a comparativ stude y of the karyotypes of some species of ... · - iv • xxx. hom!phon0txy of th...
TRANSCRIPT
A Comparative Study of the Karyotypes of Some Species of Birds from the
Sub-Class Carinatae
THESIS SUBMITTED FOR THE DEGREE OF
DOCTOR OF PHILOSOPHY IN
ZOOLOGY (Genetics) OF
ALIGARH MUSLIM UNIVERSITY. ALIGARH
BY
ZAHiD ALI
Under the Supervision of
Dr. USMAN M. ADHAMi
DEPARTMENT OF ZOOLOy ALIGARH MUSLIM UNIVERSITY
ALIGARH JUNE. 1982
T2521
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A cm^mi^im sTioir OF WB tummwtQ m w wjsm mm ms avBJChM^B emm/^m
wmm Bomr^^ ita mn
mmwscmt m mmom (cnm^msi w mioma
miG^B
m
zmm ^
mvcD. mm Qwimmms OF m» mmm H, mma
tmpmrtrnm or socLOor i^iuj-HH {fjsxiiH mvimsTT^
mtcmn
thio im to certify that "A CfsmmNSVTEi m!tm W
Ttm Kmwmm m ocm cr eiam mm WB
Bm^tMiQ CP^mmm"* is me orlglQai wotk g£
Hr* gcftiid idi is eait#>2e fc^ t ^ o£ tlie
^gesm of soctor of Phllom^hy in Sof LoQsr (0@mties)
of Muslim Mi^arts* ThiB
«oele has h&en kaat im^r ok mi^evisim*
( tBMI© I — Readiir Copurti^nt of SEooiow Aligefli Hu^la miVBSQity
- i •
fpKmmMtmmmm
fhe nuthor tflke* Gp|>cirtiitiity to «]cpr«»8 thm
a««pe«t Mnse of oratituds to Cur* UHnm M* Atti iiii «Iki
%t. «v«ry asoawnt to taring tho aissttrtation In tiso
ptmtmt fomrn Uim oritioai adrico end sugg^stioas
gemmtly adsairodU
Omtoful thanks ar« to Vcofeasoi' it« Kh«n,
Chmimm* mpm^etsamt of Itoo&ogy for ptmi^oQ oeoesaacy
CacUltivs and •tMsouragsasfit* f ha i ^ are <3u« to ttt«
Cblvaraity Cc^anisaioii ae^ Council ofi Sciaotif io
anA Iftdkiatrial. Hosoarcli foe fintfioial asotataace in
form of ra^HKStinrely.
I an graatly thaOKf ui to any vifo fira* S^at Siltia
i«ho has alv«a aM an cq^rtunit^ to utUic« tha Aaxiasia
tima in conAueting tha raaaarch i«otk witliout any
difftti£l>«f)C«* Mir eoqp«ratioR in (Shacking and cosqparing
tho ia «p[>r«ei*bla»
iio»t aineara thanka ar* aioo dua to Or* WaMin iiM«d*
Mias AnJuM Ara «id othar eolXoiiguM of tha dtpartatnt
for thair vaiuabXo auggaatiooo and holp*
u «
S f t B S - j a L S ^ a a r a Mo*
MBoiiittagpamt
of cpostsnt*
of TMw
I4«t of Vigiixtm
mtrnrSMB and tiBt!i«»(l»
«ifaaL«9
Cbmm«K$xmm ftestaits
GiseUMilMB
Fi^tira
Coaelu«ioii»
i
i i
v i * v u
2 » Id
l9-> 34
2$» 4i$
70
t i . 94
SSi
- U1 -
Pmgo Bo,
I» Oipioid chrono«oa» nunber la v«riou« of biJPdUi (Sub»olftS8 C«x>ifiiit«i»}»
IX* torphoraetry of th« scmtic oaerochronosow* 26 pugpunm munUmn»l» (Kmatmm
pueplo hoconi*
XXZ* MoKphooetry of the tuo ftros aonatle d7
lEastfrn purple imgcSTm i
iffm HocphoiMitcy of th« aimtie aacroehrofflosomes 36 of m ^ U SESXil SE1E41 ( Xttdian pond )i«coti)*
Itorpho^tiy of the t%fo tinas of acNBifttie 39 {ii«croehro^«>aie8 of ^aopiy ^aiv^^ qravil (Xndittii p&aA h»Rm)*
VX* IfoKphoawtry of tho soiiMitie oiACCochcoaiototaes % Qg ^^gjHy^lllMi, iO f f p«rtrSdg»T*
VZX* MojrphoMAtry of the two ar»0 of stffwtie 32 AacrochromDiH^s of p^wadicoriinKi
pirtridg®;.
VXXX* Mogphofli»try of tho aoaiktic ivtefochranoaoiMi* 92 ^ Qpgyp* (liou«o ccoir)*
XX MorphoiMtry of tho two «nMi of MMMtio 39 w r o c h f o i a s ^ s of StiOOUL IBUBStol. (Houao ecowi*
X* ll»xphe«otry of tho sonttie msoxochroniDtonio 34 f t f i t r
XX MorphoMitry of tho tm oma of ooaistie noeie^ 39 ehro9osoao« of Pao—r pmmmtlaam <}iou«o •parroir)*
- iv •
XXX. HoM!phon0txy of the mmmtta m»eecchwomm»im9 36 of rtlirillH l urunicapa (ll*dh»«d«d bmtinq)^
seXXX, mgphomtgy of th« tm »mm of •emtle 37 aaeeoOireao^oiMs of Msmrtrnm ha:tmicmom
bunting).
XXV* Coiv^tfrlaioit of th« q»«ii ci»l«tiv« X^ngths of 36 •emfttic ««cfoehroao«oni« of , (D«t« tosMd on
Xif* CoRparisioii of tli« centxoiaftric indices of th« 39 •oattio mafixoehxomoaoBmm o£ ^d f a purourOA
3arx« Coapaffiaion of th« mean rolative of 40 tho two lurtaa of soaistlc mmewoehf€itmmom» of
.fiSi&eiapA. itgHTOfflif ^smsls iEflXtik » V ma
36rxx , CooipariaioB of iae«n rotative lengths ^ 41 aomatio aaeroeh«(m»aoaaB of
JOtXIX c^apariaioa of tha taaan cantsoniaric indioaa 43 of tlia aonatie eaerochmioaomee of
^ l a a v r n r x w d XXX)*
XXX* Canparialoa of tha two anw of aonatie 43 attdoahjtenoaeaaa of Conrua aalandaBf^ SllliC. dowaatictta and l^jfcariaabMicapa CSita loaaad orTra rablaa
XX* Co^pariaion of tha aaan vaXaUira langtha of 44 tiMi aonatie Maeiroe xoaoaoaiaa of JUtdaa piueoyraf •aniiaaaia. fjgatrti g m i l mYlCffflBit^tffMI
» V •
XXX» Cflqparl«ioo o£ tn* iw«ii c«atraiiftrJ« indices of tii* 4$ itic vmcrochr&mmomm of Jgdea purpuiff
, , „ rgtnffgiityittf i f w a i w i . ikmm^im*
_ _ aSl|4SiE a on M,
and itm
XXXI« Coopcrision of t;ht» m^m colstivo iongtlui of tiMi tMO •ma of aomfttlc saacfoclufonow iea of
F T ^ f W f ^ f t yfty^jr m y f i
f fr>—fty ^ ^ t i c ^ ana pjBte^ifjn fcSiid on Ttttiloa IXX#V«VXZ«ZX, Xl ai^
A eoipariaion of tha atodiaa of and 81 Patnaik (1977) %dth that of preset «oxl( on tha c:|uno8ioaoia8 of Paaaar domeatieua (fiouaa
• -psT cr yiQtiuw
p«g» No.
Karyotype ahovlng th« hoaoleoou* peirs of ttw 95 Mnamtle BMier«3Chrofao»o®«« of ArOff purpugf mmllmn»im (s«st*rn haron)*
<lb) H»taiph«se piattttt shovino the soiaatic c^rotao* 9S ^ ftrafa tynrmi ai (Eastern
l>uxpX« heroiil*
<3 ) K9il !jfogcm mhemltiQ the p«r cent reiative l«agUi«9(l o£ thft aomatlc Mccrochxoiaotfoaie* ot »fdmm
iaainll«wi» (i^tmni ptttple oironU
13 } Kicyogem shotdnrr the per cem reiative 97 le>hs of the long 4H»i9 alKirt arcss of th« •cxnatic raneeochrowMtoniBa of Agtfaa puigtmraa manileaaia Ci aatere pucpia heron J,
(4a) Karyotype showing th« holtolL<»goua paira of tha 96 aoiaatic laacrochroasoaomea c^ fypdaoif oapavii grav^ (xadian pond haron)*
Nat^piiaM plataa ahowing the aonatie chronoa* 98 of Ariael* orayli (Indian pond tooson).
(5 ) Karyogrvs ahovlng th*- s air cant ealativa langth 99 of Hia aonatlc macKoc roaoaomaa of jyraaela qriiyil qgayii (M ian pooa haron>*
( i ) Karyofraai ahewing tha par cant raXativa 100 iangtba of the loog and atiort araa of tha aonatio aaejcochromoaoiaaa of MfllBl*. qravii (Indian pond hax«>n)«
(7a) Karyotypa ahoving tha hoiaologua paira of tha SOS ao«atie wacgoch«Oiiaao«aa of irraneolinua g^MPfrtailii iB t fBWi t i i «»r«y partr&dga).
(7b) Mitaphaaa plataa ahowing tha aomatic ehxeao* 101 larfnytUnvt wm^i^^mM lB$*SB2iA&ia
(dray partridgal«
- v l i •
(8 > mmcyoqtm shoving tb« p«r cmnt nlntlv* loa of the somatie mtcrochfonowataM of
, f f f . a g g ^ C T F l m ^ kmmUPm Cor ay partridgiJe
<9) Karyograiii ftho«ing the p « r cant r«i«tiif« 103 lengi^ of the img and short ants of aemmtle mtxoehxwaoaomttm oC ^itwco^inus mpp^ff^rlwiat, 4fiSSSBfiejl$$aB partriage- j
ClOa) iCaryotsrP« shoving the heoologcma pairs of 104 ^ a somatic flmcsrochronosomes of Ctefim^ sp^aoaatis (Bousa <aeow)»
(iOb) Mstsphasa piatas showing the aaaatic 104 of goi tia f^lnaaws (Houaa eeoir)
( i t ) KaryooresQ shoving tha par cent iralatlva 105 laiigttis of tha mmmtlo aoerotchrc^soaies of fiMOQi*
<ta) K«rrogr®B ot»wino the par cant ralative 106 langtha of ^ long and short ams of tha so^mtic tBaerochrotaosocias of gory^a SDlandana (Houaa crow}*
ICaryotarpa shoving tim hesnologous paira of 107 tita soraatie aacxochcotiioaotaaa of f^jsag igoaaatittis (Housa aparrow)*
(13b) Nat«[>haaa platM showing tha aoenaUe tihrosi* 107 osonas of SMUESr SESESlSBfc (Houaa yarrow)*
(14 ) SCaryogram showing tha par cant raiativa 108 langths of tha soniatie mneroeftixomtmoma of Sftlf££. ^gwfHyjlff (Housa sparrow).
<1$ ) Karyograta ahowing tha par cant ralativa 109 langths of the long and short arnw «f th^ aofliatic naefochrofliosonas of yaaaar dgwaaticua (Hbusa sparrow}*
« viii
Clia) Karyotyp* ahotring tii* tmmoX»g»vm of 110 tiitt MMBtie WMCfoelironMcmis of Wa rntmm iMPHIiftfft (MhooaMl hmtlmgh
(til>) iii««>liaM plotoo ilhmrin^ tho oooifttie ctiRMd* 110 • M f oc Mfr f f l i i iwnimfwf BIlBtlJIOl*
<17 ) K«ryogr«ai tfhowioo ttio p«r eont roloUvo t i l iOR IkO of tho ooaatics eecmchmiweoaiee of Bwptrimm tafuaieooa teiRtl«9>*
(te I ahotrifia tho por coat r « I « t iv« 113 loogtl^ of the ioB9 ittort aciMi of tho •oouitlo laieioclifonoteaiB* of ftaori— lin^ili^o., hmtimgirn
Ca9 I Coiq^ercttive lefiryegra® ohowiag tli« cent 113 c«l«tivo loagtlio of «Imi oomatio musm* of MfllBk pmroma ayilloafif
p i ^ I i ^ y o o l oaa i| i poodicagiiamii i o f taooitiif tOroy pagtridaoK
<20 } Cot pttTotivo kcryogrwa iliowtna tSw por e«iit l i t longtho of tho long «iiA aliert mxmm of tlio •OMtiO MOSOClurOMOSOMM Of
L<lo«%iuni eeeme of SfliB. eilBBillSS pucplo liiwoiiTrSpBBr
_ an poMi hvvofi) tSZ IgijfiifriimM lattTOftHm
<21 ) CoapmtXw luMrmir«M alkoiilog tlio pojr coat liS coloUim ionfiMwi of tlio ooMotie —oiotlmo-
of Qnzyt^ iBl l i i t lA ofow)* t, moiioo apiurroiff) oaA |,<IMlO|M iMRttHO).
< 22 ) Coiporativo kormrtfi ahoidiHi ••At i i « loiotivo lo»g«iio of th« Xoog oihI aliort « m of «i»o ooeifttle —oiQohffooogQw of fiilXliii fploaaiMftlBtt— etowh i i i f£L tjHHtftlffll (Stouao ep*rrow) mA t^Ti iTWynieaoo
A a 0 Ik C
mtotic chromsoma istm the prccoilebieieilft*d
^oneaarcvm cfdls o£ six of birds* ArOBm ourpur^ft
isSm^M sesKli arayli order
Ciconifortses* EUfflESlima la^os^tttff.
bnloofing to order Gallifomies^ Corros
^asaeg cea faptniccrta oS? th© or<^x
^asserifortae© t;©ir© ©ttjai^ Mter hspotcaile treetaent
wlfti sodium citrete sdutioa* ttie c ^ i e ^ r© f I k ^
in <it3K Siidcs ^ r e prep&ec^ £l&m
isqS eir drying nethoS e^d ataioed in Cored Oi^ssa
at 6.8.
f i ^ t ^ i v © ^ ^ oeta^bese piateo ^ r o ©electoa
for chrcsaoeeaai count in each aj>eoies. tio totei leog^,
the long asm lecgth on^ ^ e a ! ^ t Ikeogth i ^ e
from SO mi l ddtiottatea matqphiis© pietses.
Hieir meep relative ieogtfia m^ c^tmceric iodiocs
«cro cftlculatea. fbe chciKaosoaaes were tirrafigc^ in groups
in dftscenatog order accor^ng to tJieir siea centronieriQ
iiuUcea. Zit all th@ six aspmsiea etadiedy th© amf lhsomoaoBmm, ciaasififtd as macrochroiaosociestf mte the 4tt iargeat
pair in sias and eosprised a peir of 82 ehrosaosoaits io
«al«8 and SSM in femaiea.
Xn purpury^ fiani^cnai^ the f i r s t 12 pairs
tiav« bean desigxiataa as tha tsacrochraix^aoaies. Qroi; oai
o
eoisstltated tuo pairs &f lerge ii»-chvomDSO!9»a« ^
seeotid lies tt slnole pair of atsxhrostooGoeo*
third c^sspriMiS t^rer pairs of malZ oi^t^a € uroiaQS@ai88*
The fourth gsroap has five pairs of mrfXl fMi rcniooo'n&s*
The f i fth iccitKies th© seje^-chroaoscms io tiilch s«
chromosorzK a is ma^tw^ ^ rc^iniog
19 paira ero tho 0iiKcroeiur4^sosGciea« a v&riaticm in the
i|>ioi<3 tM «:hron»3aaQS in Md^B^ rfflrpurpa pmilmBlSi
^ Q fsoted from 94 - 66 ^i th o e^tkeil pe i^ (40?S) e t 62# t3t®
iie^ Quniber }ming 6I«44 6«604*
2© my CQBo OS ^r^a la SSiXll. SffiM. ^ ^ 12
pairs ero <2eoi<|i3ate6 ea tho taacrochjnoooseaeo. <Sro«3 00©
eonatittiited a etaglo pair of i o r ^ »-chsoiiK)socio0. Sli©
S€«on<3 cP'oup hG3 a oiia^o nair of siiiKt ?!?© chrcmmjR^e.
Qcoup third cosi>ri@eo tiu:© ? pairo of moil.
'At!© fourth grott> has a singl© pair of eai-chrotaoeo' es. She
f i f th grou^ ccaipriaoa fiv® pairs of asiaXl T^type chrofaoaorooa.
Qrot^ sixth includes t ^ sm'Cbr&noao'mB in whid) S-chsomoso*
Btes id fo-typa ^ y-chroiaosouti ia ^tjfpm* ^lie remaining
20 pairs are microchroaosoaies* fhe diploid nydt^ o^
chroaosonies in Ardcol^ ffravi^ varied from S4 • 68
with a markeu peak (S29&) at 64* the taem being 63.6004 0«68e«
First 10 pairs are designated as the iaacroe miK>somes
in prancolinaa ppp^^cfyiimyy. msssssi^a^ Group otm
t •) o
eoBStituted • siiic^* pmis of lara* mNclimiotoiMs*
<lroi|p« Meeoid* third «isd fourth also hmm •iagUm pmlem pt iwtsnpt and cfiSDaototats c«i|Mieti¥ttly»
f i f th eoai^iM* tliffte of •mtll
<lPot]ps •ixth «»d ••vttBtli iiisiud* aln^e pmirt of an* «fid
^hrotooaoiMW raspaetiirAly* nm last roMp tn^u^e
zs^zw eheemo^aes which ac« Om rwnsftintiiQ 29 paira
ar« tm laiesoehroiaeaoQias* ^Iw di^^loid ouodDar of chnBioMiiioa
j FASSS ifcJtiSSftii • Si SSBSSilSiiflB' 63 • 66 with Q nmrkftd pei^ 19291) at the laasn baing
gQgyuf yplaadans the f i r s t St prnim are aaaigiidt^
as the rsacrochiroa&aosaes* Oirousp o&e c@38titatea «
pair of lariya a!i}«ehmao803ie8« Qmap seeood also has «
•if i^e pair of fwtype ehcoiaosoniis* ^ third groi^
GO«i|^riM« «t»t3rpe of a aiUQ^o pair of chroaoaoiaas* fh«
fourth gro<Q> iooludaa ? pairs of laediuia aisad to msmXXm9t
mt0tochmmtomam last groMP inoludss ^m sax*
ia ^ l eh ss cfhroaiosoeia is ma tSTpm «sd th«
eh»oaoao«aa is »-typo« Tha raaainina 27 psirs art 0m
BierDChromoMas. His varistion in diploid maitmra of
this spscias rangsd fra-a 73 » ^ with the b luest peak *
(BM mt nsaa baiag 74.960 ^ 0*57a«
in C6S® of gaager do;a&»tica» l l largest pairs arc
designated as the eiacrochromoso'xeis. G^tip <mB coostitutes
single pair o£ i a r ^ c»«chriD3eMi& ics* ^ ^ seccmd groi^
also has e sio^e pair of st»»cticonioso-nes« Grot: third
cooprisea txm pairs of nt-ehroaoeosoBs* fcurtti geaap
include t ^ pstice of st>>type ctirotnosooies* f i f th
has pairs of smell ta-K^ooos^^s* ^bo eiKtl>
incluOes tro peiro of soallest Group
serree i ^ l o ^ s tho 6Cs>-chia33oso.'3t©s» Sstli D-anfi
Boooo in tliis CQse ere ca-chrossJS aBS* El© rcaeining
36 pairs ere the Qicrt^hrc^scnsjSA Opioid Qunibor
of chrcQosomoo varicO i rocn 6 0 - 0 2 xst^ tito hi^hoot
peak at IBe ^ o noen fcaino 76*720 ^ 0.5?S.
to ^ftteeriga II laroest p^iso er©
desifinat®^ m the maercchrcnsscuBo. oa© constitutes
t »o oairs of: largest H chroTJosonms. The pair
hes a pair of 8t«c1iroT»somcs. Group tliird conpriees a
single pair of itttdiuin sieedl ta hsornBO- BBm fmrfSn
group has two pairs of sti-chttraosoroiss. Tiso pair® of
snail in-chrowsoT* s conatitutft f i f th ^ro«p» Group
sixth contair.s t ^ pairs of stsdlc^st T-type macrochro-
nx3Soses« ^ff last iroup has the sex- h Tmosomt s to
which S;i>chronioso.ae is s>a»typfli aod ^-is 'iim
remaining 39 pair® aeaignat^ m ajycrt chsonosoiiiiea.
mm fit«it^ in ^eciea varlcid fapota 70 • ea
^ t h the I48S) 4t eOt ^
?8»400 4 0.56S.
^ ccsEiparetive eualyeis of the cssxptiolo^ end
oi ^ sesatio c^waoscaDO in ^
spcscies stuaicd has bcos lae^ estS tisoir feer^oio^^iciii
rc|.Qtioiasli^© with recjject to th©i& eloooif icatio^
ptiyXogeny lidtro beea iSiseusaeiS,
i; R T R O D If c T I o ^
MiA*« r«latioii to birds tegan in pvtliistoric tiiBfi««
imMesm men on birds wer* «iell Mtsb*
lishsd and ths nunter of flifi|hU«ss forms was f « r mora
than to«d«y* fhess birds fould b« easily ci^jturad and
man's f i rst intarast in birds %ias as food supply*
For ganstie analysis the foul %»aa the f i r s t
in which the mnOmXlm principles o£ haradity vara shorn
to hold 9Pod« Batas^*s aarly aicpariiaants attraetad tha
attantioR of other ^rScara in tha fiald with tha raault
that a baginning has bsen meda in tha analysis of aany
and variad char ta r s found ixi diffarant doiaastio beaeds
of tha fovl* Many mora y«ars would h@va to ba davotad
bafora tha analysis could ba coiiplatad avan for the a as 11 y
astaetabla charactars involving visibla diff«r»nc«s of
struetura or colour* Howsvar* tha nathods of ganatie
analysia ara now kaowii* Sot only tha various charaetar*
isties of fowl can ba analysad but manipulations could
alao ba laada for tha cacoobinatioii of dasirad ganatie
traits*
for intalligant obaanration# aecurata raeording
and authantic iataipratation of data Haadalian prinei-
plas provida awch acapa for Obtaining rasults of dafinita
and parnaofent valtfi*
IG
fh* BuaaMur ef poultry owiwrs heEv* it in tiiair powtr
to holp ia«tftri«Xly in buildino Mp that ccnpttlmtm knoirlodg*
which wt) may oo w day hope for.
Recant progress in chroa»«on«l studiea of various
animals is vary raioark^la and the probla!» of raeiai
aistinetion relating to ehronosoisal faaturas in domastieatad
aniinals is bscoming m interesting ^ i ec t attracting
special attention of the cytogsastioists* fba studies
of M^ino on nice (ig«la}» on dbld fish* and ra4i
{%94ih0 X9429m 19426« i943a) and on ungulates
(^943e, I9«4a# i94«b« 1944e) and the votk of ^ i on
axMMittitoes (I963a« i964». |96«>« |96S» 1966* 1967a
19e7b« 1968) contrihutad a great deal in tho fields of
genetics ana cytoganeties* Most of the results of these
studies did not show polyploidy or reaiarkalsle ehroffiosoraal
irregularitiea in the animals studied* Contrary to this*
as is well known* polyploidy and various kinds of unusual
phenomena concerning chronosoaes are r a t^ r conejon «aoog
the races and varietiea of plants.
Much work has tssen done on doissstic fowl in ths
fielda of eatoryology, physiology and genetic inveatigatlons*
The pioneer imrestigator i^o descritosd thr ehroaoso«»s
of the fowl for the f i r s t tiaw seens to be S^yes (i906)«
He deseribsd 6 ohroMOSoaes to occur in the primary oocyts*
o
flHRi SouMtilaroat (1908) dbssrvftd 8*19 efaoaommB In
th* prifiaJry oeeyt* fplloMiA by l4MS«Ulon (1910) Who
n^ortsd 12 chre<ao«oiMi« in p«rtlMiiiog»Mtlc •omatie
ouyvr (1909) th« iiemt tomadif th« eftronoce^ in
th« i|>«niMitog6nlal c « U » of tli« foul* H» dsecritea 1S-*19
chsrcMioa aBs in the ^^matogonial and 9 eheo!Bosoa»t
in the prisi«i;3r ^pcffnatoeyte* c l « i » i »g the aieeoveiry of en
unpaired eex—chioiiioeoae in the dotaeetle eocHi* Bowevefv
the mmltB of other workers ehowed ^ e t hoterowiitir
ttjcietea in the fenele end not in the male. Xn 1916 Gi^r
revised hie results by studying thcee races of fowl# the
I»engshiai# the Plymouth Rook end the Rhode Xslend Red ^sd
arrived at the conolusien that the ^permatogonial airono*
so^s vers Cc s pesed of 18 chrootosows «hieh include a
pair of large V shagged element considered as the amu—
ehronosoass* Culter (1918) also made observations on
gsmi cells of the Oold Caapine fowl and gave 18 • 20
ehronosomes as diploid and 8 • 10 elements as haiploid
niMbsr* All tiiese esrlier Anvestigstions dear ly
indicatsd that the t e ^ i n i ^ used was unsatisf aetory
for the precise study of ehronosows in the fowi* fhe
ChroMosoaes being clumped together did not allow accurate
stud^« The data presented by thess earlier mrlmt
sfvsn thouOh important historically* have becoias rather
outdeted as they show no oorrect picture of the chroMoeooits
either in mwiber or other aioipholoaical features*
IG
ttm •xtAiMlvtt studUs of H«)C<i (192$0 I924« 19ZU t92§h, I»2«o} maa* • sigiiif ie«at eontributicm in tli*
chrowaottcl tovestiafttioiui et fend* Hi« nock nunOm i t
e l « « r « i « t tb« diroiiocM eoaplAx of tlie fovl ooiMiUts
of • o«rt«iii nunlMir of maiemhcoiaosoiats hiciri&g « atfioitft
• h ^ ana • l « ra» tunftMiff of vary ainut* ar«8ul« l i to
iiiicroobcofiioMaiii9« Hqbc* (1984} m p o r M ao«34 cliroaoMiM*
in mm oomatie a i lK of bo^ rnxM but tfi* i «r9»«t
•hoped oie^aeot exlstftd in paired condition in the ib«|«
ceila «hiia i t was uapaioid in tiie famala oalla* ^ thia
•vidanca* aiecoJtding to hisw feoiaia h»tei:og«Mty tiaa
cytoiogieeaisr preyved. In hia aul»aequeiit work (i92ft)«
tia g m 40 a® the «a»iimn diploid nwiSsmx i^ieh eouid ba
observed in tha siat«$)h{MM of aoaiatifi oaila of chi^tan,
inveaUgatiooa of muar (1990) and YMaahina (i942«
1944) ahov tha bast rasuita that eouid ba i^taiaa with
te technique of direct fixation and aectian« tha anaiysia
of bird ehfoMosaa coitld not* howevar* oMfca forthar prograaa
uatUi tMw eytoiotfieal tKShniiitMia had baan wojdnd out*
MTooad I9i0 tlia aqaaah tai^iniqtja bagan to r«»laeo aaetioning
and a hypotonic pratraatmant of the emllm bafora fiitatioa
waa introdiiead* thia nada ooaaidarabia iayrovatnt ia
tho atudiaa of the dtgowoaawaa of irartabrataa* I»atar#
tba use of eolchlcina and othar chamieala waa aiao ooabioad
with tlia hypotoaie pratvaatoiaBt, Van irink and UabaXa
[J
(i9S6} uMa th« hypotoiv>«quMh titchiiiquft on wteryonic
tissuuB of flwy fouoia th« ehxoMoaoMi
Bunb«r to b* ilseut 18 cod Hi* •truet<ar« of
««¥«ii clixo«o«oa» pairc* dMioRsting tli* a—
chroatios€»ai p^ir nutdbcur S« findings «r* in
with the obMrvation* of Ci938) «Bd
V«iiMhifi« (1944)* Thfty did not succeed in deciding tflMith«r
tti« f«tn«I« SO or SK • V«n Brink (1959) used th«
stfM tttchniqa* ill* inwstigatttd ^ailtig miop«»
|>tt«fua q^duiatus and Pwcy ^jyaeatipiWa «itt k«ryotyp« of
Oallmt q j a i y was tlie swse le«ryotyptt of Qallua
doia6«ticu>> In M»Ioogi»t«cu« yiduiatiM tlie diploid
muflber wtf iPbout se of ia pairs wer« pointed out «0
mtcrechroaosotifits* Chre^sotM ousiber f iv « was tlia Z^
ehroaoaoiM. Pmmwer domi»«tieiif bad «9aout 76 ehfOfii0t0iD»s
of whii^ 12 pairs mm amimctaemm»omm and the 2r
thrmow&m was the 4tli iargtst* Zn Bona of ^ e ap«ci«s
eotad thay datact a ehronesona*
Vha larga mimbar of chronosonas and thaijr variation
ia aiaa in birds has ascassitatad thair balng usuaUy
clasaifiad as iiaero-and aiero-^roaoaoawa* Oha aiaa of
tha naeroehr iiaeaoispa is about and th^ir atriaetura
ia eoi parati<vaXy *asy for idantificatico* Tha miesoehro*
iBoaoMsa ara saall* 0*3 to t ^ snd thair eantronsras
ara usually inwiaibia* Moat of than taaing probably
c
ti0loe«titrle« Pr«f<fiou« imrastigat^a h«v« usually found
m dAfinit* dividing l in» botuMn saero-wd aiiore<*ehroao«-*
0069 in OalXif doawticu^ ^tme p« i r nm^r 7» M
Of thtt chroaoAonws hiive been clftsaifled «8 mievod)-
ronaoKKses (y«m«ihia«« 1944)•
mvcomx and Brant C1954) «Bd (19S7«19S9)
atattt that the difCacanea in aiaegt>- tfid mierochsoioosaMia
was Qualitativa* According to thais In Oalluf doaaatlc^f
tha nmber of aaoxoehfonoaoiwa la 12 in nalaa and 11 in
famalea and the rest of the cAicoiaoaoaiaa* eallad as diro-
iaoaom£>idt» oould laa a aort of raaarve nuelaio acid,
Katfcotaar waft unaible to f ix a dafinita nundaer of chrosio-
803iaa becauise the araail chrcKaoaomea of tern atrongly
hetarochromatic* Ooimally and I wcomfer Cl963>* with tlia
halp of autoradiography* found that the ainall chromoaooaa
aynt^aisse cna aarliar than tha larga Of^a* fhia alao
indicatad a qualitative diffaranra batwaan naorochxoaoaonaa
and ehroiaoaooioida* Utwconer i » hia aq^ort c i ^ a Hutt
(1949) who raswrtad only 6 eoiipliiig grot^a in Qallua
tor a pariod of about 10 y^ara upto 1965 aaveral
imraatigatora hava aicaminad bird Icasyotypaa to find out
whathar tha aax«-dataniining aKChaniaai ia SH SZ or ac/ZX
aad i^thar tha aieroehronoaotiaa ara raal chronoaoaaa or
only raaarva SNA* Van Brink and Ubbala (19S4) and Van
f ! i
SKiidi U9S9} fooiia that tlie Bl£ro>«hiro!ti09oaittfli bi^ft!V«d
a lit* ^diAary chronbsoMS* fhey wiNt4i« howmr* ao amall
th*t i t vfts difficult to f ix thair ttiuct niodbmr and to
diffarcntiata thaen fcom crhrooatida* ttm imrestigationa
of Ohno <i96t) during his atudiaa of mitoaia afid taaioaia
in Gallua domaa^cua^ ahowed that tha mieroehiomoaoioea
are coaatant in atisiibar* thay can ba followed throu^
cooiplatja eycl«e of mitoaia oad miomiM end that their
bahairiour doea not divarga taueh from tha raaiccoehironioaoniea*
tcriahan (1964) found during fSparmatogKnaais in males of
BBaieaoria aallopaw that the madsmv of bivelents during
the f i r s t matfitphaso was 38«4i and there ^ ao difference
lietween macro d mierochromoaoaiea* with the help of
l i ^ t eiiczoKcs^y end electron -nicroficc^y Ford and t4ioolh«pB
(1964) idiowed that the ao celled oaoro-end iQiecoc^co»>ao«
Biea only differed in aisee during mitoaia end maioaia
and Aot in hahavicmr*
lhou{(h the occurrence of « W «chronoaoiaaa in the
females waa long doubted* i t waa in 1943 that Schreid
diacoverad a aingle chromoeooi beaidea the S •chroaoaane
in the haterogwietio aeM in SnUJML dowsaticue^ i t vae
one of the mierochfonoaoaea that alao ahoMd a late and
very atrong iaoorporation of tritiated thymidina and
thia ehronoaoM he Maiked aa the
o (J
aem attappta h«v« also bmm m^OB to stuiS^ thm
ganfttic and cytologlcaX tmp^etB of hyteriaiaation in bixtSe*
SolfiOiOM^ala an axtanaive aeeouitt of tha a ac/-
XisHcad Qharactara in tha intarganaric hyturidisation togathar
with tha r«aulta of ^ a karyoIoglctaX invtatigationa of
tha hybrid bat««#n Muacovy duek (Caifiea wpachata) and the
dooaatic dacAt Iftnaa f>^atyghyfie^)> Cxwir and Koller(i936)
tsada a cytological atudy on the parantal form and their
atarila lqrt>rida* Y«Baahina (1942) ggve a hrief account
of tha chrotBCMOiiiea o£ the Muacovy duokc tha Oooaatic duck
and thair hybrid to daacrifao the huryot^a of these fonaa
and at tha a«aa tioe oorractad tha erroneous data offered
by previous inwstigatora on these fosns* CUsan (i960)
snorted succaaaful intarfanilial hybridisation of Dark
Cornish chikans and Baltsvilla snail i^ita turkeys* Ihesa
h^arida eoaibined in a single phanoi^pe* the dark plwage
and feathered neck of the l>ark Comieh chiokan* with
tha long neak, tAiita Skin and long tail feathara of tha
BaltsvUla amaXl «Aita tuil^* fheir rate of gcowth end
body weight f a l l betueen the nomala for their parantal
S!peeiaa» Although aavaral nalea lived to an age aquivalent
to maturity far either of the parental speeiaa# none pxo^
duoac any saaan or di^layed any aaxual baharioiur* Poole
(%M} atttdiad with tha help of squaah technique froa tha
tissuaa of th« taataa of hybrid asiBiKyo or from the bases
IG
of dinralcpiiig piofirathMr* fcom ptmt hybrid indi-
viau«ls betwMn MtmrU l * •mall. tmtkmy end
tutxfk Comiah ehiokaii* For ecu^arlslon of ttw chcoao*
8onfts ct tha hylirid vitli ttitltMr pavmt h« « l «e »tadi«d
th«» chcomoacwtta froW th« pinfcather of tha maitta of Stazit
Corniah ehicloan and saltarUia maXl ^itm turkay* Ha
c^aaived i4«»15 naecochfoaoaoaea in e^iiority of tha hybrid
ealla« the ran^a toaing 6 to 20* I t was diffieult for him
to diatiaguiah toetwean the gmuoma of tha parental «;>ael«9
baeauaa of the lack of Idantifyino eharactariatica in
the fiiaesroeh»^ao(aea« ihe miexoehsoaoaofaea «ore ifidlatio-
gaiatiobla frofB thoaa found in aithar of the parental iQ^aoiea*
Quita extensive nork haa been an the banding
pattern of inaeeta and mamnala (Daviaaon and flodariek^
m3f Hagala, I977a# I9771v i W i Holmquiat^ Luciani
mu 19751 Motara and Rai* 1977« I97»t PiB|>itialli at«al«*
i976| stainigar and Mukhariee* 1975» and Stoac* 197$).
In Bom apaciea of btrda mttmrnptm h«rm bean i^da for tha
atudy of interchange analyaia and aax-identification, f«ang
and Shoffnar <1974) used tha trypain banding technique
(Seabri^t* 197ii Outrillanx at, al*« I97i| ir«ng and
Fadoioff* 1972) for identifying a^eeifie chrcMoaonea and
cheomBom ama in chicken, they have deaeribad tha
devalcpment and application of the trypain 0 « banding
ia
tichiiiau« for idtt&tifyiiig individuai chxoiaosonft mgmnt*
involved in «hmBpaom«l c««crftiigtiwnt« in aoawstie
mivlicli al.* (1973) wea^log vith hmm
emm, foaea th«t l«noor tcypsio tf«fttn«iit progc«i#8iv«Iy
ivmi Q-tmioam to C « b«fid«» Stitplio» m^
lifttftrochirnKtle stalaiag swthoa of and
lf»ii (It?I) and Inscribed thm tM»«loc})r!^aatie aatuire of th* A
w «• 8Mi-ch2oaio«o(»e9 in ttMiy furthor fouaS tiiat tho
C • bgndteQ prodttee^ tiio tsy^sia «I.m> |i«od}ao«s
e h i^ ly tet/teoetiiomatio w « chxemMusm hm poaeiblo
^Ilicatlofi In aax-MiitiflcatioQ of thirds*
ftie C « lumd poaitiv® hataroehrcmtlii has liitlMir«o
beta atitdla4 only in Xliaitad ounlaiaira avian apeeiaa
via* dliidkaiit aova« pigaaci« parakaat« phaaaant^
<|uaii« Itoal an^ t»ald aac^a (sti^hoa ana
CooiCMia mad i$attoceia# m a i Stoeic at* al.* m4 i
All at* S97S}* Xt tm» baan i oaac rad Hiatt
(a) fHa M ia tha cmJly hatacocbioaatie <amm9om in tha
panileMt* phaaaant* auck and quail* In tHa dovo*
pioaon* JapMaa i|uail« l»ald aa9l« cHtekaA* tha
eaatfoaavaa of aavarn^ oiieirechxottoaoiMa are alao
c tNuid poaitiva*
(b) flia cantroraaraa €»f ilt* iiamclifoiteaonaa atw aithar
davoid of tha C wbaisd or poaaaaa it only in ainuta
^antitlaa*
Based on th^ss and thttir own obsezvatioiis on
t t u v tm in ehtelwn, Sttphoa and Anriglil (1971) htpm
»uggftst»a tliAt th« lifttBioclixoaiatiii Ifi birds is csstrictsd
to tStim w «cli«oiiiosotiis and micx m^Kwmmmrnt Also bccaus*
of thb ul»i<;uity of tli« aiemhcoiiosoaifts mC thm m ^eheoio*
soM in birds snd smtos* ^vy hsve enrisagsd m idsatlesi
situation in tbe tsio ^sgtmimm so distinetlsr different.
St, si* (1978} studied tits tietsrogsnsitsr in distrib>
ution of Constitutive tietsrochsoiaatin in four of
birds* natwiir pigeon (Coim^ba yiv|a)# dueSi (Msttap^ ooraman '
dsii&aus)^ oyna sonty. ) ^ iun^o b n ^ s r
C|&irao|ldsy stroattis)* in the pi^son^ C • bands
wers ioeaiissd raainly on ths foaeroeliroiaoso^ass i^ils in
plgMm the niieroehfoaiosomss mm prsdi^insntiy c * bsnd
positive^ In the duek listmehcomatin was restrictsd
to tha eentrooires of oniy thrsa pairs of mmsoBhK)matsm»$
bat in fiiyna i^i tlia Aaerochfotaosonss and a few »icroehxo«*
'VDSOass wrte C wband positiva* Zn tba JtaigEla b^fablar*
irraitpaetiv^ ef 8isa« almost ai l tha chroaososies mm
•aAmad with crantroaisrie hai^roohfomatin* Ihs w •chcoiBosoM A
was antiraiy hataroehronatie in al l the spsoies studisd*
yelikm mtmtXm whsra alnost ali csntroMsres are C • b«Eid
pesitiva* in birds thara is an obrious hatacogsnaity in
tha distribution of hetai»chxonatin«
IG
mk mplication pattenw o£ the •#x-chronotoaii« of
the pigeon <Caiuga>e l ivie dpaeetlce) h«ve bean s tudietf by
Oalton Bre^xy The lete ra^iicatioa patliXKCxi
of the pigeon ehxonaoeoniee wee etodietf by aeane of the I
technique of ten&inel t teke to tritieted tk^midioe ena \
eatoTi^aiogrflphy* the w •• chxronosi^ Ic the femalo coopietefi
r^lioation efter the other neersochroiaoeomes. Site
ehcoiaoeo3» in the femeie and both ss-chromoaonae in the siale
tarainated mn ayntheeia at the aane tlioeaa the cemaining
iaaegoch«omo8omea« ^ miecoeticoitioaoiaea mrm later c^Xioatlng
than ^ e ameechtomBtmm* Zt is inferred '^at iate r ^ i i e *
etion« at least in the* eaee of W •ehroiaoaoms* aignifiea
genetie inaotivaticsi in aomatio celle» The abaence of
asynchronoua raplieation of the s-chrofaoaocaea in thb homog*
ifiietlc cex auggeata that a ^ ^terminatitai in tl)e pigeon
laic^t d^^nd on a gene doaege sMMchaniam aimilar to tliat
reg^onaible for t ^ "autoaoxing** phenomenna*
the recent at^lea on aivian chroaoaoniea# though
uaing modern cytologieal techniquea* are haf^xed by certain
intrinsic inaccuraieitta* Theae centre around the validity
of auteitting coopleta cut out karyo^l^a when the enact
chroaneoMa nuntber of that apeeiea* far froa being certain*
ia only an ipproKlnatton alnee the ehromaom nmtmt ia
uaually a aodal velue in a fiequeney diatribution taible*
Moraovftft any complete photographic karyotype valuea
IG
h m to hm fitted to this rmOmr la mmy
tcchnieal difficultieti ar« involved in th« handling of
ntesechietaoaonis* Firstly ttw iikiesochfQiiO0<»ms air* quit*
likttXy to be by the largar •leaants* SacondLy
th* faint ataining of the anaiXer aie^aeota often malcaa
their detecticn extreaaiy diffic«a.t;# if not i{ii>oa8ibie*
^iffdiy* the disintegration of (siccochro-aoaonea into aagpiurate
co:i|>onent cStrot^tida my piroduce e faiae chroaioacae nuoDer*
rotirthiy* the pceaenee of artif«cta aueh aa atain ceaidue
in the ceila could erfoneoualy inflate the aiiexoc roaioaoQie
nwBiber* S«a»tly« there ia the difficulty of Obtaining
coo^letA tsiierochroaiosomal ooaiple-nent* meae factors in
eo^inatioo axe certain to introduce a large amount of
error %ihen an atten^t is neda to construct the so called
ceiq^lete karyotypes. Moreover* the arbitrary oeana of
claasifying the ^nicroehioiaoaotaaa on sise «nd danaity of
staining often yielda karyotypes which have chronoaocaea
in paira even i ^ n there ia l i tt le iq^arent hoMology
between then* a further prsble« with the eoi^plete karyotype
could be that there are« as Owen (IMS) auggested* chxoiBO-*
sonta which are beyond the reaolutien of the light sdoroseope*
A^sari and Kaul ( ifIS) atudied tranaloeation
tieterosygosity in the bird Imnfly^ t nun^ete <Ziinn) where /s
thB diploid chroaoeoiaa number waa found to be sa la one
i4
m«l« «tid on« fhcr« ar* 17 amecoctixo^sofaes in
«ith«r Mau Both th« inOlvlduAls ahoiMC m tfbbmnt
kOTotyp* in 1i«vla9 thr»« ui^aixvd «accoelicoaio«oiai« in
addition to acvta i>atix«d nmcioelisoaiot^MS. th®r®
«fi unp«ic*d li -typft autotew togithsr ^ th two «aall«r
naoroetixoQosoQies t ith their cttntronnres in the terminal
fegion* the possible natuiw of the trimelocation leading
the heteiosygiOBi^ ^ee suggested* Coi^arative Icaryologioei
studies ere nscessexy for understanding l^i^riaelples of
Icervot^^ evolution end for clarif icetion of phylogenetic
relati<mshl^ii» Etie cosqparative karyology of birds is
lagging behind similar studies of other eleedes of
vertebrates (ReptileSf nanraals). One of the major reasons
for this is lack of coiq;>arative data on groins of closely
related i|»ecies« Ihe cooperative studies on modem
tectinigtMis «sd ^nscedure have been eerried out by soma
workers (Jaiiiy* 1966$ Bloon« i9«9si Castroviejo et« al.#
i9e9f Ztnh et* al.r 1999$ JovancHTtc mA Atkins* i9ft9a«
i969bi ttanner, I970f »iecinni and Stella. 19701 R ^ i ^ i
et, al*« 19701 Bul«tova et, ai*# i971# Takagi et, al«»
19731 fMlatova and Fanov* 1973f iheodorescu* 1975f Ansari
and Kaul« i977a« 1977bi erased and Patnaik, S977i Kaul
attd ABsari, 1978)•
S#ebert <1976) etudied the soiiatic «hxo3K>soaies by
using short terq leueoeyts culture tMhnigups of
speeies of diurnal birds of prey belonging to four
IG
diffwniat of th« ordar Falconifoms* Ho ae«cjrlto*d
th« And UXuatratad k«ryoar«mi of sow ipcelfts*
MBCocaiiigiy «h»n the order i » cittMif ted into f «iitUt*«#
four karyologiCAliy diffcreat groi^a c«n be distinguished
in th« Faieonifc^naes*
( i ) CatHejrtit^t JC«ryotype» which ahov oniy 7 pairs
Of bianaed oacrochronioaomee aa^ a considarabS.* auitibar
of aaall aerocentrUrs end aiecochroroosooiaa i the
diploid sunaber being approximatiely 80)* ^ i a is the
e&Xy groMp is the order in «hieh reaily large coacro*
chro:!ios€»^s are found*
gaiconidaet ^ a karsol^}® of this groMp includes only
a single pair of bitfraed maeroehfoniDSomes* el l other
el«9ents being acrocentrics of nediuni to s ^ l sise
or microchrooiosoaies (diploid a<s«ibere of siiproxiQiately
64 and
Sftciittariid^i these are the secretary birds «ith
biemid aeorochroaosooies and 44 small acrocentrios
and mierochnxaosonss (2a * 60 «pro3ciiiiately)«
(4) ^ceipitrid^f t itie representatives of this group never
possess ^ r e then «lx»ut 8 reel aicxoehromoscmes while
^ i r karyotyp«« al ow varying aunbars of biamid
aafoeentric giaecochroriosoiies of sMall to medium sisa
(diploid auadaer raages from 40 to 76)• 9m possible
IG
rvistionihtp «rithlp •«ch of giCQap*
b«an briefly discusacd a morft mttmmivm
aiseuaaion im dmdicmtmd to tho possible rolatlonshjjp
betwestt these gro^ps# and betifeeii them and other evian
the variation in Icacyotypic structures found io
the falcoQiforaes is madh wider then that in other itipian
groups* MOMSver* i t reaains an UDaosnered questicn
ifhether this karyologieal heterogaiiiety points to a
polyphyletic origin of the diurnal birds of pre|r«
• l l y ' the chromosotte eoopleiaents of the Aeeipitridee are
taost vinootsBon among birds bmsause of their «xtretaely low
nun^r of real oiierochronaottows* ffoiimrer* of oil the
Faleonifomes only ^ karyotypes of the Cathgrtidae have
clear counter parts outside the order# since nearly
identical coqpleaents «»ere found in representatives of
the j^Qenioopterifomies and <3ruifo3Daes«
Pollock and Feehhelner (1978> studied tho chroeoo*
sooes in ^ermatogenic cells of 10 eoc3cerels (SsUttiE
by colehicine pretxeatnaent and air drying
technique* 9hey observed ^ e model nuadaer of naierochxo*
nosones or bivalents in of q;>erttatogoiiia« 9816 of
pritaary i^rmatocytes at diakinesia and 94* 3K of secondary
iq;>emiatocytes* They have estimated chiasmata between $9
iV
and i6 to oe«:ure in the blval«iita* of th«M m
«r«r«g« of at ar« •mb in thr six aaerobivaicnta*
Mr«ra9t nundbtra net* e«t* S«9* 3«#« and 3«S
for aiieffDl»itr«i«nts I thiotmii S and Hm xnapactiimly*
Tbm nurdbers of chia»nttt« dif f«x«d •Ignificcnt&y wtong
birds but ware not (Ufforontielly affected by ptett^mtm
i«nta of eolchieinft*
Anaari and Kaul (1979) je«portad the baaie
kacyot^ of coamon gf«en pigaon* yyagg^ otioanicoaf ra
having 7 pairs of oaecoctun aoaotiiea thirty paira
of ffiierocSicoaosoiaea* In the population oxaoiinad by
than* there oiciata an interaating cheoaosonal polyaor*
phiam* Chroioaotaaa on© and two both exlat in a dinosphio
foi»« Stm diiaoiphisgi in ^ach case i^peara to ba dti»
to the presence of a pericentric ioversion* ChronoeooiB
eaciata in and t^^ conditions and chroaoaoae
Yno in a*^ and ^ oonditiona* Ttioy have diaeuaaed
the aignificance of t^iia intr|i>opulation variation*
She pxeaent studies were undartaHen to .aake a
ooaparative analysia of the karyo^psa of aix speeiea
of birds fxon the aub-eiass CarSnatae# natntly yd[t§
Diaapuaes iiniliTI!t>t (Kastern puxple heron) and ju^dyoia
ara»ii qrav^ (Indian pond Imton) from tiie order
CieenifecBMe* iryiitgllgiff aondleerianua j^fi^moff^fiif
i
(any purtridgo) fron the ord«r OAllifomev, and Corv^
K>l>nd>ii» (Hous* cvow) —r dometticu* < Hous* iparrow)
brunie»D« Bunting) from the canter
i>^asMrifora»9* Umix diploid nuoter has toci«i wocksd out
in tMTms of the mean value of dis^loid counts «hich ia
diffef*nt fro^ the procadure of o M r wociiara who h«r«
given the highest pmtsk of tiie diploid count in various
laatjqphftM plates as the dj^ploid nxaSomr, a coaparetiva
analysis of different chxotiioso^9# their centcomsrlc
indiossf tnaao ralativG lonQths the two acnis has bssn
made with the hslp of taSsles imd karyo^oos*
ttm studies were undertaken to deteraiine t3sa
chrooosotac! number« laoiphoniatric «3alysis and the ^tent
of ka^ryotypic sinilarity to detemlne the phylogenatie
iwlationshlp of closely related bird species*
19
Six of birds ttaff»ly jur^f ointowemm
Mfni^mmi^ {Xasttni ^ y tMroii) MtOMlm qrittri^ ata»li
ila6im pond hsron) of th« order CieettifcMmnsi graneeli^tta
l^ondicTiiWff intMfaoaltug of tlwi ordtur GalllionMisf «9d
SSltSSSBE. eroii}» P ^mt {Homm
wpm^m} end iafegriga ^irwilcwof (itodh»«d»d Tomtit^) of
<Mrd»r 9«ii«orifo«iMMi «oloetftd foe tlio pivMut
otiidloa* birds noes |4«titiflod vith tho h»lp of
tbo K«nd Book of tlw Birds of Xiidis «Bd Pidtistso swrios*
( idi «id aipXof*
9m ohnonosoivi png^srsticos of f ivo pairs of ssdi
of tiis birds ii«ro msdo ^eeordiog to aioon (1969b} with
esrtsin modif iestieos to suit tiio soviroiMsiitsI ^nditions.
vtm nitotio eHroMosoMs mxm prspsrsd from ths bomwsrroir
eoiis* ths aost setivsiy proiifsrstiog coils of body
tlist r^rossot tiie bsst nitotie figuro for tim pmposs
of karyotipo sttady* flis siidss ii»ro prspsrsd stecordiag
to ths foUo«flJi9 prooodttro*
Ths birds wsro injootsd with 0*a «l/iOO 9bs of
body ifsifilht of O.OSK coidhieins soiation iatri^ritenisi ly
shout 90 aimitss prior to f ixstioo* «hsy mm thsn
ssfsrifiesd s»d ths log bono <Vibie»««rs«i} tfos ti^oo out*
Xn sMsilor birds ss in fpftssr dasssticuf C h o u s o spsrrow)
and Mssriss toruAiesps (Rsdhssdsd bontiag} both log sad
20
ISOfMItt iTiblo^tarsiM m^ Wmmat) mxm uatmn mt to
«tiffieiMi% Qttaii«itir <»ff IsofMUMpntiM for mtaSf* flMi
two e^eiMM wmiMt m^ of tit* horn mrm sJiitllitly cut
witti ^ liftl p of 9901S aciMors to f«eilit«to pon^atioa
of ^ ii»«dl««« noHmm mm m»am mt hoth on<te i^ith Hio
iMlp of iMMidio* «is4 tho tfomm&nttm vm flundiod out witik
tlw lid^ of • S ail, ofring* contstning aodUlam
eitvfttt ooJti&tioo lAiich h»a botn liipt tik
«r«t«<!l»«t}i taaiBt«ti)#d «it C» M aeoaios uMd witti ttM»
Ofclc^ wmm So* 2) ifid for S«rg»ir blra* and So* 26
for «o«iI«r bird*. obtaiiMia* «ftor fluttliifig
««r« onspoiiM proporly trith tlio Iwip of p%»tmw pspottm oisd lEis t for iaeolbotioii i» o i^twrnatti for tfbotit 30
atotatM «t Cm flio ooluticii tr«» tftoo contrifagtd «t
2400 for aiaiitso* IStm oui^iiotiHit wm <lioo«s<lid
and tho pollot vfts ro«iaqE»oiidia In mm tiamtivm C3fi#
•thyi ^colioX m^ oottle tola) r«frigftr«t»d
«wor n l ^ t to olJtsifi amdaun f lx*tieii* sotisfoetocy
iroatilt* Coo ol9t«iiMfl % placing nm suasioiisios for i ^ a t
tNo hmxtm i » ^ lofrigarat^* fho ooilo i0or« than
• ^ i n «Qd coiitrif«i9«d «t 3400 wpm for 10
ainatao ana ^m o^ponifttaiit «•• 4ioe«rM« flMi r*««iiiiiig
eoilo oattlod «t tho faottoi mm ouipoodMl in tho
f iacotiiro. this pxoooaa of eontrifugatioB ond daeantotioii
mpmmt0A thrieo with an intocvai of 4S aioatoo*
At tho Olid of thio pxmtmm* m minimm QUontity (2 aii}
fsc «$)( «e«tic ttOatd to t»Ni P « i i « t «nd tM
•u«p*fitioii tionogvAiMd with th« M p ef p«tttiu«r
it •iiMtU o£ th« eall •uapcnsiott « « •
tailwii out In tK« plp«t«» «tid two or ttiTM drops of tt)«
•oflpoiuilmi nero dropped from h e i ^ t of liboQt h^t m
mtmt ov«r el«tn»d« ps«^hui«du slidts tUtcd nt id
to o3»tAi» tftll i ^ a d ostuphsso figfmm* the oftll*
were dri«d pzopsvly by vwriag th« slidts In the air «»d
aiterttatimiiy pa««iti0 ov«r a l»iniar« driad alidaa
«iar« Itapt in a eoi«>iiao Sve coittaliiin? phocphata buffarad
(pH.7) Oiaawa atain aolution (0OH) for 7 to 10 oinutaa.
flia atataad a i i ^ a iiara wi^lied ifi dotiblo diatiiiad iratar«
driad and aouatad in tsmu
Rr^arad alidaa thua obtainad nara atudlad undar a
C.su Srginpal raaaarcti aiccoaeopa* tfatapiiaaa piateaa nara
eountad and na«siirad at a «agnif icaticn of 400 m 9saA m v
photegraptiad at a nagaificatioo of #40 x* fha total iengtha,
loag am iaogttia and ahort am langtiia In tHa 13 iar^aat
paira eC chroaoaonaa iaeluding tha aax-etupoaMMmaa in
igaia g m w i mntti i i ia •ao i i y ^ i tfiytt m y U ^ lo iar^aat paira of chronoaonaa ineludlng tha aax-ctirowoaowaa
i » fflBffrtlltfff Wit f Ipf l^am I f t W W l ^ i i iargaat paira of direifloaoMa ineliadlng tha aax-ehronoaooMa in ^oynp f^iandana. Paaaar dawaatieua and l alaariaft brunifapa
mrm mmturmd in id mil M i m a M mUfflHm— oiilJls
lK»th 9mmm d thm 9iM iqp«ei«« undwr ttutOym ltj«
Iftfigth ItiR) of indiyidiuOL ehromoaowKS
M p«sc«iit of tile total longfch of tlio
flMtt •NBiudiog ttio ehtomofomw vsallor than t2th pair*
iOth pair and tt-^th pair of tha thea neoupa raantionad
t^xfm xaospaetivaXy but inciotfing th« aax-ehroeioaotiiaa*
Fiaaliy tmm calatlva langtha of variotia chroaoao -aea
vara caleitlat^ raj^jeaatntad with atandara arxor
ana variation ranga* I1M19 trai and abortaxm iangtha
of a <ihirono«oM» wara oaleulatad on tha aaw baaia*
ftia e«atfoaiaric indaic vm oalcuiatsd as cant total
iangth of tha ehroaoaooa and rapiaaanted ai mhfi
eantrotaaric indait aiong with variation r«sga and
standard arfor.
Sha chromMOMa mem e l^a i f iad on the basis
of thair eantronarie poaition weeQcding to tha claMi^
fieation aada hjf tmwn mu al** (m4>« So thair classic
fieation iwran at* (i9«4) hgva daaignatad t»ia H -
«hraiioae«a having ita cantrooara in tha aiddia« tha m *
ehranosona having ita eantronsra in tha Aiddia ^
Madiw ragion tha a»»ehre«oaoflis having ita cantrotaura
in tha atilNaiadien egien, tha t»^oMoaoaa having ita
eantroiaara in Hia isxiainal ragien «Dd tha fi«chroiioaeM
having ita eantroanra at tha tatiainai nnd*
lo
fh» photinlecogri^tui of cnftlyMd ctiroaaoaooMi
spreftd n m tailH» « t • fHftgnlf tcfttion of «40 x with th*
htip Qif phote«&aroQr«>iite «ttiicliii»nt and
•itiLarotd to « tftundwra cut out and arraiMitd into
liaryotypcs witli th« «ia of orlgioftl MliOm* And int«ct
photogrit^tis*
'Sim opioid chromoeoaie mxa&mr of ««ch spAci^s of
t ^ birds KM counted in 29 tMili MtapliM pi«tit«
and in tftraw of smmk nmasrnxm
^ mtig&ntA U8«d in the prt|>ar«tiaa of tlio
cytologicai siided wmem pripttr«d « » foUonst
U 0,PS e«Ht Coichi«ia» •olutiont SO «9 of Coiehicin*
powter on m mlmtrlcnX balance and dissolved
in iOO oil of double di&tULled wat«r&wiis stsorsd in «
ffttfri^stor*
His lupotcmiic solution was prspaffsd by noi^i^ing
900 lag of sodium eitrats (M^RgO^) crystals on sA
•Isetrieai balancs and dissoivsd in 100 oA ef dsubls
diistiUsd %rat»r« this solution was storsd in isfrigsrator
than traa«f«frad to a vatsr bath at C m bour bsfor*
3. f M M f U l l fixatiira was pcapafad
by sixing aS ail of giaeiaX aostie acid ( Anala* R) with
24
7$ lyf i BMutm •thyi («DHU
if P^ m^* P** i^^tlc
nadir by mixtog i S nai of Olvclia lictttic vltli
SS 4id of eovSblm ir«tar«
fcifmp^ mv^^m* M^^tm^pn 7*0 Q$Mmm vtiiin wm miSm witSi ^m of
(U h y ^ g m !iiottophoflpli«te (il« audi j oAium hyarogen
phg^h^ imrn^ H i^^l* Wixmt th^ tsaff^r «r«» by
di«0oiirifig ^ of hydKO^n m&eGphoai?ti&t» mnA di
tt*4i gps of jaoMm hydH gim la 296 of dooibilo
imtsis' MpmtAJty m^ pl&md iti the r«f rigerator
«ft«r checkiag th# tiftl^ of iifl f^
tho iHif fcxva 7*0 (il«Mi« aoltitiofi* 1*6 isl cU ftodttm hf^xogm isciii«s^oi li«t« solutioe «f>d 3*2 a l of
liy«l >gifi pteis^ftt* aoiutloii «l(8i9 wi-tli 2 lal of f Utwcoil
stftlit «oIutt<ii m m tAm^ma togethtr 44 bA
of ^ a t U M
Mia Chmi i i a of tho tti« mico
f i r s t piecod in « hi Oaat t^ontiiifiiii^ «eia (Moneont*
•ulpliiarie «eid m^ fow crystals of potsssiun aichso*
««t«> for St iosst 24 hours* Tim s l idss mrm thsa ifsshsd
with dstwrg^t so«» sad piscsd in coi«?liiig J sis* ihsss Jsrs
tfsro plscsd usdsr cuaning tap wstsr for alboiit six hours*
Aftor wsvhingt tlis J«rs «f«rs ccmrm^ sad plsesd i a «
rsfrigsffstor for shout JO «iiMtss hsfor* using ihsai.
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47
stMi duPOiiosoflMNi Of Six t^m^imm of thm suWlftss
C«riiia%ii»# i y ^ t oiigaug»> f r n purol*
hvron) anS qraarii y y i i . CX»di«a pond bMDoal f
thm oratr C leon i f f f f t t f tC f f i l f tWi Bfffl^cfrlgifffff i m s m r Jl^ imff |>«rtrid9») froa th« offdti; «iid
ffbrvu* i^lmatng (tmm ertmU Pmmmme dommtlctx^ (ttouM
iS ttriofr ) Kaiwigtea tHmting)
fifota the ordwr studiftd* f««uXt»
of tli« 8 ttiay and 4»«erlptiosi of ««ct) ipmsimm gimn Momt
?tte totnl nuaSder of chro!ao«o>ae9 ia S5 mi l apvminS
mtephastt piatos of tho jyaff augpogta aaiiilotuiii
puvpio horoti} varied fron S4 66 with « eisrlcoa pm k
i4»%) mt 62 (TMm-'Xh a0«ti €or tiiia variation was
caXouiatad «a ^ So tha diploid count of
c^soRoaowaa io thia apoeiea may givao as Ai ^
inatoad of id ^ a* pxopoaod by othar workara (Vtk^gl
and f^iiio# i m i Bloott, iM9a« Ifi9l»i Ztolt «t«
King* i^Oi Piecinni and ataUa* i970i Huiatova «id
9mam0 1912$ Anaari «id Xaul, 1977a# I 9 m i Kaui and
Muiari* Aitaari and Ka»i* i m ) *
Ihara ara 12 paira of larga chrotaoaoawa* tha
aiaetochroaeaema inoluding the aax«eiiraioaowa# idhich
eaa bo oasily diatingulahad tKom tha anailor ehroMoaoaatf*
4C
tlw tttocoi^oiiOMM** (ViQUir* Thm mictoehjtoeiomfmm
wm Moccntrle i t ) mA vary in ship* froa an«U xed-
•h^ttd to aiaut* f t* eMecQehuoaoaanrs hnvtt
h f n clMBlMimd into five groups in ordsr
of length and Mntrontvic poaitlona (rigour** % and ) )•
Qppttp this grou^ isoltidtes tfin> £>«ir« of lacgs
wmm h&ifing timix emktxomfm in the ia«diiao rsgioo im)
CFigum I md 3). mm f i r s t p«ir i« «h« iargtst having
th« pmg c«nt r«iati<v» length varying £roa 29*178 to 2#«193«
tii« amm r«i«tiv# l«agth boiag 29«se3;| 0*l&a» itiit* th»
ceiitroiMric indsae varisd iron 4a»i€6 to 4S»00Q* tht tmtA
e«ntroMri£ indax 0*449 (Tabl* zs)* IRio sseond
ehron&soaw p«ir tsf grovip tihich is saslisr in isngtii
thsn tlie f i r s t pair l i t i*4004)# irsrisd in rsistivs isngfeh
iron 16.176 to i7*41t« th« M»«n rslative Isngth being
l«»783 • 0*30fi* esntrpjwrs in this csm siso iM in
tlis tm4lm rsgion* ths csntioMric inAtac vsrisd fros
9 « * m to 40*000 sndjuis wsn esatrossric indim vs« found
to tas 39.S90 ^ 0«2S3 ( t M * 2X}.
S^ggeJI^i A singis pair of ehro«os«mes whic^ is third
largest ceasti^itss this groop (rigiirss 1 and 3)* This
CEhrooosoMi has i ts oantronars iseaiised in thi sute'ldian
ragioR ( m ) * tlw p«r cant ralativa isngth of this pair
variad batnaen 10*903 • Ii«909* tlis mm being 11*390 ^
49
0«2tf* Coii mDtd to thm f i rst pair of cfuroaioM^ in
i«»0«h i t i « ftUiost half of tha th* ratio boMaii
tha two toaintf 2«0«9t i*00« Sha a««ii eaatroMric ioaax
ia 3i«0$4 ^ O . m , varying tetnaan ^.571 and 34.39S
(Tritda tlh
9IB$BJE11» eo!!|>riaaa tliraa pairs of ehrooioaiiafta
which ara f iftSi a i x ^ and aavanth in isngtli* tha a^hnstnoai^
not inoludad in thia groi^ b»ing the 4tli largest paiv«
All tbe thra* pairs have thair eaatronsras in tha ta»dian
ragion and are ^ r a f o r a , n « c iromoaoifiea CFiguraa 1 and 3}«
Thmy hm* thair a s a n rslativa l«igths a s t.iOii • 0*170*
€•227 • 0*034 and S*9i3 • 0«iiS and irarr in £>ar cant ralative
langths batMsan 6*028 « 7*470* 0*129 • 0*312 and S.007 * 0*312
raspectiwly* fhair lasan eantroiBarlet indieaa i i a r a fmmd
to faa 42*738 • l.UO* 39*000 • 1*200 and 44*421 • 2*039
raspaetiiraly* All tha thraa pairs ara alnoat ona third in
langth ^an tha f i r s t pair of ehromosoasa* the ratioa
baing lt3*307« It 3*774 and ii3*974 raipaotivaly,
Qtoito lyt Five siaall tsloeantric ehxtiffloaoMi paira aaika
thia group* lhaas ohroaoaotnas e«n ba idantifiad by having
thair eantcoiaaraa at tha taminal snd (figuraa 1 and 3)*
fha ralativa par cant langth of tha f i r s t pair oC this
group variaa froM 0*101 to S*31S« tha Mian baing 5*273 •
0*020* thm Man ralatiira langth of tha aacond pair is
d9
4*702 0.07>» v«ryln9 hmtmva fbft third
vari«d la Ats r«l«titre length lj»t«««n 3«870 • 4*304
«ad ttw amm for pair found to b» 4«06S • 0*072*
Th» foortli pair of thia g m ^ iiaa^M. «aa» n»lativ« iangth
« • 9«$|$ ^ 0*049 «hi«it v^iad bat«a«ii 3*322 • 3*738* Hia
last pair of the t^ich ia iiiao the anelleat naerochr^
aaomom liaa ita f&ean ealatiw iangtt) 3*i7S • 0*0SS* varying
batnaaa 2*990 • 3*3li* All the five chro^aoaoifta of thia
group are atnaller in length than tha iargaat f irat paiir
in ratioa ii4*4S7i i ; 4*998f it 9*76tt i» 6,6|| and ii7.402
rai >ae tiveXy*
aco g> yt Sex- fhroffloaooco ecfi^iad this groiip* 2» biraa«
aa ia known* the maiaa are homogmtic with chromoaoaea
while the fmatAm, having fm eonatitatioo* are heterogtfaetio
(Vigurea I and 3)* According to I «ngth,M -dhKotsemom
in owiLaa ia fourth iargaat chroooaoma* Since the eentroMure
ia in the aub aiadian region* theea are daaignatad aa *aa^
chroMoaomea* the lengtik of t^ae ehroa aoof a varied
tnm 8*044 » 8*823 in iO «et«^haaa platea and l^e mm
relative iength wa« found to ha 8*438 • 0*143* fha
Mtttronaric naaaureaeRta revealad the mmn centronaric
index aa 39*942 ^ i«34« wit^ a variation range from
32«000to 34*000* Zn fettaXaa the co^plaaant haa a pair
of 8M chroHoaoiMia* fhm 2>ehrow>aona ia of the a«ia aiaa
and type aa in aialaa* fha w-chroiaoaoiaB ia aaallar and
ia i/l«43 of the s «ehroraoaona In length nhioh
ia alitoat equal to the 8th pair of chronoioiiiaa inc luM
in 9«oup XV* i t cm b« id»ii%if i«d in
htpring m Mt«i«lXy lm%tm6 ematKommem* It9 c«R%roiMirie
i n te v«ri«d ixem 37«SOO to 41* i7S ^iieih gw m relm of S9*0ii • 0*7iS» pmr cm^ miativtt length
ir4Mri«d fco» 4*90t to and the aift«ia tvi^tiw itt i i^
emm oat to hm S»20# ^ 0«06i« Xt is « cliiomosoiei*
Hsftidtto It pairs ot tUsfmMmm ^mtm
9 i ^ge e£ 0a«ii«r c)irotao«o»»» vary
fcois ttiflut* to mti i sod>»ilii|)«d strutctur«» coil«a
miKrotShtoao»oim»» In this upoeiot M «ibout 18 In
long «rffl flUd •tiort «ffai stadioa of the Oioo«03ies of this i^ocies «iso flisae md hai e bma reported in T«hi« IXXt and figiiM 3*
HTx jyasE .gggsasiif
A stu^ of the acetic i^iro«osoa»s fros 3$ nsll
diiineated wit«>hm plmtmm fron tlM bonMiarrov pftparations
xmmtXm^ that tlia dl|>ioid ehroaiosonaa naiMr irariad CitMi
S4 M vith Adrkad pai^ CS391) at 9ha diploid niMbar
if aiQiraaaad in tafma of tfea »»«ei caii iw t i ^n as §1*60 •
0.4M is this i ^ i a s *
#gdiaaa aravi^ qyavii is aiso a Msibar of tha eiidar CleonifoCMss* ilia total nxubmt of maosochromoHaas idsntifiad ia this « ^ i a a ava ia ineluding ttm ssx - aturoMosonts. thaaa
hmm imm aif fK«iitl«t*d into « on «h« «f
thsijr iB»«Ei Mlativ* i#ogtli and e«ii^roai»rie indices
(ti^ur^ 4),
OBom this gr«Hjp cieiit«i«« titiai* ^ i r of
clurocKMiOim hwing Vtmix mntiemmm In te atiaai* ansionav
tin9 til* pair •§ lf>»ty|>«« Varying froo 48.S7I to SO.OOO
^ mean ettiitiroiii»rto indtn of cturoacMioMS 49*447^
0»3«$« par e«ii« varied tw&m 23«3$S to
M mm imimg 4 (Tame W)*
j8Ee9B-.22.* ^ sii3i92.« pmiM oi ehr«»(!io«Qcae9 conatitutRs this
ihia chr&aososie is ^ s«co&d largest in I«n«tli«
f»lativ« laaoth v«ri«« l»«tw«efi l#»00 to giving tbtt oMittEt tm 14* ZM 4 0«0SS« Sli0 mm eantroMric iadtji
found to bift ta*944 • 0.4S4 %rith m variation iron
IO*9Sa to (Tabia Baaad on tha claaaifioatioa
af W at* ai* Ci»44)* Thm pair ia ai»»t»»a# iianrino
tha eaiitroaactt loaaliaad ia tlia aWb-Hsadian ragioo
Cfiguraa 4 «id « ) •
OfttMP Thia groMp aoataiaa thiaa ahroaoaoiaa paira
ara thi^a, f i f th aad ai«th ia laagth, s-chroiM>ma
pmju^ tha 4th latgaat ia Mt iaeludadj^ia groMp. jOl tha
thcaa pairs hava thair aantraaaraa in tha aadiaa ragioa
ara tharafora i»*typ* (tigoraa 4 aad lha f i rat
^air of thia groiip haa a >a«aii ralativa iaagth $U30 • 0«a4i
which variaa hatwaaa i0*344 • i i» #27. Varying hatwaaa
Jt)
40.000 - 44*117* the meen centromeric index was found
to be 42*537 • 0.679. ISie second pair varied in per cent
rel&tive length from 7.523 to 8*637« the mean £or which
was calculated to be 8«126 • 0.197, Ihe mean centromeric
index was 43.226 if 0.961 whicii varied from 40.000 to 45.833.
The last pair o£ this group has its nacan relative length
as 6.298 4 0.102 with a variation r«nge from 6.000 to 6.644.
nie centromeric index was found to vary from 35*000 to
40.000, giving the mean as 37.145 ^ 1*012 <Table V) .
Group Wiis groiqo again has a single pair of chromosome
which have their centromeres in the sub-median region
and aire therefore of sm-type (Figures 4 and 6). The mean
relative length was found to be 5.808 ^ 0.105 which varied
from 5.462 to 5.980. The mean for centromeric index was
34.427 4 0.713, Varying between 33.333 - 36.842. Ihis is
the 7th largest pair according to length (Table SV).
Group Vt Five jsmall chromoson® pairs having their centro-
meres at the terminal end (T-chromosotnes) conprise this
group (Figures 4 and 6). They ha'/e their mean relative
lengths in the descending order as 5.550 0.122« 4.936 •
0.188, 3.894 4 0.077, 3.311 ^ 0.050 and 1.116 ^ 0.060* She
slight varjfttions in relative lengths are shown in Table IV
They occypy 8th, 9th, 10th, 11th and 12th positions
respectively according to their length*
•' 1
VXt fhft sixth lest group ioeluass the ssx*
chiroc^oaits which are s i in asles snd m in fsomlss* Ihft
gKrhroaioacms is the fourth SLorgsst i»»typ* (Figures 4 and 6)
The mm r«X«tiv« ienoth is 9*S4e • 0«ia0 varying bettrnmn
9*302 • the cantroineric inisx varied fro'a 49*333
to 46»428» ths oiiaii for which was caleulatsd to h« 44«3S04
0«6t6* M W *ehJPDQioso«i in this spseies was idantified
a» a sisail chroniosoiie is altaost aqoai in
length to tha 9th pmlr of tha pravious group* The lasaR
relative iaagth was found to faa 4*930 • 0*066 with a
variaticm range from 4*666 to S*01S XV)*
Tabla V iliuatrates tha par cent ralativa lengths
and iqe an values of two arn« ^ the chrc acMOtnea in
firdaolf ffuvfi gravii (Indian pond heron)* Kt^cyo^m
showing thr per cent relative lengths of the two arms has
bean given in figure
rhi» species belongs to the ordttr Galliforass* Zn
ordsr to study the diploid chnynoso^s* 35 good itat«phfs«
plates were counted* She chroaosoiie nuaber in the speeies
varied frosi 60 to eo with a swxlEed pei^ ($3%) at 70* The
flMan of th« an chronoaomi nustoar was ealculatad to be
76*24 • 0*622 as against 76 ^ reported in yrancolinus
dd
chroaosoona on thu b«9l» of thoir and
poaitioti of the ccntrootras h«ve b««o aividcd into 6
lAieti eonstitutft 10 paire of aaenchroiaoaows incluaino tha ««x-chxowaorae8* r«aiainiQg are th« tttecoehfoaiosoima*
Thb maecoehromosoaies tiare b»«n grovs^a as underi
^gotip.Jt tim largest i l f t pair of the caaplBmtnt conatW
tut€a thia QtOMp* ^ e contronieirs is ioealisad In th«
sub«aaat«n rttgion and ^ ehcoaoaonse is aesignatad
<Figur«8 7 and 9)« The par cant ralative length varliedi
betwe«i 20*454 - ZUQO^* tha a&an baing 20*963 • 0*247*
The ceatroiaarie Index gS this pair varied from 2S*S7| to
32*000 giving the oiean value as 39*89i 4 0*600 (ITable VX)*
Gtoup Ii;i A single pair of chro^aoaaa* the second largest
foxiBS this group* 1!he centrosoftre is in the median region*
the sia«s centroraarlc index being 37*017 ^ 0*309 %^ich
varied fre!!i 34*000 to 37*900* 3he per cent relative length
of the lAiroaieiCBie varied f m l4*S4i • 14*949 with the
awan Iwing 14*101 ^0*462 (Table vz)*
group i m thia groi^ again has a single pair of etirenosoiies
with the cantronere placed at the ter«in«l end (Viguree 7
and 9)* Zt ia the third liirgi*st pair in the oooplement
having the laean relaUve length as 12*704 ^ 0* 142 witii •
r a : ^ f ras 13*37l to 13*1B1 (table vx)*
o d\i
Group XTt this cproup also contains « sias^* st»typ«
cihxommm9 thm m»An ccntcoMrio indMc tm 21*254 ^
0*618 i^leh from 20*000 to 23*857*
the variation in per cant ralatiire ianoth n as
fsoa 10*309 to I0*909« the man baing 10*62S • 0*136
iTMm 71}* ffiis is thr f iftti laroaat pair* the ^hg&som
aomaa laeing tho fo««rtli lergeat*
SSSSLSL* yi^tti ineltiaes thcea sniaii ohcoaoaons
baring their centrotqertta at th^ tartainal «na
Sha par cant ralativa length of the f i rat pair of thia
grotip vari€ta batman • 7*727 giving the mm valua
OS 7* 163 ^ 0* 167* The mm rsiativa langth of the aaeood
pair was found to ba 6*283 • 0*099 varied from S*932
to 6*829* the iaat pair of the groi9 ha* the rmsm relative
length of S*683 4 0*094 vilh a variation range froai
$«607 to 6*185 (Table vz) , fheae ^ raa paira of chroaioaonxa
are 6th# 7th and 8th rmtpmctivmly based on their lengths*
fBBUP ¥It fhia group has a singl* pair of chr0<B0S0nes*
the 9th largeat aeong the naierechronosonsa* Since the
centrttaars of this pair is Ic .n md in the siidb-a«diaii
region i t is of lOie um»tfpm (Vigusas 7 ana 9)* oaan
caatrofssrie indmm was 36*025 • 1*113 varying batwaaa
33*333 • 36*461* the aaan relative length was 5*278 4 0*102
d i
^ i c h v«ri#d from 4«984 V2)«
VX^t This gpoup tia« the amallest (nfterochxoaaoaoiMi
p«tir hifviog mm xvlative length as 4.44S • 0«19a with a vitiriatiofi runge fro® 4*049 to 5«000«
Having the ceiitnMwrtt the tmrminal ma$ this
pair is cl«i8i£ittd as ^t^pa (Figvras 7 aftd
(appup ^ i s is t\m last gcot^ of the coi^iainent
v liCh tneiudas the sex«<tiiro'>)osc»nes« ffi io nales and ^
in iaaaiss* Both are lastacantrie homing their ca ae
eesntieomttic inOicos as ^ 0«489 ana 46*642 4 0»e43 i^ieh vatisd from 48*000 to 50*000 and 44*233 to 50*000
in z and HMShcotaosoaas rea|»«etiv«!iy VZ)« Tha
%mchxomm3m& is th» fourth largest oair t^araas tha
i«»chsows«iie is 6th or 9th aocoraing to laagth* Tha
par c^nt ralativa laagtfi of MhioaosomBs in both sajcas
variad from 10*309 to 11*343 with th(» tnean baing 10*844 ^
0*19S* Hie w-chromesome hsa a asan ralative laagth of
S*4?9 • 0*194 lAteh variafl l»twa#n 4*9S8«.S*909 (Tdbia VX}«
Norphoaatrie analyaci . of the two ar»a of all tiia
Maeroehromosones wera also aafla a&d the rasults ara
siKmariaad in Tabla VU* rigur»»9 Ulastratss tha
ftaagths of ths two arms ..of tlw aiaosochcoaosonMs*
J o
£ . MMXPTIOM OF CHROMOSOMES <G CORVUB SPLEWPKFFF
i^elfts be lo f^ to offa«r Pwwftrlfco-nes t^icti
eontain th« of inrA«ii ^yecic* studied
00 f « r « OironaDSOOftl. stisdie* tiejni ciumSk from 3S satlsfACtory
meti^hase pX«f»s» fhe diploid ehcvm&aome fnaiber« including
the Q wsro- ex»9 •aiexo-cilixioeapM^s f otmd to vary f rosii
variation waa tmlnXy due to microchxeaaoo^^i
a« nu'Sl^ aacceelicoadac^ac'a ( l i pairn) elMierved
to bc conatimt in ali, tha plates*
flvD fsmq^my iiiettsphaae ec»iiit
was liot^ 76 o€ tlie total count}* l^e niean baing
74*900 • 0*S7S« tf)^ dl|>ioid cfhxi&iiosoar »uad>iir in Cervtia
ntay m ecmaidarttd as ^ c aa^* of lihich i i pairs
ineiodiisg the emit^ttrnmrnmit ac* tha inacvochKiamosotiiBa*
Hia «i«?fOCti«oaiDSOBias beea cXassifisd into £iv«
9»oups oa tho basis o£ titsir length in dsserndina ordsr*
All the mljcvochcoaosows ot^ acrocentric* s^aall* dot*»sh^ed
Ctin>MK>SOMSS*
CiKovip X has a sin^la pair of large chro^scmas
h«ring their cantfomeies in suto**sdian region (sm)*
fhis is the largest pair* per eent rsl&tiirti length varying
froa i«*OSS to 1»*65S aftd the aeaA relative length beictg
16*730 4 0«<t«. the cmntxomtic index varied fron 23*233
to 28*S71 giving the isswi value as 3S*6€7 • l*ie3(Tafele VZIX),
geygljg^i This group h«i • •ingla p«ir of chtomtomm
i^ich is the ••eond pair having the maii
Irnnigm tm ^ 0«69$ #ii«h ^wximd fvon I3*3€0 t^
fh« c*ntiD(Mn b»iii9 loealiMd th« tacaiiial •ndt this
p« ir i « « ehsotmmimm Crigtea 10 and iaK
p^qp 1 fhia Aontaina « s in^* p«ir of st^^ipo
«9ifOiiiosom« hmvUtq th« e«atKeai»i« ia th* •y£bt»cniiii«i
c«gioii iWtgmB 10 la) , c«f}tfoiwric i»d*it Vttrt«d
fXOiB U«2dS to the m n te^ IS* 197 • 0.427
VXIX>, Zt m^mma thicd in ioagtht th« amao
mltitim haiiig i3«230 « 12* 844 ( f ^ e ^XXZ).
q!i»at> PTi ^ ia£<ge«t gsoup hvflog 7 pairs of
iwtip* «heo9»sai»9# h«vino their onitgonioxoa at the
tttcaiimi end* At mHiphase t%fD cihcoiiietids of
chsonesoaMMi iaatona of Mporetiag lonaSn attwDfiea « t ^
eeiiti!OfMc« tfid pioMnt a v ahipad pietuco* Xa ioiigth
thaaa ehro noaoiw pais* ooeupy $th to tith poaitiona
(finoraa 10 and ia) ifieiuding tha MhmMeaiaa uliich ia
4tli iar^aet pair. «ha f irat^ariea in por eoa% laiativa A iaaoth ff«M 9^992 to tha iaa«> vaiue aa
9.307 • O.iia. ihft aaeeod pair ^icfi ia 4th iargtat hae
a Man ralativa ioagith 7.449 • 0.204 lAtich irarioa ftoii
7. if3 to 0.314. «hft mm toiativ* Ua^th for tha third
pair of thia group was 4. S44 • 0. i33 iihleh variod froia
6 0
to 6.800 vn i ) « Dm fourth pair ifhleh im
•th lar^Mt mm Mmtl'wm lungith of ft.S?! # 0*071 whiai
varies h xmmn 6*137 to ««Sli« the nmt pair has a »»aii
lalatiim iaaoth S«490 ^ 0*07S «liiai variad froai S . M to
S*77«* ^ sixth pair t^ich is the 10th largest variad
in par cant calativa langth froai 4*980 to $*S04 with
iAm mm bein^ $#296 • 0*<I11* Sho last pair of ehromosooa
this group which is tha smallest ataeroehronosow has
4*6ao 0* 131 a« its aRan ralativa langth lAiich varias
sli«^tly hatwaan 4*16« » 4*980 VIXX}*
Vt fhia is the laat gro^p of th@ conplaioant CQiipri9i»9
tha sajt«ehroaD«oass #iich ara idaatif iad as a pair of SS
ohcoiapsoaies in malas sad wn ia f ai»alo«* neoordiag to langth*
tha z% chroaosotnfts ar« tha fourth largast chronosoass*
Siooa tha eantcmra is in the si^b»andian ragion ^ajr could
ha dasigoatad as s(»»t|pa* Tim langth of thasa chroooaoawa
varied from 20*791 to 11*4C6 in 10 taetiphaaa platas giving
the calativa langth aa 11*173 4 0*173* lha cantrmric
iiaasuraaitRts ravaalad the iwian cantroiiaric indsx as 31*368 •
0*981 with a variaUon range from 28*000 to 33*333 (Tal»la irxxz)*
In faflialas tiis S-ehromoso»es are of the aane aisa as in
msles* The ti»ehcomos««es is generally smaller than tha
s-chromoaomea* It ia about 1/2 of tha length of Xi>c!hsoaiosenis*
Its cantxomsric indax varied from 41*173 to 44*444 which
gave a mean value of 43*334 • 0*il3* lha ralativa length
'3
varl*d feon 6«42a to 6*66« the mm ir«lativ« length
vas oaleiaated to be 6*526 f 0»040* Die ti*ch«o«>soiae
is about tAie length of the 7tli chtoooaoiie pair bat i t
EAA easily be iaent i f in having the cem^ mere in the
(osdian region (m-ehromosente} whereas tho 7t}i pair is fwtype
(Figures 10 and 12>« Table XX presents the data to taean
relative lengths of t^e tuo aems and the sasie is ripresented
diagramatically in figure |3.
1? cEscaxpgio! CP SHt; ammQomQ w passer vo^B^xm
gasaer dotaestious is also a imsbetv of the order
Passeriforoea* Xn this species also 28 good matiipheea
plates selected for ehroaioso«al studies* as in other
species* tMD t ^ s of chromosoaiesjthc laaorochromosomes
and the mierochroBiosoass %iere idsntified* The iiiaerochro»
0090009 being larger and easily distingoishiible constitute
the f i rst 11 pairs Including thcs sexachro^aost^s irf)ereas
the mieroehroaosoaes are soialler in site and nost of then
are acrocentric* They are about S6 ih nunber and vary in
shipe fron minute granules to small rod»shaped structures*
Zt is rather difficult to draw a line between the two
types of chromosomes* Chroaosoro counts in 25 well ^read
metiphase plates showed th«t there exist a variation in
diploid count from 6B to 82 with the highest frequency
(4691) at 76* rtm sman for this variation wad found to be
76*720 • 0*175 which could be regarded as the diploid
6 2
chnimoM n\MA3«r in this 9P«ei«« i and 9lgan 13)•
Hi* Vttiriation in cdiroooaonw neiiay
in iBicffOChzoBK>soaBfts« h® {bqococ xqmmioimbs ifuiidbejr ImiiiQ
constant as 32* f1i«»a II pairs of oiacrochcoraosoais have
bs«n tD&assifiad in 7 different groups on the basis of thair
laogtli and position of tHe cantiPi^vas*
^ftom It Ihis giroMp has only on* pair of chrosaosoiass in
which the emktxomm is located in the median xegion
Bluing i t a^tjfpm c iroaoaooe (Figiures 19 end IS), ifozphoaietric
analysis o£ the cmtxmmf revealed a vaciation in ^ e
centcoansric index fsooi 36*666 to 4U6S3, tStm mm being
39.696 ^ 0«83S« This is the largest pair of chroniosoiaes in
diploid set having the mean ar^ative length ^ 0*335
idiict) varies betiieen I6«014 • 17*804 (fWble X)*
Giioyo l i t A single pair of st^chroniosoaies Coaatcoteere in
the sul^tiBminel region) forms this gxoup* ihis is the
second largsst chromeswse s>air* 9he per cent relatiire
leag^ varied fioia 13*468 to 14*243 giving the mean value
as 13*009 • 0*199* Hie msan centxomsrie index was 23*671
^ 0*797 with a variation froai 21*093 to 29*000 ( f ^ e x)*
f>pe<«» I l l t She third groMp has t«#o pairs of chromosomes
having their centxomiMs in the median legion* On the
basis of the claasifieaUon given by x«van et*al** <1964)
' s n
th«y aiMC £onK>aom««* largir of th« t«o
hcs t»te mm e«iitfofn»rie ioatx 4U29S • t»lta ifhlch
fcon 37.S00 to 49«243« cfifonoso-na ooeopieo
tlw thixd pocition lo p^r cant c«l«tiir« Ivngth iirhicti
feoa S0«e76 to iU794» thtt metin xalativa Xmmgtk
being ii*978 • 0*327 (treble Vhm aeeona p«ir of
this prGop is f i f th l a r g e s t b » i n g
4tli largest) has a mm Tslativa length of e«463 0*109
with a v&ciMion r«nga f«D» 6*308 to 8*696* ^ ratan
eootsesiaric Index for this pair was fouml to be 40«i7l •
0*7i3 varying beti^seo 38*461 * 43*eS7 iTablo
jsapoup ayi Tuo pairs o^ chroDOSonses are siic^ and
7th in their length coostitote this group* Both the
chromososies have their centJDoaisrcs in the sub»tesroinal
iragion Cst#chr0:ii080i8es} (figures 13 and 15}* fhe f i r s t
pair of this group has the msen relative length of r
6*136 4 0*106 with a variation fcon 6*011 to e*S40* fhs
osan oentromeric index of this pair was 20*611 • 0*433
varying betueen 20*000 • 22*222 (Table X)» fhe secoiid
pair of dhronosooMi has the oMian relative length as 6*006
^ 0*127 lAieh varied froai 7* 7 IS to 6*4^7* h variation
froB 19*230 to 24*000 was observed in ihe centronsrie
iadsx giving the mmm value as 21*793 * 0*966 (Table X)*
Qppqo vt ^ f i fth gro^p coiiqpriM* tm pair* of mm
iihKomaom»» fh^gm cAircraoso^s crs 8tli and 9«b largest
io l«iigtli« ^ f irst pair of this group has a par cant
raiativa laag^ varying fron 6*S26 to 7*ii7« ttie (Wtfi
for «fhich %ms fouad to ba ^.e^O 4 0« i i l * f t » men
cantxosiatie indax cornrs to be ^ 0*Sia «Hicti variad
fcoa 38«09S to 40,909* Thti aacond pair has a variation
ie par cent ralativa Imtgth from 6«23i to 6*734 with tlia
sewi ralati^'e iaxigth baing 6*628 • 0«100« The eentr^Qerie
aattsucaniBnts ravaaie<3 the oaan oantr&iEiarie index as
42«66e ^ 0*662 variad betuaan 38*09S 45*000
Xh
^ potm Vli Tm omll T-chro nosoaie palra ccHistitute this
groi^f idvosa mun caiative Imgttim ware found to b«
4*993 • 0*242 and 4*135 • 0*132 varying taatMaan 4*i66<»S*693
and 3*«46-4*626 raspaetivaly* ihaaa loth and titii in
sisa.
gpottft VIH ^ i s is tha Iftst group vhich ineliMSas flia
aanHjhfowoaomaa. In thla spaoiaa 3ffi/m aex-rnehmim
MAS 6bsarvad« tha aalas baing honogawtic* 3ha i-Kjhsotioaoass
in both saxaa ara fourth largaat and aquai in length hiring
tha Mean relative length as 10•231 • 0*11? ahioh varies
frooi 10*033 to 10*676* The naan eantrowric index for this
chrooosoos was ealeulatad to be 43*318 • 0*734 vith m
VttTlatimi betwMiii 40.000 to 43«7S0« a^ing th» iSi*ehxo(ap«oa»»
« « Dm titrndhgomaomm «hlch is «B«lI«r in Imogth than the s-chxwaoM>B» Is in X^ngtli to tSie 9tli
largnst p«ir of Hhe ^utosoims* Thm N^hcomonooB h«a its
«entco(iiertt in the taeaian xegicn hwing the mm
centronerlc inaax 39»ai»9 ^ 0.740 which varied fixom 36*642
to •Wm itie variation in par i oiit salative leo^ch o€
was frcni 6*331 to 9*05i, which genm tim
mm Mat ivc length as 6»693 • 0*l3i (f^bla XK ^
relative ler th of the two acmo o£ the maecochstMaoeosaea
are presented in Figure 15 «cid Table Mm
S> sgscRiPYicti CP THE CHRotmoms qp %tmmzzA mvii^iicBP^ mim^t> aawgwG (gAssERiFcuNBs)
species is a wmibes of the order ?«ss»riformes*
Chre^so3ial studies mrm oiBda on both of the gpee&ss
as usual* S vm a^agat the slides prsparad^25 good
mataphasa plates were salaci^d for norphooetric wialysis
of the chxoraosova* Chro^wsonal counts of the nataphaaa
plates revealed a variatioti in diploid imnber fro>'a 70 to
62 attainir.y the hicliest fraquancy at bO (4t$4)* ^
diploid mnfaer in this apecias aleo cm be aoiprassad la
tanM of the tumm which waa found to be 78*400 • 0*S6S»
MaeiochccMioaovea and nieroehroaosoaiBS do exist in this
species alao* the aaall chrooiosomes are telocentric in
6G
origin m» •Jcp^cM variation in nwAmr w«a observed
only aaiong the mie«w*moD8om*#» fittt number of fmeroehio*
tftiich mem 22 ineiudino tho ••x«ehfotaoeomee« «eiii«incd ooiuitaQt in all the twentyf ive pl«tea« titm l i paire of »aarochxoaio80B»s are «iaesl€iea into 1 groups aceoriSing to their iengt^ and oantronierie indieea,
ggt>\!P mom1* fh« f irat^has tMo pairs of large chr^aosonies
having ^ e i r cantrc^u^a in tiie n^tiim regioii im)* Ttm
f i rst pair is the iargeat# the relative length varying
from 17*102 to 80«179 end the mean relative Isngt^ being
as* 591 • 0*S44» ttm centromeric indOK variot. §mm S6.666
to 44*444 and the laaan centrosierie index iraa found to be
• l*2eo (l^dble XZiK fhe second ehroaosoiaB pair o£
this group which is siaallwr in length than the f i rst pair
(figures 1ft and 17) has a relative length varying from
11*310 to I3*94S« the asan being 13*338 • 0.S31. the sewa
eantroaitric indiM of thia pair was found to be 98*040 •
0«Sa0 iiith a variation range from 37*800 to 40*000(Ti^e XXZ><
i reuo l i t n»is groi;^ consists of a single pair of chre«o* so«es imich ae the third in length* In t^is pair tho oentroNsree are located in the attbH«edi«i region making thMi s»»typo* The p«r cent lelativti length varied from 8*980 to 11*143 with the mmn relative langth being 10*37i4> 0*344*
Tim c«Rtroai»ric indttx «r«a S2.436 • 3«2S0 tthich varied
fcom to 3T.142 <Tia»3L« XZZ)«
jppup ll|t ttie tliiJrd 0roup contains a s in^e pair ot
ehreaoaosted v^ids are f i f th l a r i a t (S^hfomoaoma
pair baino the 4th largeat pair) . Thia pair is aaaignatad
Mt m»typa having the nocn eantrcHaaric indess 49»i96^ 0«7e0
tthieh raried ixtm I7«S00 to 42,105 (l^oures and i6)« f
The per ceiit resiativo i a n ^ varljed from to
the m m tor ^ ich wm found to be ®«68S * 0*083 xixh
gapo^ gyt fhia groMp haa two pair a of chroaoaomes «diich
are sixth and o&vmth baaed ots their lengths (Figtuea
i6 ind i7)* The f i rat of thaae two pairs haa ita aieaii
centroaaeric indaae as 22.S4B • 1*047 varying from 20*833 to
26*315* On the ba«ia of the m m oantromeric the
ehromoaomea may be daaigoated as at t np * the relative
length variation being 6*163 to 6*520* the oiean relative
length vaa e«375 • 0,069* Th^ aacond pair of thia gsot^
(7th chro'iiosonD) lias alao its MntroiaBre in ^ e aub^teraainal
regicm (at^type ehroaaMotieal* She aiean relative length
of thia chroooaoma was fnund to be 6«100 • 0*046 with a
variation range front 7*956 to 6*231 XXZ). 3he
mean cantrowaric index for thia pair waa found to be
22*230 • 0*539 varying betneen 20*633 to 24*137*
l-i (1
^OUD Vt pdirs of otBQll A-ehromosooea const!tut*
thl» qroMp* They are 8tli end 9th according to length*
their rmmn relative lengths w«re found to be 6*740 ^ 0*149
end 6*374 • 0*264 varying I»»tw«en 6*402 to 7*170 and
9.S72 to 7*170 reopeetiveiy* The raeec crentronerie indices
for these pair a came out to bs 39*697 ^ 0*669 and 42»S92 ^
0*646 ««hich varied from 37.900 to 42*697 and 40*000 to
49*000 respectiwiy (^i^ble XIJ)*
groiHP VXt This groui;> has two pairs of smaiiest miHcrochro*
maamBrn *Sb»Y are iOtti and Ittli in sice* Stiese chromo*
so »8 posseas their centromeres at the terminal, ends
and are therefore T-type chroons&a&s* Itsey have their meon
relative lengths as 4*671 • 0*169 and 4*424 • 0*307 vith a
variation in per cent relative lengths from 4*496 to S*360
and 3*621 to 9*380 respectively (Table X 2 X ) *
group VXXI She sex-chroiaoso^s constitute this grov^*
Thr males toeing homogwistic and the females heteroganetic*
a typical 2SS/2M sex-astchanisai was observed in this species
also* Xb either sex the 2>.chroniosoBes are fourth largest
s«-tfpe chremosons having the sean centroaeric indsx as
36*347 4 9*000 which varied from 29*000 to 41*666* ^
per cent relative length of SB chroiiiosoBis varied betMeen
6*960 10*670 giving the aiean relative length as 9*966 •
0*280* itie w-ehroaosoms which is between the 9th and lOth
pair of the autosomss in siae has a tasan relative length
o€ S.9S7 ^ 0.232 with • varlatiofi range from 5*487 to
6*726* Thtt ccntroi»Bff« b«lng In the oMidian r«gicai« f#»
ehroR«>S4aiie nn chromoaooMi* ilm aie«n centroOf^rie
trOmx vM 38*4S7 • 1*349 varied from 33* 339 to
41*176 (Tabltt XXZ}* Th« long arm loogth^ and iftiort acn
langtha of the macrochrcmsomeo hove hmmn ehom. in Table
Xll l and figure 16 in the form of karyogren*
h eonparative karyotype study on the basic of
the laean relative lengths ot 3 species of birds (ATfSea
mnil^mli^ SESXU mJSXU «nd Fyftiffgll^T^^
PPPI^I^yltW be^ raade in Table XIV
and Figure SK>* coasterison of the mean centronneric ic^ices
of these apeelmB is given in Table XV* Table
and Figure 21 give a comparative eccount of tl^ mean
relative lengths of ^ e two arms of the macrochromosooes
of i^e above mentioned three epscies* Figure 19 gives a
cofl|?arison of homologous pairs in these species* the
results of a conperative analysis of the msan relative
lengths of the maoroehrolwsomes* their arespective arms
and eentromeric indices of C ^ m f&l«flt<?»nf £i£2t£ dSStft-ticus and Irtieriea bruniceps hmvr been sumnarised tn liables XflX, xzx end xrill respectively and are Ulustreted in figures 22« 23* end 24*
A eoQ^arfttive anelysls of all the six ^ecies
undtr study giving thslr mean relative lengfthe« eentro*
fmrUt inatccs and arm lengths has been ^own in
Tables XX, XXX. and XXZZ,
Zn th« pxvsrat vock ehronMoMs of Bix •p«el«s of
birds from thXM h«v« b » « i atudM* Hi* two
^•eitto iCdlia otirowernm tlffttfMfif <S«it«ni puaepl* IwriHi)
^ ^ ^dooiti cBravii (Zndl«ni pond horon)* ovon ttiougli
boionglng to the sftigo ord»r Cteoaifoffaie* ffovoalod notcblo
dif £««onc«s in tikoir ehronosoiM** In Ag^a fpogpuroa
wanilomiia fStm diploid ntMl or of oturonosones vi^iod f ra i
S4 to 66 witi) « a«rfc«d p««|t (4B% of tlie eoimt) « t 62
whil® in MpdOQl ffTittyii orftvii tli* amibmg^ v«ried from $4
to 60 ana the b i^est p9tk vss noted « t 64 inat»«d« &m
ey^cage or men ehremsom mtsS^t thus differed* being
61.44 in shes^ ismwm ffittf*ii»iiBit A»3 ^ . 6 in mssSbk qr^v^ gravl|* Hunger (1970) in his tmrk on the chromo90o»8 of Si species of birds reported the diploid rmtomr of ehromosomfea in Ardea cimctaaa 66 with a eontinoous redootion in length from about 6*3/1 and V9% of A • » to about
and of A • B« A Icaryotype attidy of the male
of the Sana apeeiea* Ardea einarea by ISMShioa (1990) described the an nunAsar as 76» the aeeroehromosoaiaa
nere aliagad into 6 pairs of MSta or aubHeatafeentrie and
tMO paira eC teleoeatriea* Xtoh et* al»# (1*69) studied the Ohroneaoeaa fron the feather polp eiiltures in two
birds of the s « m i species but of uidKBoim s e x * They
noted a variaticn in chronoaeMS naoAiar irm S3 to 64
M if:.
in 97 good nctaphm*. The emplmmnt tfith 62 ehronoMM*
wfts eonsliSma t^Scal d^Ioid Mt in this
flwM chrcoosoaes cosmimtma of 28 iB«erochffvmwwi»«
inciiuSing «he ••»«<jhroaiosonB« and 40 «ie»)ehroaoaomt«,
Xn th« pr««Mit •tttdlM th« MaovoeturoiDosoflita in tooth «h«
•p«ciss o£ Cteonifonnes found to b« 24 including th«
••x»<luroiao«oiMs« the nmitmr of lalcroeluroaiosoats
diff«Md« teing 3S in j ^ e ^ gmwff f j j a jU f f i f l i «
WIYU OTYll* in ddtta filfitflft CZtoh «tv ol.* and [ydo oumpufa Bfnii»n«i« as*
tn typtt utMreas in Ardsola caravi^ they arc of M-typa.
Similarly tl^ second pair of chronosones in ^ ydan amrouraa
mmiX tmim waa M-typa but in Agdaola oravii oravi^ i t vaa
ae>»typa* Third pair in the two apaeiaa« Ardaf purouraa
gpanilaaaiy and cinara^ raaacdolad i » baing mm typm
bat it ia »»typa in ^daola qravii grav^^ fha 4th pair
which ia f i f th largeat in langth (a-chroQioao» baing tha
4 th iargaat) eonaiata of la-chroaoTOaea in al l tha thraa
i^aciaa* lha f i f th m^ aixth paira in tha thraa ^paeiaa
baing coaparad «ara »>chronoaoaiaa in tha f irat tuo apaciea
hut in Arda^lf oravii oravii tha Sth pair ia o^aad tha
6tti pair ai»»typa« Tha 7th pair of Chro aoaoiata in Juc aa
y^ fp^M naUSDSiE M'daol^ fFfY^^ ffravii ara
T«ehroaioaoiaaa whUa i t ia H-typa in Agdaa cinaraa (Xtoh
•t* YVfBtlfiitig 4 pftira of ehsi03ies&ti»«
< a . i i ) in AEat* gWWtft an MflEfiU ftUXil or*vi| «ikd 3 pair* in iy^yi cinr^^ « i l
chrisnotfoiavs. S«ehroiio«o^a in Ardta c in»yf and jyfdt<y|f
ggftvii qr^vif are mmtyprn i^ila th«M ar« MKtypa in
Airaaa pugpuraa manilanai«a fha tMstironoaomes in SESlA
filQSISA asSEdS flFgYll SBticU 91- yp* i^ i l * is
in Ac.aaa putt>ar»f manUaraiif^ TaHagi and ^aaaki il974) daacribea chro nottonies o£ nine ^^aeias fcom
fciie order Cieonifoxnea out of v.tiich aeven have thair
aiploia nuQlMir as 69 • i«a. ^ IWi l i ffjK?gnla>
^Ifgptft mmH lynfqtitfRfflif
aathiopica and giidoyigiua In tha other t«o apaciaa
tlia an mnds^ waa 73 • and 70 • raiq^^ctively, Aceoc^ing
to tlia^
^Ahand** ^pa group A ia found in four out
of th» a apaeiaa f iTOBill fflffgUta fflCTBlt^ ftffWta
fceteBESlni^ I f g lBpmw
liirilltntfTrMlii C oup B OC tl^m iqpaeiaa eonaiata of S
paira of Mtacantrie or aub*a»t«eantric alarnanta* Unm
Z ia a aadiUMi aiaad oataoantrlc and tha If a Miali aub»talooantrie ehroaoaooiB. Cn cooparing tha umm raiatiira
V 4
X*tt|fth« of th« f i rst pmif of ehroflKwonBs it was obsacvsd
tt AratolA arsyii gr^vi^ it is loogsr C24*««S 0*9SS)
sua figufts 30)• The Isngth q£ th* long anas of the f i r s t
S^sir in th« tuo sps£i«s also diffcrca^betog 19*349 • 0.180
^ I f f ^ i man^iemU (Table ZIZ ana Figare 3) and
t2*46S ^ in Ardsoia aravii oravi^ C f^ l « V}* «7ttat
r^^rsG is ^ cam witiht short a r » i^ich are smollar in
lardaa purpiigef tanilanais Ct0«159 • 0*199) in j raao^a
^ay i i grayii <12.19^ * o. 231) (Table XS?X>. ms mm
eeiitro3N»ric ^saie^s of tlie tuo i^eoiea also ^oi^a eoiuii^to-
rabie aiffereoce. For MdSa fiSISsjSSa m l t e S l i , i t was
43*209 • 0*449 and for Acfleolf aravii^ <yavfi it e^ae out
to be 49*477 4 0*2iS (filxle Xfh SiraUarly* other pairs
ci ctirocDoscQies in two species also showed differences
in 0ieir aean relative lengths (V^le X2V and Figure 30)«
nean emntxemvic indices (Table JOT) «tid the two an!» tM
^ dbtrnmomm ( T ^ e lorz a»a Figure 2ll* ttisre end
srivastava (1976) reported the diploid nuoiber in flubulciy
J gtlA (Cattle egret) as CO out of whicih 7 pairs including
the sex-chrosiosoniis were classified as maereehronosoMis.
Four pairs of thm necrochreaosoaes were designated m»typ««
the Sth pair as SM*type and the «th as Ti*<3lireiaosoiiiB*
V j
Tim Hx^hxmiMm in tliis ajpmetmw W9» £oui»d trd or 4tli
iargest ai»typ« mn ^ ^ W « • « wmtAlmr ti iehiioiaosoM*
these fiadiiigt could be emwidsMd in conformity iritli
tti* prtMnt fisaincp if ttio ipoei** diff«r«iiee is tslcsn
into sccount* ttrnj slse tmport»d tlie ctifomosoaii
ovq^lsioeot of one individuai i^ich dsviated tlis
fiormtti hmemm of structural heterosygosity iinrolv^iag
a mm^x of chroniosoaiB pair on* a aiococtiroaosotas*
Ths clironosows of grsflcolinus poafttosr L^wuf
(<SE«y partridgal bsioiming to orasr
<lallifm!es nhomA a v^iatioo in diploid nmdamg from
68 60 %fit tHe ti'i^isst paidft of tlie sieti|»lioss
plates coated) at 78, falluri m^ Va^i 11909) ia s
Cotumiai eotmikisiL imot ixim. found ^ a t 9$% oi aiitotie *
isitaphasas of taot i sexrs hi^ 76 chffomo9omat tkad
7€ or 77* t^iie in m^arly cases thil tjnibar variad
fcatxisan 73 to 7S 4«id ons tatraploid call had 193
chromosoBiss* Pontan (i9#2)« in tha 9mm spscias gavs
tht diploid nonRbsr as 78 • Piccimi and Stalls 11970)
»<«»idf ffalaaoria «3uiasa fowl) rseordad 39Si of tha
nataphasaa to posssss 80 chromosonss and 20X to hiva
78* Sinca tha two diploid ntail»ara did not lAioir much
diffaranca, thay %#ara unabla to dsoids whathsr ths
V G
h«ploia amt of th« Gains* foiri U or 40* MXag (1070) r
gave an Ulyatrntid tfxoont. ot the momtiic dhMxmo&omB
^ ehixm^im (AsastraXian king qu«U)* fl»
also mmQ^ • con^oratSi* maiyals of the chromosoMS of
fiffilml^i mP^mP^ <ruiu } pubii8h«a
by Ohno mu (1964) and ^alluri and Veonl (1965).
In gMcalfactoria chinanaia he racordcd « varlatioD
fcotii 70 to ea itfith the hi^ieat {HBtik (31%) at 7e« Tidcagi
and Saaailei (S974) atudiea the diploid chronodo^a oe
3 ap%cim9 birds from the order Qaiilifonaea*
Baiearic^ pftfronia oayonie having m • chcosK soiaasi
m m OiJSU. hmim ea 1 f M i M tayMfil^i^i having 76 • ^
yffj^Ypjygtrwit feiffal^trilwR tuning 7e • i md rhiffimit
having 62 • chroaoaoraea* Xn the present
on graneolintia pondioarisMua interpoaitu^ tan
pairs of aaereehroaDSORias ineluding the wxi'<Sm>moaom9 hare been identified (Figuxes 7 and 8) out of which only
one pair (Second largeat) ia the n-ehromDaoaa* two paira
(First and ninth largaat) are a»-ehroao8oinaa* one pair
(5th largaat ) ia at«chroaos(m and the seat, (exeept
the aex-ehroaoaonts which are H»type) are T»chxoiaoaatMia«
Aay^aadlwri et« (l9C9) in Pavo criatatuf found
v v
A6 ctaomoaomn of which 9 largest pmirm iacludiag 32 pair
in !aal«9 vera the nacroehroi«o8on»»« Zo fetaaiee
ioatead of 9 pair* thay daaignatad 17 tihrotaoaonia as
taaeroehcoiaosotaas bacauaa they ^ r a uiia}>ie to identify
the li-ehroiaDsosie which is their opinion waa aioiiar
to aieroehrosioaoaes. mmg tite aacioc iDiBoaeniea tbmy
deaignated 4 paira aa ii»type# two paira at*typa and
tuo paira aa T«type ctixoraosoBMia* ZZ ehtosmao-m pair
was identified as* m-*i|areeD8omB« Owen ^1969) studied
the somatic <dh«0{B08<»»ea from the i^ieen* t ytmis and
bone Qiamnr of developing ehick (Oalius dyaeati^ya>
enbryos 14*19 days old* His f indinga ^ t h Callus doaesticuf
are quite siailar to the pxeaent findinga* exeunt for
the f i rst pair which is a»»type «od Zi diroaosoniB pair
i^ifih ia fourth largeat in f j m ^ U P B imflOAPyg^m ff
iB^gPftt lW ^ U e in mkUL HfffffiUfflW P * ^
ie f»>type «id the Smdhmomornrn ia tlie Sth largest,
A c(a«>arative study ot three species of birds
haa been aede from the order PasserifonMa, In Coryua
aplendens from the family Corvidae* the diploid chroiooaome
number was found to be 76 <S2li of the count )• Theaa
finding are in coitformitv with the findings of Hamner
7 8
(1966) in Pica d ^ a MoagAag to tha amm f wily* ^dtmxm
th« diploid nunbsr has b e « jraportftd to •xaetly tha
•an*. Howwrar, in GbqqUI fPlffM^^m tha nwAmr of
naicnxhronoMMia waa fouyod to ba 32 indtuding tt^ aa3»>
ohTOooaooiea wharaas in Pica pica tha aacjcochroiiioaoiMia
B\iii£iajr)ed only id* the dif fe««nca in tita mffiAser of
aaeroehsotaoaoMa in the two apaeiaa may be dua to tm
fact that ha oroupad a pai«s of naccochroMoaeaiea with
the miexochs&aoa&aeB* St o©n be seen fioiii the figiueaa
11 enS 12 that theca are ciaacly 11 pAiro of aeoioehi*
jRKSD^^a in Cogytia aplendwia* fhe ajrror could have baan
ceuaed due to less developed tachniquas of miccoacopy
aad mioic^hotogr^hy at that titai* As « eonsaquanoa*
i t ia no nondar that tha casults of the <$hxo^ao9ial
atudiaa of birds by earlier raseardiars ha^ baan
modified by subsequent woxkers in the f ield. Loyes <l906),
tihoaa woclc on the c;hr^x>aanea of CSellus ^teiatatieua COoaaatio
fowl} had just atartad* obsanred six ohroMOSoiaes in tha
primary oocyte of thia species* lhan •omeabcodt <1906)
obaervad 8 to chromosomes in partlienoosnatie aenatie
calls* Qksyar H m ) was the f i rst mtli»g concamsd
with tha dhnmoaomes in the spasiiateoenasis of
7 9
fevi. Me described 15 to 19 diromotone* in th«
•peroatocpnlftl and 9 chn»c»so(iie» in thm pria«ry
^pwMatecyt* el«iaing th« diseovexy of m txipmifA in « » mtlmm of 2 iUat
eon^sfiBg the saerochroaosonee of Coryue eplendene with
those of yice pice* It ves found th«tt
Ttm f i r s t pair in Conrtis spiendens is mnotypm hiK in^
the nean relative lengths of long and short anas* as
13«887 ^ 0 M $ end 4»832 4 0»24« XX and Figure la )
and «n axm ratio of about lt| irhereaa it is o^type
chroaaosoae in Pic<| ica haying the arta ratio of lil*S*
2* Second pair of ehromosoae in Coryiis sol ndens^
identified as IWihronoeonie tshile it is t*>typ« in Pica pip^
with an arm ratio of li8«S*
lbs third pair of chroaosofaes In both the species*
Coma EJm SiSA. st^type. Zn the
former the mean relative lengthe of theiuo eras were
la.ses • 0*120 and 1*007 • 0*083* the am ratio
tieing ltS*5 t^ile in the latt»r the ana ratio vas lt4*S.
4* flie next three pairs which ate Sth* Oth 7 th
eecording to their lengths sre sti»ehre«osones in Pica plea
having a n ratioa liS*0« lt3*S an^ li9*0 respectively
«^ile these peirs of chronosonss are Tutype in Corvtis
8 0
S« Ttm 6th Iftr^tt^t pair* in
CorvuB mplmndmm iM t^-dbKomotom i^iil* it im «tt»%ni«
with m armr«tio e£ lia»S in SiEaaiEl*
last mMrcoehroMosone E>«ir In Pica oiea («hicli
is ttquivsient to tlie 9th l«r«p»st pair in Cecvus idtiEBflEE.*
found to •t<»typ« %eith an ai»ft ItS.i i^Ua
its ccu^apotiding siata in ^ sgvas splatiaans is O^type*
7, ttis last two pairs of taacgochroaosoass in Cogvfi^
spianOans «hieh are absent in yira g j ^ are T«ehra»so(iias«
6« the tha SMrhromosofaa is tha
4th l8ro«»st st»»typa chromososie with an arra ratio oC
ls8»S in Pic^ Dica> 1!he s ehrcxaosonae in so l c^n^
was also s»-typa having the tnaan ralative langtha of
tha two arms as • 0*a04 and 3*Sa3 • 0«09i« liia
ar» ratio balng ii3«4«« 9h« t«-ehromosoaas in both tha
spacias artt Mhroaosoaias having tha am ratio as it 1*3
in Corviia Mplandaita end Sil,S in U s s 8lE*»
9. Stwra ara 98 nieroehroaosms in fJiSA OlSA. ^ ^
Sacoad spaeiaa atudiad in this erdsr was Pasaar
<[p^ticua (Hauaa sparrow). Xarliar wofliara h«va
daacribad tha chromosomaa of this apaciaa (Van WeUHit
19§9% Castvatriajo at« al** am) PrammA «id PatnaDt* 1977)
8 1
hat iMiith*r m eemi^mtm mwpihoamtry of chroaoaetats nor
th* dftfteriptiott of th« •eif^eluroiaoftc^s has btan oiveii*
Hmmmie ( i W ) dtacxrlM ^ d mcmommm of f—ncf
fpntmuf^ fii» obMirvea 76 • chrcmosoisM tii this upmeimm
with « cootinuDus veiStietion in Ifta^h f ro » about Sytt
and of A • 8 to i ^ t 0 »3^ and O.CK of A
Van (1999) i^octed 76 • aa tha tfipioia nunlbar
nAiUm ^aaad and £>atii«ik U977) gava 78 • aa the diploid
nuTibar of clun»aosoi«aa in Paaaer ^eatatieuf. fha praaent
«#ork aiding the 8ii mntmr in faaaar doiiiBaticuf baing
76 4 eoaforiia the findinga of %aaad iiid Patoai)i (1977)*
A coBipariaioa of the pr^sant vkxU th that of Praaad
and Pa^iailc (1977) has btm aada in the following t ^ e i
l*flibla X3CZZZt A co^ariaion of the atudiaa of ^ M a d and Patna^ (1977) with that of pcaaant tfork on the ehroBRoaoaaa of ffaiffr dcenatiofiia (House aq^arfow)*
Chrozaoaona NOrlt mm F a i r l e f f e r e d i i a l a t i v e Am C a n t r o a a r i o
t o l a n g t h R a U o Z a d a i c
Z V r a a a d a n d
t a t n a i k
« 11.00 m 36.9
f s e a a n t 01 i7.0040.sas l t l .S 39.§9«40.63S W O K l l * *
zz ffraaad a n d
l^ataaik ••3 mm 36.9
ffraaant a t lt.60940.lSS li3.2 a3.67^.7S7 ttmtfk
z u Praaad and P a t a a i l t
ta 7.6 m 36.S
P r a a a n t m ll.97640.a37 l l l . « 41.399^1.113 W O l f c
41.399^1.113
OA.
Chtomtmmm Hoik mm f ^ mirnttvm Mm Cmnuwmrio
to Imgm i^mtip laOmt
XV Pffcsad aoa « • 40.4 PmtnmiM
Pgmmnt m 0«4«S^«t09 itUi 40«ni40«7l3 mOfH
V and a • 42«S ^atiiAlk pjtmBmt St e«l$ei0,i08 ti3*? 30.etl4p«433 MOik
VX l»r«9ad •Bd Ml « , » * $U i P^tamik pgmamtt &t 6*008^.127 Ii3.$ 3t*793j0«988 wwdi
VH Prasad and Sa 8.9 • 27.3 Pataatk mernaent m 6.880«0.*ll ltl«S 39«962^,S12 mtik
VXXX Prasad and a S«0 • 39.7 Patnalii VrasttBt m MWB^.tOO tit.3 iiosk
XX Prasad «id m 4»2 • 44.8 Pataaili Prssmt t 4.993J0.242 *
Ptasad and at 3,7 • 13.8 Pataatt
SMihfOHiososa Prasad i Pa«a«lk Pr«s«it m 10.23i^.XX7 ii&,3 42*3i8j0«734
w-elvraaosMS Prasad and • « • • Pataatk Pf«asilt • 8.89340.I3S Xtl«S 3f,a89^.740
109
fh« eturonovonMi isorpholo®' of anothtr imbar of
th« <ira«r 9«»MirifonM»« btlongiog to tmUt 9XoemHm»
gaawrigfc braniatps dtudteA in th« preMmt work
feuBtS to tm quite siaiUar to that of ^ f y y ^oapaticuMi
in the following «i^«et««
the Mcond pair o£ ohro aoaoma is t»typ« in EMftSlC denfeaticua • 0«7St) whftjpeatt i t i « »»typ« in
fanmlwi C3e*6«o o.saou
8* fhiB third pair of t^emsam !»»t3fp« in faaffcr
aoinegticaa (4i,a»S • t . i ia ) but i t is ao^type C32.4a@ •
3.280) to wrvmlffiBi*
3* 3he &>caironioaoaiB in Sgmm. ^ammtlcaB ia
ooK a Ua*3ie • 0*734) ifhile i t is siftypa in Kabaris^
iSffffllWt ^ $.000).
TiOcagi (1973) dascrilaaa the rapiieation of chrooio-
msmt in 6 i ^ i e a of birds* yoachmm stfiatf, ISfiSlSBXSllft
fiiHtiifftli^ tti^ g f l i t w i ^ fWiifltetHtft wairtmra ffa^tt^ f^nrtfyiif rnmM^ Aeeordingto
htie aaich ap*«i»s limaarad to have diploid ohromosona
nuadaar of 78 to 63 includino 9 pairs of Maeroehrososonss*
•we^t Muthiid* ntf which had m ttctra swerochroeoaoe*
pair* fhs asMMBhroseaoras cmiatiMticm was i i in tha
Mais a»A in tha f aaials* tha suehronvsoms vaa tha 4tli
or the Stb iargeat la the c<»[pl«fii«nt nhcreas the
W-was anong the anaileat m«cxochjroiaoacaes« The relative
ieoQftlia anS ratioa in the eorxea^ionaing
maeroehroinoaoniB paira including the aex elenienta mxm
practically identical io Lonchura atriat§ yeenlopvoie
caatanotia oryalvora* M ^ u ^ the over all
reaetsSaleiice was evident* karyotypio detailo of iBfiOftS*
iamaE ff^l^yfgi differed fr&n f
thoae ot fcpnehura atrieti^ in alee &ni$/or eentxomvic
poaition In oerteln c^wo aoscfoea* On ^i© other hand
fetaryotypie differencea hetwttcn CardiiBlia aplnua end toina^ur^
atriata i«ere exeluaively due to fShB 'oodif led centrotserio
location in most chroraoaoiaea* St particularly
intereating to note that correi^nding paira in theae
tMO <i»eciea flowed eliaoat identical relative lengtha*
A ooBpariaion of thc lean relative lengtha of all
the aix apeciea of birda atudied XX) ahowed tdiat
the f imt pair of ctarowaocna la longaat in Jtrda^la XMiiik
ff^vii (24«MS g 0*3SS) and ahorteat ia faayfy dnaeptlcaa
(i7«0 4 0«SaS)« Second and third palra are largeat in
ftr^a pmvFf i 'BMUfMU •ba uam^Um caiSl^fyliBM
infrpoaitita C33.a03 J0»182 and 20*9«1 ^ 0.247I. meaa
tt<o palra in Corvua aplandtna and Ija^ariaa brunicepa
5 J
e e e w the 4tli Stti positioss (ti*7ao •
0*f9f «nd tt*Sti • MMlSrsi* emnttomtiK UMmmm iTMm XXX) that thiw
til* apwei** t tudM BtygMffa n i t t f a iH# f j i m donitic^ii and ffKfctrtlu truatemi) f i r s t peUr
Jtedaola ar^irii aravi^ Iim ll»i^f«aio«oait» lynA
tli» tMO ^gmWfttUilltfli poa^icwfi^ttu* ^nfgao^itoai ma
Cegfitt ftti—d—^) hmm m^tjfpm tiatmmmmm^
length of Mcond pair is ^ i «rg«»t in Ardsji
f t f j i i f i f U f o . m ) ^ n m i t i « M a u » t
Ci8»Sii 4 0«0fsi iB Cagyaa jto^yjawi^^ flMi tmwaAt tSlIrd* m
f o u r ^ «ad f i f th positions seeordiiiig to th« as«o rsistiirs
isttffttis of tlis sseond psir srs oee«pisd tqr ttlBSSlliSE.
twfllftff^tflif F tasytmsiti g USelfli OeSSS), igfllala SESIBUL fflp^if <I« . IM f »ssssr dow«stictts {i9*869^0.iSS)
1
SBA M n U i i i iMmtiinn Cis*ii« ^ o . i i i i ssspssui^oiy,
ftot •n auroHiQSOMS in tits six •pmim& 4if fscsd in
rslsti¥s isttfths ssd «s«tiwisrie ittAisss* feat i t wss
sILnsys ths fosrtli lsr«ss« psir in s l l tKs gpssiss. Wssisr
H w i in M U L iBitc^ Biimurm m d fifii^ alfia
siso rspsrtsd ths s n #irowsssts ss ths fourth isr^ist psir. ttson ( I t m ) tX— fsuna tlis s-etuewsswi ss tlM fourtfi isrusst in maiirtf Atslw. Ilsiisver, in Mcipitor
8 G
teeaSys, Uml m^ MMNwri i|i79> th* ••x«elifeiii0MMi«
to te tiw f i r s t la^gmt AMari ic»il Cl9m)
fou«4 that tiM In aafladaryai wci mm
tlilrtf Icfftst suib^tAlsGEWtrltf elifisfliMow «8d in iaat«ilM
••tiilfr/ifii ^ SMdhtenosoiM wns id«ati£ift(jl •• feiptli
l.«rg»«t Mftt«e«atrie Miiai mA aatart
l « ra» «t ^ftir IB l>i«BDitai mSi Ifieoid^*
aw^atfasm^ M m^ notmi d m ) a t t e r I M
Zrn lmmomttm tli» 4tli Itatgamt in
ftM tiiiSMi mA Bim &f ^ irartad
v i M y in ai f f « « «nt wsmcimB m atMsribta • « r i l « r «
lha MbeliraesoMi in «ri«B fMi^a mtm SamsmA for tlM
f i ra t tiaa l»r yiadrie ( I9f l l in a e m U e
iifti-isii (ii#a> dttaetad a anali iata x«spiieatia9
eluNMioaM by aaaaa ef aiitar«dlia«r«>liir ana auggaatatf i t
•a a iM^tiiwmeaa daaiufitoaa ^ ytattrie <l9«i>* 0«aa CilftS)
aktaianti nail atf inad aatupliaaa fif^iaa f ra i variena
tiaaaaa of ehi#i aalMryaa « » « rapoftaa a WH dhemoMfam
aoaatitatiaa in tlia faaaiaa*
M ia aviaant a tnietX atrian karyotypa coaaiata
af tHO tipaa af alurafliaaaaaa« «lia f irat fpm iaciaaaa a
o • I d i
c«rt«iik BiplAr of l&tgB eluDomons* ealUd th»
—jggochiowoiOTf and • I«irg» iiuiib«ir of saiallor chioaMoMt
ir«tR«iiii9 irem m M roO^htptd tttuctarcg to aiauto
CftXled tho raleeoetixotoai* tuo types
of eheomoaomM mtm studied toe the f irst tim jyy ttaiies U9a4)* m did oxtcosivs voek on tie otixooosooMMi
of the donftst&o foni. From his studies 193# «tia
|93f) i t toeciine avideot that th« chcottosoms coopisic of
fowl consists of s esfftaia nursabsr of nseroehroiBosoaiS hsnring
• dsf iiiit« slisps ftBd m IsrflfS auadbsr of wmry oAaittm grsmtis liks aiicaroehcoaosoM*
in Hswooaawr stitdisd s|>sfwit09ilissis of tito deaestie fot X m6 xsported ths bssic ehcomosotis mxataf to l»s six while the aierofihroaosoiMis wers mlsgstsd to tim status of aeesssory or supsrnuHttracy ehionosonss on
basis of thsir variaibility in sttuoturo* rosrtisr
bshariour* Nslandsr (1910) dsfinsd aoeassory ctiiowusoswis
*to dsaots such axtra disonosoMss as af>s not hoaol^gous
with thoss of ttia osdiiacir oeeplsesnt and wlioao aunter
is in o^ilibriun ia a populatioa"* Mhits 19S4)
objoetad to lha tam aiBosasary ehesMOSonas as having
baatt loosaly usad by ss«a aattas %dth Mmxmtmo to
ths saxpchroatosanaa or polysoay and aitgg«stsd M t a »
supwrwsMrsry for tha aa«s dsfinition i^ieh dsaa aat
8 8
prefttrabitt* 8tudi«» of thi tsaitotle ehroniosflaBa* which
«re always norv f cvou]ra»>itt for norphoXogical woik^
hvm vidansa ttt« ccntsoraray taatwan th« mlicvoehniaoaaBaa
or at]par»»i«rcry and naeroehrotaoaoiaaa by •liminatliig
t!ie £o«ni^ from the eateoory of chr^nosoaea* According
to thia theae oic^&ta* by their apparont origlQ*
atruetura* baha^ioor fu£M5%i<»a» do not f i t i^y
aco^ptftd difiiiitioii of e»y kind of cfhrosnoaoana* »a«fC30faar
U9S7} whiia th@ enitotio diroooaoaiaa of the
dooiaatic fowl piopoaad tiie taxsi chrK^soaoiaa for tSiaaa
elesanta. hea diacuaaea i^e opparaot origiii* »truetum»
behaviour and fuiMStion of the chronoaomoida eceording to
which the wore «caiitrio« aiiooyoiic ih A origin and behcvictir* variably heteroehn^tic and shoeed
wide verlatlon in ntiabar* l^ey ei>parently form
chrosoeentric hetarochioaMtifi and ara progreaaiveXy
dimlniahad during the raitotie wd oiaiotio oyclea* Ttm
noat piauaibla thaory for thoir function which ia atqpportad
by eyteiogieai avidanea aara that thay eonatituta
raaarvaa of noeiaie aeida for the chroiaoaon» ayntbaaia
o|ela and ara utiiiaad aa auieh. iseording to ttiia
viav thay ahouid not be givvh tha atatua of ehromoao'aaa.
^ SlUai . Coimally and ntweonRiar ( i m ) uaing autoradiography with tritiatad thymidiaa* found
3 9
that tlMi amaXlt ehiweeews SKA
Chiift l^stgut Vh^ toek thU « • tlift ttiriiliaK*
for tlM B e w t u w s w l •t«tti« of CM clivoiioMiM**
•eUnid CltiS) aUo fotnitf that DKik syaflMsis ia
<iftiiw< 4 0 — t e n i i i i A t w l ••rltor in t l » MiaU.«»
than in the Ooly ciitl^ ehteweow
in te f vnti^s* tli« ifc-etireMMCMi** w** i«ti» ia
Cttltaitea t aK>»dbiify Ci»«fl
ffiund • aif $» mm •ya^mii* imtmm tho
Md aii«3li«r ^htmmmmm* Bat contrary to othor
fifidiagv tiMi « i « l l « r t^mwrnmimB mm ImtMt in n^lleiitii^
in this ftic wuehrewew, which ia r a ^ r iarga
in liyia* iraa aiao lata in iipliaatiag*
^ M i m dtoawiaUauajaianchi and Maiiaa
aiada an important aaaXysis of tha owt uptiAo daring
fcmr dafiatd parleda ^ i«f»lMtaa. Sm tlia aaiAiaat •^paried
l a^ i i ing of tlie oioneehromaaaa aaaawd to ba of tlia
aiMa itttanai^ aa tlia laMlinff of tha •agroefcrawaaawaa*
fha liiohar tha rnmkmtt of I ^ U a d •ieieehrawaama tho
ffaatar waa tha lahaUiat of tha eaeroehiwaeaoeas*
M thia poriad thara naro alao aaaia rofieoa of mgneoehnK
aaaaa that vara atioagly lahaXlad. Maxim Uhali iA«
waa roaahad aiiMitaaaouaiy dtiring th# aaeood poriod ia
a a
teth aleMetiio«o«o»M and ii*«ieclimws6*eee Xa tlw
thiird p«rM •ow of ths aioreelurwiMcmi* and parts -wi
wcrecluEowwa ttcHpjUM tlMljr DMA «y»tli«*is* IJuiiag
tlM last pmwM thmrm was no laibsiliiig In tlis fldoiws
•3iei«>t for ths prasuese iMato^psa— in ths Csmalss* Sa
tlM naeroehKOMseass apseifie csffioas mum
this tfstaUstf m ly s i s s i sha syntlissls in tUs
siNMitie oslls of flyisui ftasmsis. ialicatss tlist thsM
is as elssr dif famities in mh ayathssis bstussa
tmoMiochtPomnfomtm sad aicaPsetiEi isafliass* Oaly ths Vmeiigom0»0i»t£
•mA.mm iQ>scifie safieas of asafaehfeassaaMs a>a
ffathss lata ia capiiaatiag tout ttis iMBliraimas dass aat
diffar aueh ffnai thtt s-ahcaiiasciis or tha autassnss ia
this rana^et^
During asiosis ia Qailm^ daaiastiatts^ fOrd aad
mtAfm Cl9i4} wiaiysad tha Mvalaats ai^aeyiag both
Jkilptt sad alaatanm aicrascspy. thsr fooad that at
taahytaas thara is as gm ia sisa bstaaaa aaarachfaassoaas
aad aiatashiaasi ansa* «hara is a caatiaasus sarias
f ra« tha Xar«sst to tha saaUast MLralaata* la Ilia
stractHta aaalyssd hy aiaetcaa aiaraacspy thay fouad aa
dif fatsaea bstaasa tha lar«s aad aaall chraaasonss. fha
prasaaca af twa* aoca ar lass distiast sisa irrai«>a a€
tittiPomwmeee in of ISift t p imit hitfi thus
tli» thiiaaftg thmt timtm tdimX^ tm ftanetioiitl
4if f4MNiiiais imtm^n iargp aaA duromsoiMM.
nmmmw» ar* AHpiMfitA AOftiiuit «lii« opinion*
fh* Qn>iip» of iMMraMn m mtmpvlym
l>i»d upwelAB to otiiy aiff«c«iit
IQ tiM fKafyoirOtleai sttiaitts i t b m
ic^ni i^mt tim nmSmvm o£ «li« eieeaif«o»»«« iigd» pttioay^
uaA ^ft^ms^ -fflg^ylli t inw ««!iy slisilacitisa
in tiMi&r «itl)
•»oii9 as «• iiith Vtm ott3«r of
«li« mmm o M w ir^portea tor otiMir Similer i *
tn* e m wim trtPfffftliiM Bsm^'tfgfi'ffliiw ^fitfypffit^f
iihieii Mlmm to mo eraor M l i f tpv i * * Muny apoei** f fwi this oraor ti«i«w mm mAfmA Umnf^wpUtiiiif md M mm ii«fv« • e%mm wmmAmBm im wmijt cHromMs* i^Mtoflfonipa i « tlie i «r#i«t o r ^ eei^riaiiMi iioariy
>>
SOU ^ th« bird gpmim mtuAfmA m f«r* €hwmommtSL Hiimmnma haww bMmriq^orM Imtmm mmsSmra of the
e r t e MoB9ii»9 to difforont fwiiiioo* tho
^ fiUXH. found to too <|ttito dif foioiit ttm
«fith th« tmti&mtB of tlMi « « « • f«iiiUy in
sica . revMlilMeiCMi .««•• ebs«rv»a between thMi
<flMn»r« 19661 • Ihe chronosemett of teaegr <yowticu«
l tAmrimm hrunicopa aftt very sisiUer in aoxphology
only ainute diffecencea l3*t»M» the tuo OR other
smbcrfi e£ the femily im diecuseeo earlier*
A re^tarkeble eiaidlerity isr a obaervea in
leag^ of aex-^tironoaoma* Zm hrcimaosm in ell tiie
apcNSi a atttdiea fourth largest pair* I^mv other
^orlcers in aifCcresit apceies reportf^d the a«m poaition
of the Dif fcrencea in tiie poaitiona oi
e@»trofarrea in tho chrDnoaoriea in aiCferetit apf»eiea may
be ea^iained tht» baaie of pericentric ifiveraiona
taking place during the evolutionary proeeaa*
M similarity of « r i « ) karyotS!pes was aiStroeated by Oliae and his Co«wettera on the baaia of dirfct karyotype analyaea (steaiua et* al*. 196if Otam, et* al*« 1964) and the imiform g mom aise in diverae ordera oi birds (Atkina et* al** i9€Sf 8ach«efm et* al** 1971)* tti iBar (1966* 1970) uee^iveeally dsnonatrated that eloaely related bird ^peciea often have identical karyotype thouQlh eaiceptiona are not rare (Ray-Chaiadhiiri et* al**19i9)*
i)3
Jttiaiysing thm nveUable a«t«. Bloom (1969) showed th«t
the mo<3o of diploid ehronoton* ouiAbtr v«a 60 in 91
«vi«ii fheifeitg» warn Sa to 94 with 77 p «r c«lit
of the chswaosoffie ataoter bolng in the to 64 r«nQ8*
Zo the present studies nseiBhleence tn lesryotypes
of eii the six i ^c ies foimd in so far as dl the
flpcseies %ihich belong to thcee diffvxent orders hoive
t«io ^ ^ s of chromosomes Hvb raactochsofflosoms and itio
foievochjcofaosoines. the f irst 10 • 13 pairs were found
to be rascrochromoaomes end the remaining as aiicfochroiaoscsaies*
They heve similar chfciaosomB morphology felth s l i ^ t
variation and the sex-ideteroining laecfhani^ in all the
i^oiea was found to be ^S/mi type* S^hromosoiie in all
the species studied was foimd to be the 4th largest*
Sven thou^ there was a close reseealalance batwesci
the species of different orders* i t was noted that the
nsmbers of one order or family show tmOh more sinUarity
of karyotype anong themsalvea «Kan betwean aembers of
other ordsrs of fMilies* for exaeple the karyotype
of lurdea purpurfft mattitlensif wa* found to be closer
to thst of Ardsola qrsvf^ oravii and other Midbers o£ the satae order (studied by other workers) than to the
of the M U i S m i ^ namely gfljaiffff
(»r*Mint a»llu« dcwticua^ Cpturniit
a;otmfniic laooaiicia (studied by othar workers)* 8i» i l «r ly
tpt^fBpfl^tVt mow clomtXy in its karyotype to raemters of its own
ordsr th«o to ths m ^ r s of othsr ordsrs* Ziilisirise
in Pssserifom^s, Corvas salsadsii^ o£ the fmi ly
Corvidse sliotra more idsntity with g ce Dicflt (Rsnvadrf
m amaibef of th« seoie fsoiiy* thsn to ^ ms&mro of th«
f emily 91oeeidae« gigaSL l^^yf^^ffuffi an^ S ^ r l l f l W f f l l ^ f f
show cioss simiXftTity anon? themseivcss* T{^rs£oxo#
ths syst&iMitie oisssificetion of di££sf«nt s,^«cies of
birds i » different taxsnoiaic gco^s< on t!tm basis of
a!»3 anstomicei strueture is iti
with that of the classification made on the basis of
their karyotype studies*
results discussed clearly indicate t^at
ooiqparative karyological studies are very helpful for
^ class if ieaticc of birds afia can be used as a
reliable tool for the s to^ of phylogeny of birds*
im lC«ryotrp« •bowing thm hooole^^u* pair* o£ thti m»mm%tQ meeochKtmomatmB o£ Atfdiiifc oug!Dur»|| miitlfMlf C'Mitttsii piuepJl* nmrm}^
figm Mst@i>hage pla^s shoving tt)e «ct:aatie durowsomcs of figagtt nuroftiwift g^^onaia (Enatttcn i»\icp|e hrnxmim
II ••
12
mA 8
4
FIG I (a)
10
FIG 1(b )
Kicywwn ^MMIimi em^ mSLt^tium Of ^ imm^ mrnmtmtfmmmmsi
g^ f f ^ l l j yy t tMBf yn^y^ i i^ iwmmmm
PERCENT relative LENGTH CLR)
i L
to
0 1 (Ji
CT fO
X o
Cf>
<o
D
K>
Jtik
w$i§m i mmfmmrn tibmfim ^ imw me t M i i i l ^ ^ ims mni rn^mt 9mm « f sti^tx^ McsfCiti i&'imwiMit ^ JWUUESIb
PEPCfNT RELATIVE LENGTH CLR5 LONG A P M SHORT A RM
JL_ I - .1 JL -L J- J L JL. 35
M
N N
O X JD
CDO (n WQ S m. Z O
Ol
a>
o
wim MMrrviyp* i(it«i»iii9 ^ tmrnrnttm
wUraMMIMMMI jyBtaJll fOMdiyL jMHMy f3MUm wmi %mmfS — - —
I f II II II
12
( Z Z )
8
F I G ^ Cca.)
tj* J
i0 5
10
F I Q ^ ( B )
t i i t i rnmrn^ tUmim ^ taiiwtiiM Qif tiM mam^t^ •MMtracliiWMMHinMi i f iMiitaia atmtU m x i l
PERCENT RELATIVE LENGTHCLR ) o'ro-^ COD o w ^ o S a J o r ^ ^ w
I I _l L L J 1 I t I 1 I
a N
O X
QQ •Jim
Z o
o>
CD
O
to
t i i * i mmnfm mm l O m ^ ^ P ^ emM mi tmmm imiillw ttMi I« i9 md siiort mm mm mmtmim 'mmmtmitmm mmB «f
PERCENT RELAT IVE L E N G T H CLR-) LONG Af?M SHORT ARM
I l _ J L - -1 I J -1- - J I L 1 L _ J I
U) Nj N
tn O X XI <T> o
112
2 m ao 2 O <o
O
M
Wi0m t « KKwr9krpm tHimimg «3i« t i m t i pmim WfliUie iMiemftims^ aiqeMMi (Ml
I
I s
0 5
i #
8
Cz^)
t i j i m 4 • %
9 fO
FIG 7a
w «
- • ' A • '
FIG 7b
«f tii« ammtirn wmmt^mrnmrnm ** ^ lESBMyttlF jfiSfflr
fO
(ti
N N
o X
OB O r m 2 O
cn
CT>
CD
V>
O
PERCENT RELATIVE LENGTHCLr)
J L O -to- w
J I... - J I I U — J
• mwmmm ^tmim tiw pm t«tt« wtO^Mm «f til* l<m$ m0 iittyt auMi
^ MKfttic of jhEwyaAigwn wmmEmR
PERCENT R E L A T I V E L E N G T H C L R D L O N G A R M S H O R T A R M
I I I I I I ' I I I I I I I 1 1 1 1
K>
O)
(O
at
o
O s moj Z o <o 5
f 1 0 • Krntymm iN«ilill9 til* HiMMlttfCMi fii»iir« ^ »m»%i€ ammdtutmiotmm ^ f^g^tt*. imrntrn mrn^m
FMh tm y^KpHese jfimm mmms «Imi AcMMtt dhemmmm ^ eagtmumM^M. HPiiimiiH iMiMiwv
iU - (zz)
1 a ft I 2 3 4 5
t j f i m 6 7 6 9 10
n
FIG 10a
FIG 10b
tt »ti<»riii^ ^ pmf mm m^trnm Icngilui of ^ mmuiMm
iVkmtm mm, m}m
0 1
N N
oi
z O oe
<D
PEPCfNT PELATiVE LENGTH ( L r ) 0 ^ cr> CP
1 . . . 1 1 _ . X ™ — L . , — 1 . , 1 - I " - ' J ~ I o M X 5 g K K S
NJ
U)
t l i f « I t Kagyagg i mumisrn 9 t r «!ciit v^gtiim of tti» Iceig «M
18 at •<niiitia tur(MM3s<iM« «f Gmmm m^mm^ttrn. irnmum mtm)m —
iOo
PEPCENT R E L A T I V E LE .NGTHCLf t" ) L O N G ARM S H O R T A R M
0> NJ O 03 Oi ^ NJ O NJ -t» Ol CP I 1- J I I L
O w -K o> J I 1 1 U-J 1 L_-J 1
KJ CO
N N" O X
s i CDQ _ V (OO s m
2 O
Oi
CD
<o
« f mmmti^ JHMlifllft
yiftt tai imrnihMm pi«t«i( 4iMwiB9 ^ • M i i i f dtmmmmm fnfiiir liiititiiMi
1 0 7
M
6
2
7
(2Z1
I I 3 4 5
m
8 t
9 10 FIG I 3»
FIG 13b
Si ~ Kttywan gNMiti^ p w C M t«iMftiw «f ^ mmg^i^ mtmmSmm&mmm ^ i M M I i S H M l M l ' * ' * ^
o t—, PERCENT R E L A T I V E L E N G T H CLR)
•u cT) 00 K E 5) 55 O K> J 1 — - I — I I
O)
X N
x> ^ O 2
ctO _ w ^ O
z m
tn
CD
8
2 O
00
<o
pmw mm fii««l!«<» «f tli» lon^ «!•« •iMwt aiMi
flMi tfiaMtlcr HtecroeiiramraMis <if I M M IHflfifill^g (iioii*» •piwrcwU
P E R C E N T R E L A T I V E L E N G T H CLR 1 L O N G A R M S H O R T A P M
jp' fr . I . i I I I • • > . P
Ok)
0 1 D o s
Ocn o u;s
ai
Z o
«o
o
Fif* « Ktryotyptt slioitln^ tiMi IMMIO^us pmUm «f th« mmmtic m atceihtomemmmm oi jifeMriffili ptvmissftim (MtosuM buetiag),
duroiioao^* of &aite»ri»i ^ayiytoftn (ll«ah«ad«(l buetiil9i>«
U l i I
•11 6
i i 3
8
fzz)
4 •
10
FIG I6b
t i « . I t K«rfieir«ii nhemim i>«r c m t ir« l «tt ir« of MttorochvcmoiiaiMis
pwfaMfi** lyuaQ^ ifyf, banting) <
P E R C E N T R E L A T f V t L E N G T H a R ) ik ar> OB
J I I — S M X 5»
J I I L _ I . N> K» M M p r ( t r
fO
iti
f m n w w o m m aSkemim th* p^r € « a t m X m U ^ %mm0lm «if tit* leaf ^ i^iart t m ^
itie niiaR»eliiono«oiM»« «f ji^tfLpg, UNiiNittM
ii2
PERCENT R E L A T I V E LENGTH a W LONG A R M SHOAT A R M
Sf> SfoOQb Oi-t'NiO ro - U o j o o o f o ^ m I I I I 1 _1 I 1 I I I I U—i L 1 1 1
lo
to
N N
0 1 OI
- n O
00 "1 Z o «
(O
i » S i i t t B 0f WtiM dwitMni llfiVPfl
I i o
o PERCEN T RELATIVB LUN6TH CLR 3
f r K) ^ Ol CO f3 Ji O 00 I. ! I. : 1 (:.• I. . . t.. ..J :-f-J
t i ^ 10 httHfoqem itiowlfig tise p*r evnt of thtt lon^ sikS sh^rt •cna of the
itie waroduweBsewe of mUSSMM " i l l . i<Mit«rti pvaplm i ^ M i S
SESXHL poaA h«ioa} mi If i>«*tri3gi7^
1 1 4
Pt-ttC£NT R f c l A r i V E L F N O T H CLR )
5 £
l.«ii9th« of am «ofSfttie ai«cscoelico-n> «Qitt«»a of imnm cmam}^ t i i i i£ tiywl^f TKs i i iFamw) iii«l I ^ B b raiBSK ibuoUngf).
l i l i
P E R C E N T R E L A T I V E L E N G T H C l R OB ,8 ,K
03 O
tt ksryogmm vhttidtnt tii« per cant rttitttive Ittn^iiw of loitg ilmit
the nai iPoetMr ia oim* of ftlffiMiftS, argw)* f a y f r ^ momm ipi^cikm
l i G
P E R C E N T R E L A T I V E L E N G T H a S " ) T - n s c r x j ^ ^ ' . r ^ ^ Cr> 05 O J^
^ C h^ ajL Q ff 8
I i )
1 1 /
Urn diploid chgoiBMOrtia in six €>f txLvds
f com thu Carinattt* vis« pmgpugea
rnaackilmtmf Nidmolm gravii pyv i i tiie erdcv
cieonifoniftsi fffftfmiytyj ggmtfUyfriaavB mmmim tMaongiog to ord«r QiliiforQ»« and •olgfid»n»^
£aiSfi£ tfOTti^^c^ ^ml f f f g i of th« ocdsr
P««««riiort!i@a verifis c»n«id*riibiy* tlie m m diploid
ehxooeaosiis mm&mv in the six ^seies found to hm
61.44 4 «3*«0 • 0.688. 76.34 • 0.623* 74.960 •
0.S76. 76*7Stt} • 0.S7S a ^ 76.400 4 0.S63 c«ii>ectively. /
Bes«d OB typical svisn ducmoso^sf tit* tm types*
tlift maiBgodtwGmosomB and the miexoclixo<aofiOiaBS» hevo Immo
in al l the spsoios imdsr s tu^ . First 10 • 12
pairs of chiioaoso^s idantified a» maoKoehsoisosoaes
including s«3i-<di«oaioso:!ias and the vest as eiesochfoaioaoaies.
maoffochroiaosoraes are classified into different
groups on the baa la of their »ean relative lengths and
ceatroneric iadieea.
« se«»Ghrenoaones in laalea ooeisiat of a pair of
chroaoso(8*s with variation in their eentfomaric position
and arc the fourth largeat nheieaa in femalea ZMf sex*
chrooosooes hcnre be«n ofcaerved. lha WiChroinosoiiie is
ssall in sise and oecupiea 7tli or 8 th position in order
of length In different species.
fh* coac>«r«tiir« karyologicfti stii4i«« «ni iracy
li*lp£uX for th« elft««ific«tloii of biffd* and e«ii
im uMd at « Mttfalo tool for th« atud^ of
phyJLe9«iiy of blra*.
i i C
1 1 9
^•••lu iPwl filttiii*
ci»ifl « ! • SMA «oa«iiit « f oiDttri 19 of fflaeastal MptU« » wdt birds* CUtoMma
jm* 8* mii itmi
iirci||ii# r*s« i m i l
horn «C til* of tnOim OMI W * S* ONfCHPd*
I«ee«liBotton of iiotofoeiunsiiioUa tn hmmn ehraeaooto* Cyto^pnotlco# lOf
im (1975)
AttiMMrit ikm mtfrni
mA sw
( a m i
SdvaUf leatioo of oo»-ehraaoooMto in tlMi bold ooi^o* €«&• Otnot* Cytol* itt S»7*t»t«
tauftic etumosoHBs o£ Ooidiii ovMm COrif^ttf fftfiiit* »aii irHeiii irrooCe ti«t* Msad* eei«p Indiot 1 •
A ottidfy of <$!« dttoaoaewMi in th« f amiiy 8tttroite C a v o o i r«»MnrifonM»l« iroe. ii«t*
Sei« iy«Si«it ZXf S01»l0i«
frtfMiaooation ti«toffo«y9ositr ia Hio bird Iiontmra nttBBtm«ta<Lion» I < n e c * i W i « S s f iSCNsSarWi * tMtff0 i (a }t S M S .
Aaurit ( I W ) •na Kaiii. Q.
Zawrsion poiyHoupliiMi in c o h k n s i gr»«s pi«MMi« Wif l f t dtiaaMfflBSKS
iirssevipiB* J T o w M E i T
^ UMO)
ma Moiiaii# 0.J*
m of Indian l i tt lo
7t T M i *
CStzeeelw «mS Fottnmo of M^i i t -otion in boM aarsow of i i l l M dBMiy«l||*
iiii
nmm (ifif^) A «iflrt«ftt lut ff «lHr«i
( SMU)
ftilslMfira* Ctfttl
I I M M i * 0«S«
MfeMTs v«rt«*Sew ffmr flvtiii SMlNelew c^riiistii**
iHM le ctircMUiiw ci M l u nmtmik S6U>i S W i *
Of tiw faittite mimm m&e^ ^ « t 6ei«» mn morntm m uvmtmi
lUf l i iYi } p i t t i t e
l i t t i l •HA »i«l0ir« M^m,
OMkiMI lltffl'ltlii CSIMnMMMMINi
M i y a i t of 1isr»Dtam*# upmUB «c f « i i l l r fiH^i«Mi
Cum}* emviiogkmt aiia>»
f j ^ l i t On tH* stwrilitir of teMta •Bd tlM M a C w i M fttiA* f t i i t ^ i *
Cfvw^ y.iwi* ( t » i i>
C«ittarl«J««f#> f lMt ) Ouristiiii* ImC.
Oiat%lo«I «ii4 Cytol^oiieal 9timm of M ftiitiit9iaftrl« ii|lMria» « t liliifcilgfiwMliUfcfc Aqv* 809* RCBBreG iS i
K«icy»im wf four « f huAf m umi f M a t M mA H/fUma* mn^u m% iM^sti*
cmim^h fiiiB* c m i ) N iMMi fo I *
MUk «f _ „ _ , MlUn* tiiH Awl** m s i s » a ]
i M a l i y * «»il. IIMS) llMTV'tllS PftttMTM i
f « id« M l HMm.
mm mmrntU
J. JL
tn^itmmm ! . « i i m i Mixjliy* 4m 4l»*f
l«»|inaMi#
I9i(irimn« ll*t« itfii CStnl lioAtrlOi*
( tMI )
Mil M tviAuiM d* l a •tnaitiiffs fin* aM «lMnMiioMn»» lMM«ia» p«r «i«Mtioii • l y i a t i e (icooaM M ptfrtieulter) C«ll« m&lU acl« (»«rj;»li m t W1UM.
Ctawwpgiafiwi iNoiaiiig pattiffiui of tiio p«r«e«atrle ^iirMrsioot la
CyteoMMitie* «Bd C«U Oiattie^ afttii i«sia«
contrlbtttioii • UtMm ^ esryo^Bse elim Im WovOmU M i w
S,ti»ik. CitM) NaoUMiRK, O.fi.11.
tii«tteiil«r chmMoaoMs of flilt 1 tfHtr III sltffwi;
(|»09> mm of th« « « U e eMciEMi* <fl||lM tHl|H|
fmmU 24$
Mm Ut i t ) sttidlM on th« eHmmMmmmm ot mm rnmmtiic emmom foid •• •Mm in maa «ilMryo»«
M U * Sli MSMrnm
MtoiM M* «Bd itfM OWi ftmpnismum p«tt«rM «f tki» MMNClironoscraM of tlM
ti a t M M *
nmnrnm^ i m i ) Chick ehr<
i#f UM9M* MMit*
K m * *
(1924) tlM iMi»tl<i eiURMDMn»« of tiMI vfilek Md tliwir pwlMle
aelAoe^ 4a4«4tS.
(ttlio) A cMiparlslen of oitooU In oliloli tlmuo oidtiuroo oaA Ia •08tlOM« OMteyOO* WMm maim, set t9 i» lS i .
Hem* M*
l l «M* Mm
Mutt*
ttiwir» H*
mirnqiaimt, O.
( l i l M
imM
(1949)
imoi
itmrn)
^mgmGmu of His ehUk ata3l« itt 44l«>«4i*
S«K «ad th» chroiiosoaM in ttm mmmtHMs foul ( 2 i l l j « flgMitSHMils
^our. Hei)^* 4fi
omt les irf tiM foiii, McMlr XcmbII Miv Havlie
hirdmrn ^ Ictfyo^pM* of nixm iSt
Karyotypes of l i birOs* flmtfitat* 44(9)1
l ^ f «r«atlal •talntfig «f polyttM duromOMi bsml* in te aiiutt baiiaing Mthodv* mfornoBoarn, 9»»
bttodiiM in po lytm ctiKoato*
S T ' c S J l ^ f S I . - ^
( i m ) CftiaraotwriBation of luitMoehrai* •tia in jgtiistocwea c—caria ky and alTlbSffia StSoaSr Clieweewk 70Ca)i I)9»IS0.
( i n t ) ma uMliaaiMi of OAeadingi Saparlaatiaa bata—liaiwatiaa*
n{z)t aos»aa4.
Ztoh« N,! Zkauehi <lMf) t,f«iliiil»a« Hit Nori* N.I N. and iiidiiaa# « «
Clfii }
A eenHrativa karyatypa attidy ia foartaap %>aalac « f birds. Jioaa* Qraaties^ 44(1) • tii-119.
•tnietura sad baligvioar at in D X jmaiuB baQdjy^
I m (Uan.) (PlSBSirBBStiiiee)* Itaa; MatU M9ad« 8«i«t lad&a*
14i»l4i«
smmmm, <itiiii) m^fmtj/gmm of two Korth iHurlciMi buBtinos Ii«>*ri«|ll«tt)* OMWtifit
JSKNTMrlo* v. Mkias* li.
( S i m ) lUiryotKM* cf ieiir Pmcr iM Moagiag to th*
Nladltfiil «Bft Conriaot, ChroMMMMb 2«t
KliMSb II.
ilMjpMfttogeMsi* of th» aoMttio turlioy* coll* lt««9t
(litOl Mitotic ^romoMMieo o€ tMi Aia»troXi«Ki tciag qtsoil (
m i j . c
C and tmtifU.
{if7S) Sooifttie chrowsoaie* of GBitio Aeeteitor Humd* WTml^Mr^
Jowavi* lUiW
CnpM N*
l i m ) ClffOODSOOt OtHdiOS ill thXOA •poeitts of »ieifof«es Ciiem)*
iSMIfl iiMstMurehtttt 8ur Im amloptanit ovoriott dM otyf« oarokiloitiw M S • ¥ic»llu« not ritif momOmtrn Arch. Mwt« miMgm*t aSK
(1910) Mm vorUtioB du Mfeto dM ObmmotM d«M oo^ewatatien dt ti* o»iif aoA fmtmnOm a* Im ooglo, C.R* 800. Bioi*« i i l
iovan*
amdberg, iwA. (iti4) NooMMioturo for coiitioiMffio
pocltieo on chcoMiooeiiiis* Nmditot ( ted ) l i t
1 L ^
Xitieiciii* %r«N» Mor««saiii* M«lli tfid A.
(1975) S4»nti€icatioii of p«chyt«M bivalmta in liuin«ei oMiiosis using CM&imding tachniQtJMi* ChronoMa. S3U)t 275-280.
tmdkmxt, e.M* (1976) Ifw soaatie chxcmoaom cwplm* msnts of 16 fpscics of Vftleoni^ fermB (n e f ) m^ karyolo* gicai nltttiootfhlps of tii« ordwr* Oftntttie«# 461 77-113*
Min«rt KK (1938) apermatogtMftis in a sex reversed female and in oonaai nales of the domeatic foiA
Atiat« Res.*
Marino* s« (l94Ui) Cytologieal imrestigations of Aic« included in tite genus Hus« Jour* Fac* 8c i* Hokki do Jn *
Sur. VX ^>ol* 7t30S-360.
HaHino* s.
^ i n o « S*
Nilcino* 8.
(1941b) Karyological study of gold f i ^ of J«;>an* Cytologia* 12 • 96*ill«
(1942a) Polyploidy in animal <In Japanese}« Sot. md 2ool*« 101 163-172*
(1942b) Chrocaosome co^parision betMeea the wild form and the donestio*
of^Bsts^aEaoEeatoE (Berlcenhout). Jour. Genet.« 161 SS.SS.
(1942c) Studies on the reurina ehroniosoaies SZ. Morphological coa|»arision of the chroaosonas between the wild fom and the doiaasticated variety of Rattus ooryeaicus (Berlcenhout). Jour. rae. sci. HMkaido 2i|>. ttiiv.« 5er.« VI. Zool.« 6i 31-43.
i IJ
(i943«> atttditts on the nurine chroaosoms X2Z« A coa|p«r«tive study of chro<» mommm in five spccifts ef RetSSB* Jour* F«e« Sci« Hokkaido lap* iaBiv«« Ser« VX« 3ool» 9$ t9»57«
MakinOf S, < 1943b) ttio ehromoAotiiea of th« liorM CBoiwy, cab«ila»)« chroEaosomt studies aHctoaTstSc raapnals X» Cytologis* 131 26*38.
Mcdcitio* a* (t943e> chrooiosoRie co{qpXex09 in go«t C£sB£B llilESMl shttcp (gslft
And thsir reletionsnl^ fenmaosomB stiadi«s in doaiestie mmmelm XX )• CytoiogiAt 13 • 39-S4*
{l944a) Hie chiroaxjsaas co3i$»ieis o€ tlis pig mtlff^rn Cbrmo»mm studies in doraestic Q9««Dmals XXZ Cytoiogiat 13«
llalciao* Bm (194«b} Reryot^es of doses tie cattle* satlaa and dotfi^stie water Buffalo* {ChgosiotKsm studies in dotaestic raitfs^s VZ)» Cytoiogia* 13*
Makino* 8*
Meiandert Y.
(1944e) Chromoscsmss of do^stic mamals* <Xfi Japanese)* siool* Mag* S6i 6*1S*
<1930) Accessory chrofaosomes ih anlnals* specially in M3S|ltiE. ISfiillE* Hereditaa* 34i 19«36*
Merrick* 8*i (1973) l*adley« ii*s* and LuiM* H*A*
vroduetion of 0» and C^bandlng with progressive trypsin treet-«snt* M i a t * Res*, 71 39*44*
Misra* N« and <197€) srivastava* m«d* l *
Sooatic aKroaoacysas of Bubulcos 4bis (It) (CatUe egret) i A case STmlproeal translocation* Osnetieet 44• 1SS*140*
m
Motera* ff«A« (1977> •Bd ftai«
Chromo»oaal differentiation in tMO apeeiaa of mamrn md thvir h ^ i d * »ev*eledW<3i«wa 0» banding* Chronosoma* €4CS)t iai»|32*
llot«ra# <1976) and »ai» ie«8.
8aiicoM« (1954) and Smmt,
Oianaa c •• banding ^attama in Aadaa (atagooyia) moaguitoea* cHmbaoaat 70(i)t Si-S8,
^MBrraatogeneaie in dooeatie foul* Jour, 451 79-S7,
rnmmme, S.ti* (I9S7) Ihe aitotie tShstmntoma of tha -dotaeatlo fowl* ll«rad»* 48I 227*334«
aa«co3»r» £«H» (I9S9) ^ Bieiotic chsomaomm of tha Cytoiogia* 241 401*410.
U960) Chicliati^tuileey hybrida. ^our* ii»rad«« Sii
Sax^hrotaoaotiea end micro*
Omo, a*f (t964) staniua* C«i Christian* I*«C«r Bac^» w. and laaak* M«U
Oiiaii« a.J.T. (194$)
Cbrf^aomal laiiformity in fhm anrian aub-cla»« Carinatata, ChroooaoBia (BtrX*) IS* 280»a8&.
Karyotyp* atudiaa on SiUllft dcia»aticua, Chromoaoraa |larX*}« Itt i n - i08.
»on«in« <r« (1942) Mraonal oo^onunioation
(1942) Cbaanrationa on tlia chxonoao* ••a of turkay^hicican tiybrida. Jour. Harad,« S4(3}f }i0i«102.
iTi
Plceiimi* t , a&a (1970) ^MXm* N.
Socw «rl«ii luiryogri^«
Santiiii* o* and (1976) Otsraeterisation ctf
hKterociuroatttiiu mm banafna* chrofiiotoa«f 97I »77*S6«,
ft* and (1977) x«ryot]n?«» of f iv « P^uMMrins birds b«longing to f w U y l>loe«id«iiB* Ceryolo0i4i» 30 O ) i
PolloOc* D.Ii» ttiid (i97e) FaehlwIinBr* ii«6«
9li« ehrotiioi»tii»8 of eoelMrela
«M&0»iS. CVtOdWMU OeiMt, 211 3«7«aei.
(1963ft) A eoaparfttlV€ gtu^ o£ fiioft<iuito )c«ryotyp9s» Amu Snt* soe* Asier.* 561 160-170*
(1963b) A cori^arativa attidy of raosqulto karyotypes* Caryoiogift* 171 S96»607*
(1964«) Cytogenetic effect of ehaaoeterHents in laoaquitoea, I Dpholftte Induced iJberr«t» ions In the flomatie chroao-
^ ^ o ^ a f d t t l O i i *
RaiA K*8* (1964b)
/ ' ' f
/ kmU K«8« (196S)
Cytogeiietio effect of chenoeter ilants in noequitoes, ZX MBchaniom of aplielata induced ehangea in fecundity and fertUity of a t ^
( D * Biol* iuia;*. 119*131*
Cytogenetiea of chemoatart* lanta iadtiead atarility in
J5: sssi^r Zntematiooal Con^aaa of BntooioXogy* lAiidon«8*16 «7uly* 1964(Proeaadinga)* Royal mtorn-elogical Soeiaty of London pp. afi»as6.
'} o X 0
ttmU (1966) Furthtr ctomrrmUmrn on tlw •otaatle ehromoAOia*** CytoXogf of soM inosi|ttito«« CDiptM:«i CuilcTidatt), Aim. Snto« soe*
SOI 26a » 266.
(i967a) iDftchoiqutts for th« study of 9i»iMitie» and CjftogVMties of v«etors* Chapter 23t 679»70i* in of Znsaet Vectors of «iri0ht and Palt a** aditora £laaviar Publiahing Cog|>aii]ri ^ fiitdiftricDida*
Cytoganattea of ^S^m MflXBSt^ Bui i . m n i ^ m S r * 15• sSi-ses.
l^hniQiaea for atudytog tha af facta of radiation on taaioaia and raiatad procaaaea In moaquitoeo with particular rafaranca to / adaa aaovpti* t&3»200 in ^Mlacrof Radiation on Haiotlc Syata:iia*« ZntamationaX Atomic Enar^ Agancy ^ a a # Vianna« Auatria.
ftay-ciiaiadhuri* (1969) sharoat tm
and RayvChaudhiiri#
la4Jabli*8«Z,f (1970) i»aBov« and Bulatova* II* 8*
A cooparativa atudy of tha chroooao^a of biJif M Chromoaonia (fiarX*)« 26t 148*i66.
Coim>aratlva atuOiaa of tha Itaryotsfpaa of two cXoaaly ralatad ipaciaa of buntinga (BtiMtitiUL
iawocaph^f! on*) hSSHaad in araai of cfvarli«>* zooi* J»« 491 i6S7»lS63 ( in Ruaaian).
RtPiaii* Jacob* MU and Sharaa»
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Comparative cytoloyicai •tudy of l^afiapus folchicuo»
laBSESL^end cus> ChrooosoiQa
Saabri^t* H. A rapid banding tactmiqua for liuman ehronosaeaes* L«Qcat« lU 97U972*
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Hatsrochromatie natur« of ehrciniCMKms in birda* E^pt» call , ftaa.* 7t 39.44,
Stock, A«D«f Arri^hi, and st^tioa* K*
<1974} ChroaosoRw hosiology in birdai banding pattama of tha ohroanaomaa of tha doMatio chickon of tha ring-nsekad dowa and tha donsstie pigson* cytoganat, call* Oanst*. i3t
8tainigwr«ai« and (197$) Kukharjaa* A.B.
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