07 Chapter 1

PART-I
KARYOMORPHOLOGICAL
STUDIES IN ZINGIBERACEAE
Prepared by BeeHive Digital Concepts Cochin for Mahatma Gandhi University Kottayam
Zingiberaceae (ginger family) is generally regarded as one of the most highly
esolved, natural and rather distinct of monocotyledonous families. The
members are distributed mainly in tropics and subtropics with the 'entre of
distribution in the Indo-hlalavan region,but extending through tropical Africa
to central and South America. They are perennial rhizomatous herbs in shad\
habitats and are characterized bv the possession of a tuberous and:or creeping
rhizome w~t h an aerial shoot (pseudostem) often covered by sheathing leaf
bases. The inflorescence is usually a spike or raceme. The flowers are
hlsexual. always epigynous and zygomo~hi c or asymmetric. Commercially
the family is of great importance, since its members find use in medicinal,
perfumery, flavour and cosmetic industries. Quite a large number of plants are
k i n g used as omamentals too.
There is considerable difference of opinion regarding the number of
constituent tava within the family. Schumann (1903) recob-ised the
zingiberaceae as having 38 genera and 800 species, whereas Bailey (1949)
recognised only 40 genera and 400 species and Sabu, (1991 ) ahout 45 genera
and 800 species. The number of valid genera as accepted today is 50 with
about l000 species (Dahl gen c/ (l / . : 11985'1. Burtt and Smith (1972) modified
Schumann's (1904) classification and proposed a new infm-famil~al
class~fication of the family Zingiberaceac and the work followed here is on the
basis of their classification. In this classification the family is subdivided into
! \ \ o sub families viz., Costoideae and Zing~heroidcac. ilnder the sub fam~l!
Zinyiberoideae four tribes are tnere viz.. Globbeae. Zingibercae, tledychieae
and Alpinieae.
In lndia the fam~l! i s \veil represented by around 21 genera. The members arc
ma~nly confined to the llinialayas. South '.Vest Indin. Ucsteni Ghat\ anil
Yorth Eastern India. Of the 21 genera present in the natlve flora of 1nd1a.I 1
arc rcpresented in South India which form aboiit .iOi'o of I UUI genera occurring
i n the \\bolt. of l nd~a. T'he 1nl.r. ilcd!ch~eac. ; S \\cl1 rcprr.seritcd i n South lrid~a
wi:h maximum number of 5 genera and 25 species. The Alpinear is the
second largest tribe with three genera and I S species. t r n ~ ~ l is the largest
genera of the family i n South India with 17 species.
From an economic stand point, Zingiberaceae form an important goup with
considerable economic potential with plants such as AfiLlmorn7mm. .-imornrrnt.
(' urcumu, Elt~rturru. Kurtnpfi ri c~ and Zingiher. Many members of the family
are used therapeutically in ayurvedic and other traditional systems of medicine
from time immemorial and some are well known spices. The
pharmacologcal, medicinal, economical and other industrial uses of various
t aw haw been discmsed in the second part of the dissertation.
The Zingiberaceae of South lndia are chosen for the present study. Till the
pioneering studies undertaken by Sabu in 1991, on the taxonomy of this
family, the kno\vledge about South Indian Zingberaceae was frarmcntay and
scattered. Nowadays most of the forests are fast disappearing due to
encroachment. Moreover,many anthropogenic activitlcs such as over
exploitation of the plant parts for medicinal use and food have placed many
taxa as endemic species. Ta.xa such as Kuempferru ~ ~ ~ l u n g u , Kocnipferru
rolundu and :2lpmru gulungu are under severe threat. Around 7 species of
(' urcumu are endemic in South India (Mangalv and Sabu, 199; I
Cyt ol o~y has been widely ut ~l ~sed in tackling taxonomical problems in the
recent past. I t is believed as a dependable tool for solving taxonomic
problems and for elucidating systematic relationships. phylogen! and
evolution of related plant groups. The informat~on like chromoa~mc number.
structure, morpholog~ and behaviour during mitotic and meiotjc di\~sions
have been of considerable value in understand~ng interrelat~onships and
delimitation of taxa [Den ilartos et (11. 1979; Yoshikane and Naohiro. 1991 1.
There haw been many cuamples \\here liner detail! ilt chromosome
morpholon ha\e yielded helpful clues for tracins the direct~on of kapot!pc
bariations. espec~ally at l owr tasonomlc Icvels.
P~oneering attempts on the cytology of the Zingiberaceae are those by
Monnga e/ (11. (1929) and Boehm (1931 ). Raghavan and Venkatasubban
(1943) determined the chromosome number of 24 species, but since their
technique was based mainly on the sectioning method, vc n little data
regarding the chromosome morphologv were gl\en. Nevertheless, this work
laid the foundation for later researchers Sharnia and Bhattachaya (1959)
carried out a detailed karyotype analv. sis of '6 species of Zingiberaceae.
Further information is available on this family by virtue of the major
c)tological surveys conducted by Ramachandran (1969a and l969b): Siew
ngo (19691, Mahanty (1970) and Omanakumari (1989). Also many other
attempts have been made in various species of the family viz., Bum and
Olatunji (1972); Prasad and Abraham (1981), Beltran and Kiew (1984),
Ratnambal(1984).
Out of the 1000 species so far around ren percentage of species are reported
for their chromosome number. It is quite remarkable that little attention has so
far been paid to the detailed cytological investigation and phytochemical
screening of members of Zingiberaceae. The c)-tological studies undertaken
so far were concentrated mainly on the determination of chromosome number
and have provided scarcely any data on the chromosome mo r p h o l o ~ and
structure in detail.The detailed karyomorphologicl reports known are centered
nlainly on a few genera such as Zlnglher and K~~c?npfc,r/ii. Moreover all the5e
karyomorphological studies depended on the convcntional methods. in
Zingiberaceae many of the members possess \en. small sized chromosome5
and hence detailed kanotype analysis is ve n dttlicult by routine procedures.
The paucity of karyornorpholo~ical data on the family Z~ngibcraccac ma\ also
he largel! due to intraclonal \anations and the \c.n high cvtoplasm~c contents
making critical analysts dilficult and time consuming.
itnall chromosomes in plants are \ e n ditlicult to idcntify h \ or d~nan
karyotype analysis. I n many cases besides the s~nallness of the chromosome
iire. chron;osomal shaprs are somett~ncs sirn~lar I,) rach other. e\en i n [he
<; xc~es h a \ ~n s large chrcrrnoson~es rnakin: the anal\s~q \c.n d~fticult Oi i en
s~~l nabi l ~t ! of chr omo~ur r ~e~ IS not al\\a\s silod 011l y N and C.'-hanti~ns
methods are presently available in many cases. These techniques ho\vever
cannot always be suited especially to the species with small chromosomes
\\here only limited success have been obtained (Gostev and .45kcr. 1979:
Ladizinsky et a/. 1979). Moreo\er, plants with cells having thick cell wall
prevent a good spread of chromosomes by squashing method. Therefore. the
ordinary karyotype analysis has provided only limited success. from the
viewpoint of chromosome identification not only in plants vith small
chromosomes, but in many other plant species as well. The quantification of
the image data of chromosomes has been almost impossible except for some
simple parameters by ordinary means. The length of the short and long arms
and their ratio represent almost all the chromosome image inforniation now
available. These parameters however, are not very accurate in identi@ing
plant chromosomes in many cases. We, need a system which can objectively.
easily and speedily provide quantitative chromosome image data based on
fixed procedure. Devices of microelectronics and image analysis methods
h a ~ e been developing rapidly in these years and now the!. can supply
indispensable tool for developing an image analysis system for plant
chromosomes.
Recent development of microelectronic techniques (Fujita. 1983) make 11
possible to analyse image of human chromosomes automatically (1.undsteen c,/
ul. 1980: Taylor and Graham, 1980). Karyomorphological studies h!. image
L'enetic analysis system (IAS) provide a better knowledge on the cyto,
constitution of various species over conventional methods. The trad~tional
basis of measuring and characterising chromoso~ne complements l . ! r~sual
e\aluatlon has been subjective and imprecise. These limitations may he
overcome by using image analysis system for direct quantitative asci.ssrncrlt of
each chromosome and this allows ior a bcner kno\vledge of thelr c!-toenstic
constitution (Verona and Galaseo. 1995: Fukui and Kaml s ~~g~. 1995)
Semiautomatic karyotyp~ng including numerical data acqu~sitlon. :ii!rir,; and
arrangement of chromosomes by digital manipulation of the image us1n2
c~mput er de~l ces results in a detallecl construct of descr ~pt ~\ e d,lrd iir.tai1c.d
studies prtalning to chromosome lengh. area. perimetc.r. \ :\ual zj.pnrrn: i!iri.r,
dimensional volume, uniformity coefficient, variation coefficient. disparip
index of chromosomes and total forma percentage (mean cenuornsric index
value or TF?/o) are made in order to throw light on phylogenetic rclationshtps
both at the intra generic and intra specific level.
Thus the various cl-tological parameters like chromosome number. base
number ploidy level and quantitative karyomorphological data are effectivel!
used as classificatory criteria in the same manncr as the n~orphological
characters; since chromosomes have a direct relationship to the genetic system
of which they are an integral part (Den Hartog er al. 1979).
Therefore the present investigation is an attempt to ach~e\ c a better
understanding of the family Zingiberaceae and kar)iomorphological analysis
conducted on 43 taxa with the aid of improved techniques (Sharma and
Sharma, 1980; Fukui and Kamisugi, 1995). The various taxa have been
examined cytologically and chemically in order to access their taxonomic
status. Further an attempt is also made based on the previous and present
cytological data to decipher the inter relationships among the various taxa to
obtain a clear understanding of evolutionary process at work in the famtly
Zingiberaceae.
Forty three tasa of Zingiberaceae has been collected from different parts of
South lnd~a. Ciermplasm collections were made from the ivild as well as
cultivated areas of tropical South India. Voucher specimens are deposited in
the herbarium of Sacred Heart College Thcvara, Cochin, Kerala, India. The
list of various taxa studied are given in the table 1.
I Castus malortieanus Wendl (Fig l a )
Perennial herb wlth root stock; errect system with highly pubescent leaves:
inflorescence in dense globose or ovoid terminal spikes: tlower large, white.
This is an ornamental species.
2. C. speciosus (Koenig) Smith ( Fig. 2a )
Perennial succulent herb with a tuberous horizontal root stock; steins spirall!.
misted so that the leaves appear spirally arranged; leaves oblong or oblong-
lanceolate acute, glabrous above; flowers white in dense globose or ovoid
terminal s p ~ kes.
3. Globba cerrtua Baker (Fig. 3a )
Herbs with creeping conical small rhizomes: manv thick tlesh! roots.
psrudostem slender. tleshy, l o~rer vegetative bracts puqle i n colour: leait.\
oblong. lanceolate, glabrous. shortly prt~oled. ligule short: inllorcscence
terminal; flowvers orange yclloiv i n colour
1. G. ophioglossa Wight (Fig. 4a )
Herbs wvith creepng small rhizome: pseudos[crn slcnder. 40-7il cm hl f l ?.
l eai t s Icntar to clliptic or oblong-!3nccoi.~ie Intloreiccncc pnr: ~ci e i \ !t i ; l>\\
bulb~ls. tlo\\ers paie to deep !ell<)\\
5 . Z cernuum Dalzell (Fig 5a)
( Z nimmonii Graham )
Leafy stem 1-15m high: leaves sessile,oblong-lanceolate: acuminate 15-3Scln
long; spikes oblong or subglobose, sessile, bracts green streaked with red;
flowers reddish yello\v, lip yellow, seed dark with aril.
6. Z neesanutum (Graharn) Rarnamoorthy (Fig. 6a)
(2 mnc&tachjum Dalzell)
Leafy stern upto 1.5m high; leaves sessile, leniar, oblong-lanceolate;
pubescent beneath, spike cylindric, 15-30cm long, peduncle 30-30cm long
with large sheathing scales, flouers white or geenish \vliite. lip pale yello\v,
striped purple: seed dark purple with white aril.
7. Z. ofJcit~ale Roscoe (Fig. 7a)
It is the common prier plant perennial herb; leaves sess~le. lanceolate, spike
oblong-cylindric, 3-10 cms long: flo\hiers greenish-!.ello\t. lip dark purple.
o%cn spottcd 1~1th yellotv.
7. Z purpureum Roscoe I Fig. 8a)
iZ cassuminnr Rorh I
Pcrcnnial herb. lea5 stem 1 - 1 Jill high; rhizome !t.llo\\iih inside, leaves
subscssile. oblong-lanci.olatc, acute or acuminate, pubescerit. spike> dcnse,
fusifonn or ohlong-cllipsoid \vith long peduncle. t!oi\ers irh~re. lip !cllo\tlsh
\ \ h ~t c .
9 2 roseum (Roxburgh) (Fig.9a )
Leafy stem l - high, l~bwle of leaf membraneous ; leaves ohlong-
lanceolate, acuminate. sessile, 10-60cm long,l0-l2 cm i n \side, pubescent
helow; spike short, dense, oblong. about 8cm long, bracts red,pcduncle short.
tlowers bright or pale red. lip whitish \sith red markings, seed dark with white
aril.
10. Z. ierumbef (1.inn.) Smith (Fig. 10a)
Perennial aromatic herb; leafy stem 60cm to 1.5 mt high. lea\c.s glabrous
heneath, oblong-lanceolate. sessile, spike oblong. dense with long pedunclt,
flowers pale sulphur yellow.
1 I . Curcuma aerugenosn Roxburgh (Fig. I l a)
Rhizomatous perennial herb. Iafi stem 60cm-lm high_ leaves, long upto 60
cms, oblong, laneolate, acuminate. inflorescence central, flowers sho\ v.
corolla tube funnel shaped, yello\v.
12 C. antadu Roxburgh (Fig 1221)
Commonly k n o ~ n as mango ginger referring to the characteristic taste of the
rhizome, large, sessile tubers present: root tubers absent. leaves four to six.
oblong-lanceolate, accuminate. iiif1orcscc.n~~ central or lateral: Sertile bracti
green; flon-ers yello\s.
13. C.aromatica Salish ( F i s 13a)
Large rhizomatous plant: up tu i 2111 high rh~zcirnc. cream!-\+!iiti. \ \ i r i i i n
aromatic. sessile tubers many. petiolc. Icing, lamina 4C~-70~10-14ci~i broadly
lanceolate. acuminate. inflorc~i.c.nce s j ~ke. lateral. coma hracts p~nl . i eni ! ~
hrac!s grceni\h-nhirc. cor ul l ~ 1di.i. l shaped. i i ~i ~ki \ h \{hi!<. l , ~ ~ ~ ~ l ! ~ i : ~ i
deep yc.l!o\r.
8
I4 C. cnesia Roxburgh ( F I ~ I4al
Perennial herb, rhizome strongly aromatic; leafy stem upto I m, high leaves
long \wth a conspicuous purple mid rib line; oblong-lanceolate, acuminate,
inflorescence lateral spike, coma bracts pink to violet, fertile bract green with
a pink t ~p. corolla tube pink, labellum deep yellow.
15. C: contoso Roxburgh (Fig. 15a)
Perenn~al herb. leafy stem 60 cm htgh. leaves, oblong lanceaolate four to five
In number. ~nflorcsccnce lateral, bracts wh~t e geen, floners yellou
16. C. decipiens Ualz. (Fig. 16a)
Perennial herb, rhizomc small. aromatic, pale yellow-white, leafy stem 20cm
high. lea\es 13, leaves broadly obovate,inflorescence central, bncts green
1~1t h pink tip, corolla tube deep purple, labellum yellow.
17 C. haritha Mangaly and Sabu ( F I ~ 173)
Perennial herb. rhizome yello\vish grey inside, leafy shoot upto 1 m hlgh:
1,-aves long lanceolate acuminate: inflorescence lateral spike, coma bracts
bnghl pink, firtile bracts green \\ith pink tip. flo~\t.rs white to light green,
\+ hite to light &Teen.
! 8 C: Iotrgu l Inn ( F I ~ I Xa)
r C: rlornesiica Valeton)
I t 1s the turmeric plant Perennial herb \vlth aromatic rhizome:leat:\ stem up to
icm high. leaves long, lanccaolate. ncurninate. 4-6 in number. lni1~ori.sceni.t.
c?ntral splke: bncts pale geen: flo\rers white to pale !ello\\.
19. C. malahnrica Velayudhan e~.irl. I Fig. I Oa)
Leafy stem up to 90 cm, long rhizome large,aromatic, leaves 4-5, long,
oblong-lanceolate, acuminate, inflorescence lateral; bracts pink: corolla tube
pink, labellum yellow.
20 C. rakracattra Mangaly and Sahu (Fig. 20aj
Perennial aromatic herb, rhizome greenish yellow inside, leafy shoot 40-60 cm
tall, pseudostem reddish purple, leaves 4-6 inllorescence lateral spike, bracls
pink, flowers, light pink to yello\v.
2 1. C zedoaria (Chnstmann) Roscoe (Fig. 2 l a)
Perennial herb, rhizome large \\ith smell of camphor, !ello\v to deep !ello\\
ins~de. leafi shoot up to l m tall, leaves long oblong-lanceolate, with purple
mid rib line; inflorescence lateral spike, bracts green with pink margin, flower
pale yellow in colour.
Herb, often robust plant, leafy pseudostem with fleshy rhizome, leaves lenlar
to oblong.lanceo1ate; flotvers in terminal spikes. flowers fragrant, pure white.
or t~nged \vith \ello\v.
Herb, rhizome thicker and harder than diploid plant. leafy shoot Irlrgc:. l e3\ ~9
leniar. oblong-lanceolale, crcam coloureJ flo\vcrs in globose s p~hz
I t 1s known as the cream ginger. Plant robusts up to 1.2 to 2m high. Lea\es
lanceolate,up to 4Ocm in length, pubescent, spike dense flowered, fragrant.
sulphur yellow in colour
25. H. spicnrum Hamilton ex Smith var. ocu
25a)
Leafy shoot up to 13m high. Rhizomes a
e r lanceolate, inflorescence terminal
fragrant.
26 Kaempferingalnngo Linn (Fig 26a)
Stcmless perennial aromatic herb; rootstock often tuberous, leaves 2-3.oblong.
acuminate, tlowers fragrant, white with a purple or l ~l ac spot on each side of
the lip.
27. Ei pulchrn K~dl (Fig. 27a)
Commonly known as the pretty resurrection lily: attractive tropical
rhizomatous herb with broad corrugated l ea~es which are flat to the ground. a
crey band i n peacock design over the bronze blade: flowers purple with broad
~ t ; l l s
Srernlrss herb: Iea\cs f eu oblong. acurnirlatr. \ar:gated grccn abo\e: i~nged
\\.:!h purple l>elo\v. t l onrrs In cro\vded radical sp1L.r.. tlo\vers t'ragmnt. \\h~:t..
lip purple or lilac.
29. ,4lpinia cnlcnrarn Roscoe (Fig. 29a)
Commonly kno\\n as 'Indian ginger', leafy stem 2 4 ti high, leares leniar-
lanceolate, finely acum~nate.intlorescence panicle, narrow. dense tlo\zercd:
flower white; lip varigated with rcd and yello\v.
70. '4. galango (Linn) S\vartz (Fig. 3Oal
Thick tuberous root stock, leafy stem 1.5-2m. high, leaves oblong. lanceolate,
acuminate: intlorescence open panicles; flowers greenish-white. lip wined
\v~th red or pink.
3 l . A. malaccerrsis (Buman) Roscoe (Fig. 3 1 a)
Leafy pseudostern 2-3m high; leaves oblong lanceolate, acuminate, the
mar~tjns of en densely ~i l l ous, Ilo\vers in simple racemes. tlowers tvhite, lip
yello\v, varigated with red, capsule 2.5 cm in diameter.
32. A. nigra (Gacrtn.) Bum (Fig. 32a)
Perennial aromatic herb Ivith thick root stock,leary stem 1.3-2n1 high I.ca\es
leniar or lanceolate-ohlon~: ~ntlorcsccnce panicle flowers small. p~nk; c;~piule
?cm in d~arnetcr
,?
2 . 4. purpurntn ( Ve~l i I K Schum hort ( r ~ g .?.;a\
Ornamental perennial herb \ \ ~ t h leaf\ stern ranpng from l-5m each endin; In
a sho\\y intlorescence. with a flower sprhe. brush-like. to 3icm long.
consisting of numerous large boat shapcd bright rcd bracts with small wh~t s
tloiver.
34 .4. smirhiar Sahu & Mangaly (Fig. 34a)
Leafy stem upto 1.5m high. leaves long margins villous, oblong-lanceolate.
inilorescence in spike with covering spathes. outer spathe shortly leafed,
quickly dscidous hracts and bracteoles: flowers tvhitc, red tipped.
. : S. ,4 .:erumber (Pers. ) Rurtt B Smith (Fig. 35a)
I t is known as the shell ginger; majestic rhizomatous plant. with long bladed
Iratheq leaves, the striking fragrant porcelain-textured flowers with bell
shaped waxy white calyx, the corolla white flashed with pink and lipped with
red and lip yellon in dense racemes to 30cm long.
36. A. ;erurnbet (Pers) Burtt B Smith var. Vrrrrgu!~ hort. (Fig. 36a)
The varigated shell ginger with broadly lanceolate, thin leathen; dark green
leaves varigated in feather design with stripes and bands of creamy yellow.
)7 Amomum cannicnrpum (Wight) Bentham ex Baker (Fig. 37a)
1-easy stern upto 3n1 high, leaves leniar lanceolate; acumrnatc. rprhe dense
flowered, peduncle short with bracts, margins of bracts of spikes villous.
flowers yello\v.
3 5 4 . I~~polrucurn l.h\vaites (Fig. 38a)
Leafy stem up to l m high l ca\ ci long-petiolsd. oblong-lanceolate. spiki: 1-7
tlo\~ered, in long underground runners pedurlcle rarely 2crn long. flo\rers
nhlte. lip with a !cllo\+ disc. tinged \rith red.
29 .A. muricatum Beddome (Fig 39a)
L.cafy stem upto lm high, lea\es small. elliptic or oblong-lanceolate.spike sub-
rrlubose and dense: peduncel 15cm long flowers white to yellowish. lip !ello\v
-
with a broad band of red spot, anthercrest yello\r.
10. .4. pterocarpum Thwaites (Fig. 4Oa)
Leafy stem upto 1.5 m high leaves broad, lanceolate, acuminate; ligule bifid.
membraneous; spike globose, floners white, capsule 9-ribbed.
11. Eletfuria cardamom Maton CV. Malabar (Fig. 3 la)
Leafy stem 6-10 A high, leaves lcniar lanceolate, acuminate, pan~cles several
up to 60cm long, arise d~rectl! from the pseudostem, prostrate; flowers \ +hi l t
lip longer white, str~pped 1~1th violet: capsulc subtrigonous, highly aromatic.
42. E. cardamomum Maton c\ Mysore (Fig. 32a)
Panicle erect.
43. E. cardanlutnum hlaton c\ Vezhi~ka (Fig J3al
Panlcle seml erect or prostrate
Table-l: List of members investigated
-- -. .-
p--- ~ -
SI.No .Name of tara
Locality of collection
2 ( ' .~pecro.\ IL\
3 Glohbu cernuu
4 (I. ~phroglv.v.~u
8. l. purpureum
9. Z rost,~rm
10. Z rerumbe/
l l . ( 'urcumu uerugmorrr
1 2 . (~'. arnutla
13. (I urornulrcu
1 1 (~'. cuesra
15. ( : L,omcr.ta
16, (~' . ~1~~crpren.v
17. '. l~uri~hcr
1 8. c'. l o n ~ u
19. ( '. n~ ~ l i ~ bc r r r ~ ( I
20 ( '. ~ U ~ I L I C U ~ I I ~ I
2 l. '. zcdourru
22. HC, L~I , C, / I I I I I ~I L,orotl~Irlilrrl
(Diploid c > ~ o t y e ~
Cochin
Kottaya~n
Calicut
Calicut
Wynad
Neryamangalam
Cochin
Vagamon
Nevaniangalam
Vaikom
Thrissur
'Fhrissur
Thrissur
Thrissur
Thrissur
Kuravilangadu
Thrissur
Vaikom
Thrissur
Thrissur
Thrissur
Cochin
---
Altitude in Xletcr
Sea lebel
so
J
45
1200
6s
Sea level
800
5 ;
Sca level
80
SO
s I!
SO
SO
h5
SO
Sea l ci cl
Si l
SO
, , ,
. , , 1
26 Kircnrpf;rru gulungu
27 E;. pulcl7r~r
18. K. rofundu
q -9. .4lprn1~r c~illc~rrnrtir
30. A. gulungu
Kuravi Iangadu
Cochin
Kothamangalarn
Thnssur
Thrissur
Kulathupuzha
Malakkapara
Cochin
Calicut
Cochin
Cochin
Sabarimala
Sabarimala
T'hekkadi
Wynad
Myladumpara
Myladumpara
Sea Irbel
8 5
65
65
200
350
Sea le\el
-15
Sea I s ~e l
Sea le\ el
800
800
900
1200
1050
1050
1050
Fig. l a - 16a : Habit of various taxa hnrestig-
l a - Costus rnalortieanus. 2a - Costus speciosus, 3a - Globba cemua, 4a - Globba o p M F ,
5a - ZingJber cemuum, Be - Znglber neesanum, 7a - Zlnglber offlcinaIe, Ba- Zingibsrpurpu~uun,
r
9a - Ungiber roseurn, 10a - Zin@bsrzennnbet, H a - Cunvrna aemgenosa, 12a - Gumme amp&,
?
13a - Curcuma aromatics, 14a - Curcuma caesia, 15a - Conwma c oma and 1 . - , Ba - Cuwma
:
decipiens.
Fig. 17a - 32a - Habit of various
17a - C u m a har i m 1 8a - Cumma long& 1 9a - Curcuma malabanca, 20a - Curcuma raktacanta,
2 1 a - Curcuma zedoaria, 22a - Hedychiurn coronarium diploid cytotope, 23a - Hedychlum coronarium
triploid cytotype, 24a - Hedyctw'urn flavescens, 25 a - Hedychiurn spicaturn var. acuminatum,
26a - Kaempfe* galanga, 27a - Kaempfena puIkchra, 28a - Kaempferia rotunda, 20a - Alpinia calcarata,
30a - Alpinia galanga, 31 a - Alpinia malaccensis, 32a - Alpinia nlgra.
I ; '
Fig. 33s43a: W of vuious tsaa iavestigabed.
33a- Alpiniu purpura(o, W- Alpinia &k&, 3%- Afpmh d&, 36a- Alpinia z m b e ~
var. varigda, 37% Amma n h y j ml wc ~ 3% A m- m y r m 40a- Amom~nn
peromrpum. 41a- E b i a c&immm W. MaZ,arl 42% Elettmio e a r & ~ ~ m CV-
Mysore, and 43a- EfdfmiQ m r b ~ CV. VazImlch
METHODS
a Mitotic squash experiments.
Somatic chromosome studies have been conducted on forty three taxa of
South Indian Zingiberaceae with the help of improved cytotechniques ( S h w a
and Sharma, 1980). Squash experiments were carried out on root tip meristem
at mitotic metaphase stage. Young root tips are collected from the rhizomes of
various species planted in the experimental botanical garden, at the periods
showing peak mitotic frequency, ie., 9.00 a m to 11 .OO a m. The root tips
collected are then washed thoroughly in distilled water and pretreated with
different pretreating chemicals. A 1:1 mixture of saturated aquous para-
dichlorobenzene solution and 0.00 1 M 8 hydroxyquinoline is found to be most
suitable for many members of Zingiberaceae. Eventhough a saturated aquous
paradic h1orobenzene solution with traces of aesculine is also used for some
genera like Hedyehim and Curcuma. The root tips immersed in cytostatic
chemicals are initially chilled at 0-SC for 4-7 minutes and then kept at 12-
l 6C for 1-4 hours for obtaining best results.
The pretreated root tips are then washed thoroughly with distilled water and
fixed in 1 :2 acetic acid-ethyl alcohol mixture (Carnov, 1886) for about 20
minutes to 1 hour, followed by 3-7 minutes treatment in 45% acetic acid, 5-10
minutes treatment in 1N HCl and stained with 2% aceto-orcien. While
staining the vial is initially warmed over a flame for getting good results. The
stained root tips are then squashed in 45% acetic acid and the preparation was
temporarily sealed. To remove the excess oil deposits, rather common in
many members of Zingtberaceae, it has been found preferably to add a little
sapunin along with the pretreatment chemicals and shake thoroughly.
b. Detailed Knryomorphological studies by image analysis system
Photographs and image processing
Well spread metaphase plates were photographed using 125 ASA 35mm Orwo
film in a Leitz photographic attachment and suitably enlarged. A 200 DPI
scanner scans each orignal photograph. The software Adobe Photoshop was
used for digitizing and Photostyler for reproduction. The contrast of each
image has been increased by raising the resolution up to 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 images were checked by the visual inspection
cornpanng with the original photornicrographs. After storage of original
digital images of the metaphase spreads in floppy discs, these images were
analyzed by using computer devices. The image was recovered from the
floppy disc and the original digtal image for the analysis was then generated
automatically. A binary image 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 grey values of the pixels that consisted of the chromosome imags were
included. Bimrization was automatically canied out by changng the pixels
whose grey values stood between the two thresholds to white and all others to
black (Fig.22 b2). The large background dust particles and other spots whose
grey values also fell between the two thresholds, were eliminated by adjusting
the g e y L-alues of the pixels outside the chromosomal regons by interactive
setting of lower and upper thresholds of the grey levels. This will result in a
binary chromosomal image with a clean background (Fig. 22 M).
Quantitative karyotyping of the chromosomes
Measurement of each chromosome from enhanced image (Fig. 22 b6 ) were
made by AutoCAD (version 14) loaded on a personal computer. This image
was automatically coloured differently (Fig. 22 b7 & b8) by the computer-
generated colouration based on the actual grey 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 cons~ctions of the chromosomes. These constrictions were
manually marked by the overlay straight lines on the pseudocoloured images
(Fig. 22 b6 gi b8). After this rnidrib lines were drawn interactively on each
chromosome. Extraction of the midrib lines, breakage at the position of the
constrictions and identification by different colours were subsequently carried
out automatically (Fig. 22 b9). The outer margn of each chromosome image
also marked by drawing surrounding line (Fig. 22 b7). Numerical data
such as the arm length, area, perimeter and an apparent visual three-
dimensional volume of each chromosome 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 al. (1983). On the basis of F% the nature of primary constriction
in the chromosomes are classified as follows:
Nature of primam constriction
Median
Nearly median
Nearly sub median
sud median
Nearly sub median
Nearlv sub tern t nal
Subterminal
Nearly sub terminal
Extreme1 y sub terminal
Terminal
The disparity index ( Dl ) of chromosomes in a karyotype is calculated after
Mohanty et al. (1991 ), by the formula:
Longest chromosome - Shortest chromosome
= -----------a------------- - ---------------m------ ------
X 100
Longest chromosome + Shortest chromosome
The variation coefficient among chromosome complements (VC) is
determined after Verma ( 1 980) as follows:
Standard deviation
VC = ------m---- ---- ---------------v- X 100
Mean lengths of chromosome
The total foma percentage or the mean centromeric index value (TF?/o) is
calculated in each taxa after Huzi wara ( 1 962), by the formula:
Total sum of short arm length
TF% = ----------- ----
X 100
Total sum of chromosome length
The arm ratio and kiformity coefficient (perimeteriarea) (Ojeda and Tomes,
1996) were also calculated. For the calculation of polyploidy, the base
number given in Darlington and Wylie (1955), Morton (1956; 1962), as well
as Love and Love ( 196 1 ), are usually followed. In those cases where the basic
num ben are not mentioned in these references, the latest literature is
consulted. All the numerical data are prepared after comparing at least five
well spread metaphase plates. The various calculations were done by the
computer package Microsot? Excel and the data used to classify the
chromosomes according to Levan et al. (1964).
Quantitative idiograms of chromosomes
Based on the data relating to the length, the idiograms were presented
combined with the resul t,s of quantitative image analysis of chromosomes.
The chromosomes were amged semi-automatically according to the length,
arm ratio, uniformity coefficient, virtual three-dimensional volume and
idiograms generated with the aid of computer software Adobe Photostyler.
22 b7 22 b8 m c:,.
' . I ,
,.-. . , . .
Fig. 22 bl-b9: Different steps of of -m d (2x1
by the image anahrsis
b 1 m - b plate, b2-OIL, b3-backgro~d b4- - h
plate with c k b d g md , b5-stlection of a ohnrmoMrme for meerrurement,
b6-mhancedimsgewith~inthecomtriction, b7Qawing marginal
OBSERVATIONS
In the present investigation forty three taxa of South Indian Zingiberaceae
including two cytotypes in Costus, two in Globha, six in Zingiber, eleven in
Curcurnu, four in Hedychiurn, three in Kaempferia, eight in Alpinia, four in
,4rnomurn and three in Eletturia were analysed. The normal somatic chromosome
number ranges from 2n=18 to 3n==63. However, numerical variations are
prevalent in some taxa. The ploidy level exhibited by different taxa ranges from
diploidy to pentaploidy. The chromosome pairs with secondary constriction are
found to range from one to seven. The karyotypes are characterized by
comparatively small chromosomes ranging from 2.81 pm to 0.3pm. The
chromosome in each karyotype decrease in size progressively and bear nearly
median to nearly sub median primary constrict ions.
The general description of the common chromosome types is gven below,
fo l lowed by a karyotype description of the members investigated.
Type A: Comparatively ldng chromosomes with two constrictions.
Type B: Relatively long chromosomes with nearly median to nearly sub-
median primary constrictions.
Type C:
Very short chromosomes with nearly median to nearly sub-
median primary c-onstrictjons.
Normal somatic chromomme number
Karyotype h u l a
Number of cfuorncwome paLs vrith secondary cmsbkth
Range of &ornosome length in pm
Totd ckmmsme b g h m pm
Arerage chromosome lengltr in pm
T.F.vekre (S)
Variation caefiiuent (V.C)
Dsparity nde x (DJ)
Total voCwneof chromosomes
in pm'
: 18 (Fg. lb)
: A2B16
1 -00
: 1.840.57
18.57
0.92
35.93
39.26
%.M
CIrmtlsome T d le@ Short aim Armratio FOh Vdurne PIA Nature of
TYW in pm length in pm (r) in pm' Primary
A 1-84 0.52 1.63 28 -29 0.076 9.71 Nearly Submedian
0.47 25.54
B 1.05 0.33 2.18 31 -45 0.023 10.25 Nearly Submedian
B 0.90 0.36 1.53 39.40 0.022 12.68 Nearty Median
B 0.76 0.25 2 .Q9 32.26 0.019 13.51 Nearty Submedian
B 0.68 0 -26 1.62 38.28 0.014 12.14 Nearty Median
B 0.66 0.30 1.21 45.19 0.012 14.47 Nearly M d a n
Normal somatic chromosome number
Somak variation number
Karyotype formula
Number of chromosome p a k with secondary ccmdMmn
Range of ehromosorne h g h in Vr n
Total chromosome kngth in pm
Average &ornosome length in pm
T.F.value (%)
Variabon ooeficient D/.C)
index (D.I)
Total vo4me of chrwnoums
in pm=
: 36 (Fig. 2b)
: 24 (Fig. 2c)
: A4 B30 C2
2 .W
: 1.814.50
32.62
0.91
39.39
3527
603
Chromosome Total length Short arm Ann ratio F% Volume PIA Nature of
TW in pm lengthin pm (r) m VITP
Primary
. .
cmhwbon
A 1.81 0.57 1.30 36.83 0.092 1 0.00 Nearly Submedian
0.51 27.62
A 1.07 0.37 ' 1.10 44.84 0.032 10.51 Nearly Median
029 27.1 1
B 1.35 0..45 2.01 37.21 0 .U36 1 2.94 Nearty Submedian
B 1.07 0.38 1.84 43.74 0.038 1 1 -37 Neady Medan
B 0.96 0.39 1.48 31 -90 0 .M8 10.25 Neady Submedian
B 0.88 0.41 1.17 39 -42 0.022 1 0.54 Nearty Median
8 0.80 0.33 1 -42 43.42 0.038 13.99 Nearty Median
B 0.78 0.28 1.77 32.46 0.020 9.42 Nearly Submedian
B 0.74 0.32 1.34 34.60 0.027 12.74 Nearly Submedin
B 0.72 0.32 1 -22 42.08 0.01 9 12.33 Nearty Medran
B 0.70 0.30 1.31 48.4 1 0.020 1 0.60 Nearly Median
8 0.66 0.25 l .64 49.52 0.015 1 2.40 Neady Median
8 0.54 0.24 1.24 44.39 0.01 7 12.34 Nearly Median
C 0.50 0.21 1.32 39.34 0.01 1 1 1 . l 0 Nearly Median
Normal somatic chromosome number
Karyotype formula
Number of chromosome pais wim secondary cmsbkkn
Range of chromosome ktgth in pm
Total duornosurne kngth in pm
Average Ctuomwxne kngh m pm
T.F.vahie (%)
Variatian coefficient (V.C)
Dq a n t y index (0.1)
Total volume of chromosomes
in vm3
: 48 (Fg 3b)
: A6 842
3
: 1.8-0.65
53.3
l .l 1
42.2
20.08
41.44
Chromosome Total length Short m Arm ratio F% Vokrrne PIA Nahre of
T y ~ e m pm length h pm (r) in pm=
Primary
consbidion
Neady Submedian
Neady Submedan
Neariy Subrnedan
Neady Median
Nearly Median
Neady Median
Nearly Median
Nearly Submedian
B 1 .l6 0.52 1 22 45.07 0 .M7 7.37 Nearty Medisn
B 1 .l2 0.53 1.10 47% 0.056 7.41 Nearty Median
Nearly Median
Neariy Median
Nearly Mean
Nearly Median
Nearly Medran
Neariy Mechan
Nearty Median
Nearty Median
B 0.84 0.39 1 .l4 46.7 1 0.049 9.00 Nearty Median
B 0.83 0.38 1.16 46.35 0.027 9.01 Nea* Median
8 0.79 0.35 1.24 44.67 0.066 8.61 Nearly Median
B 0.75 0.36 1.10 47.71 0.027 8.90 Nearly Median
8 0.68 0.33 1 .M 48.59 0.029 8 2 5 Neaity Me hn
8 0.65 0.30 1.14 46.72 0.027 8.35 Nearfy Medan
Normal somatic chromosome wrnber
Karyotype fonnula
Number of cfrarnosorne pairs with secondary mr&i&m
Range of &ornosome length in pm
Taaal chromosome length in pm
Averege &ornosome length in pm
T.F.M)ue (96)
Variation c s e h n t ( VC)
bspanty i ndex (D.[)
Totat v h of chromosomes
n V~IP
Ctumasome Totallength Shortwm Arm ratio F% Volume PIA Nabre of
T y ~ e in pm hgthingrrn (r)
in pmS Primary . .
cmdm3on
3
A 1.73 0.55 1.29 31.78 0.101 8.1 3 Neady Submethan
0.47 27.17
A 1.55 0.53 1 . l6 34 -63 0,060 7.60 Nearly Submedian
0.39 25.16
B 1 . l 6 0.51 1 28 43.87 0.082 7.38 Nearly Median
B 0.99 0.41 1.39 41 -34 0.029 9.81 Nearty Median
B 0.83 0.38 1 .20 45.54 0.028 9.63 Nearty Median
B 0.79 0.37 1.13 47.01 0.032 9.90 Nearly Median
B 0.76 0.35 1 .2Q 45.37 0.021 9.87 Neady Median
B 0.73 0.33 1.20 45.48 0.026 9 . Nearty Median
B 0.67 0.30 1.22 45 .M 0.022 10.42 Nearty Median
B 0.61 0.30 1.05 48.85 0.031 9.86 Nearty Median
8 0.56 024 1.38 41.97 0.014 10.67 Neady Meckan
Normal swnatic ctromosome nwnber
Karptypefamwla
Number of charnosome pairs with secondary mnstric
Rgnge of chromosome length in pm
TtW drromasme length in pm
A m g e dromosome'length in pm
T.F.va3ue (%)
variation ooefficierd (V.C)
Chpdy index (DJ)
Total Wr ne of chromosomes
in p&
: 22 (Fig 5b)
: A6 016
3
: 3.27-0.86
34.55
1 .S7
38.9
ctmmwmm Tdal )eneth Short arm Arm ratio F% Volume PIA Nature of
TW inpm hgt hi npm (t) in pn?
Primary
Nearty Submedian
Wady Median
Nearly Median
Nearty Me&n
Nearly Median
Nearly Median
Nearly Median
Neatly Medan
Nearly MeQan
Normal somatic chromosome n u mk
Karyotype formula
Number of chromosome pairs wiU-i secondary comtmbon
Range of chromosome length in pm
Total chromosome lenglh in Vr n
Average &tmwsome"length in pm
T.F.va& (96)
Variation coefficient (V.C)
bsparity index (DJ)
m pm3
Chmos~ne T~~ Shortarm h r a t i o F% Vokrrne PIA Nahre of
T y ~ e inpm lengthinpm (r) h pm3 Primary
A 2.19 0.79 127 36.03 0.323 6.50 Nearby Submedian
0.41 18.26
A 1.95 0.63 125 32.35 0.142 6.27 &arty Subme&
0.53 27.18
8 2.1 7 0.99 1.19 45.78 0.120 6.30 Nearty Median
B 2.03 0.55 2.67 27.26 0.141 5.99 Nearly Submedm
B 1 -88 0.80 1.37 42.1 7 0.151 627 NeariyMedan
B 1 -72 0.80 1.16 46.37 0.140 6.01 Nearly MeQan
B 1.60 0.65 1.46 40.57 0.1 84 6.48 Neatly Medan
8 1.45 0.49 1.96 33.80 0.173 5.85 Neatly Submedian
B 1.30 0.55 1.36 42.47 0.129 6.50 Nearly Median
B 1.13 0.35 2.19 31.21 0.044 7.41 Nearly Sdmedian
N m l somatic chromosome number
Number of chromosome pairs with secondary cansbrction
Range of chrornosarne length in pm
Total &ornosome h # h in pm
A = q p c h m = m e ~ i n v r n
T.F.& (%)
veriation coefhcient (V-C)
Dispway index @.I)
: 22 (Fig. 7b)
: A6 B16
3.00
: 1.49-0.81
24.66
1.12
39.07
16.38
32.00
Ctvomaslbme Totalkngb Shortarm Annratio F% Volume PIA Nature of
T y ~ e b ~ ~ n (0 in clm3 Primary
n
Nearty Sobmedian
NearlJ Submedian
Nearly Subrnedsan
Neatly Medin
Neady Median
Nearly Median
Nearty Median
Nearty Median
Neatly Median
Nearly Median
Nearty Median
Normal somak chromosome number
Karyotype formula
Number of chromosome pairs with secondary cortstrkbon
1 OM chromosome le@ in urn
Awage ctrromasome length h pm
T.F.value (Oh)
Variation coefficient (VC)
mar r t y index (D. I)
in pma
T-9: Detailed kawtype analysts of Zbgfbw prarpurun
Cbomosome Total length Short arm Arm ralk F% Vdume PIA Nabre of
Type in vrn hgih in pm (r) W Primary
Nearly Submedtan
Neady Submedian
Nearty Submedian
Neady Median
Nearty Subrnedian
Nearly Subrnedian
N e w Median
Nearly Median
Neafty Median
Nearty Subrnedian
Nearly Median
Nmal wmdc &cmcsorne number
Karyotype f md a
Number of ctvomosome pairs with secondary
R- of charnosome Length in urn
Tdchr-Lenglhirw
Ammpohromosomelengthinym
t.F.value (%)
v* c mdk k d (VC)
mqwly indcr (DJ)
Total v o h w of chromaomes
h pm=
Tabkl 0: Detailed kaqotype a n a m of Zdnglbcr roseun
Chomob;omc Totallength Shoctarm Armratio F% Vokrme PIA Nature of
Type in vrn l - in pm (r)
h pm=
Primary
n
Nearty Submedian
Nearly Submdan
Neatly Submedian
Nea* Submedian
Neatty Medan
Nearty Median
Nearty Median
Nearly Medan
Nearty Medm
Nearly Median
Normal somatic duomosome number
Karyotype formula
Number of chromosome pairs wi h secondary consbiction
Total chrommme lengih in ym
Average chromosorne"kngh in lrn
T.F.vakre (%)
Variation coefficient (VC)
Dianty index (D.1)
Total vdume of chromosomes
m pm3
: P (Fig lob)
: A4 818
2.00
: 1 AM. 7
24.99
1.14
38.16
24.1 1
39.10
Table-l 1 : Detailed karyotype analysts of Zingiber zenrmbet
Chromosome Total h-@h S h d a m A m ratio F% Volume PIA Nature of
Type in pm length in p h (r) in pm'
Primary
mmon
A 1,40 0.44 * 1.23 31.28 0.165 8.40 Neady Submedian
0.43 3021
A 1.29 0.43 1 . l 9 32.97 0.079 8.51 Nearfy Submedian
0.42 32.56
B 1.49 0.61 1.45 41.06 0.132 7.52 Mearty Medran
B 1.38 0.47 1.93 33.85 0.122 7.78 Nearly Submedian
B 1 -25 0.47 1 .M 37.95 0.066 7.66 Nearty Median
B 1.11 0.52 1.15 46.53 0.069 8.12 NeartyMedian
B 0.92 0.35 1.62 38.38 0.050 8.1 5 Neady Median
B 0.83 0.38 1 . l 9 45.72 0.024 10.79 Neaiiy Median
B 0.77 0.33 1.35 42.53 0.031 9.37 Nearty Median
B 0.70 0.32 1.15 46 -44 0.040 10.75 Near?, Median
Normal somatic chromosome n u mh
Katyolype formula
Number of &ornosome p a h wim seconday mmstidion
Range of chromosome length m pm
Total chomosome lmgm in pm
-Average &ornosome kngm h pm
T.F.value (%)
Variation coefiicient (VC)
Disparity index (D.!)
Total vdurne of chromosomes
in pm3
: 53 (Fig. l l b)
: A6 B22 C35
3.00
: 0.76-030
31.70
0.50
43.20
19.22
43.90
Chromosome Total length ShM arm Arm ratio F% Volume PIA Nahire of
Type in pm length m vm (r)
in p d
Primary
Nearly Subrnedan
Nearly Submedan
Nearly Submedan
Nearly Median
Nearly Median
Nearly Median
Nearty Median
Nearly Me&n
Nearly Medan
Nearly Medan
Nearly Me&n
Nearly Median
Nearly Me&an
Nearly Median
Nearly Median
Nearly Median
Nearly Median
Nearly Median
Nearly Median
Nearly Medan
Nearty Mr mn
Nearly Mehan
C 0 45 0 20 1.25 44 49 0 008 14 23 Nearty Medan
C 0 44 0.20 123 44 93 0.008 14 65 Nearly Medan
C 0.44 0.20 1.24 44.61 0 009 13 67 Nearly Medan
C 0 43 0 21 1.09 47 88 0008 14.33 Nearly Median
C 0.43 0.20 1.17 6 . 1 8 0.009 13 07 Nearly Medan
C 0.42 0.19 1.20 45.41 0.009 13.30 Nearly Medan
C 0 40 0 2 0 1.05 48.72 0 009 13 70 Nearly Medan
C 0.39 0.17 1.30 43.52 0 009 14.06 Neady Medan
C 0.35 0.17 1.08 47.99 0.009 14.15 NeaftyMedian
C' 0.03 0.14 1.15 46.50 0.005 17.69 Nearly Medan
Not In palr
Normal urmabc chromosome rmmber
Karyotype formula
Number of chomosome p a n secondary comtncbon
Range of chromosome ieng24 m pm
Total chromosome kngm m pm
Average chromosome length m pm
T F value (%)
Vananon coefficient (V C)
Dspanty ~ndex (D I)
Total votume of chromosomes
In pm'
Table-1 3: Detailed karptype a na w of Cucuma amada
Chromosome Total length Short arm Arm atio F% Volume PIA Nabre of
Type In vm length h vm (r) h vm' Rimay
A 1.03 0.33 l .P 32.39
0.29 28.16
B 0.86 0.40 1.14 46.66
B 0.78 0.36 1.17 46.11
B 0.75 0.35 1.14 46.65
B 0.73 03C 1 .36 42.40
B 0.73 0.33 1.21 45.31
B 0.73 0.31 1.32 43.13
B 0.72 0.28 l .S 38.99
B 0.71 0.28 1.53 39.57
B 0 70 034 1.09 47.91
B 0.69 0.31 124 U.70
B 0.68 0.28 l .U 10.90
B 0.66 0.33 1.03 49.29
B 0.66 0.29 1.25 UAO
B 0.63 028 1.26 4. 24
B 0.62 0.29 1.13 47.01
B 0.60 0.28 1.14 6.68
B 0.59 0.24 1 .M 40.60
B 0.58 0.27 1.11 47.41
B 0.56 0.23 l .45 a.87
C 0.51 0.24 1.14 46.63
early w n
Nearly Medan
Nearly M&n
Nearly M a n
Nearly MMedian
Nearly Me6an
Nearly Meda
Nearly Meda
Nearly Median
Nearly Medan
Nearly Median
Ne* Me&n
Neady Mcdrn
Nearly Medan
Nearly Medan
Nearly M&n
Nearly Median
Nearly Median
Nearly Median
Nearly Median
Nearly Median
Normal somatic chomosome rumber
SomslicvaislmMnber
K~ryotrpe lmnlh
Number cdrtmmmune psir wih secondary consbic(m
~ o f d w m m m e * n g h h p m
T o l d c t m m o s a n s ~ i n ~
Av a np drm- m h vm
T.F.va(ue (X)
vaialm cnetwud (V.C)
DapariCV indu (DJ)
Totalwolnrdchamsomes
h
Chmmoum Total hglh Short m Arm Rta F% Vohme PIA N W e of
TYPC mpm b g h n p m (r) h Vw Rimnl
Nearly Median
Nearly Me a n
Ne* Medal l
NcarlyM*
NeaIty Median
Neaiiy M e n
Nea* Mcba,
M-
Nonnal mmalic dr omaane mb e r
K - v o m -
Number of chomomne pah wih semndary congicbbn
R a w of dmmmome lengh in pm
Totaldmnw*omelenghinpm
Average ehmnsome kr& in pm
T.F.value (X)
Var i ah m W.C)
Dispa* klex (DJ)
Total voLmc af chmmosomes
in pW
Tabk-IS: L l e t e W ~ ~ o f C u c v n r a e s h
Chromosome Total length Shat ann Arm ratb F% Vdume PIA Nabb-e of
TW m pm lengthinpm (r) in pm' Rimary
A 0.90 0.29 1.27 32.56 0.035 12.08 Nearly Shrnedan
023 25.67
A 0.84 024 1 .66 227 0.033 11.01 Nearly Submedan
0 2 . 26.28
B 0.62 028 1 .20 4528 0.01 1 12.14 NeaftyMedan
B 0.59 0.26 1.32 43.00 0.010 12.87 Neafiy Median
B 0.57 027 % 1.14 46.64 0.011 13.88 Nearly Median
B 0.57 023 1.51 39.80 0.012 13.23 Nearly Median
B 0.56 024 1.33 42.89 0.015 13.79 Neady Median
B 0.55 0.22 l .S 39.31 0.009 13.10 Neatly Median
B 0.54 026 1.07 4823 0.008 13.05 Neatly Medan
B 0.53 023 1.36 42.36 0.010 13-00 Neatly M m
B 0.53 0 26 1.07 4.3.34 0.015 13.05 Nearty Median
B 0.52 0.25 1.11 47.32 0.016 13.06 Nearty Median
B 0.52 0.25 1.03 49.28 0.012 12.95 Nearty Median
B 0.51 024 1.12 47.13 0.005 13.41 Nearty Median
B 0.51 0.24 1.07 48.33 0.009 14.17 Nearly Median
B 0.50 0.23 1.14 46.76 0010 13.53 Nearty Median
C 0.49 021 1 .M 43.58 0.010 13.38 Nearly Median
C 0.49 0.21 1.35 42.49 0.009 13.65 Neatly Median
C 0.49 022 1.21 45.29 0.006 13.80 Neatly Median
C 0.48 0.23 1.07 48.21 0.012 11.86 Nearty Median
C 0.48 0.22 1.22 45.05 0.009 13.36 Nearty Median
C 0.47 0.21 1.27 U.10 0.012 12.60 Nearly Medan
C 0.46 0.22 1 .J9 47.88 0.014 13.58 Ne d y Mcdan
C 0.45 0.21 1 .W) 48.10 0.015 1325 N e a ( y Wn
C 0.44 0.19 1.36 42.45 0.010 13.30 Nearly Medan
C 0.44 0.19 126 . 44.27 0.013 16.01 N e a r l y W
C 0.42 0.20 1 .W 48.49 0.W7 13.69 Nmiy McdM
C 0.40 0.16 l -60 =.U 0.011 15.83 Nearly W n
C 0.40 0.17 1 .M 43.51 0 . W 16.33 Nearly Mcdsn
C 0.38 0.16 l . U a.93 0.009 13.94 Nearly Medsn
C 0.36 0.17 1.15 46.57 0 . W 14.57 Nearly Medan
C. 0.34 0.15 1.36 42.33 0.007 1923 Nmiy Medan
'Ndhpai
Namd m& dromomine numbec
KarVotVpsfamh
NMbaof chanaromepdrrwn, ssmdsl y~
. .
Rrpcof-winirm
Tobl dranoromel enspl n~
-c-hmnIrrn
T.F.n*c (X)
vaiacan coeRdsn (WC)
IklMm idcr @.I)
Tobl of chromosMmr
n pm'
42 (Fig 1%)
A2 B6 C34
1
0.774.31
1826
0.43
43.02
24.53
44.4
Tdc-16: D c W Lacyo(ype a na m of Cue- mmaea
Ummo s o me T o t a i ~ Shatann Arm& F?+ Vobme PIA N-e of
T w hvm hg~ hnpm (0 n pm' R*nay
A 0.77 o a 1 .m 33.70 0.023 12-67 kaly m
028 36.36
B 0.56 0 . a . 1.82 3550 0.- 13.93 Neer)y s h n d W l
B 0.58 0.23 1.49 40.14 0.005 14.65 Nedy Uedsn
B 0.53 024 1.23 44.86 0.007 13.59 W-
C 0.50 0.20 1.47 40.46 0 . m 14.72 Naa(yMedan
C 0.48 024 1.03 49.33 0.029 8-72
C 0.46 020 1 . U 42.84 0.004 15.96 Na a r l y W
C 0.45 021 1.19 45.66 0.007 14.98 NewMcan
C 0.44 0.11) 1.42 41.43 0 . W 14.80 Nea*Medan
C 0.42 021 1 .M 49.01 0 . w 15.25 WMe d a n
C 0.41 0.17 1 .36 42.45 0.004 17.11 kalyMcda
C 0.39 0.19 1.10 47.71 0 . W 15.97 NeartyMedan
C 0.39 0.19 1.01 49.83 0.003 17.22 Neatly Me&
C 0.38 0.17 1.27 44.04 0.005 15.06 Nearly M&n
C 0.37 0.13 1.88 34.76 0.004 17.43 Nearty Submedian
C 0.36 0.16 1.23 . U. 79 0.005 16.50 Nearly Medan
C 0.35 0.15 1.27 U.02 0.004 16.76 Newly M*
C 0.33 0.17 1.02 49.58 0.001 16.62 Neatly M&
C 0.33 0.13 1.49 NZ? 0.005 16.m W M e d a ,
C 0.32 0.16 1.03 49.29 0.006 15.38 Neatly Me6an
C 0.31 0.15 l .06 48.53 0.005 16.40 Nearly Mdan
Normal somatic chrornomme number
Ksryo(ype-
Mmb a d c h wn a o mc p i s wi h s e mMa r y ~
. .
R a w of drwnaurm+ h pm
TDtst chwmrPane lerglh m pm
A ~ d m m o s o m e ~ h p m
T.F.vahm W)
Va l i a h C O e ~ (VC)
M @.I)
Total d chomaansr
m pm'
chmmsom Totalkrgh Shortarm ArmraSo F% Vdum, PIA Nabre of
Tvpe inw k a h i n ~ m (r) m pm' Rinry
11.85 NeaItyMdm
11.89 NealiyMeba,
11.s ~~
11.93 N W M s d a
1l.W NmrtyMadan
12.87 Nearly-
1223 Nsaly-
11.19 Near(lMe6n
12.05 ~~
12.90 Nea(lMa.$l
12.94 NemMeaan
13.79 Nearlyhkhn
13.07 Nearly Medsn
13.78 Nemty
1339 NemtyMadan
14.88 NeaftjMe6-m
13.12 Neady Lkdan
13.80 N e w Madan
14.02 Nea)l-
14.52 Nea* Medan
ChomommsTotalsnglh Shortm h& F% VoCnre PIA NaWe of
T w hlwn -=)m (0 hlrm' Rvnay
OIL
0.17
(U1 '
0.16
022
0.17
020
0.19
0.16
0.17
0.17
0.18
0.17
0.15
0.15
0.l4
0.14
0.14
0.14
0.1 1
0.13
0.11
N W-
Nearly -
NearlV M-
NW-
NewW&n
N e w Medsn
NO* M*
Ne&yMedan
N e w Medan
Nearly Medan
N e w Medan
Nealy M e n
Newly Medan
Nemly Medan
Nestty Medan
Nearly ?&me&an
Ne- M&
Nea* M a n
N m n a l s w M C c ~ R m b e r
Somalicvai.(knnniba
KarVotVp-
Numkr of duomosome p& wih semndary mns8idkn
Rngsofdmnmomeknglhinprn
Totaldnwnmmmknghh)~
Average dnmxme *ngh h pm
T.F.n*a W)
valialbn me- (VC)
h p d y i a x (D.1)
Total &me of dromommes
:=(Fig. 18b)
: 42 (Fig. l&)
:AZ8MC41
1 .W
: 1.014.40
31 .97
0.51
44.00
20.68
45.00
V d u ~ PIA Name of
inw m
W M -
W M *
Nearty Median
Nearly Medan
'Not h pai
: 42 (Fig 19b)
:A2B12C28
1
: 0.994.35
21.83
0.52
43.39
322
43.39
Chamrswnc Total- W a r m Arm& F% V o h e PIA Nahre of
Type i rvm len@in)rm (r) h pm' Primary
c'mmcml
0.30
0.28
0.40
0.35
027
0.22
024
0.22
0.22
o z .
021
0.23
022
0.20
0.16
O M
0.19
0.29
0.17
0.18
0.17
0.14
W-
Nsafty Msdn
Nemty Msdn
W-
Nemty Ucdrr
W-
W-
W-
M*-
W M h
Ne w ShmUm
Ne* W n
N W M&
Nenty M&
Nea)y Mel%an
Nearly M&
Nearly M&
Nearly M h
Nmnslsmu(lcdmm0Samerunber
So ma ( i c Va a 0 n ~
KarptVpe-
Nmba of c hr omomv pa i . ~ s e oonda y ~ 6 on
RnosO'chmx*omaWklm
Touchr ommnbr dhkp
A v a a o c d r v m m ~
1 . F . h (X)
varisriar ccefu&d (VC)
Dispsrily*@.o
To(dvdmedchanosomes
m pm=
Chomosome Told *npm Short srm Arm ratb F% VoLma PIA Name of
Tvpe mw m m v m (0 h W
Nearly h l n
W M*
Nearly Medan
Nearly Medan
Nearly Mcdan
Nemty Me6an
NrarlVM&
W-
Nearly M*
Nearly M&n
Nesrly Medan
Nearly M&
Nearly Median
Ncarly Me&
Nearly Median
Nearly Median
Nearly Median
Nearly Median
Neariy Median
T.blc22: D e t s b d i r a y o ( y p e ~ 0 l C M n w ~
ChanavuneTMm Shmtam, Armrdo W Volme PIA Nahre of
Tvpe in *npn h rtm (0 Rimary
A 0.68 020 l .52 30.50 0.021 12.50 Nealy ShwaSan
0.16 2424
A 0.62 020. 1.16 3236 0.01 l 1221 WSu b me d i e n
0.17 27.42
B 0.55 026 1.10 47.54 0.015 12.35 Newty Meda
B 0.52 020 , 1.54 39.42 0.010 12.73 ~early~nfan
C 0.49 on 1 .S 42.07 0.009 12.31 ~ealy M&
C 0.47 022 1.14 46.68 0.011 13.49 N e w M-
C 0.46 021 1.17 46.01 0.007 13.51 Medan
C 0.48 0.23 1 .02 49.40 0.009 12.53 Neady Medan
C 0.45 0.18 1 M 41.03 0.008 13.W NealiyM-
C 0.45 021 1.11 47.38 0 . m 14.78 Nealy Meda
C 0.44 0.21 1.14 46.79 0.010 12.94 N a t y Medan
C 0.44 0.18 1 .48 40.30 0.009 13.28 Nealy Me&
C 0.43 0.17 1.52 39.68 0.WB 1 Nealy MeQn
C 0.43 021 1 .W 48.00 0.009 12.48 Medsn
C 0.43 0.20 1.18 45.83 0.009 12.43 Nealy Medan
C 0.43 0.19 1 26 44.19 0.007 14.23 NemMedan
C 0.42 0.19 128 44.32 0.008 13.41 NcaRy Medan
C 0.42 0.19 1.18 45.84 0.008 13.W Nearly Medan
C 0.42 O M 1 .ffi 48.49 0.006 13.98 Nearly Me&n
C 0.41 0.19 1.12 47.10 0.007 13.88 Nea* Medan
Contd.
C 0.40 0.17 1.31 4320 0.008 14.40 Nearly Medan
C 0.39 0.18 1.23 44.87 0 . m 14.41 Nesrly Medan
C 0.39 0.18 1.19 45.68 0.009 1322 Nearfy Medan
C 0 . 3 0.17 120 45.43 0.007 13.76 Neaty Me&
C 0 . s 0.16 1.30 43.52 0.OW I3.W Neady-
C 0.37 0.16 IZ 44.01 0.010 13.78 early ~ e d n
C 0.37 0.15 1 . 9 39.95 0.009 13.71 Nealy Me&m
C 0.37 0.17 1.13 47.W 0.006 14.04 Nf ady Medan
C 0.35 0.17 l .S 48.74 0.012 13.47 Naar)y Meban
C 0.34 0.15 l .m 43.49 0.010 14.67 NeahlMedan
C 0.33 0.15 122 45.12 0.005 15.58 NemMedan
C 0.31 0.1 1 l .77 S.08 0.- 1624 Ne~earvSUhne&n'
'Notinpai
L
4
--
l l b - Curcirmn aerugenosa (2n = 63), 12b - Curcuma amada (2n = 42),
13b - Ctrrcuma aronlatica (2n = 63), 14b - Curcuma caesia (2n = 63),
'T,b . Ci t rri ~ni a cornosa (2n = 42), 16b - Curclrma decipiens ( 2n = 42).
1 ' l .
' - I I ! ~ rrnla hartltln (2n = 42). 18b - Curcumn longa (2n = 63).
t ~ r r ! a r ~i , i l . , h, ~r ~ca ( 2n = 20h - Curcirnia rcktacanta 12n = 63;
r1rn.t / ~ - : l n . ~ r t a (2n = 63.
Fig. Id to 8d-Comparative idiogram plates of various taxa studied:
Id- Costus malortieanus (2n=l8) 2d-Costus speciosus (2n=36)
3d-Globba cernua (2n=48) 4d-Globba ophioglossa (2n=22)
5d-Zingiber cernuum (2n=22) 6d-Zingiber neesanum (2n=22)
7d-Zingiber officinale ( 2 ~ 2 2 ) 8d- Zingiber purpureum (2n=22)
Fig.9d to 16d-Comparative idiogram plates o f various taxa studied:
9d- Zingiber roseum (2n=22) l Od-Zingiber zerumbet (2n=22)
. -
l l d-Curcuma aerugehosa (2n=63) 12d-Curcuma amada (2n=42)
13d-Curcuma aromarica (2n=63) 14d-Curcuma caesia (2n=63)
15d- Curcuma comosa (2n=42) 16d- Curcuma decipiens (2n=42)
21d I , .
Fig.17d to 2 Id-Comparative idiogram plates o f various taxa studied:
17d- Curcuma haritha (2n=42) +, 18d-Curcuma longa (2n=63)
19d-Curcuma malabarica (2n* 20d-Curcurna rekfacanta (2n=63)
2 l d-Curcuma zedoaria (2n=63)
ChomosancTotalknglh Shatam ArmraW F% Vdu~m PIA Nniue of
Typd M P ~ (1) m
,wn&Mm
Neafty-
Neafty Me dan
Ne* Median
wally Median
Near+# Me b a ,
W-
E l e a r f V Md
Nearty Median
Nearty Median
Wrty Me&
Neady Medan
W* Median
Neaiiy k 6 a n
Ne s r l y?dmc dan
Neatly Median
Namal sum& chornosame nmber
KarVotVpe formla
Number of dromwrne pais v& secondary c ms k l h
Ran@ of dvomosome k@h h pm
Total dromc6ome lerrph m pn
Amage chomavwne k+h in pm
T.F.vakr (X)
Varidon coemdent &. C)
Dispady index (D.&
Tdal vokune of chomommea
in v=
Uuornmnme Total laylh Short ann h ratio F?4 Vohnne PIA Nahre of
TYW hl rm (0 in pm= Rimary
B 0.87 0.30 . 1.96 34.19 0.017 11.18 NearlySubmcdn
B 0.80 0.30 1 .M 37.81 0.017 12.05 Nearly Meban
B 0.73 0.36 1 .05 M.76 0.012 12.96 Nearly-
B 0.61 0.27 127 U. 00 0.010 12.46 NearlyMedan
B 0.61 0.28 1.14 46.62 0.013 12.37 Nearly Median
B 0.59 0.27 1.16 4626 0.015 13.89 Nearly Medan
B
B
B
C
C
C
C
C
'Not in pak
Nearly Medm
Nearly M a n
Neady M&
Naarly M*
Ncarlr-
tiearty M&
Nearly Medan
Nearly M*
ChomosomcTohlknglh Shortam Arm& F% VoCme PIA Nabre of
Type hpm langhmvm (0 h ltw h a t y
A 122 0.40 1.39 3320 0.084 841 Nearly Subme&m
028 21.31
A 1 .M 0.38 1 .W 35.38 0.072 10.36 ~~
028' 25.93
B 1.07 0.47 1 28 43.84 0.049 9.03 Nearly MedRl
B 0.M 0.38 , 1.48 40.46 0.038 9.60 Nesrly Medan
B 0.m 0.37 1.41 41.32 0.032 9.40 Neady Mcdan
B 0.85 0.37 1.32 43.M 0.051 8.75 Nearly Median
B 0.62 0.m 1.13 47.07 0.036 10.77 Naarly Medan
B 0.78 0.31 1.51 39.85 0.034 9.74 Nearly Madsn
B 0.76 0.37 1.011 48.10 0.030 10.95 Neady Medgl
B 0.75 0.36 1.09 47.87 0.028 9.58 NemtjMedan
B 0.73 0.31 1 .M 42.02 0.036 9.87 Nearly Mesan
B 0.n 0.32 1.26 44.34 0.032 9.15 Nearly Median
B 0.71 0.32 1.24 44.72 0.m t0.m early ~e da n
B 0.70 0.32 1.18 45.a 0.033 10.31 Nearly Medan
B 0.69 029 1.35 42.59 0.024 9.98 Nearly Medan
B 0.68 0.28 1.42 41.37 0.021 9.78 Neatly Medan
B 0.67 0.29 1 .34 42.71 0.017 11.19 NeatiyMedan
B 066 0 29 1.27 44.13 0.027 9.77 Neady Median
B 0.65 0.29 1.25 44.50 0.021 1022 Nearly Median
a mm0 a a mT - m Wm!cm &wtrk f% Va)unr, PIA
m h~ar (I) h e Pllimay
camwbl
aft 1M
am tab
Fig.22d to 28d-Comparative idiogram plates of various taxa studied:
22d- Hedychium coronarium (2n=34) 23d-Hedychium coronarium (2n=5 1 )
24d-Hedychium flavescens (2n=5 1 ) 25d-Hedychium spicatum
var.acuminatum (2n=34)
26d-Kaempferia galanga (2n=54) 27d-Kaempferiapulchra (2n=22)
28d-Kaempferia rotunda (2n=44)
Nmnal ma S c drommme nmber
KaptVpe-
N l b n b e r ~ f d r ~ p a i a w i h s e r md s r y ~ o n
R- of dram- h pm
T o t a l c l r r m o m m c ~ h ~
Avuwe chomoroma bgul h pm
T.F.waho (K)
v* cadkh-4 (VC)
Dspsrity index (D.0
Total v-e of cho+msonw
m w
Ur ommwToWkf @ Shorlam Arm- F% Vo*ma PIA t4abcd
TYpc hpm knghhpm in pm' m
Nmnal somalk drMDlome h
K.yo(ypc-
Nmbsr~fohomosomapa*Jwihsamndsrymnsic(ion
Rnoed-len(llhhW
Total Ehnnnaome bf@ m pm
Average dmmooome *rgh h W
T F . W W)
VaIianOn oamcia W.C)
indca (D.1)
Tatdvdunddmmaromes
h pm'
ChmnrnmTcUh~b@J~ Shatsm km* F% Vdane PIA Nabreof
Trpe inw r nmw (0 m
Near)y w n
NcarlV-
Nearly Sdmedm
Nearly-
Nearly Medm
Nearly Medan
Ntarty Submedun
Nea4yShm*
W w n
N W-
N e w Madan
Nearly Medan
Neady M a n
Near(y W n
Nearly Medan
Nearly Medan
Nearly Me&n
Nearty Mean
Nearly Medsn
Nearly W n
Needy Me a n
h a l som& d u o m m munk
KaVotypetam-
Number of ctuomcame psis with semndary comtriclm
Range of dwomomme h pm
TDPd duomusome lengh h pm
Average dwommme knglb h p
T.F.value (X)
vwiahl caffidcnt W.C)
Oaparily i d e x @.I)
Total vOLsne of dmmwms
h pm'
UromaxrmeTotallength Shatarrn Armratin F% Vokmn PIA Nabre of
Type h v m mmpm (r) h pm'
Primary
B 0.72 0.28 I .S 39.11 0.009 12.03 NWM-
B 0.86 0.25 1 .SS 37.70 0.01 1 11.78 Nearly Me&n
B 0.63 0.30 1.14 46.71 0.m 16.M Nearly Me&
B 0.62 0.24 -. 1.57 38.85 0.010 13.18 NearlyMedan
B 0.61 0.28 1.18 45.97 0 . m 13.02 Nearly Median
B 0.61 0.25 1 .U 42.96 0.007 14.34 Nearly MecKan
B 0.60 0.26 I .U 42.04 0 . m 13.19 ~esrl y M&
B 0.58 0.21 1.70 37.01 0.M7 14.33 Nearly Submedian
B 0.57 0.29 I .01 49.72 0 . m 12.86 Nearly M&
B 0.57 026 1.21 45-32 0 . m 13.38 NW ~ e d a n
B 0.58 0.23 1.43 41 .W 0.008 13.03 Nearly Medan
B 0.55 0.24 1 .26 4. 27 0.008 12.85 Neatly Median
B 0.54 0.26 1.05 48.75 0.006 13.78 Neatly Median
B 0.53 0.24 125 U.49 0.008 13.12 Nearly MeQn
B 0.52 023 126 U22 0.008 13.11 Neatly Median
B 0.51 0.23 1 -20 45.43 0.007 14.85 N e w Median
B 0.51 0.22 1.32 43.13 0.008 13.91 Nearty Median
C 0.50 0.24 1.10 47.55 0.006 14.02 Nea* Median
C 0.48 0.22 1.15 46.46 0.005 13.72 Nearty MeQn
C 0.45 0.m 1.29 43.4 0.006 13.69 Nearty Median
C 0.42 0.20 l 08 48.18 0 . W 14 l l Nearty Medlan
Charm.muTdbnph Shutam Arm- F% VoLme PIA w e of
Trpe hw W m W (1) h pm. mmay
Nearly Me&
Nem-
Nearly M&
Nearly Me6an
Nearly Median
Nearly M e n
Nearly Medan
Nearly Me&an
Ne* M&
Nearly Medan
Nearly Medan
Nearly Median
Nearly Median
Nearly Medan
Nearly Medan
Nearly Medan
Nearly Median
Nearly Medan
Nearly Median
Nearly Median
Nearly Median
N-a d dranaomc lumber
KaFptVpemJs
Numbnof dvomocomepai swi t h~eonsbi di m
Range of dmmwwme lengih h pm
T0taldvomouwnelengihh)rm
Avenee- Wbi nn
T.F.vah~ (X)
v* mdidarn ( VC)
Disparily kdex (DJ)
Total voCm of dvoma9omes
hpm'
Tme-34: ~ L ; u p ( y p e ~ d ~ p u l n n h
ChamrswToWlengih Shatm Annmb F% Vdme PIA Nalwe of
T w in rm kmg~h h pm (r)
h pm'
-W
A 0.95 0.29 l .34 30.57 0.073 14.63 ~wr ( y shmedan
027 28.42
A 0.82 028 1-09 33x3 0.044 14.57 Naa(ySubmedan
024 2927
A 0.78 0 s 123 33.34 0.058 14.44 Nearly Stbmaban
021 25.64
B 0.72 028 1.62 37.97 0.053 11.74 Near)yMcdan
B 0.67 020 + 1.U 41 22 0.058 13-19 Ne d f Medan
B 0.85 0.25 l .W 30.40 0.056 16.54 N W Mcdn
B 0.63 026 1.39 41.84 0.1721 12.94 Neattf M&n
B 0.62 0.26 1 .M 42.82 0.013 12.65 Nearly Medan
B 0.61 024 1.51 39.m 0.m 1 2 . ~ early M*
B 0.58 025 120 U.92 0.m4 13.97 Nearly Meda
B 0.57 022 1 .S3 38.13 0.01 1 15.29 Nur(yMadian
B 0.56 0.24 1.33 42.98 0.025 15.40 Nesrly M a n
B 0.55 0.25 I .n 44.85 0.010 13.22 NesrlyMedan
B 0.53 024 l22 44.W 0.028 14.54 N e w Medan
8 0.53 024 1 .20 45.37 0.014 14.28 Nearly Medan
B 0.52 023 1 24 44.61 0.01 1 15.21 N e w Mcdsn
B 0.52 022 1.30 42.03 0.019 15.05 Nesrly Meda
B 0.51 020 1 .S 39.20 0.014 12.75 N e w Medvln
B 0.51 0.25 1.04 48.93 0.016 14.06 Nearly Median
C 0.50 021 1.39 41 .S 0.019 15.44 Nearly Medan
C 0.48 0.21 1.32 43.17 0.012 13.48 Near(yMedan
C 0.47 0.18 1.57 38.99 0.015 14.65 Nearly Median
C 0.46 021 1.13 46.94 0.015 14.28 Nearly M&n
C OAZ 0.19 l .25 44.41 0.011 1520 Nearly Medan
Normal sorns(ic chrommome numbef
Ksrydyps-
Number d chromosome pak wim seadar y C m S l M M
Range of rhmmDsom IengIh n pm
Total draDswe n pm
Awagedmm6omekngIhmpm
T.F.va*e (K)
Varialion caffident &!.C)
Dispany ixLr (0.1)
Total wAme of c h r o mme s
m Mm'
PIA -of
12.93 Nearly Mcdan
14.61 N- Medan
13.90 NemiyMedan
15.49 ~e ar t y Median
13.88 Nearly Medan
14.68 NeMy Medan
15.81 Nearly M&
14.86 Nearly Medan
16.39 Nea* Median
15.02 NeMyMe&m
15.77 Nc a r ( y kdm
15.16 Nearly Madan
15.62 N W Meban
19.68 N W Msdkn,
16.53 Nearly Median
15.46 Nearly Median
15.56 Ne* Median
38.45 Mcdan
21 .M Nearlv Median
Nmndsimla&chmmoeommmlbef
Som& vaia6on runba
Kafyolpetormlla
N u mb of chom- p& vrim eecnndary comtidh
Range of chomosnm *rglh h pm
Tot al cham~l nckngl hhyl
A m - m h i n
T.F.vaCls (74)
varialan c a c m (VC)
DisprnitV idar (D.0
Total M*me of dromoswnes
h pm'
PIA
Nemly Medan
W-
-M*
Nealtj Medan
Nmty Medan
Ne&y Me&
Nmty Median
Nearhe Me&
Normal somabc dvomosm mber
Kap*pe fmdE
Number of dromosome pairs wiIh secondary omlndm
. .
Range of ckomomme lengm m pm
TcW dromoswne *ngh h pm
Avaagechomosomalm#hh~m
T.F.ua& (X)
Vaiatan cnetX4ent (WC)
ida (D.!)
Total voCHne of chrommes
m pm=
UmmommcTotalknglh Shatarm &mraSD F% VoLme PIA Nb.ad
Twe inpm knghh pm (0 hlurr. Rinu).
cm(lidm
W-
W-
W-
W-
h*-
W-
--
W-
W-
h*-
W-
N W -
N W-
W-
N W
N W -
N W -
N W Medan
Neatly Me&%
Nea* Medan
Neatly Subme-n
NamalaMIicdromasomnwnba
Kayollpsfamla
N u n b e r o f d r w ~ g o m c p e i s ~ ~ ~
. .
Rang.30fch-knghinw
Total chom~omc kqUh h pm
Average ctmmorpme fengh h pm
T.F.nlus (X)
variatbn wemed P.C)
osparitV * (D.0
Totalv&lneofdmnwaomcs
nlwn'
Uromomme Tata length Shat m Arm ratb F% Vo*mc PIA N&e of
T w hpm fenghhpm (0 h pm'
A 1 .34 0.38 l .m 2B.S O.W 13.42 Nearly Shmefmn
0.31 23.85
A 1.28 0.34 1 .W a8.82 0.032 13.75 NeadyS&mxh
0.31 2422
A 0.98 0.30 1.42 30.75 0.027 13.59 Neafly Wmdm
B 1.07 0.49 1.17 46.17 0.019 12.87 Nearly-
B 1.03 0.51 l .M 49.10 0.020 12.15 NeartyMedsn
B l .M 0.30 2.38 29.111 0.034 12.70 Neady Shmdan
B 0.60 0.34 1 .M 42.76 0.M2 15.46 Neady M&
B 0.76 029 1.67 37.52 0.016 13.49 NeadySdme&n
B 0.73 028 1 .M 37.79 0.- 15.73 Neady Mcchm
B 0.72 0.31 1.32 43.16 0.015 15.55 Neatly M a n
T 6 3 9 : D e t e l e d ~ a d p A d l k m u n ~
C h o m g o m e T ~ ~ Shor t m Ar mdo FX Vo*ms PIA Nahre af
Tvpa h) ~ m * ngmh~m (0 m@ Wluy
A 1.37 0.63 I P 28.35 O.W 13.15 Nearly S&mc&n
O X 18.911
A 1.31 0.46 1 .43 28.82 0.034 13.47 Nea* Sdmedan
033 25.19
A 1 .O1 0 . 9 1 .U 24.75 0.029 13.31 Suknedan
0.31 30.63
B 1.19 0.37 1 .U 39.70 0.044 15.33 Nesr)yMeh
B 1 .M) 9.31 1.73 46.17 0.020 12.61 Nealy Me&
B 1.06 0.36 1.35 49.10 0.022 11.90 Nee,iyMcdan
B 1.03 041 1.07 29.81 0.038 12.43 Neafty Submadan
B 0.98 0.32 1.32 3 7 3 0.016 14.35 Nea* Wmedan
B 0.91 0.40 1.23 42.33 0.015 13.02 Near)rUedan
B 0.87 0.33 1.51 41 25 0.022 12.30 Nea*Uedan
B 0.81 0.34 1.07 42.76 0.023 15.14 Near(ykSan
B 0.78 027 1.14 37.52 0.017 1322 Neatly % M a n
B 0.75 0.30 1.56 37.79 0.008 15.41 Neafty Uedan
B 0.74 027 1.43 43.16 0.016 15.23 Nearly M a n
B 0.73 020 1 .50 42.11 0.008 14.33 Nea* Mcaan
B 0.72 020 1.14 33.12 0.012 14.53 Nearly Sllbtne&n
B 0.70 027 1 .S5 35.72 0.01 7 14.1 Nsafty Wm&
B 0.68 027 1 .m 4028 0.01 7 13.98 +Medin
B 0.66 028 1.54 39.99 0.012 15.92 Nea* k 5 m
B 0.60 025 l .W 45.88 0.012 16.22 Neafty Me&
B 0.55 025 1.74 44.39 0.006 16.81 Nea*W&an
B 0.54 022 2.05 46.00 0008 17.27 NeadyMedan
B 0.53 0.22 2.45 47.93 0.015 17.89 Nearly Me&
C 0.47 021 1 .E6 4629 0.006 17.47 Nearty M e
Namal som& rhwnmomc number
'(wow fofln&
Nu- of drommwne pais A secondary combicb;on
Range of drm~some lengh h pm
T a t d d r ~ ~ h p r n
AvsnOcehanorpomcMmpm
T.F.*r*e W)
var*Son ca(lident (V.C)
Dispailyindu (D.0
Tot4 vohne ddrommmes
h p d
TabWD. D e M e d l r a y d l p e a r a t + s o f ~ ~
CIranosorne TOW- Shotterm Armratin F?4 VoLme PIA Nalue of
Type h p ~ n g h h mp (r) h i~d Rimy
A 1.14 0.35 l .37 31.11 0.070 7.61 N e w Submedgl
0.31 28.32
A 1.01 027 1.75 26.34 0.01)7 6.0s ~~
0 28 27.72
A 0.93 0.30 1 26 3224 0.120 7.92 Ne&yWmedm
0.25 26.88
B 1 .W 0.49 121 4521 0.MI) S. 7l NeaftyUedsn
B 1 .M 0.47 * 1.24 44.54 0.041 7.86 Nemly WhIl
B 0.99 0.36 1.78 35.98 0 . m 7.48 ~e&y &&wedan
B 0.91 0.38 1.41 41.58 0.027 8.50 Nearly Medsn
B 0.88 0.40 1.18 45.84 0.045 9.39 ~earlyMe&m'
B 0.85 0.37 1.32 43.09 0.041 8.14 NemlyMedhn
B 0.82 0.38 1.16 46.41 0.054 8.m NW-
B 0.79 0.34 1.31 43.22 0.052
8.46 Mealy-
B 0.77 0.37 1 .W 47.86 0.043 8.98 Ntafty Msdn
B 0 . n 0.30 I .ss 39.17 0.037 9.14 N* ~edan
B 0.76 0.33 1.32 43.14 0.050 8.73 Nearly Medan
B 0.75 0.30 1.48 40.42 0.054 9.08 Neal y M&
B 0.73 0.32 1.28 43.98 0 . m 9.57 ~ e &y Mean
B 0.71 028 1.55 39.18 0.036 8.98 N W Medan
B 0.70 0.33 1.15 46.48 0 . m 9.52 Nba)y M&
B 0.68 0.29 1.33 42.97 0.028 9.30 Nsrrly Mdan
B 0.67 0.26 1.61 38.31 0.031 9.07 N e w Median
B OM 0.27 1.40 41.M OM1 9.03 Nearly Mcdan
B O M 0.27 1.37 42.11 0.040 8.82 N e w Medan
B 0.62 0.25 1.45 40.80 0.024 9.26 NaMy M&
C 0.47 032 1.15 46.n 0.043 10.34 Nearly M*
Namalsomidrmmnnemmnbu
Kaptypa-
NMl berof obnrosomopahwi h~mrabl cbon
. .
Ran* of chnnsome lengh h pm
Tot4chwrome*nghmpm
Amaoechr ao. am* ngl hh~
T.F.vaClc (W
Varialan amRhd (VC)
m h m . 0
Total vobmed dramaromes
m pm3
Umm-Td- Shatmm Arm& F?4 Vo*ms PIA Nabn of
Trpe h a (0 h vw Rimry
Ncaly M-
M M -
Neatly Medan
Nemiy Me&m
N m t y Medm
Nesrlv-
W-
W-
Neatly Mcda
Neatly Medsl
Nearly Me&
Neatly Medn
N e w M&
Neaty Me&!
Nearly Me&m
N e w M&
Nearly Mer h
Nearly M&
Nearly Median
0 16 1 .06 UI.e-5 0.003 20.14 Nearly Me&n
Normal somalk chomasome numbn
Swnatic-nntber
KaryotVpe -
N u m b e r o f ~ p a i s w i l h s e m m & - y ~
R ~ o f c h o m s r v n s ~ i n p m
Total chomoseme length in pm
Average dxomowme length m pm
T.F.value (%)
variab;on c o t M (VC)
Dcparity iwkx @.l)
TotalvoLnneofdvorowmes
m pm'
Table-42: Detailed karyoolpe analysis of nPtlarla cvdrnomun N. b4aWw
Nearly MeSan
W-
W-
Ncaly-
W-
Nearly Medan
W M &
W-
Neartj M&
Nta* M*
NealVy-
W-
N e s r l v wn
N M Medan
Nearly Medan
Nearly M&
N e w M&
Nea* Median
Neatly M&
Nearly M&
Neatly M&n
Nea* Medan
Nonnalsonu(cchDmDonrRmber : 48 (Fa. 42b)
: A4 812 C32
N u n k r d d m m o s o m e p i s ~ s 4 c o 1 ~ b r y ~
. .
2.00
Ranaeof chomoMeMi nl l m : 0.715.33
To( ddmmosomemi nw 23.26
Awaob-mi n(rm 0.48
T.F.F.HC* (W) 42.57
Vaiacm cdki ent (VC) 18.81
-*(D4 3820
TotslvoLmeafdmnwgrmsa
V' 0.30
T W D a b d ~ ~ o f E h t & h ~ ~ . M y r a r r
ChornosarTold*n(lh Shortarm Annra(k FK V o h ~ PIA NaOra of
T w inlm m i n v m (0 in p d primay
A 0.71 024 126 33.58 0.009 16.09 Neatiy-dan
0.17 n.oc
A 0.66 0.18 1 .S 20.17 0.m 17.77 ~ e a t y ~ u b t n t ~ ~ m
0.17 25.75
8 0.6s 028 l .M U 6 7 0.011 13.79 Nearly Medsn
B 0.57 025 126 44.16 0.005 16.05 Neatly Medan
B 0.53 0 m 1 .m 47.91 0.W7 17.12 Nearly M&
B 0.53 024 124 44.61 0.m 18.38 NearlyMedan
B 0.52 026 1 1.03 49.1 7 0.007 18.10 Nearly Merian
B 0.51 021 1.41 41.48 0.003 17.33 NeartyMedan
C 0.50 023 120 45.40 0.046 1721 N e w Medm
C 0.49 021 I .S 4 2 2 0 . m 16.83 NW Me&
C 0.49 O M 1 A5 40.78 0.0% 1723 N e w Mecbn
C 0.47 020 1.37 42.14 0.- 16.98 Nearly MeGan
C 0.46 om 1-35 1259 0.m 1722 karly ~er i an
C 0.45 021 1.15 46.g 0 . m 1720 NearlyMerian
C 0.44 0.17 1.56 39.10 0.007 15.81 Neatly Medan
C 0.44 0.m 1.18 45.96 0.WB 17.82 Nearly Median
C OM 020 1.15 46.41 0.001 16.91 N W Me&
C OA3 030 1.18 45.87 0.005 17.13 Nearly Madan
C 0.42 0.18 1.33 42.83 0.007 17.97 Neafiy Medan
C 0.41 o.m 1.11 47.43 0 . m 19.27 Neatly ada an
C 0.40 0.18 1.26 44.16 0.005 18.99 Nearly Medan
C 0.39 0.18 1.21 4529 0.OM 17.97 Nearly Medan
C 0.38 0.14 1.68 37.33 0.015 2231 Nearly Submedim
C 0.33 0.15 l .24 44.59 0 . W 18.88 Neadv Medan
Nwmal sunatic dvomrmme numba
KaVotVp-
Hmbn of dmmoewne pah * secondary combdm
. .
Range ol chanoowm kngm h pm
Totd chmmogane kngU~ h pm
AwragedmmcgamekngLhhpm
T.F.rah (X)
vsriscm c o e M ( VC)
CJisparily i d e x (D.0
Totavohnneolchomoromes
h pm'
ChwrosomeTotalkngm Shatann Arm& F% ValPne PIA Nabm of
Tvpe hpm kmgthhpm (0 in pm=
A 0.86 0.29 I22 3328 0.032 10.82 Nearly Submedan
0.2 27.78
B 0.63 028 1 29 43.73 0.012 12.68 Neatly Median
B 0.62 0.29 1.15 46.52 0.014 12.59 Nearly Median
Nearly Medsn
Neady Madw
Nearly Medan
Nearly Median
Nea* Medan
Nearly M&
Nesdy Me&m
Nemiy Medm
Nearly Me6ml
Neady M&
Neady M a n
Nearly Medan
Nearly M&
Nearly Median
Nearly Median
Neady Median
Nearly Meckan
Nearly Medan
Neadq Medan
35d- AIpinia rerumbet ( 2n48)
36d-Alpinia zerumbet var. varigata
(2n=48)
37d-Amomum cannicarpum (21148)
38d-Amomum hypoleucum (21148)
39d-Amomum muricatum (2n=48)
lum
40d- Amomum pterocarpum (21148) 41d-Elettaria cardamomum CV.
Malabar (21148)
42d-Elettaria cardamonrum cv.Mysore (2n=48)
43-Elettaria cardamomum cv.~azhukka (2n=48)
a. Chromosome numbers in the genera investigted
Chromosome counts are made on forty three members of Zingiberaceae
prevailing in South India. Cytologically the family is variable. The
chromosome spectrum varies from 2n=18 to 63, with majority of the species
concentrated in the number 2n=48 followed by 2n=63,2n=22 and 21142 (vide
Table. 45, Fig. 44). The presence of identical numbers in unrelated genera is a
noteworthy feature i n this family. The presence of such widely dfferent series
of chromosome numbers in various species of the same genus and in genera
placed under different tribes,indicates that the different chromosome numbers
can be derived one from the other. However, it is quite remarkable that the
chromosome number in certain genera shows a constancy as in the case of
Zrngiber, Afpinia etc. Perusal of the literature (vide Table. 45) reveals that,
with the exception of a few genera, where the available cytological data is very
meagre; many taxa exhibit a wide range of chromosome numbers.
3
Subfamily: Costoideae
Two taxa belonging to the type genus Cosrus have been examined. The
chromosome numbers of 2n=18 and 2n=36 have been found in Coslllr
malortreanus and Cosrus speciosus respectively. Based on the previous
literature it is clear that, out of the fifteen cytologically known species almost
the entire bulk are diploid5 with 2n=l8, and the genus seems to be more or less
homogenous. However, a few stray instances of intra-specific variations are
there, including the present tetraploid Costus specrosuv (2n=36) and a
hyperploid (2n=108) are reported by Chakravorti (1948 a). Thus the genus
Cosrur shows a constancy as regards the chromosome number.
Subfamily: Zingi beroideae
Tribe-Globbeae
In this tnbe also two taxa belonging to the type genus Globba have been
exambed. The chromosome number of G. cernua is 2n=48 and of G.
ophioglossa is 2n=22. According to the previous reports the somatic
chromosome numbers evaluated in this genus so far are 211=22,24,28,32,44,48,
64,80, and % (Ramachandran, 1969 a & 1969 b). Thus the genus Globba seems
to be highly plastic in its chromosome number.
Zingiber is the lone p u s of this tribe. All the six species investigated possess
2n=22 and this agrees with the previous reports. Also in all the species no
somatic variations are found. The absence of any somatic variation numbers
suggests the stability of the genomes. The absence of somatic variations, in
numbers might have probably been due to the cytologically diploid nature and
normal disjunction of the chromosomes in these species during meiosis (Riley
and Chapman, 1958). In addition to the normal 22 chromosomes 2 B-
chromosomes were rekrted for the species Z offrcinale by Janakyammal(1945).
Unlike other species 2. officinole lacks sexual reproduction. There is no seed set
in this species. According to Ratnambal (1984) the remarkable similarity of the
chromosome data of different cultivars of Z oflcinale studied from different
geographical regions is to be attributed to sexual isolation and continuous
vegetative propagation. The chromosome number 2n=22 found in 7- cerntrum
seems to be novel. Authentic cytologic reports are not available on ths species.
Mahanty ( 1 970) after a detaled cytological study of the Zingiberales, concl uded
that the genus Zingiber should be included in the tribe Hedychieae. This was
later dismissed by Burtt and Olatunji (1972). Nevertheless, Beltram and Kiew
(1984) stated that chromosome number gives no evidence to support the
treatment of Zingiber as a separate tribe. From the present study also it is clear
that the genus Zingiher shows marked differences from the other members of the
tribe tledychieae as regards the chromosome number, and it can be considered as
a seperate tribe.
Eighteen taxa belonging to three genera have been examined. The chromosome
numbers of individual taxa ranges from 2n = 22 to 2n = 63. But inspite of these
differences t . exist a relationship between the different genera and species as
evi'denced by the frequent occurrence of inter and intra specific variations. The
chromosome numbers and the associated variations provide a clue in determining
the status of a species (Sharma, 1976).
The genus Curcunm represents eleven species. In this genus two clear lines of
evolut-ion have been noted with chromosome numbers 21142 and 2n=63. The
species with 2n=42 are diploid species whereas the others are triploid spi es.
The dominance of these two numbers in Cwczunu is very much evident (vide
Table 45). These two numbers seem to have originated from a common ancestor
or one from the other. Most of the species are not seed setting. Nambiar et al.
(1979) reported seed setting in a variety of C. aromotica having 2n=86
chmmosomes. However, Velayudhan et al. (1 990) pointed out that the species
was wrongly identified as C. aromtica. In the present study, it is revealed that
C. momfico jmssesb 2n=63 chromosomes. Ramachandran (1969) also
reported C. oromntica with 2n=63 and 2n=86 chromosomes. The inconstancy of
chmmosome numbers noticed in the same species by several workers (vide Table
45 ) points out the existence of both diploidy and triploidy in the same species.
Among the various species C. comoso, C. huritha and C. malobarica with 2n=42
and C. aemgenoso, C caesio and C. ruKtacunta with 2n==63 are reported for the
f i s t tinie.
The genus Hedychium also shows two distinct lines of evolution with 2n = 34 and
2 ~ 5 1. Of the four tawa studied Hedyehiurn coronurium shows both diploid and
triploid numbers with 2n=34 and 2n=5 1 . Intraspecific variation is thus very well
pronounced in this taxa H. $uve.~cens shows 2n=5 1 whereas, H. spit:urum
var.acuminatum shows 2n=34. Somatic variations are found in most of the
members of Hed'cltiunz which had already adopted vegetative means of
multiplication. In vegetative1 y reproducing plants variation leading to speciation
is caused by chromosome alterations in the somatic tissue (Sharma, 1956;
Konvicka and Levan, 1972). Raghavan and Venkatasubban (1943) and
Chakravorti (1948 a) reported 2n=54 for H. coromriom. The plants examined
by these authors are possibly hyper tnploids. The occurrence of different
cyotopes in the same species suggest that chromosome number in this species is
not very stable (Morton, 1962).
As regards the chromosome number the genus Kaempferia seems to be
heterogeneous with the occurrence of somatic chromosome numbers 2n=22,
2n4-4 and 2s54. A similar trend has also been observed in the previous
literature (vide Table 45). For the species K. pulchra, the somatic chromosome
number is found to be 2n=22. K. rotundo root tip cells showed 211-44 and this
agrees with the count reported by RarnachanQan (1969) and Omanakumari and
Mathew (1984). The reported occurrence of the taxa of this species with 2n=33
(Chakravorti, 1948 a) and 2n=54 (Raghavan and Venkatasubban, 1943) shows
that this species also exist in at least three different cytotypes. In K. galanga the
somatic chromosome number found with 2 ~ 5 4 agrees with the previous reports
(vide Table 45). However, Sharma and Bhattacharya (1959) reported 2n = 22 for
this species. Hence the genus Kaempferiu also seems to be highly plastic in its
somatic chrornosomk number.
Fifteen taxa belongng to three genera have been examined. The normal
chromosome number is found to be 2 d 8 invariably in all the taxa studied. This
shows the stability of the genomes of ths tribe at the inter and intra specific level.
In the previous literature (vide Table 45) also 21148 is the dominant somatic
chromosome number in this tnbe. This shows the inter and intrageneric
relationship among the members of this tribe.
All the species of Alpinia studied were found to be with 211-48. Of these A.
purpuratu, A. zerumhet var. vurigu/a, A. smithiae are new1 y reported for their
chromosome number. The chromosome number was found to be constant with
very rare occurrence of somatic variation among these species. All these species
79
are seed setting and propagated both vegetatively and by seeds. Interestingly the
two exotic members studied also show 2n=48. This shows, though the species
are separated geographically that the chromosome number is the same. So these
species are genetically more or less stable.
In the genus Amornum also the chromosome number was found to be constant
with 2n=48 in the four species investigated. In A. muricaturn the somatic
chromosome number 2n=48 is reported for the first time and this seems to be the
predominant number in the previous literature too (vide Table 45) This shows
the stability of the genus at the intm-generic level as regards the chromosome
In the lone species of the genus Elettaria, three cultivan of EIettarza
cmdamomum showed the chromosome number 2n=48. This is in conformity
with the d i e r reports made on the same taxa (vide Table 45). However, 2n =50
and 2 ~ 5 2 were also reported in these taxa by Chandrasekhar and Kumar (1 986).
This may be aneuploid numbers. In the present study somatic variation with
2 ~ 5 0 has been noticed for the cdtivar Malabar.
Table 45 : Index to chromosome numbers in zingibereceae
Taxon Previous Freeat Reference
Count Count
P) (2d
Subfamily: Costoiedea
Gemls: Gmts
Venkatasubban ( 194
6)
Venkatasubban ( 1 946)
C. cyIirsdnms (phrn.) 16 Boehm ( ml )
Jacq
18 Simmonds (1954)
C. elepns hort
C. jhedncknii
C. peiersen
C: . i g m ~ Brown
Lucmsionus Braun et
Schum
C. dwt i eamr s wend1
C. musaicus hort
C. niveopurpt~reus Jacq
C: pidus
C. ~ p e c i ms Smith
Tribe : Globbodeae
18 GeOrgor~ (1 936)
Venksrtasubban (1946)
Simmonds (1 954)
Venkatasubban ( l 946)
Sato (1948, 1960)
Simmonds (1954)
Raghavan,
Venkatasubban { 1 943)
Chakmvorci ( 1 948)
36 Shma, Bhattachary a
(1 959)
Genus: Globba
(; . bulblfiru Roxb 44 S h m a & Bhattacharya
- (1 959)
G. albijluru vllr aureo 32
@dl) Holtt.
G. clourhi Baker 24
G. b k e r i clarke 22 (24)
24
24
G. ophioglossa Wi& 22
22
G. racernoso Smith 24 (23-25)
24
32
G. schomburgkii Hoot 48
F. 48
c.64
G. cemua Bak 32
48
G. cernua ssp cornea, 48
Lin
G. patens Miq S.S. 32, 48
G. wznzt ii C .H. Wright 32
G. pendula ROM-S. 32, 48
G. leuwtha Miq.
G. f&ata Bid1
. Sub-family:
Zingiberoideae
Chakravorti ( l 959)
S h a m & Bhattacharya
(1 959)
Raghavan &
venkatasubban ( 1 943)
Ramachandran ( 1969)
Mahanty ( 1 963)
Mahanty ( 1965)
tarsen ( 1 972)
Sharma,Bhatfacharya
( 1 959)
Sachava ( l 977)
Meha& Sachdeva (1979)
Mabt y (1965)
Bhattachary a ( 1 968)
Lirn S.N Siew Ngo
( 1 972)
Ramimchandran ( l %9)
Shanna,Bhattacharya
( 1959)
Mehra& Sachdeva ( 1 97 1,
1976)
Sarkiu er ai ( 1974)
Siew ngo ( 1 972)
Siew ngo ( 1972)
Lirn ( 1 972)
Mabanty ( I 965)
Lim ( 1972)
Tribe: Zingibereue
Genus: Zingiber
boehm
Z. cemum (2- ninrrnonr~ 22
(Graham Dalzell)
2. cylindricurn Moon 22
m 2. naesanum 22 22
2. oflcinale Rosc 22
2. meum Rosc
Z robens Roxb
Z wjghtianum
Thwaites
Z z e d e t Smith
cassum- Roxb)
Tribe: Hedyc hicre
Genus: Curtumorpha
A-S.Rao &
D. M. Vma
C longflora 0.
kuntze
Genus: Curcuma
C. am& Roxb
C- mgusrfolia Roxb. 42
C aromurica Satisb 42
63
Mahanty ( l 970)
Ramachandran ( l 969)
Sagiura ( l 928)
Raghavan &
C h a r a d (1948; 1952)
Sharma & Bhattacharya
(1959)
Ramachandran ( l %9)
Datta & Biswas (1977)
Krharaer 01. (1931)
Bisson, S. et.01. (1931
Raghavan &
Venkatasubban (2943)
Chakravorti (1 948; 1952)
S h a m & Bhattacharya
( 1959)
Rarnachandran ( l % 1,
1 969)
Chakravorti ( 19481,
(1952)
( 1959)
Ramac hmdran ( l 96 1 )
Raghavan ( 1959)
Raghavan &
venkatasubban ( 1943)
Chakravorti ( 1948)
( 1 952)
Ramachandran (1961.
C'. caeia
4
, C'. coniosa
C. harifhu Mangely
&Sabu
C. longa Linn.
wight.
C petlolota Roxb. 63
&l
Tribe: Hedyehieat
Genus: Curprrwrpikn
A.S.Rao ' 1 &
D.M.Verrna
Genus: Curcuma
C. amada Roxb.
1 969)
63
63
42
Ramachandran ( l 96 1,
1 969)
42
42
Sato (1948, 1960)
Chakravorti (1952)
C. anpst$olio Roxb. 42 Chakravorti (l 948; 195 2)
Shanna & Bhattachartya
( 1959)
63
64
Ramachandran (1961)
Raghavan ( 1 959)
oroma/ic Salisb. 42 Raghavan &
Venkatasubban ( 1943)
-p-
63
Chakravorti ( l 938. 1 952)
C- caesia
C. contosa
C decipiens Dalz.
C. harifha Mangaly &
Sabu
C. Ionga. h
C molabaric~ velay
C. neil@rrensis
wighr
C- petiolata Roxb
K. gbsonii
K. gilberti
Ramachandran ( i 96 1,
1969)
Sato ( 1948, 1960)
Ckakravorti ( l 952)
Raghavan &
venkatasubban (1 943)
Sharma & Battacharya
(1959)
Chakravorti ( 1 948)
Ramachandran (l % 1)
Sugiura ( l 93 1, 1 936)
IRam;sch;mdran (1961,
1 969)
Ramachandran (1 96 1 )
Venkatasubban ( 1946)
Chatrravorty, ( 1 948,
1952)
Shanna and
Bhattacharya, ( 1959)
Raghavan and
Venkatasubban, ( 1 943)
Raghavan and Arora
( 1958)
Ramaehandran
( 1 969)
Omanakmari
( 1 989)
Raghavan and
Chahavorty ( 1 952)
Chalcravorty ( l 948) &
Mahanty (1 970)
Bhattacharya ( 1968)
Bhattacharya and Sharma
(1 968)
-. -. - -, - -
---p
36 Raghavan and
K. involucralam King 22
ex. Bak
K. ovairfoIia Roxb. 22
K. spciosa Baker
Genus : R a m
Smith.
R a l p i ~ Royle.
R pu~puma smith. 24
26
Var. gganleo wall. ' 26
Tribe : Alpiacae
Genus: Aframaatum
K-Schum
A. iatfiIium K. chum 48
Genus : Alpinia Roxb
A. albo - line& 33
A. niqa ( g m ) Burtt 48
A. aqualica Rosc 48
A. braclearn Roxb 48
A. calcoratu Rosc 48
A. chinsensrs
A. fbrmo.sam
Bisson el al. ( 1968)
Sharma and Bhattacharya
( 1 959)
Bhattacharya and Sharma
(1%8)
Chakravorti (1951)
Chalcravurti (1948)
Bhattacharya ( l 968)
Bhattacharya and S h a m
(
Mahanty ( 1 970)
O m d k (1 989)
Raghavan and
Vd-bban(1943)
Venkatxtsubban ( l 946)
Sharma & Bhtxharya
(1 959)
Malik (1961)
Mahanty (1 970)
Mehra & Schadeva
( 1 976)
Mebran& Sachdeva
(1971, 1976. 1979)
Sachdeva ( l 977)
Raghavan &
Chakravorti ( l 948)
48
Sato D. 1948. 1 %0)
Chuong er al. (1 %3)
.4. japonica
A. malacceusis Rosc.
A. purpumfa (veil])
K. Schum
A. raflesiuna Wall ex.
Bak
A. sanderiana hort
A, rerumbet (pers.)
Burttt & Smith
(A-nutans Rosc.)
Var. variga fa hert
Genus: Amomum
Linn.
A. connicarpum
A. hpIeIucunt
Thwaiter
A. involucmtm Benth
A. mognljcum ~e ni h
& Hook
A. nturica turn
Beddome
A. pterocupm
Thawiter
A. sabulaium roxb
Genus: EIeftaria
Maton
E- car&momum
Maton
48 Raghavan &
Venkatasuban ( 1 943)
48
48
Sato D ( I 948)
48 48 Chkravorti ( l 948)
Raghavan &
Venkstasuban ( l 943)
Venktasubban ( l 946)
Ragavan &
Chakrvorti ( Z 948a
1952b)
Ramachandran ( 1 969)
48
(1 959)
48 Gregory ( 1 936)
( 1 959)
Sat 0 (1960)
Bhattacharya ( 1968)
Bhattacharya and Sharrna
(]%g)
Bisson et al. (1 968)
Ramachandran ([ 1969)
b. Basic chromosome numbers in various genera
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, 1 970; 1974; 1 978). It is known
that many basic chromosome numbers are involved in the origin of the polyploid
series in Angiosperms (Grant, 1 982).
The chromosome data so far known in Zingiberaceae roughly cover sixteen
genera in five tribes. Based on the previous reports and the present investigation
of somatic chromosome number, an attempt is made to discuss the possible
direction of basic chromosome number evolution in this family. Investigations
are conducted on forty-three taxa belonging to nine genera. 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. Cytological data on
diEefent genera of the k i l y Zingiberaceae show an array of basic chromosome
numbers such & 8, 9: 10, 11, 12, 13, 14, 16, 17, 21 and 25 of which X=12 is
predominant being present in nine genera (Hedychium, Roscoeu, Kuempferiu,
Globbu, Amornum, Alpiniu, Phaeomaria, Efettaria and Zingiber). It may be
possible that X=12 itself could be a secondary condition of polyploid origin of an
ancestral X=6 state. While X= l l is present in four genera (Zingiber, Kuempferia,
Globbu and Reneulmia). The genus Costus shows X=8 and X=9. Hedychium
and Cuutlqvo show X=17 and in Brachychilus X=16. In Curcumu and Hitchrniu
X=21, while Bosenhergiu shows X=25. It is rather difficult to determine the
family's ancestral basic chromosome number with absolute certai ni ty
(Omanakumai, 1987). The majority of the taxa shows X=12, while in some
others where the basic chromosome number is X= 1 1 there are variations with
lower (X+) and higher (X= 1 3, 1 4, 1 6 and 1 7) constitutions. However, the genera
Curcumu and Bosenhergirr show still higher numbers with X=21 and X=25.
This suggests that X = l 2 being the possible earlier evol ~ed lines in such tribes.
The other lower and higher numbers could be the derived ones from this by
descending and ascending aneuploidy (Omanakumari, 1989).
In Zingiberaceae the secondary base numbers might have evolved from the
primary numbers X=6 to 9. Grant ( 198 1 ) proposes that the orignanal primary base
numbers of angosperms range from X=7 to 9 and that they are ancestral in the
phylogenetic sense. From these primary base numbers many polyploid series
developed both by autopolyploidy and amphipolyploidy. The latter involves both
hyper and hypopolyploidy (Femandes and Leitao, 1984). Thus it can be
confinned that polyploidy has influenced and played a role in the evolution of
base numbers, mostly of the higher order usually by 'p1 yploid drop' (Dal tington,
1 956) and occasionally by ' polyploid lift' (Jones, 1970).
Among the South Indian taxa reported here X=l l and 12 are predorninent, and
the other numbers exist singly or in combination with these numben(vide Table
46). The besic chromosome constitutions of bibes with special reference to the
genera belonging to these tribes are briefly discussed here.
Subfarni ly : Cmtoideae
t
Under this subfamily only type genus Costu. and Tapienochilus are reported on
their chromosome number. The known chromosome numbers inbcate that
species of these two genera are based on three numbers namely, X=8,9 and 1 1 of
which X=9 is the dominant one. Most of the researchers are agreed on the
possibility of X=9 being the earlier evolved condition in the genus Costus. The
hgh levels of polyploidy in this number, known in a few species, and the infra-
specific polyploidy on this number (2n=27, 36) known in C specrosus.
(Subramanyam, 1 978; Nagendra and Abraham, 1 98 1 ; Subramany am and
Khoshoo, 1986) appear to strengthen this possibility. Regarding the species with
2n=44 based on X=l l (Chakravorti, 1948) we can find that it may be evolved
From the union between a normal gamete of a diploid species (n=8) and an
unreduced gamete of a tetraploid species with 2n = 36.
Subfamily: Zingiberoideae
Tribe: Globbeae
This is a small tribe with four genera. Chromosome reports are available only
with the type genus Globba. Chromosome numbers are known in eight species
from India as well as a few taxa from Malayan peninsula. The chromosome
numbers reported for the Indian species show that they are based on X=l l and 1 2
almost evenly. Malayan taxa show 2n=32, 48 and 80 (Chong and Siew-ngo,
1972). Among the reported Indian species those based on X=12 are mostly
plyploids while X= 1 I ones are all diploids with 2n=22. This situation appears
to suggest that X=12 could be early-evolved one and the other X=l l could have
evolved by descending aneuploidy. Concerning Malayan species Chong and
Siew-ngo (1972) have suggested X -8 as their basic number. In some taxa they
have reported allotetraploid, hexaploid and decaploid races on this number. In
the present study, the most possible basic number in G. ophioglossa (2n=22) is
X=l l , and in G. cernua (2n=48) is X=12.
Tribe: Zingi berae
The monogeneric tribe with genus Zingiber is known chromosornall y for about
eight species. The basic chromosome number falls under X=l l and 12 of which
X=11 predominates. Kihara et al. (1931) had reported a different count of 2n=24
for 2. of i c i de and this could be an aneuploid cytotype with X= 1 2 as the basic
number.
Tribe: Hedye hieae
This is a very large tribe with fifteen genera of which nine genera are
cytologically known. The chromosome data indicate that the various genera fall
under the basic numbers of X=9, 10, 1 1, 12, 13, 14, 16, 17, 2 1 and 25 of which
X=12 condition is the predominant one followed by X=l l . Most of the
cytologically known genera are monobasic, while a few others with dibasic and
polybasic conditions are also found. Out of the nine cytologically known genera
two have predominantly X=17, two have X=2 1 and one each has X= 10, 1 1, 1 2,
16 and 25.
In the genus Hedychium, which is a polybasic genus with X=9, 1 1, 12, 13 and 1 7
all the four taxa studied here show a basic number of X=l7. It is considered that
X-12 is the earlier evolved condition in this genus. Most of the previous workers
are also of the opinion that X=12 is the original number (Sharma and
Bhattachafya, 1959; Mahanty, 1970). Sharma and Bhattacharya (1959) have
considered the basic number X= 1 7 as being derived from 1 2 by p01 yploidy and
aneuploidy. According to Mahanty (1970) X=17 condition originated as a
secondary basic number from X=12. However, Raghavan and Venkatasubban
( 1 943) suggested that the genus has evolved along with the X+ and X=13 lines.
In Kaempferiu also the available data indicate that various species of the genus
exhibit several basic series with X=9, 11, 12, 13 and 14, of which X=l l is the
most frequent one. In the present work, the three species studied also possess
X=l l condition. Thus it is clear that the genus is p01 ybasic as regards the basic
chromosome number is concerned The predominance of X=l l condition
together with the occurrence of s pi es with different levels of polyploidy of this
number may be su&estive of X=l l to be the earlier evolved chromosome
constitution in the genus (Sharma and Bhattacharya, 1959; Mahanty, 1970). The
concentration of X=l l among the relatively less advanced group of species of
this genus provides support to the possibility of X=l l being the original
condition in the genus. It may be possible that the higher basic numbers X= 12,13
and 14 are the derived conditions from X=l l by ascending aneuploidy
(Omadumari and Mathew, 1984).
Interestingly, in the case of Curcumn still a higher basic number is noticed, ie.,
X=21. Such a high basic chromosome number in C~urcurnu might have been
derived either by dibasic amphidiploidy ie., the combination of lower basic
numbers 9 and 12 found in some genera of the family, or by secondary
p01 yploidy. (Ramachandran, 1 96 1 ; 1 969 j.
It is quite remarkable that polyploidy and, aneuploidy could have played a
prominent role in the evolution of basic chromosome numbers of the tribe
Hedychieae. Based on the previous reports (vide Table 46) one cannot disregard
the possibility of considering X=12 as the earlier evolved condition in many
genera from which descending and ascending aneuploid trends have occurred
resulting in the X=l l , 10 and 9 and X=13. The X=12 condition itselfcould be of
a possible polyploid origin from the primay condition of X=6. This possibility
appears to be stronger in the light of the basic chromosome number of the type
genus Hedychium in which different basic numbers vary from X=12 condition
both in descending and ascending series (ie., 9, tl1+-, 1243). In BrachyChilus,
Koempferin and Cautleyo al so ascending aneupl oid l ines from X= 1 2 to higher
numbers are prevalent. The X =l 7 condition in Hedychium and Cautleya is a
frequent and well-established condition. But it seems that, the X=17 condition
W uld have evolved by descending aneuploidy from a probable X= 1 8 polyploid
state, which could be a hexaploid on X=6 or tetraploid on X=9 (Ornanakumari,
1989).
*
This is also a very large tnbe comprising 22 genera, of which 6 genera were
chromosomally reported. The present study includes three genera. A11 the three
genera presented here possess X= 1 2 as their basic chromosome number-
However, the previous data (vide Table 46 ) indicate that two basic numbers exist
in this tribe such as X=l l and X=12, of which X=12 condition shows striking
predominance occurring in most of the genera either singly or in combination. It
is also quite evident that, most of the genera are polyploids based on X=12
condition. Even though polyploids are more prevalent in this tribe, it seems that
polyploidy has very little role in changing the basic chromosome number from
X=12, which is considered to be an early evolved condition in the family
Z i ngi be raceae.
Table 46 : Basic chromosome numbers found with different genera of Zingiberaceae
Genera Basic
chromosome
number
Subfamily :
Costoideae
Topinochzlm
Subfamily:
Zingi beroideae
Globba
Bosenbergiu
Bruch ich ilus
Hitchenia 21
Kaempfer ia 11,12, 13, 14
Roscoea 12,13
AIpinio 11,12
Aftamomum 12
Amomum 12
Eletturia 12,13
Phaeomaria 12
Renealm ia I l
c Polyploidy in the family Zingiberaceae
Poliploidy is a wide spread cytogenetic phenomenon found in over 3096 of
dicotyledons and 50% of monocotyledons (Love and Love, 1975). It clearly
plays a part in initiating discontinuity both within and between species, by an
increase of gene number and variety, producing radically different we1 l-adapted
genomes. Various aspects of this phenomenon, such as the mode of origin and
development, its cytogenetic behaviour and significance in speciation and
evolution have been discussed and reviewed @e Wet, 1980; Jackson and Casey,
1 980; Goldblatt, 1980; Tal, 1980; Ehredorfer, 1980; Sharma and Shma, 1984).
Polyploidy is one of the best genetical and evolutionary processes, which has
greatly contributed to speciation and the evolution of higher plants (Gottschal k,
1985). This is mainly due to the ability of polyploids to increase the chances of
fertilizab on by breaking the reproductive baniers, which permit natural selection
and establishment of species even under adverse environmental conditions.
(Stebbins, 1974). In the family Zingiberaceae the incidence of polyploidy is
fairly frequed,being prevalent in about 500/0 of the taxa spread in 16 out of the
128 cytologicdly b w n genera. Atthough this much incidence of polyploidy,
which obviously does not exceed the gross incidence in the angiosperms at large,
apparantly gives an impression that this phenomenon has played an active role in
speciation and evolution in the family.
Studes conducted on 43 taxa reveal the dominance of polyploids over diploids in
South Indian Zingibemceae. Out of these 27 taxa are polyploids at different
levels. There are 15 dploids, 8 triploids, 1 8 tetraploids and one pentaploid.
It seems that the life form and the growth habit of the plant are correlated well
with polyploidy. Growth habit is one of the factors which influences the
frequency of polyploids in angiosperms (Bquar, 1976). According to Stebbins
( 1 97 l ), the highest frequency of polyploids occurs in perennial herbs, the lowest
in annuals and the intermediate in woody plants. The origin of perennial habits
as well as accessory methods of vegetative reproduction is considered to be a
direct consequence of ployploidy (Gustafsson, 1948). This has been found to be
true of several angiosperm families. In some monocotyledonous families such as
Lill iaceae and Amarillidaceae, studied from South India (Vijayavalli, 1 986)
pol yploid species predominated in herbaceous perennials. But in Zingberaceae
which comprises of mostly of perennial herbs such a distinction as in the other
angosperm farn iles may not be we1 l evident (Ornanaliumari, 1 989). However, in
the present investigation polyploidy is more frequent in species of herbaceous
perennial genera such as Hedjchiurn. KaempJeria, Curcurno. Alpinia, Amurnurn
and Eleltaria.
Another notable feature of this family is the intraspecific polypolidy and
aneuploidy prevalent in some of its specices. Such difference in chromosome
numbers within species complexes is sometimes correlated with notable
difference in morphology and it can be considered that the phenomenon brings
about speciation. In the present study intraspecific polyploidy was observed only
in one species (Hydychium coronorium) The previous chromosome reports ( yide
Table 45) of many other species from diffierent geographical regions show that
many of them exist in two or more cytotypic forms. Many of these species
possess considerable difference in both morphologrcal and karyotype , details as in
the case of the diploid and tri ploid taxa of H coronurium. In H. coronurium the
triploids differ recognizably from the diploids with thicker and harder rhizomes,
larger leaves and pseudostern. Moreover they differ in chromosome size too.
The trends of anenuploid changes have been in different direction, chiefly
k n c h n g and some times ascending or both ascending and descending in
different plant groups. As discussed elsewhere in the dissertation it seems that
the aneuploid changes have resulted in the plybasic conditions in Hedychium
(X+, 11, 12, 13, 14, 17) and Kuempferia 0(=9, 11, 12, 13, 14) and it is rather
clear that in most cases the change was ascending from X=12 which itself was
considered to have originated from a lower primary number X=6. In the genus
Hedychium, the incidence of aneuploidy at various p1yploid level S has occurred,
both in ascendng and descending direction. In the present study aneuploidy
though resulting in different basic numbers, has almost been nil in genera like
Costus (X+), Zingiber (X=I l), Alptnia and Amomum (X=12). However, the
situation is different in many other genera of the family elsewhere in India, as
noticed by previous authors (vide Table 45).
Sub family: Costoideae
In this only the type genus Cosru.~ is available with chromosome reports and the
data show a striking instance of verq little polyploidy. Out of the 15 cytological
known species almost the entire bulk are diploids with 2n=18 except ( '.
speciosus. This species is reported for diploid, triploids, teraploids and a
h yperpl oid (2n= 1 08) including the present tetraploid taxa with 2n=36 (vide Table
45). Thus intra specific polyploidy is well established in this species. Meiotic
behavior revealing the absence of quadrivalents in the tetraploids (Ramachandran
1969; Omanakumari, 1989) and nne occurrence of trivalent in tnploids
(Chattopadhayay and Sharma, 1981) suggest the possible allopolyploid nature
and the occurrence of structural alterations of chromosomes in the evolution of
this species. Anyhow, one cannot neglect the idea that polyploidy has played
only a very little role in the evolution of the genus Costu~.
Tribe: Globbeae
The present study includes only two species of the type genus Globba viz., G.
ophioglosso (2n=22) and G. cernua ( 2 ~ 4 8 ) . F m the previous reports it is clear
that in the genus Globba there are two different lines of evolution in basic
chromosome number based on X= l l and X=12. In most cases X=12 condition is
predominant. Out of the thiry three cytologcally known taxa in the genus
Globba four were studied from India by various authors (Chakravorti, 1948;
Sharma and Bhattacharya, 1 959; Mahanty, 1963; Ramachandran, 1969;
Omanakumari, 1989)' The remaining taxa were reported from Malayan
peninsula and Europe (Chong and Siew ngo 1972; Mahanty 1970). All the alien
species are polyploids at different levels (tetra and hexaploids). Shama and
Bhattacharya ( l 959) and Mahanty (1963) support the X=l2 line as the truly early
developed condition in this genus. After the meiotic analysis of the Malayan
species Chong and Siew-ngo (1972) suggested that most of the taxa with 2n=48
are auto tnploids based on X= 1 6 which itself arose themselves from the original
basic number X=8. They have described allotetrsploid G. cernzua with 2 ~ 3 2 and
hexaploids with 2n-48 based on X=8. But a critical study of their work threw
doubt upon the validity of this hypothesis, as the chromosomes in the genus have
been found so small and uniform in appearance. Hence its importance as
evidence to support the hypothesis would seen dubious. G cernua seems to be
an auto tetrapolid based on X= 12.
Tribe: Zingi bereae
In contrast to the other tribes this rnonogeneric tribe shows only few stray
instances of polyploidy among the members studied In the present study five
species are worked out and all are diploids with a somatic chromosome number
of 2n = 22. From the previous literature (vide Table 45) it can be seen that only
Zingiber mioga (2n = 55) is pentaploid taxa based on X = 1 1 . Kihara et al.
(1 93 1) had repted a different count, 2n = 24 for Z oflcinale and this could be
possibly an aneuploid cytotype. Thus it is very well clear that polyploidy has
very little role in he evolution of genus Zingiber.
Tribe: Hedychiese
This tribe was found to attain very high degrees of polyloidy in many members
and out of the eighteen taxa studied tenexhibit polyploidization. Most of these
members are resorted to vegetative means of reproduction, an adaptive feature
exhibited by p01 yploidy.
Polyploidy has played a substantial role in the evolution and the speciation of the
genus Cwmma. Out of the eleven species studied six species were triploids with
2x143 based on X=2 1. These triploid species are not seed setting. From the
karyomorphological analysis it seems that these triploids are possibly auto-
pal-yploids. The chromosomes are very small and more or less similar in size.
Kqomorphological details have been discussed elsewhere in the dissertation.
The genus Curcurno exhibits a high basic number, which is considered as a
secondary one. Such a high basic number might have originated from lower
number such as X+ and X=12, which is rather common in many other related
genera. Rarnachandran ( 1 969) supported the dibasic amphidiploid origin of the
genus from two forms which have a lower basic number of X=9 and X=] 2 or by
secondary tri ploidy.
As mentioned elsewhere in the dissertation the incidence of polyploidy is very
much striking in the genus He4chium. In H. coronaritrm two cytotypes, one
diploid (2n=34) and the other triploid (2n=51) were found. This genus also
exhibits a high basic chromosome number with X= 1 7, which is considered to be
secondary in orign. The rneiotic behavior of the species H. coronarium very
often shows its auto-plolyploid nature (Omanakmari, 1989).
Karyomorphologrcal details also agree with this situation. For this species only
diploid and triploid species are so far known from this region. Thus intra specific
polyploidy is very well established in this species. It is likely that the triploid
forms must have arisen naturally from the union of reduced and unreduced
gametes of the diploid taxon. The triploid nature is also found in Hedychium
jlavescem .
The genus Koempferiu with three species studied showed a polyploid series
(diploid, tatraploid and pentaploid) based on X= l l . The exotic species K.
pulchra with 2n=22 is a diploid. While K. rot& with 2n=44 is a tetraploid K.
galanga 2n=54 is an aneuploid w'th a pentaploid status. The metiotic studies
(Ornanakuman' 1 989) and the karyomorphological analysis (vide Table 45)
indicate that the species K. rotunda is probably an auto-tetraploid. From the
t
earlier reports (vide Table 45) it is clear that the various species of this genus fall
under different basic numbers with X=9, 1 1, 12, 1 3 and 14 of which X= l l
predominates in many species. It is possible that the higher basic numbers
X= 1 2,13 and 14 are the derived conditions from X= l 1 by ascending aneuploidy.
Tribe: A l pin ieae
The incidence of ployploidy is very much stnking in this tribe. All the members
studied showed a tetraploid condition, with the basic number X- 12. The various
species of Alpinin, Arnornurn and Elrtluriu showed 2n-48 which support thc basic
number X ~ 1 2 . A perusal of the literature makes it clear that another line of
evolution with X=l l condition in a few members of this tibe which might have
arisen from X= 1 2 condition by descending aneuploidy.
The genus Al'pinia with al l the eight species studied shows 2n=48 based on X=l 2.
All the species are thus tetraploids. This agrees with the previous reports (vide
Table ). Meiosisisregularwith24bivalenceinA.calcarutaandA.galonga
(Omanakumari, 1989). This results in normal anaphase separation and high
degree of pollen fertility and thus good seed setting. This clearly indicates their
possible allopolyploid nature.
The genus Amornum exhibits the tetraploid condition with 2n=48 based on X=12.
All the four species studied with their regular meiosis and seed setting indicates
their possible allotetraploid nature. The karyomorphologicd analysis of species
also give support for the view.
The three cultivars of Elettana mdmomum studied are also tetraploids with
2n-48 based on X=12. Chakravorti (1 948a), Chmdrasekbar and Kumar (1986)
had reported 2 1 ~ 5 0 and 2 s 5 2 conditions in this species. This could be a
tetraploid with X 4 3 condition (in 2n=52 taxa) or possibly an aneuploid with
X= 12. Meiotic studies by Umanakumari (1 989) prove that this species is an
allotetraploid
d. Ksryomorphometrlcal analysis
In angiospems the species of several families both dicotyledons and
monocotyledons are found to exhibit a direct relationship between their
phylogeny and the chromosome constitution The chromosomes being the
carriers of heredity, both structural and numerical changes in them can influence
the genetic evolutionary process at work than do any other type In
angiospems,the species of several families both dicotyledons and of changes. A
detailed information regarding the chromosome arch tect ure in higher plants may
help us to understand their systematic relationship and to trace the trend and
direction of their evolution (Love and Love, 1975).
Some of the major karyotypic characteristics of considerable evolutionary and
taxonomic significance are ( l ) difference in the absolute chromosome size (2)
difference in the position of centromere (3) difference in total chromatin length
(4) difference in karyotype formula and ( 5) difference in number as we1 l as in the
position of satellites.
Since, most of the members of Zingiberaceae possess very small chromosomes
the detailed karyomorphological study by conventional methods is not easy. So
far only a few plant species were reported for their karyomorphyology. An
image analysis system provides a better opporturuty for studying the various
chromosomal parameters. This method has only been used recently in plant
cytology and chromosome studies (Fukui, 1985; 1 9 86). It appears to be a. very
powerful tool can 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
ha became possible to obtain various kinds of qutitative data on the chromosome
images. The length of the chromosomes and/or arms and the ratio between the
short and long arm values have represented almost all the numerical information
available on the chromosomes for the past decades. It has become however
possible to obtain not only one dimensional data of the chromosomes such as
length, but also two and three dismentional data of area and volume within a
limited time by using image analysis system. Aiso it is possible to make an
accurate pairing by PIA ratio mainly in those cases where chromosome sizes are
really small. In addition to this imaging techniques provide pseudo colouration
which is a very usehl tool for identifying the primary and secondary
constrictions of small chromosomes very clearly. Idiograms can be prepared
semiautomaticaliy and by using computer devices. By the treditional 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 usefbl information
for chromosome analysis and may be helpful as an essential tool in chromosome
research (Fukui, 1988; Fukui and Kakeda, 1990; Iijima et al. 1990 Fukui and
l ij i ma, 1 990; Kamisugi et (11. 1993)
The general feature noted in the family ~in~ibeniceae is the wide range of
chromosorns with very small sized chromosomes in most of the species.
However, the chromosome complements in the various members differ in minute
karyotypical details (vide Table 47). With regard to the gross morphology,
chromosomes are nearly sub median to nearly median in nature. The
chromosome range from 2.68 to 0.24pm in length. The chromosomes with
secondary constrictions ranges fiom two to fourteen in number. The average
chromosome length (ACL) varies from 0.39 pm to 1.69 pm. The total chromatin
length (TCL) shows a very wide variation with 16.21 pm being the minimum and
88.56 pm being the maximum value. The disparity index values ranges between
27.9 to 64.7 the coefficient of variation ranges from 12.76 to 41.27 and total
chromatin index UP!) ranges from 35.5 to 44.65.
These variations found in the karyotypic parameters
Zingiberaceae members are characterized by S ymmetri
karyotypes. An increase in the range of chromoso
increase of sub meytacentrics at the expense of ma
our increase in the coefficient of variation leading to assymmetry (Stebbins,
1998). Thus karyomorphological studies are of considerable importance in order
to throw light on the telationship among taxa of flowering plants (Iwatsubo and
Nanrhashi, 1991).
Subfamily: Castoideae
Genus: Costus
Two species of the genus represented in the present study, Comprising with other
genera Costus exhibits longer chromosomes. As regards the chromosome with
secondary constrictions C speciosm possesses four chromosomes, while C
mdor/ieunm has only two. The karyotype in both species is somewhat
symmetrical and thus shows primitiveness. In C. maIortieanu.~ there is no 'C'
type chromosomes. Also the higher values for average chromosome length,
Chromatin length of haploid complement and range of chromosome length are its
primitive features. However, comparatively higher variation coefficient and lower
TF9h are advanced featms. In C'. specioL~u.s, which i s a tetmploid plant, the
average chromosome length is smaller than C: mulortieanus. The range of
chromosome length is also lesser than C. maIort~cunus. Also a high TF% with
low variation coefficient show its advanced features than C. malortieunus.
Subfamily: Zingiberoideae
Tn i : Globbeee
Genus: Globba
Genus Globbo species in this study, the displayed G. ophioglosso and tetraploid
G. cernua. So far no report is available on the karyomorphology of ths genus.
Karyomorphologicall y this genus stands in a we1 l-advanced stage. According to
Sharma (1984) the genus Globba is considered to be more advanced than Costus,
Zmgiber and Kuempfiia. Chromosomes are moderately small with syrn metri cal
karyotype. In both cases the absence of "C'type chromosome is a speciality. In
G. ophioglossa their two pin of chromosome with secondary constrictions.
While in G. cernua there are three pairs of satelite chromosomes. From the table
it is clear that the higher values for average chromosome length, range of
chromosome length, chromatin length of kaploid complement along with a low
variation coefficient and high TF% reveal the Orimitiveness of G. cernun over G.
Tribe: Ziagi bereae
Genus: Zingiber
The genus Zingiber with 6 species studied for their detailed karyomorphology .
The genus is characterized by a general homogeneity observed in the minute
details of karyotype. Eventhough all the species are of 2n=22 the number of
chromosomes with secondary constrictions are only four in Z ofjiciole. The
. di~erencesobservedinthekaryotypefomula(KF),averagechromosomelength
and chromatin length of haploid complement (CLHC) among diKerent tarn might
have been probably due to minute structural alterations of chromosomes (vide
Table 47 Fig 46 ). Speciation depends more on chromosomal rearrangements and
mutation of individual genes, than on changes in the total amount of genetic
content (Stebbins, 1959).
The high value of average chromosome length and chromatin length of basic
complement shown by Z naesanum and Z cernum (vide Table 47) probably
show their primitiveness, whereas the lower values for these parameters found in
Z oficinale and Z zerumbet denotes their 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 Qsparity index (Dl) value found in Z cernum followed
by Z rosewn correspond to the heterogeneous assemblage of chromosomes in
these taxa (vide Table 47, Fig. 46). But comparatively lower values of D1 found
in other members point towards the general homogenity found in various species
of Zingiber. Normally a low disparity index value corresponds to the
homogeneity of chromosomes in most of the higher as well as lower plants
(Mohanty et al. 1991) The high variation coeficient exhibited by Z cernurn
undoubtedly confirms its primitiveness.
The Laryotype is seems to be more or less symmetrical in all the investigated
species. A symmetric karyotype is considered to be a primitive one (Stebbins,
1959). Eventhough the karyotypic features of the six species 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 intrakaryotyjx size
differences (vide Fig. 46). It is remarkable that only minute structural a1terations
exhibited by the genus. In the light of the chromosome data so far available on
the genus Zingiber and the karyomorphological information of six species
reported here, it appears that both numerical and structural alterations in
chromosomes have not played any major role in speciation and evolution of this
genus.
Tribe: Hedyehiae
Genus Curcuma
Karyomorphology of l l species of this genus have been studied. Detailed
karyomorphology studies of this genus were not tried so far, possibly due to the
very small size of chromosomes. The hfferent species of Curcuma are
exceedingly variable both in chromosome numbers and in morphometric
characters of their karyotyj~s. This variability of the genus evidently reflects an
important side of its evolution, apart from its morphological differentiation on
which classification is based. The number of chromosomes with secondary
constrictions varies in different species. In various diploid taxa they are 1-2
pairs in number. The triploid species also possess 1-3 y r s of satellite
chromosomes (vide Table-47). The various species are characterized by their
homogeneous symmetrical karyotypes with lesser range of chromatin length, a
primitive condition. However, the higher number of chromosomes with small
size and lesser values of average chromosome length, chromatin length of basic
complement, disparity index and TTh (vide Table-47) advocates their evolved
nature. Among the karyotype of disploid species C. horihu exhibits a higher
number of short sized chromosomes (Type C) , comparatively low values of
*
average chromosome length, total chromatin length as well as TP! along with a
high value of variation coefficient. Thus C. haritha seems to be more advanced
than C. amado. These two species stand in extreme poles, as regards their
chromosome evolution. Amidst the six triploid species studied the high values of
total chromatin length, average chromosome length and TF% exhibited by C
raktahnta put the species in a lower position in the trend evolution. Whereas the
species C. ~edoclria seems to be more advanced karyomorphologically, among
the triploid species. This species is characterized with lower range of
chromosome length and higher number of type C. chromosomes. In general,
karyomorphologically the genus C u r c m is more advanced one comparing with
other generas of family Zingiberaceae. The morphological character of these
plants is also in well agreement with this statement. Besides the primitive
characters such as symmetric karyotypes with low range of chromosome length.
they exhibit advanced features like low values for average chromosome length,
chromatin length of basic complement and TF%. So the exact evolutionary status
of the genus appears to be enigmatic (Table-47, Fig. 47 &48).
Reduction in chromosome size is apparently a consequence of polyploidy, since
it is an adaptation to a decrease in size of the cell or to an increase in number of
chromosomes @arlington, 1956). The very high number coupled with small size
of chromosomes may indicate a higher evolutionary status (Sharma, 1 984) of this
genus. Also the triploids are considered to be more advanced over diploids, in
many cases.
The genus exhibits difference as regard the chromosome morphology. The
hyotype in various species studied is found to be more or less homogenous and
symmetric. The chromosomes are found to be very small in size. Chromosomes
with secondary constrictions are varying from 1 to 3 as seen in different taxa
(vide Table-47 Fig. 49). The differences observed in the karyotype formula (KP)
average chromosome length (ACL) and chromatin length of haploid complement
(CLHC) among different taxa might have been probably due to structural
alterations of chro~osomes. Speciation depends more on chromosomal
rearrangements and mutation of individuals genes than on changes in the total
amount of genetic content (Stebbins, 1959). The species H. fkrvescense is
characterized by high values for the major karyomorphometrical parameters and a
comparatively low variation coefficient. Thus these observations are in advocacy
with the primitive status of this species. Another notable feature of this species is
the absence of "C' type chromosomes. Except the high TF% and low variation
coefficient all other karyomorphometncal characters show the species H.
Spicotm var occumimtum is more evolved than other members. In H.
coronarium if we are critically analysing the various kar);ornorphological data
and the respective idiograrns of the diploid and triploid species, it becomes clear
that the polyploid species is an autotriploid (occurrence of three similar
chromosomes in each set). Meiotic studies (Omanakwnari, 1989) also proved
this statement. Moreover from the data it is clear that in the triploid species of H.
cornnorrum even though there is an increase in chromosome number, no
appreciable decrease or increase in chromosome length or size comparing with
the other genera in karyomorphological details geneus Heafvchicum stands a
ahead in the trend of evolution and it is more evolved one (Sharma, 1984).
Karyomorphologcally this is a very inadequately investigated genus with the
improved techniques and the data in the three species reported here are too
insufficient to make a consideration on the nature and extent of structural
alterations in chromosomes that have contributed to speciation and evolution in
the genus. The data (vide Tables-47, Fig. 50) indicate that, all the three species
have more or less symmetrical karyotype, which is considered to be primitive.
On a close examination of the karyotype of the three speciese it reveals that
karyotype asymmetry is progressively greater among the higher polyploids, as
they show higher frequencies of submedian chromosomes. The change of
centromere from median to submedian ps i tion, and increasing difference in size
between chromosomes of a complement are two basic factors in the evolution of
chromosome morph01 ogy. Similarly intra karyotypic size difference of
chromosomes which is brought about by differentia) deletion of segments of
individual chromosomes as well as through occurrence and establishment of
unequal translocations between nonhomologous chromosomes (Mathew and
Thomas, 1974) is also seen to be greater among the polyploids. The evidence
thus indicate that the above factors which lead to karyotype specialization also
have played some role, along with polyploidy, in the evolution of the genus
Kaempferia. In K. gnlunga even though the karyotype shows some trends
towards asymmetry the higher values of TF%, disparity index average
chromosome length, chromatin length of basic complement reveal its trend
towards primitiveness. In K. pulchra, the lesser values for all these parameters
show its rather evolved nature. Such type of variations is important from the
evolutionary viewpoint. When comparing with other generas of the family
Zingiberaceae genus Koempjkria represents a primitive line of evolution
(Sharma, 1984). Lung chromosomes with symmetrical karyotypes arc further
evidence of primitiveness (Sharma, 1984) of this g n u s .
Tribe; Alpinieae
Eight species of this genus were studied karyornorpholog~cally. So far no
authentic detailed reports available on the karyomorphology of this genus.
Karyotype in various species seems to be formed of very small chromosomes. All
of them are in tetraploid condition. Reduction in chromosome size is ap&rently
a consequence of polyploidy since it is an adaptation to a decrease in size of the
cell or to an increase in number of chromosomes (Dartington, 1956). The
karyotypes of various species are found to be more or less symmetrical and
homogenous. Even though the lesser range of chromosome length and lesser
variation coefficient value denotes, primitiveness all the other characters
including a low TF?h (vide Table.47) shows the probable advanced status of A.
smithiae. The centromeric index is a karyoiype characteristic independent fiom
chromosome numbers. A low means ceatromeric index value suggests a highly
advanced karyotype and a high value represents a primitive karyotype (Vasil eva
et al. 1985). The lesser range of chromosome length and low variation
coefficient with higher values of average chromosome length, chromatin length
of haploid complerneqt, disparity index and TPh (vide Table 47, fig. 51 & 52)
pointing towards the more primitive condition of the species A. gaIanga. The
chromosome pairs with secondary constrictions are varying fiom two to three in
various taxa. The karyomorpholog cal differences found among these taxa fully
justify that speciation and evolution in this genus has been principally effected by
mi nute structural alterations.
The different species of the genus Amomum are characterized by the more or less
homogenous symmetric karyotypes (Table47, Fig. 53). Despite the constancy in
chromosome number, the kmyotypic differences among the species were quite
distinct. Karyomorphometrical surveys on the genus are rare, probably due to the
small size of the chromosomes. The karyotype of the A. hypolrucum
characterized by a higher variation coefficient and a lower value for chromat~n
length of basic complement. These two parameters are sufficient to place it
among the advanced members of Amomum along with comparatively lesser
values for average chromosome length, total chromatin length and TF%. The
high disparity index value in A. hypoleucunl shows it's more or less
heterogeneous karyotype, showing some trends towards karyotype asymmetry.
Whereas the high values for all major karyotype parameters along ~ i t h a low
variation coefficient confirms the primitive status of A. cunnicarpurn.
Eventhough a general karyotype symmetry is noticed in various species with
small chromosomes, it is evident that minute structural alterations are prominent
in the karyotypic changes and thus causing he evolution of t h~s genus.
Elettaria
Concrete conclusions cannot be drawn From the karyomorphological features
(Table 47, Fig. 54) exhibited by the closely related genetic strains of E.
cmdamomurn (cultivar Malabar, Mysore, and Vuzhukka). So it seems likely that
%represent three closely related lines as has been found in the evolution of
their morphological features. However, at the infra specific level E
cardamomurn CV. Malabar is characterized by high values for major
karyomorphometrical -parameters such as total chromatin length, average
chromosome length chromatin length, of haploid complement except F h and
variation coefficient and are considered as primitive characters. Whereas the
Cultivartor Mysore stands wide apart from this condition. From the
karyomorphological analysis it is somewhat clear that the cultivar Vazhukka may
be a natural hybrid in between these two. Karyotype is found to be homogenous
and symmetric invariably in all the cultivars.
e . Cytological evolution in Zingiberaceae
From the cytological observations made in the present investigation it can be
concluded that speciation and evolution have been possible as a result of increase
in variability through changes in the base numbers. as well as numerical and
structural changes in chromosome number. 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
chromosome numbers observed in many genera (vide Fig. 44) in the present
investigation mark a significant role that aneuploidy and ployploidy have played
in the evolution of various taxa at the generic and species level. It also appears
that various kinds of abenations have played a vital role i n the evolutionary
diversification of the family. The mitotic and meiotic irregularities qight have
lead to structural and numerical variations in the chromosomes of a species.
Individuals with same chromosome number but with difference in
karyomorphological details reflect the ongoing evolutionary process at micro
level.
It seems possible that in South Indian Zingiberaceae, 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 sunive in nature
because they express various degrees of weakness and chromosomal aberrations
leading to genetic sterility. However, some individuals carrying the changed
chromosomal constitutions are well within their range of tolerance. This was
confirmed by the occurrence of normal meiosis and seed setting in some
polyploids. Moreover, most of the taxa belonging to this family have efficient
means of vegetative pr%pagation. This ensures the survival of these genetically
altered !ypes which otherwise would have faced extinction 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. h that respect meiotic accidents may proved to be more
useful than mitotic aberrations from the evolutionary point of view, since the
meiotic abnormalities are hereditary and likely to multiply and establish in a
population.
The order Zingiberales, or in a wider sense Scitamineae including Musace.de,
Zingiberaceae, Cannaceae and Marantaceae has been regarded to have originated
from Bromeliales (Shanna, 1984; Huchinson, 1959). Takhtajan (1969) however,
placed it in the super order Lilionieae of Lilideae along with 1,iliales and
Bromellales. Zingiberales is a distinct assemblage with multiples of X = S. 9. l I
and 12 chromosomes.
l 09
The very high chromosome number coupled with small size, may indicate a
higher evolutionary status of this family. On the basis of cytological data, taken
in conjunction with progress of complexity in flora biology, Costus, Zingiher and
Kaempferia may represent a primitive line of evolution as compared to Globha,
Cautleyo etc. It is not unlikely that the group is directly derived from the
Musaceae or its immediate ancestor (Sharma, 1984). Comparing with other
genera Costus. Zingiber and Kaempferia possess large sized chromosomes, which
are few in numbers. But the chromosomes of Globba, Hedychium, Curcumo etc.
are small sized and more in number. The relationship of chromosome size to
increase in chromosome number or polyploidy has been pointed out in several
genera. (Sharma 1984). The diminution in size often noted along with this
process has been attributed either to elimination of heterochromatin or
differential coiling of chromosome segments. Similarly there are genera, where
no such diminution is recorded, e.g., Kaempfrio. Possibly, an answer to this
interesting behavior may have to await a complete analysis of the extent of
repetitive DNA or redundant DNA occurring in such taxa at the diploid level.
After a critical study of the morphology of chromosomes as well as their meiotic
behavior in different species of the genera like Alpinia, Phaeomeria and
Hedychryum Chakravorti (1948) suggested that they are neither polyploids nor
arnphidipioids. According to him the increase in number of chromosomes of
these genera has been brought about by hgmentation of some chromosomes of
other closely related genera at the region of supernumerary constrictions which
are so prevalent in the chromosomes of the species of Zingiber and Kaempfrio.
Thus the latter two having longer chromosomes but smaller in number with
unusually large nwnber of secondary constrictions have very likely contributed to
the origin of the different species of Alpinia, (:urcuma. Hedychrum and others, all
of which have comparatively high number of chromosomes which are small in
size and with fewer supernumerary constrictions. However, this suggestion was
ruled out by Sharma and Bhattacharya (1959), because no such accentric
fragments can acquire a centromere of their own In the present investigation
also no such evidence of fragmentation at secondary constriction region was
evident.
I l0 Prepared by BeeHive Digital Concepts Cochin for Mahatma Gandhi University Kottayam
The very high chromosome number coupled with small size, may indicate a
higher evolutionary status of this family. On the basis of cytological data, taken
in conjunction with progress of complexity in flora biology, Costus. Zingrber and
Kaempferia may represent a primitive line of evolution as compared to Glohba,
Caufleya etc. It is not unlikely that the group is directly derived from the
Musaceae or its immediate ancestor (Sharma, 1984). Comparing with other
genera Costus. Zrngiber and Kaempferra possess large sized chromosomes, which
are few in numbers. But the chromosomes of Globba, Hedychium, C'urcuma etc.
are small sized and more in number. The relationship of chromosome size to
increase in chromosome number or polyploidy has been pointed out in several
genera. (Sharma 1984). The diminution in size often noted along with this
process has been attributed either to elimination of heterochromahn or
differential coiling of chromosome segments. Similarly there are genera, where
no such diminution is recorded, e.g., Kaempferia. Possibly, an answer to this
interesting behavior may have to await a complete analysis of the extent of
repetitive DNA or redundant DNA occurring in such taxa at the diploid level.
After a critical study of the morphology of chromosomes as well as their meiotic
behavior in different species of the genera like Alpinia, Phaeomeria and
He&chij.um Chakravorti (1948) suggested that they are neither polyploids nor
amphidiploids. According to him the increase in number of chromosomes of
these genera has been brought about by fragmentation of some chromosomes of
other closely related genera at the region of supernumerary constrictions which
are so prevalent in the chromosomes of the species of Zingiber and Kaempfer~a.
Thus the latter two having longer chromosomes but smaller in number with
unusually large number of secondary constrictions have very likely contributed to
the origin of the different species of Alpinia, Clurcuma, He+chium and others, all
of which have comparatively high number of chromosomes which are small in
sue and with fewer supernumerary constrictions. However, this suggestion was
ruled out by Sharma and Bhattacharya (1959), because no such accentric
fragments can acquire a centromere of their own. In the present investigation
also no such evidence of fragmentation at secondary constrict~on region was
evident.
\\o
Considering the evolution of basic chromosome number most of the genera of
tribe Hedychieae are plybasic, whereas, in tribe Alpinieae the monobasic
condition (X=12) predominates in many genera (vide Table-46). So in this
respect the tribe Hedychieae is more advanced than tribe Alpinieae. Moreover,
the various morphological charactem also support this view. The reduction in
size of rhizome (e.g. Bmenbergra and K~emp~r r i a ), the non-frondose lea6
shoot, the reduction in number of locules of ovary from three to one and ovules
from many to one (e.g. Bosenbergiu and Purucouf~eyu) are regarded as advanced
characters. But in tribe Alpinieae the large rhizomes with frondose leafy shoot
and trilocular ovary with many ovules are considered to be primitive (Sabu,
19911
Ployploidy is said to have ancient origin if a taxon includes only polyploids or
very few diploids, and relatively of the recent origin of deploids are more
frequent than polyploids or both are in almost equal proportion (Stebbins, 1950).
On this context among the members investigated, based on the previous and
present studies, in the various genera such as Cosfus, Zingiber and Kaemp+riu
ployploidy assumed to have a recent origin. Whereas in many species of tribe
Alpinieae including AIpinia and Amomum, polyploidy seem to be of ancient
origin. However an effective conclusion cannot be reached in these genem since
many of the species are still to be investigated cytologically. There are still
enormous gaps in our knowledge as regards the cytologxal evolution of
Zingiberaceae and much still remains to be done before a major cytotaxonomic
review may be attempted. The X=12 condition is predominant among the
members being present in eight of the genera (vide Table-46) followed by X=I I .
It is considered that the X=l2 condition itself could be of polyploid origin from a
possible X=6 primary condition.
Table: 47 Summary of the Karyomorphometerial analysis on the forty three members of Zingiberaceae investigated
SI.No. Name of the taxa 2n PL KF CPSC
G. ophioglossa
Zingrber cemuum
Z. neesanum
Z. oflcinale
Z. purpureum
Z. roseurn
Z zerumber
Curctrma aerugenosa
C amada
(7. aroman'ca
c'. caesia
c'. comosa
(I dectpiens
(: harrtha
C . longa
C. malabarica
C: rakracanta
C. zedoaria
He&chium coronarium (2x)
H. coronarium (3x)
RCL ACL TCL CL.HC
(in pm) (in pm) (in pm) (in pm)
'Total
volume
H. I ~~a~~cscr ns
H. spicurr~m var.acuminaltrm
Karmpfiria galanga
K, prtlchra
K. rorrmnda
Alpinia calcarala
A. galanga
A. malaccen~i.~
A. nrgra
A. pupttroa
A. smi~hiae
A. zcntmhe/
A. zrrumher var. varigara
Amomzrm cannicarpum
A . hypoleucum
A. muricalum
A. plerocarpum
1:'tettaria cardamomum cv. Malabar
K. cardamomum cl,. Mysore
1;. cardumomirm clv. Vazhukka
formula, CP-Chromosome pairs with secondaly constriction. RCL-Range of chromosome lenglh. ACL-Average chromosome length.
chromatin length, CLHC- Chromatin length of haploid complement, DI-Disparity index. VC-Variation co-emc~ent.
TF%-Total forme osrcentaaa.
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Kaempferia galanga K. pulchra K. rotunda
Fig. 50 Comparison of major karyotypic parameters in
Kaemp feria
Alpinia calcarata A. galanga A. malaccensis A. nigra
Fig. 51 Comparison of major karyotypic parameters in Alpinia
DTCL
001
VC
OTF%
Etettaria E. cardamomum E. cardamomum
cardamomum CV. Mysore CV. Vazhukka
CV. Malabar
Fig. 54 Comparison of major karyotypic
mTcL
B DI
nvc
OTF%
parameters in Elettaria
SUMMARY
Forty three taxa representing a total of nine genera of Zingiberaceae found in
South India are studied for their detailed karyomorphology. The chromosome
spectrum i n South Indian Zingiberaceae ranges from 2n=18 to 2n=63 with
majority of the species concentrated in the number 2n=48 followed by 2n=63 and
2n=22. 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 variations. Ths wide range of chromosome
numbers may be due to the difference in number of chromosomal biotypes
belonging to different groups. From the previous literature and the present
investigation it is clear that many genera like Costus, Globba, Curcurno.
He$ychium Kaernpfria etc. exhibit inter, intra and infra specific variations
among chromosome numbers. The presences of such widely different series of
chromosome numbers in the species of even the same genus and in genera placed
under different tribes and, sub tribes, indicate that the different chromosome
numbers may be derived ones from the other.
The basic ch~mosome numbers are found to be varied in South Indian
Zingiberaceae. This &eat variability in the number of chromosomes at the basic
level could possibly be the result of aneuploidy at generic level. Out of the nine
genera investigated three (Globba, Hedychium and Kuempferia) have been found
to be polybasic. Whereas, Costus, Curcuma and Alpinia are dibasic. Zingiber,
Amomum and Elettaria are more or less mono basic. Both primary as well as
secondary base numbers are found to be involved in the evolution of forty three
tawa investigated. The family is a distinct assemblage with multiples of .X = 8, 9,
11 and 12 chromosomes. The X = 12 condition is predominant among most of
the members followed by X = 11. The genera Costus XI= 9, Globha
(X1=11,12), Zingiher (X1=1 I), Kaernpfer;~ (X1=1 l), Alpinia (XI=] 2), Amomum
(Xl-12) and 1~;le~roriu (XI-12) exhibit primary base numbers. While the basic
numbers of the species of Curcurno (X2=21) and Het@chrurn (X2 = 17) are
secondary in origin. Thus it is likely that protopolgloidy plays an active role i n
the evolution of these species.
The family Zingiberaceae is characterized by a relatively high frequency of
polyploidy. Studies conducted on forty three taxa reveal the dominance of
polyploids (62-79%) over diploids in South Indian Zingiberaceae. In the present
investigation polyploidy is more frequent in species of herbaceous perennial
genera such as Hedychium, Kaempferia, Curcuma, Alpiniu, Amomum and
Elertaria. Among the polyploids the majority are triploids but some tetra and
pentaploids are also present. The role of both polyploidy and aneuploidy in the
mechanism of speciation is obvious in South Indian Zingibereaceae.
The computer aided image analysis system provides a better opportunity for
studying the various chromosomal parameters especially when the chromosome
size is very small. Since most of the members of Zingiberaceae possess very
small chromosomes the detailed karyomorphological study by conventional
methods is not so easy. The method of using conventional numerical parameters
such as length and arm ratio is insufficient to distinguish the chromosomes of
many members of Zingiberaceae. 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.Idiograrns cart be prepared semiautomatically by using computer devices.
Hence this method would provide useful information for various
karyomorphological analysis.
The general feature noted in the family Zingiberaceae is the wide range of
chromosome number with graded symmetrical karyotypes. However, the
chromosome complements in the members differ in minute karyotypeic details.
With regard to gross morphology, chromosomes are nearly median in nature.
The chromosomes were found to range from 2.81rn to 0.3rn. The total chromatin
length shows a very wide variation with 16.53rn being the minimum and 88.56m
being the maximum values. The disparity index values were found to range
between 27.9 to 6473 and total centromeric index value (TF?6) from 19.8 to
44.65. The coeficient of variation ranges sfrom12.76 to 41.64. Thus the various
micromorphological details of the karyotype like, differences in ahsolute
chromosome size, Qfferences in position of centromere, differences in total
chromatin length, differences in karyotype formula and differences in the number
as well as position of the satellites, vary from cytotype to cytotype. These
variations found in the karyotype parameters suggest that South Indian
Zingiberaceae members are characterized by symmetrical to slightly
asymmetrical karyotypes. The karyomorphologivcal 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 chromosomes found in many genera mark the significant
role that both aneuploidy and polyploidy have played in evolution of various taxa
of the family at 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 abenations have played a major
role in the evolutionary diversification of the family. Individuals with same
chromosome number but with differences in karyomorphological details reflect
the on going evolutionary processes at micro-level. It has also been found that in
South Indian Zingiberaceae, Robertsonian changes or mutations might have also
played a major role in the evolution of karyotype.

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