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A Cc!iiip^riti¥e Stody ©f teie Karyotypes of
Coin^bidie
Dissertation Submitted in partial fulfilment of the requirements for the
Degree of Master of Philosophy in Zoology of
The Aligarh Muslim University
By
ZAHID ALI
DEPARTMENT OF ZOOLOGY ALIGARH MUSLIM UNIVERSITY
ALIGARH
1979
DS55
Oeilioofeed
to my
A COriPARnTrJE STUOy OF THE KARIfDTVPES Of THREE DIFFEI EIT
8r»EGiES cr eiRos mm THE rmivt cQUimtmt
Syboitted in poefeioi fulfilooot of tft© !?®t itQi!s©nte fo& tli®
Oegpe© of d^tci^ of Ph&loeoptiy in 2f3olooy
of
Th@ Al igo th {%j©lio Univorc i ty
By
Zahid Al i
OapartmcPt of Zoology
AUg.^rh ?ljsli@ Unlvereity^ Aligarh
1979*
I certify that *A coaparatitf© study of the karyotypss
of threa dlffarent spsoias of birds froci th« f^milf Colu»bid«»»
£e the otjiglnol uork of % . Zahid AI4 end i s suitable for par t ia l
fuifllmsnt of th© rec^ulrciaefits for the degree nf f1a®ter of
Philosophy in tml&qy of th© AliQorh fkmlim Unitferelty, Allgarh,
The yorte ho® been don® tinder lay eiiporvlaion*
(Uemr.n (1« Adhasji) Ph«D* (^otr© Dane) Reuder^ Oapartraent of ZccXoqyf Aligurh PJuelin Uniw0!?sityt Aligarh*
( i )
A C K ^ 0 U l . E 0 6 C ? l £ : i S T S
Th® out'-pr tdkca an op30rtunity to express tfte deepest
sons© of grotitud® t o Or» t'tnoii {1» Adbaal uhc helped at avery
portent to oospicte tti® di©eoi?featicr} i n the precont forra^ lUs
o rd t iou l advice and ^aluzblm sugies t icne ure qrs;!tly admired,
Gr-t@ft!l tfif nk® or® ^Ivo dii® to Profot:sor S«3, Aiu®,
Mead, Oeporttisnt of Zoology for proyJLding n^oescory fooi l i t ie®
und onoouruQGJJcnt* Thv-nJi© apo cico due to tha Otilvercity Gr^nt©
Co!n?!i9t ioo for fincnoioi cooictcoo© durAng tho terajre of ttiAs uork«
I uBj grea t ly thonfeful to ny elf© rir®, Tolct Eehid uho
h(5s given m@ :;n opportunity to u t i l i z e the woxlsiuc} time in
cnnr'uctlnq th® res© ^rch ticrk wit'ioot any d is turbance . Her
cooper-t ion in oheckiw^ >in6 orciparinq the the« is i s hiqiily
apfjrsoiable*
l^oat s incere thurtke are .Iso tlue to a l l the cnlleaguee
of iRy lab* for t h e i r oriticis«B .nd valuaMe sugieetione*
<u)
TAste or cofiiTEfrrs
Aoknicyl@tlr|@®@nte • • • • • • ( i )
Table of oontente ••« • • • Hi)
t4®t of Tables «.« • • • (iH)
List of riQUrOS e«e »•» (iV •* M )
In t roduc t ion „«o e«e 1 » 1 1
I^atoriale end ^ t h o d s «»« 9«» 12<»16
Oliaorvatlcns and Btsul te • • • «•« 17 • S I
Tabltts • « • • • • 22 « 30
Olseuttslon ••« • • • 3 1 - 4 3
Conclusions ••« ••« 44
Suanary • • • • • • 48 • 48
Raftttamsas • • • • • • 47«*S4
riguras • • • • • • SS «* 9H
iiii)
iisT or rmim
l« ShMO|tii| {fiploid shroKp^o®8 mifflbef in thtBm speolss of l*i®
order c:oly©b4fotffl®s»
! ! • fei^ticsett^ of ch£i3fflioG©BJe© of th® SndiGO docicstie pi§son
IS la notphcja©tr^ of oh?o®oooac@ of th® IndSors blu® roc^ pig@en
ltf» ne^tioisotiy of ohFKaoccffio© of Indian l i t t l e btoun or
^« Cofflpati®ion of th® Pfeafi Ralotiii® iangfcti® (t^} of Indiou
dcwestie pifson (Cf lyBil3t| JLiwla ^oiaes^fqa). Indian felo©
tod< pigmsn fe^t^H^ AluiQ |f^1^qnn^lo, end lodian 11 t t l s
btoi^ dov@ Cet,ff,ff f p, l,i„q flfn»ly^,«liJ|, ,g*ftaYjnali)»
tfl* ConparitiloR of tli@ »@ar} folotivt length of the longciimi
of tl)» chromoscnes of Indian doffi@stlQ piQeoity Ittdian blum
rodk pl^son and Indl^iin l l t t l o btoun timtem
tfIX* Co«parl@10fi of the Pis@n r@lati¥B l@ngth of th« shoirtaxms
of thtt ohtonoscffies of Imriafi doa^stic plgeooy Indlatt blu«
ro(^ pigeon and Indian l l t t l « bm%m dovs*
unx« CUswnarittlcin of tha m^^n centroMsrle lndac«» (X.) of the o
ohrcmosoiRiie of Indium dowestle pl<}«on« Indian blue rook
pigeon ^nd Indl&rn l l t t l a hroun dove.
UST or FIGURES
! • • Ustaphgse plote ehoulng th@ soisatio {naoroshromnstsiM of
eial« Indian domestic pigeon*
lbs Lutd diplctene of tho prinerjf spofputooj^te ef Colu^ba ^iufa
|doffia»tioa ehoying the tmn lavg@9t pairs of (OvmiroolircfiiC80R«»«
SI* MetaphcGs plote ohouing the sofsatio oaoroohromoGcass of the
f^iaie Indian do@8@tio pigoofi*
III* (lataphas® plote ehouing the sosotio oooroohrcsooeomo® of mole
Indian blye rook pigeon*
X * :%tapha8« plute shoying the eofsetio mooroohromocc^es of
fsmalQ Indian blue ?oete pigeon*
II* f%tapha@e plate sho jii g t e eomutio mocroohroraostntee of
eale Indian l i t t l e broyn dove*
VI* !1etaphaee plate shouinQ the ioitatio sacroohrofnosotnee of
fefftale Indian l i t t l e broun dove*
tfll* Karyotype of male Indian doaiaetio pigeon uith zy of fenale
in the ineeot*
Uni* Karyotype of «<4e Indian blue rock pigeon yith ZU of fenale
in the inseot*
IX* Karyotype of male Indian l i t t l e brown dove uith 1%! of female
in the insect*
• • y M
Km Itiloqt'M of th» sonatio naorochrooosonsv of Col^wba ^f^jlft
XI • Idieg^aiB of tH« «o«iiUo isaoroehreifioeowtt* of Colupiba 1,4MAI»I
XSI« IdioQr :!!! Of the scmmtio eaorootitomosotBee of M££&l£EiMS
XIII• CoBjpai&Gtii;© idiogrua of th© ©caatio tasorochrooiosoQes of
.Sfcg.@{a.t.q@mll. g.ef>aQc.l0flQi.B fitMt3.a^q.ogla>
lengths of the ocsotio ooorcahrciiioouisee of CoAtif8fc((| AS igla
XV. CoeiptiaStd idiogr-3a of the ordat GoXumbiftitmmB (.Hffe©if Ray-
Chaudhuri et al*, 1969)*
JSJ.JfLfl
Th© recant ptrcgrofis in karyoloqy cf unimcX» i s vary
rem^rk^.b-'-a .Jid the prnhlsm of rsoA. 1 d i s t l n a t i r n re l .ting tn
chrcBiosrm-il features i n dtmoetio un io . le i s baavming .n in teres t lnr i
(>hioc3t c-ttr^ctiiiQ thB speaitil a t t cn t i cn cf cv'toic'iiitte. The vovk
of Hafeino en mia& ( i 94 t c ) , ptl '"eld flKlipcn crSc© unci euts
Ci941b, 1042% b» £3, 1943Q) -nd en ynqulotca (19431:, o, 19n4a, b , n ) ,
ccntribirtcd c o-o-^ cJe-:;! i n t h i s f iold of biclo:}y» The r c j u l t s nF
thoce etuCieB did not c'loy polyploidy or cny bind of £r^80"A^-^^ty
i n chrcoooctnoe i n the cnicaols otudicd* Contrary to thio^oo i© u c l l
ftnown, polyploidy -nd vctiruo kindo of unucycl phencsono of c h r c n o
cooes ore Fwthor ccD'non cnong ttm !?cooo uid vfcpietics of p l^n to .
Provirusly aoet of york uco dono en the dcmnatio few!
fhinh tj-s !;up^GS0d to b© o favourit© ©vstcri*;! fcr res: -rch in f-is
f i e lds of embryclcgyt physiology *nd gcnntie invi=stir}>.tlrns» The
picnper in^estiqwtfers uho desoribod tho ohrntnnsnTnr.s rf tfie fnyl ft»r
tha f i r s t t i i© soems tr bo Usyez (1006). I'm desaribad 6 chromrBDw-E
to occur in the nicl5 ' s-y nocyta. Than Scnnenbrndt (irCIP) Pbserued
W - 16 chrcnnaofii0s in tha primary oocyte fnlloued by LoiCiiillon
(1910) who rsportad 12 chrcmosoraea ih pc^rthanrgenetie aomatio o a l l s ,
fijyar (1909) i s tha f i r s t uthor ct nnermtd vith Hha chromrsowaa in
tha aparm i togrnial ca l l a -nd ^ chroproscmoa in the primury sperwato-
cytet ci-iinting tha dlaotjvr-ry of . n unpaired sex chromoarrat in the
wale of donaatio fou l . Houatier, tha rasul ta cf the qanatio exper i -
raanta nbt»»in«d .»t that time, pl*«inly showed tha t het-rogamat-y
exiptad in the f«w.*i® -nd fitit in the m .le« In i^ iS Guyer ^ttsrapted
a rcviseti rtudy ucing t h r s e r >C88 of foul» th© icnqahsJfH thK Pi.ymcuth
i?ock ..nri the Rhfid® Ssi^nd Had und nstf© the fcl louing oGtrenticn*
The upmtm^tv^rtni''-X chrvrnvw-mnB ^^tm oafnpns«d nf IB chrompstinios which
include ^ p^ir of Jt-'irgs V ©hc.'jed eles^cnts onnsitlered am th® sex
ohn^nnrGfses* Cofcier (iCiG) <)l50 m<:ti@ obscpwctims ttn germ •» CBUB
cf th© Gcltj C«-npino foul ;.nd QC^© 10 « 20 ohrcr;oonr;j>n co e^lplnir:
ond n » 10 clcTscnto no hapAoitJ* Thnso QA^C 5.Q.:I inves t iga t ions
aXocvly ioc:loc;ted thut tho technique ysod uuo tinswfeicfootospy fcr
tho rirooir-o oturfy cf chpcoooonss in tho fCL*3.» Tha chrcaoGrmcG
being ciynped toQCthcp ^nd not clAooino GOCUP^^O otocIy» The 6^tc.
prooontod by thcco - *?lior uoEtts^ ewcn though ic:'30i?t::;nt cti t hc t
otirao huw© bocmc iv^ftor cut G!®tod co they ohcu no corrcot p ic tu re
of tho chrtsrocrioo citho£? in naobor or othop oorphoioQiool foc5turse»
St i ' oerti-in thot tho chfonoocR© otutly of tha doosctic
fryjt adti.nsed on© step by the imtonsiv^ forfc of H..nce (2.923^ 1924,
1926 a, b , c)« By h i s e f fo r t s i t bea^iue avidont t h - t the chrnmo??0?nea
ocwolex nf foul c rnnis t of « CB^taia numbov cf m.^oro^chro icjicf»08
huvint? s Cefini te sh„n« ~ncJ a luvqm riuphsr cf very mi nuts gr'^nule
l i k e wioro-ohrowcsofflaa. H® reported (1924) tba t the re were 3Q -• 34
ahrc?i»J0HrRi8s in tht? sora^itic c a l l s cf cods ^ nd hen ^nd tha t the
larg«8t \t shaped el^went e r i s t a d in the n^sired condition in ^ e
naJLe CBXIR while i t w- s unpaired in the fsm^^le c e l l e . By t h i e
evidence as ho thinks ftfii.ile hotronemehy i s cytrlnqi0'4.^y proved.
In hie s'jtoaesaiue '^turiy (1926) he q-'ve 40 ^& the l a rges t d iploid
nuiabor ihJSch h» anuld rbaerved in the netanh^aee nf the soB»«tic
cBllm of c^ijck^n. The Invest jgat inne nf Miller (l'"'3«^) and
3
¥»..Bt»ishin.Ji ClW2 nti 1944) mhi^u t,h« fee»t renul t s tha t octjld bci
obtuifiit'ti ' i t h the tachniqu© of diiroot f ixa t ion and sec t l cn . The
vH^lysls of bi rd ohrcnostro© tjculd nnt^ housver, ta^kii furthur
'!rn*^"cns u n t i l nmv -sr©p^r..tii n tsichnSque h d bteo ir rkpd out ,
Around I9S0 tJio ©qsiaeh-tsofwa" qu® b©gw!n to pepluO® sscttioninq, o
hypctcnlo p rc t re - tncn t of th© o@lls before f ixa t ion u^s introduced.
fhim Quured a csanoldorablo itapstswcoent i n the pcsc i fe i l i t ies of
GtodyAng th© ctircecsnjans in wortefcrwtBs* Lcter colchicine nnd
e ther chonicols how® JLGO been ocssbiniHa u i th th© hypotonic pr®»
trcwti3ont« Won Brink und Obbol© (1966) o®od th© hypotof»»squQ®h
tcchnic3,'o en ambryonic tioruG© of SalltiR rernestlcti.s» Thoy found
tho ohrccoccoo mobor to be obowt ?Q end csnoiyocd tho ott?uctu^€s of
ecven blggoct chrotscocse puirs» dooicnoting th© E chrcacooia® »o
pviir nunbcr B« Thca© findin§o oro in ugroGDont u i th th© ofcQcrwo^
ticn© of i l i l l a r (1930) end Ycscohinu (1044)* They did not ®oocesd
i n deciding uhother th© fen.;!© yag 20 or 2U» V-n Brirtfc (19B9) used
the Sv-roa teohnit,u» t,'h©n she inuostigi^tsd Gallmj gallog^. t'lglcpsitt^a-
a m ttP%,^.vli,^. -^'^ ?.MM^ IHSSMJSSm* ^ ® k-ryotyp® of S i i | | J£
oallug u.?s th© s <P! • lis the feoryotyp® of S.mojs rfcwieaticua.
rielopgitt^iOt^s isntS^X ^tu^ h,*d c chrcscscfso nurafeer of .fcr«ut "SS of
yhich 12 puira yero pointed cut <m (B<^cro-chrotao8C»e«« Chrfmcsoma
number B u39 the Z Ghretttrscfne* PiisRgg doweaticua had about 76
chronrycraea, 12 p j i r a tfsr© macro-ohromnscniaa - nd the Z chrnrnnartwe
u*J8 nuwh'-r 4» In nnne of th© apeoiaa oculd aha d i s t inouish a
U ohrofnoBona*
Tho 1 rga nuiabar of chrt3Bjr!sr»«aa In birda ^nd tha iwrga
v a r i - t i n n in aiza h*4We e n t a i a r i t h a i r bolng usual ly cl<^aKifi©d aa
maorr >.inci jaicro-nhrontttsaroes* TfiB Rixo of the R)aoro>chrc!Bi08riBii8
i s ^bnut 3 « 7 /^ jnd t h s i r »truotur« i s coop .ratiwoiy mmy t o
anivilysB* Tho raicro-t^romosaaso at© Bm.sli, n#3 t o about 3/*^ and
t h e i r R«ntrc3ro©ra i a t a u j i l y i n v i s i b l e ! most of thmm a re probably
t©lncentrio» Protfioui ini#e8tig*itor8 hc\t& u3u-41y frund a d«f in i t«
dividing l i na &©tu8®n caaorce and laicro-ohrcsiaoecaje® o*g* i n Sallu.a
dcmgaMcua a f t e r pcip nuabsr f tho ohrcmoccsea or© aiorowohronroscBies*
f%ucit a©r -nd " r . n t (I9S4) vjnd ^^iswocmorllSS?, 1980) s t . tti
tfi«jt the difforonc© in racoro end niorr!«ohreB!oecmo0 i s quclitc^tive*
According to thcia in Ga itJ _ domratious tfs© miiaber of ©core-
ohrcooGC5ii@s i® 12 i n tho ool® .m6 11 i n th© female cind oil o ther
o^riMrsoacisng- cul led es cSiroscsc^ida, ci;tild be Q Dort of rooorve
mtoleio coid« ^eyocsaor v<is un«-!bl© to fiis Q de f i n i t e ntimbet of
ahrrmoccraes, heo -uca th© small chrcncscsiea uore often etrcnrjly
h8t9roobrc«wtio« Donnally .nd 'h\jotm&T (1*?63) h->v© t.-ith th® help
of autoradiography found tha t the small chrnmnaoraaa eynthaaiza OM
e a r l i e r th^n the l a rga ono©« Thia ^^lao should indic^ ta a q«d.:.ltatiwa
differenca betuaon wacro-chronoKcraaa ^nd chromosoRioida* Sawcnwar
u i sc rom-'rkQd tha t Hutt (1949) had raportad only 6 coupllnQ gxoupa
During a period of about 10 yaara upto 196S aevoral
invast lQatora hava axamined bird Karyotypaa to find nut uhathar
tha 558x-»datBtwining neohaniafa i a ZU/ZZ or ZO/ZZ nH thothisr tha
nicro«chrow0f?f»Bi©B .ire ra^sl chrowoaowaa or only reaurvs Of A, Man
Prink >na Ubb«<?« (19S6) >inti Van Brink (19S9) found tha t tha wicrn-
chroBOBOWaa behawad l ika ordin«»ry ohromoBCwaa* Th«y uara, houavar*
c
0
1 tht^t i t M«» d i f f i c u l t t c fiK t h e i r ©s^-ot mimbotp .mi! ife
ut*« d ipf iou l t to daterminw uhioh yers cfiroisatid® and uNich wer*
chfofi?rso»8s# The ifi¥08tig..tSof»8 of fshfio (IfSlJ in raltosis &nd
m®i^ii9 if* CaJ ltiB, dcaai i ^^qufs. ehnu t ha t tfi8 ©icprs-c^irowosoEift® ar®
ccsnst.tnt in t h e i r nosber and tfmt they o- n fo© foliouitd through
eo^'-jliitii nitoei© end oelooie^ thmit b&h^micut net diirerging frcii
thm s:»*oro»ohffoooGCi3e8» Krisharj (i964) fcufid in the n-JL© neicei©
f* >'tel@0QgiB. OQAXOO. tfo tli.it the nyiabef of 6iiw«4ofit© dos-ing dlp^.ntono
of Gotaphase onm ^VB 38 •» 41 cnci there i e no diffrrenc® between
aaoto iiRd oicFO oicBtaento* With the telp of l i g h t ©ictcseop-^ unci
el©otrcfi niotoecop^ fotd i..r}d t'ooitics (1984) ahoyod tha t th@ so
GQiidd ncoro -i nd olas«*chrcc30ocooa onijf di^og-gs in e i se tSorino
raitocio -od ooiooi© ond net in bcf^o^icye*
The nocyrcoo© of o y»cfircrnocooe in tfso fcjocii© yos Arng
dcobtcd tut In 1982 Cohnid fojnd tha t besides the Z tfier® i s ..n'^tfint
e ingie c!iroQor.c8io i n the he t rcgans t io ©©» i n Oal^Iys t^pffie^tlcus» Tine
of th© wioitJ-chromcjjrfnes nhoued ei latm .^d very s t r rng incorppr- ' t ion
of t r i t iv i ted thyiaidin© -^rt^ t h i s chromiscsa® h@ narks tm the ?-'•
Fey attarapts haya alac bsen raade en t he gsnatlcrfl -and
cytcloQio.^1 inyastigftt ion* on biri! hybridisE«ticn, Th® papera of
SokQloyskaja (J193S@) published *n <i3«t»nsivs stud^ of th© nmx l inked
oharoctsr© in th® int©rg@h®*ic hybr id iza t ion , togwthar with th©
restflt© of th© tearycslogieal in t fss t igat iont of the hybrid b«twetm
the Pluacovy duck (Cuirine* fno«<; hata) md the dniaastic duck (Ana©
nlal^yfhvncha). Ctmu - nd Keller (1936) m^d9 a cytological
rn the parenta l fnrm© and t h e i r e t a r l l e hytSrlds. Tdniuehinu (1942)
6
h^n qiv®n a brinf ucccunt on the ohroracsCBre© of the Huaoc wy dtdk,
th© OoBostlc cfuc3k nd theiv hybr id , to dasoribe tho kciryety-jcs cf
these fovm9 e>nii ot the Bera© titm onrreoted th© arr^nefjus d^^ta
nffprad by ptm/itnim 4nv»«tig..tors en thsati f o r a s . fllssn (1960)
reported s!ieons©fui in te r f^mi l io l hybr id i i a t i cn of Dark Cornish
chickens end B©it®wiilG sianil uh l t e tusfrcys* Thooo hybrids cctnfcinct}
in a r4n«3l® phonotyp©, the darfe piucsog© end f9JthcrQd ncofc cf th®
f'aife Cornish ohidien vith tho long ncok, uh i t e rMin .md long t « i l
fect'.'JcrG of tho Coitsvil^© email t^hito tus^cy. Thoig ^-^to of
Orouth ^nd body u©Aoht ?cll bcttj®cn ths ROKIOI© for t h e i r parent. .!
cpsoie©* Aib?icLigh ooirorci ooios litiod t o un ago equiw-Jiont to
oa tu r i ty fog' oithoi? of tho poi?cntol cpcoicog qtrw perJditssd any
OTOcn c r di©picyud ony ©QKUOI beC^cwicus^
t"n in tcnc i^e fork hcs bc^n dono on tho DoncJing putteim cf
n.-:r:':^2i .n chrcffirscmc©. In fetj cpeoics nf bird© otter.nte hnv© hj»* n
P!-»d8 fcr the study rf i n t e r oh.mg© c n ^ y a i o ^ nd ®8js i d s n t i f i c a t i ' - n .
Uang tind Shofffier (1974) ad.^ptsd tho t ryps in bimdim tyohnitty©
Csnabriqht, l^lf Outr i i lans e t t 4» , l ^ l , Ki^m ^^fH fedorrff , 197?)
fnr ident i fyinq spetsifie chrcmrsnme© . ntl chrnnnspaie <dtm9 ft th©
chldkan. They huv© dascrl^sd the devolepsBstit n6 ^pplic-Jtion of the
t ryps in fi-banding tnGhnqvm for ident i fying indirfidu 4 chrcstnrscjne
cagntante invoivet' i n induced chrcjtnoecfaol ro-rr» ngmsants in th«
ehicMcan. t larrids e t ^1.(1973) tcrfcing u i th huin* n ohrcposnuies,
fnund th«it longirr tryt3sin t r e tmBrtt u i l l proqrsssitfrly it: d frnti G
b.^ndm to C fo'^nds, Stephoa -nd rtrrighi (1971) used the heterr^
ohrcM«atin st<4ning wathrd nf wrriqhi jnd Hmu (1971) *nd described
7
80X chfnmosowa in birds* Tho
Obanding producsdl by the t rypnin taehniqu* j l to produoM a highly
heternohrnmatie U ohrcmosone ani3i horn po t r ib lo oppl icat ion for nm%
i d e n t i f i c a t i o n In birds*
Th® &»bund pos i t ive hetaroohroeiatin has h i t h e r t o benn
etudled only In a l i ts i ted number of o^lan ep@ol&« \flzm chicken^
dQtf@f pigeony purakeety pheoo^^nt^ duok^ Sapunece quuil^ quail imd
bold ec^Q (StefOQ cnti Arr ighi , 19719 Ccmingo end J^uttoocic, I977t
Stock ot ol»|, 1974s ^ (3t ol«f 107S}* I t has been observed th^^t
(o) The y 1© the only hotoroohrcootie chrcraosocsiG in the
parakeet , pheoccnt, due* end q y a i i . In the dovog plfjeon,
3epcne©e quoil^ bold ougie <ind chiclton» the oontrcnorfiS
of 8©vorol ©lotos or© olso Obond pooitlV0«
(b) The ci0nt--"Trr:r~s of tho ouoro chrcmocotnc® or© e i t h s r devoid
of the C b<jnd or possess i t only in minute quont i t les*
Based on these im6 t h e i r own observations on r e p e t i t i v e OiA in
chitken Stefos and Arrighl (19?4) have sugqested tha t the hetarc^
chr'^watin in b i rds i s r e s t r i c t e d to the U ~^t\A wioro-chrc^tosrfSKie.
Alsoy because of the ubiquity of the Riloro«*chrcmreDirae <^intS the U in
b i rds as yall cas sn-^es^ they huve envisaged ^n iden t i ca l s i tu i i t ion
t o occur in snakes o lsn , nemurtf 3ucQb L>n6 Sharna studied (1978)
the heterogeneity i n d i s t r i b u t i o n of cons t i t u t i ve heterochroiidtin in
four species of birds n-mely pigeon (C^ltyba l i v l a ) . duck (Xtettao^s
cogo«andelianue)> nyna (, ti.P!Mff BfMm) ^^ bungle b<ibhler (Tur^oides
s t roa t i i s )* Dccept in ths pigeon Ob^nds uere Inealized mainly an
8
the m..oro»chrcmosnm88 where js in ths piqatin the mjlcro**chramo80m88
are nredr minantly &»bunti positive* In the t&jck h«terochrr»«ati n
ytt0 rsetr ieted to the eetttronere* cf only thrse paire cf naeiti«»
chir£mosoaMi8» but in oyiia e l l th© isaoro^ohroaaRtratefii and only a few
micro-chrrnRosoisest uere D-hend positive* 2n the Jun^e b^^bblett
irrespective cf the oliet «^®f st J l l the chrcisosom©© wet© ©ndr u'Sd
ui th centtoaierie hsterocrfircitatin. The y ohrcmescsje yu© entirely
hetarnchroaatio in o i l th© opeoic© utudiec!* Oniike ooinale yhere
almost ull th@ dentrotser^sure C b und positv®^ in biird® thore ie ^ n
cbviou© hoteroQenoity in the dietr ibyticn of hetoroshETSiatln,
D{3A repliocticn pottom© of the sea chrcocsoocs of the
pigeon CfoloPba AivA-fi ftpmi @.t.lGc>) hoc boon etwdicd by Gciton end
Oredbory |1S66). The i c te re:3liGQtion pattern of the pigeon
ohrcncocEics uas studied by csocn© of th© teohnicm© of torainol tipt'k©
to t r i t - i c t e d thymidine *.>nd ojtorodiogtophy. The y chroocscstri in
the fttfflule ocfoplct©d replication oftor th© other muoro-ohrontosowee*
The t chr(Moacrae« in the fesj^le « nd both t ohromosnrase in the m-ie
terrainsted DHi^ eyntheais ut the B^mm time <»e the retnc-ining ffi««Qro*
ohrnntoaoises* The oiiorcchrnffiosniaee uere la te r replic^itinQ than the
»aorQ»chro»ti8cnee» I t i e inferred that l a te replioiaticn^ ut least
in the c<ji8e of the U chroinoeomet eignifiee gsnetie inactiv^tion in
eonatio oelle* 'nie ubeence of asynchronous replication of «t Z
ehronasone in the honogaKotie sex suQqeete that eex determination in
the pigeon might depend on « gene doeage iseohanian similar to that
reepcnsible far the •eutoeexing* phenomenon*
Aneari and K^ul (1978) studied the Tr<<ineloaaticn hetero
zygosity in thi» birdf Lonohura ounctuldita (Unn) uhere the diploid
9
ohrcmosofio mmiaar y« » found to im 62 in one iialu und on« fmn<AX9m
c m ti8vsnt»80n n.^OfoohrooososAs i,n both thssaxsii* 6oth thd
inditfidutilt fihau en abbavont karyotyp« in having thr«« unpaired
Hiaero-ctifOiineonee in addition to the seven paired «a0]^o»ohro•losolle8•
There i s an unpaired 01 t^pe autoerne together uith ti.10 entailer rnaerc*
chromosciaee yith thai r oentroneree in the temtinal region* The
poenible nature of the trunelooati^ni leat^ing to the hetero^y^eity
i e ouQgeeted*
Th© recsist ©tudie© on cvicn chr®Boscisee though using
modern oytologiccjl tochnSc:u©» nr© hcepored by certtJin ontrinsio
Inaccyrcoies* Thceo centre oroond tho validity of subiaittino
ccisplete cut«out Itoryotype® yhen infcot the Bn&ot- chraaoscso mjisbor
of thut speoioB i s olaoet«»c©rtQiniy only en epprosiatation eirsse the
ohrciaoacKE nuobcr i s usurjly a sodel ^aXae in a frequenoy d i s t r i -
bytion tablo* 'Icreovar^ any oofflplet© photogr-iphio karyotype value
hQve to be f i t ted to thie nuffibar* In addition, ra.^y toohniu ;.
diff ioul t iee av involved in tho h^ncfjing of luioro-chroBiosoaMJe.
f i r s t l y the micro-chrowosoaee are quite l ikely to be over-lapped by
the larger elenente* Secondly the faint s tainin > vf the em^Xler
elemente often n^ikee the i r detection extremely difficulty i f not
iapoesible* Thirdlyt the dieintegr^tion of fRicro-ohromostinee into
eepdriite oonponent ehroiiatide may produce a f&lm» ohronosoiie number*
FcHjrthlyy the presence of artif<3ete etmh ds stain residue in the
ce l l s ct?t;Id erronetaysly inf la te the nicro^ehromosowB nunber*
Lastlyt there ie the diffiowlty of obtUning complete »ioro»chro-nosom& i
ooiaplement* These faotore in omibination are oert^iin to introduce
a large *i«iount of error when an attempt i s made to ooruitruct the so
10
cal led oowpleto kuryot pas* t^tmnvmt^ th» urlJlty'^ry wi?««n« cf
tiXusBiffinq th© CTicrn*chrc!«ii09r)B©» on »itu .*nd danslty of stdJlning
cftert yiaXti* fc^ryotyptta uhich hay« ahrtsnossfmes in p^ i rs yH«n th«r«
i s a l i t t l e iipfsarent hcitetioqy bQtu«nn thwm* A ftjyth^r pr^^ieBi u i t h
th« cot!ij3l&t» k ityotype ssotiid IJ» t h a t tt^er© **re, ws Cu«n (1565)
sugqeetad, ohyoaoscfass uhich .-le beycnd th® reso lu t icn ef the l i g h t
rjiornoocipe«
otanding th© ppincipic© cf koryctyp© otfoiution und for o l a r i f i op t i on
cf pC>yioc©iietio S'ol*-»ticr5ehip« Itie ccapurctiva kucyr»loQy of birdn
i e i^.ii99ir;j b&hind Gitaii<at» otudic© of cthas" OIOGCCS cf ^crte^^rutt*®
(t^Gptilio, nGG^ulo)* Cno of thQ Oojog- reccon fo© t h i s io c l..dk of
cosspafutiyQ (Si^lQ on gffcupe of oloooly to io ted cpeoiea* The scrpara^
tlv© 0tyc2i<3S on ocdsrn l eve l of prooodor© h-v© t:e©n oosrlod out by
d i r ro ren t authci's C^ailty, 1966| ulcers, ie69f Custtu^iojc e t a l . ,
1969| ItGh c t a l . , 1969f 3ciianoyia anC Atkinsf 1969a» IS i ibJ Hay*
Zh.mdhuvi ©-ti v l»» 19838 Hum>ier, 1070f P ias ln l .•nd S t e l l a , 1970|
PsdJ.J'ai e t *il . , 1970} Buldtowa et wl«, 1571| Tok s S a t ^il», 1972|
Pui .tc*/.:: Pit Jim9 19731 Thaodcroteu, 1978f Anauri s t u l , , 19771
Haul a t a l . , 197«=5),
Th« presont work yas undertJkan t^ T; k- a coiapati^tiv*
study of thu karyotyp»fi of tUrtm opaciss of tha ord«t Colunbiforaet
baionging to tha family Cnlu«ihida«, n..RiaXy C<? wb„ H v l a dowaatica
(Indian dcmi^atio nioaon^.CDluiBba l i v i i | irtl^Bimadia (Indian blue reck
pigmon) .md JUtg^fffff^l"^ IfnMa-^ffnfflff ^i^iXIOiiU (Indian l i t t l v
broun at aanagal dova)« Th«ir difsloid Qhronosoma nunbsr hat bean
11
qiv@n in tersit of th@ oiean v dun cf tHo tiiplGiii atmnt in eontr-i«%
to oth0r ynrkure uh© hiiw© olvnn tfm highest p©*ak c»f th« dipiniif
count in v^rit^io ii©tapha«# f3latee as tho diploic! f^sibttr. An effort
has &een eadB to determine ths imti^t of karfotjrpio similarity in
t' rtiQ threa olcrely ralatiKt 6p@oi@«»
nALf^Ri ^ A n iLM, nj.Lnj.p,.s
1
Ttireo d i f fe ren t epeoies of th© orc^or ColumblfcnsB® t»i2»
Coluuba •^iv;l,s dowgsUca (Indian dr^raestio pigacn) , Celuwba Ziyfa
4,nton .g jl,js. (Inc!iv>n bio© reck pigeon) ond p,fc,; ,opt,q.pfil4„ ft^M<3,1^,OT§Aff,
qawb.ayf?n8A,.s. (Indico l i t t l e broyo or eenegol dove) were s tud ied .
These spooles yop® i d s n t i f l c d u i th tb© b®lp of tbo Mend Took of the
Blrc:© of India end Peklstcn (All end Oil lcn, 1960)»
th© chrtjnoccoe propofution© of teenty flv© pcir© of etioh
of th® bird® tjsre nude cooopdlng to OIOOQ (1009) u l t b oortoin pcdi -
f l ec t ions to cu l t tb© onwlfforiDontal conditions* Tho o i t o t i c
ohrcffioscKcsB yers prepoired froa tbo bone«»oos'Fcy cei le^ tho ooot
cctlwoly prol i fopct ing c o i l s of tbo body tha t ffcpteoonto the best
foitotie f igure for the pyrpcso of feoryotype otudy# Th© s l i d e s sacro
prepared according to tho follouing prooodure*
Th© birds y©r© injeotod u l t h 0«2 nl of 0,0S^ oolohioln*
so lu t ion ner humired gr«4S8 body uoSght i n t r a p o r i t n n i a l l y <.boot W
iBimttos p r io r to fix«iticn» They yore thon eecrifiswd < rjd the thiqh
bona (femjr) y^e tcJcen out* Tha tyo ca lo i f i sd roundsd ©nde of the
bone uere elirjhtly nut u l th the help of qo€d sciosore t o f i ic l l i twto
peni ta t ion of the needle* Holee yere m^e at bcth ends y i th the
help of needlae und the bone mar toy c e l l s uere flushed nut i^ith
the help of « S ml* syringe having No* 23 o r ^Om 24 neudle oont^n-*
ing Um^% SQdiu« c i t r a t e (Hypotonic) eblution* uhich had been kept
in udtter bath maint<^ined ^t 37°C*
13
CvlJL*^ thus obtainedf «ifttir f lush ing uers suspandntf
pronar ly y i t h th« h»lp of pastuer p ipe t ta and kept f o r Ineubation
In a yatfiV bath f o r abmtt 30 iirinut«« £tt 37°C« Th9 aol t i t ian uac thsn
o»ntiriftit}«c} s t 24@0 tpm f s r 10 miny t i ^ * The supornatant uma d i t oa rd *
ed and tha poll@t ya« resuspunded i n the f i x a t i v e (3 s I f @thyl
u lcohcl and olacia j , oeet io aoid) >nd s>efriget>^ited cvot n ight t o
obta in etaximura f i x o t i o n * Sati@footory rss t j l ts ocn be hud by p iec ing
th© stispanaion f o r obeot two houre i n the i refr i oei^-^^or* The c e l l e
uare then ag.^in resuapended onti oentr i fuged at 2400 tpm f o r 10
minutea i2n6 the cupomatcnt uca dieoc^Fded* The rsmcining c e l l e
s e t t l e d ot the bottoss tiere suspanded i n the f i x a t i v e * Thie pcceas
of oan t r i foga t ion end deocntct lcn yea pep^oted t h r i ce u i t h un i n t e r *
vo l o f 4B o inu tes . At tho end of thi© piroo0oo» o oiniisuts q o o i t i t y
(2 ml) of 4B^ eoet ic ocid uo® oddod t o th© pe l l e t end the auapension
uae hcjuogenieod y i t h the help of paotuor p i p e t t e . l\ email qu^mtity
of the c e l l suepenaicn yaa taken out i n the p ipe t te end tyo or thrae
dropa of the cuspenaion yere dropped fro© a height of ebout ha l f a
nater rvar oleaned, pre^ohi l led^ t.'et s l idea t i l t e d at un angle to
cfot<iin uall apre «d wetrfphcise f igurea* The ce l l a uere d r i ^ ^rmperlf
by y a v i m the al idaa i n the « i r i^nd a j ternut ivtBly pas ing over «
burner* The dr ied al idaa uere kept i n a coupl ing j a r containing
phoaphate buffered (pH 7} Gimiaa s ta in so lu t ion (BOH) fo r ^bout 5
ninutaa* t>ia at<^inad mXitSmm uara uaahed i n double (sil&tili.9(t uatar*
d r ied 4nd mounted i n DPX*
Baaidaa the m i t o t i c ohronoscmaa frciB the bone marroy ce l la^
ntaotle ohromeaomaa frnm the taataa usra - Iso made frnm the ac«a
1 4
colctilcinlated m^lm bird® by th© wethcif describad ahrvn. Prior t©
hypotcmie tr©*tffl©nt th© tesl^es u&tw obt^^ined after opsnin^ th©
i3@rltoftiu@» Th@ tuftloa of the t e s t i s uaci removed und pl<«e«d in &
patr i dist* ccnfe«ifiific| pham^Hatm feuffet saiin© CfiS) at a pH of ?»Q»
Tti© tubule® yor© t©w-igad out ^od sHaksn uith th© tt©lp cf i* fin®
foroejse to obt-lfi tfie o®ll®« Ifm cel l eyspefinicn wa® e®Rtslftjg©d
end th© cypoffiotsRt UL>® dieoowdsd. to tho polAot tiom uddod hypotrnis
Golwtiofi and recsft of tho proosdur® uos the OGO© a® tti-jt foi? tti©
nitotio Ghro®ooc®c8»
Prepared oiido® thw® obtQAnod uero acceu^d oitdot o C,Z#
EfQc iX fstoroeoc^© ot o Qagnifiootien ef 400 H» ^nd yore photcorcphet!
ut a aagnifioatiofi of S40 K© Th© totoi icfigthSf iong assi icngtfiSi
ohort am Xongthe end tli© cer»trooorici indac©© io the 10 luff^est
p^iir® of the ohrc®3se®es inciwding the oeK c^rossoef^o^ were s^r-cured
in iO uell delin®^t©d ©otaphaa© ©oil© firora both th© ©s^ee of th©
thro© opooies uocfer ot«dy« The reictl»@ length (tM) of iRc'ini^ual
QhroEiC8Ci3o ya® sKprescsd e® percent of the tc to l length cf th»
hjplcid met exeior^iog th® ohrcJispscii©® sRiaJtier than the iOth pair*
FlUw-Xly th© w@af» r«l-2tiv© length yae calcifl.-.ted and represented uith
stiifidard errors -r^ v^riatiun rang©# Lung arw . nd short ««wi langthe
of a ohrofsioeaffle yere eaieui^ted on th® n^:mm hoeie* The eentromeric
index y«iB ojicul-^ted ue percent to t^i length of the chroiaoeoiRe and
represented in mm^tx Ofntroraeric index uith waristion range and
st<*ndard er ror .
The ohromo^owee were clascif iee on the basi© of the i r
centromerie position according to the ci^iesification made by L«ivan
JL 0
fredgc;£ ^nd Sandberg (X@64}* In t h e i r cX-stSBific^tion Utv<ift a t ^il«
(1^64) had dQ8ign.^t«d the P! chro®o«ow« h.-vlng i t s centrtmerQ in th«
ridc!Xey thu m cbttmosmm having i t s centrooere in th« midt X« nr
i!t«di.»n refiion, tha sai c^rc^oaciao h -ning i t s centrr^wura in the mib-
metiit^n r©qion, th© t chrcmcscme having i t s oentromsro in tha tf?rini«»
n-1 rogicn .nd the T chrcsiosciss hewing i t s centrcmsr® -it the tsiMBin.;!
The photcciioroQS»ophs of cuolyRed nhrr^rsois© 8p f?v:c* vwre
tcken ot (i fBcgnlfioaticn of 640 H u i t h the help of o G - 2 Crqatfil
photcsaiorogiruphy ottoohcacnt und cnlopged t c u standard ©izSe ciit nut
end QFffwfjgcd i n t o tcorsfoti^pe© u i th the eld of oriQin^.! o l idcs , rid
i n t c o t photoQroph©.
Th® ffoogent® used i n th© prepuroticn of th© oytoioQioci^
e l i d e s ysff© pfferj*>red o@ folloyeS
X. 0>05 nqgo^nt _Seighieif^y Aolutign : 50 mgs vf s o l s h i d n s
pt .x'ur ufe.s disenltf©d i n lUO lais of do^jbl© d i s t i l l e d u.-.tct <^'rtti kept
i n a tuftiq&S'^.tov*
Tho hypotonic solut ion yas prsp^ieed by disssolving 900 mgs cf sodium
c i t r a t s cvystuls (HaC^H^Qy) i n 100 ml of double d i s t i l i a d yatsr*
3 . r^Hi^UVf JK I ;? ^fff^*y¥,^??W),,l ^ » f ix«t iv« uas prsparsd
by mixing 25 n l s of g lac ia l aootie .ioid (Analu, R*) v i th 75 ml nf
iibaoluts ©thyl alcohol (BOH).
4« 4S narcsnt Aestic ^<eid> 4S petcmnt a e s t i c .jeid una matim bf
mixing 45 M1 of Gl^ieial Hcstio iCid with 55 ml* of doubls d i s t i l l s d
-f, fV
5. Phofinh«=<ta Suffer Salinm PH»7 (PBS) BolutAont This so lu t inn
uas prepi^eci by d isso lv ing 0*109 ^ n of sodium hydrogen isonophosph t e
(Ra Hg Po^)p 3»2 gra© of eodiua hydtogfifi diphosphate (Jtog M Po^) and
i«7 g{58« of eodiuet ohJtnride (!Sacl) i n 200 Q1^ of douhla d i s t i l l e d
uater*
6 . Buffered tyH 7>0 Siamaa a to in an lu t ion 8 Buffered pH 7.0
Giensa Etti in co lu t io f i U*^B ©ado wi th tha help of eodiuo hydrogan ncno«
phoaphata (Wa Hg Po^) and ecdiu® hydrogen diphosph^ita {Ua^ H Po*)#
F i r a t the buf fer ooa o^de by dis'-olwing 6.0S goa. of oodiora hydrogen
oonophosphct© ontS 13*41 gtiia* of oodiuo hydrogen diphoaphnte i n 2S0 m l ,
o f double diBtillod wotar flreporotely end pluced i n the r a f r i g a r - t o r
u f t o r ohcQking the pM u i t h the help of pH pupor* To prcporo the
buf farad pH 7*0 Oiemaao a to in ao lu t i on , 1«0 al o f aodiura hydrogan
QonaphoaphQta co lu t ian end 2*2 o l of oodium hydrogan diphoaphota
so lu t ion ulong u i t h 2 cil of f i l t e r e d Giesao o to in ao lu t ion (BOH)
UQS blend together i n 44 isl of double d i e t i l l e d uutar*
7» Claunino and Ch i l l i no o f the Sl idest The e l ides uere f i r s t
plueed i n a beakar ccnt^4ning chromiG aoid (ooncantr^ted sulphur ic
ooid and fay c rys ta l s of potassium dioromate) f o r a t laaat 24 hours*
The e l ides yore then uished u i t h detergent soap c nd pli^ced i n
coupl ing Jars* These ja rs uere pl<}Oed under running tap uater f o r
about e ix houre* Af ter uaahing, the j^^re uere oonared and placed
i n r e f r i g e r a t o r f o r ubtiut h<ilf an hnur*
A 0 D n 1 . S U, , i
I?
The chroraosc3fli08 of throe ctpsoXes ©f tho nrdar Coluwbifonses,
n.kwn.ty Coluinha livA.q (?emostlco« CoXombAj l i v l a ifit,6rmBd4ii .md
*itreDtopelAa arnaoalenaia ccitnbuvfloais uoro s tud ied . Th® chror.osofs
numbar® in tho three spwoies wore counted in tycntyPive aoXl eprr ccj
nataphon© p i s t e s in both th© OORCS*
geaogjotion of the ohgcfnoGprngs of the membegEi Qf tha ordar
Tho t o t u i nuDbop of chrosoooeo® in tuontyfivo ue l l 0pro::d
octc^hosc plutea of th© dcnootic pigoon voriod froQ 69 to 02 u i th o
ncrfeed riBtJk (46'/) ot ^ , The ©eon for th io wupiuticn uu© cotejl^jted
cm 7f1«06 * 0*497« Oo th© diplo id cctint of th© chtcnoscnos in th©
dcrasstio pigeon ujy b© giwen us 70 ^ ins tead of BO • os ppoporsod by
ct^'Cf t'OrkeralTak.gi i Rakino, i 9 6 6 | Olocts, i969f Itoh c t a i . , 1969f
King, 197©! Plccinni ^nd S to i i a , 19708 Bulatcwo cntS Pcnou» 1973?
^nsc-ri M6 K.jul, 1977(«), 1977 ( b ) ) .
Thsr© are i^bout 10 pa i r s nf la rge chrcmoBoawe, the macro-
chrcmoscraee inoJudlng the cex chrnroosorass trhlch c jn bo eas i ly
dlet inquished frcsi the ewaller chromoaofnest the ajicro-chromrjsoiBes.
The micrrKchromosomee umtm t e l o c e n t r i c (T) ind v<jrled frcw email
rod shaped to iRlnute granulee in ehape* The rar^oro-chromonnfflee are
ol.jS'^ifiBd in to five nrnupe jocerdlng t c t h e i r lengthe <md cen t re -
mario posi t ione (Karyotypee 7, 8, 9 ) ,
f Q
Gyt>yff I*. Group f i r u t compriRas tuo rj*.irs of l a rga chrt3fBf?som«s
hawing t h e i r csntroraeres in tha n©di«n region (m). The f i r s t pa i r
i® the l a rges t having the r e l a t i v e length u j r i a t i nn frnm 22.34 it
2S.45 qiuing the ne..n r e l a t i v e Icnqth a© 23*52 * n,53 nd i t s
csntrcmeric indost vuried frsm 30 to 40, The m -n cantromcirio in6t2X
for ohton'^scfm on® y^s calcula ted to b© 36«66 • 1.6B (Table ? ) •
Sacrnd chrom^cpfso pui r of t h i s group uhich i© cntriior in lenqth In
a r-it in nf 0.95 t 4« l i to the f i r s t p a i r , hi^d o r e l a t i v e length
waric^ticn frcro 1S»03 to 17.G7 end the oecn r e lu t ive Icnoth (t^) as
16*43 • 0»44# I t has ^-ioo i t o centroQoro in the nedion region
howino the ru-nga of wcriat ion bett'Scn 30»6Q ^nd 47»3G ond the nr .n
ocntroEsorio iodea ce 40.IS • i«8S (Tobio 2)*
GgcfijaL.lJ.» Thie giroop ccEJoriooo of o oinglo pui^ of chscnccct'oc
tthich or© hotyeen th© f i r s t tyo poiro of the chrcnoocma© rf the
group one in l ang th . This chrcnoonno hoo i t s centrcnore Icool.^; ^d
m the tetRsin, 1 region ( • ) • I t i o ^boot 3.2/B.CJS of the nhrtTmcKoras
one in l eng th . I t s rolc;tive length verir . t ion uas found t c be 11.27
t c 14»Q8 r i t h a we«n of 12«ni ^ 0»3S, Thie chrcrarsrtn© i s u t
type nhr'^«"srwB,
Gi oup III< This group h.» t t c paire of small sized chrf i nnnmr^a
uhich are f i f t h »nd s ix th in nosi t ion «® nhcun in the karyotype 7
i r r e spec t ive of t h e i r lengths* Hut of these two p a i r s , the f i r s t
one has i t » mean centrnmeric index 40.43 • 1.S9 which \t.^Tin^ in mm-
di f fe ren t nl^^tee from 36.«4 to 47 .73 . This nean centrrmeric index
designate* the chromnsoBie am ^i m type chrosirsrwe. The rei^-tiwe
length voi-iad frow ^.IS to 11.27, tf e wean rel^ititfe length v.4lue
hminq 9»2S • O S*?* The st.oond p^it of thi® qroup ('ith chrrajojsrme)
hus .>ASO i t * cantrnmers in the Biedian raglon (a typ« ohrrwosnBie©).
Tho mean r e l a t i v e length of t h i s ohvamt^sprm U^B found to h« 7.92 •
CJ.S'I Mitfi a i»«ir4^.tSnn SviJicj© fees© S.77 t c 9«02» Thoy f 41 in d
r - t i r cf about 5>»32 8 B.8S . nd 1.98 i S»9S respeet lvs ly to th©
f i r s t :i.4ir t:f chromocosos*
Grpup IMo, Thio r!J?cii? inoiudec fcup pi^p© cf nn^lJl s ised
ohrcnt>uot3co. They h w© thoi^ o®r»tto©0ro iceol lgcd at the tefnSnol
cno'o They oro cbcolutoly T type chroEcccisss* '^t cetophoso th® tuo
(jhpoffiutiCs of th© chronoccQo® inotc-d of ceporuting 'sa^ain attaohod
vt tho C8ntroaoPG end prooent o U ehcped pioture* Their r.aon
rcl^'-titf® length doorooeoD frcn 6»2D * 0i»23 (7th chccnrtoooo) to
3*70 «> O.IB (10th ohffoooooso) graduoliy (Tcbio 2)#
Groifc V. This orcup cona t i tu tcs tho CCK chrf-nocoGses. In b i r d s ,
-o u© Glrov.dy knou the ncloa ore hcsogcroctio c i t h 22 chrrmosniBCs
t;hil8 tho fcci^ies, hawing ZW c c n s t i t u t i c n , era hctrogafis t ic . Accrrd-
ing to length, th© ZZ ohrcmoecffiss i n sa l e s cto th® fcurth l.»»c|S8t
chrnmnftcfflos. Sinno t h e i r centtcisere i s in tho middle they could be
daaitin tted a® H tyno chroBinscfflS. Th© length of these chtonrscmes
wariac* from ^1.13 tc 11»27 in ten met^^phaoti pl-^taa with a mQnn
rs la t i t fa length 10«1S • 0«34* Tho centromeric moasurements revedletd
tho m&Bn o«intrc?»erio iniiwn as 47.73 • 0#fSJ2 with a vvifiution r^ni^o
from 4S t c 50. In fem-lan the ociaplsfaont has a pwir of Zlf
chrfsraoscmo. The Z chtownsrao being cf th© & MB nizm sa in malm
of these speolee, the U chroa'scme i s generally aroaller th..n the
Z DhrowosDBw. I t in ^bout 4/S of the length of the Z chromoafrae.
20
I t s centroner ie intSmx varied frosi 42»@S to S0»0 uhich givao a mean
value of 47#1S • 1#30« Th« rel^itiva length Uiariod frcan 7»30 to
9*D2 and the taeaft for th@ ir@lfititt» length yas ca lcula ted t o b&
tl,39 • 0#409« Tho W chrosionniia i s s tou t th« length ef th» 6th
0hrciiRO8O«a but i t o^n b© ©aeily i d e n t l f i ^ i n haulng ^ gentr«l ly
iocutsd ocntrciaoi?®* I t l e <il80 o ^ type chroiio©oiB®,
Th© clirofflosofs®© of th© Indian blu@ focfe pi§eon and Indit^n
l i t t l © broui) don© hcv® besn g^otiped in th@ ®im& {i»nn©r« XhGf present
Q homology to th© ahroisosciiQs of the doses t io pigeon* Moy@ver» thsy
d i f fe r in th© chfoaoQCfae nuiBbor, moan rslat iv® length end m®Qn
oentrcmerio inddM* The mmcn tow the diploid Gsownt in th© Indii^n
bXu® toali pioson oomoQ to b® 7S#26 ^ 8«©39 y i t h o nt-pked poek(46^)
a t ffl ^nd c va r i a t ion i-ango fpc® 60 to 83 CTsbl© l )« Thye tho
d ip lo id ocynt in e p j ^ g ^ MMM. iM^m¥^ "i®o cmjld b& given m
70 * ins tead of 93 «; e© eugq©st©d by r.ooi© cthep uctkors Cstsnlus
ofc c l e , 1563? H ifaP!er» 1S?S6| I teh «t ul», 19@9)y fjhci conrid^sr®^ cnly
th® highest pmM of th© diploid a?unt« the fs®.45 for th« diploid
count lit Indian l l t t l ® brown drve Mds found to be 77»42 * O«420
u i th o marked p®^ (44^) a t 7f?. So th« diploid chroH!oi?ntw« nomber
in th® Indian l i t t l o broyn riov« can im given as 77 • • The rang*
of va r i a t ion of the d ip lo id chropiosoiaa counts in 2S msteti^aae pl<^t«8
yae fnund to be 68 to 82 (T«ible l}» Xn both the speoise , Indian
blue tock l iqeon and Indian l i t t l e broun dov@« the diploid oooht
contains 10 puire of siacrochroiiiosofiee including the SVK ohroffiosoeiee
and the remaining are wiero-'cHrtsinnsioiree*
Ths i!!s»n ral<:^tive lengths <ind «e»n centronoric indeeee
vf tho v«irlouB chropcsonse of the Indium blue rock pigeon «im! Indian
21
l i t t l f i hroundotfs a rs given in TatitXe® I I I and lt/«
oentromeric indeC8S» long rra® Icngt? », shcr t utmn lengtlia has been
ffiad8» Thi« oewpirision has baon mjisi'sarizad in Tahla» U, VI, tfll
^nd Jlllm Comp.*rotiw® idiogrw-ms chowing the oentrGmerio p a i i t l c n s
v.r»d Ipnqthg r f the too eras of th® fehro© cpeoies hav© olso bmn
©wcJ® to o ^ e tho obsceViJtions racr© oev^ningful (FiQur© 14)•
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A compar^tlva study cf th© karyotype* of throe soeoisa,
Cc luwba If via tjotUBBtioa (lndi„fi tirmBatic plqeon), Col^mba ^iviia
intfirwgtfla (Indian blue rook pigccn) i.nd Str^ptooolfu
BonaQaloTisim Q^B^buvonsAa (Indlon l i t t l e brctrn or eonegi'l dou©)
of th© r rde r Coluo>blforra©8, rcivevjled u not^blQ UQrlr.tino in th®
fHjtabnr of cbrcsosfsras© of , ; l i the thr»9 ©p©oics. for Cclunba
MyAa S!£B£SUsa <-^ ggluwhQ | | ju4£ ial££2S^iif «i»i0 tfcriaticn was
Prcia 69 to 02 ond 06 t c 03 t©opectS,wely» So both Cwcec^ hououer,
th© QUQffcn© chrciaoooBo number tju© Pound to b© ?0 4-» In eft
,nfe,y,R,P.tePfii A . RQR®J3.9i£PI>.l,§ OGmbuye^Ma tho Ruober wug-iod botueen
66 v-nd B2 but th© {itfor< (5® nuobcr of chrtJDOGCDc® ua® found to b©
77 • • Thaso findings ur© in acccrdanoo u l th the studio© cf th©
chrofBoscraes of pigeon ;nd dowss by rocwns of the Ql^.&rAo t e s t e s -
sec t ion nathod undertcksn by f^ntashina ^nd Hakino (1946) . nd
riakinc pt 1*C19S6)* Hou««ar, these findings - re s'-meuhit
d l f feront buoauso they noted 18 a jora-ohroi!Jr<r»«}0p in bnth the
specimens uher©a» our »tuc1i(i» cia-srly revealed th« presenao r»f
20 Bt-oro^chrowrsnmwj including th« sex-chrownsono, Th« rs^aon
for t h i s r 'ifforence miqht b» tha t they gtoupad th« r8m*iining tuo
p«iir« of c l s a r ly victibls Ri^cro-chroroarrfn«« wlonqyith th« mlcro-
ohroncnniRoa* I t c-'n b« eaen from th© figuroa 7, " ^nd 9 t ha t
thoro Arm 10 nv^irs of eS' cirtj ohromnsfitaBO in -11 tha thr«6 spscies*
In gsner '1 pigoona h-vo tuo lanra chromoBtinsa th.^n tha dova hut
th ia differanca i a only <i«ioni tha »icro*ohro»o®oB8B not in tha
0iaaTa->ohro»o@owi&. Gulton .ind Bredb«iry (196P) h wa rBcognizod
0 n O 6
1«? macro-chro«o80»«« In the pigeon. The expianjiticn of th ie
tfarl j t l en .iinonq the macro-ohroroosrmee muf be given ae adv ^nce-
ment c$nd ecfiiTistication cf the Jaethnda in cytolcqy .nd cytrqone*
t i c s .*8 valX as of the t c c l s of niorosanpy .nd photDqr.:phy*
yh©n the s - r ly ©tudi©© of b i rds esoeoisjlly in fjailjugfj domesticue
{dnrjectio foyl) yor© oturted^ i t uo® toyez (1006) uho observed
r ixr chromoooKce in th© priD.:^y occytf?. Then ncnnonbroritU^OB)
cbserwrd f) to 16 chrcjcjocrraos in porthenogenrtic scmctio cell j^.
Guycr (1909) tir:® th© f i r s t tiorkor concorned u i th th© chrcfncscncs
in Bportaatogonesie of tho fOijl» ft© doooribod IB to 19 chrcnc"
ocBJQ® in the GpoBfJotocjoniul coll© ood D chgcnoocno© in tho
pvifSi^ty Dporoctocyto olninino tho dioeovory of un unpaired SGK«
chrnnciSGBe in th© male of Hallup, Mmn^titkmB. (dcneotio fctjl)*
All thee© ofocottfiJticn© cupport tho cbowo ©Kplonution. * '
I toh ot cJL« (196D) DQtla o oomp-ratiuo k- ryo t 'po otu(fy in fcurt -n
species of b i r d s . In these 3t:jdic8 they dosoribed th© tn t . 1
number of chrofflcjromee -.ts 80 • -nd obsorwed o wariation r^nrje
from 69 to ^2 und 60 to PS in the domestic ^^li^pcn „nd t.;ild piqarn
respeotively* The ohrfimnsriaee of Colyw l?. ^ pHjlymfyg, l» have been
studied by HdWaor (l^SS) fro» the embryonic t isyoee obt^^ned
frr«i pgne incub^^ted for B to 15 duye* He counted 78 • as
d iploid ©hrcmofiomr mimber. L'dagjua|[1054) et^udied tho chrr'nosnmes
of Colurebu fanthinu and i*tfe the 2n numbrr us 00 • ,
Similarly the chjromns' mee of Streotopelid epncoalengie
c>«wbtjyeni i8 gave a v>*ridticn r^nqe from 6B to P2 with a tn-rked
peak at 7f^ (about 425^ of the t o t ^ i pl^^tee e t td ied) ^nri a l ees
3S
oonspiquous poak a t BQ (abcut IA% of the pl<^t«8 etudJLed}* The
mean for diploid nmnt in t h i s ^peoisa comas to 77*42 *, These
s tud ies ..re in .»greoPi8nt with th© sttidiee of Srivastavct ^nti
Ml era (1971) uho studied the chroiaosnroes cf StraDtopGl;f>ti dec«iftctc«
They deecrihgd the 2n uc 7^, the highest p0 4« ^nd the n©xt to
thiB pevife .^oaording to theo yte 76 whsre^-s ye h..ve recnirdBd TO
chroranccmes «-s th® nort highest po«4r, Odog^yu(10S3) studied the
chrcmosrme® of thre© opeoiee belonging to i:t)iuiibic*ac, Urdtjid^^e
^n6 Alcidas* H© described th© chpcoocoines of .flohiemi. pu
MsfesJLSlli "S 7f^,*. fitJgi2^,E£M^i ,afiiFW,°. .t?. ^s 76 <^ end nU.^p,tp?gaa
cffientcilis as 76 ** Itoh c t wi»|[i9S9) Qtuc'icd the fearyctfpcs
of dioiBcnd dcvo CS.aciMli/, c ,n@at@) > r ing dom (r.tj?e!at,oonAin
riepga) end uh l te dcwe (StreptopGli, ' .r,i.o„ef.G tf«ral|ba) and acntici>»
ed t h e i r diploid ohrcmoccrae coaplemsnt© *-o 72, 74 and 74
r8BpcotiWQly#
I t i s to fco nnted th>it the wariat icn in number of
chromcscwea u i th in the SUBJO cpecie© <m t-ell .»s in diffeir'^nt
p l a t e s of a tjlidee u«is only .iwonq the micrc-ohrnnjnsfjraes* The
number cf sioerc-chro-'onrctee remains the s-ifno» This u s r i ^ t i cn
among inioro«chrowosn«ee Bi«*y be tiuB to ieiprnper sprs'-sding und
s t a in ing of the miero^chrowosomest due to hypotonic treatment
^nd dropping the aall suBoension on the s l i d e s from a consider-*
able d ia t jnce sn t ha t the tuo chromatids of a miarc-chromfsomos
which are very ewall -nd t e l o c e n t r i c wiqht get detached frow
each other ^r^ ^re crunted as separate micrc-chroronnnroBs qiving
more ccunt thdin the uc:tu<d nuiaber* The cnse nt y be revoreod in
Improper s p r t . d i n j uhore one microchroraoanm® ovorltipe the o ther .
0 4
rto i» fsvident a typio.4 avi^n kuryotyp« onnsist nf
tine typaa of chromrgonie©* The Pipst typo inciiides & o^ttAn
Lumber of large chrrnrormeo aulXmfi ttt© n«^erf5-chroniononie8 v*nd a
l - r o e number of esmuii©!? chrowcscras® r-nginf) frofsi ais^ll red
fjhaped to oiruito grafiules» ocl led tho rsjiorG^chrrnoEtJiBes. These
twc typas nf chrcncccmos uer© otudicd fcr the f i r s t time by
Hunc» (1524). H© ho® done ©Ktensiv© wsrli on th® chrooo'jrmes
cf the donootic Pottl* Ftco hio ctisdie© (1923, 1^24 ->nd 1926)
i t bcouD© oulcJcnt tha t th© chroraof30G!e ooicpIeK of foul conrint©
of G oer ta in nuafoor of occro-ohromoccfsco h^.wing o dof in i to ehupe
end a locQO nuabcif cf wory a inuto grcnuio lib© olcrochrcmoof^ssG*
Tho Qaerc»chrc£!>coc43C© of the thrco opooic© otudied chou ronark-
ablo r l o l l a r i t y ctaong theooclwo© u l th olig*'^ d i f ferencos . They
h^wo hp.on diMidB6 i n to d i f fe ren t grcupa cccording to t ho i r
lenRthe itid cen t rcser io indeoco a® desoribod ©ar l ior .
Cn comparing isong th© f i r s t p-*ir of ohromocOB© hciong*
iOQ to gr^i:p nn© In the thro© spsc i s s ( f i g . X3), i t i s Inngaet
i " PC,;y"»fe,ri i i j d i i dofttcstiy (23.62 • 0 .S3) , sra-llor in C^.l^fapA,
^.1,,^.^.., nt 8 iqgcj_ , ,. (23.49 • G.!'32) .nd Btn^XXmt i n Ht£,tR|,t?,pjM.<?.
eenaq^ilen^ia pi^wbavanais (2t .03 * 1»5S)« Th« short . rm lenqth
(T b i s 7) of CclufBb ^ litijw intcrroedifci i o longast (9*63 • 0.25)
ajsiong the three Gf38Ci0« uh©rti.»» Colmaba Aivia domeetica h. 8 «i
shc r t rw length interfliediate between the l«ngth« of Co^uwba
ILiiA L2%»m%4k§. -^ HtraotODBli^i MSPm.L^mf^M &i^^Pmk§. ^^^^^
h«« shnr taa t langth (7 .65) . Th« Itsnq drffl lengths (Table S) cf
CoiuBba^ lltfi . antamteriia nd S^lffptopf;^ , , JiaQlfli*ij?2ll;&
carobdyonBls ara alwoat aqu^l (13.72 • 0.71 -'Od 13.67 • 0.97
S5
rospecUye iy ) . ^ i Ift CoAtiBiba liv^a dCTSsatlo® i t i o iongei' than
th« tuo (X4«94 • 04i618)» Coluwba li \^i4 ^ntermedjl^ has g r t a t e t
omnttommvie Index (41.?4 • l«6 l ) than COj ofiibQ l.iv.ij? doaiitstioa
C36*@6 • 1*66)• Th® Stf^gptopella eptrciea h-s th© ioyeot
oentroaer ic index C3S,04 • 2 . ^ , Tafele R)« Seccnd pait of thl©
©roup in th® three epocios hon® t h e i r {soon r e l a t i v e longthi i n o
cJ©cre«ieiR§ ord^r oP 16*43 • 0«44« 16«S8 • 0.478 end 1S.3P ^
C3»32? (Table S and Fi§# 13) • Th© two respect!v© uros of th@
tyo ®p60i®s Cq^illiliJ, MM^ JBtorsiodiQ an4 g.tx.^p^ii,^o,l,io ooneoalen*
®,1.@ Gorotocveneia howe o laes t th® ©a©© lae^n relotiv© length©
{P«50 • 0#30 ond 9«40 • 0#29) und (5«02 • 0*32 <jnd S . ^ • 0«24)
but i n eo®® of Cel^iabQ l iv iQ doroeetioo which i® lonQor^ tho I t no
und ohort otB@ (Tobl©© 6 anti 7) or© s l i g h t l y l©rQ©r then th©
two ooeole© (©•87 ^ 0«34 und 6»63 ^ 0.34)* Th® c©ntroraoeiG
indeo©© of th© ohrcffliocoQ© I I ya© found to b© i n a d©oreasin9
order from Coltiroba Aitiii.. domm&tiQu (40*15 • 1»SS) to CpJ umba
li tf io interwedia (3?*51 • 1*4) to Streotooel ia sgoeoalenaia
oamfaAifcnsia (86«8 • 1*23) • Th© chro8!o?.o!»© thrs© uhich bcong©
to the qroup IX i© longer i n Colyaba liMhi doaaatica (12*S1 •
0*3S) thtin In th© other two ©paoie© which haw© dtlmoat mqu-al
l8n§th» (li,0f^ <!fiti 11 *H7)»
%a8uroa©nt8 r0\f©^«d tha t the ohrnnosoeio tf of th©
grc^p I I I i n the tyo ColMWbat 8p©oi8s ar© almost equal i n length
und the ohrom^sone of fitreotooel^id specie© i© sm 41©r* Out on
ooi^drinigi th© ©me of the two Colmabd srjeciea, t h e lonci ana of
Coluwfaa Ut^iJ^ ^mmMB^ ^> em^ller than th^ t of Cofuwb^ livAa
L ? 0
Intatwadla (5»37 • 0*26 ntd 5.89 • ^'^^ f tsnttct iucly) whrre^is
th» short rffl of Coluwtia Zivi^ dowgst^ca i s l a rqa r than tha t nf
,^Ati?#,^ l l ^ A i If' f ' irf *^ O^*^^ • 9*21 end 3.19 • f i a ? r©ao8Ctitf0ly),
Tha ssacond p .4r of th i» nrnup (Ul chrcransmr©) i s einjllur to tha
f i r s t p-4r In the thrsa ssecios u i th the only diff0t©nc8 that the
chrcsQsomeo of SoluCTtjiQ l i i i i a intogna,dia 4© th© longest eicong the
thto© ODOOioo with t h e i t r©op^ot4«® ottas lonfer than th® respact**
i v s utms of th© ohfcmoscfBc® of the tuo refsainihg opecles, Chrc?-
ocsosioo yiJf tflllf IK end X oocspriee® th© foarth rjrnup. A f©murk-
ubie point t o be noted u i th t h i s group of ohrosioscraos i e t h a t fche
iongths of the feus pair© of tho ohyoooooctc^ in the throo cfsjciss
uro in inereiiolog order frora ColMfnbq 11 vio {rtonostio}^ to Colunba
l i y i o inlfcrmpdia ^n6 StrcntooaliQ opooiee uhoro^o a l l other
chrcfliocofflos of the othor groups (I to US) h^»® t h e i r nscn r s l o t i v e
lengths in deorOv:oin0 ordar in th@ Bomo oor iss*
Ray-Chdudhuri ot al«(1969) h^v© described the ooiBp«rs-
tiwe Keryotypes of 11 birds* They cotaparod th© chrcfsrscwee nf
o r l e n t u l j e ^nd f'tind much difference in t h e i r l engths , nmong
th© f i r s t p^ i r of chrofsoQOQes of three spnoies^^ the chroiBoenme
of Coluwbe o.aluf3buf i e nuoh sa.al lsr (about 3/k) of the
o ' Cdlmsba livi,.^ d iieetiOHei anti S t reptooal ia qriBntai;i.^B. which
i4r© alre-^at equal . The chro«oacwe m»»bsr two of Coluwba oalowbue
i e longest Jwong the three speoiee whereas Colt^aba l i v i s dowesMca
. nd ,S^rfot^ooelii^ aneciee have almost aqu^ ^r«i niioilar ohromosomae.
The chroBioeeBf) H I of those thre© sDeoiea studied by R4ty«-.Chaudhuri
ot ^ 1 . (1969) thay doslQnutod th« chromosoK© ae »t typii and aa
in r i3ur«/5" thay mtmberad I t nm a lx th c*»rt3«oso«a» Thay hawa
Jirrangad the v-»riDua ohroiaoson^ in r e l a t ion t o t ^ a i r eantrch*
raaric p o s i t i c n , Tha chrowoaoaie of Straototyaliy o r i c n t a l i e i a
sia-illast and thwt of tho Colufiba 11 via dofaaatica i a larrjfi^^t.
Tho ohrotsoGOiBo of Cfti ^foa oolotajbus i© i n t e m o d i a t a batyoen the
two, Chno ot cl« (1964) dassribed th© chroraoccmol uniformity
i n tho ijivion sub»claso Corinatea* They hcwe given nine p^ira
of iaGorc5~chrctaoscc»88 including th© ocx-chroisosctaes, in uhlch
thoro are tuo pci re of Iwrge eub-raotocantric ahromosoroo, a s lnqlo
p^it of to looen t r i c chroooocaoe, two pa i r s of email eub-aetooentr ic
c^ircmocsoisssy threo poire of t c loocn t r io Gmoller ehrofsoccmaa
inctoad of four puira oe dosoribad by other uorkere oni^ th@
present cuthor, end Ica t ly c OIRQIO pu i r of ooK»ohro: OGOiai3o«
Th© f i f t h p j i r of chroaosomoa in Columbe l i v i u riofaGstica,
Coluntbu ptJurebua. ucre deecribad in the ©vtaa yay by Ray^Chaudhuri
a t ..^(1969) but they ysra un^bl© to find such chrcmo^ewaa i n
GtreptooBlifcs o r i a n t - J i a . thoy huv© fjiacad th ia chrcferKowa in
8treptupeli«=> spaoiaa .m Mil ohrorootjctBa (^ifjuro 1 5 ) . Tha ©eornd
n i r of t h l a qroup {Ml chrnmnrome) ia sicill >r to the other qrf»jpa
i n a l l the three speoiaa t i t h the only dlffarpnca tha t tha
chrof9030«e of Cpluwba ^ iv i a dowtrntici^ i a tha longaat aaong tha
thraa r^pooiaa u i th i t a raapaotive araia longer than th© other
t%iv chrrimnsonea of the rem-ining apaoiaa* In tha 1 mt group
they h< ve dasoribad three paira of ohrnmcsoaaa (tfXIXy 2X iind X)
Df the spaoiaa and noted in qfiAMWfr Ikiki llffifffl A,ff, , « « " ^ i
chrnwoooBso about 2/3 of tha s ize of the chrownaotiea of Cpluwba
paluBibya and Struotopali'* o r i an t a l i a (tflll chrowosowa) • Thia
3S
chroii08Ofli« i s t s l o c s n t r i o whils tha &^mo tshtomtsnvam pair i n
Coluwbaf aaJ uEtbuq has i t s oentmwars lacAized in the terminal
region -jnti i© about 3/2 of tho si t© of the ohrnmoscroo of Columba
l i v ^ a dowgstica. The cjhrtMnosoae of Strootpoeli-i Drient i l i t t i»
mqutii in s iza ti i th tha t of Cnlowba oalumfaua with tha rniy
tJlffarenco i n hituioQ the cantroaer© ^t tho termin:;! enc!# Chrr'ucj"
co©o IX (fit^ure 14) in Col^mh^ liul,^ domostigo jnd ColutBb^ pal ffmhus,
-re e i a i i - r to the Mill chrrraocaiso but th® chrcnor,c!!!® of
StraotQosjt^^u p r jgn t l i a h,;® i t s oentrcaer® near the tcri»in;;i
region l , e . ohroiaoocso® of Cqlytubfa p.ilumb«j<; and Str |9^t tnolia
o r i e n t o l i a wro Qioilur in eiHo end ehupo. In tho K puir r.il
the throa cpeoios h^VB T chrocioocaos in uhich Cclumb/i t {4lufal3U|3
haw© cocseuhat l a rge r ohrcaocorae© tbun tho othoff tt^o spcoics
uhich how© .JlBic::t ©quui length .
Rrcup f i f th GCjnprissB the sc»»chr-ocscme. In scrpho-
Metric t ab les tZ chrocsascjniea occupy fourth pcai t inn wCcortJinq t o
t h0 i r I t nq th , Tha 2 chmwoarnea in II tha t ' raa apacias ..re vlu-ys
tha fourth Iv^rgeat p d r . Hairier (1965) i l se Found in An^c]:' . • -^or.
Cclumba pfc.l wbua and ,HijP.fi picu tha sax ahrr rannfHsa - a the f' ur th
lurgaat n^it» Bloom (1969) «iso round the fourth l a rgaa t ohro-
woanme aa tha aax c^irnmnspina i n . l i l la rd ducka. K<»ul rtnd Anrart
(197S) found in Aecioftar ^a^^uf the f i r a t lurgaat pa i r aa tha
sex-chroiiioaoma, Anaarl ' nd Kaul(l977) found in Sturnua paoo-'
daruf^ tha th id aub- te locen t r ic ehroBosowaa aa tha Z chrciiBoaowea
- nd i n Sturni^ fflalabarica^ tha Z ehrranoKrina u -a idan t i f l ad a the
fourth l«trgaBt ne tacant r ie chromosome* Kdul ^nd Anaari (197P)
have given the 11 l<jr§aat p*iir of chrtmoaomBm -^a th« 2 chrcwoaomaa
39
i " f^tt^l^l*!^, f?afii aefB»* |a AnjUgj* and th» f i r « t l a rges t p^tir of
th« coaplamanfc a» tho 2 chronDsois© in Pinppiytn banq»^« lttr yi «ind
Tha V chrotRosoiBo tshiah I s . I s o mataoentrio has a nean
tmXutivB length mqoal tne Ul chrcEJoocoe* I t 4s a l i g h t l y lutgc?!?
The U chto»os£3CJ9 in avian fcaJ lss was observed f r r th©
f i r s t tiro© by freder io <19@1) io tho denest ic fowl. Sohemld (1962)
detected e scroll lut© repl ioot ing chrosjoecaae by Docn© of outc-
rodicQraphy ^ nd miQgeated i t oe o y chrctMieomo dssoribed by
feder io <iS61)« CuBn (1960) obtoir»c3d u e i i defined tsstcphas©
f iguros from various t i s s u e s of chick essbtyoa <:nd reported u
ZU ohroiKJ^ass cons t i tu t ion i n th© female*
Ths sraalier chroraosoncs (raiciro'chrcnospfnes) uhich vary
from sfs^li rod shaptsd to mimite granulss were d l f r i c i i t t o
measure* They l i e bstisom 0»Sy^ to 3 > ^ ^^ l ength .
In 1954 %ucf ra€jr ^UsdiBd soeim-togsnesis cf t h s
drmsst ie foyl ^nd rsported ths h.*s4o chrrracjscffls nucibcr tn bs
s ix uh i I s ths fflicrn-chronosrji^s wsre rslstj t@ri to ths s t ^tus cf
aCGsrriory or suoernuwary ohrcnor.nmes on t h s bas is nf t h e i r v<sria«»
b i l i t y in ntnjct i j rs , nu«b«r and bshjvimr* ^ l a n d s r (19S0)
dftfinsd aecsssory chrnwcsciwis *to dsnr t s njoh sx t ra chromosrinss
as a r s net hoMologn- s with thnss of t h s fjrdinary cowplsmsnt and
yh0s« nuiRbsr i s in squ i l ib r iun in a pRpulation"* I t i i t s (1^49
1954) objscted to ths t s m aooss^ory chromosoms 'JS having basn
40
looaaly usaci by soitse ^^ojthora in raf@rsnoe to tha sax-chrnnosome
or polysemy and «ut}'^88t8d tha tern auparnuiBary for tha e. sia
da f in i t ion which dc«8 not mmem preferable* In the sttidy of
mi tc t io chroiBorcfliae yhioh a re olyaya more fovourJrle fnr morpho
logica l study hue uidened the genetic and oytogenetical r i f t
betueen the rolcro or superwiaory end n^orc-ohroiBoscmce b^ reffloi^
ing the forraor frosB the outogory of chrofao6ocie®# Those elenentef
by t h o i r cpporcnt o r ig in , ©truoture, beh^wicur ^nd function do
not f i t uny occspted def in i t ion of .ny teind of chromoeniBee.
flcyooajsr (19S?) described the© i n the otudy of c i i to t i e chromosotfjes
of the dcaes t ic foul end proposed the t e rn chrofsosoiaoid®* 2n h i s
pupor ho has dieoueoed tho apparent origin^otnfcturo* {aehaviour
t-nd function of th© chromooc^oid©» In h ie opinion tho chrciaosc-
ooids yore coent r io , ol looyol io in or igin c-nd behowiourf var iably
h©terachv'cm..tic v:nd ohou oid© woriution i n miober* Thoy opparont"*
ly from chroraocentrio heterochrcraotin and ore progreesitfoly
diminished throughout th© enitotio ^nd no io t i c oyoloe* The most
p laus ib le theory for t h e i r function uhioh i e supported by cytolo*
Qio evidence seams thutthoy ccno t i tu te reeervee of nucleic ^cide
for the chrowoaomo synthsais cycle and i^ro u t i l i z e d ae such.
They ire not ohronoacinee* Oonnelly ^nd Neycomer (1963)f wfter
autoradiography with t r i t i e t e d thymidine found in Callua doateaticue
t h a t the entailer chromosoiiee eynthesized NOA e^irlier than the
l a r g e r once. They took th i e ae the evidence for the nonchromo-
srsial e t ^tus of the ets.iller (^rf^mntomes*
Schmid (1962) J ISO found tha t DNA ayntheeis in Gallua
dnwaaticua terminated e a r l i e r in the smullar chromfiaomee th.jn in
4
thm X irgmw cHrotsr9@cmo8# Only vno m*ill chfomcficmm in the fem^dcie
prB«uw.jhly thm U chitjw "Soa» w » i a t» in repl ic^t inq* In Cc,3,uinba
A^y^ftf ^«itnn . nd Brcdbury (1956) Iso fruind a dlffrrrence In DMA
synthesin between the .lander .nd the srauller chrnmnsomeB* Put i n
th i« spaciest the sraaJllet chtciBc::nroc8 u@re l a t e s t in rep l icnt in i} .
a lso the U chrcmoeoiies 4n Coliiwfetg li.tfi,a uhich i s r ah te r X, rg© tia©
l . t e in ropl la i i t ing .
S* t Q - "3 domoBtiims. Bicnchl ^nd Ilolina (IDS?) ta .els -n
ItapcrtwOt onalyoio of tho OOA uptuko duping fcur dsfif^ed periods
of 8-phv.c3» Xn tho Gor-iicst J>por4od IcbefMng of tho o^orc-
chrccoooBB© soerocd to bo of the octsB intensity c.o t to lobe l l ing nf
the mliorc>»ohroaiocoaca» 1"ho highrot tho miabot of i^bolicd aioi^c*
chroiTOBcmes^ tho oroo'oi? uc© oloo tho iobol l lng of the rav^opo-
chronc ciBes* In th4© period thciro UGP© - loo eom© roginne of
fnaorc-chronr;<jcj!i88 thu t uesr© stpcnQly Icbol lod . nusimuo ItiballAnn
%:VB reaohad slnultansouaiy in the cccrnd period by bnth n i c v s
,.nd faacrns. In the t h i r d period ocm® of tho raioros * nd p^^rts r?
itBCtcm c^pplsted t h e i r Df?A ©ynthoois* During tho l - r t period
ths r» ua« no l a b a l l i n g in the miotce oxa©pt for tha i^rssuwsd
U chroansows in the fein-4e. In the n<.crn8 cpedifio reqicng uer©
lubellttd*
This ds t i i ied ^4ndly8i® of Of>IA cynthosis in the aoinatlc
c e l l s of Gallua tfowaatioue ind ic^ tas thut there i s no ola^r
d i f fe t snce in DNA syntheeis betwcan n!«tcro8 ^nt' a i c roa . Only the
I. chrownsowe -^nti »n»e epeci f ic regions of ffiacros are r^ither
l a t e in r8plic«itinQ« but the y ohrcfTtoaoBie i s not 1 i t s r than the
2 ohromnsone or the autoBoiie«
42
In maicBis in Gallu», dowaiit^cuii Ford mti Uoollas (1964)
an^ilysed tha b iya len ts enployirtg both liqht .Md eleotrori mioto»
ecepy* Thay fcund tha t i n p^ichytenai t h e r s i e no gap in mif
batutun «aert}« anm mloroa* Thera i s a oontimicsus aar ia* fttm
th« l a rges t to the smfailest bivalent®* In the e t ruotur* an^lyned
by ©idotron nicsrosGcpy th®y found no di f fer , nee batueen the l^rge
^nd stt<all chrenosQines* The preeenoe of tuo^ more ot l e se die*
t i n c t s ize groups cf ohrciH^sonee in eos t invse t iga ted bird epecies
h-^m thue e v c ^ ^ the thmight Vhat th@»i uoiil6 ci>@o be funotionai
differenoee batyeen the icirge ond email ohromoso^ee* But the re are
ceverul orgumente coQinst t h i e opinion* The tyo s i ze grctipe of
ohroiaoocfiic® are on ©Hoeptioo ra ther then ihc ru le yhen ooapuring
the karyotype® of bird epeoies belonging to ocny difforcmt orders*
Chrcssosooe e i se forcis for tsoot of the speciee cnolyced^
B Qontinuoue s e r i e s in nitosi® v i s i b l e in several of the reocnt
uorke on bird karyotype (Krishan ot ol.g, 1965t Krishn n <jnd
Shoffnerf 19661 H.-^serf 1966{ Ronzoni v nd UeQnl-»Tallurif i986f
fJayChaudhuriy e t a l * , 1966$ Kay*»Chmjdhuri e t al*t 1969)*
Th^re are no a t ruc tu ra l differencee between l-a^t^m and
sffi^l ehromosoMee (Ford and toolXom, 1964) and no differencee in
DNA oyntheeie (Sianehi and f1olln«<, 1967)* Thue the re yould be
no reaenn t o eceept functional differencee betyeen latQa und
efitiill chromoaoiiee in <^ -jiucn isi^ncies*
In the present karyoloqioal etudy i t hae been found
tha t the three d i f ferent epeeiee of the ord^r CoXuabifnresee are
4 ^
vmtf Q1CS« t o one <ifioth(,r» Thay h^.vm s i i s l l a r ottrofflcsoeitt otsuotyrop
sh^ipti and miM9 bfith nift i t t i t d i f f« r«ne««, Th« tablac tsiffialeel t f i . ' t
^ I f a f t lm t l t * ^ mimiw and M i a t Z S UMkL ll^^«!I««#i,f^, ^i^« v«ry olc}@«
t© rtn» another than the giylywfc^^ My ^ .a do«f«tio«> Th4» oloso
asBQisi c'tjlcn msf bm <lu6 t o tha fac t that thay l i v e ^tlnost i n ths
e iias aeolot i ioal niefist yhiJls gplu.^ba l i i i i a dtp,f^gti.oa. i s ii p@t and
dosisstioated Qnisal*
Th© katyotypa enelysio of i i t d s raay bo wary psyoh usefu l
i n syfittmiatio oJlaasi f ixat ion and i n th® etudy of tha b i r d
phylc§any.
p p n. s, .i I? 8 I 0 0 8
4 4
1* Th« diploid chro«oso«8 nimbttr in ths thr«i« 9p«cia« of
CoXunbidaii vlx. Mii^^ HOElJl flP^^%^9^f SsklSM UiUA Intfffyy^^iy
<>ntl „§, ,r«B, offiMl . fff"^g^^y?^ffll Cdi8b4VBfipii varies ornsidsrably*
Tha m&an chronoscHse miiaber in the thre« cp&oies was found to h»
78»06 •» 7B*36 • v.nd 76«06 • feepectitfoly*
2* Based on typicul uvian cHroraoscaest ^^^ tyo typas
saost^^ohroaocoaea ^nd mioro^sHros^jscmiaa have bo@n fciund* Ail
tha thraa cpeoiaa atudiee' have 10 pairs of 6iaoro»ohro^oscsffi8S
inoXuding tha sex ohromocooee* Ttia reisuining ^re th© i3ion>»
ohroi!iocc3tiics» Tha ©ooro»ohi?oaocomo8 have been grcupcd in five
different oat^gories aocording to their lengths und centrcmario
indacss*
3* S8x>*cbromosciBos in eialee oonsiet of a pcir of Z
ohro<RosntR«B tihidl ars natacentrio < nd are tha fcurth lurgast
uheraaa in famalea tU sax»chtoi&pso«v.a have been obeerved* Tha
U-chrnfliGHORa inete^d of being ciatueentrie ia small in mizw and
oeeupias 6th position in rrdar of Isnath* Cotspurativa analysis
ravealad similar karyotypss in tha three epeoiee*
s » r? R A R Y
4
In th« prssont yerk a cnapar^tiv* study tif th«
k ryotypa of thr»» di f farent speci«»g Golu»»b^ 11 via ifowif tiot^.
fe.^«fl»^ i i ¥ 4 a m%^mM,% '»"<3 .mm^.qfeff^.'i senegalenoslg belong-
Jlng tn t?rd«r Ccslumbifoifsu® oa« ro«id«# Th« chrc®csrra© preparation©
uers m.;d« u i t h th® help of prsvicusly colchioinlzsd bene fnarrow
celiB of the adul t indiwlduQls^ The c e l l s uam Qivsn hyptitonio
trectment t i l th 0«9 percent ocdiuio c i t r a t e Golutlon^ fiK:>d in 1 s 3
^aotc-loohcl v,nd then f laae end c i r d r i ed . The e l ides were
ct^inod i n fcuffcrad G4®asa8tcin ©oluticn c t pH 6 .0 .
Chrosioscaie© woro otudied end ocuntod frcsi 2S cQoti
netvif^es© plutee cf both ©OK©8 for 0 ;oh on@ of the three epcc ies .
The t o t a l len'^th, the ehort arm length rrtd th© Icnq <-Tm length
usr© measured. Th^ir reean r e l a t i v e lengths (ind pentrofneric
indeoee U&T® c^^loulated. A ooispartitive v:nulyais nf a i l thsise
f indin ie uaa ^Iso f^iade. The cVircmnsomee uore arranged in f ive
grnupe vn t he hasie of t h e i r ©iua '^nti csntrofflerio indtces*
Group one conat i tu tae two p - i r e cf lurge » type chromosrmee.
Secrnd grftip hae s in ?le na i r of t «:hros!nsrtraB. Xt i s t h i rd in
poei t ion i n respect to i t s length , nroup t h i r d ooaiprises tuo
pai re of KfttiXl ohronosonee having t h e i r centromere loca l ized in
the median region. They occupy f i f th - nd s ix th poeit ione «iccord-
ing to t h e i r l eng ths . In the fnurth gr tup, the re -^re frur pa i re
nf wm^ll chromosowae, which have t ' e i r centromeree *st the terBiin**l
end (T chronnsoiiiee) • They «re so effi< ll <nd i den t i c a l i n e l s e
t h a t i t i s d i f f i c u l t to d is t inguish between them. The l<ist
'•a
grc»jp Jl« th« oim pa i r of 8«x chromn«oms«» The maXem H JWB a p«»ir
of Z2 chr'-moaowes, u l th cantt->Ily laCited oentrcffieri^ i1 chrrHBosnpr»»
Tft«y ttra tt^e f©tfrth l a rges t eHre o©»«>@o in th© e«Bpl©»tf Ht i « a i l
the thr«» spsoJlss* Tim fern >1B B@K chromosnffiee includu Zl?
chroii08f^®»« The 2 belni af tho sua© eiza as tha t in pjwles hut
th© U chrofsnr.nm® i® o i i - l ier than th© 2 ahromccmie. I t i s ubout
the sisr© of the n i^ th chromoccm©, but i t o jn he eaftily idawt i -
fiad frcra the n thsr smaller chttiooncraes being taet^ioantrie in
oriQio*
Th© cotapi^rotiw© ©tudy of the three ©peoies of the crdcr
Coiucsbiformes rovsaled tha t thcao cloooly sol^tod cpocies uhich
haw® tacny © i a i i ^ r l t i e s t-moiti theioa^lv®© UIBO posso® ciccc
tesctabl^nco in t h e i r karyotypo.
Thua u© c^n eay tha t the c lass Atf»® i s OR© of thrj
woat ccnservatiua groups in the ver tebra te ©ories.
£JLJLL£AJX£^
7
ni'jlBy, S,D« (1969)
Ally f^.f Ffichhtinar* N,S» and Soufcup,S«UW5)
H8U, ?.&• (1971)
0,(1077a)
Kuol, D. (1977b)
Knuit 0, (197n)
BlanohJLy ^.O. cind Jlcllfic:, 0#3, (1967)
eiocTiB, S*E:*(1969)
eiooRy S*C«(1969)
Panov. E«H* and fl«dJabU« 5.1 ,(1971)
Bulatotfoiy N*8» and Panov, E.N. (1973)
Cutl€r« 0«U.(1918)
M«*ndbonk of the bird* of India Pakistan, tfnl. 3 , Oxfox^.
ami
Xdsntification of ths S»x ohroainsftiiias In ths bttld sagltt. Can.3.n»net.Cytcl« 17 t 187-191•
LooalizJtion of hotercchrciaatin in humr.n ohro<3occTi808« Cytcqsnetios. 10 t ei*es» Sosatio ohtcmosrmcs of noldon cr lo ie -qg^olue* oriolua (nriolidaBf P««^eti« fortnes) * Proo.Nat.Houd.Soi. India, 47(e) S t 64*67.
A oomporotiife o^dy of tbo chrcffiosomas in the fo@ily Sturnidae (Avest Paeseri-foimes). Proo.Not.Aood.Soi.f Indi^. 47(e) XI t 101»105.
Tran^looation heterozygosity if% the ^^^^ t>ooobugQ ounptul^tii (linn) (Ploeeidae t f\\t@B) Nat. Aood. Sol. Lott©r8f 1 (2) 8 e3-»8S.
Chronology ~:nd p,:tt@rn8 of replication in thfgf bone marrou chrofflnsomie of t alJ tig doraestiouf. ChrCTioaoaia, 21 s 387»397.
Mitotic ohtofsnsceses of Hullard Duck. 3* Hered.y 60 (1) t 3&*39.
A current l i s t of ohronosomi numbere ifuS iiiiriationa for species t3f the avian aub^olass Carinat^uB. 3» Hered.y 60 f 217-220.
Oesoriotion of avittn karyotypee of the tlSSf? fauna. Proo.Aoed. of Sci.* 199s 1420-1433(in Ruesi^m).
Compeirative analysis of k^ryotypMi of 18 speoiee of family Turdidae. (Aves) Caryologia, S6 (2) t 229-244.
On the s t e r i l i t y of hybride betueen the Pheaeant < nd the Gold Csmpine fowl. 3our.tienet. 7 t 1S&-16S.
48
t t h r i a t l i i r i t U C . and
Coo ing* , 0«E, and Pta t toec ia , £• (1072)
(Sfouchy, 3« cf©| f i n s f , n» and t e j e u n e , 3 , (1971)
* F c d e r i o , 3 . (1061)
fo td . E«M.R» ond I'eolJttfflt 0*H»^» (1964)
Ouyer, K . r . ( 1 ^ 9 )
Goycr. ««r» (1016)
f l a i t o n , H. and Btsdbuty t P.R.^1966)
Hanc», R»T* (1923)
M^nCB, R»T. (1924)
Hanc», 8»T. (1926a)
G0n»tl8<»l dnd C<|ftoiogia«4 s t u d i e s o f tho i n t s v g s n s r i o h y b r i d s o f
rtcha* Pfac*J^y«5a0« Edinburgh. 56 t 210-24t«
KatyeNtypaii o f f ow t sp9di@t o f b i r d s o f t f i « f a m i l i e s Pltaesidas ^nd Par idas* ^•H@rs» 60 t 134-136*
OfiA o f Jsaainallafii and ^Mian hmtmvo>» ohttmating Ciip«Csll*R®s«p 71 t 113«»131*
ii i jfcoraclicg»aphio p a t t e r n s in c u l t u r e d l#i idcioytes o f t f i e ctofRs^tlo fciyl*Exp« O s l l * % & * y 30 8 353*3 6@»
nin en &t#id(3nc« de l^ @trueitur@ f i n s d«® of)r(3nosoi!©s huisains p«ir d i g s s t i c n smryts^itlo (Ptonase @n p a r t i o i l i e r ) C»K.Aoad»Sei»(Pot io) t 273 S SR7«»Sn8»
C o n t r i b i i t i e n a i e t u d e c^ ouiryotyp© Qhm 1@ pot3 lo t«Af i :h»0 io l»{L i©go) , 72 § 18S-289.
Tssfeioalor cfisciaoEoises o f GtUlus ^OjBegtiaua, CfiroTOsom®, IS t B i S ^ 7 0 »
The spoRBstogeneei© S3f th® dowest ie sshickena (% j | | | a , a.41p|i 0,Mf. | | .mif fJ» rtn<at«An2«« 24 i 22-'24*
' t u d i e s en the ohrctBcsrmes o f t h s doff isstie eonnon f n u l ^s sssn in t s s t s s sn& mt'SfQBm 6 i c l . e u l l » , 31 t 221-2fiR«
OM r e p l i c ^ t i e n p a t t s m s a f t h s ssx ohriswowe^ss n f t h s p igeon (Colustos 1 ^t3Bi^»tii8s) Cytng©n«t ios, 5 t 295-31
Ch i t ^ ohromr^enffi^s *mi$ ssx ds topa ina* t i o n rtnattRss* S6 t 348*
• j -
Ths sonat ie Ghrnmonnnss of t h s cshck <ind t h s i r poss ib le ssx r e l a t i o n s * ^ Seisnesy 69 t 424«>42S*
A coi^3dri8if3n 0 f n i t o s i s i n ah ick t i s s u e oyXtures and In ssc t i oned embryos* 6ie»l«6yl l» SO t 155-169*
4.^
HanoB, R.T, <1926b)
Hutt, r.B, (1949)
Haiamor» 3« <1©66)
Hawser, 8# (19TO)
Itoh. fl«{ Ikeuchi, ?•$
Sasakiy fl* and Slg^inoy S* (1969)
3a i t ly , P. (1966)
^ovunorioy tl« and Atkins, L« (1969a)
3ovanoric» If* and Atkins, t , (19691))
Kriahan, a. (1964)
Kri«h«»n, A, I Haidcn. G«3«. and Shoffnar, R.N* (1965)
KrisHnt^n, K« nd Soffnar, R*N. (1966)
King, « . (1970)
Th« chroii090«a« of tha chick scma, Biol*eull* SI S 443-*449«
Sax <infi tha chrrmneomea in tha domastie foyl (Gallua dnnoaticua)* 3out«norph« 43 t 119-14S.
^anatiea of the fcwl Hc^Graw-Hili Book Co«, Hevt York*
The karyotypaa of nina birds* Harcditlka, 5S t 367**303*
Kuryotypaa of 31 birda* Harcditast 6S t 2B-58#
A conparativa kurtypo study in fourtean apaoiaa of birda* ^uoan* 3* Genatioa* 44(3) t 163«>170*
Struotyro end behaviour of 0hro«BO8otBQ« in Dinooiuw , t2,qte a,y». (Unn*) (Pic idaafPic i forSaaTi r>roo*{9at*Aoad* Ooi*, India, 36 t 24&»248*
Karyotypea of t i^ north Amariean {nintinqa (Avasf Eaberisinue), Qenotika (SaonrQd), 1 t 165-175*
Karyotypaa of four P^aaarina belonging to the fonil iaa Turdidae, 'lisiidaa <ind Corvidaa* Chroncaoma, 26 t 38S-394*
tliero-c^'rowoaniBaa in tha apamato* ganaaia of tha donaatic turkey* txpm Cell*l?a8*, 33 t i*»7*
l i t o t i e chronosopa < d tha U-aax ohronoaofia of tha great horned owl Ijkibo U viroini ami a) • Chrnaoaoma (BarT*!, If t 588-263•
Sax ehrn'nnaoiie in the donaetic fcul Turkey
ind the ohineee ilea)*
!^tctie ohromoaonaa ef the Auatralian king quail Caryologii .i«,cM.u(§sjsius^j5jSj3,a).
5 0
A n » a r l , H^ll . (197S)
A n a a r i , H«A» (1978)
* Li iy««, «• (1906)
* i e c a l l l c n t A. (1010)
and Sandberg A«A«, 1964
n i l l e r , R,A, (1930)
n^k i nc , S. (1041a)
Ptiikino, S, (1942a)
.^cikint?, 8 . (1942b)
«1aklilOt S. ( i942c)
Hakino, S. (1943a)
ScMKitic ehronoat^ i s of m^lm Aco la | t t i y taUiiiUt* ^outmHrntti^ 66 t 175-176.
Chronosona s t u d i a s I n t h r s v spaciva o f P i o i f o r a a s (Av8s)» Ganat lon, 48 (3) t ISCKl fd*
HsoharoHas But i * davalopment ova r ian daa oaufa marobl^^atlquaa a tfltallua n0 t r i t i f abnndant* Aroti« d ' Snat . miorosf 8 t 239*
i s v a r i a t i o n du ncHibrc ds?s chrojanaefaa dawa l a aegmcntat ion de g» oeuf non faoonde ^m l^ pou la* C«R*Soc«6iol«9 69 i 34«36*
^Qsienolatura f o r csn t romer ie p o s i t i o n on obrofliosofiioa* Harad i taa (Ujnd) 32 t 201»220«
Spai^otogenosia i n u a e x r a v c r a e d fsKav:!© ond i n oonaol taalaa of t h e doeiast ic f o w l . Gal XMS flcwpp.tiCMS » Anut .nec . 70 t 155-1 f ^ ,
C y t o l o g i c u l i n v e e t i g a t i a n a o f micu i n c l uded i n the cjenus i]si&» 3cur«fao« So i * Hokkaido Imp. U n i v . , Sur« V2 2 c r l « 7 s 305"3a3,
A k a r y r l o n i c a i s tudy o f go ld f i s h o f 3apan« Cy to log i c 12 S 96«1I1«
Po l yp lo idy i n un iT i - l ( I n 3dp<jni!taa) Bot» and Z o o l . 10 s 1S5-172.
Chrowosowa cnrnp^irlsion betuapn/ .u l ld form ^nd tha dofRestioatad tf.jriaty o f 5afciMa» n ,yygq, ,9M| Serkanhrut. 3ap. 3cur»Genet. IP » S3-»5*5• .
l U Studlaa on the murina ohrowcRnmra norpho log ioa l cnmpar is ion n f tha ohronooonea hetuaen tha u i l d f o r a ^nd tha domest iodtad V i t r i a t y o f Rat tu f nory^g icua Barkanhout. l ou r . l acSdT , kokkdido l a p . U n i v . , Ser«, V I * Z o o l * , B S 31 -43 .
Studlaa on the murina ohromoaonai I I I A conp<ir«itiva study of chromnaoaaa. m f iwa spaoiaa o f Rat tua . 3 o u r . r i i c . S c i , Hokkaido lap.Unlv .Sar . t fX Z o o l . 9 i 19 -57 .
51
nakino, S. (19431})
nMim^ S* (1943c)
^akino, §• (1944a)
The cHromosoiRss of th» hor«« (tmtup o^atoallum) chr^oscflMi atudisa in doiHiestie m^mmalB^ I* Cytologla 13 t 26-38.
The chrowesfais ccwplaxsii in ge«t "Ci^six M.gmi)^?'^ ®!!?®p, <M„i asiaa) and th®lir riilationahip(chi'ewoEO«« studiim in dcmesttio rtiam'ttols 11) Cytolociia 13 t 39-S4* Th® cshtriBOaomo cojspl»x of th® pig (Sufi a S S U ^ (chrr-mnnris® studie® in doffiestic «'aamal9 H i ) Cytologia, 13
fl^'klnet S. (lS44b)
* {^okino, S« (l§44o)
WelarJdar, ¥• (1950)
tieklno^ 8»y l-ldaqaue^ T» and Yymashiniig Y* (i9S6)
%7rick» S«y l e d l t y . R«S* .jntl iuhsg H.A» (19?3)
Newcoffiet-y E*H* and Brant, 3.y«A.(19S4)
NoydQ»»fg E,H, (1957)
Nnuocwai', E»H« (19S9)
Olaan, H»U« (19S0)
Ghno, 5» (1961)
^aryotvp©® fif domostid e e t t l e , tntsts anti dewttstic y^tor b u f f ^ o (cshrcnosfwa s t o i i s s in df^ncstic mwffl'-iol© tfl) dytologiuy 13 i
Chrcsioscme® of domestlo ©om'SQle (In 3apLinm@) ZodlcBag* S6 t B»IS,
Accessory ohrosjo©cra®s i n anim, ls , epsoiul ly i n ^ " difeas, 36 s 19»'< epsQiuliy i n golyoe^ii^ I».nyi..^# Here-*
"" ''SO*
IC«ir)t:iypG etudiee in b i rds 11 , A oo«p^!ar-'titf» ©tudy of chrcimsowe® in the Cclumbiddte* Caryologi@, % t 21^2S3»
Ptcduction of G -.nd C bunding y i th proQres'^jive tryp.»i'n tre^^twant Pediat.Rs®. ? I 39-44•
Sparmatoisnesis in domomtic foul 3eur» Msred* 45 t 79»B7«
The la i to t lc chrowosnraee of the dopestlQ foul ^ u r * Hsred* 4B t 2?7-23i#.
The n e i e t i c chroMoeoaies nf the foul CytnloQia, 24 I 403-410*
ChickenitHcsy hyblrtds* 3eur«Hered« SI s S9 : 73*
Sex-cHronosomes and mioro-ohrt3mner>nee
11 t 494*498^
52
Ohno» S«, Stsnlus. C;
Urn iTtti Bacdky I1*L«(J1964)
Ouan, 3.3.T. (19IS)
Pieeiniy £• and S t e l l a , n* (1970)
Renionl A* uod tfngni-TRlIuri, ?l.(1066)
Ray«>Chaudhurl, S.P«| Ray-Ch^udhurl. 8* v nd Rhv'iWii, T« (196C)
Ray-Chaudhurif H»|
Ray**Chaud!u!rit S#P»(i969)
« Rodjabli, S»l#} Psncw, E,8, and Bulatouvi, ^US* (1970)
Raean. R«s 3aocb« t «s and Shsjrraa, T,(ig70)
» Sonnenbtodt, R«(1909)
SokolowilkaJ<a. 1«3« (19353)
(193SI})
Schmldy U* (1962)
Stsinuty C«t Christian. L«C* and Ohno, S* (1961)
ChrcNBosan*! unifor»ity in t<is avian aut}»olaaa Carintitaa* ZhTomomoma (Barl)) 18 t 2aK28ff»
Kdtyotypa atudiaa on Sallua ffmmfXl^m Chro«r>«aiia (Barl*), 16 i fiOl-fiOB*
Scmte avion k<iryogr ffia« Caryeilcqia« 23 t 189»2n2*
The k^ryogfama of »o«a raloonlfonRaa <^n^ Strigiforaias. Chromosoma, 20 a 133*1SD»
The U»Chrt5mo8orao in tha fera.il«8 of tim Indian apGoiea ef birds* Chtemaiioma^ 20 t 1S1«154»
A Cospar-^tiva study of the chrcsmrBamas cf birds* Chromosoffia (Berl*)* 26 t i4eoi6a»
Cc5{apar:ititf8 studisa of tho karyotypea of two closely rel-ted Gpecie® of buntinga (Em!?flrAli?g. QAW^Of „}.?. * ^ nd ,f ,fefMlf ^effi«^ffP^^^aa»,ga.> hybridiasd in aroaa of overlap•fool*3*, 49 t 105^1663 (In Huasian)*
Hatarogananeity in distribution of oonatitutivo heterochromation in four sp^oioa of birds* tjenetioa«48(l)*6t»6S*
Dia Uaahata;m8p3ri od^ dar Oooyta daa Huhn08*Aroh«fHrikr*Aniit«9 72 t 415^4^*
£a(pari«ants on hybridiaation of birda* I* Sax^linkad charaotara in hybirda of
im^m ^n^^fp '^Am^i nMn^i^m I a l t s zooi<
tfargleiehanda untaraushung dar karyotypan Uon Cairin^ atpaehata. Anaa
3l«it|vrhvnc».und dar dureh Krauxuno lasardtrtan arb^^ltanen bastards.
ZBit8*Biol.(?toak^M) 4 t R93-904.
OWA rsplio>.ticn p^ttorna cf tHs hatsrocbroffloaciBaa in ggiljia. d,«ffllB lffyff* Cytoganaticat 1 I 344-352*
Cowparativa cytnlofjical study of ^ffli^y^^t ff^^^jAfVftt "t^ffiqrtff
cnromoaona (bar l *^ 13 t 515»520*
t) '1
St«pho«f K« and Arrlghi, r.E. (1971)
tlisra, I1#(i97i)
Sstibright, fl* (1971)
Stock, A* 0»f Attlqhl r»E» iifiti StefoSy K* (1974)
Tafcugl, Urn and •lakinOf S« (286S)
und Saaakit (1* (1072)
ThBrCP'-Bscu R«C»(197S)
Udagaua, T* (1953)
Udagawa, ?• (1954)
• Wan Brink, 3.PI. , 8 t L ' b h e l s , G.A. ( 1 9 5 6 )
* Van Brink, 3 ,n , (1959)
Uhit«, n. 3.D« (1949)
uhitt, n^a.o. (19S4)
H«tBrt7chronatle natura of yh chvofioaoiHi m blrd«. Cxpt.Cvll.Ras* 7 t 39-44.
dreapcto (Coltmt3lforiRM7 3,lf«wid.f fi2CS} I 373-374•
A rapid banding taehnlqua for fmman ohromottosws. Laneet 1971 I I , 971-972*
ChrrtiiGsois* homology in birdsf banding piittarns of the ohroraoaoiias of tha doaestlo ohlekana of tha, rlno-naekad dova and doweatle plgaon. Cytagenet. Call.Ganat*, 13 t 4t0-418*
A revised study on tha ohronososaa of thraa saaclaa of blrda* Caryologla, Ifol. 1 9 | 4 ) | 443-455.
Chronososaa studies In four species of Hatltua (Att68)» Chromosoiaa, 36 t 281-291.
Tha kuryotyplo evolution In two Palaoanlforiusis saQCles lAvas). Caryologla, 2 6 ( 4 ) | 459-466.
Karyogran studies In blrda. 11. The Qhroaosomaa of thraa spaolsa belonging tn tha Colunbidaei Ardaldaa and Aloldaa.Annot.Zool.Japdtn., 26 s 2B-31.
Karyograa studlaa In blrda. Itf.Tha ohronosoffiis of four pasaara and tuo Colunblna apaolaa. AnnQt.Zaol.3apan., 27 t 20B-214.
La ouaatlon daa hataroohrotROsomes ohax laa aaurnpsldaa. Olaaaux. Exparlantla^ 12 t 162-164.
t.* expression morphologlque da Im dlgoMatla char laa aauaopaldas s t las nonotreaaa. Chromosona (Bsr l . ) , 10 f 1-72.
Chronosomis of tha vartabratea. A ravletiit Evolution, 3 t 379-3B1.
Anlnal oytology ^nd Evolution. Caabrldge University Preea, New York. 2nd Ed.
li 1
U&n^m H«C« and r«doroff (1972)
Shaffn»t, R»N« (1974)
Yamaehinuy ¥• (1942)
Vaeashina^ ¥• (1944)
tft isi^ehina, ¥ • and n ^ i n O f S« (1946)
Sanding In hif«««fi chromssoiMis tmatad uith trypsin, ttatuir* (Lund*) N«u 6iol»» 23S t S2->S4.
Trypnin G»and C-b. ndinQ for intarchdngi analyni* .Hind «0K-idflntlfieation in th« chicken* Chrc^osiHia (8«rl«)» 47 t 61«»69«
A revissd study »f the chroMtssmiitB of the iButoovy duoky the donsstio duck tind their hybrid* Cytologic 12 t 16^169*
Karyotypo stitdies in bimJo* ! • Coffiparativtt laoi^hology of ehronosoMut in um»mn%mm% raco« of dnmestio foul* Cytologiuy 13 t 27CI->296«
A study on thn chrotscBO»e@ of piQuon @tfld d0V8a« Soifoutuy 2 t 92-*100«
* Referono38 not consultnd in original*
r 1 G u n E 8
eale Indiafi donestio plgoon*
b . Uat« diplotena of the pr4©aipy spernotooyte of Cclyaba lltffat
dnwesticQ 8f»oyJin3 the t&n lopgest pair© of caaoroohrcsosaas*
FIG. la
FIG. lb
I I • fWtaphasc plats •hewlfiQ tH« •onatie mtteroehronottOiHM of
thti f««al« IficSian cftMKKitio pigeon*
58
I
FIG.2
I l l , «»t«ph«»« i>l«t* «hoyii»9 th« •o«»tla •ac»o©hrfi«MJ»is«»« of
• • I * SndiAH ^lum took nigMfi*
60
FIG. 3
Itf« nfttaphas* platd •houifif thii aonatle naefeohromsBOiiM of
f«iAl« Xniiian hlvm took pigson*
sz
FIG.4
V« notaphaae lalat* •liowin« th« •onatie aaeroehroffiosoiiM of
•a2« Xntiian l i i t l s totoun dova*
84
FIG. 5
VI* Retaphatiii p la t * •hewing ili« acNittiie •aeroohroaosoiias of
fMia l * Indian l i t t l * brown dov»»
63
FlO.6
VII. Karyetyp* of m»l» Indian dowiatie pitmen with ZU of fomao
in tn« insset*
6S
II II • • zw
1
II 3
5
7
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mm
• IP 8
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FIG. 7 • I
V I I I * K«ryotyp« of aa l * IndiAR 6Iu« rocsk piQvofi with 2y of
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72
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74
PERCENT RELATIVE L E N G T H ( L R )
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PERCENT RELATIVE LENGTH ( L R )
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21
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