stuoy of the 'g' banding patterns of the chromosomes … · 2018. 1. 4. · in the...
TRANSCRIPT
STUOY OF THE 'G' BANDING PATTERNS OF THE CHROMOSOMES
OF ALBINORAT (Rattus-norvegicus) AND Funumbulus
pennantj wroughton
Dissertation Submitted in part ial fulfilment of the
requirement for the Degree of Master of Philosophy
ot
The Aligarh Muslim University Aligarh
By
ANJUM ARA
DEPARTMENT OF ZOOLOGY ALIGARH MUSLIM UNIVERSITY, ALIGARH
February, 1982
< TT
A. ::SH
HWil
/J
>
CHEi::::n--3002
DEmCATED TO MY PARENTg
IN
GRATITUDE
I certify that 'Study o^ G Banding patterns
of the Chromosomes of albino rat (Rattus norvegicus)
and Funumbulus pennant! wroughton', is the
original work of Miss Anjura Ara and is suitable for
partial fulfilment of the requirements for the
degree of Master of Philosophy in Zoology, of the
Aligarh Muslim University, Aligarh, The work has
been done under my supervisJron,
Usman M. Adhami Ph.D. (Notre Dame) Reader, Department of Zoology, Aligarh Muslim University, Aligarh,
n
Aolc&Qv2.94geaes'l8 • • • «• •
TabSbO of Oeatttats • • • • * •
I4.at of Sabl#» ••• •••
Mat of iX^iixmm ««• •** >
XntsodUiOtloii ••• ••* 1
Hftt«simJL8 and H«tllo4B •*• 10
IPiiotogimflia aM HiStOil^MiS • « • 05
^vsmsaiy ••• • • • 105
KP^^^^^J^JiJJ^^^^^^^^^^^^^
mko h&9 «mf«irfl0*d tliis wosk* Mis iralimbl« mum^stlouti,
e r i t io s l a4iri09 and eoim^mxkt mmQwm^^m&at imra of
gseat laporlanee tos the ec^apXatlom of tMa
astttcfuXly a^tooirX«dg«aioiit i s «ad« to txof • lla«ab
Eaoan JOia t H' ad of tli« BtpaxTteiiaiit # < 3 jpjpofildlii
iiae«00a£^ fa0l]Ul.titts«
I as aleo vaa? thaoJsfitX to ay Xab* aatae Mr* MMd
&34» Hss* Baepi fandon* aM Mlaa imadiat faae^n MaXllE,
for thalj; @ooi «£atlo&«
airida2!« thaiil^s aa?a dua to Bs« Vaatwa Ataaad faifidit
Hap* Afaal A»SlddiqixL and Hiss MB&Q iaidul^Ab foi? th«l£
half dmein^ the oouxaa of prase&t atod^*
if
( -1 )
!• TABLE lA - lorphoaetry of ttoe atLraiaoa<»ii€is of
yimaiiife»Xiifl geimimtl (Wroiaghton ) (Hal® )
Diploid Ho* ( an ) « §4*
2* 1!/UJLE I IA > Mo£ph(»et2ry of th« ObrcHiosoaee of fumimtottlm^
pcmoaaia^ Wrom^litoa ( Male ) showing uxrea ioduoftd
3» tkam Tllk • Hosphoi^tir^r of the QhxomoB&mea of gmawabulao
peaaaaati tfrott^ten ( feiaale ) showing U£ea iMuoed
Q biucids*
4* TkdhB I f 4 "• ShowiSfg Moxphtmetry of the QkrQmo@&&9® of Fmaaatottlue
i>simaiit|>i yroufhton ( tem&le ) in trypsin induotd
Or htm&Xog pJDtpiurations*
3* TA3IM Y 4 «> Showing Morphoaastrj of the 3hroskOS<»ie8 of Fianaitemltte
jsemiaiit^ ayoiMhton ( sa le ) in trypsin indueed
Q btiMing p3»ipiyrations«
6* TAiiM VI Jk * Showinif Morphf^etry of t^e ahro&osomea of lianaabulttB
sennaati Wroughton ( feaale ) in \irem indaoed H
banding preparations of Anaphase stage*
7« TMiLB IB - iihowing , v.-•- . .,.:,.. ,^'fHorphoiaetry of the ahrc^aosomea of
iiattua norveigiema ( female ) ( 2n • 42 )
d« TAMIS 11 a " Showing Morphometry of the QhsemomtmeB of itatttis
noryeglous ( fesjsle ) in trypsixi induced a
banding preparations.
( 11. )
TmM XIX B - ^howixig Morphoa@t£y of the Ohromosc^ea of iiattue
noxvm^ouB ( male ) In trypsin induoed 0 banding
prepaaaatious*
I'AbliE IV B - showing Moarphometxy of "ttie ob.£(»Qoacmes of Eattue
nojsve^cms ( Male ) in urea induoed a banding
pxreparations*
tA^LB V B • uhowlng; Morpiie»]aet£f of l^e clisomosomes of i^ t t i is
aoaryealo'tui ( female ) in urea induced Q banding
preparations*
tAjdiS V'X b • Shovixiii Morphometry of the obromosc^ea of female
aat tua norveidLoua in urea induoed ir banding
preparations of Anapliaae stage*
(ni)
urn Qi WliMiii^>i l a
X* HI3T0&EM lA - liepresentg the normal Kazryotype of t h e
fiuiODbulus p@nimnti(tfrougliton)( iml® )
Diploid no* 34*
2* kil.ymmJM XXA- Eepresents the Q bm^ding p a t t e r n IMuoed by
urea In the karyotype of yvmuabttlua pentiaati
( male ) •
5 . la^foaiiM I I I A- liepreeenta the a baiading p a t t e r n induced by
urea i n the dhroiaoscKses of funuabulus peananti wm
wrou^tofi ( female }
4« iilafoaiya^ XVA- ilepreeents t he a bandliig p a t t e r n induced by
t ryps in i n the ehrc»ioaoaie@ of guimiabulue
pent^nt i wroughtcn^ ( amle )
^» tilsfoaiiAM' VA • Eepresents the 0 bioiding p a t t e r n induced by
t ryps in i n the ohr(»u2eoi&ea of Funumbulus peniiaiiti
wrou^^h^an ( female ) .
6 . lUSfoaiAM VIA* Eepresente the (I bandiz^ p a t t e r n induced by urea
i n the Anaphase s tage ohromoeoiaea of female
funuabulus i^ennanti.
7 , maiXiailAH IB - iJhowlng a r ep re sen t a t i on of the aoisttai is^ryotype
of Eat tua nonre^icua (female) 2a « 42,
8 , iiIdI!O0EAM IliJ - Showing a r ep re sen t a t i on of a banding pa t t e rn
inducea by t ryps in i n the ohroiiussomes of fenale
&attua norveitiot^
( IV)
9* iilUSOOliAM I I I iS «• ii9p£es@n%& the CI banding pattern Indmoed
by txsfpmlu in the ohroaosomes of mj&Xe
XO* llIofQ0liAl'! XV B • Eepzedeats the a baiidl3:ig pattern induoea
b^ usea In the ohromoeoiaes of umle Mattiata
11* iHi fua ily^ VJ « &epx9B9m%& Vote Q banking pat tern lodueed b "
ucea in iiie chrososcmee of fessale ^a'^tua
X2m .ll^WtiMM VX M * &ep3Mieezite t^e Q banding pattern IMuoed 'by
VLS@& In the Jkxm^g^me etage QJaXQmosomea of
( V)
F l a t e Ik t • 3hov the noziaal kasyotype of male I'miuabulttji laemianti
lar^o-fc^p^IA wgowgfttoiit 2ii « 54.
P l a t e I I k
E a r y o b ^ ^ I I A s - 3how th® © bsi^dlng p a t t e r a Induoed lay tirea i n the
ohro»osomi»e of male I W H ^ ^ ' ^ a^^,!^IA J££SBglllai*
f la t© I I I A +
K&ryQHY< I^I As- ahow the Q bandieg pa t t e rn induced by urea i n the
ehroaoaosea of feisale jr'^ft»i^f3.\ii, gfOM^^^ Hf9M^1ffi'
P l a t e I ^ A
Karyo^wH IV A s - iihow the Q baadliig p a t t e r n iMuoed by t ryps in in
the ohrc»aosMtes of male jrunumbuluB penoant i .
P l a t e 4 A
Karyof^p^ /A : - ahow Q baiiding p a t t e r n indaeed by t rypai i i i n t ae
ehromoa^iE^a of female gunimbulua pemi^aiti.
P l a t e VI A
£aryot\Hpe ?I As- dhow Q banding pattern induoed by urea in the
Anaphase stage ohromosoaea of female funumbulus
Plate I i;
t.&ryoh\^ I ji : - dhow noj^a l karyotype of female Hattim norve^ioue
P l a t e 11 a
Karyok)^ U ij j - Show the Q banding p a t t e r n indai^d by t ryps in in
tne ohiiramosaaeB of female i i a t tus norvegious
( v/)
Plat« I I I B
Kajcyokt^ I I I B s- ahow the 0 bandSjo^ |Matt®£a Iziduoed b^ trypsii*
in the oh£C»Bos<»ie@ of sml» Uf&ttuB aomreaiotts*
j^ la te I¥ Ji
Iaryo+«^. IV Bs- s5Jio« t&e S baiadli:^ pat tern Induoed fey urea in
the Qhr9i»>s{me@ of stale Eattus noir^igioue
y ia te V B
Ea£yo/H|p< f£ ; - 8ho« the & banding pat tern induced by urea In the
ohroaosoaes of female Eattme jaorve^oma*
Plate VI ^
^aryo-HjIf^ /I xij- Show the Q baadiiig pat tern induoed by urea in
the Anaphase ohrosiosoiMits of f <m!iale ^at tus
(v//)
iJ^jiji^JUJ^^J^^^
JL
m e deTo^Lopi^iit of bmiding p a t t e r n s nas followed
b;^ %h& dtsooveae^ In 1970* a tliat ythma. Qlxmmomom^& a3sm staixied
with the flaosooiiroae qidnaoriae or t t® de id i^ t iYee aad
Tlewed imder tU-tamirollet l i gh t* they fluoveaoe o r glov Ui
alterxi&te l i ^ ^ t mmd dark bande* l4,ke f l i iger p£ln%B these ua&da
are oharaoter ls 'y .c fo r e®.oh ohromosoae* yul lke fi i iger jprlnty*
howeirer t h e bands a re geu&XBll;^ ooiaeistent fo r a taxon, except
fo r minor ' r a r l a t ioas* A number of otner oh^ii(m3jt oaa a lso
prodao® banda e i t h e r i d e n t i o a l with or d i f ferer i t from the
fluoreaoeiit ones based oti d i f f e r en t pr lnoip lea* amoh
d i f f e r e n t i a l baadincs imtteamn ttsually observed e t spec i f io
re^^oas on p a r t i o u l a r ohromosomest &x% beia^ i i ioreaaii tgV
used for t he i d e n t i f i o a t i o i i of ohrooosoaess. 2h©s© &a*ids ar© tasiae
viM.bX@ as d i f f e r e n t i a l l y s ta ined a reas uride? the l i g u t
aioroscope*
A *band* i s defiaed && a p a r t of ohroMOsoae wMch i s
c l e a r l y d i e t i n ^ l s h a b l e frois i t s adjacent s e ^ e i t t s b /
apjpeariai: darker o r l i f t e r with a» ii» Cl» or a etaiaift,^
methods• i>y t h i s d e f i n i t i o n there a re not iu terbands*
The hydrigem botids Unking a iJ«l»A double h e l i x may
be brokea b / the hi*^h inteiiniity of t eopera ture ( l» u®v ) ,
r a d i a t i o n s aiid aoae chemicale «jf the i m r t i c u l a r ooaposit ian
eto« th@ euBulng s tpesa t la r i ^£ the a tx^idat asaocia ted
witii an inorease in u .v afesairbaocy a t 26uU^ A, i s toown as
denatiue&tlon* fh© teapersitur® requi rea foir ooapl«te
ienntmrt ' t ion i a Mgher fo r a « v2 tiia» for A • I
Suiisequeat &lQv cooling r e s u l t e im ro t ta tura t loa or
annealing: of two oompXei&esitHr^ a t raads* fh i s piieaoiifsiion
of deaatox^tloia and r ena tu ru t l o^ hae to@exi u t l M a M for tlm
banding pat texn aia&l^ais of oliroaosom<ia through dlffere£i.t
processes of denaturatioxi a/id reoatuastiosi* the d i f f e r e n t i a l
corapoaition of ciiroaosoaaX s o ^ e n t s t (xiiitaiiiiug Af/a%) xat iua
causes d i f f e r en t r u t e s of reaimealiiig b^ imLXiu/, pri£iaad.ly
a t a |>art ioular etagot o@rt£iixi segsteiita able to renature
f a s t e r* or i ^ s i s tmut t o de»mtu»ation» e i ^ b i t ba»d@*
aasperd30ii€<>^' JL97^ ) conceived t^e i a e a t i ia t irariou®
Qh:eoaommm.X segstents ma^ be eeXeotivel^ etidtied idLtn au
appropr ia te fluoresoeiut ^ei&t t h a t i^taM bind eeieotiTeXj to
baaea i n otakroaoeoiiaX D.IUAt £tie f i r s t , fXuorooiiroiae stwiied
was quinacrii ie siustai:^ ( ^.H*) whicit t^vm In tense fXi>^reaeeiioe
i n t^e heterooliroma^c r e^o r i a of dir^ckoaomes* Jueh b&zid&i formed
the laud marka i n a de ta i l ed ana l^e ia of the liimafi ^aryot^'pe.
wther a lkjXat iag fXttoroGliro'.}e©, iaoludii-ic; aai i ioaorldi i ie ,
•iXtrogeii mustard aud a oislnenaiiaidaEoie d e r i v a t i v e a l so
produce siaiXiir brnds i u pXajat ohrooiOBameo. 411 tiiese u^es
Bhoyi a pronouaeed speo i f i o l t ^ fo r nuoleio aoids e i t h e r
beoaya© they r e a c t witii ii^h^A* by a l ^ ^ i a t l o a of oeutaiu
purine bases ( a»H# ) or btoause they feear poalt lYe oharc:©a
( AUt p ro f l av in , aorofiavizi ) wMoh allow thea to I n t e r a o t
with tae D.l4«&» phosphate baekbone.
'ihe f a c t o r s influeacljcm th© degre® of co ioura t ion
dep@M@ oa the ohOEoiQal a f f i n i t y between the ooi is t i tuenta
and t h e dyet the dens i ty of the ooiist i tueii te and i t a permea**
b i l i % h'j the d @« Qhromoeoaai s t a l . i s @re tersied ac id i c or
oaaio on the baaia of thedr ohfisaicaL imtmre and beha^our*
X:i some cases t as i n oroein« the^ a r e amphoteric* fo r
d i f f e r e n t i a l a ta i i i in^ i t i s the p r ac t i c e to overs t a in the
ohroiioaome» followed b^' the remcml of the exoee® s t a i i i , a lao
kaowu :ia differ©atiat ioB« J t a l o i a j ca» al@o 00 performed
progjeeeeiTely by grawiual iuoreaee i a the ia te i ia i t i r of the
colour C liaker, I9&0 ) • Han^ chroaioscae @tairA& are avelMble
with varyiiig degrees af spedLficit^- fo r iimti»A» A widely uaed
cl-iromosome a t a i u l a orceiii* l i ta ih iag i n t h i s aye iaTolvee i t s
coacensation a t the polypeptide l inkage of t h e ehrofflosooal
px^ t e i a . Another utaln ueed extenaiTeljr in trnxmBiULmi ooromo-
aoees i s aieiasa •• a mixture of aereraJ. dyee* inc lud ing
ae th j l e ae blue Siiid i t a oacidation products • assures * A* a.id
*i3* lijid eoaiaa X, file q-aaMt:^ of the a ta i i i mi r ies vt%n -jeegnx-ik
to tiie propor t ion of the d^e used*
Methods to demuoetrate i^iemaa staiueo, oai^dini; pa t te rua
'4
of maamalituiL aturomosomea wese© deaortbed from d i f f e r e n t
laeoratOEles ttsiag baaio p r i n c i p l e s of i n vitaeo iiyi^ridissa-
t lon of J«4.,A» Ta® e a r l i e r methods depended on aUialixe
deuaturat ioi i followed by r e n a t u r a t i o n i n an adeqaate buffer .
Pardue arid a a l l ( 197U ) t i n prepai^t iSt ts f a r i n s i t u ayferi-
d i l a t i o n wiiih oouse s a t iJ»k»A», noted tiiat a f t e r mutoradiogray%
Odiutraaerio regiaiiS af mou@@ olircfflO@oa@a nhoired mar® de^.a^
f^raiaa tiiaii oiiraaasoae h3smm uad w@re @ou&id@£ed to ^m a@t@ro^
cfiromatie* *£he same procedure ^a& used to ali@¥ ta@ preaei^ae
af auali reijious km hw^axi c^iroisosoQea wita modifioatiQiia* i.lie
areaa of oen t r io iieterooiiroiiatla a ta iaed mQ3:% i i i tensel^ t^i&u u. fe
r e s t * I t was eu^^gested t h a t tliese regitstfia probably/ repreaeiited
repet i t iT© U.fitA^ Th& preauii. 9d bas i s f e r d i f f e r e t i t i a l atairilii^
under t h i s teolmique was tiiat re^iatured D«B«4« appeared b e t t e r
ab le to aoabine witii s t a i a ^ a a p a r t i a l l y denature^ j*A*k%
4ft@£ denatura t ion by the d i f f e r e n t age&te used i t was apparex.t*
Xy tiie r e p e t i t i v e l)»i,A. whieli reaa tured most r a p i d l y , fhe prooeau
of renaturatiojEi was Imlted optimally a t a t i a e wben repe t i t i i re
Jmil^k* hM. n o t . Tlbiia epeoi f ie a ta i sdng wae a t f i r s t a t t r i b u t e *
to oona t i tu t iTe beterooliroiaatin. tiowever, l a t e r co i i s t i tu t ive
heterociiromati.i , though genera l ly aasooiated with sate^juite
u«li«A* ta the mouse ( t h a t i s repetiti '^re J«iuA«, Q.Q% the J.£i«i^*
of e a t e x l i t e e ) t was aeeu to be a e t u a l l y a d i f f e r e n t e a t l t /
aad 4id ao t neoeasar i ly oons ia t s af ehor t r e p e t i t i v e i m i t a .
AGaordiiig to oosiiiit;s ( 1973 ) tiie ©eiitr^-ujerio aettfro-
ciroifiatia appe^^rs i n the Q bmi^B ^uid the io te roa la r j ' neter^*
oarometiii i n a waady. 'Ihe oei i t rauer ie aeteroohraaat i i i i s £A.aa
v>
ill hi^-^hly r e p e t i t i v e eat t iULte 4i«iUA« wMie tne iutbsoaloj^
iieteracisxom-ti.* i s reg:ar4e4 to have aon r e p e t i t i v e » poo£Xj
i4tJUler ( 1975 ) obs#rv®«i t h a t D«i»«A«, tlie ia^or ta t i t
Oriromoso^l oons t i tuen t La oaiidiia^ doee not aocoutit fo r a l l
trie variOiits* J ta i t i ing witfa ^liemaa aad o ther tiaai© d^©s dfejieaue
t a a Qouaideru&l# ex ten t o& t^e bitidir^; s i t e s pneueut lu tiie
i}»:»i,A, Axi@ a v a i l a b i l i t y of tlieae ait@a i® oout ro i led by tfi®
f a c t o r j i t a e i r naiiig olooked by pxotelia.
Mu e t al« ( 1972 ) obat r red the a b i l i t y of t ryps in
:u^ a oliymotrypelfi to produce baadiag p i t t e » i s i n iiuaan
oh£omosom©0 au d i^imllai? pat t®me were a l s o obtained witu p@p»iii.
•iiiey ooaeluded t i iat the aiid.l*irity of if©suit3 witu d i f f« rcu t
©naya#e suggests t a a t the bloQheoiaal aaapos i t ioa of aasooir^ted
iilataa©e does imt deteraiu® tae baadiiig j ^ t t e m .
iironn e t al» ( 1 9 7 P ) presented a oytoohemioal data
fsoai "^e ohiaea© hamster f l b r o o i a a t lljae» i)on aaa the ofeHs af
4>erog.?8oa^ ereairOus to show t h a t the feistone f r a o t i j a s f_
aiid f .,a were iuvolved i n ta® iaduot ion af oarajsosoaai. \» ttaads
Wiiereus the f ^ ^ * ^5 fa^totions a re not involved* Aesoval
of the f and fg a fraotiui ia probably QOCUZU duria^^ fixati.^ii
u.id I s neoeasary fo r the i t iaact ion of y baiids,
iieoeat aethoda for sti*dyia,t| ir hmv^ln^, a re a i e p l e r , waaed
on asBumed d i f f e r e n t i a l dena tu ra t iua oiily, 'A'ae mg@nt3 for
proincing bands a re a l l : a l i uaed i a a i f i ' e rea t t eape ra tu rea ,
-me of severa l p r o t e e l y t i c engymes, aad pro te in deiiaturiiig
v>
siibstanoes ( iirea )f tacLiai-ig ci«t©r4,eiit8 usually a.iioriio
ott©a» 2h& pi'oo^aa of djLffer ta t ia i preceaainci use<i i n tliese
s impl i f ied teoaniques aeeas to depend on taae temperature, pu,
i o a l c s t r e n c t a , l i v a l e n t c a t i on coutent aii4 p r o t e a i / t i a ©.issy J®
soute.i t of %h® aedium. A jpittejcsi jt furi'StlB jtolalu^ i© awt»e£-
ved uJ'ter tiiee© t res tees i t J u«id af t t i" staijd.a*f, ^ i t i i y ieosa .
a* vmiAa pr&dmoe a d i s t o r t i u n af ta# riO.Ka4ii c i i a t i i u a t i . , .
•A tii3 ouroiaoaaaai. f ibe rs* f!i« dividLefit c a t i j u s amj pla^ u
r o i e i a oaaaiii *• f ii© dajck aad l i t i h t baiAO, ret-iou0 a re ifegarUei
to r e f l e c t ttt@ coi ieeatra t ion litid amiiic®®^ ' " ^ ^ iiuoleoproteiii
f i b e r s ( nsHT, 197> ) .
2he ro l e of p r o t e o l j t t o ©liajaea i n proaixoing oiUiaiUf
I n d i c a t e s ta« imjp^rtaiioe of oiisoiaosoaai pj?oteiii8 l a wariaiag.
lliG teoiiiii^aee ma®d i a p j^ees t atudy are simple usiat; tr^^.ji*-
uad mreu as denaturat i t i ' : agents separatel^r fo r the stiwi;/ af
si bands i a jii&tttta <iorvegioa@ u id .^wnqabi^iie fcuaa^lfl. tfrQu..iitM.^'j
chromosome ooapl#iieut3«
uenera l l^ onXj t^ie ai^,;«3tive ac t ! an of t r i p s i n a^m ^«ea
coas id t r ed . ii^erteps t ryps i» Sias ween M't^e e f f i c i e n t taais o taer
t rea taeat© i a produoini! baiittla/, beoams© i t e ac t ion oa protein©
proQ«@da itt tvQ a i a t i n c t s%^£itB, i«e«» Jexit^tur^tioi* mid
d inea t ion ( O h i a r e l l i , -. 1972 ) •
Trypsijti i » i t l a l l j aei iatures a p ro te in ambatr^it©
( i i laaeretor i i , 1952 ) , a f t e r wlilca Uim pjK>teiiis axe laore
seuslt iv© to the hydra ly t io a c t i on jf %a& eussyme* £&ia ae«a-
t u r a t i o n a c t i v i t y iiivolveis oliita^ieB i n ui® aecoiidur^ a d
t e r t i a r y baadinif i a th© pro te lu a o l e c u l e s . ih@ ©iiaiitjes i a
thea© biiride K weak bonds } cause an aitex-ation in tae tjeuerai
oonJti^^ttiimtiau of t^e pro te in molecule* k,i case of tki& fiberooa
p ro t e in s ttie &o^o& of the dena tura t ion prooeea Im^olvee
breakiiQig of the seooxidlary li^dr^g^a toonas tmcl re@tilti» in tae
leagtJEiiHQdLxi of tlie molecmles*
The second s t a g e of t i ^ f s i a ac t i on i s a digest! ire one 1 .
which the primary boEds are brokea and t h . p ro te in i s euo*
ai'Tldeci i n pept ide eegmimts* fhe otiier pK»teol^t ic e n e ^ e a
proceed d i r e c t l y i n d iges t ing the p ro te ins*
file f l r a t s tage i s the i^aX ttanding p rodudng aotiOQ
aad the lengtheriiBg of p r o t e l a s i s observable on the tiioros*
cope i n p repara t ions in vhich a s o r t of over laps of proteliris
of ohroaosi»ies a r e demonstrable a f t e r tr:^psin treataaent*
fhe i n t e r a c t i o n of t ryps in v i t h o ther f a c t o r s must a l so
be considered* Apart fr<»a t i o p e r a t u r e and coneentratdLon,
t r / p s i n i s found to be acti-^ated by c e r t a i n cat ions« suoh aa
calciimy manganese and magneslim and t o be inhlbi teci by o the rs
( Sine* mercuric e to*) t ( a h l a r d l l l » 1972). Moreover i t i s
i n t e r e s t i n g to note l ^a t some substances such as versene or
ethylenediamine - t e t r a a c e t i c ac id reduce the ac t i on af
t r y p s i n ,
Extended ac t ion i d . l l r e s u l t i n complete d iges t i on of
the ohromo3omea« tiie presence of versene or ethylen^usdne
t e t r a a c e t i c aoid w i l l probably i n h i b i t d iges t ion*
6
j j i l t f aotlom tlrae alioMQ th© tr^-psiii to aot a s a deuatttririir
agent and not QM a liigeetiv© ©ug -ia.©* iJsu'idiiig ^aa^ thus invoive
only the dena tura t ion prooess exteiiding p ro t e in i n o#r ta ln
a-usoeptible reg ions and, r©3ai t i»g iwt a weak e t a i a l n g a r ea .
aoxisidesiiig a l l ^ o s e po in t s t^e baxuls cm 111® olixrcmtouc^eii
appear to be the product @f the i n t e r a o t i o n aiaofig time/
tempejeatiase/ oozioeutsatiOQ aM preaenee/ absenoe of other
fh i s sime o r l t io le ia mid tlie aaiae ^sokieideratioua iiiay oe
v a l i d fo r the banda psodixaed by the otfeier agents suoh a@ h@&t»
u l t rase i i io waTe©» aoid/b&se/tirea tsemtiei i t eta* wliioii pix)da.m
denaturatioi^ of f ibesous psoteina*
lle.tl've glo<.\bular p ro t e in s imdergo denatura t ion to y ie ld
unfoMed, raMoE ooufijsaations of t h e i x polypeptide ohains wheti
t r e a t e d with atromg so lu t i ons ( 3 M ) of urea* Altiiough taXm
ohange i e aooompaBi<^ by l o s s of t h e i r b l o l o g i o a l a o t i v i t y t
some deaatured p r o t e i n s spomtaxieouely reoover t h e i r b io log ioa l
a c t i v i t y and thus t h e i r o r i g i i m l natyLve coiifismatiofi, eometLiaea
very rapidly* Olasaioal expexisent^ c a r r i e d out by ^itum Mkltn&(Sii
{ X9T} ) on r ibonuolease f i r s t showed the importaii,ee of
aminoaoid aetuence i n the detexmln^ition of na t ive coijfirmation*
i'reatsient of n a t i v e r ibonueleaae with a M urea in tlie
presence of the reducing agent ^ meroaptotxianol oau^ea complete
•ojifoMin^ of the r ibonuci«i«e ao leou le , to y i e l d a randc^
farm*
\ 'l
knXtmmmk ( 1975 ) ahovesi tliiit a naoieae® fxoa sitajphytooocams
oexla loses a l l i t s biological ao t iv i ty and i t s native
oonfi^ia'y.oii wiiea aotdifi©dl to pii 3«0 wiiaa the pH i a restored ta
7» the ao t iv i ty t e quiokly regained*
The 2»epO£t of olark ajad if'elsentfeld ( 1972 ) OB asaooiation
of arginine * r io^ Metonee witii G Q rlob. U*'&»A» may i>e au
iia|)ortcfuitt kei* to oux underetmidii^ of cliromosomd atruetuse
di8t:dLI>ution of tlie T&xdoae t jpes of maoirsmoleottles tmevet^l^ &lau£
the otirfmoec^Bes*
Qoiisidenug a l l ^ e s e poiii.ta the buMs on Va% ohzroiBosoaes
appear to be the prodmot of tbe ji«I«A* froteiu iateraotion*
'i?hla view explaino almost a l l the a^milable eTld@Eoe@« eiuoe
Ijraiae rioh histoaes appear to bind prefere^^tiall^ to kt rioli
'ilfmUrnkm
In tae preaeat i^ric att<rapt has been made to dovelop and ..
obaerre the 0 baMing j ^ t t e m a iaduoed b^ two deaaturatiag <:gsEta
t r jpain aad urea in the karyotype of ftyi^mbttlttg geaac^fiat^
oeaaijaiiti has beeii diaoueeed in deta i l*
mmm^M MM Mi^m
10
Mbox&tos^ hve^ albino xmts l»e» ,§f;|^tti ^oirrt^ewa of
l>oth aexe@ and f leM o€tXleot«d si&a.e i^at feniale flire 6txflpp«4
sqttlrrcie ( ImJ i l i lWI gfHIMl^ Iie9^?^l« )««»« «s«d for th®
fO£ c^£oaQ802»kl Btudl«» hone masvoit was eel«et«d as i t i s
aB 143^1 aate£i&l for the s t i^y of a i t o t i o ohrcaosciiiies* for tli@
m«lotle Qhroaoa^ae preparatiooa* tes tes of sa le 3eat@ and amle
eqiarrela were used* Ti^ a i r dried aetiiod ( £«a« Hen and
Patton* il«B* 1.96 9) wsui adopted for t^e preparatioii of siidea
beoaitee the flails dried elidee are r e s i s t an t to tbe treat&ieat
given for iadaai»g baxidiag*
fhe aniai&le were iajeoted with 0*2 al» of 0«0y peroeat
oololiioiife eoiutioia/ 100 DSIS of tood^f weight» iut@ra$erltQ#eall^
one hour prior to dieeeetion* After Qn» l&our the eiiiaaX vsm
aaorifieed by an oTerdoae of #iXo£ofliam* flam aarrow irae a b t a i n ^
froei femur* Sbe heade of the baae were emt obli.tttteii'* JloXea were
bored with needle no* 24 a t e i ther end* the boi^ aarron irae
flushed out by foree siith 0»%% peroeat poteaeitim ehloride at
31^'<** Qells thus oolleoted with th ie hypotonic solution ( kcl)
were brought in to suepenidLon and were t&&\i^&t®d for th i r ty
aiitiutee* In oaee of laale animals t ee t i a was removed and the
euapeneion of t iseue s^de in leal and iiioubated for half an hour
in the inoiibator*
After iaoubatl&ia both t^pes of uMup&iBtonB pese
oeiitnfiiii®<i a t 2400 si^ for t©ii atiiut©s# aaperiiatatit waa
diaoarcied mud Jainsojr's f ixat ive ( 3 j ^x t s ©©timnoi, j 1 part
£:laolai aeet ie aeid ) was Ed4©4 to the r@@ldue luid &mp% for r*
!aialm»»a oi oae hour i a ref£ig«3eator. I t was agata CMin'&etftt,:ts4 far
ten aliiut«a and th@ supeimataat dl3Qai?4#4« the mhowe fixation
proc94m?e was a?©p#at©i twio@ with aa tn'tmv9&l of a t l^ ia t 1/2
hour and tii© susipe&sioa was topt in r«f£lg©rstor f o:r 24 hours.
4ft©r i a s t o@iit3dlfugatlou the suponiataat was disoai»i@4 and
a miziimtaii suitou&t of fixmti're was added to the roaidue* i'Ms ee i l
suap@n©ion was dropped on pre'Tiousl^ eliilled gXtmm s l ides wita
the help of faateur pipette*
^499%^f|, ^ g i J§y^^^ h^^mu
tlie pji^oedure adopted to indue© 4 liaMe in the eliroQiosoiieo
of botb rodent apeeiea was tbe same a s timt desoribed by ae&i^ri.-ht
( 1971 ) with a l ight modifioationa.
( i ) IMij gJ > ^i^iOi ^^milQif ; - G,20 i>er©eiit or y«23 percent
stook solutioB of tr^paiii wea prepared lif disBolirla, 2u- mc
of tryptsift or 250 m^* of trypBin/JAiO lal* of leotoai© ph^siolo-
g iea l ealift© ( the sa l ine free from Ja"*" aay %"*" ioas ) .
2hi3 oolution i s stored a t 0**J i a freeiser.
(2) 'iCiii ^^J >i;uLai'l,uii /w v Xl 'liMJlijtj i, u^^ 5— ihe stook trypsii.
solution wiAis diluted to Is 10 bj addinci lu parte of ifoster sua
^ragert isotoi*tc pti^'sioloijiaal sjr-li.-ie t,-' one mirt yf atook
l :
t ryps ia solutloii* fkila soJUyktion nas st@billss«d to 4^u )>efo£@ ms€*
Ba oXt 0*3a6 ilm Kol* 0*0^9 m ^£^^4 ®^^ 0*420 ^ at MaMGv
dissoXired in one Utara of distiXXsd imtdi: to ototalia t&e jrequlired
(4) 95 pereent Ithanol.
(i?) ii'iiwa iy>'i:i& .iiUf fM $« 0.442 ^ of aodtaa phospliate oibasio
( Mol* wt» «• X77»99 ) Has disaoXired in 2^0 ml. of d i s t i l l ed
water @ d 0*390 ^ * of aoditM plioapl^te aoaobasio was dissoiTed
in 2 ^ ml* of d i s t i l l e d water separately* tl he 0*01 M £loreniioi:i* @
phospliate I»i2ffer wa® obtained by adding 10 •§ ml* of aonobaeio
eolutioa in 2 ^ ml* of dibaeie*
(6) a iMM m^MTlQU. $» 2 ml* of diemsa eolution m@ added to
the 93 al* of Soreiisoii*9 bi:^fer to obtain Cfi^ssa solution of
m 7*0
frypsin eolation ( IslO ) wms prepared in a Oaplin jar*
Uaing th i s Jar i s mQim oon*ve:^ent tbian flooding tue s l ides with
the eolmtion*
the elide© were dipped i n th i s trypsin solution
< a t 4^Q ) for 30 seoongs.
iiiiising was done with ia^tonia ph / s io lo^oa l saline two
or three t iges
1 " J, <.)
Ih® ne%% £iti9ing was dotm toy St) roeat etiianol mid tne
slides w®re dried*
After i&xylng, these 83i.d@s were p^Ueed in 2 peroent buTt'ered
iliemea Et&lutloii foi? 4 minutes for staiM-zig*
JoatroX slides ( without t&e treateei^t of trjpaia etc* )
irere also etained in 2 pereeiit aiestea eolutien side hy aide bat
in separate Jars*
washing of tiseee slides after atminini^ ^m& done by delanised
water and the slides were kept in the ineutoator for inoubation
at i7^U for one hows* Buparol •m.e used for aotinting*
>lany faotora mi y' influenoe the suocess af a band staltiLui;
by trypsin t^^atient* fhe suooess of i» banding depends pxlnariJ^
on tbe oonoentration of the trypsin solution and tlie duimtiou ul
the treatment* TempeaEtiture of trypsin solution also plays important
role in induoing ^ bcmds* ( tk@ higher tne temperature, the
faster the reaotion)* The solution ehouM therefore be atabiisea
at 4 0 prior t^@ use* The stosk trypsin solution should be Eept
in freeser ( at Q^o )•
'l!h@ prooedure of induc4nig il bayc^s, given by Klato txad
losida C 1972) was followed with slight aodifioations*
tl) jj.|,4 ,^*^^kji.It),.: jw d M aq^ueous urea solution was prepared
14
by dlssolirlng 4B.048 ^9» of ts»«« ( Hoi. wt. • 60*06) i a 100 al«
of di@tiia.«d wa'fees* fhis s t oA solulioii was k«]^t i » £9fxrlge£at@£.
fhsee pavts of t i l ls a t o ^ soXmtlos V9X» 3^x«d t i i t^ 3. past of
0«01§ II 3o£en90ii*8 fhosfiiat« l>iiffer to dia^t* i t« fli# d i ^ t e i
urea aaS l>uff«£ nlxtuse was @tateills«d a t ^l^Q l»efoir« tb«
tjpea^ent of s l ides*
<2) 9f0,ii If .jc^iiif^is'f f»ff?MiM pmm «*• ^ • ^ t ^ 8 . of 80di\n |^osphat« dlbaalo ( Hol* wt* « 177*99 ) was diasol'ved in
IDO nl* of (iiatlll«dl vat«£ oM 2*^40 gm®« sodium pliosphate
ffionltl^iaaio ( Mol* irt* • 156*01) vaa also dissolved in 2Qy B1* of
dis t iUt td water sej^rately* The required 0*01§ H buffer of pH «
6*3 vas obtained by addisg 22* § a l of manobaaie solutioa to IQO
ml* of dibasic solutioa*
MSSKmrnmimmMti
th9 s l ides vere kept i» urea aad «iorea@os*s buffer mixture
for 10 miiitttes a t 3t^o •
iif t e r 10 a iautes these s l ides were xinsed witli tap i^ ter
and without drying dipped l a hvi£t«s a i^ i sa solutioii ik pRreeat)
of pE * 7 as i a the oase of frypola t reatoeat for 4«*6 miautee*
file oontrol s l ides were also staiaed aide b^ side but
separately 4 "• 9 aiautes*
jurter s ta ia la^ t r las iag was a^aia doae l a tap water* fbe
alidee irare tliea didi^ and oouated l a l apero l .
ifouatla^' material plaore ^^ importmit role* beoause ii.P*! eto
lo
iir band 9llil«(S aM tlie 8iia.®@ i^s t a ln 'msg mmu^ altMou^a aouati^;!
16 not »#o«a»ar| i t @li(ml4 ton doae fos prsstnrlciii to« s l i d t s fox
2ii# @a#pij^ aM. studi€i» &£ l^aadln^ pattern of a H
sbx0moa©ai)@ wmm and© ua4«» te# yairl* 2i©aa ^. CtjavaJ. Mgfet
miQX0&mpe» M4e®opJ»otoip?apto ir®r® also -te^^a b - tli« 8a»« »l«»o«-
l a th« <»o&$t3»6tioii af thm QhsmkQ&^m^ eap3» ^mo'ii band wmm
£>ef«£i?@d to to^ it@ is4dXliie load act b ^ l t # m&^^tm* k ebxsoma&omB
* Laiii vmtk* was @ojn@idUiir«d as a oousistant aM diatiBOt i»«-i;| h6«»
lo.flssl f«at«tr0 tliat im mi iia^ortaat filaipioette aid in id©iitif„.-
t=0loRglssg «ntii3©i|' to th« »#gio». h i a t a l to tfei© i&iid iiitXM aatt ifas
aoooMM iMB bmid ri*»fe«» of •I* IK the i?#gior»»
o@mtra»@ir@ @ut ward aleiig 9s@M olucosaoS'Cms ai^* A U&A4 M3#ot«&
b^ th% iM&iitxomere UMI Qvm»iM^^4. m& two ImndSf aa b«ir4{: Xato<illoa
aaaEEtfAfiQM^ j ^ POTM§
i:>
%^ '•i,^*^^' *•*• „ » , ^ * ^ ^ i ' - i ••••'™r' *^ " « « . ' ' « . * ' '^«w,^ '^ , '^«,* '« . • *"*»„*•« . ' * *
Til® t eoha ioa l quality' of the oaroBsosoaea i d of tiie
ufeoet Importaao® for cha rac t e r i z ing di i^aosoaes toy t i i e i r
bandlstg pa t t e rns* I n faot t i t i@ mudi more important taati
morptoolog^ wiiloii Was pre-rioasi^ csoasrfldered as th® aole c r i t e -
r i o a for pf@p&ad»ag the kar^et jpe* llevertheXese Morpholo^oauL
oon€iidem,tioiis of s i s e wad cen t r^aes io posittost s t i l l reinaiii
o r i t i o a l parameters used i n the ide ja t i f i ca t ion of ctiroraosoise;^*
III til® presen t work karyotypes of two rodent spec ies | ia t tu^
at ' t^ied with the he ip of 0 baading pa t t e rn as veil as by
employing %ne olaeeioaX aiorphological techniques In an e f f e r t
to d ia t i i i^o ish i n d i v i d u a l ohromoaoiae.
aomio^s ( 1973 ) has suggested t h a t i&teroalary
heteroohroEiatin appears im the a bands» irhioh i s regarded ta
have rn&G. r e p e t i t i T e poorly methylated i)«ll«A. I n ^ or a baauiii<sa
n e ^ t i v e or pale atai t i ing reg ions are dLxm t o the presence of
non h ie tone p ro te ins* binding i n the eiaall groove of D«lu4.
aad preventif^ t i ie i n t e r c a l a r y binding of ohrososoaea* i'he clfir^
bsndsa Slid l i g h t or negat ive bande a re jL«@garaed to r e f l e c t tri®
oonceatr&tion aiid arrangeii«it of nucleoprote in f i b e r s .
the r o l e of cheeraloale or organic compounass l i k e urea
and p r o t e o l y t i c ®ai.yae, Mite t r yps in in producing baadia^
i ad iop te trie importij^oe of chroaosoEiaJL p r o t e i n s in banding.
4 r-'^
MMf^J^Um iM. SP il.iiaUsl9il^.„ yl^
f}i« d ip lo id UQ* 2n a* 54 was d,ete3rai».#ii aft@r obs«zrvliig
s e v e r a l metapliase p l a t e s of t i ie i ad iv idua l e of ea@ti sex*
Sliree shmpes .of the ct^xoaosomes oisous i n ttie eomplemeat
accordlag to the p©ai^oa of eeiitroaei?© oa tae caroaosoaie.
( 4ft@r Albert LtYaii, KmxX fredga asai A^esy A^Jandserg (19fe4).
(1) Sutaetaeent ido (2) Meta@«iit£ici O) and Aoxoeexitirle.
^ex ohromoaoaee x and jr a re idemtlfla,ole eaiai.li'* i. I s
a l a r g e aubae taeent r io chroBoaome liaeluded i n group I while
y i s the smal les t aoroeen t r io ohromoaoiie o l a s e i f i e d in group
\ r i l l t I t i s t h e sfflalleet oliromosome of tlie oompl^tinit also*
tiie aa tosoa te caa be arranged i n t o e ight e a a i %
d i s t i agn i shab l e groups socordiijg to t h e i r length,, oeiitromeide
poe i t i oa sjad pa r t io t t l a r type of banding pa t t e rn •( After Ali^ert
LeTan, Ea r l i'"redga and Arer^ 4»iiaBdberg» 19t>4 ) •
0roa.p f i r a t otaprise® of two p a i r s of autoseotee
^ pa i r a I •*• I I ) and one- pa i r of X elir0ttosoffie8 i a ease of
female or s i n g l e % i a ease of s a l e aaliial*
iiitOMO tlfi l liu« ]L » QhxomoQQSke oae i e tlie l a r g e s t eliroaoeome
of the eomplem^it* I t s r e l a t i v e l eng th rangea froii 6*66vta'l»^.# I )
to 8»70 ( fable I ¥ . ) percent of the to t i i l imploid leiii?th. i t
Geatroaer ic inde^ riuiges frcm 2Q ( fab le III,) to 50 (fable. i»'. )
I t 3Moif.8 t ha t tiie ol'iroaosoia® i s subaetsaoeritria, -sltk proaiiiect
m
s h o r t aaoBS* The r e l a t i v e ieagt i i of shor t axma rsmges from i « 4 l
( Tatol® i n . ) to 2•75 ( fabi® IV« } peroent of th© t o t a l riaplold
lengths I*
In both t^p®a of pipepaxatlons ( mrea tzreat^d and tsypain
t rea t^ i l ) whole shor t arm regloB aiid t2i@ oeatramerio ceglan
show inter^se darii: s tsdola^* X^ the long asm p£oM,m&ll^ or In
the regloE f i r s t s &. l i g h t 'bajad and a imxxQv darj£ hanci ar@
preseii t t Xn the region seooM» one torosul dark band (laiuimiirk)
i s aefmimted fron another dark band di&talljr by another l i g h t
s t a i n i n g band*
Cil&QM03Ql jp Ha.* 2 8- AoaordLlEg t o the ©eatroiaerie index value
whioh xmigBB from 20,40 ( fab le IV,> to 32,0 ( f ab le I . ) t h i s
ohromoaome p a i r tm a l s o en^^e'l^oexitrio i n shape* She size
con t r ibu te s abomt 5.95 ( Table I^ ) to 7.76 ( fable IV^) pereeat
of the haploid se t* Here a leo the s h o r t arms a re very prominent
and t h e i r r e l a t i v e s i s e ranges froa 1*41 (Sable H i A) to 1.91
( -fable ' ^ I A ) peroeut of the haploid s e t lexigth.
In urea tre^^ted prepara t ions ( f ab le 11^ + XII^ mid
idiogriffii 11^ * lll^ ) @ d a leo i n t rype in t r e a t e d prepara t loae
( Tahls IV^ *• VA a»«i idto<gj»a ^^A, ^ ^z ^ ^^^^ darM: and three
pale or l i g h t bands are t i e i b l e on t h i s ohromoeoiHi •
jpi^sdjaal region of ehort asm poeaeasea a l i g h t band while
d i s t a l region of t^e same haa a darK: one* Oentroaerlo region
i s Mgiit ly s t a i n e d . In the r e ^ o n f i r a t of the long ana a dari.
baud X& p resen t prosdmal to the oentromere* Remaining port ion
of the proxiiaal region ( region two ) showa a iiroad pule band*
i:
A amaXl dark baM ttieo a narrow HgHt and again one
bvoad dark bsKid air® prestmnt In the d i s t a l region of tae i,ofi,:
X OiiiAO iSQMI ;-* Tti© 8iE# of x ohromosoa® raagea from 5.6>
( Tabi® 111^) to 7*44 ( T&hU I¥^^ ) peareettt of %h% t o t a l
haploid length 0 tM.& i s a lso a smbaetmodntidLo ohi omOBoa®*
$h« e9»tceaerlo iiid«z miogea fsoa 22*22 (fa)»Xe ¥^) to ^2*00
(fabl« I^) wltb laxg« mA ]^iemainen% ohgvt arms aliout half
th® l«]Q||th of lOEg ^cem* Th9B9 short asae oontxibute about
1*44 (fabX* V^) to 1«9B (Sal»I« ^^^4) ptcoont of the haploid
80t«
A daxie baad l a prolm«ixt d l s t a l l ^ on the short am*
proxlfflall;^ ^^ ahOMS a 34ght bs^e^ only, a^sitroaesdLo rogLozi
ahovm dark staining band whioh ooTors a anal l portion of
proiimaX r e ^ o a of the l^ag aim of t h i s ohroaos^ie also*
Xa the proximo ail r e ^ o a of the sayRe a m a pale ba»4 1@
preeexit of t e r the dark one* Oieta l region of t h i s long Q3M
shows only 0]»i broad dark bittd*
^he eaae type of banding pa t tenw were obaorved
i n bot^ urea aa iml l tvfpeiM t reated preparations
(Tables ZI^ -¥ III A and Idlograias II k 4^ III A and tables
IT A 4- YA and I d i o g r a ^ XV A and 7 A )•
Ohr^aoacme pairs 3 to 6 are inoluded In th is group*
fhe Q banctinj^ patterns are s i i ^ l a r In urea t r ea t td as well
as trypsin fkreated pi^paratlona*
dd^i^lQaai^ m*3tm Aoeordlng to the oentr(»aejdLo position which
'Ihls dteoaoaoae I s aexroQ«iitad.e l a slMi|)€t and Xargest of all
the aei:ooe&trio@ of ^aM 09spl«s#ntf with a 8iss@ raage of
§•22 (falaXe XV A ) to 6,19 (faille XX A ) poroftnt selatiTe
length*
fhe proximal cogioii of long axm shows a darB: and then
a l i gh t ba id and one narrow dark band af ter the pale one*
In the aiddle region one narrow pale baM i s present*
i>iatal rOf^on shows two dark bauds with one pale h^aid in
hetveen*
aH&OHOaOMi ly*4s* fhis ohromosoae i s also aorooentric tn
shape with very inoonspiomoue short arEas* She siEe ra.4ge
froa $.22 (!]^able I f A) to 6«02 (Sable I I I A) peroent of
the haploid len^^th*
fhe short mma are l igh t ly stained • aentromerle
re^<m shows a dark band vhioh covers the small portion of
proximi'.l region of 3Long &rm also* A l ight band i s alao
present p ro^aa l l ^* ih^ cdustal region af losmg am shcvu
two broad dark bands with a narrow pal® one between them*
CHiiQHOi yME MQ.^t* Shis another acroeentrio ehroaiosoae
var ies frs^a 4*74 (fable X A) to 5.57 (fable I I I A ) peiOSBt
i n i t s eelativ® si20*
Uentroaerio region and «ia»ll portion ei"' proKim'. .;i
region possesses a single dark banc(s» re .it of the proximi^il
region shows one l igh t and one dark baad r s speo t ive l j .
2
I n tti© d i s t u l ree5to» on© pale hand afspax-atee two ir@i?y
proatnent ia^k baaaa.
OiEiyMQi?ailE. MQ>fes-> fh l s ohromoaoaie l a a lao a larg® aoirooentjcio
ohroaosom® vlth a r e l e t t r e si®© 4.74 (fabl© i A) to 4.5?
(Uabl© i n A) p«i'o©iit.
aentroe@£'io rei:i.on and .a mm.ll, por t ion of p£oad.iml
region a re ooirered by a axroiil dark baaiit One paie and one
dark band i s a lao obaarvffd i n ta@ pi.'QXi@al r#e;ion* I n the
d i s t a l re^jion of tk© Bt%sm axm t«o uax'ic fe&jds aj:© obaerved.
with a l i g h t band between thea .
fh i s grgmp haa only a. ^ n t r l e p a i r of st tbaetaoentrio
ohroaoso®© (ahroffloaoa® io«7) as ind ioa ted a^ the ceiitr0Eei.'ic
index arltioh ra,'iir@s fa^oa 23tOQ (Sable I A) to 31*^ ( fable I I Aj
pereerit .
Th-: sho r t uxm i s ireor o i a t i n o t a id a lso about auM the
a i ze of long aisa* clisse i^aiiger. for t h e short aim i s 1«19
(Table ¥ A ) to 1*85 ( l a b l e I I I A) ptroeti t of t o t u l haplotd
length* fri© r e l a t i v e also of whole ohroffiooome rcaiges from
4 . M ( fable V A ) to 5«y5 ( fab le I t A ) peroeut .
She iffhole sh:;-i.'t atm i e aackllpr sts*lnt:a, aeatroaifci-ic
region shcsws pale stainiae:;» j^roximul por t ion of the lon^t:
&im ahotfo £1 broad 4.BX1S, band and a l i f ;ht band while i n the
22
d i s t a l segXoxk one broad dark baM was obseznred*
fhia group taeXudes four p&Xsm of aerooentcic ciiroaosoaea,
from BOS* S to I I*
g.iaaMfta^Ml MQm 3 **» tio tra<9« of short arm vas ©bB«rved her©
and th® eeatromer® i s localised termlaall^*
*£h® relatiT© eis6 of %hi& Qhsow^&mie rsag#s froa 4*20
( fabl® V A ) to 4«59 ( 2aDl© IV A ) p«re«at,
fhe oenxroffleiio region and a small portion of proseHial
region are ooyered by a dai^ barid« Uext to th i s a l i ^ t baud was
observed and af ter that a seooad dark band i a freae&t i a the
same prosElaal region* £wo dark biuads are prosiinent Xn the d i s t a l
region of long aiia Mith a l igh t one in between*
QxiliOM-QSQItE IfO* $ s- the r e la t ive l e n g ^ range., of tliia
ohromoa^e i s from 4»20 ( 2able V A ) to 4*43 ( fable IV 4 )
percent*
fhree dark and two l i i ^ t bands were observed here* iM&ll
poaetion of the proximal rea:ion v i t b oentromere i a darkly ataineo.*
One li#(.t band in tbe proxiaal region i a eep^rated from aiioiber
pale band in ^ e d ia t a l region by a broad dark bm&d*
9 -
thoui^thi aOsatitlaiQei i t i a aoafmsed with a ©^^aetaoeatrlo bee;.:.i.3®
of the preaenod of v@r; islnutt) 0h.)£t ax^a i«itli a amUL sa te H i t«
t » on® ©£• tHe skor t ax*;u«
iielativ© l©»gth of t ae cli.i.-oaosoae x'.a>^ea firoii 4«y:
( fab le ¥ A) to 4.55 ('X'able I I A ) poirceiit. ^ a t e l M t e e , mxy
minute shiort .aj3a@ .md t h e oeiitro;«e£"io s?#gioii» i.iJ.1 a re aarkl^
ataiii©4« ffals broad 4£i-;k baaa exttiiob mpto ta© sfa&JJi. port ion
of proxliaal regioii of long a i a a lao* fkoaesfoire I t i e ooaol-
derod as tro darlc bands*
A bxroad pal© hmtd l a v«i.y ooaapiOtous i n th® ©iddlo
por t ion of loiit: a»:i# I n the^disital region two dure b&yado JAT©
pre a© a t with v« r j uuari ow paJi^ bcaads i n between.
aiiiiyMy^ai BQ.-llt - fh© irelativ© l e i i g ^ of tb© ohrosiosorae
raagea firoa 4.Q5 ( fab le V 4) to 4*43 ( fable IX A ) percent .
mre© dark and' two Hf-ht baiido a re qu i t e oonsplmma, unc- dark
ijand 0-avers the oentromeric i3ortion and s a a l l p rox iaa l por t ioa
of tiie long arm. A l ig i i t band aeparatea i t froia another dark
bmid ill tik© proximal reg ion . Uhe o t a t a l r eg ion of m& stiiroaosoM
shows a prominent l i g h t aad a broad dtirk band.
TalB group haa onl^- on© p a i r of oiiroaiossoaes tt iat i a
nu • 12.
9 '
gijEQiOjQHI S0« Xa J - AeiioMlag to the a,jcia 2?atlo wmeb i s
aeacli' 1:1 » tb is ehsomo&Qm® pale i@ i&#taoeatid,o la anap®*
7:m iftlatlire aiE© of the &lms% mm i s aliout 1«§4 ( fable I I . ) ti
to a*12 (. fdtbld I I I . ) p&itQm.% mM of the lonc^ ai» i s alao 1»'}4
{ fabl® I I , ) to 2t l2 ( '^tol© I . 4- I I . ) i>©£o«at.
0#jQit£ome] <!i iregion an ^@ll QB proiimal re^slons of botii Um
asms a»i l i§ht3^ stained* Dista l seglctB of eaeli iixm sl&ows on®
fills gz^ttf tuQlvt&em m%'W»a paiss of medium else
aerooemtsio Qhroaosomes ( 100* 1> to 19 )
OiiBQ^Oi llI 10 # 13 i# fhe 33«lstlv© s i 51® of th i s oteoffloae®®
ranges fsom ] *11 ( fable II14 ) to 4*05 C i'able 1 4 ) p@roeiit.
flu^ee pxro^neat dax't 'tends and tif0 34gtit bands vece
o 'been^ heire* On® dsafk teM im. tlie oestroaej^o vegioB suid
psroaiiml p>Qrtio» of long a i s i s sepmi^ted b^ a rmxy naxxow ILgbt
baiStd, froit a&ol^eir broad dark baad in Ifoe projslsal portion of
tbe long a ^«
yue bx-oad Mglit baod i^d syaother da£k bai d ase pxfeseBt
d i s t a l l s*
QiiaQlyJCMi; B0« M j - fae relati-r© iei:igtii of tiji® oiirosioao»e
Tariea from 3.04 1 table V4 ) to 3.59 ( fable i h ) peceeat.
Oii@ dask and &ne l i ^ i t band^a^e pxeseai: i n tHe psoxiaal
iregioii of the long aa^« Geatir^aexlo region i s also darkly 8ti;iii<.ea,
9-
iHro dark bands with a naj row i l rJ i t oixe Iti between ars v«£ '
grosilneat in th.© d i s t a l a^gion of '^© loiif.:
qj,ili( MQa3g .ilQ« 15 s- file re la t ive leiigfe of m i e Qlaoffloe*
Vi>£le@ from i.04 ( fable VA ) to ^t^d C fable X 4 } pe ro^ t*
a<^it3^Qeno region aiid aaal l proxiaal ^ r t i o s i are eovered
by a dark b£md* A l igh t hmad. i a also present in tHe pmsiaa l
r©.gio».
Sia ta l regidsi toegiua witii a e^ai'row daric ba..i<l aad eiids wit.u
a broad l i ga t baul eituateil i a t^e tej^lsml region of t^ie leng
ana*
aiiiiymiayMl au* 16 j - 5:he re la t ive eia® ef thie aoroeentrio
oiiroaoQoae i^ns«s ^ » » i#8t ( fable 14 ) to ^«16 ( fable IV A )
peroeiit*
4 dark bani eovere eentroaere and wtall portion of tiie
proximal region. A li4,1it biua4 i s in tlie middle and a darfc band
dietall;^ ia also ver ^ proalxie^t*
aifaQaa3QMI io.* 17 s* fhie aeroeeatrio ohroaaeoiae raages from
2.75 ( f&ble y A ) to 5.11 i fable I I I A ) peroeiat of the
iiaploid leo^tli lE e i s e .
fwo darli band®, one in tbe oeatroaerto and proxi«al regiori
and til© o ^ e r lia the d i s t a l regioa are separated b^ a broad
M g l i t baiQtd.
j iMjauaaMii m* IB «• :£iie reluldLve lei^gta af i^ia aorooea1»i©
oaroaoatme variee froa 2.7Si ( Sable V & ) to ;$.ll C Sable ill A)
pereent. .
4 broad daxk l and eoveim the oeiitR^aeailo as well as tiit
ifiioI« fsoxliB&I i^gioi^* l a %h0 d i s t a l £@glon one &arK a£id tii«
otbes M ^ t band V9se obstrtred, the dask o&« i s si tuated
QHRQMyiMJME ^0 , f§ «• fhe i?®latlir© length of th i s lemll ao«®-
eent£ie ehjrciaioeotte jp^gee from 2*57 ( fable IV 4 ) to j<*ll
( tumble HX 4 ) p9T@mktm
A eiuill da^k ^M LB present in %m oentsoaexiie and
jproxLiaal ^egiona* A M ^ t band was also obeerred im the p«»ji.fflal
region* Dlatall^ there l a a |»roiiine».t dai± hand*
aroup eeT0)ith oc^aprlaes onl^ a ©in^le eia&ll pair
( ho* 20 ) of ehromosoffiee*
gJllgiaai^Ml fio* 20 8 - iieeauee the mmk xm.tlo i e nearly 1:1
arid tlie oentromerio IMen^ i s ahout ^ ^ereent the ohrc»oeasi®
pa i r l e ©eteeent^e type* Shie pmlr i e ©iatalcea as wa aoro-
oeatxie 0£ie» ( B*&*?« Eao asid Sharda 1964 ) beommee the two
short aj^e are geaerall^ not Ter^' olearly eeen in poorly etained
preparatioBS*
fhe relati ire s iae of the ehort era raage® from 0*79
i fahie IV A ) to 1.43 { table i 4 ) peroeat a*id the whole
rel&tiTe length of eiiromoecme 'V&riee from 1«!»4> ( 'liable Iv & )
to 2*36 ( liable £ A ) poreent*
fhe ehort arme are ©taiaed l igh t ly iliotfing no dark baud*
A broad dark band ooyero oeatromesio region aa neH a® the whole
loa^i aim*
£ifl^h or tHe l a s t gromp includes six pa i rs of -rer^ small
aeroo^atxlo oftiroaosoises a@ imll aa tti© sex Qlirom0sc»SM j in oad@
of th9 i&al9 anlMtl.
g ^OMOiliHB. 1I0« 21 £• fills aaa l l aor&o«atiele oi)£<»sos9ate v&riee
itt »©lative l«ngtli from l.^S ( fable I¥ A ) to 2.38 ( tabl© I A )
pero@»t» fh« oetttKOM«rio mglQu i s l igli t i^ stained* A uxomA 4srk
band eovers t^e pEoxXtml a® well as tHe d i s t a l segioua of tlie
long
^.ii^:iQaaip UQ*_ZZ I » flie re la t lTe leiigtii of th® elijeoiaoaoffie 22
ranges fron 1*^8 ( fable I¥ 4 ) to 2*3B ( fable X A ) peroeut*
.4 siogle l i # i t band oorers t^te oeatroaexlo regioa and a
oaiall portion of tlie proximal regioa* East of %m perHon of
proximal region and iibole d i s t a l region i s oorered by a single
dark band*
Q aQMOiJaiB MO* 25 : - The e lse of t l i i s o^eaosoae ooatributes abou
1.58 ( fable I¥ A ) to 2*53 ( fable I A ) peroeat of tiie iriapioici
set*
A dari£ band eovers the wiKile prosd^ial re^o& atid the c^iitro*
{fisrio rsisioa also* Distal l^ taere i s onl^' a broad, l igh t baz»d*
Qmx^.mME. MQ. 24 j - file r e l a t ive sis® of tiie oiiroaosoae varies
frosi 1*19 ( fable la ) to 1*73 ( £able f4) peroent*
ijandirig i s aot clear* Some plates siiow eeatroaerlc region-
l igh t staicved aad the whole loot; arm dark*
9 0
%% 1«73 ( ^ b l » IV 4 and fA ireapttdtif^ljr ) p«s««iitt of %txM
HapidM set aa far as i t s s i zs l a oixieo^iied*
ftis osatxoasrle i^^gion i s l l ^ ^ t l ^ stidLaed aad immX Qt tits
IsBgtliL of li&m& mJ» shovs Sasl^ stmlsilmg*
aiHiOMCiaoMI iQy ^6 : - $lis ceXative al@s of t h i s mmlX st^
( fabXs ¥ 4 } j^esQsatt
staining*
.1 ^ 4 P ^ a ^ i | s f ^ s i s tH© sliortsst @l»saos9iiie ©f Vam
esapX«ie&t with m seXm^re s i s s v&£i@tioii f£0@ X«if ( fable lA )
to 1«73 ( fabie ¥ A >.
ftie vholtt ¥ slivoaosaiae i s ligiitXir stained siiowiiig iis#^tiire
heteropf oacxdty* In siost of tlie smM@ lisj^^ss ant ejd^eiiets tlie
A oii roaosoffie shovlxig si.«gativ® iietezropycmoaity i » spexnatogonial
^etaplmses was reported bf illiiie« H»^*i)« ( X977 ) • In Aoheta
domestlstts £ shxoaosome ah&ws negatlTs tietevopyouositjr ixi epersa-
togOBial metapliases** MoweveSt i n soemtie oe i l s and Oogonia %m
X omronos^^e «&@ xrepoirted to show isap^r^snoedty ( Sliez* Ail 1979 )
Altliougli Tajdlatiozis Xa %tk® mm&mx of sly^daososMis of
^at-^:ttf nQfysffimtf have been i^epo^ted ixi Uteiratuurt the 2u iia«
of elm>«9seaes l a tlie pjfeseut s tudies was detezmiiied 4B axi tm
9 0
tiasla of Z'y v e i l spseadi ndtotlo plates*
the aulosomta ««£« sctielall^ »i«li«£«d fjeom 1 <» 20 In a
deaeendi&g ord«r of l9zigth EB4 the sex clirimosoaea ase sefei^ed
to as X and y« Mem^le milm&X l a x x wiiile oale I s betesogattetie
and sliows x aM j ali£oaos(»Be@ i n i t s 6Jfy?cttosom« oamflcoie&t*
flies® 42 oiii:tomosQae@ Haire Iteen o itogeidLaiMl to s ix gso«#e
aoeeMlmii to t ^ i s etojpe ( eentrosieirio f03ltl03&)« aJlse and i»
emudiiig pat tern iMuoed liy uurea and t£^peixii« I'M two types of
treateenta l&dueed alaost eiallair type of d ^aadla^g imttexii
( Xdldgsaiis I B *• II ^ and Til ^ •¥ tV B ) tn a l l the etooaiodeaes
ae in the oaee of aqtiiixel*
asoup X ooapxiaed only one p i i r of ehx^moeoaeet abj;<moa<»e
atpoiQaOME ii&* It s- me «e»tieoaeEio index value 'miriee tmm
14.28 to 20*0 ( fable H I B XV ii -»• V is ) showing ttoie ohie<»ao-
aoaie as eubaetaoentacio type* the i l l a t i v e length of etiozrt axm
varies from 1*19 C fable I B ) to 1.90 ( fablatXI^ ) pesmat
euid tiiat of tlie long aasm fTOa ?«1§ ( fable X B ) to 6*04
( Sable XI i» } pecd^it* fherefojce t^ae irbole ohvoaoaoae
oontxiljutes about a«^§ ( fable I i> ) to 9*7§ ( fable XI a )
pesaent of the haploid set*
The proximal regioa of aliort axsa peeaeaaea a li l igh t btuid*
!l^er!3iiially there la a proeiiiieat da.rk btuid mid tHe oeDtrotterio
region l a l igh t ly stained* flie proxim&l region of tae lon^:
ana show© one da3?kt one ll^t$ one dark mid on® l i g h t baud
sespeet iTely* D i s t a l l ^ two daxk bands wexe obseirred with a l i g h t
one i n between*
This group iiiolud^s l a rge aorooent r io ehromosoie no* 2«*9
( autosoiies ) and one p e i r of x ohroaosoeies i n f^oales or a a ing le
X i n males*
u ^ u a y j j i ^ riQ*^ £- Shis ehroaosoae i a the l a r g e s t aoroeMiiitcio
ohroQtoeome havisig a r e l a t i v e l e n g ^ vmriation fro@ 7 * ^ (Sable
IV ii } to 8*58 C Table III U ) poroea t .
A dark band Im p i^eea t In the oetttxomexXQ j^gioi^* '£he
proxLmml region ahoira tvo l i g h t bands @@|>arated h:y a dark one*
lui the d i e t a l region two 4.m^ baada a re separated by a l i ^ t bandt
aiiaaMQaOMS no* 3 8- Ohromo©«me oo* 3 has i t s oexitromere a i tna ted
almoet terminally ' aeeiayiaing mx ao roeea t r io ^ a p e * fhe r e l a t i v e
J^ngtii of t h i s ae rooen t r io oharoffiosome ranges from b*Q9 (i 'able I I B
to 8*58 ( f ab le I I I i> } percent*
'ikw pentroiaerio region ehowe a dark baiid* l a the pxt^^imal
regioii tvo l i g h t bajada are separa ted b ~ a cLdL>::i" one* i ) i s t a l l ^
there are two dark baxida separated b - a broad l i g h t ba^id*
9iiJ^Wi'^^M.MQ, 4 ; - fhe r e l a t i v e e l s e of t l i i e l a rge aorooetttr ie
ohroisotdsoBe ranges fa^m 6.09 ( fab le II JH ) to 6.90 ( Sable I a )
peroeiit*
'i?hree dark aad two iijfht bauds were observed on theee
31 cliiroi6ososs©0. CSentroEierio regioa showa o»® daric band* i'ra:3£laaJ.
region has one l igh t aod on® dtirk band respectively Mid th# d i s t a l
regioE also ahovis on® l i ^ t audi one ^mxU hmdm
A aiiEullQJQMS. i- The % ohroaosoa© oo£itributes about 5»56 to
6 * ^ ( Table X\r i» ) p^coe&t of tii© Haploid s e t .
fhe Ijcuidiiig patteim i a so t ver^ d i s t i ao t iiere* i^^oxiaal
region aho^o onl^ a narrow l i - ^ t baad* flie eentros@rlo i^gloti;
re^aaliiiiig proadUial region md. ir^ole d i s t a l reg^ony a l l are darkly
stained ahowitif two daric aM a aijigle l l ^ t ^axid o^ tM.s
ohromosome*
a.,4iiaaOJQMI i<0> Ss.*' the relat i i re leugte of t h i s ohroaoeoae varieg
froia 4«77 ( fable I B ) to 5»72 ( Sable H i ii ) peroent, Tm
c&'ntroaere la looalieed terminally giTi&g tlie o^onoeoae an
aoroeeiitzlo shape*
fliree d^rk baads are res-g froslaeftt One i s %m the oentro*
^erio regiont one l i | ^ t and otie dark MtiA i s looalieed io tiie
proximal r e ^ o a of loiig as^ wblle the d i s t a l region shows one
M ^ t aad o;^ dark tend respeot^irel^*
g iaoaaacMii iiQ> fet.*' fhla ohroaoeome oontributee mismxt 4«77
( 'fable I ii ) to 5»72 ( fable l i i B ) gereeat of t^e hapioid se t ,
the oentroffierie regiom &Mwa oi e darlc baM* fro<^®all^' one
l i gh t liBMd was obaerrt4« the d i e t a l r e ^ o n ebows tuo <lu rk baads
separated by a U g a t one*
ifiiiiOm . Ml l u» .7 J- fbie obromoaoae ooatributes mbaut 4.77
( fable I i* ) to 3*12 { Sable I I I ^ ) feroesit of tlie impxoia o®t«
3,
tix£@9 dax^ bimds are psoaineat heie®* One i s in tiie eeatro-
merio jcegion separated froa another in the pxoxlsmX region hy
one l igh t band* ^tBt&X region of ttie 2a&g a ra poaseeses two dar&
bandit, oiie l igh t aM theii msiotlier dark respeotiirel^*
aijauaQJOMfc fiu* i. s«> the ramive siee of t&is ohroaoetMe varies
from 4*77 ( fatol© I ^ ) to 5.40 ( fabi© If a ) percent.
flyree dar^ and two l i ^ t bands were obeerred here. One
dfirk band in the eeiitrosierio region i s eepaji^ted from another darM
btmd ill the p^coxlmBl x^e^on, hy a l i gh t band. In the d i s t a l
re^Ofi one l i # i t mnd one daric bond i a@ observed reapeotivel^ •
g AiiOSy Mli MO. Si" i'his ohromodome ooiitcibutes about 4.77
( fable X ii ) to ^.40 ( fable IV a ) percent of the haploid aet .
fhie ohromoso^e also shows three daxic lyiid -^o l i i h t
oands. One dfixk band i s i n the oi^ita^omerid re^em. X^ the
proximal region tliere i s one l igh t and one dark band reepeotlvel^t
and a i a i l a r l y in the d i s t a l region ime Ught a ^ one dark band
was observed.
Shore are ^ r e e pairs of mediua sized metaoimtria
ohroaosome ( 10#12} In thim group.
UiiiiUIIQ^Mi; My. M i» She a m r a t i o of tMe ohriMoaoae i s
neara^ 1:1 and oentroiserio index i s about 5Q p .o . indieating
the ohroaososie to be ^etaeentr id . Eelative l^ igth of l&e eliort
ajasa ranges fro© 2.09 ( Sable XII B ) to 2.40 ( fable I I x> )
pere«it and that of l&e lon^' arm ranges from 2.09 ( fabl© Hi a)
to 2*40 ( fable IX B ) peroent.
k broad da£k band biseoted by emutmmewe was £tga]:d«d
as two daxk bands C After Qaepereeofi e t al« 197^ )• ^ ^ psoxlm&l
regions of botb tiie anse ahow Uglit basde* Diatal l^ or ra ther
tesdnaXlsr m.9h axm uhisma one dark band*
a mOMQ^MB,, 9. 14. s - fHis oUroaosoiBe a lso haa aa a r a rati©
eq imJl to Is I , aad a oentromexlo iMex value of about ^Q peroent
Indieating the #ir(mo©aae to be aetaeemtrio type*
fhe re la t ive length of eaeh asm rangea from 1«90 (fable
I£X i ) to 2.3a ( fable X B ) pere«:it.
fhe oeatromead^Q region and tbe fXN»Jslnal r edone of betb
the arms are l l # i t l ^ etalned* In the d i s t a l region of eaah a m
one daric band iiae observed*
QHaoifQpCMl HO. 12 s«» She a m ra t io i s a ^ d n I s l here* She
short arm length e^atzlbutes about 1*90 (fable I I I B ) to
2*19 ( fable XI B ) peroent and ^ e long arm also about 1*90
( fable XXI & ) to 2*19 ( fable XI B ) percent of the t o t a l
haploid length*
She oentroserio region shoirs a broad dark band biseoted
by oentairoaeret therefore o<msidered as two dark bai»ds*
Proximal regions of both the arms are l i # i t ] ^ stained while
the d i s t a l region of eaoh arm i^oirs one dfirk band*
fhree pairs of mediiua s i sad aorooentrlo chrososoae
( noa* 13 to 15 ) mm categorised l a tM.0 gromp*
31
g iiiUMyaail ,li « 13 s- th@ r e la t iTe XmigVaL of tUla ctijsoaiosoa©
vari®® from 3.80 ( i'abl© I I I ii ) to 4.59 ( f« I I ij ) pej?o©iit.
i'tiree dar^ and two l l # t baMa ir@£# obserred on tbis
ohrosiosoae* fhe eeiitjcomerlo x@^on possesses a dmck band
wbloh i s separated from the o-^er dax'& hmd ia the proximal
region hy a l i g ^ t band. 'Bte d i s t a l region of loa^ mxm shows
ono ligltt BXiA 0»s dark baud rs8psoti7@lj*
g.ktoay^yl'iii; J^Q* 14 s- fto® re l a t ive Isagtfa of tli is o&raaos<si®
ranges from ji«73 ( fable X¥ 1$ ) to 4*^9 ( fable I I B ) peromt*
fbe oentromerio region ataitia l ightly* Que dari: band and one
l igh t baM i s looarlised in the proicimal ce^on* the other dar^
and l igh t band i s preseat in the d i s t a l wmgJLQtk of loa^ arm*
gAJtioyaKJIll MO. p I - Uhis morooentxlo ohroaosoae oontributes
about 3*33 ( fable I ii ) to 3*90 ( Sable I I ii ) p e r o ^ t of the
hsploid length.
Here also two dark bmide were observed* On® oovers the
oentxomerio re.gioii as well as the whole proxiaal region axii.
the other i s i n the d i s t a l region* i^etween these two there l i e s
a narrow l igh t band*
^roup f i f th includes four pai rs of ohroffiosoaes »
nos* 16 •» 19*
mimmmjViM KU* M J - I'he ara r a t i o being I j l aad the oeutroaerio
iadex vmlue ( ^g peroe&t) show that th is i@ a aetmeeAtrls
80
to 1*701 ( fabl« XX ii ) pes&ent aad tiao JU»Bg aa^i about 1*32
( f&bX« XXI M ) to 1*707 ( mhU XX B ) fexe«nt of tlie t o t a l
hajplold l6»gtli*
fh^B oe&tromtxdio re^^oa sibtova one bSQoA ^ask IMXA kia«ete€
in to tuo das^ bands bar #»nti:offie]?«« Fsoximal x«giona of both tlie
acao utoif l i gh t bani whllo thes* l a onlor one daslc baM in ^ a
d l a t s l zre^OB of aaob asn«
q£a4llQ;l0p ao# 17 s- fUe r e i a t l w lengtli of tlia mhowt asm varies
fson 1.33 ( fabla XXX i> ) to 1*67 ( fabla X M ) jpajrottit of t&e
t o t a l liaploid l e i ^ ^ a»d tbe loa^ aim also vaa^ea fsom 1«^>
( fabla XXX ii ) to 1«67 ( fabla X B ) ^eso^ t*
Swo 4aa^ baMa asa ps<mitt«at in lAia t asa lna l za^ona of
both tha asvst ona in aaahf aantrotaexia and psoadaal saglona ica
l i gh t ly atalnad*
airnQMoaQIH MOm m i* tha idea of ahoirt am of thia sMtaeantslo
ohrosioaoaa imngaa f:eoai 1*03 ( fabla X9 B ) to 1*46 ( fabla XX M )
fexraaat of t o t a l haploid langth and tha long asm alao oontsibmtaa
about l«oe ( fabla XV <i ) to 1*46 ( fabla II 3 ) fa]»»ant of t o t a l
haploid eat* aant3?oaa£io saglon and tha ^i?03ciaal portiona of
both tlia asma as* l igh t ly atainad* fha d l a t a l leagion of aaoh mm
ahova &m 4i&xk band onily*
atjiiUMaaftMI Ha.* if- t# fhe ralati ira length of ahost a»a of th ia
aataoantida ohz^aoaoaa xan^aa fson l«Oi ( fabla Xf ^ ) to 1*46
( fabla XX ^ ) j^araant a id long aasm alao jrangaa firom 1.06 (fable
IV B ) to 1<.46 ( 2al3l« IX B ) peretnt*
fwo iBisk 1»aads are p£«8fiiit on lafte tiihsc^oaciie In %h«
t«£iitlniil ireglon of 9m^ earn* m&t of l^e oluKmosoffle lnoJudliig
oentar^nesie sw^om shows l i # i t ba&4*
- iix:m.p s ixth ooapdLsss of oas pais of elixoiiosdisies i'Eo* 2y )
and ths y ohxomQm&mi i a aalji ra t s*
9aMii/^9il Pft .g^ , |* fhs «mtroiiis»« 1© ioeaiissd ImxAxmll^
giTlBg tlie ohiPOffi>os<»e an aocoesntrio skiaj^* fhs ehroaosoMt
oontid^&iitss about 1«@0 ( faliJls I ? Ii } to 2*43 ( fahXs IX M )
fsx^ss&t of t o t a l hsplold length* the m&t&smmsiQ s e ^ o n possosaes
l igh t haad aa4 s s s t of th« %0m^ mm shows dasie s.taiii4i&iE*
JU2i iMjML.»* ^ ^ 9 ehromosoai© vajeiss twm a#30 ( fahle X i^ )
to 2*45 ( tah ls ZX B } posoo»t of ths hsfloid longtb Isi sies*
Ixospt the sofitv(^Msie ssgioa ths whols c^mornQmrntt shows
datk stidLnii^*
In gsaojeml ths masbsr of 4 i s t i ao t bsnds insrsasee with
the ineifeiuie in lexi^th of tiie ohronosoae* t h i s was observed l a
AaaphMie pla tes ( liable ?I Aso^ tabis fX ii ) aM also Xn prophase
of the meiotio ps^spajmtions C fhotogi^ai^ VIZ A mA VXX i; )•
the msaber of bto^s tm a ohjeroBK»s«;»se was obserred to rmsg
with the s ta te of ooati^otioiif the tmalit^ of prepaidtioa and
the t^^es of treatment aad stain* In the present study trypaia
t rea ted preparations showed more shrinkage and oontraotiony
87
while wmB. t sea t td prepai^tlo&s did w&t mh&v aueli #ff«et of
urea OKI the e irciBiesQrae Xeagl^*
A ato^dy of the meaa selatiire lengthy aean oeuti^^aezlo
indieeef %JSO^ assi l^o^tiis &i^ shoirt ftim Xisng'^s of the
oh£oao8(»Mis have heea eunMi.sieed in fablee Ik to tl k BM
I B to VI B» Xdiogimiaa ehonisg the ee&tsoaesio peeitioiui>
Q B-an^fig pat tern aad Xea^thJi of the ttto ame of the ohroaoao*-
ae of both the animal epeoiee have also iMien aade to aalce the
ohaervmtioae laore aeaziin^Tttl*
Pk g M th
u 6 m o «
§
es 4»nd is ^ Ol-I su. • »"Sv » K • .
"3 %<§ 4»4» » o oeq P fl p <8 93
H M IS M P
g® gS*o
^ H « i QQ 4f tt tse o -fi P > 13 O
sas" «»«»«r«»i | | i« '«> • > » * * » «
« - B A O 4» m
la « l4 psi!;
M tt IS o<H tt ^
« e o u §1 h ^m g
M M b * M H *^ N
C O O O
> ^
H H <y» si €^ m tes ch
• • • 1 1 1 1 • f t * •
o o o o •H *! M "H
^ S g 8 Hi * * * * i ' * ' ' o o o o o o o o o o o o
• « • • • • • • • • • •
o o o o o o o o o o o o
m o ^ 1 1 1 1 1 1 1
•1
o O 1 1 1 1 1 • 1
O O O O O O O O
O O O O O O O O
-H +1 -H -H "H -H +1 +i -I-I -H 4«| -H 41 +1 +1 -H 41 44 • ! "H
• • • • • • • • • • • • K N O t A » A © e O < D e O
^ "^ m tr\ O O O O O O O O O O O O O O O O O O O Q I •
o o o o • • « o o o
• * o. o
o • • • o o o
o o o o o o
^ » o o o « ^ - ^ ' ^ - K l l ^ i < ^ i < ^ K ^ H O l A u ^ b b b e o
K> O • o
• «
e4 m • H
• H
U> o
• o
»*% Ol O • o
•1 41 H m « H
•
W
^ •
H
• H
1
1
« l«^
1
1
« •#•
1
1
« u \
1
1
• ^
H - •
i o 41 Ch fi It
H
-• t -
1
1
* <0
1
1
-« Oh
1
1
• o 9"^
1
1
•
a
GO O
o 41 M H
• «M
• CM H
H M O • O
41
tf\ O
« ^
• K\ H
1
1
• ^ H
t
1
• U\ H
1
1
« ^ H
1
1
• ' C*-H
1
1
• €0 H
1
1
• cr* H
HI ^
g '
o I I I I i I I o • I 8 2 I I I • I I I
23 gj <?\ «A £^ « ^ •'N CM M l^» <M % o o o o Q o
o o o o ^ o o
(is pN 9s HI ,1^ ,»| H
o o o o o o o o • • » • • • • •
o o o o o o o o
• * « * •
HI M M <M '"T f**? i"? J'^
O I I I I I I
• t i l l
© H W l»N 'tf' tf^ « f t f ( M f t i f > t C V < M M > - (
%4 O
m
I «
M f l o i d no* §4«
40
X
«
s m
w
K
it
lO • • •
S
2
M
Ul
1 O W)
O
2 O a: X
o
•A
f I I T - r " r — r • N «) lO ^ pj f^
H i O N 3 1
• - r I f 1 1 -T—1—[ 1 1 T f "1
3AllV13d I N J O M J d
41
4» 0
o o
o
i f M 5 s 1 1 I I I fi I i I I I I I I I I I i I
8 ' * SJ
I I
I I
o «
+1 tA
I I I
t I t I
o • I o o
i tA M O
o o 0%
o 0%
o o o
M -^ M is* *^ p : A* S4 o f >
] i •M
"* CO • •
CTk O H • •
o o
•••t 4-1 4-1 • ! HI i>* ^ SI «*! o
• • • ^ \>0 ^
o o
-3 <3F o
80
o
in *» +1 "H +1 • ! • !
5 O
M O
o
M »A O o
2?
H tA O o
tA O *
^1 • !
<M O
o
« tA Ch lA O • •
" * lA
•l-l +1 *l M +1 4-1 -fl ••'I "l-l +1
o H tA O
o
0% o rA
«
o O tA (A lA Ch (X%
i • • lA ^ ^
* lA m lA
O eA (A 4
tA {A
^1 • ! 4.| +1 eA 5A
tA < ^ tA tA
• • m o tA
?A iA !A q o o • • • I
O O O
' " • • • • • • • - • • • . . • 4
«A lA iA ® C0 CD * * •
0 \
o I I
t i l l +1 tA
4
o
I I I I
I gg
• • I
tA
r4 CM Mk 9 • tA \0 CO m t% H
H H fA
o o Cf d O
S I S I I I I S I I I I I S ^ i i 8 i i i ^ ^ ^ " ^ '
o
o
a
s o
g 3 » y o H S ^ <4 <«j s? «3J «ij '«^ S "S "^ -- -^ is<j <*5| ^af
I I I t I I
I I I I t I
nj
o •I
• I
I I I I I
R t I t I I
O o
{ A
O o
t
o
m ri 0%
o 3 H *
O o
t
o
r-o H O
• I -H "M -l-l 4-1 "*•! +1 4.I • ! +1 +1 4»J 4-1 +1
42
<y> {*• p * o m as • • •
?^ CI OJ
ini w a \ Q c* ys>
• • • M 0 * «M
O O Q «
o o
o
lA M
a 3
«^ il? ^ d} i 9 CO
f t • f^ <*i <^
t ^ t«% !«N M CH W
• . « • H H r l
HI O
£9 M
a
0»
«
r4 HI
o o
•H 4-1 • ! 4-1 * ! 4-1 4-1 -1.1 4'| 4'! -H 4'| • ! +1
• • It '* * 0 \ q) f »
lA H tA 0^ ®
O * •
r4 H
KN l<\ r s l A e4 (M d M
4 ft . • *
f""§ i^*^ I " T (
1
1
• •
i4
1
1
•
19
1
3 •
S
t
1
•
a
1
1
•
S
- • o 4-1
•
• o CM
1
1
• H
1
t
•
<M
t
t
• tA Al
» .
t
•
1
1
• tA
1
1
•
CVI
1
t
• !N
m $
8
$
Hia?O0MM IIA* M9pT9&9ntB the 9 \mnM,ng pattsza liidue«4
li^ UV9WI, l a the kaz^rotype of gimismbiiltia
4.-;
>CZI >
X
in
It
o CM
o>
S
r>.
Sfi O
z
S O CO
CO O - 5 « 2 — Qc
I _ u o
C3>
GO
rv
(t>
in
TO
CM
oo — I 1 — r CO i n «<•
— I 1 r TO CM — O — CM TO
—I 1——r i n «3 f^
00
H 1 9 N 3 1 3Al lV13a lN30d3d
a
9
9} 4
I -'5 .'3
H
3
I
^
<M
H M
43
o M
a ra ja "3
^ •»
M
^ ^ ' » g t4
64 ta '^ -o ':^ *3 -" O -••I
, r.> ag -5 o »-3 «<4 '^ > 5?. -23 <4
M.I sa
re?
''A 'A ,"3 O "4 ';» C* M W "- ; i •<«", JH f'sl
tij ;5| 8t| '2 -;5-,.5: m o *i '0 S-1 lag
O sa .;33
ai Q n»
CI 9 O l>
O O «>
^ | i -P c> ^ o
ittttttitttt iiiiisiliiH i i i 8 2 s « •§ I I I I 35 as 0J «3j -<ij ^
K\ OS O^ M «h H o rv o o » • • • •
o o o o o 4>i 4^1 4>1 t I i t 4>t I t I t 44 » O N O O M O
• 4 • O M UN P5 M ir\
f I i A
I t I
I I t
t I t
^
o o o o o o o o o o o o o o o o o o o o o o o o o o o o +1 44 +1 '•'I 41 +1 4-1 44 "H 4-1 4't 4-1 4'| 44
<rti-l H t f \ < M H i n f i h « l » r 4 f l h i < %
o o o o o o . o o o o o o o o o o o o o o o o o o o o o o 44 44 4'| 4>l 4-1 4-t 4-1 44 +1 44 • ! 44 4.| 4-1 ^ ^ i > i A t A t < * e > - r 4 ^ 2 r < > t < 4 > < > l H
• • * • » • • • « • • • • • •
o •
o 4-1
• KS
o •
o 4 1
9
o « • *
o o •1 4-1
O HI • •
o o 4-1 4-1
• •
O • • •
O O O
o O « o
o
«4>l 4-t 4-1 I 1 1 I 4 1 f t I I ^i t
• • • r4 H ri
9. I I I i t « » ' 'c4
t I i
» t I
CM • • • «» ir>
m q r4 cS H H H H r4
•n 3 3 S4 l a *
3
m «M»w<i»ttF4!K>
•I
• I 0?
2 M ^ SS M M aS JH
i S f I I & I Q § § ! • § § § §
I I I ^ , i t I •
o O
1 1 1 * 1 1 1 1
€h t - «M N N m «
o o o o o o o o H -H 4>i -i-l -H • ! 4-1 -t"!
d d d s f^ ^^ 1 ^ w^
i«\ K% m ^ ( 9 vO
0» £^ W O* « -«• © <P< K\ t«%
o o o o o o o • • f • • * •
'O O O O O O O
0% 1 1 JA H Q
O
d d ^ 0> ® l*% 1*^ <f* CI'* 0 * <o CO 'i©
K S f ' \ « n o H I ' - « r l H
I t I * I I I I O
• I I I I O I I t I
o
• • •
u s s i d tM US CM 04 M
I I
I !
I I
I I
tTk
o o o
• •
a o
• I +1 • ! * l <R. • ! •••I -H •••I 4-1 0% 0% M» «4I>
t •
f I
I I
(A \ 0 fvl (M
43
%4
o m 41
8
o
1
4» sS
43
to
(9
n
S
St
«»
CM
'T" T r" r K «» tf> "»
MiON n 3AI1V13W IN30W3d
e I s
I I U It
5 S
47 o 3
0 ®
o I s
fa M M M
o
us
o o * •
o o
o o • •
I I I I • I o
I I
s ^ S 0 o •
its <•
8
8 S • •
o o •H -H •»§
O o
2 •
M
0 H
« 0
•H UN
• ^
fV 0
• 0
M
S •
«
o o • * o o
l> irt 1^ H H «^ 0 0 0 0 0 0 0 • • « « • • *
0 0 0 0 0 0 0
¥\
s o
in rv
r» o o
• « • 0 0 0 ' H ' H ' H ' H ' H ' H ^ ^ ^ ' H ' H ' H
» CM CM 11% a «^ 1^ in CM W O •<•' »A « • •«#
• • • • • • • 1 0 lA 0% rv •# ««• • r
1:? 2R ^ • « •I r\
»_J
4* q4» SAir fi « N pQfl • H O H .
• » O
r O M
o
lO O • •
10 o
* • •
CM
o , o •I
I I
f t t I
O o I I I I
l i l t
^ I o
4
H Hi H i«4
H %M •
1
1
H IV> •
O •
o
H H •
t
1
H O »
t
1
<^ •
1
1
0 •
1
1
-4 *
*
o • o
<f> •
1
1
vn •
1
1
4k • •
1
1
VJ) •
1
1
M •
1^ •
o « o
IV} •
•
o •
o Ml
H «
o •
o
VM • *
0^
! •
o • o €B M
m H
o • o w
•«l
N o • o 0* VM
© o © • • • © o o 00 ^ C39 V) Vt "U)
o i
8 O * P H
1 IT >• o » <» o o o
o
€•" M € T H »* « J* CI I *
M Ml
\ i i Mt «i> • • •
t^ t^ H OS <Vt * 4
!• N- I* N-©
** %Ji Mi \n %M • • • • • M l # j O W W v|» G8 M l IM M W H* f^ I4' W* !• §•
O O * • o © S V0
o o o o o • • • « • o © o o o H H ftl M3 Ml V£» 0« Ml Mi
• • •Sfc M l * • V« O MS Ml
M! Ml M« « tt • O IM M M l IM N l H ! • !<»• N> N>
©
I I
I I
o © © ©
MO
• M M l
© © ft IM
© o © p o © ^ M l M l O * M l M l
I I
3 » f *
jJB,
0
i 0 «»•
^
»
8 3 0 «*•
f»
c^ ^
i m
s § CI
I I
s & &
o
V O
»< o
I
f o jar «•> ^_
8 0' H # H « SI
• ' ^ laHui
4 :
«c»
10
m
M
f^ m m I III | i iiiiiiii^ii|i»> r|
tf> « fs 19
H10N31 3AI1V13W lN30a3«i
HI3fOQEAII ZXZA- Ettpmsents the 6 banding pattern iMuo«ct
hf uxea ia tli« GlisociosoBes of faaf iapbltta
iMHtnai t waMitifilitoa ( f«iBale )
t B B B B ^ B t t t t t t t % 4»
i 8
5 " I
4» «l 4» 4* S *3 S 0 "" 9 «
I t
a 8
I t
8 4«
I I I I I t
O
8 a
I I I I I I ^ 4» 4» 4» 4» 4» « 0 • e f « O II 9 O O O s g g e 8 e *«
B S S S 44 << •< «$
I t I I I i
I t I I I
o •4 - I | i % <f tl> u>
» O o » o s • • • • • w
> o o o o o
H r4 O ffl i'^ S* *»
S 1 K ^ ^ ^ 5 o o o o o o o
• • • • • • •
o o o o o o o
• « • • • rk rk K% ft! w
« £ » » s a •"f f"! <P*f (P"! |'''f f f
o o o o o o o o o o o o o o o o o o o o o o o o o o
• • t • • ifs it\ if\ C4 e4 IV • • •
o f-i H • • • •
I""? I'"f #•'1 ( F T
I I I I i I I t l i l t
t I I I I t I I I I I f I
H H » - l H « - I H M M M < M C V t ( V < M
43
%4 0
lan
1 e
HZSfOOHAH TA • fiepxesenta th« 0 banding pftttoia iaduoed by
txiyp9JUi ifi th9 ehxososoffids of fanqabujlufi
» I
8 CM
CO CM
CM CM
CM
O CM
95 O .. z
JO
tn
£2
O
c»
QO
lf>
CO
CM
2 O if)
o
O Q: I o
i I I I I I 2 o) m rx (o ir>
1 r f TO CM ~- O O _
H19N3T 3AllV13d
I I I I I I I I I C M P o ^ i n u > K » < » 2
XN30M3d
m c
i ^ .'SI ft
S ^ • ^
s 2
i
S3
H
3 a £ 4 |
i' i
»••»*•«>«••••'
3 41
» » 4 :r! 3
.15 * v > ^ 3 t -4 > ^
IB ;i Q S3
3 1 ^ 2 5^313 0 t'-Ni PI* fe^ '^ &* r .:>
< o
^ ^ --s? 2 ^ •'4 ^^ . i^ 83
'.•3
4^
••| aj q i sa "JO >
fil M jfl *» KJ -f« /~|
8 S4 "ie<
'•?3
O^j
r> c-i »
o o o
H I I
t
I
I I O O O O P t I C I O O O O
§1 1 t i tt 1111
3 g si d g g d « 18 I I I I
<4 Vi m-^ '^ ^ M • 'm
K>
O •
o •1 e«-M •
e* CM
m H o • o
m H •
c# •H M <M
• M M
« «A O •
o
l<%
3 •
o 4-1 CSl <M • Oil
IM
H K\ O
ft o
9 ^
I I I ^ I • I I ^ t
• I
t i l l K " ' ' 5 '
• » •
o d •H -i-l 4-1 +1 H
ft • <0 vO
. * •
ft • •
e4
O Q • *
o o
H M O* 01 tf\ lA y> t» tA ff^m nts tf\ m m O Q O O Q O O O O
ft'ftftftft'ft ft ft ft
o o o o o o o o o •H 4>| -H 4-1 4>t4| 4-1 ••t 4-1 ^• SB - * Q O ' 4 A lA «^ 99
o ^ rk Hn o o MB f-l ft ft • ' • . • • ' • ft ft
0 * «t> « • ' • «# {r% m
+1 -l M H • !
o o ft ft
l A tIN
<5\ 0\
ft ft lA a\
ri , «VO» M m lA <iA ^ . H K% uf>tf% l A l A »n o o o o oo o o o
• • ' f t • ' • • ft ft ft
o o o o oo o o o #1 •! tt 4-1 HH 4-1 •! 44 ft ft- ft- ft- • •' ft •• ft
Ol *<• ' T ^ ''9' f^ ft
O - O »
o
„ . o ft ft
o o I t
H -H 4-1 t * «* «* '*• "** '"'M"
N r4 H
i f I I o I o o o • I
I t
4 • • » • • •
_S JI J ! ^ • • ^ • • ftOHfSfrv
I l l J I l S i l i l l B B •S Ha Sc IK S5 Ea SS -P ^S 2! 53 s i s i i i s i S i s
t i l t
I i I I I I
H O
-••I
§
I I I I I I I
I I I I I I
o o o o o o o o o o o o o o o o • • • • •
0 0 0 0 , 0 • o
«
o o
0 ft
o o 0
o «
o • i • !—H -H -H 44 4-1 -H 4-i +1 +J • ! 4-1
o o ! i J r - r - S 8 8 S 8 P P f * t -J J J f"! ini ^TI
f f ir*f f^ #"1 r» rv CM M «M w
Cf% &% <^ ri C4 HI
o o o o o o o o • • • • # • • •
o o o o o o o o
H ^ m m ri \t» # fvi K\ t tn M o o 0 0 o o • • • • « «
0 0 0 0 0 0 • I #4 • ! 4-t • ! 4>| • ! -H +1 44 4-1 • ! • ! 4"!
K% r» « W ftl IM ^1^ 1** #"f f r 1'"^ I**! H I 1"^
in o I I I I I I • I I I I I I I o 44
t I t I I i ^ I I I I I I I o
<*4
"»4 9t«
3£S
^4
HISfOHBAH XVA- llp£9a«iit8 tlie @ bm&dlug pftttom induottd by
tsifpsin in tilt Qlizo»080ii«8 of faaiaalitiltia
wsovmhtoa i oal© )
O t.
KUD
I r 1 1 r cp t^ u> in ^ CO CM —
H19N3T 3AliV13d lN30b3d
1
H
»
{III ^
1 o
i3
• %
o 3 m
3?
3 '' a ^
^
8 fi
Q
d
^ ft
&*
i n 41
4.1
w w w
I I I i 8111 i
•HI S ** f! T!
•^ 59 "*!
H Art
» I I I
4i ^ M
Its hi &\
o • I
i • I
*.» • • » • •«•
S3 I f £•* ;
a i M ., t W a * ;
as
r4 t!f\ its Mi
t •
o o
I I
o o o • • • •
o o o o
I t
o
• I
-!*% ^
§ d n « r4 H KN ON O O O O O O 53
• » • • • • * O O Q O O -O O "N • ! • ! "rt 4.| -H +1 • ! 4-1 - l • I H ^1 +1
^ H f4 r^ r ll IA q w ^ «s t* r» H • « «
f r* t*
ri tfV
O 3 8 S tA *»< 0« H
•o o o o • • • t
o o o o
• • o o
• • o
-H • ! * | 44 H H M M
t •
O O * •
3 p i«% a \ in t * r\ i*\
• • • • in JA a \ lo
s
l A o « I t
• I +1 • ! -<0 H i-l- O
* • w Hi
«M
H
H 00 O
o
*
I I
I I
1
t
m o • v J
-H g #
1-1
fA
• I • ! • ! • ! *t • !
<y <V «y t* . 89 H ® (S as lA tr^ iA • • • • • •
m r \ tA ?«% r i «*N
& 0\ n m ^ r-i
OK}
S I I I I I I I I I S I I I I . _
§ I § I § I 8 8
< SB
I I I I I
I I I I I I
H O o o o o
l i f t
I • f I I
W M H K\ HN ©
o o o o o o o o o o o o o o o o o o
H Ch M» t» er\ <^ «M o o o o • • • •
o o o o • I -H M • I • ! +1 • ! ^ 4*1 "H > l +1 4-1 3 3 S S •^ M CO 00
CM W « t -€0 CO Ou ^
• • « t t • E*> JA N IM Ol «
' ^ H HI H |(^ mt K\ O O* 0% <n lA U> iA
• • • • • • * Cif f*^ 1** f^ #^ f ^ ^ f
o o
o o,
<v r* Or» VO t - m « M
o o o o * • fl •
j ^ ^ ^ ^ w r ^ ^ w O O O O
• I +1 +1 +1 4-1 H H 4-1 H H • ! 4-1
H KN m a o o 3 • •
o o o
0 * <f»
o o * •
o o
o o o
0 g c Q W M O J t ~ « M H H H K \ r \ l A HI f^ CO CO CO ^
• • • • K\ |f\ «M CM CM
H H K\ K\ o\ OS o\ ve\ \ss MS
• • > • • • • • Ol 1 ^ f'f f^ f^ fH ^T r^
I I I I
I t I I I I
H O
O
4-1 M
2 •
I I f I I I
• • • • • • • ;5 «? :$ J «| ^ g
(M ^ ( M ( M CM CM
t I f t I
• • t • t^ ir% '^ if% u>
On
I g
45
IQ O O
MffiMaa r^ ^ ^ ••g vj>
W O K » 5 «*•
t ) 0
** fl s 4* 4J> 4*
^ tf g ^ S i 8 S S o o « o & i § 8 8 8 8 5
o
m * I
I I I I I I I
t i l l
^ ^ m o o o • • • o o o "H -H -H a\ vrs us 1^ ^ H • • •
«o r - f -
0\ ^ |r\ H PN «r^
o o o • • •
o o o
E* IfN VO !*• e» w o o o • • •
o o o -H •• -H (V u» if\ 0v m <M
• • • vo ir\ lA
fc: £! ^ ST fc !^ o o o • • •
o o o • I "H -H >*| •! -H u\ »\ *p» H %0 r l
• • • !*• ( ^ f -
a « " tfV « l
o o • «
o o
o o o o
W O t
o • I
r4
I I I I
I I
I I
ti H rs 8 o o
• • • o o o •H •! •!
o o o • • •
O Q Q OS w^ K
§ S ^ tf% P o o o o o • • • • • o o o o o •H -H "H •! •!
P f: t t CVJ t • •
si ri r* «3h i*N ^ « r - « 'W o o o o o
• • • • •
o o o o o *l -H •! -H -H f P R IR S
• • • • • «r ^ fM C4 CM
O «
o
I I I I I i I I I
o o • I 41
1^
#4 CV fN M lA vo CO m O (pl fV H H H
5 I)
mi «! 4» 4* 4»
1 i f i # ^ 4S 4» X K K iS
f4 M ¥S it\ i^ Q ¥\ VS 0 0 0 0 • • • «
0 0 0 0 •H 41 41 44
8 8 8 3 • • • •
^ u% ^ ^
0
t 8 3
1
1
I I
I t t
o o d o o o o o • • • • « • • •
0 0 0 0 0 0 0 0 4 | 4 | 4 i 4 f 4 | 4 | 4 f 4 |
• • • • • « # •
O O O
i %
d A "
0 9^ ¥S $i fi ^ 0 0 0
• • t 0 0 0 41 4| 4i l» t * t^ rs M» vD • • • l<% H H
S S 3 • t •
0 0 0 4| 41 41 r »<% ffl 1^ <• !«N
CM
I I I
I I
S I 2 # o o o 4| 4| 41 41
HO
o o • •
o o
• * • • • • • I *« (h* ® ??• o H H H H H cS
•
^
fl
s
CD N <0 U> CO
— I r
CM —
T X 1 T
O _ CM PO •*!• i n t o
H19N31 3Al iV13a lN30d3d 00
0 o
I • PH as
o o 7} 3 • ^ "5 D
s nj
«4
3 • ^
M
5^
cS ;3
mm
o
- 1 ^
a HI
m
t
&4 ««*»
4* 9 ® # ® 9 9
« o o g o I o
s i o o c i o a e i o 9 e f ) ® #
£4
'J
'33 &*.!*«
+1
» O «< O -^ M
to :4 M ^tj O M flk .'s; w -3
C9 I
i 4 « Q ; t5 J4 H -^
nt ;^ ^ O 1 > &«
i |< r*4 >..•> __
H ( - : l K g fe«J ^ r l fq O M 9 £4 4 5-3
i^ 'n •r
&»o ;$
Mm: P4 O '• V, fc4 Si? T ! O f i t 5> 72 + 1
?:.5 &» £M O • aJ
34 f^ IS*! ' 0 A p: ca
> " 3 «
3 o
•
O
t I f I t • I I
I I I t I I t I
M r4 VS ri O O
• • • 0 0 0 "H "H -t-l o o ^ O O 3 • • •
»l% 5 ^ l A
•4> K\ Q
# 0
f ^ i n 0 •
0
e* e*» o • 0
r t * 0
• 0
^ ei 0
« O'
H l * \ 0 •
0
tf\ 0 •
0
r4 r4
O O O
0% \e%
* o o
«
o D t
O
• I • ! 41 -ft +1 +1 ^ l •••I 4-1 4-1 • ! 4.1 +1 tPk ^ iA <M \ 0 l A !*• ir> MO Hi €S 0% W P*
• • • • • * • © C* O »« lA if* <•
f«- e** I * p* H* §(>
•
o o
r> *« o O
«
H l A tA
«
ri H g s o o
H <M fA t * 0% ^ 0 0 0
• • « o o o
44 *\ 4-1 4'l 4-1 4*1 • I 44 4-1 44 4-1 4-1 4-|
«
JA
O •
o 4-1 2?
{A iA (M sO fcA r -vd r4 <y> Ch €M t »
• • • • • • r«- !*• MO lA lA «(|>
P r» c* ^ ^ ^ • • • •
M CM W
I I I I I I I o O* «A
o «
o
o o
4-1 4-1 44
I I I I S S S tA tA H
I I I • • • t f<i (^ m
• • • • f • Al (A <^ H l A £> t<- ® 5 c ^ d ^ ^
e 9 o o o i^W ^9 ^ * )^rw 4 |w
8 •
•4 <
t i m
t
1
H
O •
o +1 H SO
• K\
r l WN O
• o •1 I*"?
CO •
t
1
• r* H
t
1
o • o •« H CO
• K\
^ M O
• o •1 H eo •
1
1
• "t H
f
i
n o • o 4-1 tf\ tr\
* trs
CO M O
• o 4-1 t»> t<\
•
1
i
• ^i\ H
H r\ o • o M o o * ^
d o « o •fi IfS ^ «
»**»
0^ ¥\ o • o ^ (
c* M>
4
^ o • o 4.1
^ I W
vf» «
r*
• t # I***!
H O o • o •1 o a • 9
H
O •
o +1 r% ¥S
• K%
8f\ i A
•^m^
• O
•H
p* sd
•
<& ii\
a • o +1 t * va
• r*
« C-H
K\ f ^ O •
o 4.1 O o • o tA
JA O
• a 4.1 ^O 09
* M
<3> 0% o • o 4.1 tf\ ^
•
<y* o> o • o 4-1 »A ««»•
• H
•
n
l A K\ O
• o +1 o o • R
s 3 • o 4.1 M» m • CM
l«% ^ O •
o •1 «*% •• 4
H vo o • o +1 m
t i fT
• 3% H
1
1
H O
» O
4>t
R »
CM
H O •
O
4-1
R •
t
1
• o CiJ
HISfOdEAM IIB • Showing A ]?dp]?®9oatatioii of § baiidiiig
pfittoiea iaetmooa hf txyp@iM la %}m olisoao*
G(
r T T f 1 1 1 r
O M
2
S
b
<o
tf>
S o
S2 Z CM Ul
- 2 = 8 2 2
o
OP ^
<X>
in
CO
CM
HiON31 3AllV13y J.N3DB3<i
IE p*
IS m&
IkiO
o g I
lis IS
as M
9 ©3
M H
I f B I I I J ? i I t I « n I § 11
8 g g J 2 f ® o o - § § 8 S ^
<< «< 4 •< <<[
4» ••» * • 4» 4»
s 8 It s $ 3 8 §
« 5 5 5 4 a 5 4» m
I I • « t I I I 1 o o ,
t I t i l l
j o o o o o o o S g g g g d
yes M
#% St <§ o o o
# •
o o o o • I
t * <M 1^ tf\ * • r* « \ ifk
^ 41 41 41 4f 4i 41
5 {: «
CO • •
1^ ¥\
s t
o o
S Fi ^ o o o • • •
o o o ^ 4 f 4 | 4 | 4 i 4 | 4 | 4 i # |
o o J? • • o o o
3 o
£r ttj N w *© *d t«- r» • * t • sa so ig% us
I I
-N 41 41
I I I I
ri O
o
I I I f I i I
J O o
4| 41 41
s a g • * •
<M »4 r4
m UN m s a HI
G '
I g ^ I I I I I J I g
^ 4P •«* 4» i v
§ 1 1 1 1
I I
th f4 V* ¥S )h» m OS ci%
o o o . o . o o o o 41 • ! 41 -H
. o o o o I • • • • I
1 § i u
f-l P US ^ O r4 O O tf\
O^ CM u\ q^ \0 H ^ o o o «v
•
o •1
8 •
i*\
in o •
o •H
8 *
¥\
1
1
• *#* H
•
•H
8 •
r».
a «
o "H
8 •
rs
1
t
• if* H
•
"H
o «
m
g •
o •M
•
o • o
•1
St •
H
• MBI H
• O
•f
• CM
St o •
o -H 1^
•
O • O
•H
• H
• t * H
• O 4t
• IV
o •
o •H
H •
o • o •1
• H
• CO H
•
-H
* <M
o «
o •1
• f"!
H O •
H
« H
• <^ f**f
•
^
g •
H
• O
•1
g •
H
i
1
• O {V
•
o •1
00 •
H
t •
o * l
m •
H
1
1
>*
G2
9 «
8
I
HXSfOiMR lit B - B«f3»s«atf tli« & I)aii4ins I»Att«a i»dao«i
^ ti^rpaia in tlie ehsosMiaosies of maid
f—I I — I 1 1 — I — - T — I I 1 — 1 1 1 1 — r — I — I
H19N31 3A(lV13a lN3Da3d
e.
|L *»4»
B mm
1' 0 • M p M Or-t 0 NCQ
« « !
§4 o O O _
I a P a 9
r4
CD
M O «
o
HI
t i-f
S 1 as s^ « "• o o o q
o a S5 N o § o «<
o a t; «* o i o <
c» iQ
O o i o • <
o M an i 4
sa? o s o •4
o u e @9 c» s o <
o 3 S4
j ^
o ^ @9
O
§4
o
3 SC
o a &( S3 Q < t4 m
I I I I I I I I
I I I I I I s s ^ ^ ^ M ^ KS p^ O O D O O
• • • t •
o o o o o
• t" H m r» 11% ^> ^ b o d
• «
o o o o
H H O o
s« ;§ o o
o
t i I
I I
i«r V^ M>
o o o i • •
o o o
g UN d S? SD £r t- r\ O O m
• • • * • m f- VO vO "^
1 ^ j *» ^ » ^i» (0 m S3 CD • * « «
< i ^ - ^ « * ^
!*• flh <f* & ¥S ¥\
t • •
0^ O * • •
" j" ^ l'^
W M O O o o
« o
o o
91 o o
o o
^
Ir o • o
H 11% O
• o
H
S •
O
m w o • o
s a> o • o
KN V\ o • o
3 O
• o
"t (A O
• o
H 4»
O •
o • I "H •• -H •! -M • i TFf •*Tf ^ 1 HKf • I #1 4 | *4|
t' ^ m •of
>4
4 !
«
I t I I
I I I t t
«SI <M CM
0* H r4 rv ^ R # o o
• • •
o o o
(N
r4 H CM
I i
I I
• • • lA ve r» CD
« ON
« * O H
WN
• « • • « i*% «• m f H Cf I*""! ^*|
C)''l
g <4
i r4 O •
o ••
8 •
^
o • o •1
• rv
o • o •1
1 * p t -•
»4
a •
o •1 1 * g
t H
• NO H
O
1 •«4 #4
i rk o • o •H
S •
8 eh
• o •1
Si •
<M
o • o •1
le "• *
H
Si O • o •1
^ •
H
• e-#4
o
8 < p3
HI wr\ o • o H
S •
^
OH
o • o •1
0 1 *
M
Ps O •
o •H
<4» '(f •
H
• o •H
^ •
H
• m H
o
i 4
O
g4 fia K
KN O •
O
•H
8 •
^
^ o « o • •
u •
<M
St o • o •1
t *
H
• o • 1
t •
H
* m H
O
i
a «*
1
f
a o • o •1
s *
w
l*"f
o • o
M ¥\ ^ •
<M
1
1
• o CM
O
m
s m o <<
1
1
o « o -K
s •
w
«M
• o
•N • A ^ •
<M
1
1
b*
H
J
1 %< 0
(13 ^3
€)
e tf^
i c «
i0 •jO
«4
9 w
SI3f00141! X? B • Rtpxttsoiits ttm § banding pattoxn Induotd
^f ttj»a in til* oliiKinosoisve of aaXo Battua
Go
I r CD rv
—r r
>
X
o CM
QD
h;
(g
!£>
CO O
~ Z C>< 1,1
°i O
a> a. I
00 O
CO
CM
H19N31 3Al lV13a lN30d3d
t i 5 B H I S I I « « U «
8 M i i 9 § i i P O « fi S t # O _ _
M i MMt ^ ^
J11 i i i 11 i IBI -
i f
» I I • I
in o o
o o
o o
•• -N •!
H i " |8 8 8 « R S IM ?iM o lo^ t- t^ Jt J .1 *0 lA
IfN ON tf\ , 1 ^ !««« |r\ • O O O
' • • • o o o ^ n H
I 8 8 8 a i i
K £ rf H c* o S S i 8 g o o o o o o •« "N •! -H +1 •!
? 8 S S S g li »Jt »z ,jt • • • • tn «» lA if ^ , ' •
o 41
? | g 8 ? 5 ? l « o o
o o • •
o o o o
o d o o
* o
r\
lA tn O
o
^ ^ u\ u% w^ UN 4*V A » - T • v\ m it* US N IV
< I i I I
l i l t
• S i s O CI •
m> irs ^ fm.
*H *•! 4|
**» €^ H
« • • • • © <y> o HI Jv
H ri H
G7
o o o o o o o o ^ n n n i s t i i i i i i I i I i I I I
f
1
Pi o • o ••
% • rv
fA o • o •nl
t • KN
1
1
1
1
t • O •1
e • rv
i o • o -H P •
trs
1
1
1
1
¥S
O • O •1
s •
¥\
o * o •1 o S0 • K\
1
1
r4 O • o 44
8 • s
A o • o •1
• KS
o • o +1 M
• HI
r ^
•« iO
-H 3
Pi o • o "H 8 •
@
« o « o +• CM «i l»\
O • O •H
• H
M rs o • o •1
O • o •H 8 •
S
St o • o •1 so H • M
S^ o • o ••
s • H
o% «• o • o •H 8
S o • o 41
8 • o
P. o • o H
r4 •
m
^
o « o -H S « H
0\ ^
•
o "H ISO
o
1
1
o • o •4
8 • •-4
i o • o •1
w « H
•
1 • • • •
H H r4 H
tf) ^ tr ^ ^ g ri ri H *^ H ^
G v i
% B
I o 3
RX3f0aE^ ?B «• i«ps«@«iit8 tli« Q baadlBg pft%t«m iMiie«d 1^
U£»a in the okmmQ&omm of f«mdL« lattiia
G :
x 00 r>s u> in
— I I I I I r
cocM — o o — C M c o < « r i n <o
H19N31 3Al lV13d lN30d3d 00
•SS8S irMI»»*iil>
S O
o o o
si-
S»'S€**
a l l * ^
e 9 6 "
.' I i
o s ^
g o «e
o S
n O ««
o s o o O "4
o a g g s o ««
o ^ £! PS o 6 eg D -«
O S g e S o «<
o
i-t
e
m o <
o y m o o m o • i ^
o
o o
o
I I I I I I I I I
t I I I I I I I I I
4«> R r4 H
o o o o • • • •
o o o o
o o
O o • •
o o o o
o
o a o f-« j ^ fa O
64
§4
^ u s
i 4 *^ c4 o a
o
I i
s <VI o
• o •M
« CO
p o
• o •1
• t>
lA O
• o -H
w» •
r>-
C5
o •
o •H
• P»-
s •
O •H
& «
lA
Pt O
• o +1 &
• \t\
5t o t o +1 s;
• v\
• I +1 • ! 41
1^ (H i"! P* • • •
lA lA i n
lA P o o
• t
O O
• •
I I
H lA lA ^
o o o • • •
o o o
lA
•H "H
CM MS
CM
l A
« tK K $ o o o q • • t •
o o o o
i P St o o o
• • • o o o •H • ! -H 41 41 41 4t w »*• r- r-m ^ ^i^ s6
M9 r* t * t * "^ tf> ''^
lA HI ^ lA ftf 1*4 O O O O • • « •
o o o o
0> lA o o o • • • o o o
3 5t S O O o o
4 1 4 1 4 1 4 1 4 1 4 1 4 1 4 1 4 1
lA tA lA fSi M M «A lA #rk
m o
• o 41
z •
H
•
1
1
• CM
1
1
• l A
I I I I • 3 I S • o o o
• • •
o o o • I 41 4|
I I
I i I I I t R S • • •
«M M AS
I I
lA vo t*. CO <y» O H f"! ^9^ f f f«f P" f"
• • • W lA tA
o ^ #4
m o < St fifi
S ^ o • o • i
g •
^
o •
o 41
o » »rs
g «
O 41
• H
O m W^
o «4
1 ^ o • o •1
9 * E
B •
o H ^ « IM
B •
o •1
a •
HI
O
IS E*4
§ O
2 S
o • o •1
8 •
^
o •
o •1
o t
w
rf •• o • o -M
o • H
» u N S8I £4 O <
1 O •
o 4 8 • a
O • € * l
s •
ev
g O •
o H O
• H
O M
^4
pl O
s
1
1
• o 41
9. •
C"!
H O •
o 4|
• f^T
r
•^ ^ "• H CM fM l A r -
o o o o . • • • • I
o o o o •"^ 4 i T f • ! fS «^ M M tA O O O
• « • • I H H p>l <H
• • • • •
*o r- «o 0 a H H H H M
M
m
C-4
lISfOtlAif ?X 1 - £9fx«9«iit» tlM t ^aadiiic ^a t tva iiidme«d
of f«B&3.« jiaiiaa ftg,iiiMlfiig
>c
r"' 'i 1 'I I ' I I «D ^ <o in ^ o) <N
"1 I I — I I O — <M 00
-, , , , , r u) <o fv 00
O CM
S2
in
£2 CM H M B
—
o a»
CO
t^
(O
in
"V
CO
CM
o z UJ
2 O u> O 2 O X O
HJ.9N31 3AllV13y lN3oa3d
Flat® lA J- Bhow the norsaX karyotype of male
73
'i is
KBxyaiOf}^ lA : • 3liowsthe normal karyotype of male
fimmabttlua permantl wrow^hton* 2n « 54.
II 2
fli Ik G I I'
« l G 2 3
II G 3 7
G 4 8
G 5 12
G 6 13 14 rS 16
9
II li ID II
4 i A% i f 17 18 19
6 7 20
f I I G 8 21 22 23 24 25
• m 26
• X
A Y
Plate II A ;- Showsth© G bandijog pattern induced toy
urea in the ohroaosomes of male
Funumbulua pennaati wroughton.
Karyor^^|?& I I A t* 3how the Q banding p a t t e r n induced T>y
orsa i n the ohromoaomes of male
Fimuiabulus pemiantl wrought on.
U (} G 1
G 2
G 3
G 4
G 5
G 6
G 7
1
u 3
<i 7
' 'w
8
I f
12
If 13
f t
20
2
ii 4
if d
i» 14
'
h 5
n 10
{ >
15
(
h 6
tv
II
'
19
T6
•
'
« •
l'7
r
G 8 21 22 23 24 25
II 18
4C
26
I. X Y
7C
19
Plate I I I A:- ohov^the Q banding pat tern induoed by
area in oiiroiaosomea of f eiaale
Fimumbulua pemianti wrowthton.
'7
• ^ *
^ ^ y , . . ___ ^~ji'^ ;^^w% ^«*J/K( iJUMl
Karyot|*^ III A :- Show the G banding pattern induced hy
urea in the ohromosomes of female
ifunumbulua pennanti wrought on*
6 I
6 2
G 3
G 4
G 5
G 5
G 6
G 7
Si I
if 3
f* 7
ii 3
11 12
« i Id
• ^
20
« i t 21
II 2
4 1 4
If 9
i i 14
i^_^ 22
^1 5
It 10
• f 15
• #
23
'
f l 6
II II
t l 16
^ ' ^ ,
24
•
• #
17
! • Ik
25
'
* ; • 13
* .^
26
i i X X
19
Plate IV A ;- Showi the Q banding pattern induced by
trypsin in the chromosomes of male
•g\inumloulu8 pennaiiti.
Plate V A :- ShowSG"banding pattern induced by trypsin in
the ohroraosomos of female Punumbtilua pennanti»
7a
'w^
# -
K.axyo^^pi' IV A J- Show th« Qt banding pattern Induced by
trypsin in the ohromosomes of male
FunurabiJLus pennanIA»
ir II Gl I
G2 il u If
6 7 20
^ 0 tr(
G3 7
If ft 4 G4 8 9 10 II
If G5 12
i i l i l i M Bi I I «# G6 13 14 15 16 17 18 |9
G8 21 22 23 24 25 26
I I X X
K:aryo:|&JE> I B ; - Show normal karyotype of female
Hattus norve^otia»
Karyotype VA :~ Show Qt banding pattern induced by trypsin in
tii« chromosomes of female jfunumbulua pennanti •
H ft GROUP 1 I 2
G2
63
G4
G6
II It li M
M 7
II 8
• 1 9
11
10 ii II
G5 12
M 13
v« 20
"^Ji
21
f t 14
# ^
22
i 15
^
23
M 16
- t
24
G7
G8 21 22 23 24 2B 26
81
4 •• '''i^
I I X X
Plate VI A:« Shows G banding pattern induced by urea in
the Anaphae stage ohromoaoaea of female
|funumbuiua pennan- i wrought on»
Plate VII A :• howa ia banding pattern Induced by urea In
the Prophase stage ohromosomes of female
Funumbulua pennantl wroufchton*
k
82
]
83
N G I I
X X
8 1
G 2 2 * • * • • ! M 4 5
» »
6 2 6 7 8 14
6 3 10 II ,2
4« ^ G 4 13 f4 15
6 5 16 a t
i7 18 19
G 6 20
Plate II B :- tJhow the Q banding pattern induoed toy
tr;ypaln in the ohromosomee of female
Plate III 3 :• Showsthe 0 toandlng pattern induoed by
trypsin in the ohromosomes of male
itattua norve/Aons.
85
Kaxyol^pe XI B ;- 3hovs the Q banding pattern induced by
trypsin in the chromosomes of female
KattUB norve^icua*
G I
11 8G
m^t
X X
e 2
li 2 3
k
4 5
G 2 6 7
:'TS?
9M'
8 9
G 3 10 (I 12
G 4 13 14 15
G 5 16 17 18 19
'a^^ili
G 6 20
Kaxyot^: XXI B:- Shows the Q banding pattern indaoed hy
trypsin in the chromosomes of male
Kattus fiorve/doua.
II G I t
» t X Y
87
Al 1% • « 11 G 2 2 3 4 5
G 2 6 7 8 9
I I "A M G 3 10 II \'2
G 4
G 5
4r 13
il 16
fel
14
• k. 17
«t 15
1 * 18 rg
G 6 20
flat© IV i> :• ;jhowsthe Ur banding pattern induoed by
urea in the ohromosomes of male
Plate V Ji;- Showj the & banding pattern induced by
urea in tne ohromosomea of female
liattua norve^ioua*
, / 8a im
Karyot^fje IV B;- Shows the Q banding pattern Induced by
urea In the chromosomes of male
Rattua norvegicus.
G r
6 2
h f
fi 2
14 3
80
ii G 2 6
II 1 4 S
l l l l Q 9
X Y
G "3 10 11 12
G 4 13 14 15
Mm . 1 «> «4 G5 re 17 le 19
< 2^
G 6 2D
Kaxycb^i^e^ VB :-> Bhows the G bandix g pattern induced by urea
in the chromosomes of female Kattus norveKicua.
li G I I
r r.o
11 X X
U ll *< !• G 2 2 3 4 5
^1 ll 4i I I G 2 6 7 8 9
li li M G 3 10 11 12
i« • •• G 4 13 14 15
G 5 fB
ah 17 \8 19
G 6 20
P l a t e VI B : - ^ihowgthe Qt loaxidiuQ p a t t e r n induced by
ixrea In the Anaphase chromosomes of
female xjattua norve^iouft.
P l a t e VII iJ:- Shows the G banding p a t t e r n induced by urea i n the Prophase chromosomea of female
i iattua norve>glou3.
91
*U^JUUUUJ>^
The eirolutloii of 1»aadlBg p a t t e r n tec^toJlqu&s has aaile
tii« i4«x i t i f ioa t loa of iadlTldiJ^il ohrcHsos^m® ^ i l s s pc»@@lblet
evea to tlie extont of 3 l3 toc oibii^omiitid extitoiigea» &M -^e
ixiY«8tlgation3 of iaa»y elu»»Boaomiil ^(maMe©* Siie teeimiques
fo r a* d«M*0« and G hoMXngf So^reJlopod l a rooe&t yoaxa* Pi.'«.'.)eut
tlio aiiQtgJle d*!*!!.* 4ombl@ hellK aad it@ aaii@eiate4 psotliia
v i t h l B a i>l&^l« oiisooatlii as tigtita^r ool led and foMo4 tn
the aetapliase oiufoffiosooies* Hetei?o.^ohro©ii,tl& » s ta^ons r ^ i a l n i n
t i l ia ooapaot s t age Csitotflug aUlooyoIir or l ietero p;^03:iooit^)
i n intexplmsef a re t i g h t l y hMk^ by miWt Isistoae pi^otelnst
undergo a l a t e B«i«A* repUoc-.tion ^od appear aa poa l t ive ly
s t a l l i n g ba£ida I n metapbase with 4^<l and Q t@olinl-a®s imd
nega t ive ly s t a ined E btmde (£van@ 1973)• l he (band regions
oorrespond ni t l i ' ^ e oen t roae i lo type of lioterouohromu£lin*
wMoh may be loo&t@dt i ^ regions o ther tlia^n t^e eentramere
a l s o t the t? bmids '0 baids oorrelfi te ^ i t h the intex'onliijcy type
of heteroohromatin and with the H baiv-a aa wen*
the oonat i tu t i i re heteiroohrom&tin was ^ph4&iz«^'d by
ooabined app l ioa t i on of t he t r y p s i n treatoa^it and aiemmi
s t a i n i n g methods a t pii 9* Sinoe the Q b&xds oorreiap0nd to
the heteroohrofflatio re ,^ons of the ohr^isiosaiae i t has been
proved t h a t l^e O'^nstltulive h<eterooh^'Offlatin mxy be ^iphasi&ed
t h r o u ^ both banding typea» the meohant^ts of U R M %i
bcmding of ohrooosoae*
Xt l a po00lbl9 that hoiu>l@gou3 oiiro&oao&ies of tine
mme speoies iiro stjcuotvtzrig&d l a tlw sese vci^t with %m
aeone proteins and Euolelo adld Qc»Bpo»itlaa aiia* t t@i?«fo£e»
they wtll ty^rm ^9 ssae ba&cis wmtk tire&ted uodezr i d e n t i c a l
oacpedmexital ooii4ltio»3«
l a masmals the Q bandi&g l a elmraoteadlsied by heavy
staixiln^ of the ee&ti;t^©£lo regions of the eeoomdary
eonstrlotions ma^ by the ooouso^eroe of ohroBiatiae fre®
a ra >re r l^sa ston^ily stlned a l ternat ion of hands* the
ocoeiparative analysis of the ohromosose eowplemeats of
aany maiamals through Tazlous teohnlquea show that i n the
oase of oertaln ohx^aosoeal eones there i s a^eeem@Bt
between the Qt aM G banding*
She present study also oonflsas th i s faot* In aany
ohrofflosoa® p^ilrs of .IffiMHUJ MWmP^U ^^ M^^M mM^m^i^^m
aentroiaeric r edone show tntetme s t a ln l git but here I t i s in the
oentrome^o r e ^ o n or in prlmai^ oonstjdLotlon ru^on*
£he absolute s l se oi a ^ ohi^moaoiae -raries vi th the
stage of mitosis* Qhroffiosoiaes are longer mx& l e s s coiled
i n propha.se and shorter and more oompaot at the end of
metaphaae* She ntMsber of d i s t ino t banks inoreaaeii with
inoreaaing le i^ th of the oiirotaosome* tha t metaphaae ohromosoiues
alfjo show a fin© ik beaid pattern or pro£ btaa@ ohroiioso;.jes of
guni,aablue pennsmtl and Eattus norve^dous was obsenrod in
Q -
pxesent study* tht aajdlffiiai i ]iil3@c of bai^a &X9 Bhonn by
t h t otiroaowemes i n p3»»]^aa9 ( fte:* VIX A 'i* VXX ii ) beoauae
the ohroffioaome oontsaotion l a l eas t a t tM3 sts^e* fhe
latexmediate atmgee ef eoati^otloii «• l.ate prophase ttM
early aetaphaset tiraoe the path ways of s p i r a n t b^id
fusion me the o e l l pxrogreeaee to metapha^Hi. She pm^ha^e
bands om laaiay ohxomosomeo tend to ooous tn d i s t i n c t
groupat (Mgfer e t al« 197§> the nimbeir of whioh i l i iaatel^
mesg9 to give l^e d a ^ a bai^s of aetai^imse oh.i'omoeomes.
Sv^xty a band of the eteiida£d »@tapha@e pattejm i s oompo-
uaded f roa two os i^se leophase baiids. l a g^e l@aet
ooatraoted prophase ohjrojsosoiaes exasiaeili some bands a:re
seen to have ao ob^ous aMtaphaee eowi^terparta* fhex-e
are laarkSii s i&l la r l t i ea b^tireea baaded pirophase and the
oh^onoscNiie pat tern seen a t raeiotle prophase*
fhe different banding pat tern teohniques ha^e ses'^ed
to enphmslse ^ e oonsenmti're nattiui^ of e'^olution of the
ohrcmosoaes of higher m^^nals* Within aost rodsuts aod
primates the bandicig pat terns are rwsy siinilar inspi te of
ohrcK»&soiBal alter^atloxis* the prinoipal or Main ohanges
appear 1» be in the isaaterial oontalnsd i n %h» Q baadin^
regionat whioh are regajeded to preseat oertain redaadat^t
regions in the gencN e* Thm banding pattern ahwon by
aorooeiitrle ohromosoaes of |^ t l f i^ fioyge^oaii and laaaa^^ia^
pemmatic* the two rodents of different families are
v ' v J
alaost lil^allAr wltH a h»&r:y »%ai&l&g xregloii In tli« eentr *
merlo sjreas and tlye«« os fotts dasfc & t^aads on tbe Mng eLxam
of tft© <^iroooaoae3. l!«taeeBtric pai«» of ^iljfeas i^yYf^4l^W
show aiialXaxr bandiog pattesa as tiiat of fimuatolas pm^aa^^MtlM
motaoo&tirio pais? CgJ?oup V) i«@* Unstained o#ntcoMeiflo xBi^oxm
with dark bands on ithe tsxminal poartlone of loth mxam of the
ohrcoiosoiies* 4 s^npueisoa of ths ti»» kaj^otypes ehows that
frcffii ly o eToXutiomaxy point of Tlew ths kiy^otype of
£3l&ifi iy[M£ J£Si ttBdyi£ having 2M£S9 nushsc af 0h£OffiO3OBi9S
quite a few of whloh are aoiNlOf^tseiOf i s ra ther mozw
primitive than that of M$^m i<?Fff^ffli vhore the diploid
number of ohroaosomeo (42) I s leas than that of the for ler
and aorooentrio oh'jroitiosomes are almost equal i n numher with
other shapes* Xt has hmen demonstrated by Matthey (19^1)
and white (1934) that generally anlaals with hlg;her
ohromosoQs nombers posses a Mrge nuabsr of aorooentrios
while those id th lesser n\Mber of ohromaaoffie have fewer
aorooentrios and the karyotypes shawlug large niMber of
acrooeiatrios are more primitive then whlc^ have less nuaber
of aorooentries*
the findings of Ohu and Bender (1961»62)» Mamerton
e t aJL» (196?) t aiid kllnger (1963) also g^poiet t h i s view*
fhe asstiDptlon i3» tiiat the oeatr io fu^oa* special ly of
iiObertsonian type (Mobertsonlant 1916) has played an
important role in the evolution of different karjrOtyi>e •
fQxt$t two i s %h» mo&% fxedofflin^at diploiol nutabsx f oui^
lagt&us Eai^ttttt. «]iicsh l a ^eaesal oottprisem o t 7 pairs of
iHitaewEitriosy 3 palirs of 9iil>teloo«ntrio8 oxA U aocoooatrioe
iaoluding sex ohxemomomea* A oo&sidssabXs dsirl; a t ioa
iadiploid aiai1»«r lia@« hov/eirert been reoorded i a aiffexrent
speol^Sf the extareae low beitsg 32 i a aa;lti«i qea^tna
(to@ida e t &1. 196f )aa4 high as 52 i a i i H i i Smrifeg
(yo\iaglf60)« Although a OIMBS out Eobestsoaiaa vaciat ioa
has dhl^ hsea obsenred Sja ^ t t m s ratt ias. the reduoticoi i a
ohromosose atmhes, l a geaeimX i s due im deorease of
aevooeatrlos aad iaerease of hiax^ed ohscmosomes*
4s fas as the aoa oeatxomerlo hete£» ohromatia i s
ooaeesmedf the ii^Uloatioas fo£ the pj^eseaoe of diff exeat
oategoiie^ of hete3?ochromatla should he mais^atzeA as
revealed hy the Q haaiag patte^m l a t h i s study i a oase of
% t t a a &^SgMkMM* ^^^ ^ ® species shows a dimlauti&a of
oeatr<»ierio hetero* chromatia materialf aad aa iaorease of
hetero^,oh£cmatiLo fflstexlal ( losida aad Hagctif 197'i)*
f asaXlel flndifigs i a faoily fioTidae miA l^uitbe are also
reported whose eirolutiona^ ohanges terid towai:i{s a
diadautioa of oeatrcm0£ie heteso chsoaatia (fiuoJclaad and
aad 'Mm-mm, 197S) aad (ilydeir e t aX« 197S).
iiao aad ^eshadxl (1966) showed i a the amtoradlograpiiic
studies of squ i r r e l tha t x ohromosome repl loates latex>»
Bxperimeats with oert>';iia agents whioh block D.ll.A* syatlieBla
i f (
l ik« malmo pxtateiuB &u the hvm&sk chronoaomes (Pet©ru3oct»1964)
a&d deQ3qrfiu9£orliiae on hamster oella (Hsutl964) ^ti^ested
tidat 1 ohrcMosoae in these sp«ol@a i:eplioate3 lat@« Tt&ae
o^servratloas apart frcm showing a^syuohxo!^ of x in femax&B,
help i n the Ident i f loat lon of the x chrcmo&ome in th i s
asiaal* Qhoprm and Fai (196^) fxfoa studies of malest
ident i f ied the £ Qh~Qmo&omm i n the sa»e maiigidt as a meaivea
slsed aarooentrio* In an e a r l i e r Bttjuiy» tiaing hoth liaXes ud
tmmX»m of funttaM.\i^ peiqayntif liao and h is msaoQlate® (1964)
shoved "^e X ohromoeome to hft suhmetaoentde* Jbater voi.*k by
the saae worker© shown tliat the X;-«.te sepMoatin^- chriMoaome
i s not aox'ooentrio as studies hy Qhopmi mud ^ai» but i s
olieasl^ eubmetaoent&o« £he detaiXi^ studies of the X
ohscraiosoaes 0t s<|UisireJLs hy Qth&s woskers (Madler 19671
Hadier and Suttan 1967 • JIadXes and Blook i962} have also
shoim the X eh£<»ios<»ie to he subsetaeentcio*
fhe r e su l t s fresented li^re ax9 also i n oonforait^f vith
the studies of Kao and ids associates (1964) su: BImiasa
and Qa£g (1971)f shoving t^e X ohrosiosoae as a sulwetaoentado
ah£omos<»e i n t h i s speeies«
f ief lo e t s i* (1967) eomld not find i n i ^ t and aouse
bone inai^oifs oh£OfflOsosie@» asjrnohros^ among t^e X chxomo&omeB
i n Vivo syatesiSf but they irexe able to demonstrate i t i n
t i ssue oultuire oe l l s in cat studies in ^ v o Mao e t a l (1968)
found in an Indian gerbi l f e t r a indioa fesf le t a mruc ed
as^r&ohrc^ l a hoiiio3 ip»si X chr^tt08c^6*iiiail&rl^ they tovmA
asynohron^ amo&g tl%@ X chromosome of feat Xes of FainiBbiiJma
XJ e I oh£e»aiOsoa@ Xn t h i s fpeaies l a am-ilXBBl amd
negatively hetejcopfenotic.
O-axg eM Shaasfta (1971) olassif ied tli« fim-gabulijis
B»afii^ti*s obroiaoaoiiiea into fou£ groups* four pairs of
fflotcieen-lxlas l a gxoikp hf ^ i x ' 3 a t e l l i t ee l a gcoup &, two
pai£8 of substetaoontioe i a gxoup Q aaS 14 paiira of aoxooeritries
i a sx MP &• Eao e t al* (1964) olasiiifieA these autasomc^s i a
Jix g£:mpB* Tw4» pairs of aaliaetaoeiitios i a group If tiaree
pairs of large aorooeatrios i a groii|i £I» Six pa i r s of
fflediim sized aoi^oentrios l a gromp XIXt oae pa$^ of large
aetaoen^lo i a gromp IVt seiwa pairs of small aerooeatioe
i a gSQUp V aad sevea other ^airs of amall acrooentio i a
group tfl or i a tlie l a s t group.
Shese tvo studies shewed 20 aad 13 bicirmed pairs of
autosoiies respectively* l a the preseat study 21 paisn of
biazmed autosomes werefouad froa the aaimf^ls oolleoted i a
Alisarh area* Aooordiug the sia^e of ohrcfflosmef the oeatrosierio
iadioes aad def ia i te Q hoMlng pat teras t these Autosomes have
heea oiasaified i a to eight groupa*
fvo piiirs of larg:e su%>aetaoea1iioe i a groSLp I , oae l^xrge
- ' • ^
1 ' , >
sul»metaa@iitrlo i n gx<mp XXI and f ouc pai rs of large
aerooenlilros showing a diot inot a baMliog pat tern imre l)een
pXaoed i n groups XX* Four pairs of meditai aissed aorooentlos
acoortllng to t he i r Xtm^tli and Q banding pattejms mx9 placed
in group I f 9 Hhese are two d i s t l n e t aetaoentio pairs i n tlie
ohr^esosone ooapl^ient* £lie large one Imving i t s aras
intensel^r stained d i s t a l ! ^ and skewing thie oentroaerlo
heteroQlirosi^tin in the oentroaerio regiont whioli rem ins
unstained* i s plaoed in group V instead of group I f or
groups X as pnJiiipusly done hy other workea^* Another
aetaoentiie grup uhlan i s jlonfused generally as aii aorooentlo
pair i s plaoe4 i n group VXIf aoeording to i t s sise* th i s
9IE otetcioea^o pai r has also been reported by iiharma and
darg (1971) l»at they plaoed i t with otlner metaoentlo ohrooioaoaea
i n group X* iiao and h is assooiates ( 1 9 ^ ) reported i t as an
aoroeentio pa i r and grouped i t with other email aorooentlos
i n graup V* fhe tiro short aras of ' ^ i s ohroiulsome pair
s t a in IXgh^l^ while the oentromerio region and lower arms
hare dark staini>t0 in & baM preparations* shoving the
presenoe of in te rea la r ly heteroohroaaatin in aentroaerio as
well as long ara regions* '£h@ group VX has seven pairs of
asi&ll aorooentlos cmd i n the l a s t or VXXX group sijc paixrs of
sna i l rod shaped or a.rooeatleie ohrosi^somes are plaoe^i*
fhe resu l t of the present study as well as the f i n d i n g of
e a r l i e r •»Q'££&£B (Shares and Qarg 1971^*Chopra a ;d Mvil 1965
1
3»ii«V« iiao 6 atiar^ia 1964 etQ« ) shoii tiiat ttie 2u tiymh&L- la
thlK speol«a le ;?4« dlowev^r* Mansia and : aXui£da£ ( X96^ )
have r®port«d tb« diploid auratoer as §6 fi?o@ Elal^axii* 411
these maloe i t a vei:: i s t e r e s t i ng Kms^ol&^o&l prol^leti
and i t wom34 be worthwhile to iB^reBtigate i f there i s wide
spread c^romosoa^l pol^moi^hism i& th is species or a problem
of to.r^otype divesgenoe i a iiiYolved*
•:J
^^^.J^J^J^^J-J^^
W W i^ W A« U fci 'X ii... -<
(1) She 1 ,1 .A, p ro te in I n t e r a o t t o a s a^e involved in
developing banding on the ohrimQa^ma*
(2) the i a t e r o a l a r y he t e roe tooaa t i a whieh i s noa r©f@titiv€
poorly aet i i^ la ted i>»H«A», genera l ly aggtara t n si bands, i'he
Qentsomexle hete:roo^roQati^<m appears i n G bands* I n tiie two
karyotypes presented here a bandisg p a t t e r n s ladueed b^
t ryps in aiid urea ar© ot almllsx type ehowiag var ious omoxmta
of i n t e r c a l u r ^ lieteroo&rosatiii i a i^e two kaa^otypes* Maa^
olircmosome |imirs In botli ohroaoaome Gc»!ipl.eaeiit3 ahow dark
stairiit ig i n the eentromerio region a lso*
(3) Tim X ohromosom^t of both apeeies ahov as^BOhroinr i n
feualeau The y chroaoaome i s negsttivel^ heteropyonotio i «
m i t o t i c metaphase p l a t e s of Funuabulua j^g^HMlil*
(4) i'he X ohroaosome in .^^uauaibulua i^eanaati i s eubmeta-
c e n t r i o i n shape while y i s an aorooent r io oae«
(^) . 1*1 i iat tua a o r v n ^ e u a tooth % and y chrtaaosoat® ar© of
aorooentr io type*
(6) fhe autoaomes of Battue aorveifdlouft a re c l a e e i f i ^ i a t o
s i x groups while t h a t of j?^uftuabulu9 p«tiaftati i.u e i g h t groups.
(7) wroup I I I i n t^se of I'uiiuabialiif penaaoti ooaprises a
a subaetaoe. i t r io ohromosome patir reported ae aii ajeroeentrlc
one by e a r l i e r workers, fh© ^ o u p ¥11 ©oapriaee of a 3i^.i i a e t a -
c e n t r i o chroaoaoEi© which i s genera l ly ©oafused as aa acrocent r ic
chromosome. I t shows pecu l i a r baiidiag, which i a not siailtAr
1 ^. '^
to the baaciiag pat tern of pair In the emkplBmBXit tliat i s
l igh t ly stained short arms and brigbtl^^ latained oentapomexio
seglOB aM loti£; cum*
(8) ao]i|»arl»;:«roii of the two Icaryotyj^a xe'reala that
the ohromo0<me 0<MpX<»»ent of gimMuabttli^ peaoimati i s Mtthes
i^ose pzri.mitiYe than that of ^ 6 M^ttug novoizieaft , tsom
the eniutionaxy point of view because i t has a gapeattr
niuBber of diploid ohroaoaoaie© ( 5 4 ) q,uit@ a fei« of which axre
aorooentxricB*
^J^^JLAJU
10 3 ',-i. ci A d i
Keo®nt stArtBMeea in ©ytological tee to iquea hair® aade th©
study of ohsomoaoaea eas ier* l%e development of banding
pa t t e rn teaJ:mic|u®e were the r e s u l t of n@m disooyex^es t&
t^ie aeireiities In t h i s f ie ld* 0iich di f fcotent ia l banditig pattexiia,
usual ly otosenred a t apeci f io regions on pf i r t ioular olircMoa^^es
are 'ueimg ine reas ing ly ueed fo r the idei i t i f ie&tioB of
iudivldiml ohx&moBmmB*
In the preaeiLt work the Q bai&diBg mae iiadueed In tae
cliroiBosomes of Indiaii f i ve s t r i p e d s q u i r r e l , I'UBuabaluii
peaaaat i aiid a a t t u a aorve/^iotMi. with the help of a p ro teo ly t lo
eiiayme- t rypa i a aud the organic eompouad urea^ u t i M s i a g ta®
d i f f e r t i i t teclmiquee.
The a i r drying mettwd of s l i d e preparatioaa naa adopted.
i i f ter the t reatment of these s-lides with t rypa ia or wlta .
urea s e p a r a t e l y , staiiiiiJM:; ^^^ done in 2y percent buffered
aiemaa s o l u t i o n .
l%e karyotypes nud the baiidiug |>atterns were ol*.served
oarefu:i.ly« a tud ies of ohromosoBal types ( shape ) by taJsiiig
oeatromerlo i n d i c e s , ohroiaosojae. m&pptuxg » ttie baadiag pa t te rn
aaaltV'sis and the aicrophoi»graphy of d l f fe re i i t ae t ap tese p l a t e s
vere aside under the Oarl ^^iess £a rgav i l r e sea rch a ie rosoope .
i t was observed tha t the Q eaadi&:. patterii© obtained
from both teoi-mlquea a re a l aoa t simil^ar as shown by the
cl;iroinoaoiae coaploments of the two apeoies*
1 f]
fw@ii% pairs of autOBomeo cwd one paii; of eex
ohirouosoaes ( x y ) have been elaaalfled into six groups
in aattiM aorveigioujB. Mo@t of the autosoaea siioved dark
bands in the Qentroiiexlo regio&s* £he bandirig pattern were
poorly <l®ve34»ped on the x ohrofflosomes* 2M.3 chrosoaoiie shows
a8yQ.ohrcin^ in females* She y ohroaoeome i e a amalX aorooeutrie
showing dark s ta in ing except in centromerio region*
fwenty s ix pairs of autosomes were placed in to eight
groups in the case of FttnurabttXtts yennanti* She sex ohrosioeoiae@
X and y showed poorly developed banding patterns* Hhe x i s a
large submetacentrio chromosome shoving aaynohron^ in ftmialee*
The y ohromosome i s a siaall aoroeent:dLc one saowing negative
heteropyonooity in setaphase p la tes of bone laarrow eella*
Here also most of the autosomes show dark bands in t he i r
oentromerio regions*
fhe karyotype of Jfunuiabiiilua oennant;!. having greater
nufflber of diploid ohromos(»aes ( §4 ) » quite a few of *<chiCii
are aorocentrio, i s ra ther more priiaitiim than '^e karyotype
of Eattms nervefylQiia whage the diploid number is 42 aiMi,«al|'
about ^ percent of the ohromosoQes are aorooentrie*
103
Albert Iteraxkt Karl fvedga a»d ATecy AtJai^dberg ( 1964 )•
iioffioiaGlature for e«iitroa@rlo post t l on oa
ahi»>aM3@osiea -
Meredita@ §2 s 201 • 220.
Aaifenseia, 0«B,( 1975) »B«inelfl®s tiiat gevtras the foMtag
of pQl^|»eptlde ohalns*, 3aLe&i3e> lSlia2>-230(1973)«
A Hobel pr ise lecture reoapi tula t iag tlie earl^
history aad reoeat progress*
Baker, J*B. ( 1966 ) Ojtologioal Teohniqi^e, London : lethuen.
Baks, a«F. ( 1973 ) Oorrel&tes of ehroisoeoffial Banding a t the
le<9el of u l t r a etruotiire*
Tutorial froo* Int* Aoad. of a y t o l o ^ 2, 33*7;^.
Bigger, t»'iULm and J«li«£* lavage*
Mapping CI bands on hvm&n propliitee ohroaoaomee*
ayto^enet. a e l l O^enet. 1^(2) $112*121 iUms. 1973•
i^rown, Eon&lA, It* , sl.fatimK and f«a* Ham*
the Possible Eole of Mistones in the Meohani^esji
of aurofflosoraal a iJsndlng
^eienoe (wash, m) 139 (420d)s 1Q9Q«1091 illU8.1979.
BttG^^«4and, li.A. and H«J. Bvans ( 1978 }•
Uytogenetio Aspeots of Phylogefly in the i^ouidae
IX a-banding. OytoMnt*, Cell aenet. 21$ 64-71
OEU, £*H»¥ and J^eMer* M»A. 1920.
QytogeneUds and evolu^on of j^simates*
haxixm I»X» Aoad* iJoiOfiQei 102 ; 2yi«266«
Oliopra, V.L* and Pa l , E.4, X96^«
Ch2 mos0@@s of the Indian Fiiro atxlped sciiairzeX
Chaoipaka aeahadrl* 3«E»?« Hao and f«a* Shah*
3tudl«a on rodent ohj^oiiosomes X- Tl « sax ohsoffiatia
and oh£O£io@ome0 in giantaalmXaja p e n n ^ t i
ohroffioscm® fodayt ToXuma 2 : X4@«
Caapasaaon, f*» Zaoh* !•«» and Joliiineaonf C» ( X970 ,, )
AnaXysia of tiiiaian aetaphaaa oiuroooaoma aet bf
aid of B«H.A« binding fXuo£«83«nt a^nta«
%•%:$* aaXl Aea* 62» 490 • 492*
Oaaparasonr f«, Saoh, Lf and Johanaaont 0* ( X970 b )
Xdentdfiomtion of tiHa ?hXXadaXpMa ohroiioaoaa ae
a nuBbax 22 b^ qiilnaoidlna anataxd fXi»3i!^aaant
anaX^siat
£xp. OeXX % a . 63, a3S*239«
Claapeiraaon» f«, I^makka* 0, aiid £aoh, li»( X97X )•
fba fXiu^i^aaeant pat tarn of himan matapbaaa
ohroiao0om@d diatlng\ilaM.ns obaraatars and
variabiXity.
liaraditaa 671 89*XD3»
l o :
Caspersson, f«, HuXteia, M»» Linda ten» S*, w&d 2^oh, L« (1973)*
Id tn t l f ioat lon of ohroaosoa© blvaienta in hwmn male
melaals hy qulnaorlne mustard f JU&oreeoeno® anal^eie.
iiei?edlt&8 671 147*149.
Clark and F«laeixfeld llatur« Hew M&logy 240 # 226 1972.
Oomings, D#B« ( 1972 )# !2li8 s trueture and famotion of oUroaatla
In adYanoee In Hvamm aenotios* •
( U.Harris and K.HlrselihorB, eda« )
Vol. 3 , pp 237-431.
I'leama Wse&ei, li«v lo rk .
Oiit®r«lli, B . , a a r t l OhiarelU, M# aad 3haf©r, D.A. 1972.
denstioa 43 : 190-194.
Goings $ ^*B« (1973) Iz^ ahromos<»&e Xdeatifioation eds. f.
Oasoerssozi and L.^dh» lew Xork $ Aoad^ai®.
OhiareUlt B« 1973. E^fOthetioaX meeliaxilaia of aotlon involved
in producing baada witli trypsin and other
eubstanoee on chromoaoiies. Maii»m3Jlaia, ohrom@@ame
levfi l e t t e r , Volume 14 • number 1 , 1973«
mmm, fl.J. ( 1973 ) B r i t . Med. BuU.. 29 s 196
aargf S.3. aad Sharaa, f. ( 1971 ) Karyotype of the iadian f i w
striped squi r re l fiiaua^bujlui P J S M M M liStMl^Mfe
( aoiuridae ) Hodeatia.
Oiaromoaome iafoaaaatioa aervioe Mo. 12 , 1971*
Haaertoja^ J«L» Mng0ir» M*f • l^ttttoat i>«B« arid h&Mdf l.M.t 196; *
fli« iJcmatio QhircMB08ome@ of the HoitloToidea.
aytogdiaetles 2 i 240 «> 263*
Hsu, T«a* and J*B« i^attoa* X$6$*
Bone aarsow psep€dimti<m for ehacomosoae studies*
aomparatlTS aa»eia34m& O; tog0&etl«i@»
@d« JLurte. Boiiiysoiiksd 4S4«>460*
KliAger* H*P« 196^ l»« flie Sooatie ohrfmoscmss of aose prisatfs*
EatOt H« aiid fd8ida» f«H* ( 1972 ) • Baxidli&g pattsime &t Qhln9s%
liaaatsr ehxoiBosoiaes r^vaaXsd X^y new teohiiitues*
Oli}roao3€»ia 36 fl 272-280*
Lae, a«L»¥, J,f* WeXoh aM E«J*S* Minsev ( 1972 )•
iiandij^ jpattems in Mtfflaii QhxQmo&Qm9& ; Product!on
hy protool^'tio ansimaa*
MAi(lii«ir» E« 195X* flie ehromoaoawa of irt£tib3»t6«
Mir. (^anat* 4t 159 - ISO.
Hazma a.K. and M.faXukdax ( X96§}.
Manallan etoomaaoBa U9w® l a t t a r 17• 77.
ajelCYaag* Aaaseiy Ka^ahallt Joa, and acopp, Alfa3:d.
dhxroeoaomaa of rt ta m-A lalaa of f ijdland •
MaaaXiaii oliroi&osoiae nawaXattax (1973)
¥oli»ffla H, Bo* 3* 91-92-.
•-...' , i
Miller, 0«J« • 197^ • l a 9l»«oaioso®© Identl t ioat lon*
Bobdl eymposia 23 t Mevyosk : Aoademlo*
lladler* u.£'« -iiid button* D*A.» 1962, i!i1»}tic oliroaosoaee of
some Nortli Asieelaaii OMpuriks ( Jotilrldae ) belongJUQg
to geaom feolao aad Butaatas.
Qhromoeoaetf 1^ s 1 • 15*
lad le r , Otf* and UuttoE* i)«A* 19<I2« Mitotlo oliJ?oao80ii«3 of @osao
Hortli AjiMi£ioaii kioulridme
l'iroo«f aoe*t Bxp* ^ l o l . M«l« 110 i 3h»3&
UodlBX, a*F«» B*M»Jbay C 1967) Ohsonosomes of some spoeiee of
Mecltmea ( Hasmalla t liodes^tla )
£• ^a%e1:i«ri:* 32 t 2B5 - 291-
Petezraon, A*^« ( 1964 )• B«M#A* Syntlieale and ohrosioefiiBia a aynensoii^*
4»0ell* J iologr 23f» 6§1 • 6§4*
i'airdue, M*L. and 0alX «f«(l« ( 1970).
ahasfGs scHQal loca l i sa t ion of mouaa a a t « l l i t a D»ii»A.
Soieao«. I«I« 168 s 1^§6«
Baria QonSeisen^m { 1971 ) Stax^axrdisation is. Eiman aytoga^atios
( 1972). Jiirth JDafeote : Origlmal Art, iier* VXII 7»
Tha Bat* Fomidatioii Mareb of Plm«8» tlav Itork*
Eobartaon, tf«H«B,t 1916• QJtiroaosoffle 3tudiaa tsuamasmlo
rolatLouahips aliowu in fettigida® and Aorldidaet
Looustidae asid aryll ldaor ehroiioao»ea a&d
im£iatioiia«
^•MorpH, 27 : 179-331.
11
Eaot S»E*V*» aeahadidt Champaka and ^ihahf V«c;. X9$Qt
atmdles on rodent ehx&mo9m»& of m«Jbta<i* lllXXagai..&
fl^eltada ( ^ray ) aM an aooeimt of aia a b t r r e a t
^as jo typt i n a mal«r»
a\»rr. i^oi. 57 ( 11 ) t 50© - 309.
E^der, o«4«» lUO* Bpel i K.Bdndrsohkd ( 197B)
Chromoaom® i^andlmg 4ltudl«3 of £t^<^&®*
Uytogtwt* G e n . QeneU 20 s 52>-550»
a.E*V«£iao Bd ^harda ( 1964)*
aoaat lo ahroaosoeits of I M l a n f i ve a ts lped aqtuirrdl
GytQgenetLG&, 3§ 342*
Shaxaia, V.K*, Eao 3«A,V« and Qlmh V.O* 1970
Masia3J.aii ohroiioaome*
I n d . J*B3Kp, Mol# 8 J 279.
3«a>^rig]at, M# ( 1971 ) .
A rapid b&'a.dlng taohaique for liumaa ahroeoaoaes.
fhe l anee t I I , 971-972.
aaarma, A. 1976. * She ohros^isoffiea* Mohm. f s l a l a M ,
Oxford and I B H l^ubUshing Uo. India 1 « 286.
ilh«r,Ali C 1979 ) Ohrosoaoaaa of the iiouae c r i eke t A e h e ^
doiaesticaB ( $r^a.Iidae : o r thop te ra )
I M . J . of Exp. i s io l . Vol. 17 : 1030-1040.
11
^Hite, M.J.D^ ( 1977 ) Ajainal Uytology and evolution, (aaaoridge
Onivesridt^ press* Cam0rldge» Xio doj )•
XoBs, 'd%J* X969* karyotypes of Ha la^a r a t s ( fiodentia -
riurid&e, aen\^ Eattue Fiaiier ) •
ioaidat S«H«t ^suohi^ai IC*, Xmal* H«S«« Moxziwaki, K aiid
Udgawa, £• 1969*
ahxomosoiiie BiimbexB of rodent epeoles in aoutb
Bast Asia aad yoeania*
M g M 10, 217 - 219.
^ miSf J*tr« ( 197?) Oonstltutlve heteroelirom&tlii md tbe repeated
D»1«A* gequenoee of aasasi&ls* lu ^Modera aspeota of
e;ftQgeiietiQ3« Uonetltutive lietero-olirosiatln in miia' •
dympog^a ffedloa Uoeohat 6, ^ 1 7
( Htk* f f e i f f e r , ed. ) F*IC« aotiatlaiwr Verla^^,
dtuUgartf UBM ¥ork«
tvmla ^a» and danohea, 0 . ( 1973 )•
Of ijimding and ohromososie s t ruotore vJliroatoecMa ( iserlia ) 44t 1S«23«
¥imiSf Jeorge, J« ( 1974) - Husiaii ohromoaoiie MethodolOc^^*
AoadeiBio ^resa Hew lork axid l«ondoa«
Xosida, I.H, 5 i*.iiaga4 ( 1975 )•
Variation of 0 <* bmn&u Xu Ifoe ohrcxsioeiM@is of
aeTeral eiatospeoiea of iiattu© ruttua*