome pairs with secondary constriction are varying from one to
four in various taxa. The kar)~omorphological differences found among these
taxa fully justify that speciation and evolution has been principally affected by
minute structural alterations.
Tribe: Helinieae
Concrete conclusion cannot be drawn from the karyomorphological features
exhibited by the closely related genetic strains of Tagetes erecta (cultivars
orange, pale yellow and yellow) and Tagetes paluta . So it seems likely that
they represent three closely related lines has been found in evolution of their
morphological features. However, at the infra specific level Terecta cv pale
yellow is characterised by high value for major karyomorphometrical
parameters such as total chromatin length, average chromosome length.
chromatin length of haploid complement are considered as primitive
characters.
Tribe: Anthemideae
Chrysanthemum parrhenium is the lone member studied in the tribe. The
higher karyomorphological values exhibited by it denotes its primitive status.
However the vast difference of chromosome length and a high disparity index
value denote an advanced heterogeneous nature.
Tribe: Senecioneae.
Three genera were studied karyomorphologically. The number of
chromosomes with secondary constriction varies in different genera. In
diploid taxa they are one pair in number. The tetraploid species possess three
pairs of satellite chromosomc:s. Ernilia and Notonia are characterised by
higher values for the major karyomorphometrical parameters and
comparatively low variation coefficient. Thus these observations are in
advocacy with the primitive status of this genara among all the members
studied in this present investigation. Another notable feature of this species is
the absence of 'C' chromosonles.
Sub family: Liguliflorae.
Tribe: Cichorieae.
Sonchus oleraceus, the lone plant has been studied in this tribe is diploid in
nature with low chromosolne number (2n = 18). In this genus, the low
disparity index (Dl) value reveals the homogeneity of the chromosome
complement. The high 'IF% found in conjunction with a low variation
coefficient value reveal that S. oleraceus is a primitive plant. A high TF% and
low variation coeficient ,falues correspond to the primitive status in the
evolution of flowering plans. (Stebbins, 197 1 ).
e) Cytological evolution in Asteraceae
From the cytological observations made in the present investigation it can be
concluded that speciation and c:volution with this family has been possible as a
result of increase in variabilit) through changes in the base numbers, as well
as numerical and structural changes in chromosome numbers. The various
cytological phenomena like protoautoploidy, amphiploidy, ascending and
descending dysploidy might have resulted in the variability of base numbers in
the family. The wide range of :hromosome numbers observed in many genera
in the present investigation marks a significant role that aneuploidy and
polyploidy have played in the evolution of various taxa of the family at the
generic and species level. It also appears that various kinds of aberrations
have played a vital role in the evolutionary diversification of the family. The
mitotic and meiotic irregularities might have lead to structural and numerical
variations in the chromosomes of a species (Roy, 1998). Individuals with the
same chromosome number but with differences in karyomorphological details
reflect the ongoing evolutionary processes at micro level.
It seems probable that in Asteraceae, Robertsonian changes or mutations
might have also played an important role in the evolution of karyotype.
Drastically mutated individuals are usually unstable and unfit to survive in
nature because they express various degrees of weakness and chromosomal
aberrations leading to genetic sterility. However, some individuals canying
the changed chromosomal constitutions are well within their range of
tolerance. This was confirmed by the occurrence of normal meiosis in some
polyploids investigated. Moreovc:r, most of the taxa belonging to this family
have efftcient means of vegetative propagation. This ensures the survival of
these genetically altered types which otherwise would have faced exlinction
on account of sexual sterility imposed as a result of these changes.
Accumulation of such small changes can sometimes lead to a taxonomic
divergence in a species during the process of evolution. Thus meiotic
accidents may prove to be ir,ore useful than mitotic aberrations from the
evolutionary point of view, sirlce the meiotic abnormalities are hereditary and
likely to multiply and establish in a population.
M e very high chromosome number coupled with small size may indicate a
higher evolutionary status of !:his family. Comparing with other genera Emilia.
Notonia and Blumea posses3 large sized chromosomes which arc fewer in
numbers. But the chromoscmes of Agerafum, Euparorium, Spilan/hes and
Wedelia are small sized and more in number. The family Asteraceae seems to
be in a fairly active state of t:volution because of the quite common occurrence
of polyploidy and aneuploidy. Mutation of genes, structural changes in
chromosomes, non-disjunction, chromosomal rearrangement and several other
abnormalities are those mec.hanisms leading to the development of polyploidy
and thus to the differenti>dion of new taxa in Asteraceae (Femandes and
Leitao, 1984). Recent molecular and cladistic analysis has revealed that the
tribe Eupatorieae originated from the tribe Heliantheae (Ito et al. 2000). Based
on the available data from chromosomal and ntolecular phylogenetic studies of
the tribes Helinieae, Heliaitheae and Eupatorieae show that a polyploidization
polyploidization event occurred during ihe course of the divergence of the
tribe Helinieae and Heliantheae. Changing the chromosome base number and
the one of the polyploidy progeny was the ancestor of the tribe Eupatorieae
(Ito et al. 2000).
There are still enormous gaps in our knowledge as regards the cytological
evolution of Asteraceae, and much still remains to be done before a major
cytotaxonomic review may be attempted.
SUMMARY
Forty seven taxa representing a total of thirty four genera of Astcraceae are
studied for their detailed karyctmorphology. The chromosome spectrum of
Asteraceae ranges from 2n = 10 to 2n = 78 with majority of the species
concentrated in the number 2n == 18 followed by 2n = 24,2n = 36 and 2n = 40.
In spite of the wide range of chromosome numbers, there exist a relationship
between the different genera and species as evidenced by the frequent
occurrence of numerical variatton. This wide range of chromosome numbers
may be due to the difference in numbers in chromosome hiotypes belonging to
different groups. From the prt:vious literature and the present investigation it
is clear that many genera like Ageratum, Blumeu, Sphaeru~~hus. Cosmos and
Tagetes exhibit inter, intra and infra specific variations among chromosome
numbers. Presence of such widely different series of chromosome numbers in
the species of even same genus and in genera placed under different tribes and
subtribes, indicate that the difcerent chromosome numbers may be derived one
from other.
The basic chromosome numl~ers are found to be varied in Asteraceae. This
variability in the number of chromosomes at the basic level could possibly be
the result of aneuploidy at g.eneric level. Both primary and secondary base
numbers are found to be ~nvolved in the evolution of forty seven taxa
investigated. The primary base numbers (x 1) range from 5. 7. 9 and
secondary base numbers (x 2 ) range from 10, 1 1, 12, 13, 1 5, 16, 17 18 and 20.
The basic number x = 9 was found in majority of the numbers (30%) fbllowed
by the numbers x = 10, x = 12 and x = 11 with percentage 19. 17 and 13
respectively. The ancestral bzsic chromosome number of Asteraceae appears
to be x = 9.
In the present investigation 55.31 % are diploids and 44.68 % are polyploids.
Among the polyploids the majority are tetraploids. The various genera which
show a predominance of ployploidy include Adenostemma. Ageratum,
Chromolaena, Eupatorium, Cw~yzu, Blumeu, Bidens, Parthenium, Spilunthes
Tridau, Wedelia, Tagets, Cr~~socephalum and Notonia. The role of both
polyploidy and aneuploidy in the mechanism of speciation is obvious in
Asteraceae.
The computer aided image analysis system provides a better opportunity for
studying the various chromosomal parameters especially when the
chromosome size is very small. The method of using conventional numerical
parameters such as length and arm ratio are insufficient to distinguish the
chromosomes of many memlxrs of Asteraeace. By using image analysis
system it has become possible to obtain various kinds of quantitative data such
as length, area, perimeter and visual three-dimensional volume on the
chromosome images, idiogram can be prepared by using computer devices.
Hence this method would provide useful information for various
Karyomorphological analyses.
The general feature noted in the family Asteraceae is the wide range of
chromosome number with graded symmetrical karyotypes. However, the
chromosome complements in the members differ in minute karyotypic details.
With regard to gross morpholc~gy, chromosomes are nearly sub median to
nearly median in nature. The chromosome was found to range from 2.54 to
0.41pm in length. The total clromatin length shows a very wide variation
with 14.82 being the minimum and 90.32 being the maximum values. The
disparity index values were fout~d to range from 18.13 to 48.63 and the mean
centmmeric Index value (TF%) from 40.3 to 50.74. The coefficient of
variation ranges from13.94 to 35.97. .Thus the various micromorphological
details of the karyotype like, differences in absolute chromosome size,
difference in position of centromere, differences in total chromatin length
difference in karyotype formula and difference in the number as well as
position of the satellites, vary from cytotypes to cytotype. These variations
found in the karyotype parameters suggest that Asteraceae members are
characterized by symmetrical to slightly asymmetrical karyotypes. The
karyomorphological diversity found in the family shows that the group is still
undergoing active speciation.
The presence of a wide range 'of chromosome numbers, numerical variations
and structural changes of chromosome found in many genera mark the
significant role that both aneuploidy and polyploidy have played in evolution
of various taxa of the family a1 the generic and species level. The variability
in base numbers might have been resulted through protoautoploidy,
amphiploidy, ascending and descending dysploidy. Both mitotic and meiotic
aberrations have played a ma-or role in the evolutionary diversification of
family. Individuals with same chromosome number but with differences in
karyomorphological details reflect the ongoing evolutionary processes at
micro-level. It has also been found that in Asteraceae, Robertsonian changes
or mutations might have also played a major role in the evolution of
karyotype.