in the sheep , bu t that th ese di fferences were very small . The accuracy of vi sual e s ti ma ti on of chromo s ome l ength and p o s i t i on , as shmm in karyotypes , proved to be low at p o s i t ions b el m·r chr omos ome
7.
There was frequ ently di ssimi larity ir1 l ength b ehJeen homol ogues , especi ally in chromo some
1 .
Gimenez-Ma r t in and Lopez
-
Saez( 1966 )
found that the larg e s t chromos ome had a subm edian centromere , t h e s e cond chrom o s ome of almo s ttrw s ame l ength a subm edian o r medi an centrom ere and the thi rd; vri th a
median centromerc) d i ff ered in si z e by a greater amount than the difference ivhi ch exi s t ed b ehreen the fir s t hvo chromosome s .
and NcLaren
( 1 967 )
and R o ca and Rodero( 1 971 )
hm·rever , found tha t the s i z e o f chromo som e 2 uas mi dvray b etvreen that- of chromosom e1
and3 .
Dain( 1 972 )
claimed that th e long arms o f the fi rst and second pai r so f m e tacentri c chromo some s and the fourth pair vrhi ch a r e acro centri c ,
were longer in rams than i n ewe s , and di s cu s sed her theory tha t these
observati ons a r e c o nsistent vli th the p r e s ence of male-determining genes
on the long arms of the fir s t t1vo pair s of auto somes . Dain' s
o b s e rvations wer e n o t in agreement 1·ri th , an<1 no reference vras made by h er t o , the 1>10 rk of Bru
�
re( 1 966 )
o r B ru�
r e and l'kLar en( 1967 ) .
There was some variatj_ on b o th in position and morpho l ogy o f
s e condary cons t r i ct i ons . The maj ori ty v1e re found on aut os ome pai rs
1 ,
2 , 3
and4
and_ the X chromosom e . Half o f the recorded s e c ondary con s tri c ti ons o ccu rred on the three pai rs of metacentri c chromosomes .I t vms suggested
(
B ruh
e ,1966 )
that th e s e may be th e main nu c leolarorgani z er chrom o s omes in the sheep . Sub s equent evi den c e from
aut oradi ographi c and Gi emsa b anding s tu d i es has confirmed this suggesti on
(
Zartman and B ru�
r e ,1974 ) .
The various constrictions r e c o rded on the remainder of the a cro cen tric chromo some s did not suggest any means of di fferentiating these chromo s omes . Bru�
re and McLaren( 1967 )
noted an increase in the inci dence of s e condary c ons tri ctions when hypotoni c s o dium ci tra t e s o luti on was u sed ins t e a d o f Hanks balanced salt solution and suggested that thi s could be ass o ci a t ed vli th removal of cal cium from'"'
the m edium during the hypotonic treatmen t .
The c entromere posi tion was n ever seen v ery c l early i n ei ther the metacentric o r acro centri c chrom o s om e s , althou gh i t app eared t o b e more central i n chromo s ome
3
than i n1
or2
(
Bru�
re and llicLaren ,1 967 ) .
Bru
�
re and McLaren( 1 967 )
s tressed two morphologi cal feat�resin the karyo type of the sheep not emphasi z ed by previ ou s au thors ; firstly , the p res ence of c l ear, sh ort arms on th e X chromo s ome , and s econdly the su'ometacentri c nature of th e Y chromo some . The X
chromosome whi ch was the largest of the acro centric chromo somes had mu ch more c l early defined short arm s than those o ccas i ona l ly s e en on
the other a c rocentri c chromos ome s.
B1�
�
re(1 966 )
found that the modal diploid numb er o fchromo some s
( 54 )
was pres ent in87 . 44
p e r cent o f cells c ounted from a total of1 831 .
Later , in a compari son of chromo some counts o f22
normal sh eep wi th five fr eemar ti n sheep ,Bru�re ( 1 967 )
r ep o r ted a signi fi cant increas e in bo th hypomodal and hypermodal ce l l s "l'ii thincreasing
age .
He su gges t ed th at simi lar biologi cal m echanisms tothose whi ch contribu t e to age aneuploidy in man (Jacob s , C ourt B rovm and Dol l ,
1 961 )
could a l s o app ly to the sheep .Gerneke
( 1 967 )
found a diploid numb er o f54
chromo somes inthe Merino , Karaku l and Afrikaner breeds . H e <ms unab le to decide whi ch o f the acrocentri c c hromo somes was the X , bu t his des crip tion of the Y chr oJLo some wa s in agreement wi th that of Bru
�
re(1 96 6 ) .
Althou gh no features comparab le to the sat e ll i t es s e en in human acro cen tri c chromo some s ( Ferguson-Smith et
1 962 ;
Fergv.s on Smith and Handmaker ,1 961 ; 1 963 )
w ere record ed hy Bru�
re( 1 966 ) ,
what he interpreted as ass o ci ati ons b e tween acrocentr i c chr omo s omes were seen. As soci ati ons b etween acrocentri c and metacentri c
chromosomes Here also re co rded by Bru
�
re(1 96 6 )
but he c ould not dravr any c orrel afi ons between th ese as s o ciat ions and s e c ondaryconstri ctions . Ferguson-Smi th and Handmaker
( 1 963 )
found thatexcessive treatment 1-ri th hyp o tonic solution appeared to reduce the frequency of chromosome ass o ci a tions . Likev1i s e in the she ep , B ru
�
re( 1 9 6 6 )
suggested tha t the s a t e l l i t e associations in the sheep mi gh t b e more compl ex and de licat e than i n man and mi ght b e di s rupted by techni cal manipu lation .The diploi d chr omosome numb er , s ex chrom o some complement and chromosome morphology of the dome sti c sheep ( Ovi s ari e s
)
have b eene s tab li shed and are independent of the b reed studi ed . Using
conventi onal method s , i dent if i cation of the individual chromo some s in the karyotyp e i s di ffi cul t a s many are morphologi cally simi lar . I t i s
now po s sib l e t o identi fy every chromosome in the s heep' s complement u sing Quinacrine fluores cenc e t echni qu es
(
Evans , Buck land and Sumner ,1 973 ;
Hansen ,1 973 ,
a ) and Gi emsa ban ding techni ques ( Evans e t1 973 ;
Zar tman andBru�re , 1 974 ) .
Th ese techni cal advanc es shoul daid the de tection o f chromosomal abnormalities i n sheep . suggestion is di scusse d fu rther i n Chapter
Vo
I I . 2 TAXONOI'U C Al\TD EVOJ_,U'J'I OEARY IW.lESTI G AT I ONS IN SHEEP
This
Chromosome studie s have recent ly been applied to taxonomi c and evolu ti ona ry i nvestigations i n sheep ( Hurster and Beni r s chke ,
1 968 ;
Dain ,1 970 ;
Nadler ,1 971 ;
Nad l e r , Lay and Hassinger ,1 971 ;
Evans e t1 973 ;
Nadler , Ho ffmann and Vloolf,1 973) .
Di fferent t ypes of wi ld and dome s t i c sheep have been shovm to b e characteri z ed by di fferent diploi d chromosome numbers ( 2n ==54 , 55 , 5 6 , 57
and58 )
-.ri th in all cases a fundamen tal number or the numb e r of chromosome a rms ( Na tthey ,1 945 )
of60
(Nadler1 971 ; 1 973 ) .
Examinati on of two other speciesb elonging to the family Bovi d ae , the goa t hi rcus ) and the ox
( Bos as lre l l as
the
sheep has sh mm tha t although the chromosome numb e r is variab l e , the fundamental numb er ( N .F. )
is remarkab ly constant ,varying behreen
58
and 62 in th e ma .jori ty of species, (l·lurster a ndB enirschke ,
1 968 ) .
. This has b een regarded as good evidence tha t the chi ef mechani sm o f karyo typ e evolution in the Bovi dae is by Rob er tsoniantranslocation ( Vlu rster and Beni rschk e ,
1 968 ) .
This vi ew is suppor ted by Evans et al .( 1 973 )
who used modern ban ding techniqu8s to s tu dy chromosome homologi es as well as c entromeri c hetero chromatin i n the ox ,goa t and sheep . These authors consi der hovrever that band i ng t echniqu es
are not ab l e to gi ve an unequivocal indication of the direction of evolu tion ( Evans et
1 973 ) .
Nadler e t a l .
( 1 973 )
suggested t hat although Rob er tsonian changes du e to centric fusions b ehreen acrocentri c chromosomes migh t b e responsible for changes in karyotyp e i n sheep from 2n ==58
to 2n ==54,
i n certain popula tions centri c fission could have occurred to raise the chromosome number .I I .
3
SHEEP xGOAT
HYBRIDSThe p rob l em of hybri di sa tion of she ep and goa ts has int erest ed many vrorkers in the fi elds of a ni maJ. bre eding and reproductive physiology
(Hancock , McGovern and S tamp ,
1 968 ;
r1cGovern ,1 969) .
Inab i li t y toLIBRARY MASSEY U �-l:VERSITY.