Genetic analysis of growth and its relationship with lifetime production in Rambouillet Targhee and
Columbia ewes under range conditions of Montana
by Juan Francisco Chavez
A thesis submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in
Bilolgical Sciences
Montana State University
© Copyright by Juan Francisco Chavez (1986)
Abstract:
Growth characteristics of sheep from birth to maturity and the evaluation of the genetic and phenotypic
relationships between growth traits with lifetime lamb and wool production were studied using data
from 302 Rambouillet, 338 Targhee and 175 Columbia ewes born between I960 and 1976. The Brody
(1945) growth model was used for the derivation of growth parameters mature weight (A) and
maturing rate (k). The Fitzhugh and Taylor (1971) equation-free model was used for the estimation of
growth statistics; Absolute Growth Rate (AGR), Relative Growth Rate (RGR) and Absolute Maturing
rate (AMR) for five intervals from birth to 18 mo. Genetic and phenotypic parameters were estimated
by half-sib intraclass correlation using Harvey (1977) least-squares method for each breed and for the
pooled data. Targhees were superior to Rambouillet and Columbias for weight of lamb at weaning
(ATWW) and efficiency index (El). ATWW was the lifetime yearly average of kg of lamb weaned and
El was ATWW per unit of ewes mature weight (A). Columbias were superior for yearly average grease
wool produced (ATFP). Age at maturity was estimated on 39 mo, 38 mo and 41 mo for Rambouillets,
Targhees and Columbias, respectively. Columbias had the highest A and the smallest k, Targhees
matured the fastest. Ewes born twins had the highest El. From the pooled data, heritability estimates of
average total of lambs born (ATLB), average total of lambs weaned (ATLW), average total weight of
lambs weaned (ATWW), average total grease fleece produced (ATFP) and El were .43±.15, .33±..15,
.11±>15, 68±..16 and .15±,15, respectively. Genetic correlation, between ATWW and A, indicated that
ewes with high additive genetic potential for ATWW will have high genetic potential for larger A. The
genetic correlation between ATWW and k was zero. For the three Fitzhugh and Taylor (1971) growth
statistics, highest heritabilities were obtained for the weaning-12 mo interval: AGR2 (.80+.16), RGR2
(.76+.16) and AMR2 (.81+.16). RGR2 had the highest positive genetic correlations with ATWW
(.95+.81), El (.77+.S2) and ATFP (.39+.19) among growth statistics. Inclusion of AGR2 and RGR2 in
the construction of selection indexes would improve accuracy of selection for ATWW, ATFP and El.
However, the contributions of A and k were inferior to AGR2 and RGR2. The use of any of the growth
traits studied in selection indexes for improvement of ATFP and El simultaneously gave no advantage
in improving efficiency of selection. GENETIC ANALYSIS OF GROWTH AND ITS RELATIONSHIP WITH LIFETIME
PRODUCTION IN RAMBOUILLET, TARGHEE AND COLUMBIA EWES
UNDER RANGE CONDITIONS OF MONTANA
by
J u a n F ra n c is c o Chavez
A t h e s i s su b m itte d i n p a r t i a l f u l f i l l m e n t
o f th e re q u ire m e n ts f o r th e d e g re e
of
D octor o f P h ilo so p h y
in
B io lo g ic a l S c ie n c e s
MONTANA STATE UNIVERSITY
Bozeman, Montana
J u ly 1986
©1987
JUAN FRANCISCO CHAVEZ
All R ights R e serv e d
ii
APPROVAL
o f a t h e s i s su b m itte d by
Ju an F ra n c is c o Chavez
T his t h e s i s h a s been re a d by e a c h member o f th e t h e s i s com m ittee
and h a s been fo u n d to be s a t i s f a c t o r y re g a r d in g c o n te n t, E n g lish u sa g e ,
fo rm a t, c i t a t i o n s , b ib lio g r a p h ic s t y l e , and c o n s is te n c y , and i s re a d y
f o r su b m issio n t o th e C o lleg e o f G rad u ate S tu d ie s .
r?£C
-
Date
'
C o -C h airp erso n , G rad u ate Committee
«7.
)9 f O
•
C o -C h airp erso n , G rad u ate Committee
Approved f o r th e M ajor D epartm ent
/ffL
Date
Head, M ajor D epartm ent
Approved f o r th e C o lle g e o f G rad u ate S tu d ie s
J L ^ . ,
Date
^
/o,
G rad u ate Dean
ill
STATEMENT OF PERMISSION TO USE
In
p re s e n tin g
th is
th e s is
in
p a rtia l
fu lfillm e n t
of
th e
re q u ire m e n ts f o r a d o c to r a l d e g re e a t Montana S t a t e U n iv e r s ity , I a g re e
t h a t th e L ib ra r y s h a l l make i t
th e L ib r a r y .
th e
U .S .
b o rro w ers under r u l e s o f
I f u r t h e r a g re e t h a t copying o f t h i s t h e s i s i s a llo w a b le
o n ly f o r s c h o la r ly p u rp o s e s ,
in
a v a il a b le to
C o p y r ig h t Law.
c o n s i s t e n t w ith " f a i r u s e " a s p r e s c r ib e d
R e q u e s ts
fo r
e x te n s iv e
c o p y in g
or
re p ro d u c tio n o f t h i s t h e s i s sh o u ld be r e f e r r e d t o U n iv e r s ity M ic ro film s
I n t e r n a t i o n a l , 300 N orth Zeeb Road, Ann A rbor, M ichigan
48106, to whom
I have g ra n te d " th e e x c lu s iv e r i g h t t o re p ro d u c e and d i s t r i b u t e c o p ie s
o f th e d i s s e r t a t i o n i n and from m ic ro film and th e r i g h t to re p ro d u c e
and d i s t r i b u t e by a b s t r a c t i n any f o r m a t ."
To my w if e , my p a r e n ts ,
my c h ild r e n and my
c o u n try .
To M artha, my s i s t e r
V
VITA
Ju a n F ra n c is c o Chavez C o sslo was b o rn t o Mrs. M artha C o sslo and
Mr. G erardo Chavez i n Lima, P eru , on O cto b er 8 , 1948.
He a t t e n d e d C l a r e t i a n o H igh S c h o o l an d w as a d m itte d t o th e
F a c u lta d de Z o o te c n ia , U n iv e rs id a d N a cio n a l A g ra ria La M olina i n 1967.
He g r a d u a t e d a s a n I n g e n ie r o Z o o te c n is ta i n 1972, and a s M a g iste r
S c le n tia e i n Animal S c ie n c e i n 1979.
I n May o f 1975» he a c c e p t e d a p o s i t i o n w ith INIA ( I n s t i t u t e
N a c io n a l de I n v e s t i g a c i o n e s A g r a r i a s ) o f P e ru i n th e E s t a c i o n
E x p e rim en ta l A g ro p e cu a ria La M olina, in Lima.
F our y e a r s l a t e r , i n J u ly o f 1979, he became A s s is ta n t P r o f e s s o r
o f Animal B re e d in g a t U n iv e rs id a d N ao lo n al A g ra ria La M olina w here he
i s now A s s o c ia te P r o f e s s o r i n th e Animal H usbandry D epartm ent.
I n December, 1982, he was aw arded a f e llo w s h ip from S h a ll Ruminant
C o lla b o r a tiv e R esearch S u p p o rt Program , B re e d in g P r o je c t, to u n d e rta k e
g ra d u a te s t u d i e s a t Montana S t a t e U n iv e r s ity , Bozeman, M ontana, U .S.A.
He i s m a rrie d t o S usana Ram irez C a s t i l l o o f Chimb o te , P e ru .
They
have two c h ild r e n , M a r le lla and J u a n i t o .
Vi
ACKNOWLEDGEMENTS
I
e x p re s s my s in c e r e
th a n k s to
Dr.
D.G.
Cameron and D r.
R .L .
B la ck w e ll f o r t h e i r p a tie n c e , encouragem ent and g u id an ce th ro u g h o u t my
g ra d u a te program .
-
A p p re c ia tio n i s e x te n d e d t o th e members o f my g ra d u a te co m m ittee:
Dr. D.D. K re ss, Dr. R.E. Lund, Dr. H .J. Cameron and D r. E.R. V yse.
S p e c ia l
th a n k s
a re
e x p re s s e d t o
B u r r is and Dr. J.G . B e r a r d i n e l l I ,
D r.
P .J .
B u rfe n in g ,
from whom I re c e iv e d
a ll
Dr.
M.J .
k in d s o f
encouragem ent and f r i e n d s h i p .
I
am g r a t e f u l
to
my f r i e n d s
A n to n io Dona do,
R odolfo C a n te t,
D a r r e ll N evins, Matthew C ronin and S h e lly W a te rs, who made me f e e l a t
home.
I
a p p re c ia te
th e
d a ta
p ro c e s s in g
an d
c o m p u te r
p ro g ram m in g
a s s i s t a n c e p ro v id ed by K. H anford and h e r husband S. Kachman, and to
Joanne J e n s e n who a s s i s t e d i n e d i t i n g t h e f i n a l copy o f t h i s t h e s i s .
I a p p r e c ia te th e econom ical s u p p o rt p ro v id e d by th e Sm all R u n in an t
C o lla b o r a tiv e R esearch S u p p o rt Program , B re ed in g P r o j e c t , th ro u g h fu n d s
p ro v id e d by th e U n ite d S t a t e s Agency f o r I n t e r n a t i o n a l D evelopm ent.
I o f f e r my s in c e r e g r a t i t u d e t o a l l th e members o f Montana S t a t e
U n iv e r s ity who h e lp e d me.
The s u c c e s s f u l c o m p le tio n o f my G rad u ate
Program was due to th e w illin g n e s s o f th e s e p e o p le t o g iv e f r e e l y o f
t h e i r tim e .
V ii
TABLE OF CONTENTS
Page
LIST OF TABLES............................................................... ..................................................
ix
LIST OF FIGURES...............................................................................................................
x iv
ABSTRACT......................... ......................................................................................................
XV
IN TR O D U C TIO N ..................................................................................................................
I
REVIEW OF LITERATURE..........................................................................................................
3
G ro w th ..................................................................................................
Growth and lo n g e v i t y .................................... .....................................................
Growth and e f f i c i e n c y ........................................................................................
Models f o r grow th e v a lu a tio n ................................. ................................... ..
G e n e tic s o f grow th p a ra m e te rs and t h e i r r e l a t i o n s h i p t o
p ro d u c tiv e t r a i t s ......................
MATERIALS AND METHODS......................................................................................................
A nim als and e n v iro n m e n ta l c o n d itio n s ....................
D ata......................................................................................................................
S t a t i s t i c a l a n a ly s e s ................................................................
E s tim a tio n o f g e n e ti c and p h e n o ty p ic v a ria n c e and c o v a ria n c e
com ponents....................................................................
S e le c t io n in d e x e s ............................................
Summary o f th e s te p s fo llo w e d f o r h a n d lin g and a n a l y s i s o f
d a ta ..................................................................................................................
RESULTS AND DISCUSSION....................................................................................................
B reed and group d i f f e r e n c e s f o r p r o d u c tiv e t r a i t s .........................
L ife tim e p r o d u c tio n ................
F i r s t r e c o r d ......................
Second r e c o r d ...................................................................................
T h ird r e c o r d ..........................
F o u rth r e c o r d ................................................................................
Brody e q u a tio n p a ra m e te rs A1 B, and k and th e e f f i c i e n c y
in d e x ( E I ) .....................................................................................................
B ro d y 's grow th e q u a tio n s and age a t m a tu r ity . . . . . . .
C m p a riso n s o f AGR1 RGR and AMR f o r v a r y in g tim e i n t e r v a l s
betw een b i r t h and 18 m onths o f ag e................................................
AGR c o m p ariso n s....................................................................
RGR c o m p a riso n s...............................................................................
AMR c o m p a r is o n s ...........................................................................
E f f e c t o f type o f b i r t h and r e a r i n g on grow th and p ro d u c tiv e
perform ance o f e w e s . . . . . . ...........
6
7
11
19
26
26
27
29
33
33
35
35
35
36
39
41
41
44
46
46
46
49
49
53
V ili
TABLE OF CONTENTS (Continued)
Page
E s tim a te s o f h e r l L a b i l i t i e s , g e n e tic and p h e n o ty p ic
c o r r e l a t i o n s ...........................................................................................
G e n e tic p a ra m e te rs from th e p o o led d a ta
(R a m b o u ille ts , T arghees and Columb i a s ) ................
H e r i t a b i l l t y e s t i m a t e s ......... .....................................................
G e n e tic c o r r e l a t i o n s . . . .................
P h en o ty p ic c o r r e l a t i o n s ......... ....................................
S e le c t io n In d e x e s ............................................
C onceptual a n a ly s is o f r e s u l t s ...................................................................
68
73
SUMMARY AND CONCLUSIONS..............................................................................................
76
LITERATURE C IT E D ..........................................................................................................
80
APPENDIX,
85
55
57
57
64
67
ix
LIST OP TABLES
Page
I
R ep ro d u ctiv e in d e x (A fte r Holmes,
1 9 7 7 ).........
12
2
H e r i t a b i l i t y e s tim a te s f o r grow th
c u rv e p a ra m e te rs ..........
20
3
H e r i t a b i l i t y e s tim a te s f o r grow th
s t a t i s t i c s ........................
21
4
G e n e tic c o r r e l a t i o n s betw een grow th p a ra m e te rs and
p ro d u c tiv e t r a i t s ...................................... .......................................
22
G e n etic c o r r e l a t i o n s betw een grow th s t a t i s t i c s and
p ro d u c tiv e t r a i t s ...................................................................................
23
D i s t r i b u t i o n o f th e ewes by b re e d , group o f p ro d u c tio n
and y e a r o f b i r t h . . . . . ......... .........................................................
27
7
EWes1 p r o d u c tiv e and grow th t r a i t s a n a ly z e d i n th e stu d y
30
8
D i s t r i b u t i o n o f s i r e s and o f f s p r i n g p e r b reed and y e a r . .
32
9
Mean s q u a re s and l e a s t s q u a re s means f o r th e l i f e t i m e
a v e ra g e s o f p ro d u c tiv e t r a i t s o f ewes (H arvey l e a s t
s q u a re s a n a ly s e s , model A ).................................................................
37
Mean s q u a re s and l e a s t s q u a re s means f o r th e c u m u la tiv e
av erag e o f p ro d u c tiv e t r a i t s o f ewes th ro u g h t h e i r f i r s t
re c o rd (Harvey l e a s t s q u a re s a n a ly s e s , model A ) . . . . .........
38
Mean s q u a re s and l e a s t s q u a re s means f o r th e c u m u la tiv e
av erag e o f p ro d u c tiv e t r a i t s o f ewes th ro u g h t h e i r
second r e c o r d (H arvey l e a s t s q u a re s a n a ly s e s , model A ) ..
40
Mean s q u a re s and l e a s t s q u a re s means f o r th e c u m u la tiv e
av erag e o f p ro d u c tiv e t r a i t s o f ewes th ro u g h t h e i r t h i r d
re c o r d (Harvey l e a s t s q u a re s a n a ly s e s , model A )..............
42
Mean s q u a re s and l e a s t s q u a re s means f o r th e c u m u la tiv e
a v erag e o f p ro d u c tiv e t r a i t s o f ewes th ro u g h t h e i r
f o u r th re c o rd (H arvey l e a s t s q u a re s a n a ly s e s , model A ) ..
43
Mean s q u a re s and l e a s t s q u a re s means f o r B rody’ s grow th
curve p a ra m e te rs (A, B, k ) and th e ewe e f f ic ie n c y in d e x
(H arvey l e a s t s q u a re s a n a ly s e s , model A ) . . . . i .....................
45
5
6
10
11
12
13
14
X
LIST OF TABLES (Continued)
Table
15
16
17
18
Page
Mean s q u a re s and l e a s t s q u a re s means f o r a b s o lu te grow th
r a t e (AGB) from b irth -w e a n in g (AGBI)f w eaning-12 m onths
(AGB2), 12-18 m onths (AGBS)f b ir t h - 1 2 months (AGB4) and
B irth - 1 8 m onths (AGB5) (H arvey l e a s t s q u a re s a n a ly s e s ,
model A )...................................... ..................................................................
48
Mean s q u a re s and l e a s t sq u a re s means f o r , r e l a t i v e grow th
r a t e (BGB) from b irth -w e a n in g (BGBI)f w eaning-12 m onths
(BGB2), 12-18 m onths (BGB3), b ir t h - 1 2 months (BGB4) and
b ir th - 1 8 months (HGB5) (Harvey l e a s t s q u a re s a n a ly s e s ,
model A)........................................................................ ................... ..
50
Mean s q u a re s and l e a s t s q u a re s means f o r a b s o lu te
m a tu rin g r a t e (AMB) from b irth ^ w e a n in g (AMBI)f w ean in g 12 months (AMB2), 12-18 months (AMB3), b ir t h - 1 2 months »
(AMB4) and b ir th - 1 8 m onths (AMB5) (H arvey l e a s t s q u a re s
a n a ly s e s model A )........................................
51
L e a st s q u a re s means f o r b i r t h - r e a r i n g e f f e c t s on a l l th e
s tu d ie d l i f e t i m e t r a i t s o f th e ewes u s in g th e b re e d s
p o o le d in f o r m a tio n (H arvey l e a s t s q u a re s a n a ly s e s , model
C ).......................................................................................................................
54
19
E s tim a te s o f h e r ! L a b i l i t i e s , g e n e tic and p h e n o ty p ic
c o r r e l a t i o n s f o r l i f e t i m e p ro d u c tio n and grow th t r a i t s
(p o o le d d a ta from B a m b o u ille tf T arghee and Colum bia
e w e s)................ ...... .......................... ............ ........................................ . . . 5 8 - 6 1
20
D eriv ed s e l e c t i o n in d e x e s f o r th e a n a l y s i s o f th e e f f e c t
o f A, k, AGB2 and BGB2 on th e im provem ent o f a v erag e
t o t a l o f w e ig h t o f lam bs weaned (ATWV) and th e
.
e f f ic ie n c y in d e x (E I) by s e l e c t i o n ....................................................
70
D eriv ed s e l e c t i o n in d e x e s f o r th e a n a l y s i s o f th e e f f e c t
o f Af k, AGB2 and BGB2 on th e sim u lta n e o u s im provem ent
o f average t o t a l o f w e ig h t o f lam bs weaned (ATWW)f
e f f ic ie n c y In d ex (E I) and t o t a l g re a s e f le e c e produced
(ATFP)................................................... .........................................................
72
21
xi
LIST OF TABLES (Continued)
Page
23
24
25
26
27
28
29
A verages, v a r i a t i o n and ra n g e s o f l i f e t i m e a v erag e o f
t o t a l o f lam bs born (ATLB) f o r ewe g ro u p s w ith in
b re e d s ...........................................................
85
A verages, v a r i a t i o n and ra n g e s o f l i f e t i m e a v erag e o f
th e t o t a l number o f lam bs weaned (ATLW) f o r ewe g ro u p s
w ith in b re e d s ...........................................................................
85
A verages, v a r i a t i o n and ra n g e s o f l i f e t i m e a v erag e o f
th e t o t a l o f k ilo g ra m s o f lam bs weaned (ATWW) f o r ewe o f
g ro u p s w ith in b re e d s ..................................
86
A verages, v a r i a t i o n and ra n g e s o f l i f e t i m e a v erag e o f
k ilo g ra m s o f fle e o e produced (ATFP) f o r ewe g ro u p s
w ith in b re e d s ...................................... ................... ...................................
86
A verages, v a r i a t i o n and ra n g e s o f m atu re s i z e (A) f o r
ewe g ro u p s w ith in b r e e d s ......................................................................
87
A verages, v a r i a t i o n and ra n g e s o f B rody’ s c o n s ta n t o f
i n t e g r a t i o n (B) f o r ewe g ro u p s w ith in b r e e d s .........................
I
'
A verages, v a r i a t i o n and ra n g e s o f m a tu rin g r a t e (k ) f o r
ewe g ro u p s w ith in b r e e d s .....................................................................
88
A verages, v a r i a t i o n and ra n g e s o f e f f i c i e n c y in d e x (E I)
f o r ewe g roups w ith in b r e e d s ......................... ................... ...............
88
30
A verages, v a r i a t i o n and ra n g e s o f a b s o lu te grow th r a t e
i n th e i n t e r v a l b irth -w e a n in g (AGBI) f o r ewe g ro u p s
w ith in b re e d s ..................................................................................... ....
31
A verages, v a r i a t i o n and ra n g e s o f a b s o lu te grow th r a t e
i n th e i n t e r v a l w eanlng-12 m onths (AGR2) f o r ewe g ro u p s
w ith in b r e e d s . . . . . ........................... ............................ ................... ..
32
A verages, v a r i a t i o n and ra n g e s o f a b s o lu te grow th r a t e
i n th e i n t e r v a l 12-18 m onths ( AGR3) f o r ewe g ro u p s
w ith in b re e d s ..............................................................................
33
A verages, v a r i a t i o n and ra n g e s o f a b s o lu te grow th r a t e
i n th e i n t e r v a l b ir t h - 1 2 m onths (AGR4) f o r ewe g ro u p s
w ith in b re e d s ........................................................ ...............
87
X ii
LIST OF TABLES (Continued)
Table
34
35
36
37
38
39
40
41
42
43
44 -
45
Page
A verages, v a r i a t i o n and ra n g e s o f a b s o lu te grow th r a t e
i n th e i n t e r v a l b ir th - 1 8 months (AGR5) f o r ewe g ro u p s
w ith in b r e e d s .............................. ...........................................................
91
A verages, v a r i a t i o n and ra n g e s o f r e l a t i v e grow th r a t e
i n th e i n t e r v a l b irth -w e a n in g (RGRI) f o r ewe g ro u p s
w ith in b re e d s ........................................................................................... ..
91
A verages, v a r i a t i o n and ra n g e s o f r e l a t i v e grow th r a t e
i n th e i n t e r v a l w eaning-12 m onths (RGR2) f o r ewe g ro u p s
w ith in b re e d s .................. ...........................................................................
Q2
A verages, v a r i a t i o n and ra n g e s o f r e l a t i v e grow th r a t e
i n th e i n t e r v a l 12-18 m onths (RGR3) f o r ewe g ro u p s
w ith in b re e d s ..................................................................... ..
02
A verages, v a r i a t i o n and ra n g e s o f r e l a t i v e grow th r a t e
i n th e i n t e r v a l b ir t h - 1 2 m onths (RGR4) f o r ewe g ro u p s
w ith in b re e d s .................. .........................................................t . T. . TTt
Ql
73
A verages, v a r i a t i o n and ra n g e s o f r e l a t i v e grow th r a t e
i n th e i n t e r v a l b i r t h —18 m onths ( RGR5) f o r ewe g ro u p s
w ith in b re e d s ............................................................................................ ..
Ql
7J
A verages, v a r i a t i o n and ra n g e s o f a b s o lu te m a tu rin g r a t e
i n th e i n t e r v a l b irth ^ w e a n in g (AMRI) f o r ewe g ro u p s
w ith in b r e e d s . . . . . . . . . . ................ ..................................................
Oil
A verages, v a r i a t i o n and ra n g e s o f a b s o lu te m a tu rin g r a t e
i n th e i n t e r v a l w eaning-12 m onths (AMR2) f o r ewe g ro u p s
w ith in b re e d s .....................................................................
Qll
A verages, v a r i a t i o n and ra n g e s o f a b s o lu te m a tu rin g r a t e
i n th e i n t e r v a l 12—18 months (AMR3) f o r ewe g ro u p s
w ith in b r e e d s . . . . . ......................... ................................... ..
QS
A verages, v a r i a t i o n and ra n g e s o f a b s o lu te m a tu rin g r a t e
i n th e i n t e r v a l b i r t h —12 m onths (AMR4) f o r ewe g ro u p s
w ith in b re e d s .............................................................
QS
A verages, v a r i a t i o n and ra n g e s o f a b s o lu te m a tu rin g r a t e
i n th e i n t e r v a l b ir t h - 1 8 months (AMR5) f o r ewe g ro u p s
w ith in b r e e d s .................................................... ...........
Q
6
7V
P r e d ic te d body w e ig h ts and p e r c e n t o f m atu re w e ig h t (A)
f o r each b re e d and ewe group by B ro d y 's grow th e q u a tio n
d e riv e d i n t h i s s tu d y i n s i x m onths i n t e r v a l s . . . ............
97
X iii
LIST OF TABLES (Continued)
Table
46
Page
L e a st s q u a re s means o f b i r t h - r e a r i n g e f f e c t s f o r each
b reed (H arvey a n a l y s i s , m odel B )....................................................
g8
E s tim a te s o f h e r i t a b i l i t i e s , g e n e tic and p h e n o ty p ic
c o r r e l a t i o n s f o r l i f e t i m e p ro d u c tio n and grow th t r a i t s
o f R am b o u illet ew es................................................................................99-102
E s tim a te s o f h e r i t a b i l i t i e s , g e n e tic and p h e n o ty p ic
c o r r e l a t i o n s f o r l i f e t i m e p ro d u c tio n and grow th t r a i t s
o f T arghee e w e s . . . .................................................................................
E s tim a te s o f h e r i t a b i l i t i e s , g e n e tic and p h e n o ty p ic
C?r ^ ef a ^ 0nS ^ o r l i f e t i m e p ro d u c tio n and grow th t r a i t s
o f O olunbla ewes.....................................................
107_110
I
Among s i r e s com ponents o f v a r ia n c e and c o v a ria n c e f o r
pIh0 dUc t ^ f n and Sr0w th t r a i t s ( P°o le d d a ta from
R a m b o u ille t, Targbee and Colum bia e w e s ) .............................. 111-113
W ith in s i r e s com ponents o f v a ria n c e and c o v a ria n c e f o r
x iv
LIST OF FIGURES
F ig u re
1
2
3
Page
A r e p r e s e n t a t i v e o f B ro d y 's e q u a tio n (and m o d if ic a tio n )
f i t t e d t o a n Angus h e ig h t-a g e cu rv e ( A f te r N e lse n e t
a l . , 1982) .....................................................................................................
15
Flow c h a r t i n d i c a t i n g th e s te p s fo llo w e d f o r h a n d lin g o f
d a ta and s t a t i s t i c a l a n a ly s e s .............. ..............................................
34
Growth c u rv e s and e q u a tio n s f o r R a m b o u ille ts (R );
T arghees (T) and Colum bias (C )f d e riv e d from th e L e a s t
s q u a re s means from t a b l e 1 4 . . . .......................
47
---- —
MtBC •
db
10/01/86
10/01/86
—— /— / —
I (WoOLN)4 0 5 1423
IC h o v e z , Juon Froncisco.
IGenetic analysis of growth and its relationship with
lifetime production in Rombouillety Torghee and
Columbia ewes under range conditions in Montana / by
Juon Francisco Chavez.
Io
IBozeman, M T ,, 1986»
Io
Ixv, 116 leaves
IThesis (PhD*)— Montana State U ♦, Biological Sciences
---- David Cameron. ' .~
'951
LON
MEP.,
T I L .0
IMP.
COL
Ia
Io
Io
XV
ABSTRACT
Growth c h a r a c t e r i s t i c s o f sh eep from b i r t h t o m a tu r ity and th e
e v a lu a tio n o f th e g e n e ti c and p h e n o ty p ic r e l a t i o n s h i p s betw een grow th
t r a i t s w ith l i f e t i m e I a s b and wool p ro d u c tio n w ere s tu d ie d u s in g d a ta
from 302 R a m b o u ille tl 338 Targhee and 175 Columbia ewes b o rn betw een
I960 and 1976.
The B rody C1 9 4 5 ) g ro w th m o d el w as u s e d f o r th e
d e r i v a t i o n o f grow th p a ra m e te rs m a tu re w e ig h t (A) and m a tu rin g r a t e
(k ).
The F itzh u g h and T a y lo r (197 1 ) e q u a ti o n - f r e e model was u sed f o r
t h e e s t i m a t i o n o f g ro w th s t a t i s t i c s ; A b so lu te Growth R ate (AGR),
R e la tiv e Growth R ate (RGR) and A b so lu te M atu rin g r a t e (AMR) f o r f i v e
i n t e r v a l s from b i r t h to 18 mo. G e n e tic and p h e n o ty p ic p a ra m e te rs w ere
e s tim a te d by h a l f - s i b i n t r a c l a s s c o r r e l a t i o n u s in g H arvey (1977) l e a s t s q u a re s m ethod f o r each b re e d and f o r th e p o o led d a ta .
T arg b ees w ere
s u p e r io r t o R a m b o u ille t and C o lim b las f o r w e ig h t o f lam b a t w eaning
(ATWW) and e f f i c i e n c y in d e x ( E l ) . ATWW was th e l i f e t i m e y e a r ly av erag e
o f kg o f Ia n b weaned and E I was ATWW p e r u n i t o f ewes m a tu re w e ig h t
(A ).
C olum bias w ere s u p e r io r f o r y e a r ly av erag e g re a s e wool produced
(ATFP).
Age a t m a tu r ity was e s tim a te d on 39 mo, 38 mo and 41 mo f o r
R a m b o u ille ts, T argbees and C olim b i a s , r e s p e c t i v e l y .
C olum bias h ad th e
h ig h e s t A and th e s m a lle s t k . T a rg h ees m atu red th e f a s t e s t . Ewes born
tw in s had th e h ig h e s t E l. From th e p o o led d a ta , h e r i t a b i l i t y e s tim a te s
o f av erag e t o t a l o f lcm bs born (ATLB), av erag e t o t a l o f lam bs weaned
(ATLW), a v erag e t o t a l w e ig h t o f la n b s weaned (ATWW), a v erag e t o t a l
g re a s e f l e e c e produced (ATFP) and E I w ere .4 3 ± .1 5 , .3 3 ± ,1 5 , .1 1 ± ,1 5 ,
68±..16 and .1 5 ± ,1 5 , r e s p e c t i v e l y .
G e n e tic c o r r e l a t i o n , betw een ATWW
and A, in d ic a te d t h a t ewes w ith h ig h a d d itiv e g e n e ti c p o t e n t i a l f o r
ATWW w i l l have h ig h g e n e ti c p o t e n t i a l f o r l a r g e r A.
The g e n e t i c
c o r r e l a t i o n betw een ATWW and k was z e r o .
For th e t h r e e F itz h u g h and
T a y lo r (1971) grow th s t a t i s t i c s , h i g h e s t h e r l t a b i l l t i e s w ere o b ta in e d
f o r th e w eaning-12 mo i n t e r v a l :
AGR2 ( ,8 0 + .1 6 ) , RGR2 ( . 76+ .1 6 ) and
AMR2 ( .8 1 + .1 6 ) .
KJR2 h ad th e h ig h e s t p o s i t i v e g e n e ti c c o r r e l a t i o n s
w i t h ATWW ( . 9 5 + . 8 1 ) , EI (.7 7 + * 5 2 ) and ATFP (.3 9 + * 1 9 ) among grow th
s ta tis tic s .
I n c l u s i o n o f AGR2 a n d RGR2 i n th e c o n s t r u c t i o n o f
s e l e c t i o n in d e x e s w ould im prove a c c u ra c y o f s e l e c t i o n f o r ATWW, ATFP
and E l.
However, th e c o n tr ib u tio n s o f A and k w ere i n f e r i o r t o AGR2
and RGR2.
The use o f any o f th e grow th t r a i t s s tu d ie d i n s e l e c t i o n
in d e x e s f o r im provem ent o f ATFP and E I s im u lta n e o u s ly gave no a d v an ta g e
i n im p ro v in g e f f i c i e n c y o f s e l e c t i o n .
I
INTRODUCTION
T he
la rg e
n u m b e rs
of
sheep
b re e d s
and
th e
in d iv id u a l
c h a ra c te ris tic s
o f th e w ide ra n g e o f e n v iro n m en ts t o w hich th e y
a d a p te d
a f u n d a m e n ta l
p la y s
ro le
in
th e ir
p o t e n t i a l r e l a t i v e t o o th e r d o m e stic s p e c ie s .
u n iv e rs a l
a re
p ro d u c tiv e
T h e ir a b i l i t y to s u b s i s t
and produce under d i f f e r e n t e co sy stem s a t t e s t s t o t h e i r b ro ad a d a p tiv e
c a p a b ilitie s .
in d ic a to r
R ep ro d u ctiv e perform ance o r f i t n e s s h a s been u sed a s a n
o f how w e l l
a
p o p u la tio n
e n v iro n m e n t. F it n e s s h a s a l s o
by g e n e tic
e ffe c ts
o th e r
is
a d a p te d
to
a
p a rtic u la r
been shown t o be in f lu e n c e d im p o r ta n tly
th a n a d d i t i v e ,
th e
la tte r
b e in g
th e
raw
m a te r ia l by which s e l e c t i o n a c c o m p lish e s th e g o a ls o f im provem ent.
Some advance h a s
been a c h ie v e d i n
th e
g e n e tic
im p ro v e m e n t
of
s t r a i n s o r b re e d s o f sheep f o r r e p r o d u c tiv e perfo rm an ce and a m ajor
gene f o r m u ltip le b i r t h s h a s been d is c o v e re d i n th e A u s t r a lia n s t r a i n
of
M erinos c a l l e d B o o ro o la .
Y et,
i n many o f
th e w e ll
e sta b lis h e d
b re e d s o f sh eep r e p r o d u c tiv e p erfo rm an ce i s th e m ain l i m i t i n g f a c t o r o f
p ro d u c tiv ity .
I t h a s been o b se rv e d t h a t r e p r o d u c tiv e p e rfo rm an c e, e v a lu a te d i n
many d i f f e r e n t w ays, does n o t re sp o n d r e a d i l y t o a r t i f i c i a l s e l e c t i o n .
R easons f o r l i m i t e d re s p o n se t o a r t i f i c i a l s e l e c t i o n may be due t o o u r
la c k
o f knowledge a b o u t
th e many p o s s ib le
b io lo g ic a l v a r i a b l e s
and
t h e i r i n t e r a c t i o n s in v o lv e d i n th e e x p re s s io n s o f re p ro d u c tio n and ( o r )
t h a t we have n o t y e t d is c o v e re d th e m ost p r a c t i c a l way f o r e v a lu a tio n
and a p p l i c a t i o n o f In fo rm a tio n i n s e l e c t i o n .
2
Growth h a s a ls o
been a s s o c ia te d w ith f i t n e s s and i t
is
p o s s ib le
t h a t a g e n e tic r e l a t i o n s h i p e x i s t s betw een th e s e two b a s ic t r a i t s .
th a t i s
tru e ,
re p ro d u c tiv e
th e use o f grow th a s a n i n d i c a t o r
p erfo rm an ce would be im p o r ta n t i n
If
of fu tu re lif e tim e
d e v e lo p m e n t
of
b re e d in g p la n s t o im prove r e p r o d u c tiv e perfo rm an ce and e f f i c i e n c y
of
p ro d u c tio n i n sh e e p .
re la tio n s h ip s
c o n s id e r a tio n
to
A d d itio n a l in fo r m a tio n i s
b e tw e e n
g ro w th
th e f a c t
th a t
an d
th e
needed r e g a r d in g
life tim e
p ro d u c tio n ,
grow th and l i f e t i m e
th e
g iv in g
p r o d u c tio n ,
a re
c u m u la tiv e p ro c e s s e s i n developm ent.
The purpose o f th e p r e s e n t s tu d y was t o e s tim a te th e g e n e tic and
p h e n o ty p ic r e l a t i o n s h i p s
among m e asu re s o f
e a rly
g r o w th ,
ra te s
of
m a tu r ity , m ature s i z e , r e p ro d u c tio n and p ro d u c tio n o f ran g e sh ee p , and
to id e n tify
t r a i t s t h a t can be m easured e a r l y i n l i f e
t h a t w i l l have
v a lu e f o r p r e d i c t i n g g e n e tic m e r it o f l i f e t i m e p ro d u c tio n .
3
REVIEW OF LITERATURE
Growth
G row th h a s alw ays been lin k e d t o
p a rtic u la r,
g a in s ,
body
when developm ent i s
t h a t developm ent i s
g r o w th .
a s s im ila to ry
(n u trie n t)
d e fin e d
u n s ta b le
B ro d y
w hich u n d e rg o e s
g ro w th
as
th e
e q u ilib riu m
s u rro u n d in g i t .
e v a lu a te d i n
c o n s id e re d t o
(1 9 4 5 )
s y n th e s is o f
p r o g r e s s iv e d ev elo p m en t.
d e fin e d
g r o w th
at
d is s im ila tio n " .
b e tw e e n
in
th e
of
body w e i g h t
be a consequence o f g e n e r a l
one s u b sta n c e
in c re a s e
te rm s
In
th e
as
"C o n stru c tiv e
th e
expense
E a rlie r
number o f
o rg a n is m
and
of
B ro d y
c e lls
th e
or
a n o th e r
(1 9 2 8 )
due t o
an
e n v ir o n m e n t
The approach to a s t a b l e e q u ilib riu m i s c a l l e d grow th.
Bonner (1 9 5 8 ), in d i c a t e d t h a t a c c u m u la tio n o f p ro to p la sm by grow th
i s a consequence o f d evelopm ent.
He w ro te ,
"The t e l e o l o g i c a l p u rp o se o f developm ent i s t o c r e a t e a n
i n d i v i d u a l t h a t i s w e l l a d a p te d a n d s u c c e s s f u l i n i t s
e n v iro n m e n t". "A nother im p lic a tio n i n t h a t th e r e h a s been an
in c r e a s e i n d i v i s i o n o f la b o u r , f o r c l e a r l y t h i s i s v i t a l f o r
a d a p tiv e s u c c e s s .
I n te r m s o f d e v e lo p m e n t t h i s m ean s
d iffe re n tia tio n .
In c re a s e i n s i z e and d i v i s i o n o f la b o u r a r e
th e two prim ary g o a ls o f d e v e lo p m e n t.." .
" In l a r g e m easure
th e s u c c e s s o f o rg an ism s i n n a tu r e depends upon t h e i r b e in g
w e ll k n i t and c lo s e ly c o - o r d in a te d .
I t i s n o t enough t o have
a la r g e organism w hich p a r c e ls o u t i t s a c t i v i t i e s ; i t m ust be
a d i s c r e t e , sm oothly f u n c tio n in g u n i t , p h y s io lo g ic a ll y w e ll
b a la n c e d w ith in , a s w e ll a s a d ju s t e d t o i t s e n v iro n m e n t.
It
m ust have sy stem s o f p h y s io lo g ic a l r e g u l a t i o n , c o r r e l a t i o n and
c o n tr o l, so t h a t i t i s s t a b l e even i n a d v e r s i t y . T h is k in d o f
s t a b i l i t y by c o - o r d in a tio n p la y s a n im p o r ta n t r o l e i n th e
s u r v iv a l o f th e f i t t e s t and m ust be c o n s ta n tly im proved by
s e le c tio n .«
From th e p re v io u s p a ra g ra p h i t m ig h t be i n f e r r e d t h a t grow th i s a
r e a c t i o n o f th e an im al t o
id e a l,
its
a b a la n c e d s u c c e s s f u l
e n v iro n m e n t.
s iz e fo r
Im p lic itly
th e r e
a s p e c i f i c e n v iro n m e n t.
is
an
That
4
i d e a l could be c a l l e d
p r o p o r tio n a l ity
th e f i t t e s t
th e optimum
Thus, th e tr e n d s o f d i f f e r e n t i a t i o n co u ld be a n a ly z e d
i n te rm s o f grow th p a t t e r n s .
b a la n c e
c o rre s p o n d in g t o
and d i v i s i o n o f la b o r among a l l th e c o n s t i t u e n t p a r t s
i n th e organism .
would be th e
s iz e
An im p o rta n t v a r i a b l e w ith in th e system
tim e r e q u ir e d f o r an in d iv id u a l t o re a c h
its e lf,
which w ould be r e l a t e d
to
fitn e s s
th e
an d ,
p o in t
of
im p lic itly ,
s tr o n g ly in flu e n c e d by n a tu r a l s e l e c t i o n .
I n sh ee p , B u t t e r f i e l d e t a l . (1 9 8 3 a ,b) d e m o n stra te d t h a t betw een a
la rg e
and sm all
s tra in
o f M erinos ,
p ro p o r tio n s o f m u scle, bone, f a t ,
o rg a n s r e l a t i v e
t o w e ig h t.
t i s s u e s and o rg a n s
th e re
e x is ts
a w id e
ra n g e
of
a lim e n ta ry t r a c t and o th e r i n t e r n a l
However, when th e
p r o p o r tio n s o f
th e s e
w ere compared i n r e l a t i o n t o m a tu r ity , a r e d u c tio n
o f th e s e d if f e r e n c e s was o b s e rv e d .
M cC lelland e t a l . (1 9 7 6 ) re a c h e d a
s i m i l a r c o n c lu s io n i n a s tu d y on th e d i f f e r e n c e s i n body c o m p o sitio n o f
fo u r b re e d s o f sheep a t m a tu r ity .
These s tu d ie s s u g g e s t t h a t w e ig h t a t
a s p e c i f i c age i s o n ly an i n d i c a t i v e p o in t w ith in a w hole b io lo g ic a l
p ro c e s s
c a lle d
g row th.
That p ro c e s s in v o lv e s a n im p o rta n t group o f
v a r i a b l e s , and i s m ean in g fu l i n c o n s id e rin g c o m p a ra tiv e re s p o n s e s a b o u t
anim al e f f i c i e n c y .
The p a t t e r n s o f grow th a r e s i m i l a r f o r each o rg a n , how ever, th e y
d i f f e r i n e a r l i n e s s o f m a tu r ity
(co m p lete d e v elo p m en t).
in
n o te d
body
w e ig h t w ith
s ig m o id a l
c u rv e
tim e ,
w ith
" s e lf-a c c e le ra tin g "
tw o
p h ase,
as
w e ll
b e fo re
fo llo w in g " s e l f - i n h i b i t i n g " p h a se .
by Brody (1 9 2 7 a ),
d iffe re n tia te d
an
The in c r e a s e
in fle c tio n
The shape o f th e
p ro d u c e s a
s ta g e s :
p o in t,
th e
an d t h e
grow th c u rv e f o r
5
each c o n stitu e n t body p a rt w ill tend to have a sim ila r p a tte rn to th a t
of the whole body (Hammond, 1932).
The
s e lf-a c c e le ra tin g
p h ase,
s ta rtin g
at
c o n c e p tio n ,
shows a
broken p a t t e r n o f a t l e a s t th r e e c y c le s c a l l e d i n f a n t i l e , J u v e n ile and
a d o le s c e n t
(R o b e rts o n ,
1908;
c ite d
o c c u rs t h a t o th e r s have c a ll e d
le a s t
fo u r
(1 9 2 8 ).
of
th e s e
by Brody,
1 9 2 8 ).
A phenomenon
"m etam orphosis" (D a v e n p o rt, 1926).
c y c le s w ere
c o n s id e re d t o
These could be r e l a t e d t o
ch an g es i n
be p r e s e n t
p ro p o r tio n
At
by Brody
due t o
th e
d i f f e r e n t s i z e s o f th e com ponents when d i f f e r e n t i a t e d i n th e embryo, a s
w e ll a s
th e ir
d iffe re n t
a s y m p to tic
w e ig h ts
an d
g ro w th
c u rv e s
at
d i f f e r e n t p h a se s , a s p o in te d o u t by Fow ler (1 9 8 0 ).
These
c y c le s a r e
i n t e r n a l o rg a n s .
e v id e n tly
p ro d u c ts
of
th e
developm ent
of
th e
They p ro c e ed to w ard t h e i r m atu re w e ig h ts e a r l i e r th a n
does th e body a s a whole ( B u t t e r f i e l d e t a l ,
developm ent a r e a ls o in c lu d e d .
1983b).
M uscle and bone
The amount o f f a t , th e t i s s u e o f l a t e s t
developm ent ( B u t t e r f i e l d e t a l . , 1 9 8 3 a), i s a n i n d i c a t i o n o f m a tu r ity .
T a y lo r
( 1 9 8 2 ),
s u g g e s ts
th a t
m a tu r e w e ig h t can be d e fin e d a s an
e q u ilib riu m w eig h t c o n ta in in g 15% o f chem ical f a t .
T h u s,
g ro w th
is
a
g ra d u a l
p ro c e ss
of
re a c h in g
e q u ilib riu m o f a n organism t o a s p e c i f i c e n v iro n m e n t.
th a t
th e
fa s te r
g ro w in g a n im a ls w hich re a c h m a tu r ity
p ro b a b ly be th e f i t t e s t .
a
I t is
b a la n c e d
p o s s ib le
e a r lie r w ill
I t would be im p o rta n t t o know, a s r e g a r d s th e
m atu re anim al p erfo rm an ce, w hether th e r e i s any s e l e c t i v e ad v an tag e f o r
th e f a s t m a tu rin g a n im a ls compared t o
th e
slow m a tu rin g o n e s .
TWo
a p p ro a c h e s can be ta k e n t o lo o k f o r a p o s s ib le an sw er; a n a n a l y s i s o f
6
lo n g e v ity
an d
an
a n a ly s is
of
p ro d u c tiv e
p e rfo rm a n c e ,
in c lu d in g
e s p e c ia ll y r e p r o d u c tiv e p e rfo rm an c e. .
Brody (1928) s u g g e s te d th e a p p l i c a t i o n o f th e e q u a tio n Wfc * Aek t
f o r r e p r e s e n tin g any o f th e segm ents ( c y c le s ) o f th e s e l f - a c c e l e r a t i n g
phase o f grow th.
(t);
(Wfc) , r e p r e s e n ts th e w e ig h t o f a g iv e n an im al a t age
(A) c o rre sp o n d s t o
th e a d u l t w e ig h t (m a tu re w e ig h t) ;
(e ) i s
th e
b ase o f th e n a tu r a l lo g a r ith m ; and (k ) i s th e r e l a t i v e r a t e o f grow th
w ith r e s p e c t t o m atu re w eig h t (A ).
Growth and lo n g e v i t y
A fte r th e o n s e t o f p u b e rty , th e r a t e o f grow th d e c r e a s e s g ra d u a lly
u n t i l th e maximum s i z e i n a l l th e o rg a n s i s re a c h e d and th e m a tu r ity o f
th e in d iv id u a l i s a t t a i n e d .
Along w ith d ev elo p m en tal gro w th , a g ra d u a l
change i n th e c a p a b i l i t y o f th e organism t o overcome d is e a s e ( v i t a l i t y )
o ccu rs.
The maximum v i t a l i t y
S e n escen ce,
th e o re tic a lly ,
is
s ta rts
re a c h e d a t
at
p u b e rty ,
p u b e rty
when
(B rody,
sexual
1 9 2 8 ).
a c tiv ity
b e g in s.
A ccording t o
Comfort (1 9 6 1 ), a g in g o r s e n e sc e n c e means a d e c lin e
i n v ig o r o r v i t a l i t y w ith th e p a s s in g o f tim e , and i s c h a r a c te r iz e d by
th e In c r e a s e d p r o b a b i l i t y o f d y in g .
He f u r t h e r s t a t e s t h a t lo n g e v ity
o r th e lo n g d u r a tio n o f l i f e h as been fo u n d to be c o r r e l a t e d w ith s i z e
in
d iffe re n t
e x is ts
a n im a l
b e tw e e n
l o n g e v i t y 's
s p e c ie s ,
lo n g e v ity
c lo s e s t
s in g le
an d
but a
net
c lo s e r r e la tio n s h ip
re p ro d u c tiv e
c o rre la te
seem s
to
ra te .
be
th e
a p p a r e n tly
H o w ev er,
" in d e x o f
c e p h a liz a tio n " w hich i s m easured by th e e x c e s s o f b r a in s iz e o v er t h a t
e x p e c te d
from
(C om fort, 1961).
th e
g en eral
p a t t e r n o f mammalian o r g a n ic p r o p o r tio n s
7
N orm ally grow th and se n e sc e n c e a r e e x p re s s e d a s a consequence o f
tim e , b u t f o r grow th and s e n e sc e n c e , tim e i s somewhat r e l a t i v e and can
be c o n sid e re d a s a c o n ju g a te e f f e c t o f a l l th e f a c t o r s in v o lv e d i n a
s p e c i f i c e n v iro n m en t w here th e in d iv id u a l grows and re p ro d u c e s .
m ore
im p o rta n t
n u tritio n
e n v ir o n m e n ta l
fa c to rs
and i o n i z i n g r a d i a t i o n
appear
(C o m fo r t,
to
be
1 9 6 1 ).
The
te m p e ra tu re ,
The e f f e c t
of
n u t r i t i o n on grow th and se n e sc e n c e h a s been d e m o n stra te d i n r a t s (McCay
e t a l . , 1935; B erg and Sim s, 1961).
fla ts have a much s h o r te r l i f e span u n d er optimum fe e d in g f o r r a p id
grow th th a n r a t s
under a system
o f f e e d in g w hich
a v o id s any v ita m in d e f ic ie n c y .
ju v e n ile ,
and d is e a s e
a p p e a rs
n u tritio n a l
th a t
ra p id
le v e l
grow th r a t e .
bu t
fre e
th a n r a t s
h a v in g
a
When th e d i e t r e s t r i c t e d r a t s r e c e iv e a d d i t i o n a l fo o d ,
th e y m ature and d e v elo p n o rm a lly .
it
grow th
The r e s t r i c t e d r a t s a p p a r e n tly rem ain
and a r e more a c t i v e
norm al d i e t .
checks
g ro w th
seems to
From th e s e r e s u l t s
te n d s
to
be th e f a c t o r
s h o rte n
(C o m fo rt, 1961)
life
sp an ,
and
o f g r e a t e s t im p o rtan c e on
T h is id e a was a ls o m a in ta in e d by Brody (1 9 4 5 ).
Growth a n d e f f l n i A n n v
E ffic ie n c y
e n erg y
u tiliz a tio n
fa c to rs ,
in v o lv e d .
to
in
a n im a l p ro d u c tio n
or in
te rm s
of
can be e v a lu a te d i n
n e t m o n e ta ry
te rm s o f
in c o m e w hen o t h e r
i n a d d i t i o n t o th e b io e n e r g e tic a s p e c ts o f p r o d u c t i v i t y ,
I n some c irc u m sta n c e s b io e n e r g e tic a s p e c ts a r e
c u r r e n t m onetary
e ffic ie n c ie s
v a lu e s .
among s p e c i e s ,
However,
b re e d s
a b e tte r
or
tra n sfo rm e d
b a s is f o r
s tra in s
en v iro n m en ts w ould be i n te rm s o f energy u t i l i z a t i o n .
in
a re
com paring
p a rtic u la r
F lu c tu a tio n s i n
8
economic f a c to r s a re l e s s s ta b le fo r e v alu atin g animal e ffic ie n c y , and
th e re fo re of lim ite d value except fo r sh o rt periods o f time.
W ith r e s p e c t to grow th, e n e r g e t i c e f f i c i e n c y can be e v a lu a te d
by
th e p r o p o r tio n o f grow th a c h ie v e d (work p e rfo rm ed ) r e l a t i v e t o th e f r e e
en erg y expended (B rody, 1945).
e x p re s s e d i n te rm s o f grow th.
In g e n e r a l, an im al e f f i c i e n c y can be
However, two a s p e c ts m ust be c o n s id e re d :
( I ) th e grow th w hich i s r e l a t e d to developm ent (assu m in g a maximum i s
re a c h e d a t a d u l t a g e ) which i s c u m u la tiv e ly in c r e a s e d u n t i l a d u lth o o d
and l a t e r m a in ta in e d
(m eat p r o d u c t i o n ) ,
( 2 ) o th e r
and
a s p e c ts
of
grow th, n o t d i r e c t l y a s s o c ia te d w ith dev elo p m en t, which c o u ld be c a l l e d
c y c lic grow th, o c c u r r in g a t any p e r io d i n th e l i f e
c y c le o f a n a n im a l,
and r e l a t e d t o a c e r t a i n p ro d u c tiv e component ( i . e . m ilk , e g g s, w ool,
e tc ).
R ep ro d u ctiv e t r a i t s would be a p a r t i c u l a r case because o f t h e i r
i m p l i c i t dependence on o th e r f a c t o r s b e s id e s th e an im al i t s e l f
m a tin g ).
(i.e .
In g e n e r a l, th e s y n th e s is o f any p ro d u c t w ith in th e o rganism
co u ld be c a ll e d grow th.
E ffic ie n c y ,
a c h ie v e d
over fr e e
a c c o r d in g t o Brody
(1945)
en erg y expended.
Some o f
n o rm ally d i s s i p a t e d a s h e a t .
and e v a lu a te i n
g ro w th ).
is
th e
th e
ra tio
o f grow th
expended e n erg y i s
E n e rg e tic e f f i c i e n c y i s e a s i e r to m easure
o rganism s a t
e m b ry o lo g ic a l
s ta g e s
( d u r in g
p re -n a ta l
However, a f t e r b i r t h , due to th e in f lu e n c e o f many v a r i a b l e s ,
t h a t ta s k i s more d i f f i c u l t (B rody , 1945) .
A n im a ls u s e
fo o d
e n erg y
to
m eet
s u rv iv a l
re q u ire m e n ts w hich
in c lu d e m a in te n a n c e , d ev elo p m en tal g ro w th , r e p r o d u c tio n and p ro d u c tio n
a fte r
m a tu rity .
Holmes (1977)
s ta te s
th a t
th e e s tim a tio n o f fe e d
9
e f f ic ie n c y sh ould be done i n r e l a t i o n to th e l i f e t i m e a n im al y i e l d and
th e t o t a l in p u ts o f fe e d r e s o u r c e s .
D iff e re n c e s i n an im al e f f i c i e n c y a r e r e l a t e d i n v a ry in g d e g re e s t o
th e g e n e tic make-up o f th e a n im a ls .
g e n e tic v a r i a t i o n .
For exam ple,
T h is i s q u a n t i f i e d i n te rm s o f
s e l e c t i o n f o r c le a n wool w e ig h t p e r
h ead r e s u l t s i n a c o r r e l a t e d in c r e a s e i n e f f i c i e n c y (T u rn e r and Young,
1969)•
There a r e ,
e ffic ie n c y .
g e n e tic
how ever, few e s ti m a te s o f g e n e ti c p a ra m e te rs f o r
T e rrill
d iffe re n c e s
n u trie n ts ,
( 1975)
in d ic a te s
th a t l i t t l e
is
known o f
among sh eep r e g a r d in g r e q u ir e m e n ts
of
th e
s p e c ific
and t h a t th e a v a i l a b l e in fo r m a tio n comes from r e s e a r c h w ith
d if f e r e n t o b je c tiv e s .
Most s t u d i e s have u sed u n i t s o f fe e d consumed p e r u n i t o f w e ig h t
g a in ed a s a m easure o f e f f ic ie n c y and h e r i t a b i l i t y e s tim a te s c i t e d by
T e rrill
(1975)
w ere o b ta in e d
by B o tk in
(1 9 5 5 ),
W itt e t
V esely and R obison (1968) and E rcan b rack (1 9 7 4 ).
a l.
(1 9 6 7 ),
L a s le y (1 9 7 8 ) g iv e s
.20 to .2 5 a s th e a v erag e h e r i t a b i l i t y f o r t h i s m easu re o f e f f i c i e n c y
i n sh eep .
I n o th e r s t u d i e s ,
due t o th e d i f f i c u l t y
o f m e asu rin g d a i l y
food
consum ption p e r a n im a l, th e a n a ly s e s o f e f f i c i e n c y f o r w e ig h t g a in have
u sed body w e ig h ts g a in s i n s p e c i f i c p e r io d s o f g ro w th .
has
been made
th a t
th e
g e n e ti c and p h e n o ty p ic
The a ssu m p tio n
c o rre la tio n s
b e tw ee n
av erag e d a ily w eig h t g a in s and f e e d e f f i c i e n c y i n sh eep a r e h ig h (-.7 3
and - . 6 0 , r e s p e c t i v e l y ) a c c o rd in g t o L a s le y (1 9 7 8 ).
and d i g e s t i b i l i t y
a r e h ig h ly
p ro d u c tio n e f f i c i e n c y
has
c o rre la te d
a ls o
been
g e n e tic d i f f e r e n c e s d e m o n stra te d .
Feed e f f i c i e n c y
( P r ic e e t a l . ,
s tu d ie d
and
th e
1 965) .
e x is te n c e
Wool
of
Even a t low l e v e l s o f fe e d in ta k e
10
T e r r i l l (1975) c o n s id e rs th e p o s s i b i l i t i e s of e f f i c i e n c y im provem ent by
s e l e c t i o n t o be f e a s i b l e .
S tu d ie s r e g a r d in g th e e f f i c i e n c y o f r e p r o d u c tiv e p erform ance a r e
sc a rc e .
Yet th e m ain l i m i t i n g f a c t o r i n
in d u s tr y
is
th e low r e p r o d u c tiv e r a t e
Holmes (1977) i n d i c a t e d
im p o rta n t t r a i t ,
which
th e
( B la x te r ,
th a t re p ro d u c tiv e
can be m easured i n
success of
th e
sh eep
1968; T u rn e r, 1969) .
p e rfo rm a n c e
te rm s o f
is
its
a v e ry
c o n s ti tu e n t
p a r t s ; age a t f i r s t b re e d in g , le n g th o f r e p r o d u c tiv e c y c le , r e g u l a r i t y
of
b re e d in g ,
litte r
s iz e
and p e r i - n a t a l
d o u b lin g th e number o f progeny i n
50$.
A re p ro d u c tiv e
in d e x
m o rta lity .
He s t a t e s
th a t
sh eep w ould in c r e a s e e f f i c i e n c y
(H o lm e s,
1 9 7 7 ),
by
a n n u a l mass o f l i v e
o f f s p r i n g born d iv id e d by th e m ass o f t h e i r dam i n te rm s o f m e ta b o lic
w e ig h t
(W 7 ) ,
s u g g e s ts t h a t
sh eep a r e
among th e
s p e c ie s w ith
th e
p o o re s t perform ance (T a b le I ) .
However, R obinson (1 9 7 4 ) p re s e n te d th e
fe a s ib ility
in d o o r
of
an i n t e n s i v e
lam bing i n t e r v a l o f 205 d a y s.
p ro d u c tio n sy stem ,
u tiliz in g
a
The c e n t r a l em p h asis was l i t t e r s iz e and
fre q u e n c y o f b re e d in g , m a n ip u la te d by c o n t r o l l i n g n u t r i t i o n , l a c t a t i o n ,
and day l e n g th ; w ith in d iv id u a l f e e d in g a c c o rd in g t o
p ro d u c tio n o f th e ew es.
I n t e r e s t was a l s o
th e
le v e l
of
c e n te re d on r e p r o d u c tiv e
e f f ic ie n c y i n t h i s i n te n s iv e sy stem .
From a p r a c t i c a l p o in t o f view , th e com m ercial p ro d u c e rs ' i n t e r e s t
i s t o m axim ize n e t r e t u r n s
( e f f i c i e n c y i n te rm s o f in c o m e ).
At t h i s
p o in t, b a la n c e m ust be fo u n d among th e more im p o rta n t p r o d u c tiv e t r a i t s
a s to t h e i r r e l a t i v e im p o rtan c e ( i n c r e a s e i n n e t r e t u r n p e r u n i t change
in a s p e c ific t r a i t ) .
I h a t b a la n c e u s u a lly in v o lv e s r e p r o d u c tiv e r a t e ,
wool and m eat (R ae, 1982).
The i d e a l c o n d itio n ,
in
te rm s o f grow th
11
would be t o
produce f a s t e r
grow ing a n im a ls w ith low m ature
red u ce m a te rn a l c o s ts (R o b e rtso n , 1 9 8 2 ).
be an e a r l i e r
age
o v u la tio n s
th e
M ight,
in
1967).
at
p u b e rty
s iz e
A c o r r e l a t e d a d v an ta g e
an d h i g h e r
p ro p o rtio n
ewe lam bs t h a t re a c h e d p u b e rty
to
would
o f m u ltip le
e a rlie r
(Lang and
T ie rn e y (19 6 9 ) a ls o s u g g e s ts t h a t s e l e c t i o n f o r
ewes
h a v in g f i r s t
e s tr o u s a t a young age
can r e s u l t i n in c r e a s e d o v e r a l l
fe rtility .
A s im ila r
a ls o
re s u lt
has
been r e p o r te d
in
c a ttle
( L e sm e iste r e t a l . , 1973) .
A n e g a tiv e r e l a t i o n s h i p e x i s t s
betw een s i z e and wool p ro d u c tio n
,■ I
:.
(T u rn e r and Young, 1969) due t o th e p o s i t i v e r e l a t i o n s h i p betw een s iz e
and s u r f a c e a r e a o f s k in .
i n wool p ro d u c tio n .
R ie r e f o r e , sm a ll s i z e r e s u l t s i n a r e d u c tio n
However, a g r e a t e r number o f sm all a n im a ls c o u ld
be r a i s e d p e r u n i t o f la n d (o r fe e d a v a i l a b l e ) which w ould com pensate
f o r th e n e g a tiv e e f f e c t o f in d iv id u a l s iz e on wool p ro d u c tio n .
M odels f o r grow th e v a l u a ti o n
The r e p r e s e n t a t i o n o f grow th i n te rm s o f body w eig h t was d is c u s s e d
by Brody (1 9 2 7 a ).
r e p r e s e n te d :
He i n d i c a t e d
th r e e ways t h a t
th e s e
c o u ld
be
( I ) a v e l o c i t y c u rv e ; ( 2 ) a c u m u la tiv e cu rv e r e p r e s e n t i n g
th e sums o f a l l g a in s ; and ( 3 ) a r e l a t i v e - r a t e
c u rv e ).
g ro w th
He recommended th e
th re e .
The a d v a n ta g e
c u m u la tiv e
cu rv e
( o r p e rc e n ta g e
c u rv e a s th e m ost u s e f u l
t h a t B ro d y
saw f o r
th e
c u m u la tiv e grow th r e p r e s e n t a t i o n was m ain ly r e l a t e d t o i t s
in te rp re ta tio n :
( I ) fo r e a rly
grow th o f d i f f e r e n t a n im a ls ;
com parison o f th e r e l a t i v e
use
of
of
th e
b io lo g ic a l
p ro g r e s s o f
( 2 ) r e p r e s e n t a t i o n o f grow th s ta n d a r d s ;
and (3 ) f o r p u rp o s e s o f i n t e r p o l a t i o n , because a c c id e n ta l v a r i a t i o n and
e x p e rim e n ta l e r r o r s can be m inim ized .
12
TABLE I .
C hicken
Turkey
R ab b it
T y p ica l
m ass o f
dam
kg
2 .5
10.0
W-75
kg
Progeny
m ass a t
b irth
kg
No. o f
progeny
per y ear
1.99
.05
100
5.62
4 .5
REPRODUCTIVE INDEX3
.10
3 .0 9
.05
Ewe
Cow
170.0
70.0
500.0
47.1
2 4 .2
105.7
a A fte r Holmes, 1977.
.
R e p ro d u c tiv e
In d e x
kg
kg* 7 5
2.0
2.51
3 .7 7
6 .2 8
150
250
5 .0
7 .5
12 .5
50
80
5 .0
.5
8.0
.8
1.0
2.0
.44
20
40
Sow
Progeny
mass
per y ear
kg
3 .0
5 .0
.22
1 .5
12
1.2
24
4 .5
4 .0
3 .0
2
4 .5
.06
8.0
.12
3
9 .0
.13
I
42 .0
7 0 .0
.08
.14
42.0
35.0
I
2
18.0
2 8 .8
.11
.17
.89
1.4 2
.32
.6 5
.3 8
.61
.1 9
.33
.37
.4 0
.66
13
Ih e use o f th e c u m u la tiv e cu rv e gave r i s e to te rm in o lo g y which i s
b a s ic t o
th e
u n d e rs ta n d in g o f
a n a l y s i s o f grow th.
th e
g r a p h ic a l r e p r e s e n t a t i o n a n d
That te rm in o lo g y ,
th e
a p p lie d t o th e model u sed by
Brody ( 1 9 2 7 a ,d ) f o r th e d e f i n i t i o n o f th e p o s t- p u b e r a l phase o f grow th
(s e lf-in h ib itin g ),
in v o lv e s
th re e
p a ra m e te rs .
IVo o f
them have
Im p o rta n t b io lo g ic a l m eaning: ( I ) m ature s iz e (A ), w hich i s a ls o c a ll e d
a sy m p to tic w e ig h t; and ( 2 ) th e s lo p e (k ) which i s th e r e l a t i v e v e l o c i t y
o f grow th o r th e r e l a t i v e - r a t e o f grow th w ith r e s p e c t t o th e g row th y e t
t o be made (A - W), r e p r e s e n te d by:
dW/dt
* 8
W here; W = W eight a t age t
t = Age i n tim e u n i t s
A = M ature w e ig h t.
'
I b . t h i r d p a ra m e te r (B ) , which h a s no b io l o g i c a l a l g U f l c a . c e ,
a g e -p a ra m e te r r e p r e a e n tln g th e I n t e r c e p t o f th e c u rv e
I, a.
(A-M) when t =0
(a t b irth ).
O th e r
m o d e l,
w e re
p ro p o se d
fo r
th e
post
in fle c tio n
(s e lf-in h ib itin g )
phase o f grow th p re v io u s t o B ro d y 's m odel.
th o ro u g h
B ro d y
s tu d y
c o n c lu d e d
th a t
th e
fo llo w in g
A fte r
n o n -lin e a r
th re e -p a ra m e te r e q u a tio n was th e m ost a p p r o p r ia te m odel:
Vt
S
A - Be“ k t
W here:
Wfc s W eight a t tim e t
A s M ature w e ig h t
B
s C o n sta n t o f i n t e g r a t i o n
e
s Base o f n a tu r a l lo g a rith m
k
s R ate a t w hich th e c u rv e a p p ro a c h e s th e a sy m p to te .
T h is e q u a tio n can be tra n s fo rm e d t o ;
14
Wt - Wt e = A - A e - k ( t - t " ) ,
i n which th e grow th cu rv e h a s been f o r c e d t o a s p e c i f i c p o in t (Wfce,
t » ) . A p a r t i c u l a r case o c c u rs when th e o r ig i n o f th e c u rv e i s s e t to
th e i n t e r s e c t i o n o f th e o r d in a te and a b s c is s a
(t* = 0 Wfce=O).
In t h i s
s p e c i a l c ase (F ig u re I ) , B e q u a ls A and th e f u n c tio n i s re d u c e d t o a
two p a ra m ete r e q u a tio n : .Wfc = A -
Ae~k t . N elsen e t a l .
(1982) u t i l i z e d
t h i s approach a s an a l t e r n a t i v e f o r a d j u s t i n g by B, f a c i l i t a t i n g
th e
d i r e c t com parison o f th e k v a lu e s among a n im a ls .
M odels o t h e r
th a n
B ro d y ’ s h a v e
a ls o
l i t e r a t u r e f o r f i t t i n g anim al grow th c u rv e s .
of
fittin g
th e
w h o le
s e l f - a c c e l e r a t i n g phase
g ro w th
c u rv e ,
(E ise n e t
a l.,
p ro p o se d
and a p p lie d
in
th e
d is c u s s e d
in
th e
Many have th e c a p a b i l i t y
in c lu d in g
th e
p re -p u b e ra l
1969; Timon and E is e n ,
Brown e t a l . , 1976; DeNise and B r in k s , 1985).
been
b een
1 969 ;
O th e r m odels have a ls o
a n a l y s i s o f grow th
c u rv e s ,
bu t
a p p a r e n tly have n o t been compared w ith o th e r s ( S p r e n t, 1967 ; W arren e t
a l . , 1980; F le tc h e r , 19 7 4 ).
The i n i t i a l th e o ry ab o u t th e m a th e m a tic a l
r e p r e s e n t a t i o n o f grow th developed by Brody (1 9 2 7 a , 1927b, 1927c, and
1928) h a s a l s o r e c e iv e d new im p o rta n t c o n tr ib u tio n s (T a y lo r,
1 9 8 0 a ,b,
1965, 1968, 1982, 1985; T ay lo r and F itz h u g h , 1971 ; F itz h u g h and T a y lo r,
1 971).
E ise n e t a l .
(1969) made a com parison among th e grow th f u n c tio n s
( L o g is t ic , Gompertz and B e r ta l a n f f y ) i n m ice from b i r t h t o 54 d o f a g e .
T h e se
g ro w th
m o d e ls
c o rre sp o n d
e q u a tio n s w ith f i x e d i n f l e c t i o n
to
th e
p o in ts .
g ro u p
of
th re e
p a ra m e te r
B ased on th e d i f f e r e n c e s i n
r e s i d u a l v a r ia n c e s , th e y found t h a t th e L o g i s t i c f u n c tio n gave th e b e s t
fit.
A lso,
th e c o e f f i c i e n t s o f
v a ria tio n of
A, t» (ag e a t p o in t o f
height
( c m)
15
HiP
A - Be
o
t
m m t's
(months )
^z:, ^ im
16
i n f l e c t i o n ) and k , e s tim a te d from th e L o g i s tic f u n c tio n w ere s m a lle r
th a n
th o s e
a c c u ra c y
c a l c u l a t e d from
of f i t
fo r
th e s e
th e
o th e r
th re e
two f u n c tio n s .
fu n c tio n s had
However,
no e f f e c t
on
th e
th e
c o n c lu s io n s a b o u t th e d i f f e r e n c e s w ith in and betw een th e t h r e e l i n e s o f
m ice s tu d ie d .
Brown e t a l , (1 9 7 6 ), u s in g d a ta from b e e f c a t t l e , made a
c o m p a riso n o f f i v e
G e n e ra liz e d
n o n - lin e a r m odels
L o g is tic ,
th re e -p a ra m e te r
and
fu n c tio n s
(B rody,
R ic h a rd s ).
w ith
fix e d
B e r ta l a n f f y ,
T he
firs t
in fle c tio n
G cm pertz,
th re e
p o in ts .
a re
The
G e n e ra liz e d L o g i s t i c h a s a v a r ia b le p o in t o f i n f l e c t i o n and R ic h a rd s i s
a f o u r p a ra m e te rs f u n c tio n w ith a v a r i a b l e p o in t o f i n f l e c t i o n .
concluded t h a t th e R ic h a rd s ' f u n c tio n gave a more a c c u r a te f i t ,
c o m p u ta tio n a lly more d i f f i c u l t th a n th e o th e r s .
They
b u t was
R ic h a rd s ' m odel, due
t o i t s v a r i a b l e p o in t o f i n f l e c t i o n , was m ost a p p r o p r ia te f o r f i t t i n g
c a t t l e w e ig h t-a g e d a ta p r i o r to 10 months o f a g e .
B ro d y 's model gave
r e s u l t s s im ila r t o t h a t o f R ic h a rd s f o r w e ig h t-a g e d a ta p a s t s i x m onths
and had th e added a d v an ta g e o f b e in g c o m p u ta tio n a lly e a s i e r .
However,
c o rre la tio n
th e a u th o r s
b e tw e e n t h e
c ite d
v a lu e s
above
re p re s e n tin g
e s tim a te d from th e d i f f e r e n t m o d els.
f i t v is u a lly ,
d id n o t f i n d
th e
a
n e ar-p e rfe c t
sam e p a ra m e te rs
They e v a lu a te d th e g o o d n ess o f
p r e f e r r i n g t h i s ap p ro ach to th e u s u a l t e s t o f g o odness
based on r e s i d u a l v a r ia n c e s .
T h is was due t o th e i m p l i c i t in a c c u ra c y
e x p e c te d from th e c o r r e l a t e d e r r o r s (dependency) among t h e l o n g i t u d i n a l
grow th d a ta .
(1 9 7 6 ).
The same l i m i t a t i o n was a l s o
in d ic a te d
by F i t z h u g h
These a u th o r s con clu d ed t h a t th e s e l e c t i o n o f any one o f th e
m odels would depend upon th e n a tu r e o f th e stu d y t o
w e ll a s , th e in te n d e d a p p l i c a t i o n o f th e r e s u l t s .
be p erfo rm ed , a s
17
Timon and E ia e n (1969) o b se rv e d i n m ice d a ta t h a t th e f i t o f th e
R ic h a rd s and th e L o g i s t i c f u n c tio n s w ere v e ry s i m i l a r .
However,
th e
a sy m p to tic w e ig h t was u n d e re s tim a te d , m ainly by th e L o g i s t i c f u n c tio n .
B oth f u n c tio n s
gave a mean p o in t o f i n f l e c t i o n
at
n e a r ly
h a lf
th e
a sy m p to tic w e ig h t ( A /2 ), which i s c o n tr a d ic to r y t o th e v iew s o f Brody
(1945) and T a y lo r (1 9 6 5 ).
G re a t d i f f e r e n c e s w ere o b serv ed betw een th e
k
each
v a lu e s
o b ta in e d
fro m
of
th e s e
c o r r e l a t i o n was z e r o w hich s u g g e s te d
r e p r e s e n te d d i f f e r e n t t r a i t s
fu n c tio n s .
th a t
th e
The
g e n e tic
tw o e s t i m a t e s
by th e two f u n c tio n s .
of
k
T hat o b s e r v a tio n
c o n tr a s te d m arkedly w ith th e v e ry h ig h g e n e tic c o r r e l a t i o n s betw een the
o th e r
v e ry
d e riv e d grow th p a ra m e te rs .
c lo s e r e l a t i o n s h i p
( r g=.91 and r p= .9 5 ) .
A nother im p o rta n t f i n d i n g was th e
betw een th e
R ic h a rd s '
The m p a ra m e te r i s
k a n d m p a r a m e te r s
c a l l e d th e shape p a ra m e te r,
r e p r e s e n tin g a p o in t i n th e c u rv e w here i n f l e c t i o n ta k e s p la c e w hich i s
e x p re ss e d a s a r e l a t i v e m easure t o th e l i m i t o f th e c u rv e .
DeNise and B rin k s (1985) compared th e g o o d n ess o f f i t o f B ro d y 's
and R ic h a rd s ' f u n c tio n s u s in g d a ta from b e e f c a t t l e .
They co n clu d ed
t h a t R ic h a rd s ' f u n c tio n had s m a lle r r e s i d u a l v a r ia n c e and a b e t t e r f i t
to th e a c t u a l d a ta p o in ts , how ever, B ro d y 's c u rv e was f a s t e r and l e s s
c o s tly to com pute.
P itz h u g h
and T a y lo r
(1971) d e v e lo p e d a g e n e r a l method f o r
th e
a n a l y s i s o f g e n e ti c r e l a t i o n s h i p s in v o lv in g m a tu r ity r a t e ,
s iz e ,
grow th r a t e w ith o th e r t r a i t s in v o lv in g d e g re e o f m a tu r ity .
They used
a sim p le tw o-com ponent,
e q u a ti o n - f r e e model w here th e
and
two components
a r e s ( I ) m ature s i z e (A ), and (2 ) p r o p o r tio n o f m atu re s i z e re a c h e d a t
a s p e c i f i c s ta g e o f grow th ( u ) .
The F itz h u g h and T a y lo r m odel, u sed
18
f o r a n a ly z in g
g e n e tic r e l a t i o n s h i p s
among m a t u r i n g r a t e ,
s iz e
and
grow th r a t e , i s :
* t - ^ t* ,
w here,
Yt = s i z e a t age t (e x p re s s e d a s a f u n c tio n o f d e g re e o f
m a tu r ity u and m atu re s i z e A),
u s
p r o p o r tio n o f m atu re
s ta g e o r age t ,
s i z e A a t t a i n e d a t a g iv e n
and,
Ufc = YfcZA.
The grow th
s ta tis tic s
T a y lo r (1971) a r e :
p r o p o r tio n a l
(I)
d e riv e d
o r a p p ro x im ated
A b so lu te M atu rin g B ate
change i n
th e
d e g re e
consequence o f
th e m a tu rin g r a t e
or
an d
(AMR), d e fin e d a s th e
o f m a tu rity
e x p re s s e d a s p e rc e n ta g e i n a g iv e n i n t e r v a l :
by F it z h u g h
per
u n it
AMR = d u /d t.
change i n
u over
of
tim e
AMR i s
tim e f o r
a
th e
t r a i t , and can a l s o be e x p re s s e d a s :
AMR = 1/A (d Y /d t) = 1/A (Yfc2 _ Yfc1) / ( t 2 - t , ) .
(2 ) A b so lu te Growth
R a te
(AGR)
is
th e
change
per
u n it
of
tim e ,
e x p re s s e d i n k ilo g ra m s and commonly r e f e r r e d a s a v e ra g e d a ily g a in :
AGR = dY /dt = (Yfc2 - Yfc1) / ( t 2 - I 1) ,
(3 ) R e la tiv e Growth R ate (RGB) r e f e r s t o th e change i n th e p r o p o r tio n
o f s i z e p e r u n i t o f tim e i n a g iv e n i n t e r v a l ,
a t t a i n e d p e r u n i t o f tim e i n a g iv e n i n t e r v a l
a ls o
o r p e rc e n ta g e o f s i z e
: RGR = l / i d Y /d t. May
be re g a rd e d a s A b so lu te Growth R ate (AGR) r e l a t i v e t o s i z e o v er
th e p e rio d o f i n t e r e s t :
RGR = [(Y fc2 - Yfc1) / ( t 2 - t , ) ]
(V Y fc2) .
19
S to b a r t (1983) makes a c l e a r I n t e r p r e t a t i o n o f th r e e b io l o g i c a l l y
I m p o r ta n t
g ro w th
s ta tis tic s
d e riv e d
fro m
t h a t m o d e l,
and
th e ir
a p p lic a tio n i n th e a n a l y s i s o f grow th and c o r r e l a t e d re s p o n s e s on ewe
p r o d u c tiv ity ,
He s t a t e s t h a t ,
"The m ajo r a d v a n ta g e s o f F ltzh u g h and
T a y lo r (1971) method o v e r th e f i t t i n g o f grow th c u rv e s i s t h a t th e r a t e
o f m a tu rin g i s n o t fo r c e d t o rem ain c o n s ta n t o v e r tim e ; b u t r a t h e r both
g e n e tic and e n v iro n m e n ta l v a r i a b i l i t y i n m a tu rin g r a t e can be a s s e s s e d
f o r any a g e i n t e r v a l ,
and d e g re e
o f m a tu r ity
can be e s tim a te d from
few er and l e s s u n ifo rm ly sp aced m easurem ents th a n r e q u ir e d f o r f i t t i n g
o f grow th c u r v e s ."
S.snstls3_0f
grow th
tra its .
param ete r s and t h e i r
re la tio n s h ip
to
nroduntlvm
There a r e no p u b lis h e d s t u d i e s f o r sheep a s s e s s i n g th e r e l a t i o n
betw een e s tim a te d v a lu e s
su b seq u en t
p ro d u c tiv e
of
th e
g ro w th
p e rfo rm a n c e s.
r e g a r d in g th e a p p l i c a t i o n o f th e grow th
e q u a tio n
fre e
p erform ance i n
m odel
of
F itz h u g h
ewes ( S t o b a r t ,
p a ra m e te rs ;
T h ere a r e ,
s ta tis tic s
and T a y lo r
1983; S to b a r t ,
A and
how ever,
k,
s tu d ie s
d e riv e d from
(1971)
and
th e
and p r o d u c tiv e
1 9 8 5 ).
S to b a r V s
in c lu d e d a p o r tio n o f t h a t u sed i n th e p r e s e n t s tu d y ,
d a ta
He fo u n d t h a t
a n im a ls more m atu re a t any s ta g e o f grow th w ere m ore m a tu re a t l a t e r
s ta g e s o f g row th, and a l s o l i g h t e r a t m a tu r ity .
The f a s t e r m a tu rin g
a n im a ls w eighed more th a n th e o th e r s up t o 12 mo o f a g e .
g e n e ra l c o n sta n c y i n th e grow th r a t e s up t o m a tu r ity .
T here was no
A nim als grow ing
f a s t e r i n a g iv e n tim e i n t e r v a l te n d e d to grow s lo w e r i n th e i n t e r v a l
im m ediately fo llo w in g .
S e v e ra l
h e rita b ility
e s tim a te s r e p o r te d i n
p re s e n te d f o r th e grow th p a ra m e te rs k ,
th e l i t e r a t u r e
m, A and V
( th e age a t
a re
th e
20
i n f l e c t i o n p o in t) (T a b le 2) and th e grow th s t a t i s t i c s d e riv e d from th e
a p p lic a tio n o f
th e f r e e - e q u a ti o n model o f F itz h u g h and T a y lo r (1971)
(T a b le 3 ) .
TABLE 2 .
HEBITABILITY ESTIMATES FOR GROWTH CURVE PARAMETERS
-
Source
Modela Methodb S p e c ie s
E ise n e t
a l.( 1 9 6 9 )
Timon &
E ise n
(1 9 6 9 )
L
G
Be
R
L
FS
Mice
FS
Mice
Age
B-54d
B-54d
B-54d
5-9 8 d
5-98d
E s tim a te s
A
t'
k
.50
.7 4
.81
•30±,1
.7 6 ± ,2
.47
.40
.51
.6 6 i> 2
.7 6 ± ,2
Brown e t a l . (1972)
B
PHS
C a ttle
B-109mo • 33±>2
B-109mo .7 5 ± ,3
-34± ,2
.2 1 ± ,2
DeNise &
B rin k s
(1985)
B
R
PHS
C a t tle
B -4 y r
B -4 y r
.4 4 ± ,3
.4 4 ± ,3
.2 0 ± ,3
.3 2 ± ,3
.80
.7 5
.85
1 .17± ,1
I .3 2 ± ,2
■m
.5 3 ± ,2
•
.2 1 + .3
J B=Brody, L = L o g is tic , G=Gcmpertz, B e = B e rta la n ffy , R =R ichards.
E s tim a tio n m ethod: FS=F u l l - s i b s, PH S=Paternal h a l f - s i b s.
G re at
v a ria tio n
e x is te d
among t h e
h e rita b ility
e s tim a te s .
D iff e re n c e s due to s p e c ie s , b re e d and e n v iro n m e n ta l c o n d itio n s co u ld be
re a so n s fo r th a t v a r i a b i l i t y .
The d i f f e r e n t m e th o d o lo g ie s a p p lie d i n
th e e s tim a tio n o f th e h e r l L a b i l i t i e s a r e a ls o p o s s ib le f a c t o r s .
Bie
e s tim a te d h e r i t a b i l i t i e s f o r a l l th e c u rv e p a ra m e te rs , a s w e ll a s th e
grow th s t a t i s t i c s ra n g e from in te r m e d ia te t o h ig h v a lu e s , s u g g e s ts t h a t
th e s e t r a i t s a r e s u s c e p tib le to change by s e l e c t i o n .
S to b a r t
(1983) fo u n d
th e
h ig h e s t
h e rita b ility
e s tim a te s
fo r
A bsolute M atu rin g R ate i n th e i n t e r v a l s b i r t h - 1 2 mo, b i r t h - 1 8 mo, and
w e a n in g - 1 2 mo,
( .6 6 ± ..1 2 ,
.59±..12 and
.64±_.12,
re s p e c tiv e ly ).
h e r i t a b i l i t y e s tim a te f o r AMR i n th e i n t e r v a l 12 - 18 mo was .3 2 .
H is
21
TABLE 3 .
HERITABILITY ESTIMATES FOR GROWTH STATISTICS8
E s tim a te s
Methodb
S p e c ie s
F itz h u g b
& T a y lo r
(1971)
PHS
C a ttle
S n ith e t
a l.( 1 9 7 6 b )
PHS
C a t tle
B-200d
2 0 0 -3 96d
396-550d
5 5 0 d -3 .3 3 y r
.5 7 ± ,3 0
.4 2 ± ,3 0
.6 9 1 ,3 1
.1 9 1 ,2 8
.6 7 1 ,3 1
- .0 7 1 ,2 5
.6 9 1 ,3 1
.1 9 ± ,2 8
.6 2 1 ,3 1
.3 1 1 ,2 9
.4 4 i,3 0
•09± ,27
S to b a r t
(1983)
PHS
Sheep
B-V
M 2m o
B - 18mo
W-12mo
12-1 Smo
.2 1 i,1 1
.2 9 1 ,1 1
.3 5 ± ,1 2
. 5 2 t ,1 2
.4 3 1 ,1 2
.2 8 i,1 1
.5 0 i,1 2
.4 4 i,1 2
.4 8 i,1 2
.3 9 1 ,1 2
.1 9 1 ,1 1
.6 6 i,1 2
.5 9 1 ,1 2
.6 4 i,1 2
.3 2 1 ,1 1
Source
Age
P r e n a ta l
B-6 mo
6-12mo
12-18 mo
I 8 -m atu re
AGR
RGR
AMR
.38
.40
.45
.3 5
.48
.27
.47
.24
.42
.22
.42
.46
.24
.42
aA ll p a p e rs u sed F itz h u g h and T a y lo r (1 971) m ethodology.
^ E s tim a tio n m ethod: PH S=Paternal h a l f - s i b s.
In a d d itio n
to
th e
a p p lic a tio n
of
th is
in fo r m a tio n
th r o u g h
s e l e c t i o n t o d i r e c t l y change m a tu rin g r a t e , m atu re s i z e and g ro w th , i t
is
d e s ir a b le t o have knowledge o f th e e x p e c te d c o r r e l a t e d re s p o n s e o f
such s e l e c t i o n i n o th e r t r a i t s o f b io l o g i c a l and eco n o m ical im p o rta n c e ,
p a rtic u la rly
re p r o d u c tiv e
p e rfo rm an c e.
T a b le s
4 an d
5 s u m m a riz e s
p u b lis h e d e s ti m a te s o f g e n e ti c c o r r e l a t i o n s among th e s e p a ra m e te rs and
grow th s t a t i s t i c s , r e s p e c t i v e l y , and t h e i r c o r r e l a t i o n s w ith p r o d u c tiv e
tra its .
The r e s u l t s
in
T a b le s
4 and
re la tio n s h ip
betw een m a tu re s i z e
re la tio n s h ip
has
been
5 in d ic a te
(A) and r a t e
h ig h lig h te d
in
o th e r
a
n e g a tiv e
g e n e tic
o f m a tu rin g ( k ) .
s tu d ie s .
T a y lo r
T h is
and
TABLE 4 .
GENETIC CORRELATIONS BETWEEN GROWTH PARAMETERS AND PRODUCTIVE TRAITS
E s tim a te s
Source Modela Methodb S p e c ie s T r a i t Age
E ls e n e t L
a l.( 1 9 6 9 )
FS
Timon A
E ise n
(1969)
FS
R
Mice
Mice
L
k
A
.55±>5
- .5 0 ± ,5
.54±>5
k
AGR
B-54d
B-54d
k .
A
k
5-98d
5-98d
5_98d
k
A
B-109oo
B-109mo
—. 62±_,3
- .2 8 ± ,5
- .52±_, 6
Brown e t B
a l . (1972)
PHS
DeNise e t B
a l . (1983) R
PHS
C a ttle
MPPAc B-TSmo .6 8 ± j,T
B-TSmo I .3 2 ± J .
DeNise A B
B rin k s
R
(1985)
PHS
C a ttle
k
A
k
A
B
C a ttle
'
B -4 y r
B -4 y r
B -4 y r
B -4 y r
B -4yr
- .2 9 ± ,3
B
m
.9 1 ±,0
- .1 1 ± ,2
-.34± > 2
• 9 8 ± j.2
-.5 0 ± > 4
-1 .1 6 ± 2 .
- .8 4 ± j.
.8 2 £ l.
-.8 4 ± ji5
-•9 T ± 2 .
• 31±..6
^Growth F u n c tio n s : B=Brody, R =R ichards, L = L o g is tic .
" E s tim a tio n m ethod: FS= F u l l - s i b s, PHS= P a te r n a l h a l f - s i b s.
0Most P ro b a b le P ro d u cin g A b i l i t y f o r w e ig h t o f c a l f a t w eaning.
.
■ '
1 .1 0 ± ,3
- .5 0 ± 1 .
- .9 8 ± 2 .
23
TABLE 5 .
TRAITSa
GENETIC CORRELATIONS BETWEEN GROWTH STATISTICS AND PRODUCTIVE
E s tim a te s
Source
Methodb S p e c ie s T r a i t
Smith e t PHS
a l . ( 1976)
S to b a r t
(1985)
PHS
Age
AGR
C a t t l e Growth0 B -20Od
- • 5 9 ± j.5
ra te
200-396d
.80 ±»5
396-550d
- . 1 1±#4
5 5 0 d -3 .3 3 y r .3 6 ± ,7
Sheep
RGR
AMR
- .3 4 ± ,4
- .0 2 ± ,4
.6 3 ± ,7
-• 3 5 * ,4
•94±,6
- .0 6 t* 5
.80 ±3.
Average*1 B-W
f le e c e
W-12mo
w eig h t
12-18mo
B-12mo
B-ISmo
- .8 9 ± ,4
.6 3 ± ,2
•36±>3
•09± ,3
•27±>3
- .7 8 ± ,4
.68± j,3
-•31±>3
- . 18±j 3
.3 8 ± ,3
-•74± j,5
•64± ,2
-•38±>3
•2 4 * ,2
- .0 2 ± ,2
Average
number
o f lam bs
b orn
B-W
W-12mo
12-18mo
B-12mo
B-ISmo
• 19±>3
—.06±#2
•3 6 ± ,2
-*22±j,3
.1 0 ± ,2
- .3 7 ± ,3
- .0 1 ± ,2
-•32± #2
- .2 6 ± ,2
- . 1 8±j>2
- .1 1 * ,3
- .0 2 ± ,2
•40±j,3
-•0 7 * ^ 2
.1 6 ± ,2
Average
number
of
lambs
weaned
B-W
W-12mo
12—18mo
B -I2mo
B - I 8mo
- .3 0 ± ,7
.0 0 ± ,4
.1 7 ± ,5
- .20±>6
.0 0 ± ,5
- I •05±#9
.0 6 ± ,5
.0 4 ± ,5
-.68± > 6
- .7 7 ± ,7
- . 1 1±>7
•02±,4
.0 8 ± ,5
-.0 2 ± j,4
•04± ,4
Average
w eig h t
lambs
weaned
B-V
W-12mo
12—18mo
B - I 2mo
B-ISmo
1 .1 1 ± 2 .
- .0 5 ± ,7
•98± 2.
• 59± J.
1 .3 2 ± 2 .
- .9 7 ± 2 .
-• 1 7 t# 7
.4 7 ± J .
-•8 3 * 3 e
-•7 7 * 3 .
1 .0 7 * 2 .
.01 ±,6
.9 3 * 2 .
.44*^9
. 1 .7 7 * 2 .
-
^From a p p l i c a t i o n o f F itz h u g h and T a y lo r (1971) m ethod.
“E s tim a tio n m ethod: PHSsP a te r n a l h a l f - a l b s .
C o r r e la tio n s a r e betw een grow th r a t e a t d i f f e r e n t a g e - i n t e r v a l s and
th e grow th s t a t i s t i c s a t p u b e rty .
C o r r e la tio n s betw een p r o d u c tiv e t r a i t s w ith grow th s t a t i s t i c s a t
d iff e r e n t a g e -in te rv a ls .
24
F itz h u g h (1971) r e f e r s to t h i s c h a r a c t e r i s t i c r e l a t i o n s h i p a s th e b a s ic
d e te rm in a n t i n
th e
g e n e tic change i n
fle x ib ility
ra te
shape o f
th e
which i s
th e
grow th
c u rv e ,
in d ic a tin g
th a t
th e
shape o f th e grow th c u rv e w i l l depend on i t s
based on th e d e g re e o f in dependence among s i z e ,
o f m a tu rin g and i n f l e c t i o n
t h a t i n H e re fo rd fe m a le s 78% o f
p a ra m e te rs .
For exam ple,
th ey fo u n d
th e a d d i t i v e v a ria n c e f o r
th e tim e
ta k e n t o m atu re was in d e p e n d e n t o f m a tu re w e ig h t.
There a r e two b a s i c a l l y d i f f e r e n t p a t t e r n s o f grow th a c c o rd in g t o
( 1 972 ) :
( I ) th e
a n im a ls have s i m i l a r m a tu re w e ig h ts bu t
k v a lu e s ,
and (2 )
th e
Brown e t a l .
d iffe re n t
d i f f e r e n t m atu re w e ig h ts .
a n im a ls have s i m i l a r k v a lu e s bu t
In both c a s e s , th e v a r i a b l e w hich a c c o u n te d
f o r th e d if f e r e n c e s i n k o r A i s th e le n g th o f tim e r e q u ir e d t o a t t a i n
m a tu re w e ig h t.
The k v a lu e i s i n d i c a t i v e o f d i f f e r e n c e s i n grow th r a t e
o n ly when two a n im a ls re a c h s i m i l a r m a tu re w e ig h ts .
w e ig h ts a re d i f f e r e n t ,
When th e m a tu re
th e k v a lu e m easu res th e d i f f e r e n c e s i n grow th
r a t e r e l a t i v e to m a tu re s i z e .
When th e R ic h a rd s m odel was u se d ,
th e cu rv e shape p a ra m e te r (m)
showed a v e ry h ig h p o s i t i v e g e n e tic c o r r e l a t i o n ( > . 90 ) w ith th e r a t e
p a ra m e te r (k ) ( Timon and B is e n , 1969 ; DeNise and B r in k s , 1985) .
s u g g e s ts t h a t many o f
a ls o in f l u e n c e
th e
T h is
th e gen es w hich in f lu e n c e th e m a tu rin g r a t e (k )
shape
or in fle c tio n
p a r a m e te r
(m ).
p a ra m e te rs may r e f e r to th e same u n d e rly in g g e n e ti c t r a i t ,
a h ig h c o r r e l a t i o n e x i s t s
betw een k and t*
T h e se
s i m ila rly ,
(ag e a t i n f l e c t i o n
when th e L o g i s t i c model was a p p lie d (E is e n e t a l . , 1969) .
two
p o in t)
25
Assuming t h a t k and m a re in flu e n c e d p r im a r ily by th e same g e n es,
fo r
p ra c tic a l
m odel
(i.e .
fa s te r,
a p p lic a tio n
in
anim al
b re e d in g ,
Brody) would be a d v a n ta g e o u s.
more econom ical d e r i v a t i o n
th e
Its
in fo rm a tio n
use
of a
s im p le r
use w ould p e rm it a
th a t
is
e a s ie r
to
in te rp re t.
There a r e
c a s e s i n w hich th e co rre sp o n d e n c e betw een th e grow th
p a ra m e te rs was n o t a s c lo s e a s e x p e c te d when two d i f f e r e n t m odels w ere
a p p lie d to th e same d a ta .
The g e n e ti c c o r r e l a t i o n s betw een th e A, B
and k grow th p a ra m e te rs e s tim a te d from Brody and th o s e e s tim a te d from
R ic h ard s by DeNise and B rin k s (1985) a r e a t y p i c a l exam ple.
However,
in
g e n e tic
a n o th e r
p u b lic a tio n ,
co rre sp o n d e n c e
betw een th e
( Timon an d
Bi s e n ,
1 9 6 9 ),
th e
grow th p a ra m e te rs A and k e s tim a te d from
R ic h ard s and th o se e s tim a te d from th e L o g i s t i c model were s i m i l a r .
A ll th e p ro d u c tiv e c h a r a c te r s a n a ly z e d by S to b a r t (1 9 8 5 ), e x ce p t
th e
av erag e
num ber
o f la m b s
b o rn ,
w e re
p o s itiv e ly
c o rre la te d
g e n e t i c a l l y w ith th e grow th s t a t i s t i c s AGR, RGR and AMR f o r th e 12 - 18
mo i n t e r v a l .
in d ic a tiv e
Growth perform ance i n
of
th e
e w e 's
g e n e tic
in c lu d in g w e ig h t o f lam b w eaned.
th e i n t e r v a l
p o te n tia l
fo r
12
to
18 mo w as
la m b p r o d u c t i o n ,
Growth perform ance betw een w eaning to
12 mo was more h ig h ly r e l a t e d t o th e g e n e tic p o t e n t i a l f o r wool p ro d u c tio n .
26
MATERIALS AND METHODS
A nim als and E n v ir onm ental C o n d iM n n a
The d a ta f o r
th is
s tu d y
came from th e l i f e t i m e r e c o r d s o f 815
p u re b re d R am boulllet , Columbia and T arghee ewes.
The sh eep w ere r a i s e d
by th e Montana A g r ic u ltu r a l E x p e rim en ta l S t a t i o n ,
Red B lu f f R esearch
Ranch, N o r r is , M ontana, betw een I960 and 1981.
The Red B lu f f R esearch Ranch, lo c a te d a lo n g th e West s id e o f th e
M adison r i v e r , c o n ta in s a p p ro x im a te ly 5 ,0 0 0 h a , w hich a r e m o stly ra n g e
w ith some meadows.
The e le v a tio n s e x te n d from 1 ,4 0 2 to 1,890 m.
The
an n u al av erag e p r e c i p i t a t i o n i s 47 cm, and th e a v erag e te m p e ra tu re i s
a ro u n d
8
C w ith
maximum av erag e
minimum o f -5 C (D ecem ber).
o f a p p ro x im a te ly 21 C ( J u ly )
and
The u p la n d zone i s composed o f b u n cb g rass
type v e g e ta tio n and w ith b lu e bunch w h e a tg ra s s ( Azroovrum s o l o a t,m, I a s
th e dom inant s p e c ie s .
The n o rth s lo p e s a r e c h a r a c te r iz e d by san e a r e a s
o f b ru sh and t r e e s .
'
The a n im a ls w ere managed and h e rd e d on th e ra n g e a r e a s o f th e
e x p e rim e n ta l s t a t i o n and on th e h ig h m o u n tain n a tio n a l F o r e s t S e rv ic e
a llo tm e n ts .
They were g a th e re d and b ro u g h t to th e s t a t i o n h e a d q u a r te r s
o n ly d u rin g b re e d in g , lam b in g , s h e a r in g and d a ta c o l l e c t i o n tim e s .
ew es w e re
p e n -m a te d i n
s in g le -s ire
g r o u p s o f 20
to
40
each f o r
a p p ro x im a te ly 20 d i n November and December and th e n r e tu r n e d t o
ra n g e and m ass m ated t o b la c k fa c e ram s f o r 15 d .
The
th e
T h e ir lam bs w ere b o rn
i n A p ril and May.
D u rin g
th e
b re e d in g seaso n ,
mixed g r a s s and a l f a l f a
hay were
s u p p lie d f r e e c h o ic e and a t o th e r tim es d u rin g th e w in te r o n ly when th e
27
snow c o v e r was to o deep f o r g ra z in g . At th e end o f th e b re e d in g s e a s o n ,
th e
ewes w ere h e rd e d to g e th e r
lam b in g se a so n .
on th e ra n g e
u n til
th e
s ta rt
of
th e
S h o r tly a f t e r p a r t u r i t i o n , ewes and lam b s w ere p la c e d
i n 1 .3 m2 lam bing pens f o r 24 h o r m ore.
A ll th e lam bs w ere e a r - ta g g e d
and t h e i r w e ig h ts re c o rd e d w ith in t h a t i n t e r v a l .
Then, th e ewe and h e r
o f f s p r i n g w ere moved t o sm all p e n s, w ith c a p a c ity f o r a p p ro x im a te ly 8
dams and t h e i r lam b s, and m a in ta in e d th e r e f o r 2 t o 3 d a y s.
The la n b s w ere weaned i n th e f a l l
a t a n a v e ra g e age o f 128 d,
w in te re d on th e ra n c h , and fe e d .23 k g o f p r o te in su p p lem en t p e r day.
D ata.
T able 6 p r e s e n ts th e
number and d i s t r i b u t i o n
of
th e
ewes t h a t
p ro v id e d d a ta f o r th e s tu d y .
DISTRIBUTION OF THE EMES BY BREED, AGE PRODUCTION GROUP AND
B reed
Group
Year o f b i r t h ■
number o f ----------------------------------------- ---- ----re c o r d s
60 61 62 63 64 65 66 67 68 69 70 72 73 75 76
R am b o u illet
3
4
5
T arghee
3
4
5
Columbia
3
4
5
T o ta l
9 4 11 I 3
6 11 2 16 3 3 2
7 7 1 1 6 1 8
3 4 4 21
12 12 5 13 12 20 13 17 15 13
10 15
4 I 3 6
2 10 6 6
6 6 3 12
6 3 3 6 4 4 4 3
5 2 3 I
17 10 15 15 27 30 21 16 10 14 19 12
5 4 2
4 7 6
1 3 5
1 2 2
13 7 13 14 9 4
6
2
9
5
5
2
8 10
5
3
5
3
I
7
5
I
3
2
3
2
I
7
2
2
8
55 56 52 62 73 83 70 73 42 68 66 51 30 29
T o ta l
74
64
164
74
47
217
54
21
100
5
ii?
28
O nly
ew es h a v in g
at
le a s t
th re e
c o n s e c u tiv e
y e ars
of
lam b
p ro d u c tio n , fo u r y e a r s I n th e case o f wool p ro d u c tio n , w ere in c lu d e d i n
th e s tu d y .
The norm al c u l l i n g ag e f o r ewes was done a f t e r th e 5 y r o f
p ro d u c tio n ,
th u s
th e maximum age f o r
ewes i n
th is
s tu d y was 6 y r .
Ewes t h a t p ro v id e d d a ta f o r th e stu d y w ere c l a s s i f i e d a c c o rd in g to
w h eth er th ey had a t o t a l o f t h r e e , fo u r o r f i v e cum ulated r e p r o d u c tiv e
re c o rd s in
re m a in e d
o rd e r t o
in
th e
stu d y
flo c k
th e l e v e l
fo r
th e s e
of
p e rfo rm a n c e
d iffe re n t
of
p e rio d s o f
c l a s s i f i c a t i o n i s r e f e r r e d t o a s "g ro u p " i n th e t h e s i s .
th e ewes w hich l e f t th e f lo c k w ere c o n s id e re d f o r s tu d y .
ew es
tim e .
th a t
T h is
Only d a ta from
For exam ple,
f o r R am b o u ille t ewes born i n I9 6 0 , 9 l e f t th e f lo c k a f t e r 3 r e c o r d s , 7
a f t e r 4 re c o rd s and 12 a f t e r 5 r e c o r d s .
V a r ia b le s w ere
c a lc u la te d
p ro d u c in g a b i l i t y o f each ewe.
number o f lam bs
born
(ATLB),
to
e v a lu a te
th e
c u m u la tiv e
life tim e
These new v a r i a b l e s w e re ; A verage t o t a l
Average t o t a l
number o f la m b s w ean ed
(ATLW), Average t o t a l w e ig h t o f lam bs weaned (ATWW) and A verage t o t a l
w e ig h t o f g re a s e f le e c e produced (ATFP).
A minimum o f sev en body w e ig h ts w ere r e q u ir e d f o r ewes w ith th r e e
cu m u la tiv e p ro d u c tio n r e c o r d s .
The o th e r ewes w ith f o u r
and f i v e
c u m u la tiv e p ro d u c tio n r e c o r d s had e i g h t and n in e w e ig h ts , r e s p e c t i v e l y .
The w e ig h ts w ere re c o rd e d a t b i r t h , w eaning and 12 mo d u rin g th e f i r s t
y e ar,
th e n i n
th e f a l l
fo r
th e su b se q u e n t y e a r s .
The c o rre sp o n d in g
a g e s w ere re c o rd e d i n m onths.
These w e ig h t-a g e r e c o r d s f o r each ewe w ere a n a ly z e d t o e s tim a te
th e p a ra m e te rs o f B ro d y 's (1 9 2 7 a) n o n - li n e a r g row th model by u s in g
a
m icrocom puter program w r i t t e n s p e c i f i c a l l y f o r t h i s s tu d y (S . Kachman,
29
p e rs o n a l com m unication).
Those a n a ly s e s p ro v id e d e s ti m a te s o f A, k and
B f o r each ewe from w hich grow th c u rv e s f o r each b re e d and group w ere
d e riv e d .
The p a ra m e te r v a lu e s f o r A and k w ere added t o
v a ria b le s
(tra its ).
The B p a r a m e te r w h ic h
is
a
th e s e t o f
c o n s ta n t
of
i n t e g r a t i o n , h a v in g no b io lo g ic a l im p o rta n c e , was n o t in c lu d e d .
A new v a r ia b le c a l l e d Lanb P ro d u c tio n E f f ic ie n c y In d e x (E I) was
c a lc u la te d from th e c u m u la tiv e l i t t e r w e ig h t a t w ean in g and th e e w e's
m atu re w e ig h t.
T h is in d e x e x p r e s s e s a v e ra g e a n n u a l Ia n b p ro d u c tio n a s
a p ro p o r tio n (* ) o f m a tu re w e ig h t (A ).
E w e's a v erag e a n n u al l i f e t i m e lam b p ro d u c tio n , kg
EI =
— ------------------------------------------------------------ 100
E w e's m a tu re w e ig h t, kg
Growth s t a t i s t i c s
(AGR,
( t r a i t s ) pro p o sed by F itz h u g h and T a y lo r (1 9 7 1 )
HGR and AMR) were c a lc u la te d f o r each ewe.
o b ta in e d from B ro d y 's model was u sed to
s ta tis tic s
(tra its )
Jfe tu re w e ig h t (A)
d e riv e AMR.
These g ro w th
were c a lc u la te d f o r f i v e a g e - i n t e r v a l s :
( I ) B irth
t o w eaning (B-W), (2 ) Weaning t o y e a r l i n g age (W -12), ( 3 ) Y e a rlin g t o
18 months (1 2 - 1 8 ), (4 ) B irth to y e a r l i n g age (B -1 2 ), and (5 ) B ir th to
18 months (B -1 8 ).
These same i n t e r v a l s w ere s tu d ie d by S to b a r t (1 9 8 3 ).
T able 7 shows th e t a b u l a t i o n o f a l l t r a i t s s tu d ie d ,
th e i r d e s c rip tio n
and u n i t s o f m easurem ent.
S ta tis tic a l
.
Maximum, minimum, a v e ra g e ,
s ta n d a rd d e v ia t io n and c o e f f i c i e n t o f
v a r i a t i o n w ere c a lc u la te d f o r each t r a i t by b re e d -g ro u p s u b c la s s .
a d d itio n ,
th e
c u m u la tiv e a n n u al av erag e o f ATLB,
In
ATLW and ATWW, f o r
f i r s t , f i r s t two, f i r s t th r e e , f i r s t f o u r and f i r s t f i v e r e p r o d u c tiv e
30
TABLE 7 .
EWES* PRODUCTIVE AND GROWTH TRAITS ANALYZED IN THE STUDY
T r a it
symbol
D e s c r ip tio n o f th e t r a i t
ATLB
Average o f th e t o t a l number o f lambs b o rn to
th e ewe p e r y e a r
n
ATLW
Average o f th e t o t a l number o f lam bs weaned
by th e ewe p e r y e a r
n
ATWW
Average o f th e t o t a l w e ig h t o f la n b s weaned
by th e ewe p e r y e a r
kg
ATFP
Average o f th e g re a s e f l e e c e w e ig h t produced
by th e ewe p e r y e a r
kg
A
A sym ptotic w e ig h t (m a tu re ) o f th e ewe d e riv e d
from B ro d y 's grow th e q u a tio n (m odel)
kg
k
M atu rin g r a t e c o e f f i c i e n t , o r r a t e o f d e c lin e
I n grow th a s th e a s y m p to tic w e ig h t (A) i s
ap p ro ach ed , d e riv e d from B ro d y 's model
%/mo
EI
Lamb P ro d u c tio n E f f ic ie n c y In d e x , th e r a t i o o f
ATWW d iv id e d by (A) e x p re s s e d a s k g /k g x 100
AGRa
A b so lu te Growth R ate, body w eig h t g a in s f o r
s p e c i f i e d i n t e r v a l s from b i r t h t o 18 mo
RGRa
R e la tiv e Growth R ate, body w eig h t g a in s f o r
s p e c i f i e d i n t e r v a l s from b i r t h t o 18 mo r e l a t i v e
t o f i n a l w e ig h t f o r th e s p e c i f i e d i n t e r v a l
%/mo
AMRa
A b so lu te M aturing R a te , body w eig h t g a in s f o r
a s p e c i f i e d i n t e r v a l from b i r t h t o 18 mo r e l a t i v e
to m a tu re w e ig h t: AGR d iv id e d by (A)
%/mo
a
<
( E 5£
H
^
U n its
%
kg/mo
31
r e c o r d s a s th e ewes p ro g re s s e d th ro u g h t h e i r l i f e t i m e p ro d u c tio n .
In
th e case o f wool p ro d u c tio n (ATFP), th e c o rre s p o n d in g number o f re c o rd s
w ere f i r s t tw o, f i r s t t h r e e , f i r s t f o u r , f i r s t f i v e and f i r s t s i x
re c o rd s.
The DNIVAHIATE p ro c e d u re from SAS (1986) was a p p lie d i n th e s e
a n a ly s e s .
B reeds and g ro u p s w ere compared on th e b a s is o f a l l t r a i t s l i s t e d
i n T able 7 and f o r t h e i r f i r s t , f i r s t two and f i r s t th r e e c u m u la tiv e
r e p r o d u c tiv e r e c o r d s f o r ATLB, ATLW, ATWV and ATFP.
S im ila r ly , a
f i n a l com parison was made o f th e g ro u p s w ith f o u r and f i v e c u m u la tiv e
r e p r o d u c tiv e r e c o r d s .
L e a st s q u a re s m ethod (H arvey, 1977) was u sed t o
perform th e s e a n a ly s e s u s in g a m odel (m odel A) w hich in c lu d e d y e a r o f
b i r t h o f th e ewe, b re e d o f th e ewe (B ), group o f th e ewe (G ), B x G
i n t e r a c t i o n , age o f dam i n y e a r s , ty p e o f b i r t h - r e a r i n g o f th e ewe, day
o f b i r t h ( c o v a r i a t e ) and r e s i d u a l a s s o u rc e s o f v a r i a t i o n .
A lso , th e
grow th s t a t i s t i c s (A, k, AGR, RGB, and AMR) d e riv e d from th e com plete
in fo r m a tio n o f re c o rd e d w e ig h ts w ere a n a ly z e d u s in g model A.
Ewes b o rn
a s t r i p l e t s a s w e ll a s th e o n ly one o b s e r v a tio n i n 1976 f o r C olum bias
( t o t a l o f 11) were n o t in c lu d e d .
A lso , th e a n a ly s e s in v o lv in g th e
grow th p a ra m e te rs and grow th s t a t i s t i c s , a s w e ll a s E l, d id n o t
in c lu d e d a Columbia ewe w hich had a k v a lu e more th a n 3 s ta n d a r d
d e v ia t io n s from th e mean.
T hat ewe was n o t in v o lv e d i n a l l th e
su b se q u e n t g e n e tic and p h e n o ty p ic a n a ly s e s .
E s tim a tio n o f g e n e t i c an d p h e n o ty p ic v a r i a n c e an d c o v a M annm
-
com ponents.
L e a s t- s q u a r e s a n a ly s e s (H arvey, 1 9 7 7 ), u s in g a m ixed
m odel, were perform ed f o r a l l t r a i t s l i s t e d i n T a b le 7 .
A n aly ses w ere
made f o r each b re e d s e p a r a t e l y (m odel B) and f o r th e combined d a ta s e t
32
(model C )e
The e le m e n ts o f m odel B ( s o u rc e s o f v a r i a t i o n ) w ere b i r t h
y e a r o f th e ewe, s i r e o f th e ewe w ith in b i r t h y e a r , ty p e o f
b i r t h - r e a r i n g o f th e ewe, day o f b i r t h , age o f dan i n y e a r s and an
e r r o r term ( r e s i d u a l ) .
Model C in c lu d e d a s s o u rc e s o f v a r i a t i o n b i r t h
y e a r -b re e d o f th e ewe, s i r e o f th e ewe w ith in b i r t h y e a r - b r e e d , ty p e o f
b i r t h - r e a r i n g o f th e ewe, day o f b i r t h , age o f dam and a n e r r o r term
(re s id u a l).
For b o th m odels, s i r e s and th e r e s i d u a l e le m e n ts w ere
c o n s id e re d random v a r i a b l e s and a l l o th e r e le m e n ts f i x e d .
V arian ce and
c o v a ria n c e com ponents among and w ith in s i r e s w ere d e riv e d from th e s e
a n a ly s e s .
The d i s t r i b u t i o n o f s i r e s and t h e i r o f f s p r i n g p e r y e a r i s
shown i n T ab le 8.
H e r l t a b i l l t l e s o f th e t r a i t s s tu d ie d , and th e g e n e ti c , p h e n o ty p ic
and e n v iro n m e n ta l c o r r e l a t i o n s among them were p ro v id e d d i r e c t l y by th e
H arvey (1977) a n a ly s e s o f m odels B and C.
The fo rm u la s f o r c a l c u l a t i n g
th e s e e s tim a te s a s w e ll a s th e s ta n d a rd e r r o r s f o r h e r ! t a b l l i t i e s and
g e n e tic c o r r e l a t i o n s a r e p re s e n te d i n a p p en d ix A o f th e H arvey (1977)
U s e r 's g u id e f o r LSML76.
TABLE 8 .
YEAR
DISTRIBUTION OF SIRES (S ) AND OFFSPRING (o ) PER BREED AND
R a m b o u ille t
S
O
______________________ Y ear
60 61 62 63 64 65 66 67 6ft 69 70 7? 73 7ft 76
10 3 7 6 6 3 7 7 6 8 7 7 12 7 I
28 23 17 15 21 21 27 31 21 33 24 13 17 10 I
Targhee
S
O
7 4 6 6 5 7 7 8 7 7 8 8
27 14 21 27 33 44 31 25 21 22 25 25
Columbia
S
O
T o ta l
S
O
B reed
6 7 5 6 8 6 8
19 14 20 19 18 12 17
6 4 3
13 17 13
5 5
9 10
3
4
3
4
4
9
17 13 20 17 17 18 20 23 13 21 19 18 20 16
55 56 52 62 73 83 70 73 42 68 66 51 30 29
T o ta l
97
302
93
338
66
175
4
5
256
815
33
S e le c tio n
In d e x e s.
S e le c tio n
I n d e x e s w ere d e riv e d from
th e
g e n e ti c and p h e n o ty p ic p a ra m e te rs e s tim a te d from th e p o o led a n a ly s e s I n
t h i s stu d y u s in g th e m ethods p re s e n te d by F a lc o n e r (1 9 8 1 ).
fo r
w h ic h
s e le c tio n
in d e x e s w ere computed w ere ATWV,
ATWWtATFP, ATFP+EI a n d
ATWW+ATFP+EI,
u s in g
v a rio u s
The t r a i t s
E l,
ATWW+E l,
g ro w th
tra it
c o m b in atio n s a s p r e d i c t i v e e le m e n ts i n th e in d e x e s .
Summary o f th e s te p s fo llo w e d f o r h an d l i n g and a n a l y s i s of* d a t a .
F ig u re 2 shows th e
h a n d lin g th e d a ta ,
a n a ly s e s em ployed.
flow
c h art
of
a ll
th e
s te p s
fo llo w e d
in
c a l c u l a t i o n o f th e new v a r i a b l e s and th e ty p e s o f
34
DATA
f
-------------------- H
DERIVATION OF NEW
VARIABLES ATLB,
ATLWt AlWW AND ATFP
I
-
DERIVATION OF A, Bt
AND K USING BRODY'S
MODEL
DERIVATION OF
FITZHUGH & TAYLOR
GROWTH STATISTICS
AGR AND RGR
DERIVATION OF EI
AND AMR TRAITS
USE OF MODEL A FOR
ANALYSIS OF GROUP
AND BREED EFFECTS
DERIVATION OF
BRODY'S GROWTH
CURVES BY BREED
USE OF MODEL B FOR
ESTIMATION OF GENETIC
AND PHENOTYPIC
PARAMETERS BY BREED
USE OF MODEL C FOR
ESTIMATION OF GENETIC
AND PHENOTYPIC
PARAMETERS FROM
POOLED DATA
*
CONSTRUCTION OF
SELECTION INDEXES
raE m p s F0LL" ED 11
HANDLING IHB
35
RESULTS AND DISCUSSION
D e sc rip tiv e
s ta tis tic s fo r a ll
tra its
by b re e d and
p re s e n te d i n Appendix T ables 22 through 4 4 .
num ber o f r e c o r d s ,
v a ria tio n
av erag e,
s ta n d a r d
and ra n g e v a lu e s f o r
s u b c la s s .
T hese
d a ta
a re
u n d e r s ta n d in g o f
th e
m agnitude
a b s o lu te
of
th e
g e n eral
c o e ffic ie n t
each t r a i t w ith in each
le v e l
v a ria tio n
u sefu l
fo r
o f m e rit i n
an d
a re
These ta b le s show th e
d e v ia tio n ,
c o n s id e r e d
g ro u p
of
breed-group
o b ta in in g
each
tra it,
an
th e
th e r e l a t i v e
v a ria tio n
( c o e f f ic ie n ts o f v a r ia tio n ) en co u n tered i n each d a ta s e t .
S p e c ific
com parisons among b re e d s, groups o r o th e r f a c t o r s a re o b ta in e d from th e
more complex a n a ly s e s o f models A, B, and C.
High c o e f f ic ie n ts o f v a r i a b i l i t y were o b serv ed i n th e re p ro d u c tiv e
tra its .
T his was due to th e f a c t t h a t z e ro v a lu e s w ere in c lu d e d
as
re c o rd s f o r th e ewes th a t f a i l e d to g iv e b i r t h to o r wean o f f s p r in g
any y e a r.
Breed and group d i f f e r e nc e s f o r
The fo u r p ro d u c tiv e
analyzed u s in g model A.
( li f e t im e
p ro d u c tio n ),
th A
tra its ,
DrorinnHv*
ATLBt
ATLV,
ATVW and ATFP, w e re
T his was done f o r th e average o f a l l re c o rd s
and th e average o f
the cu m ulative p ro d u c tio n
through th e f i r s t , second, t h i r d and f o u r th re c o rd s .
These r e s u l t s a r e
p re s e n te d from Table 9 t o 13, r e s p e c tiv e ly .
U f r t U s B-BItinMmi.
Table
9 p re s e n ts
th e
le a s t- s q u a r e s means f o r th e fo u r p ro d u c tiv e t r a i t s .
breed and groups w ere h ig h ly s i g n i f i c a n t f o r
e x c e p tio n o f breeds f o r ITLB (P < .0 5 ).
a ll
mean
sq u a re s and
The e f f e c t s o f
th e
tra its
w ith
The I n t e r a c t i o n between breed
and group was g e n e r a lly u n im portan t e x c e p t f o r ITLW and ATWW (P < .0 5 ).
36
T arghees w ere s u p e r io r
to
Colum blas f o r
ATLB and ATLW.
For ATLB1
R a m b o u ille ts d id n o t d i f f e r from T arghees and Colum bia s , and f o r ATLW,
T arghees and R a m b o u ille ts had s im ila r perform ances#
The l e a s t s q u a re s
means f o r ATLB w ere 1 .3 8 , 1.43 and 1 .3 3 , and f o r ATLW 1 .1 4 , 1 .22 and
I .0 5 , f o r R a m b o u ille ts, T arghees and C olum bias, r e s p e c t i v e l y .
T argbees
were s u p e r io r i n ATWW to R a m b o u ille ts and C olum bias ( 4 8 .6 9 , 4 2 .3 9 and
41.19 kg o f lam b produced p e r y e a r , r e s p e c t i v e l y ) .
C olum bias had th e
h ig h e s t ATFP, w ith T arghees b e in g s u p e r io r to R a m b o u ille ts ( 4 .9 1 , 4 .7 3
and 4 .4 5 kg,
r e s p e c t i v e l y ) « The f i v e —re c o rd
g ro u p w as
s u p e rio r
to
th r e e - and f o u r - r e c o r d g roups f o r ATLB, ATLW and ATWW, w h ile th e t h r e e and f o u r - r e c o r d g ro u p s showed no d if f e r e n c e .
T here was no d if f e r e n c e
betw een th e f o u r - and f i v e - r e c o r d g ro u p s f o r ATFP, b u t th e t h r e e - r e c o r d
group was i n f e r i o r f o r t h i s t r a i t .
Fi rst record*
means f o r
T able 10 shows th e mean s q u a re s and l e a s t s q u a re s
th e a v erag e p ro d u c tio n f o r
d ^ ’^ e re n t f OI* a l l th e t r a i t s ,
th e f i r s t r e c o r d .
B reeds w ere
b u t th e d i f f e r e n c e s w ere n o t so d i s t i n c t
a s i n th e p re v io u s a n a l y s i s o f com plete p ro d u c tio n f o r ATLB and ATLW.
For ATLB and ATLW, T arghees were s u p e r io r t o R a m b o u ille ts, b u t s i m i l a r
t o Colum bias and C olum bias w ere n o t d i f f e r e n t from R a m b o u ille ts.
For
ATWW, T arghees w ere s u p e r io r t o th e o th e r b re e d s w hich w ere s im ila r t o
each o th e r .
C olum bias and T arg h ees showed s i m i l a r perfo rm an ce f o r ATFP
and w ere s u p e r io r t o R a m b o u ille ts.
There was no group e f f e c t f o r ATLB,
ATLW and ATWW.,
The f i v e -
re c o rd group was s u p e r io r t o th e th r e e - r e c o r d g ro u p f o r ATFP.
h a v in g f i v e
perform ance
re c o rd s
w e re i n t e r m e d i a t e .
( f i r s t re c o rd )
T h is r e s u l t
The ewes
from
e a r ly
when compared to th e r e s u l t s from l i f e t i m e
37
TABLE 9 .
MEAN SQUARES AND LEAST SQUARES MEANS FOR THE LIFETIME
AVERAGES OF PRODUCTIVE TRAITS OF EWES (HARVEY LEAST SQUARES ANALYSES,
MODEL A)
Source o f
v a ria tio n
Mean Snnan as
df
Year o f
b irth
14
Breed ( B)
2
Group (G)
2
B xG
4
Age o f dam 5
Tjrpe o f b i r t h re a rin g
2
B ir th d a te I
R esid u a l 773
ATLB
ATLW
.4607**
.3588*
3.0920**
.1329
.0267
.6999**
1.1132**
5.5445**
.2883*
.0849
.5692**
.0107 .1006
.1762
.2844
.0958
ATWW
ATFP
1 ,7 8 6 .1 * *
2 ,9 1 9 .7 * *
8 ,4 0 8 .2 * *
414.0*
9 7 ,8 7 3 .0
1.5411**
8.4934**
3.3332**
.2391
.6116
335.4
405.3
124.0
2.4522**
.3328
.2822
L e a st S a u a re s Means
B reed :
R am b o u ille t
Targhee
Columbia
N
n
n
kg
295
335
174
1 .38±_.02ab
1 .4 3 ± .0 2 a
1 .33± .03b
1 .1 4 ± ,0 3 a
1 .22±>02a
1.05 ± ,0 3 b
4 2 .3 9 ± ,9 5 b
48.69± > 91a
4 1 .1 9 * J .2 b
4 .4 5 t.0 4 c
4 .7 3 ± .0 4 b
4 .9 1 t,0 6 a
198
129
477
1 .30± ,03b
1 .33± .03b
1 .5 0 ± .0 2 a
1. 02± , 03b
39.50±..97b
4 2 .4 4 ± J.2 6 b
5 0 .3 4 tj.7 8 a
4.57±>05b
4 .7 2 t,0 6 a
4 .8 0 t_ .0 4 a
kg
Group:
3 re c o rd s
4 re c o rd s
5 re c o rd s
1 .10± ,03b
1 .30±>02a
a , b, c , a r e LSD com parisons (P < .0 1 )
*P< .05, **P<.01
38
TABLE 10.
MEAN SQUARES AND LEAST SQUARES
MEANS FOR THE CUMULATIVE
AVERAGE OF PRODUCTIVE TRAITS OF EWES THROUGH
THEIR FIRST RECORD (HARVEY
LEAST SQUARES ANALYSES, MODEL A)
Mean S q u ares
Source o f
V a r ia tio n
df
Year o f b i r t h 14
B reed (B)
2
Group (G)
2
B xG
4
Age o f dam
5
Type o f b irth r e a r in g
2
B ir th d a te
I
R esid u a l
773
:
B re e d :
R am b o u illet
Targhee
Columbia
ATLB
ATLW
ATWW
.8134**
1.0492**
.1811
.0512
.1034
.7026**
1.2170**
.1638
.3851
.1107
1 ,6 2 7 .3 * *
3 ,5 3 0 .4 * *
3 2 0 .7
575.9
111.9
5.5370**
4.2920**
1.1413**
.5860
.7529**
472.5
27.6
264.7
5.4354**
4.1140**
.2580
.4219
.0866
.1853
■
• .2489
.0145
.1943
ATFPa
L east S a u a re s Means
N
n
295
335
174
1.14±..04b
I .2 8 ± ,0 3 a
1 .19±»05ab
198
129
477
1 .2 2 ± ,0 4 a
1 .1 7 ± .0 5 a
1.2 3 ± .0 3 a
n
kg
kg
.88±j.04b
1 .0 3 ± ,0 4 a
• 91jL.05ab
32.95±.1.4b
4 0 .9 3 ± 1 .3 a
3 4 .8 9 t1 .8 b
4 .2 2 ± ,0 4 b
4 .4 3 ± ,0 4 a
4 .5 3 ± ,0 5 a
.9 3 ± ,0 4 a
•9 2 ± ,0 5 a
•9 7 ± ,0 3 a
3 5 .8 7 ± 1 .4 a
35.30± .1.8a
3 7 .5 9 ± i.2 a
4 .3 4 ± .0 4 b
4 .3 8 t.0 6 a b
4 .4 6 ± ,0 3 a
Group:
3 re c o rd s
4 re c o rd s
5 re c o rd s
aSecond reco rd
a ,b , are LSD comparisons (P<,01)
** P<.01
39
r e c o r d s , s u g g e s ts t h a t g re a s e f le e c e p ro d u c tio n I s more I n d i c a t i v e o f
life tim e
p ro d u c tio n
th a n
th e
o th e r
su p p o rte d
by th e h ig h r e p e a t a b i l i t y
tra its .
T h is
r e p o r te d f o r
s u g g e s tio n i s
th a t
tra it
in
the
l i t e r a t u r e (T u rn er and Young, 1 9 6 9 ).
T able 11 shows th e mean s q u a re s and l e a s t s q u a re s
SfiQQBd .r e c o r d .
means f o r
th e
av erag e
p ro d u c tio n f o r
th e f i r s t
two r e c o r d s .
Breed
d if f e r e n c e s a t t h i s l e v e l w ere more m arked th a n f o r th e a n a l y s i s a t th e
second r e c o r d .
b u t w e re
For ATLB, T arghees perfo rm ed b e t t e r th a n R a m b o u ille ts ,
not
d iffe re n t
R a m b o u ille ts.
C o lu m b ia s w h ic h
w e re
s im ila r
to
T arghees w ere s u p e r io r to R a m b o u ille ts bu t s i m i l a r to
Colum bias f o r ATLW.
change i n
fro m
R a m b o u ille ts and Columbian were no d i f f e r e n t .
ra n k o f R am b o u ille t from ATLB to ATLW c o u ld
The
be ta k e n a s
i n d i c a t i o n o f a h ig h e r lam b l o s s from b i r t h t o w eaning i n R a m b o u ille ts
which
h av e a h ig h e r fre q u e n c y
(1984) r e p o r te d
of tr ip le ts
th e lo w e st s u r v i v a b i l i t y
R am b o u illet dams.
a t lam b in g .
fo r
p ro g e n y
S u b an d riy o
o f 3 -y r-o ld
He a ls o r e p o r te d low s u r v i v a b i l i t y to be a s s o c ia te d
w ith sm all b i r t h w e ig h ts .
T arg h ees w ere s u p e r io r t o R a m b o u ille ts and
Colum bias f o r ATWW, but no d if f e r e n c e was o b se rv e d betw een th e s e l a s t
two b re e d s .
F or ATFP,
C o lu m b ian w e re
s u p e rio r
to
T a rg h e e s
an d
R a m b o u ille ts, and T arg h ees had g r e a t e r ATFP th a n R a m b o u ille ts.
The t h r e e
g ro u p s w e re
not
d i f f e r e n t when compared f o r
how ever, th e y were d i f f e r e n t f o r each o f th e o th e r t r a i t s .
f i v e r e c o r d s had th e h ig h e s t a v e ra g e p e rfo rm a n c e s.
t o th o s e h a v in g t h r e e ,
but
ATLB,
Ewes w ith
Ciey were s u p e r io r
n o t d i f f e r e n t from th e f o u r - r e c o r d ewes.
The tr e n d tow ard h ig h e r perform an ce f o r th e f i v e - r e c o r d ewes began t o
emerge more c l e a r l y a f t e r two r e c o r d s th a n a f t e r one r e c o r d .
40
TABLE 11.
MEAN SQUARES AND LEAST SQUARES MEANS FOR THE CUMULATIVE
AVERAGE OF PRODUCTIVE TRAITS OF EWES THROUGH THEIR SECOND RECORD
(HARVEY LEAST SQUARES ANALYSES, MODEL A)
Source o f
v a ria tio n
Mean S a u a rm a
df
Year b i r t h 14
B reed (B)
2
Group (G)
2
B xG
4
Age o f dam
5
Type o f b i r t h r e a r in g
2
B ir th d a te
I
R esid u a l
773
ATLB
ATLW
ATWW
.5237**
.4616**
.4153*
.2547
.0630
.6930**
.7510**
.9450**
.1488
.0313
1 ,7 8 8 .1 * *
2 ,5 3 2 .9 * *
1 ,2 4 3 .9 * *
173.7
56.7
5.4310**
6 .0 880**
1.2550«*
.3660
.5151
.5337*
.0059
.1298
.2440
.0514
.1330
381.4
131.6
174.1
3.7370**
1.4420*
.2610
ATFPa
L e a st S q u a re s Means
B re e d :
R am b o u illet
Targhee
C olim bia
N
n
295
335
174
1 .2 3 ± .0 3 b
1 .3 2 ± .0 3 a
1 .24±,.03ab
198
129
477
I »26±.03a - 1 .00± .03b
1 .2 3 ± .0 4 a
I .0 2 j> 0 4 ab
1 .3 1 ± .0 2 a
1 . 10±_.02a
n
.99±,.03b
1 .1 0 ± ,0 3 a
1 .02±_.04ab
kg
kg
37.4l± .1.10b
4 4 .3 1 jJ .1 5 a
3 9 .9 4 jJ .4 b
4 .3 2 ± ,0 4 c
4.56±>04b
4.70±,.05a
G roup:
3 re c o rd s
4 re c o rd s
5 re c o rd s
m0
a t h i r d re c o r d
* b, a r e LSD co m parisons (P < .01)
(P < .0 5 ), ** (P < .01)
3 8 .6 5 t1 .1 b
3 9 .6 5 t1 .5 a b
4 3 . 0 3 t .9 a
4 .4 6 ± ,0 4 b
4 .5 3 t* 0 6 a b
4 .6 0 t,0 3 a
41
Th i r d
re c o rd *
s q u a re s means f o r
T a b le
12
p r e s e n ts
th e mean s q u a r e s and l e a s t
c u m u la tiv e perform ance o f th e f i r s t th r e e r e c o r d s .
B reed e f f e c t was v e ry im p o rta n t f o r a l l
th e
tra its
e x c e p t f o r ATLB.
The R a m b o u ille ts and C olum bias had s i m i l a r p e rfo rm an c es f o r ATLW and
ATWW.
T arghees showed th e h ig h e s t p erfo rm an ce f o r th e s e t r a i t s .
b reed d i f f e r e d f o r ATFP.
Each
Colum bias had th e h i g i e s t ATFP fo llo w e d by
T arghees , w ith th e R a m b o u ille ts h a v in g th e lo w e st p erfo rm an ce.
W ith r e s p e c t to th e group e f f e c t , th e d if f e r e n c e s f o r th e a v erag e
o f f o u r r e c o r d s were c l e a r e r th a n f o r th r e e .
The f i v e - r e c o r d ewe group
showed th e h ig h e s t p e rfo rm an c es f o r th e fo u r t r a i t s .
re c o rd g roups were n o t d i f f e r e n t .
to
th e
th re e -re c o rd
f o u r - r e c o r d group.
group f o r
T h ree - and f o u r -
The f i v e —re c o rd group was s u p e r i o r
ATFP1 b u t was n o t d i f f e r e n t from
th e
B reed group i n t e r a c t i o n s (P < .0 5 ) w ere o b s e rv e d f o r
ATLW and ATWW.
fourth—record*
The a n a l y s i s o f av erag e p e rfo rm an c es o f ewes a t
t h e i r f o u r th r e c o rd i s d i f f e r e n t th a n th e a n a ly s e s u s in g th e a v e ra g e s
o f o n e,
two and th r e e r e c o r d s .
T h is
com parison in c lu d e d
b re e d s but o nly th e f o u r - and f i v e - r e c o rd g ro u p s .
Colum bias and R a m b o u ille ts .
R a m b o u ille ts and C olum bias,
th r e e
T ab le 13 d i s p l a y s
th e mean s q u a re s and l e a s t s q u a re s means from th e s e a n a ly s e s .
d i f f e r e n c e s w ere found f o r ATLB.
th e
No b re e d
For ATLW, T arg h ees w ere s u p e r io r to
For ATWW, T arg h ees perform ed b e t t e r th a n
which w ere n o t d i f f e r e n t .
The C o lm b ia s
a g a in had th e h ig h e s t p erfo rm an ce f o r ATFP, w ith th e T arg h ees second
and R a m b o u ille ts t h i r d i n ra n k .
Ewes h a v in g f i v e re c o r d s w ere s u p e r io r
to ewes h a v in g f o u r f o r a l l th e t r a i t s e x c e p t ATFP w here no d i f f e r e n c e s
42
TABLE 12.
MEAN SQUARES AND LEAST SQUARES MEANS FOR THE CUMULATIVE
AVERAGE OF PRODUCTIVE TRAITS OF EVES THROUGH THEIR THIRD RECORD (HARVEY
LEAST SQUARES ANALYSES, MODEL A)
Source o f
v a ria tio n
Mean S a u a r e s
df
Year o f b i r t h 14
B reed (B)
2
Group (G)
2
B xG
4
Age o f dam
5
Type o f b i r t h r e a r in g
2
Date o f b i r t h
I
R e sid u a l
773
ATLB
ATLV
ATW
ATFP
.5365**
.2837
.6948**
.1069
.0648
.7436**
.9830**
2.2330**
.3314
.0707
1 , 878 . 8**
2 ,9 1 7 .9 * *
3 ,1 7 7 .8 * *
431.2*
7 7 .0
3.4430**
6.8130**
1.1910**
.2824
.5813
.6015**
.0055
.1130
.2930
.2815
450.3*
431.8
140.7
2.9930**
.7328
.1100
.2726
L e a st S o u a re H Maana
B re e d :
R am b o u illet
Targhee
Columbia
N
n
n
kg
kg
295
335
174
1 .3 2 ± .0 3 a
1 .3 8 ± ,0 2 a
1 .2 9 ± .0 4 a
1 .0 9 ± ,0 3 b
I . 1 9 + .03a
I .0 3 ± ,0 3 b
40.85±_1.0b
47.53dL.97a
4 0 .5 8 t1 .3 b
4 .4 1 + .04c
4 .6 7 t.0 4 b
4 .8 5 t» 0 6 a
198
1 .3 1 i.0 3 b
1 .28±,.04b
1 .3 9 ± .0 2 a
1. 02±,.03b
3 9 .7 9 ± 1 .0 b
4 2 .5 2 t1 .3 b
4 6 .6 3 t.8 3 a
4 .5 8 t.0 4 b
4 .6 6 t.0 6 a b
4 .7 1 ± .0 3 a
Group:
3 re c o rd s
4 re c o rd s
5 re c o rd s
129
477
1 .09± ,04b
I . 20± , 02 a
aTourth record
a , b, c, are LSD comparisons (P<.01)
* (P<.05), ** (P<.01)
43
TABLE 13.
MEAN SQUARES AND LEAST SQUARES MEANS FOR THE CUMULATIVE
AVERAGE OF PRODUCTIVE TRAITS OF EWES THROUGH THEIR FOURTH RECORD
(HARVEY LEAST SQUARES ANALYSES, MODEL A)
Source o f
v a ria tio n
df
Year o f b i r t h 14
B reed (B)
2
Group (G)
I
B xG
2
Age o f dam
5
Type o f b i r t h re a rin g
2
Date o f b i r t h I
R e s id u a l
578
ATLB
ATLW
ATWW
ATFPa
.3835**
.1586
1.1512**
.2047
.0674
.4710**
.5142**
1.9944**
.3967*
.0638
1 ,3 0 4 .9 * *
1 , 716 . 6 **
3 ,1 9 0 .3 * *
544.3**
5 2 .9
1 . 6060**
5.3390**
.1444
.2668
1 . 0280**
142.8
227.4
114.9
2.6980**
.1443
.2720
.4589**
.0429
.0950
.1052
.1929
.0933
L e a s t S o u a r e * M eans
B re e d :
R am b o u illet
T arghee
Columbia
N
n
n
223
262
121
1.39±>03a
1 .42±,.03a
1 .3 4 ± .0 4 a
1 .l6 ± ,0 3 b
1 .2 4 ± ,0 3 a
1.10±,.04b
43.58± .1.l3b
4 9 .7 4 ± 1 .0 8 a
4 3 .0 6 ± 1 .5 b
4 .5 3 t.0 5 c
4 .8 lt.0 5 b
4 .9 8 t.0 7 a
129
477
1 .3 3± .03b
1.45±>02a
1 .09± ,04b
I .2 5 ± ,0 2 a
4 3 .2 7 ± 1 .3 b
4 8 .6 5 t.8 5 a
4 .7 5 t.0 6 a
4 .7 9 t.0 4 a
kg
t
kg
Groupz
4 re c o rd s
5 re c o rd s
aY ifth record
a» b, c, are LSD comparisons (P<.01)
* (P<.05), **(P<.01)
44
were d e te c te d .
I n t e r a c t i o n s between b re e d and gro u p w ere o b se rv e d f o r
ATLW (P < .05) and ATWW (P < .0 1 ).
ErPdY equation p a ra m e te rs A. Br and k and th e E f f i c l e n n v I n d e x fRTl
T able 14 shows th e mean s q u a re s and l e a s t s q u a re s means f o r th e
grow th p a ra m e te rs A, B and k and th e e f f i c i e n c y in d e x .
had th e lo w e s t m a tu r e w e ig h t
( 7 0 .0 4
kg)
w h ile
th e
R a m b o u ille ts
T a rg h e e s and
Colum bias w ere s i m i l a r (7 2 .4 2 and 73.11 k g , r e s p e c t i v e l y ) .
The g ro u p s
o f ewes d id n o t show d i f f e r e n c e s i n m ature s i z e s and no i n t e r a c t i o n was
d e te c te d betw een b re e d and group f o r t h i s grow th p a ra m e te r.
The th re e b re e d s had d i f f e r e n t m a tu rin g r a t e s .
C o ltm b ias had th e
s lo w e s t (.1 1 0 5 ) w h ile T arg h ees h ad th e f a s t e s t (.1 2 0 0 ) and R a m b o u ille ts
had an in te r m e d ia te v a lu e ( .1 1 5 9 ) .
m atu red f a s t e r
th a n th o s e i n
EWes i n th e group o f th r e e r e c o r d s
th a n th e group o f f i v e r e c o r d s ,
th e group o f f o u r .
Ewes i n
b u t w ere no d i f f e r e n t
th e f o u r - and f i v e - r e c o r d
g roups had s i m i l a r m a tu rin g r a t e s .
The s i m i l a r i t y i n m atu re w e ig h ts among th e g ro u p s, b u t d if f e r e n c e
i n m a tu rin g r a t e s s u g g e s ts t h a t w ith in b re e d , w h a tev e r th e p ro d u c tiv e
lo n g e v ity o f th e ew es, m atu re w e ig h ts would te n d t o
be s i m i l a r ,
bu t
d if f e r e n c e s w ould e x i s t among them w ith r e s p e c t t o m a tu rin g r a t e s .
T a rg h e e s
had
th e
h ig h e s t
e ffic ie n c y
R a m b o u ille ts w hich w ere s u p e r io r to C olum bias.
in d e x ,
f o llo w e d
The l e a s t s q u a re s means
f o r th e s e th r e e b re e d s w ere .6 7 8 1 , .6099 and .5 6 7 1 , r e s p e c t i v e l y .
w ith
fiv e
re c o rd s
in te rm e d ia te
w e re
and th e
th r e e - r e c o r d g ro u p .
s u p e rio r,
p o o re s t
th o s e
w ith
fo u r
re c o rd s
perfo rm an ce was shown by ewes i n
L e a s t s q u a re s means w ere
by
.6 9 8 3 ,
Ewes
w e re
th e
.5941 and .5629
45
TABLE 14.
MEAN SQUARES AND LEAST SQUARES MEANS FOR BRODY'S GROWTH
CURVE PARAMETERS (A, B, k) AND THE EWE EFFICIENCY INDEX (HARVEY LEAST
SQUARES ANALYSES, MODEL A)
Source o f
v a ria tio n
M ean S n m i r e s
df
Year o f b i r t h 14
B reed (B)
2
Group (G)
2
B xG
4
Age o f dam
5
Type o f b i r t h r e a r in g
2
B ir th d a te
I
R esid u al
772
A
424.34**
450.17**
53.97
3 2 .7 5
60.96
B
3 35.01**
385.40**
15.35
23.86
3 0.64
414.11**
27.44
3 3 .5 8
5 7 .3 9
5 .8 4
3 0 .9 9
k
EI
.011054**
.003381**
.003037**
.000301 .000437
.1955**
.5051**
1.3374*«
.1036*
.0314
.010965**
.006441**
.000605
.1828*«
.0520
.0276
L e a s t S q u a r e s Mmana
B reed:
R am b o u illet
Targhee
Columbia
N
kg
kg
KZmoa
% o f Aa
295
335
173
70.04±,.53b
7 2 .4 2 + .5 3 *
7 3 .1 1 + .6 7 *
6 2 .6 9 ± ,4 8 b
6 4 .9 5 ± ,4 5 a
6 5 .4 7 ± ,6 0 a
11.59±,.2b
12.00± ..2a
1 1 .0 5 ± ,3 c
6 0 .9 9 i1 .4 b
6 7 .8 1 i1 .3 a
56.71±J .8 °
198
129
476
7 1 .2 3 + .5 4 *
72.29±,.70a
7 2 .0 6 + .4 3 *
64.03±_.48a
64.66±..63*
64.41± ,.39a
1 1 .9 7 ± ,2 a
1 1 . 3 9 ^ 3 ab
1 1 .2 8 i,1 b
5 6 .2 9 i1 .4 °
5 9 .4 1 ± 1 .9 b
6 9 .8 3 ± J .1 a
Group:
3 re c o rd s
4 re c o rd s
5 re c o rd s
fit
Values a re expressed a s percentage
a, b, c, a re LSD comparisons (P<.01)
*(P<.05), *•(?<.01)
46
fo r the groupa of f iv e , fo u r and th re e re c o rd s, re s p e c tiv e ly .
The
breed by group in te r a c tio n was s ig n if ic a n t (P< .05).
B ro d v 1S grow th
e q u a tio n s and age a t m a t u r i t y .
Brody1S grow th
c u rv e s and e q u a tio n s f o r each b reed and gro u p a r e shown i n
From th e s e e q u a tio n s ,
F ig u re 3 .
th e tim e a t w hich each b re e d o r group a t t a i n e d
m atu re w e ig h t was d e te rm in e d .
A ppendix T ab le 45 shows th e e s tim a te d
w e ig h ts g e n e ra te d from 6 to 84 mo f o r each grow th e q u a tio n .
For th e p u rp o se o f d e f in in g th e p o in t a t w hich m a tu re w e ig h t was
a c h ie v e d ,
an a r b i t r a r y
c r i t e r i o n was u se d .
The a s y m p to tic n a tu r e o f
B ro d y 's e q u a tio n does n o t g iv e a d e f i n i t e p o in t o n th e age s c a l e when
maximum w e ig h t i s re a c h e d .
Age a t m a tu re s i z e w as t h e r e f o r e d e fin e d a s
th e age a t w hich 99 % o f A was a t t a i n e d .
ages
a t m a tu rity
w e re
a p p ro x im a te ly
U sin g t h i s
c rite rio n ,
th e
39 mo, 38 mo and 41 mo f o r
R a m b o u llle ts , T arg h ees and Colunb i a s , r e s p e c t i v e l y .
ggfflp a ri sons _.Pf AGR. RGB an d AMR f o r v a r y in g tim e I n t e r v a l s
betweer)
b i r t h and 18 m onths o f a r a .
Afiii—SPfflP a rl 8009»
T ab le 15 p r e s e n ts th e mean s q u a r e s and l e a s t
s q u a re s means f o r th e e f f e c t s o f b re e d , group and t h e i r i n t e r a c t i o n f o r
AGR i n
th e f i v e grow th i n t e r v a l s
s tu d ie d .
b re e d s w ere o b se rv e d f o r a l l i n t e r v a l s .
D if f e r e n c e s i n
AGR among
For AGRI and AGR3, T arg h ees
and C olum bias w ere s i m i l a r b u t s u p e r io r t o R a m b o u ille ts .
T arg b ees w ere
s u p e r io r to R a m b o u ille ts and C olum bias f o r AGR4, and R a m b o u ille ts w ere
s u p e r io r t o T arg h ees and C olum bias f o r AGR2.
fa s te r
g ro w th
ra te s
th a n
e ith e r
For AGR5 , T a rg h ees h ad
R a m b o u ille ts
or
R a m b o u ille ts had slo w er grow th r a t e s th a n C olum bias.
more
s t a b l e p e rfo rm an c es a c r o s s
C o lu m b ia s
and
T arg h ees showed
th e d i f f e r e n t i n t e r v a l s .
T here
was
47
70
-
60
7 3 . 1 1 - 6 5 . 4 7 e~
50
--
LU
7 2 .4 2 - 64.95 6
40
.iiost
-
7 0 .0 4 - 62.69 6
.1 2 0 0 1
1159t
C
6
12
18
24
3O
36
42
M O N T H S
48
,54
60
66
72
48
TABLE 15.
MEAN SQUARES AND LEAST SQUARES MEANS FOR ABSOLUTE GROWTH
RATE (AGR) FROM BIRTH-WEANING (AGR1), WEANH G -1 2 MONTHS (AGR2) 12-18
MONTHS (AGR3), BIRTH-12 MONTHS
(AGR 4) AND BIRTH-18 MCNlHS
(AGRS)
(HARVEY LEAST SQUARES ANALYSES, MODEL A)
S o u rces o f
v a ria tio n
Means S n u a re s
df
Year o f b i r t h 14
B reed (B)
2
Group ( G)
2
B xG
4
Age o f dam
5
Type of b i r t h r e a r in g
2
B ir th r a t e
I
R esid u al
772
AGRl
AGR2
AGRB
AGR4
AGR5
24.284**
18.247**
.1 3 7 "
.179
3 .591**
6.438«*
4.015**
.008
.496
.402
107.718**
14.191**
.912
.479
.399
7.482**
. 706 **
.093
.158
.315*
4 .0 6 9 * *
2.199**
.024
.053
.089
89.481**
10 . 261**
6.336**
.998*
.228
1.863*
2.972**
.035
.128
.5 1 3 * "
.186
.075
.830
.722
.544
Means
B re e d :
R am b o u illet
Targhee
Columbia
N
kg/mo
kg/mo
kg/mo
kg/mo
kg/mo
295
335
173
7 .9 4 ± .0 8 b
8 .5 4 ± .0 7 a
8 .2 7 ± .0 9 a
1 .4 5 ± ,0 4 a
1 .2 3 ± ,0 4 b
1 .1 5 ± ,0 5 b
2 .5 9 ± ,0 6 b
3.03± ,.06a
3 .1 0 ^ .0 8a
3 .7 3 ± .0 3 b
3 .8 l± .0 3 a
3.68±_.04b
3.28±..02c
3.49± ..02a
3.40±_.03b
198
8.22±>08a
8.29±,.10a
8 .2 4 ± .0 6 a
1 .2 8 ± ,0 4 a
1 .27±,.05a
I .2 9 ± ,0 3 a
2.99±>06a
2.86± ,.08a
2 .8 7 ± ,0 5 a
3.72± ,.03a
3 .7 6 ± .0 4 a
3 . 76± , 02 a
3 .4 l± .0 2 a
3 .3 7 t.0 3 a
3 .3 9 t.0 2 a
G roups
3 re c o rd s
4 re c o rd s
5 re c o rd s
129
476
a , b, a r e LSD com parisons (P < .01 )
** (P < .01)
49
and Columbiaa w ere s im ila r b u t s u p e r io r t o R a m b o u ille ts.
T arg h ees w ere
s u p e r io r to R a m b o u ille ts and C olum bias f o r AGRU, and R a m b o u ille ts w ere
s u p e r io r t o T arghees and Colum bias f o r AGF2.
fa s te r
g ro w th
ra te s
th a n
e ith e r
For AG16, T arg h ees had
R a m b o u ille ts
or
C o lu m b ia s
R a m b o u ille ts had slo w er grow th r a t e s th a n C olum bias.
more s t a b l e
p erfo rm an ces a c r o s s th e
T arg h ees showed
d iffe re n t in te rv a ls .
T here was
ev id en ce o f com pensatory grow th i n a d ja c e n t i n t e r v a l s from
w eaning
(AGR1) ,
from w eaning t o
(AGRB);
h o w e v e r,
th e
th re e
b irth
12 mo (AGR2) and from 12 to
b re e d s
d id
not
and
re sp o n d
th e
to
18 mo
sam e.
R am b o u ille ts had h ig h g a in s i n AGR2 and low i n AGRI and AGRB; w h ile th e
o p p o s ite was g e n e r a lly t r u e f o r th e o th e r two b re e d s .
d iffe re n c e s
d e te c te d
among g ro u p s nor was t h e r e
I n no c a s e w ere
a
b re e d
by group
in te ra c tio n .
SSiR comparisons.
T able 16 p r e s e n ts th e mean s q u a r e s and l e a s t
s q u a re s means f o r RGR i n th e f i v e grow th i n t e r v a l s .
e x c e p t from b i r t h t o w eaning (RGRI) and b i r t h
e ffe c ts
w e re
s ig n ific a n t.
RGR2 ,
For
In a l l i n t e r v a l s ,
t o 12 mo (RGR4), b re e d
R a m b o u ille ts
had
s u p e rio r
perform ance and T arg h ees and Colum bias h ad lo w er b u t s im ila r g ro w th .
H ow ever,
T a rg h e e s and C olum bias w ere
s u p e r io r
to
R a m b o u ille ts ,
and
Columbias s u p e r io r t o T arghees f o r RGRB.
Group d i f f e r e n c e s w ere n o t
d e te c te d
T arghees and C olum bias had
fo r
an y
in te rv a l.
For RGR5,
s im ila r grow th r a t e s and b o th were s u p e r io r t o R a m b o u ille ts .
No b reed
by group i n t e r a c t i o n s w ere o b se rv e d .
AMR COfflParl BOng.
s q u a r e s m eans f o r
T able 17 d is p la y s th e mean s q u a r e s and l e a s t
AMR i n
each
d i f f e r e n c e s w ere d e te c te d f o r a l l
of
th e
in te rv a ls
th e i n t e r v a l s .
s tu d ie d .
Breed
For AMRI, T arghees
50
TABLE 16.
MEAN SQUARES AND LEAST SQUARES MEANS FOR RELATIVE GROWTH
RATE (RGR) FROM BIRTH-WEANING (RGRI)t WEANING-12 MONTHS (RGR2) 12-18
MONTHS (RGRB) 1 BIRTH-12 MONTHS (RGRiJ) AND BIRTH-18 MONTHS (RGR5)
(HARVEY LEAST SQUARES ANALYSES, MODEL A)
Source o f
v a ria tio n
df
Year o f b i r t h 14
B reed i b ;
2
Group (G)
2
B xG
4
Age o f ram
5
Type o f b i r t h r e a r in g
2
B ir th d a te
I
R esid u al
772
RGRI
RGR2
RGRB
RGR4
RGR5
.000223
.002114**
.001867**
.000381
.000213
.000071
. 001836**
. 000006**
. 000001**
.000059**
.000005**
.000166
. 000222 *
.020499**
.001958**
.000173
,000113
.000169
.000002
. 000010**
.000000
.000001
. 000002**
.001135**
. 000618 *
.000104
.000088**
.000497**
.000003
.000035**
.000184**
.004851**
.000432*
.065677**
.000140
.000002
.004537**
.001047**
.000080
.000001
L east S a u a r e s M eansa
B re ed :
R am b o u illet
Targhee
Columbia
N
%/mo
%/mo
%/mo
%/mo
%/mo
295
335
173
21.6 ± . 1a
2 1 .7 8 ± ,1 a
21 . 65± . 1a
3 . 01± , 1a
2.52*.. Ib
2 .3 9 ± ,1 b
4 .1 8 ± ,1 c
4 .6 4 ± ,1 b
4 . 86*..I a
7 . 86* . 0 a
7 .8 9 * ,0 a
7.87*_.0a
5 .4 2 * ,0 b
5.45± ,.0a
5 .4 5 * ,0 a
198
129
476
21.80± ,.1a
2 1 .6 3 ± .1 a
2 1 .5 6 * . I a
2 . 66± , 1a
2 .6 3 ± ,1 a
2 .6 4 ± ,1 a
4.67*.. I a
4 .5 1 * ,1 a
4.51*.. I a
7.8 7 ± > 0 a
7 .8 8 * .0 a
7 . 86* . 0 a
5 .4 4 * .0 a
5 .4 4 * .0 a
5 .4 4 * .0 a
G roups:
3 re c o rd s
4 re c o rd s
5 re c o rd s
a RGR expressed a s percentage r e l a t i v e to Yt9
a, b, c, a re LSD comparisons
2
* * ( ? < .0 1 )
51
TABLE 17.
MEAN SQUARES AND LEAST SQUARES MEANS FOR ABSOLUTE MATURING
RATE (AMR) FROM BIRTH-WEANING (AMR1), WEANING- 12 MONTHS (AMR2), 12-18
MCNTHS (AMR3), BIRTH-12 MONTHS (AMRlt), BIRTH-18 MONTHS (AMR5) (HARVEY
LEAST SQUARES ANALYSES, MODEL A)
Mean S o u area
Source o f
v a ria tio n
df
Year o f b i r t h lit
B reed (B)
2
Group (G)
2
B xG
4
Age o f dam
5
Type o f b i r t h
r e a r in g
2
B ir th d a te
I
R esid u a l
77 2
AMRI
AMR2
AMR3
AMR4
AMR5
:
.00363**
. 00122**
.00013
.00007
.00026
.001550«» . 01838**
.001053** , . 00152«*
.000004
.00031»
.000089
.00013
.000098*
.00015
. 001980 "*
.000242**
.000006
.000024
.000015
.000417**
.000117**
. 000060*«
.000029
.01080**
. 00132**
.00017
.001575**
.000141
.000044
.000217**
.000048
.000025
.000026
.00084**
.00014
.00009
.000010
.000007
.000011
L e a st S q u a re s Mmana
B re e d :
N
R am b o u illet 295
xargnee
335
Columbia
173
%/mo
%/mo
%/mo
%/mo
%/mo
11.39iL.1o
1 1 .8 4 ± .1 a
11.41± .1b
2.09±j»05a
I .73 ± ,0 5 b
1.6l± ..0 7 c
3 .6 6 * ,0 8 b
4 .1 4 * .0 8 a
4 . 16* . 10a
5 .3 6 ± ,0 4 a
5 .3 0 * .0 4 a
5 .1 1 * .0 5 b
4 .7 1 * .0 3 b
4 .8 3 * .0 3 a
4 .6 8 * .0 4 b
*'
1 1 .63*. I a
1 1 .5 3 ± .1 a
1 1 .48*. I a
1 . 83±,.05 a
1 .8 0 * . 07 a
1 .8 l* .0 5 a
4 .1 4 * .0 8 a
3 . 88* . 10a
3 .9 4 * .0 7 a
5 .2 8 * .0 4 a
5 .2 5 * .0 6 a
5 .2 5 * .0 3 a
Group;
3 re c o rd s
4 re c o rd s
5 re c o rd s
198
129
476
a AMR a re expressed a s percentage r e l a t i v e to A
a, b, a re LSD comparisons (P<.01)
*(P<.05)I •*(P<e01)
4 .8 l* .0 3 a
4 .6 8 * .0 4 b
4 .7 2 * .0 2 a b
52
were s u p e r io r
fa s te r
th a n
to
R a m b o u llle ts and ColumM as,
R a m b o u ille ts .
H ow ever,
s u p e r io r t o T arghees and Columb ia s .
Colum bias i n th e i n t e r v a l (W-12 m o).
fo r
but
C olum bias m a tu re d
AMR2,
R a m b o u ille ts w ere
T arg h ees w ere f a s t e r m a tu rin g th a n
The AMR3 p e rfo rm a n c e s o f T arghees
and Colum bias were s i m i l a r and h ig h e r th a n R a m b o u ille ts .
w e re
s im ila r
C olum bias.
fo r
R a m b o u ille ts and T a rg h e e s;
AMRH v a lu e s
b o th w ere s u p e r io r
F in a l l y , f o r AMR5 T arg h ees w ere s u p e r i o r ; th e
to
R a m b o u ille ts
and C olum bias showed no d i f f e r e n c e .
Groups w ere d i f f e r e n t on ly f o r AMR5.
The th r e e - r e c o r d group had
f a s t e r m a tu rin g r a t e th a n th e f o u r - r e c o r d g ro u p , b u t was h o t d i f f e r e n t
from th e f i v e - r e c o r d group.
Growth s t a t i s t i c s
b re e d s w ere
s im ila r.
e s tim a te s fo r
th e
r e p o r te d
by S to b a r t
(1983)
The m ain d i f f e r e n c e s w e re
b irth
to w eaning and
f o r each o f th e s e
o b serv ed
fo r
RGR
12 to 18 months i n t e r v a l s .
A p p a re n tly , t h a t d if f e r e n c e was cau sed i n p a r t by th e d i f f e r e n t fo rm u la
used.
S to b a r t u sed a s r e f e r e n c e
( i n i t i a l w e ig h t + f i n a l w e ig h t
in s te a d ,
th e mean w e ig h t f o r
d iv id e d
by
tw o ) .
th e f i n a l w e ig h t was u sed a s r e f e r e n c e .
In
th e i n t e r v a l
th is
s tu d y ,
The f i n a l w e ig h t i n
th e i n t e r v a l was used a s r e f e r e n c e i n o rd e r t o m in im ize th e r e s i d u a l
e f f e c t s o f th e p re v io u s i n t e r v a l .
A lso ,
i n t h e case o f AMR v a lu e s ,
d i f f e r e n c e s w ere o b se rv e d i n th e 12 t o 18 mo i n t e r v a l , w hich would be
caused by th e d i f f e r e n t m ethod u se d by S to b a r t ( 1983) f o r e s tim a tin g
m ature s i z e .
53
E ffe ct
of
ty p e
of
b irth
and
re a rin g
on
g ro w th
and
p ro d u c tiv e
perform ance o f ewea.
Hie e f f e c t s o f th e ty p e o f b i r t h and r e a r i n g o f th e ewes on t h e i r
su b se q u e n t grow th and p ro d u c tiv e t r a i t s w ere o b ta in e d f o r each b re e d
from
th e Model B a n a ly s e s u s in g th e l e a s t
(1 9 7 7 ).
The l e a s t s q u a re s m eans f o r a l l
b re e d i n A ppendix T a b le 46 f o r r e f e r e n c e .
s q u a re s m ethod o f Harvey
tra its
a re lis te d
f o r e ac h
Hie a n a ly s e s u s in g Model C,
f o r th e combined d a ta s e t s , p ro v id e d th e e s ti m a te s o f th e l e a s t s q u a re s
means p re s e n te d i n T a b le 18.
The g e n e r a l r e s u l t s o f th e s e a n a ly s e s i n d i c a t e t h a t ewes b o rn tw in
and r a i s e d a s tw in s o r a s s i n g l e s had th e h ig h e s t r e p r o d u c tiv e r a t e ,
weaned th e
re la tiv e
g re a te s t
term s
w e ig h t o f lamb i n a b s o lu te
( E l) .
te rm s (ATWW) and i n
They produced s l i g h t l y l e s s wool and a t t a i n e d
s m a lle r m atu re w e ig h ts (A ).
EWes born and r a i s e d a s tw in s ap p ro ach ed
m atu re w e ig h t a t slo w er r a t e
( s m a lle r k ) th a n e i t h e r tw in s r a i s e d a s
s in g le s o r s in g le s ra is e d a s s in g le s .
E vidence o f c y c l i c a l grow th from
b i r t h t o 18 mo i n d i c a t e s some com pensatory grow th (O lso n e t a l . , 1976)
fo llo w in g
w e a n in g
e n v iro n m e n ta l
(AGR2).
T h is
a p p ea rs
c o n d itio n a s s o c ia te d w ith
ty p e
to
be
re la te d
of b irth
to
th e
and r e a r i n g .
Twins r a i s e d a s tw in s grew slo w e r p r i o r to w eaning b u t f a s t e r fo llo w in g
w eaning.
The o p p o s ite was g e n e r a lly t r u e f o r tw in s r a i s e d a s s in g le s
and s i n g l e s r a i s e d a s s i n g l e s .
A fte r 12 mo o f a g e , th e a b s o lu te grow th
r a t e s (AGR) f o r th e th r e e b i r t h and r e a r i n g g ro u p s w ere e s s e n t i a l l y th e
sam e.
The grow th r a t e s
d u rin g th e
combined th e in f l u e n c e s o f
e a r l i e r grow th segm ents.
lo n g e r
th e d i f f e r e n t i a l
p e rio d s
(AGR4 and AGR5)
grow th r a t e s
d u rin g
th e
The e f f e c t s o f ty p e o f b i r t h and r e a r i n g w ere
54
TABLE 18.
LEAST SQUARES MEANS FOR BIRTH-REARING EFFECTS CN ALL THE
STUDIED LIFETIME TRAITS OF THE EWES USING THE BREEDS POOLED INFORMATION
(HARVEY LEAST SQUARES ANALYSES, MODEL C)
Tvoe o f B i r t h - r e a r i n g
T ra its
S ig n if ic a n c e
No o f o b s e r v a tio n s
**
ATLB (n )
ATLW (n )
ATWW (k g )
ATFP (kg)
**
A (kg)
k (*/m o)
*»
EI (%)
**
AGRI (kg/m o)
**
AGR2 (kg/m o)
AGR3 (kg/m o)
**
AGR4 (kg/m o)
**
AGR5 (kg/m o)
**
RGRI (%/mo)
RGR2 (%/mo)
**
RGR3 (%/mo)
*#
RGR4 (%/mo)
**
RGR5 (%/mo)
Si
AMRI (%/mo)
**
AMR2 (%/mo)
AMR3 (%/mo)
**
AMR4 (%/mo)
*
AMR5 (%/mo)
• (P < .0 5 ), ** (P < .0 1 )
S/S
T/S
T/T
411
1.38±>03
1.1 7 ± ,0 3
45.72±>90
4 .8 1 ± ,0 4
73 .7 9 ± ,5 0
11. 66±,20
62.33±J.O
8 .8 1 ± ,0 7
1.09± ,04
2 .9 4 ± ,0 6
3 .7 9 ± ,0 3
3 .4 4 ± ,0 2
21.5 7 ± ,1 0
2 .1 7 1 ,0 7
4 .4 9 ± ,0 8
7 .7 5 1 ,0 1
5 .3 9 1 ,0 0
1 1 .9 9 t,1 0
1 .491,06
3 .9 3 1 ,0 8
5 .1 9 t,0 4
4 .6 8 i,0 2
65
1 .4 6 ± ,0 4
1 .2 3 1 ,0 4
4 8 .1 7 i 3 .7
4 .7 3 1 ,0 7
69 . 80±,86
1 1 .7 0 i,3 0
6 9 .2 5 t2 .0
8 .3 8 ± ,1 3
1.2 2 ± ,0 7
2 .8 4 ± ,1 0
3 .7 2 i,0 5
3 .3 6 i,0 4
2 2 .1 5 1 ,2 0
2 .5 2 i,1 0
4 .5 5 1 ,1 0
7 .8 7 1 ,0 2
5 .4 7 1 ,0 1
1 2 .0 3 1 ,2 0
1 .7 6 1 ,0 9
4 .0 4 i,1 0
5 .3 4 i,0 7
4 .8 2 i ,0 5
327
1 .4 7 1 ,0 3
1 .2 1 i,0 3
4 7 .6 3 1 ,9 2
4 .6 8 i,0 5
7 1 .3 7 1 ,5 1
1 0 .5 2 i,2 0
6 7 . 0 2 t J .0
7 .7 1 1 ,0 7
1 .3 9 1 ,0 4
3 .0 2 i,0 6
3 .5 9 1 ,0 3
3 .3 8 ± ,0 2
21 . 671,10
2 .9 7 1 ,0 8
4 .8 li,0 8
7 .8 3 1 ,0 1
5 .4 4 i,0 0
1 0 .8 5 1 ,1 0
1 .9 6 i,0 6
4 .2 0 1 ,0 8
5 .0 7 1 ,0 4
4 .6 9 1 ,0 3
55
c o n s id e ra b ly
dampened,
b u t d if f e r e n c e s
c o n tin u e d to
be
s ig n ific a n t,
w ith th e tw in b orn ewes h a v in g a n a p p a re n t e n v iro n m e n ta l d is a d v a n ta g e .
The e f f e c t s
s ig n ific a n t.
of
ty p e
of b irth
and r e a r i n g on BGR and AMR w ere
!W ins r a i s e d a s tw in s o r s i n g l e s had f a s t e r grow th i n th e
w eaning t o 12 mo i n t e r v a l .
The c y c l i c e f f e c t w hich had been o b se rv e d
when grow th was e v a lu a te d i n te rm s o f AGR a lm o st d is a p p e a re d betw een
th e s e two i n t e r v a l s when e v a lu a te d i n te rm s o f RGR and AMR.
The need
t o a d j u s t grow th and r e p r o d u c tio n r e c o r d s f o r type o f b i r t h and r e a r i n g
f o r m ost t r a i t s
b irth
s tu d ie d i s
c l e a r l y e v id e n t.
The e f f e c t s o f
ty p e o f
and r e a r i n g on r e p r o d u c tiv e perfo rm an ce in c lu d e , i n a d d i t i o n t o
e n v iro n m e n ta l f a c t o r s in v o lv e d i n th a t p a r t i c u l a r re s p o n s e , th e e f f e c t s
o f genotype o f tw in s compared t o
s in g le s .
E vidence f o r t h i s can be
drawn from B asu th a k u r e t a l . (1973) i n w hich th e ty p e o f b i r t h o f th e
s ire s
in flu e n c e d
th e
life tim e
re p ro d u c tiv e
p e rfo rm a n c e
of
th e ir
d a u g h te rs .
E s tim a te s o f I i e r l t a b i l l t i e s f g e n e tic and p h e n o ty p ic c o r r e l a t i o n s
L e a st s q u a re s m ixed-m odel a n a ly s e s w ere u sed t o o b t a i n e s ti m a te s
o f th e g e n e ti c and p h e n o ty p ic p a ra m e te rs f o r each b re e d (model B ).
In
th e a n a ly s e s o f th e p o o led d a ta , w hich in c lu d e d b re e d a s one o f
th e
s o u rc e s o f v a r i a t i o n ,
each
b re e d
a re
model C was em ployed.
p re s e n te d
in
A p p e n d ix
G e n e tic p a ra m e te rs f o r
T a b le s
47,
48
and
49
fo r
R a m b o u ille ts, T arg h e e s, and C olurnbias, r e s p e c t i v e l y .
Low h e r ! L a b i l i t i e s f o r
p ro d u c tiv e
tra its
and some n o n -e s tim a b le
h e r i t a b i l i t i e s (ATLW, ATWW, E l, RGRI, RGBS and AMR3) w ere o b ta in e d from
th e a n a l y s i s
of
th e
Targhee
d a ta .
T h is r e s u l t e d
g e n e tic c o r r e l a t i o n s in v o lv in g th e s e t r a i t s .
in
n o n -e s tim a b le
N e g a tiv e s i r e com ponents
56
o f v a r ia n c e w ere th e d i r e c t cause o f th e s e r e s u l t s .
Very low a d d i t i v e
g e n e tic v a ria n c e f o r th e s e t r a i t s i n T arg h ees and poor p r e c i s i o n i n th e
p a ram eter e s tim a te s
could be a re a s o n f o r
th a t
p a r t i c u l a r re s p o n s e .
However, sam ple s i z e was n o t c o n s id e re d a s a s p e c i f i c f a c t o r because
th e
Targhee
C olum bias.
sample
s i z e was l a r g e r
th a n e i t h e r
th e
R a m b o u ille ts or
I n each b re e d , th e e s tim a te s o f h e r i t a b i l l t i e s and g e n e tic
c o r r e l a t i o n s had r e l a t i v e l y la r g e s ta n d a rd e r r o r s .
I n g e n e r a l,
th e Columbia h e r i t a b i l i t y e s tim a te s w ere h ig h e r th a n
R a m b o u ille ts e x c e p t f o r some i n t e r v a l s o f grow th e v a lu a te d i n te rm s o f
RGR a n d AMR.
G e n e tic
c o rre la tio n s
showed more m arked d i f f e r e n c e s
between Colum bias and R a m b o u ille ts th a n t h e i r h e r i l a b i l i t i e s .
G e n e tic
c o r r e l a t i o n s in v o lv in g ATWW w ith ATFP and k i n C olum bias w ere low , but
p o s itiv e in
A ls o ,
c o n tra s t to
th e
g e n e tic
R a m b o u ille ts w hich w ere h ig h
c o rre la tio n
betw een
and n e g a t i v e .
ATWW and E I was h ig h
and
p o s itiv e ( .7 9 ) f o r Colum bias compared t o R a m b o u ille ts w hich was low and
n e g a tiv e ( - . 03 ) .
G e n e tic c o r r e l a t i o n in v o lv in g grow th s t a t i s t i c s w ere s i m i l a r f o r
Columbias and R a m b o u ille ts.
The grow th s t a t i s t i c s f o r th e w eaning t o
12 mo i n t e r v a l showed h ig h and p o s i t i v e g e n e ti c r e l a t i o n s h i p w ith th e
m ost im p o rta n t r e p r o d u c tiv e t r a i t s
c o n s ta n c y
of
re la tio n s h ip
th a t
a s s o c ia tio n
p ro b a b ly e x i s t s
(ATWW and E l) i n both b re e d s .
in
th a t i s
b o th
b re e d s
The
s u g g e s ts a g e n e r a l
not p a rtic u la r
to
a
s p e c ific
b re e d .
The g e n e t i c
p o s itiv e ;
g e n e tic
in
c o rre la tio n
th e R a m b o u ille ts ,
a s s o c ia tio n
of
b etw een ATFP and A i n
th e a s s o c i a t i o n w as
ATFP w ith
th e
C o lu n b ia s was
n e g a tiv e .
grow th s t a t i s t i c s
The
was q u i t e
57
d iffe re n t
in
fla m b o u ille t,
w h ile i n
th a t
th e
th e s e
two
even f o r
b re e d s.
A lm o st
n e g a tiv e
in
th e i n t e r v a l s w eanlng-12 mo and B ir th - 1 2
mo,
Colum bias th e s e a s s o c i a t i o n s w ere
c o rre la te d
s ta tis tic s in
re sp o n se
in
a ll
w e re
p o s itiv e .
ATFP fro m
T h is s u g g e s ts
s e le c tio n
fo r
th e i n t e r v a l i n d i c a t e d would be d i f f e r e n t f o r
grow th
th e
two
b re e d s.
D i f f e r e n c e s w e re
betw een A and E l.
a ls o
o b serv ed
fo r
th e
g e n e tic
T his c o r r e l a t i o n was n e g a tiv e i n
c o rre la tio n s
C olum bias w hich
means t h a t s e l e c t i o n o f a n im a ls f o r l a r g e r m a tu re s i z e s w i l l r e s u l t i n
progeny w ith lo w er
e ffic ie n c y .
The o p p o s ite would
be e x p e c t e d
in
R a m b o u ille ts w here th e g e n e ti c c o r r e l a t i o n was p o s i t i v e .
fisae tl c paraa ste r s from th e pooled da t a
and Col umbi an).
( R a m b o iin ie ts . T = ,.,* ...
T able 19 shows th e h e r i t a b i l i t i e s ,
and g e n e ti c and
p h e n o ty p ic c o r r e l a t i o n s among th e t r a i t s s tu d ie d from th e p o o led d a ta
u s in g model C.
These a n a ly s e s were perfo rm ed i n o rd e r to o b ta in more
g e n e ra l e s tim a te s o f th e g e n e tic p a ra m e te rs w ith th e a d v a n ta g e s r e l a t e d
to
th e g r e a t e r sample s i z e .
The sample s iz e
c o n s id e r a tio n p r im a r ily
a f f e c t e d th e e s tim a tio n o f th e s i r e s com ponents o f v a r ia n c e .
He r i t a b l l i tY
e stlm m W P .
The h e r l t a b i l l t y
e s tim a te s f o r
th e
p ro d u c tiv e t r a i t s ATLB1 ATLWt ATWV and ATFP w ere .4 3 , .3 3 , .11 and . 68 ,
re s p e c tiv e ly .
S to b a r t (1983) fo u n d v a lu e s o f .4 2 , .0 8 , .03 and .31 f o r
Timon and E is e n ( 1969) r e p o r te d
s im ila r to th o s e found i n t h i s s tu d y .
e s tim a te s o f h e r i t a b i l i t i e s f o r A, e s tim a te d from R ic h a rd s and L o g i s tic
m odels i n m ice,
t h a t w ere l a r g e r .
The h e r l t a b i l l t y
e s tim a te s f o r k
w ere s m a lle r when k was d e riv e d from R ic h a rd s e q u a tio n ,
when d e riv e d from th e L o g i s t i c e q u a tio n .
but g re a te r
58
TABLE 19.
ESTIMATES3
OF HERITABIL!TIES, GENETIC AND PHENOTYPIC
CORRELATIONS FOR LIFETIME PRODUCTION AND GROWTH TRAITS (POOLED DATA
FROM RAMBOUILLET1 TARGHEE AND COLUMBIA EWES)
ATLB
ATLW
ATWW
ATFP
A
k
EI
AGRl
AGR2
AGRj
AGR4
AGR5
RGRI
RGR2
RGRj
RGR4
RGRt>
AMRl
AMR2
AMRj
AMR4
AMR5
.4 1 + .IR
.9 1 ± ,1 2
1 .1 3 ± ,5 4
.20±>24
.2 3± j.j0
. 13±>26
. 82± ,3 0
.37± ,43
.09j> 23
.3 1 ± ,2 8
.2 3 ± ,2 8
.4 9 ± ,2 9
.0 3 ± ,2 5
.08± ,23
.1 5 ± ,3 0
.01 ± ,30
.04± ,24
.0 4 ± ,2 9
. 09±,22
•30±,33
•05±,25
.2 6 ± ,2 5
ATLW
ATWW
ATFP
.72
.65
.3 3 + .IR
.95
1 .1 3 ± ,3 0
.1 1 + .IR
•20± ,28
.2 7 ± ,5 0
• 17±,35
• 51±,73
.0 9 ± ,2 9 .004±>51
.9 2 ± ,1 0
.75±>32
-.2 7 ± ,4 9 -.5 4 ± ,9 1
•45±,27
.8 6 ± ,7 4
- .1 4 ± ,3 3 - • 16±,56
•38± ,32
•75±,72
•30±,33
.6 2 ± ,6 5
-.2 4 ± ,2 9 - .4 2 ± ,5 9
.5 1 ± ,2 8
•95±,81
-.2 8 ± ,3 4 -» 4 3 ± ,6 5
»26 ± ,34
•64±,73
.05± ,27 - . 1 1±,46
- .3 5 ± ,3 3 -.8 0 ± ,7 7
.46± ,27
.82 ±,71
-.2 3 ± ,3 6 '-• 3 9 ± ,6 6
.23± ,27
.3 3 ± ,4 9
•14± ,29
• 12±,49
.02
.01
.04
- 6 8+ .16
• 04±,25
- . 20±,22
.0 9 ± ,4 0
-.8 1 ± ,4 3
. 38±,18
-.0 9 ± ,2 4
.0 5 ± ,2 3
- .1 2 ± ,2 4
- .4 2 ± ,2 2
•3 9 ± ,1 9
-.1 4 ± ,2 6
-.2 8 ± ,2 7
-.4 3 t,2 2
-.5 8 ± ,2 8
»34±,18
-.1 7 ± ,2 9
-.0 4 ± ,2 0
“ *24±,23
A
k
CJ
O
ATLB
.12
—•11
—.07
.27
.4 2 + .IR
-.6 8 ± ,4 4
- . 29±,62
.07 ± ,44
• 15±,22
.49± ,26
.29± ,26
• 55±,21
- .2 4 ± ,2 5
.07 ±,23
.38± ,31
-.2 1 ± ,3 1
- .0 9 ± ,2 4
-.7 4 ± ,4 7
-.0 3 ± ,2 3
.2 5 ± ,3 6
-.5 7 ± ,3 7
-.5 6 ± ,4 1
.17
.17
- .0 9
- .5 3
.R6+.16
.4 6 ± ,3 8
.0 6 ± ,3 8
.3 2 ± ,1 9
-.38± > 30
.3 6 ± ,2 2
-.0 2 ± ,.2 7
. 29±>21
. 36±.20
-.4 3 ± ,3 4
.6 4 ± ,2 4
.4 6 ± ,2 0
.6 2 ± ,1 5
.4 4 ± ,1 7
- . 1 9±,30
.8 9 ± ,0 5
.78±.,09
3H e r i t a b i l i t y e s tim a te s a r e u n d e rlin e d , p h e n o ty p ic c o r r e l a t i o n s above
th e h e r i t a b i l i t i e s , and g e n e tic c o r r e l a t i o n s below h e r i l a b i l i t i e s .
59
TABLE 19..
T ra its
ATLB
ATLW
ATWW
ATFP
A
k
EI
AGRI
AGR2
AGR3
AGR4
AGR5
RGRI
RGR2
RGR3
RGR4
BGR5
AHRI
AMR2
AMR3
AMR4
AMR5
(C o n tin u ed )
RGBI
RGR2
RGR3
RGR4
RGBS
AMfll
-.0 0 2
.02
.01
.06
.06
.05
- .0 8
. 06
.03
.0 4
.05
.06
.06
.06
.0 9
.0 8
- .0 5
- .0 4
- .0 9
.006
•I^
- .0 4
•58
- .0 9
- .1 8
.16
•05
J i S + .15
•23±>20
'.02± ,26
•31±,23
•18±,20
.8 0 1 ,1 4
.1 7 1 ,2 0
.0 2 i,2 8
• 20 t ,20
.0 8 1 ,2 2
.04
.05
, .02
.13
.05
- .4 3
.97
-.3 4
.48
.14
- .1 3
.7 6 +.16
- .7 2 ± ,3 8
.5 3 ± ,2 0
.2 4 ^ 1 8
- .3 6 t,3 0
.9 9 i,0 1
-.6 7 ± ,4 0
.7 6 i #13
.3 0 1 ,2 0
—.28
.02
- .1 8
- .5 0
.93
-.5 5
.20
-.21
- .4 3
.40 + .1 6
-.1 7 ± ,3 6
.13±>24
.0 4 t,3 1
-.7 3 i> 3 9
. 95i ,03
—.6 5 1 ,4 4
.1 1 i,2 7
-.11
.01
.23
.05
.1 8
.32
- .2 4
.46
.22
.34
.31
- .3 3
.4 0 + .1 6
.7 0 1 ,1 6
.2 7 t,2 9
.6 lt,1 9
.0 2 t,3 6
.7 7 1 ,1 8
.5 4 i,2 5
.06
- .1 4
.01
.15
.05
.05
.12
.26
.19
.36
.11
.12
.23
.52
.70 + .1 $
.2 1 1 ,2 4
.3 0 i,1 8
.2 5 1 ,2 5
.5 0 i,1 9
« 6 4 i,16
-.1 0
- .4 0
.67
.1 9
.70
- .3 4
-.21
.1 8
.01
.57
- .4 3
- .2 3
.17
.04
.4 4 + .1 6
- .2 8 i,2 6
.2 2 t,3 4
.3 0 1 ,2 1
.3 8 1 ,2 1
60
TABLE 19.
(C o n tin u ed )
T ra its
EI
ATLB
ATLW
ATWW
ATFP
A
k
EI
AGRI
AGR2
AGR3
AGR4
AGR5
RGRI
RGR2
RGR3
RGR4
RGRs
AMRI
AMR2
AMR3
AMR4
AMR5
AGRI
AGR2
AGR3
AGR4
.11
.10
.0 9
.09
-.0 0 4
.27
.05
.60
.09
.04
.92
.95
.02
.01
.04
.05
.11
.11
.12
-.0 6
- .3 7
.3 2
-15+ .15
-.5 7 ± ,8 3
.6 3 ± ,4 8
-.44± >52
.4 2 ± ,5 2
• 13±,50
-.2 0 ± ,4 2
.77± ,52
-.6 0 ± ,5 7
•71±,59
.1 8 ± ,3 9
-.2 0 ± ,4 5
•73±,47
-.4 5 ± ,5 3
.66±>41
.4 9 1 ,3 6
.36
.26
—.08
.1 9 + .1 5
- .4 5 1 ,4 4
.5 1 1 ,4 8
- .1 8 1 ,3 9
- .0 3 1 ,4 7
.9 0 1 ,2 5
- .4 8 i ,51
.5 9 1 ,5 9
.1 7 1 ,4 3
.2 5 1 ,3 5
.6 1 1 ,2 4
-.4 8 ± ,4 7
.7 3 1 ,6 6
- .2 7 1 ,3 4
- . 061,38
.1 9
.17
- .2 5
. 80*.16
- .4 7 1 ,3 0
.9 6 i,1 0
.431,21
- . 20t ,20
.991,01
- .6 7 1 ,3 8
.5 2 i,2 0
.2 7 1 ,2 0
- .4 0 i,2 9
.9 8 t,0 1
• - .6 li,3 9
.7 0 1 ,1 2
.2 5 1 ,1 9
-.21
-.002
- .0 0 3
-.3 6
.4 4 + .1 5
- .2 4 1 ,3 2
.7 2 1 ,2 0
- .0 9 1 ,2 5
-.5 2 1 ,3 1
.9 5 1 ,0 4
- . 10i ,3 2
.1 5 1 ,2 3
- .1 0 i,3 1
- .5 4 t,3 2
.9 6 i,0 4
- .5 8 i,3 8
.1 3 1 ,2 5
.10
.20
.41
.3 8
- .0 4
.51
.65
- .2 7
.4 8 + .1 5
.5 5 1 ,1 9
.0 4 t,2 4
.9 4 i,1 4
-.4 4 i,4 3
.6 7 1 ,2 0
.5 0 i,2 1
- .3 1 i,2 6
.9 0 1 ,1 2
- .3 2 i,4 2
.6 2 1 ,1 5
.2 5 1 ,2 3
AGR5
.17
.11
.15
.1 8
.61
.15
- .0 5
.48
.28
.50
.67
.4 0 + .1 5
- .2 0 i,2 5
.3 9 1 ,2 3
.5 6 i,2 9
.2 8 i,2 9
.5 3 1 .2 0
-.4 8 i,3 2
.3 4 1 .2 2
.7 2 1 ,3 0
.0 2 i,2 6
.3 7 1 ,2 3
61
(C o n tin u ed )
T ra its
AMR2
ATLB
ATLW
ATWW
ATFP
A
k
EI
AGRI
AGR2
AGR3
AGfW
AGR5
RGRI
RGR2
RGR3
RGR4
RGR5
AMRI
AMR2
AMR3
AMR4
AMR5
.04
.0 4
.07
.0 4
•*.08
.31
.10
- .3 3
.97
-.4 1
- .5 6
.14
- .0 9
.97
-.5 1
.33
.12
OX
i
ro
TABLE 19 .
. 81*.16
-.6 5 + .4 0
.8 1 + .0 9
.3 6 + .1 8
AMR3
AMR4
AMR5
.11
.08
.13
.14
- .0 5
- .4 4
.85
.27
.24
.24
- .1 4
- .5 0
.81
.3 8
.14
.12
-.11
- .0 9
- .0 3
.14
- .1 3
- .4 3
.92
- .4 5
.2 8
- .1 8
- .3 7
.95
- .2 7
.25
- .0 7
-.4 1
.I ? * .is
- .4 5 + .3 8
.4 0 + .2 5
.16
.20
.11
.5 3
- .5 0
.62
.13
.16
.46
.15
.12
-.61
.14
.45
.18
.52
.63
- .3 8
.6 4 *.16
.6 7 + .1 2
.22
.22
.24
.37
.06
.3 9
.4 8
.2 5
.41
.66
.5 6 * .1 6
62
each of these t r a i t s , re s p e c tiv e ly .
ATLB and fo r ATWW.
The e stim a te s were sim ila r f o r
However, the d iffe re n c e s were la rg e fo r estim ates
o f ATLW and ATFP.
H e r i t a b i l i t y e s tim a te s f o r A and k w ere .4 2 and .5 6 , r e s p e c t i v e l y ,
in d ic a tin g
th a t
A and k e a c h would re s p o n d
re a d ily
to
s e le c tio n .
H e r i t a b i l i t i e s r e p o r te d f o r A and k i n m ice (E is e n e t a l . , 1 9 6 9 ), when
e s tim a te d from th e grow th p a ra m e te rs d e riv e d from th e L o g i s tic grow th
e q u a tio n ,
w ere v e r y
s im ila r
to
th e
v a lu e s
fo u n d i n
th is
s tu d y .
However, h e r ! t a b i l i t i e s f o r th e k p a ra m ete r d e riv e d from Gcmpertz and
B e r ta la n f f y m odels gave h ig h e r e s tim a te s .
H e r i t a b i l i t y e s tim a te s f o r
A, f o r a l l th e grow th m odels used by th e s e a u th o r s , gave v a lu e s v e ry
These two s t u d i e s d e riv e d grow th p a ra m e te rs from d i f f e r e n t grow th
e q u a tio n
m o d e ls
h e rlL a b ilitie s .
and
used
th e
f u l l —s i b
m e th o d
fo r
e s tim a tin g
T h is c o u ld be a cause o f some o f th e d if f e r e n c e i n th e
e s tim a te s o b ta in e d .
T here i s
a ls o a s p e c ie s and m ethod o f r e a r i n g
e f f e c t in v o lv e d .
The h e r i L a b i l i t y
found by Brown e t a l .
e s tim a te s f o r
A and k w ere h ig h e r
( 1972) u s in g Brody* s model i n c a t t l e .
th a n th e s e
However,
th e av erag e o f th e h e r ! L a b i l i t i e s fo u n d by th e s e a u th o r s f o r k was v e ry
s im ila r.
DeNise and B rin k s (198 5 ) r e p o r te d e s ti m a te s o f h e r ! t a b i l i t i e s
f o r A, u s in g B rody’ s and R ic h a rd s '
m odels,
i n c a t t l e t h a t w ere v e ry
s im ila r t o th e v a lu e s fo u n d i n t h i s s tu d y , b u t t h e i r e s tim a te f o r th e
h e r ! L a b i l i t y o f k was s m a lle r .
The h e r ! L a b i l i t y e s tim a te f o r EI was .1 5 .
T h is low v a lu e s u g g e s ts
t h a t th e d i r e c t s e l e c t i o n by E I w ould n o t produce a r a p i d re s p o n s e .
No
o th e r r e p o r t s i n th e l i t e r a t u r e f o r t h i s ty p e o f in d e x c o u ld be fo u n d .
63
H e r i t a b i l i t i e s o f th e e s tim a te d grow th s t a t i s t i c s w ere i n g e n e r a l
l a r g e r th a n th e grow th s t a t i s t i c s fo u n d i n sheep ( S t o b a r t , 1983 ) and i n
c a ttle
( F it z h u g h
h e rlta b ility
an d
T a y lo r,
e s ti m a te s found
1971;
S m ith
by S to b a r t
et
(1 9 8 3 )
a l.,
1976 ) .
The
t h a t w e re i n
c lo s e
agreem ent to th o s e found i n t h i s s tu d y were f o r th e 12-18 mo i n t e r v a l .
For AGR, RGB and AMR, th e a u th o r r e p o r te d .4 3 ,
alm ost i d e n t i c a l
th e s e
tra its ,
s ta tis tic s
to
.4 4 ,
.3 9 , and .3 2 which a r e
.40 and .3 3 found i n th e p r e s e n t s tu d y f o r
re s p e c tiv e ly .
The h e r l t a b i l i t y
e s tim a te s
fo r
th e
i n o th e r i n t e r v a l s w ere d i f f e r e n t and d id n o t fo llo w
th e
same p a t t e r n s .
The l a c k
e s tim a te s w ith
of
c lo s e
used i n
o b ta in e d
set
of
of
th e AMR v a lu e s fo u n d i n
c o m p u ta tio n al d if f e r e n c e
v a lu e s
a g re e m e n t
S t o b a r t 's
th is
AMR h e r l t a b i l i t y
stu d y m ig h t
u sed i n each case a s n o te d e a r l i e r .
c a lc u la tin g
AMR v a lu e s f o r
th e
p re se n t
by a p p ly in g B ro d y 's e q u a tio n in d e p e n d e n tly
a g e - w e ig h t
be due t o
d a ta .
S t o b a r t 's
d e te rm in a tio n o f m atu re s i z e ,
a p p ro a ch
w as
The A
s tu d y
w e re
to each a n im a l's
b ased
on
th e
which c o rre sp o n d s ro u g h ly to B ro d y 's A
v a lu e , u s in g th e mean o f w e ig h t d a ta from 42 and 54 mo.
These d a ta
re p re s e n te d 9 4 .9 and 97.6% o f m atu re mean w e ig h t, r e s p e c t i v e l y .
Ih is
r e s u l t i s i n d is a g re e m e n t w ith th e f in d i n g s i n t h i s s tu d y w here 97% o f
m ature s i z e was found a t 30 mo and 98% a t 36 mo o f a g e .
M ature s i z e
was re a c h e d i n R a m b o u ille ts, T arg h ees and C olum bias a p p ro x im a te ly a t 39
mo, 38 mo and 41 mo, r e s p e c t i v e l y .
With r e s p e c t t o th e h e r l t a b i l i t y e s tim a te s o f RB*, th e d i f f e r e n t
fo rm u la used by S to b a r t (19 8 3 )
c o u ld
be th e r e a s o n f o r p a r t
of
th e
d if f e r e n c e . For «08, th e r e m ig h t be th e o th e r c a u s e s ; t h e sam ple I t s e l f
64
and th e d i f f e r e n t model u sed f o r
th e
e s tim a tio n
o f th e
g e n e ti c and
p h e n o ty p ic p a ra m e te rs .
t l C—correl a ti ons,.
im p o r ta n t
p ro d u c tiv e
p aram eter A w ere .5 1 ,
G e n e tic
tra its ,
c o rre la tio n s
ATWW,
ATFP,
betw een th e most
and E l,
.0 4 , and - . 2 9 , r e s p e c t i v e l y .
and th e
grow th
The c o rre s p o n d in g
g e n e tic c o r r e l a t i o n s w ith k w ere .0 0 4 , - .2 0 and .4 6 , r e s p e c t i v e l y .
The
g e n e tic c o r r e l a t i o n betw een A and k was - . 68 , a v a lu e v e ry s im ila r t o
th o se p re v io u s ly r e p o r te d i n o th e r s p e c ie s : m ice ( E is e n e t a l . , 1969)
and c a t t l e (Brown e t a l . , 1972; DeNise and B r in k s , 1 9 8 5 ).
The g e n e tic c o r r e l a t i o n s fo u n d betw een th e p r o d u c tiv e t r a i t s and
the grow th p a ra m e te rs A and k i n d i c a t e t h a t ewes w ith a h ig h e r a d d i t i v e
g e n e tic p o t e n t i a l f o r p ro d u c tio n o f Ia n b a t w eaning (ATWW) w i l l have an
a d d itiv e
g e n e tic
s e le c te d
fo r
advantage
ra te .
in
nor
p o t e n t i a l f o r l a r g e r m ature
lamb p ro d u c tio n
d is a d v a n ta g e i n
s iz e .
H o w ev er,
ew es
(ATWW) would h av e n e i t h e r a c o r r e l a t e d
th e ir
g e n e tic
p o te n tia l
fo r
m a tu r in g
The sm all but p o s i t i v e g e n e ti c c o r r e l a t i o n betw een ATFP and A i s
agreem ent w ith
T u rn er and
f in d in g o f S to b a r t
(1 9 8 3 ).
r a p id m a tu rin g r a t e ,
Young (1 9 6 9 ),
but is
The c o r r e l a t e d
e ffe c t
c o n tr a r y
of
to
th e
s e le c tio n
fo r
k, would be a r e d u c tio n i n m atu re s i z e ,
h ig h e r
lamb p ro d u c tio n e f f i c i e n c y ( E I ) , r e d u c tio n i n g re a s e f l e e c e w e ig h t, bu t
no e f f e c t on a b s o lu te w e ig h t o f lam bs w eaned.
The
d is a d v a n ta g e
o f u s in g e i t h e r
c r i t e r i a r e s t s on when i t
th e a n im a ls u n t i l
A or
can be e s tim a te d .
k a s a m ass s e l e c t i o n
I t w ould r e q u i r e k e e p in g
th ey re a c h a d u l t o r m a tu re s i z e s w hich i s to o l a t e
fo r e ffe c tiv e s e le c tio n .
BGR2 would p e rm it e a r ly
However, s e l e c t i o n on th e b a s i s o f AGR2 or
s e l e c t i o n a t 12 mo.
Both AGR2 and RGR2 have
65
h igh g e n e tic
c o r r e l a t i o n s w ith
ATWW (.86
and
.9 5 )
and EI
( .6 3
and
.7 7 ) . AGR2 and RGR2 a ls o have r e l a t i v e l y h ig h g e n e tic c o r r e l a t i o n s w ith
ATFP ( .3 8 and .3 9 ) .
The c o r r e l a t e d re s p o n s e s from s e l e c t i o n on th e
b a s is o f AGR2 o r RGR2 would be p o s i t i v e f o r th e s e t r a i t s bu t w ould show
l i t t l e e f f e c t on A.
G e n e tic
c o rre la tio n s
among p r o d u c tiv e t r a i t s
w ere a l l
p o s itiv e .
ATFP had low g e n e tic c o r r e l a t i o n s w ith ATLBt ATLW and ATWW. ( . 2 0 ,
and .27» r e s p e c t i v e l y ) .
ATLB was h ig h ly c o r r e l a t e d w ith ATLW and ATWW
(.9 1 and 1.13» r e s p e c t i v e l y ) .
w as
h ig h
( 1 .1 3 ) .
T h e re
c o r r e l a t i o n s r e p o r te d
.20
The c o r r e l a t i o n
w as p o o r
betw een ATLW and ATWW
a g re e m e n t
by S to b a r t (1 9 8 3 ),
w ith
p a rtic u la rly
th e
fo r
g e n e tic
ATFP-ATLW,
ATFP-ATWW and ATLW-ATWW w hich w ere - . 4 4 , - .0 1 and .8 4 , r e s p e c t i v e l y .
The v e ry low p o s itiv e g e n e tic r e l a t i o n s h i p o b ta in e d betw een ATFP
w ith
ATLB and ATLW c o n t r a d i c t s th e f i n d i n g o f a l i f e t i m e p r o d u c tiv ity
s tu d y
in
R a m b o u ille t
c o rre la tio n s
o f - .1 3
by
S h e lto n
and - .2 5
an d
H e n z ie s
betw een th e s e
( 1968 )
tra its ,
who
fo u n d
re s p e c tiv e ly .
However, th e s e c o r r e l a t i o n s w ere v e ry s im ila r t o th o se o b ta in e d i n th e
p re s e n t stu d y f o r R a m b o u ille ts. . P erh ap s th e e f f e c t o f th e p o o led b re e d
a n a ly s is c o n tr ib u te d t o t h i s r e s u l t
s in c e R a m b o u ille ts w ere d i f f e r e n t
from Targfcees and C olum bias i n t h i s s tu d y .
Very h ig h g e n e tic
same i n t e r v a l s
c o rre la tio n s
s u g g e s ts t h a t
betw een AGR, RGR and AMR i n
RGR a n d (o r)
AGR v a lu e s
c o u ld
th e
be used
a d v an ta g e o u sly f o r e a r l y s e l e c t i o n i n s t e a d o f AMR when s e l e c t i o n f o r
r a p id m a tu r ity i s d e s ir e d .
and AMR w i th in
th e
The g e n e ti c c o r r e l a t i o n s betw een AGR, RGR
same i n t e r v a l s r e p o r te d
by S to b a r t
(1 9 8 3 )
w e re
66
g e n e r a lly lo w e r.
T h is was e s p e c i a l l y
tru e fo r
b irth
t o w eaning and
b i r t h to 18 months i n t e r v a l s , where n e g a tiv e e s tim a te s w ere o b ta in e d .
G e n e tic
s ta tis tic s
c o r r e l a t i o n s r e p o r te d
and p r o d u c tiv e
e s tim a te d v a lu e s i n
c o rre la tio n s
of
th is
by S to b a r t
tra its
w ere g e n e r a lly
s tu d y .
ATFP w ith
(1985)
AGR2,
betw een g ro w th
d i f f e r e n t w ith
th e
Good ag reem en t was fo u n d f o r
th e
HGR2 and AMR2 where th e h ig h e s t
p o s itiv e v a lu e s w ere fo u n d i n both s t u d i e s .
However, th e c o r r e l a t i o n s
betw een th e grow th s t a t i s t i c s w ith th e r e p r o d u c tiv e t r a i t s ATLB, ATLW
and ATWW were d i f f e r e n t both i n v a lu e s and p a t t e r n s .
The d if f e r e n c e i n
t h i s case c o u ld be due to th e f a c t t h a t S to b a r t u sed d a ta from ewes,
some o f w hich d id n o t com pleted t h e i r l i f e t i m e p ro d u c tio n .
The g e n e tic
T a y lo r (1971)
c o rre la tio n s
grow th
g e n e tic r e l a t i o n s h i p
o f ATWW and E I w ith
s ta tis tic s
e x is ts
suggest
th a t
betw een grow th i n
p e rio d and p ro d u c tiv e p erfo rm an ce.
a
th e
F itz h u g h
p o s itiv e
and
a d d itiv e
th e w eaning t o
12 mo
The g e n e ti c c o r r e l a t i o n s
betw een
th e s e grow th s t a t i s t i c s and ATFP w ere a ls o p o s i t i v e b u t lo w e r.
From a
p ra c tic a l
p o in t o f view ,
BGR i n
th e w eaning t o
12 mo i n t e r v a l
age
sh o u ld be u s e f u l i n s e l e c t i o n .
The p o s i t i v e c o r r e l a t i o n found betw een th e grow th s t a t i s t i c s f o r
w eaning-12 mo i n t e r v a l w ith ATWW and BI a p p e a rs to be b ased i n p a r t on
th e a n im a l's g e n e tic c a p a c ity f o r overcom ing s t r e s s a f t e r w eaning and
d u rin g th e f a l l - w i n t e r grow th p e rio d and i t s
e f f i c i e n t l i f e t i m e p ro d u c tio n .
a s s o c i a t i o n w ith a more
However, th e g e n e ti c c o r r e l a t i o n s o f EI
w ith th e grow th d u rin g th e b irtb -w e a n in g i n t e r v a l w ere n e g a tiv e .
T h is
l a t t e r o b s e r v a tio n s u p p o rts th e id e a t h a t e a r l y grow th from b i r t h
w eaning sh o u ld n o t
be u s e d
as
s e le c tio n
c rite ria
fo r
to
su b sequent
67
p r o d u c tiv ity .
S h e lto n and H enziea
(1968) i n d i c a t e d a
p re fe re n ce f o r
u s in g y e a r l i n g w e ig h t a s a m easure o f th e p r o d u c tiv e p o t e n t i a l o f th e
ewes i n s t e a d o f w eaning w e ig h t.
P h e n o ty p ic c o r r e l a t i o n s . Ih e p h e n o ty p ic c o r r e l a t i o n s o b ta in e d from
th e p o o led d a ta (model C a n a ly s e s ) in d i c a t e r e l a t i o n s h i p s among t r a i t s
due t o a v a r i e t y
o f f a c t o r s t h a t a f f e c t th e in d iv id u a l a n im a l.
The
c o r r e l a t i o n s betw een ATLW-ATFP ( .0 1 ) and ATWW-ATFP ( .0 4 ) i n d i c a t e t h a t
lamb p ro d u c tio n ,
u n d er
th e
ra n g e
c o n d itio n s
in
produced t h e i r lam bs, d id n o t a f f e c t t h e i r ATFP.
w ith B asu th ak u r e t a l .
(1 9 7 3 ),
but
w h ic h
th e s e
ew es
T h is re s p o n s e a g re e s
does n o t a g re e w ith
S h e lto n and
M enzies ( 1968) .
S to b a r t (1983) r e p o r te d p h e n o ty p ic c o r r e l a t i o n s t h a t w ere somewhat
h ig h e r, b u t had th e same p a t t e r n .
between th e
c o r r e l a t i o n s ATLB-ATFP and ATWW-ATFP.
w ere .13 and . 17, compared .02 and
s tu d y .
There w ere s u b s t a n t i a l d i f f e r e n c e s
S t o b a r t 's
.0 4 , r e s p e c t i v e l y ,
v a lu e s
fo u n d i n
th is
A lso, th e c o r r e l a t i o n betw een E I and ATFP ( - .0 6 ) i n d i c a t e s t h a t
lamb p ro d u c tio n r e l a t i v e
t o m ature
s iz e
d id
not
a ffe c t
th e
e w e 's
perform ance f o r ATFP.
On th e o th e r hand, th e c o r r e l a t i o n o f -.3 7 found betw een E I and A
in d ic a te s
th a t
th e m ost e f f i c i e n t
ew es t e n d
to
be
s m a lle r.
The
e s tim a te d g e n e tic c o r r e l a t i o n and th e e s tim a te d p h e n o ty p ic c o r r e l a t i o n
betw een E I and A w ere s i m i l a r ( - .2 9 and -.3 7 »
r e s p e c t i v e l y ) s u g g e s ts
t h a t p a r t o f th e r e l a t i o n s h i p betw een th e n may be due t o th e e f f e c t o f
re p ro d u c tio n i t s e l f . P a r t o f th e re a s o n s m a lle r s i z e ewes p ro d u ced more
lamb a t w eaning p e r u n i t o f w e ig h t may be becau se
c o n tr ib u te s to s m a lle r m atu re s i z e s .
t h e i r p ro d u c tio n
T h is s u g g e s tio n i s r e in f o r c e d by
68
th e en v iro n m e n ta l c o r r e l a t i o n ( r e= -.4 2 ) found betw een th e s e two t r a i t s
w hich i n d i c a t e s t h a t th e environm en t c o n d u c tiv e t o a h ig h E I te n d s to
have a n e g a tiv e e f f e c t on A.
P h e n o ty p ic c o r r e l a t i o n s
betw een grow th s t a t i s t i c s and p r o d u c tiv e
t r a i t s te n d ed t o have low n e g a tiv e and p o s i t i v e v a lu e s ,
.2 4 .
from - .1 4 t o
The lo w e st v a lu e c o rre sp o n d e d t o th e c o r r e l a t i o n betw een ATFP and
HGRS1 and th e h ig h e s t t o
th e
c o rre la tio n
AMflS w ith
ATLW.
low
p o s itiv e
p h e n o ty p ic
c o rre la tio n s
V ery
betw een AGfl,
betw een AMRS w ith ATWW and
v a lu e s w ere o b se rv e d f o r
HGR a n d
w eaning-12 mo w ith a l l th e p r o d u c tiv e t r a i t s .
AMR i n
th e
th e
in te rv a l
S to b a r t (1983) r e p o r te d
e s tim a te s w ith s im ila r p a t t e r n s , w ith d i f f e r e n c e s m ain ly in v o lv in g th e
AMR r e l a t i o n s h i p s .
P h e n o ty p ic a l l y ,
th e r e l a t i o n s h i p betw een grow th s t a t i s t i c s b e fo re
18 mo o f age and th e p ro d u c tiv e
in v o lv in g AMR te n d ed t o
t r a i t s had low v a lu e s .
be s l i g h t l y
h ig h e r,
b u t AGRS a p p e a rs t o hav e
g r e a te r p r a c t i c a l a p p l i c a t i o n f o r e a r ly
s e le c tio n
p h e n o ty p ic
p ro d u c tio n .
p o te n tia l
fo r
su b seq u en t
C o r r e la tio n s
o f a n im a ls w ith
H ow ever,
a
th e
p h e n o ty p ic c o r r e l a t i o n o f AGflS w ith E I i n d i c a t e s t h a t a n im a ls s e le c te d
on t h e
b a s is
of
th a t
c rite ria
w ould n o t
be n e c e s s a r ily
e f f i c i e n t and a l s o would te n d to be o f l a r g e r m atu re s i z e s .
th e m ost
The most
p r a c t i c a l grow th s t a t i s t i c i n p h e n o ty p ic te rm s a p p e a rs t o be AGR2 which
h a s no n e g a tiv e c o r r e l a t i o n s w ith th e p ro d u c tiv e
t r a i t s and a ls o i t s
c o r r e l a t i o n w ith EI i s p o s i t i v e ( . 17) .
S e le c tio n ln d e y e s
S e le c tio n in d e x e s w ere d e riv e d u t i l i z i n g th e grow th p a ra m e te rs A
and k o r
th e
g ro w th
s ta tis tic s
th a t
show ed
th e
h ig h e s t
g e n e tic
69
c o r r e l a t i o n s w ith e i t h e r ATtfW o r E l.
in c lu d e d .
In some in d e x e s , ATFP was a ls o
The among s i r e s and w i th in s i r e s v a ria n c e and c o v a ria n c e
com ponents fro m w h ic h
th e
g e n e tic
and
c o v a ria n c e s u sed i n t h e
developm ent o f
p h e n o ty p ic
v a ria n c e s
and
th e in d e x e s a r e shown i n th e
Appendix T a b le s 50 and 51.
T a b le 20
p re s e n ts
th e
22
in d e x e s ,
th e ir
re s p e c tiv e
s ta n d a r d
d e v ia tio n and e x p e c te d re s p o n s e s to s e l e c t i o n r e l a t i v e t o s e l e c t i o n f o r
th e s i n g l e t r a i t s ATWW o r E l.
in c re a s e d
The i n c l u s i o n o f A and k , i n in d e x tw o,
th e a c c u ra c y o f s e l e c t i o n f o r
ATWW; 1745.
From th e
o th e r
in d e x e s , f o r ATWW t h a t in c lu d e d A a n d (o r) k , i t was o b s e rv e d t h a t th e
m ost i n f l u e n t i a l p o s i t i v e e f f e c t was due t o A.
The in f lu e n c e o f ATFP
when in c lu d e d gave more a d v an ta g e th a n th e i n c l u s i o n o f k. When AGR2
and RGR2 w ere in c o r p o r a te d ,
in d e x n i n e ,
a v e ry
s ig n ific a n t,
380%
re sp o n se was o b s e r v e d . compared t o th e use o f m ass s e l e c t i o n f o r ATWW
a lo n e .
The m ost i n f l u e n t i a l e f f e c t i n t h a t re s p o n s e was due t o RGR2.
The i n c l u s i o n
of
A and k i n
th e
li m i t e d a d v a n ta g e , o n ly 21% f o r in d e x 13.
in d e x e s d e riv e d f o r
BI gave
Thus, th e r e a p p e a rs to be no
p r a c t i c a l use f o r A and ( o r ) k a s com ponents f o r an in d e x o f s e l e c t i o n
fo r
e ffic ie n c y .
The e f f e c t s
o f AGR2 and RGR2 o n th e
s e l e c t i o n f o r EI was more b e n e f i c i a l th a n A and k .
a cc u ra cy
of
In d e x 20 shows an
ad v an tag e o f 117%, w hich r e p r e s e n t s a s u b s t a n t i a l in c r e a s e i n e x p e c te d
re sp o n se o f E l.
When ATFP was in c lu d e d i n th e in d e x e s f o r ATWW and E l, t h e r e was a
v e ry s l i g h t p o s i t i v e e f f e c t .
e ffe c ts
of
a n ta g o n is tic .
ATFP on
T h is f i n d i n g shows t h a t th e i n f l u e n t i a l
re p ro d u c tiv e
p e rfo rm an c e
a re
at
le a s t
not
70
TABLE 20.
DERIVED SELECTION INDEXES FOR THE ANALYSIS OF THE EFFECT OF
A, k, AGR2 AND RGR2 CN THE IMPROVEMENT OF AVERAGE TOTAL OF WEIGHT OF
LAMBS WEANED (ATWW) AND EFFICIENCY INDEX (E I) BY SELECTION
N
T ra it
-■ '
In d ex
'
In d e x a
S in g le b r e l a t i v e 0
s ta n d a rd t r a i t
re s p o n se
d e v ia t io n re s p o n se
ATWW
1
2
3
4
5
6
7
8
9
10
11
62.56
6 2 .3 2
5 9 .1 2
40.12
4 3 .8 0
111.48
101.24
108.72
109.16
100.88
107.80
. I 29EI+. 0 I 4ATFP+.0006A+.764k
.131E I+ .001 A+.785k
.147E I-.0005A
.123E I+ .670k
.157EI+.013ATFP
.131EI+.019ATFP-.196AGR2+15.13RGR2
.1 5 1 EI+.005ATFP+.079AGR2
.144EI+.008ATFP+4.99RGR2
. 12 9 E I-. 175AGR2+14 . 11RGR2
.150EI+.080AGR2
.142EI+5.02RGR2
1.02
1.00
.8 5
.9 8
.8 8
1.8 2
1 .4 8
1.66
1 .7 9
1 .4 8
1.6 6
S
EI
12
13
14
15
16
17
18
19
20
21
22
2 2 .7 2
. I 02ATWW+.292ATFP+.278A+30. 0 Ik
•105ATWW+.286A+30.1 5k
.113ATWW+.222A
.107ATWW-.3.22k
.0 9 3 ATWW+I .09ATFP
.076ATWW+I .76ATFP-6.33AGR2+676RGR2
.088ATWW+.512TFP+6.03AGR2
.0 85ATWW+.8 3 1ATFP+350 RGR2 '
. O96 ATWW-4 .4 3 AGR2+5 82 RGR2
.093ATWW+6.09AGR2
.094ATWW+352RGR2
fln d e x s ta n d a rd d e v ia t io n : a T
0T r a l t r e s p o n s e : i h 2a
1
cR e la tiv e re s p o n se o f Pln d e x u s e : i c r j / l h 2(jp (F a lc o n e r,
1981)
2 .7 5
2 .7 4
2 .6 0
1 .76
1.93
4 .9 0
4 .4 5
4 .7 5
4.8 0
4 .4 4
4.71
1.23
1.21
1.0 3
1 .1 9
1.06
2 .2 0
1 .7 9
2 .01
2 .1 7
1 .7 9
2.01
71
A d d itio n a l in d e x e s w ere e s tim a te d f o r th e s im u lta n e o u s im provem ent
o f ATWW, ATFP and E l. These in d e x e s a r e p re s e n te d i n T ab le 2 1 .
s e t o f in d e x e s
(I
to
8,
except 4 ) ,
AGR2 was fo u n d t o
In th is
h av e a h ig h
p o s i t i v e in f lu e n c e on th e s im u lta n e o u s im provem ent o f th e th r e e t r a i t s .
AGR2 e x p l a i n e d
a p p ro x im a te ly 60# o f
g e n o ty p es t h a t in c lu d e d
th e s e
tra its .
th e v a r ia n c e
of
I n c lu s io n
AGR2, w ith
of
th e a g g re g a te
RGR2,
r e s u l t e d i n an a d d i t i o n a l 10# i n a c c u ra c y .
The p o o re s t e f f i c i e n c i e s
w e re
o b serv ed
fo r
im provem ent o f ATFP and E I (in d e x e s 4 and 9 ) .
alo n e
(in d e x
9) o r
combined w ith
e f f i c i e n c i e s (R ^=.18 ) .
had a v e ry
th e
s im u lta n e o u s
In th e s e
SGR2 (in d e x
4)
gave
cases,
th e
AGR2
h ig h e s t
BGR2 when c o n s id e re d a lo n e (in d e x e s 10 to 13)
low o r a lm o st
n u ll
improvem ent o f ATWW, ATFP a n d
e ffe c t
EI
or
(from R2s e OOS to
th e ir
.0 1 ) on th e
c o m b in a tio n
in
p a irs .
I n c lu s io n o f ATFP a s component i n two o f th e in d e x e s (5 and 18) f o r th e
improvement o f ATWW and E I r e s u l t e d i n a n I n c r e a s e o f alm o st 5# i n R2
when a s s o c ia te d w ith
AGR2 and RGR2,
and 20# when ATFP was in v o lv e d
a lo n g w ith A and k .
D epending on th e o b j e c t i v e s o f s e l e c t i o n , In d e x e s f o r im p ro v in g a
s in g le
t r a i t o r th e c o m b in a tio n o f s im u lta n e o u s t r a i t s w ould r e q u i r e
c e r t a i n s p e c i f i c grow th t r a i t s f o r m axim izing th e e x p e c te d re s p o n s e to
s e le c tio n .
I n th e case o f s in g le t r a i t in d e x e s , RGR2 p ro v ed t o be th e
most u s e f u l ; how ever, when
more th a n one t r a i t was th e o b j e c t i v e o f
im provem ent AGR2 was th e m ost a d v a n ta g e o u s.
I n g e n e r a l,
s e ts
of
in d e x e s .
th e
c o n tr ib u tio n o f A and k w as s m a ll i n th e s e
T h e ir l a t e
a v a ila b ility
fo r
s e le c tio n
two
p u rp o s e s
72
N
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
T ra its
to be
Improved
ATWW,
ATWWt
ATWW,
ATFPt
ATWW,
ATWW,
ATWW,
ATWW,
ATFP,
ATWW,
ATWW,
ATWW,
ATFP,
ATWW,
ATWW,
ATWW,
ATFP,
ATWW,
In d e x
ATFPt
ATFP
EI
EI
E Ib
ATFPt
ATFP
EI
EI
ATFP,
ATFP
EI
EI
ATFP,
ATFP
EI
EI
E Ib
EI
EI
EI
EI
In d e x
v a ria n c e
A g g reg ated
genotype
V a ria n c e a
-4.08AGR2+591RGR2
11914
-3.89AGR2+576RGR2
11578
-4.32AGR2+587BGR2
10936
• 044AGR2+19.6RGR2
3 8 .8 3
- 6 .1 5AGR2+677BGR2+1. 6 5ATFP 11677
6 . 60AGR2
10123
6.52AGR2
9878
6.29AGR2
9172
•398AGR2
3 6 .8 7
7.338082
225.1
7.23RGR2
219.1
7.00RGR2
2 0 5 .7
• 422RGR2
.7459
.287A+36.9k
2271
.287A +36.0k
2273
.292A+40.3k
2356
-.0 0 5 A -2 .4 k
.571
.246A+37.5k+1.67ATFP
3103
16622
16216
15480
209.1
15480
16622
16216
15480
209.1
16622
16216
15480
209.1
16622
16216
15480
209.1
15480
A ccuracy
R^a
.72
.71
.71
.1 8
.75
.61
.61
.5 9
.1 8
.01
.01
.01
.003
.14
.14
.15
.01
.20
aMethod o f e s tim a tio n a s p r e s e n te d by F a lc o n e r (1981)
to 1U ts s
r
^
i s
1?lyT u r : s ^ i ^ r . r 9 ? . ln3tead o f Airp due
73
f u r th e r lim it th e ir use in programs of s e le c tio n as p r e d i c t o r s
of
g en etic p o te n tia l for productive e ffic ie n c y .
C onceptual a n a ly s is o f r e s u l t s .
From th e g e n e tic a n a l y s i s o f a l l
seme s p e c u la tio n
co u ld
be made i n
th e in f o r m a tio n i n t h i s
te rm s o f
th e
p o s s ib le
s tu d y ,
b io lo g ic a l
b a s is o f th e r e l a t i o n s h i p foun d betw een grow th i n th e i n t e r v a l w eaning
to 12 mo and r e p r o d u c tiv e e f f i c i e n c y .
The p a r t i c u l a r
re sp o n se
fa v o r in g a s p e c i f i c
co u ld
g e n o ty p e .
be a
s e le c tiv e
The i n t e r v a l ,
e ffe c t
of
w eaning t o
sea so n
12 mo,
Im p o rtan t from an a d a p tiv e p o in t o f view f o r th e p o p u la tio n .
w eaning,
th e a n im a ls
b e g in t o
i n t e r v a l when p u b e rty o c c u rs .
depend on th e m s e lv e s
an d
it
is
A fte r
is
th e
As th e a n im a ls a r e r a i s e d on t h e ra n g e ,
grow th from w eaning t o 12 mo m o s tly o c c u rs u n d er th e h a rs h e f f e c t s o f
w in te r , c a p a c ity f o r e f f i c i e n t w in te r grow th sh o u ld be a v e ry im p o rta n t
c h a ra c te ris tic .
An a n alo g y betw een grow th p erform ance from w eaning t o
su b se q u e n t a n n u al r e p r o d u c tiv e e f f i c i e n c y co u ld be made.
of
g e s ta tio n
o ccu rs
in
w in te r.
12 mo and
Almost h a l f
T h is p re s u p p o se s t h a t
a s n a tu ra l
s e l e c t i o n h a s alw ays s e l e c t e d th e p o p u la tio n to w a rd s th e i d e a l f i t n e s s
fo r
t h i s p a r t i c u l a r environm ent and t h a t r e p r o d u c tiv e p erfo rm an ce i s
th e m ain t r a i t a f f e c t e d by n a tu r a l s e l e c t i o n .
e ffe c t
of
n a tu ra l
a lw a y s
re p e a te d
s e le c tio n
I t i s p o s s ib le t h a t th e
on ach iev em en t o f
e n v ir o n m e n ta l
event
(w in te r),
g e s ta tio n
has
c o r r e la te d re s p o n se on grow th s p e c i f i c a l l y from w eaning
(so u rc e o f g e n e tic c o r r e l a t i o n ) .
d u rin g an
d e v e lo p e d
to 12
a
mo o f age
74
W inter e n v iro n m e n ta l c o n d itio n s I n Montana a r e h a r s h .
S u c c e s s fu l
grow th and r e p r o d u c tiv e perform an ce c o u ld a ls o be c o n s id e re d a c a p a c ity
f o r overcom ing s t r e s s .
S t r e s s and a d a p t a b i l i t y a r e two o p p o s ite te rm s
t h a t e v a lu a te th e same th in g .
a
c o n ju g a t e
of
two g e n e tic
I t a p p e a rs t h a t l i f e t i m e p erfo rm an ce i s
com ponents; one r e l a t e d
to
th e
g e n e tic
p o t e n t i a l c a p a c ity f o r p ro d u c tio n and th e o th e r r e l a t e d t o th e g e n e tic
c a p a c ity
fo r
e n v iro n m e n ta l
seasonal
overcom ing th e
ch an g es.
o n s e t.
p e rio d ic a l
s tre s s
p ro duced by s e a s o n a l
The r e p r o d u c tiv e r e q u i r e m e n t s
In
th is
s itu a tio n ,
th e
a ls o
s u p e rio rity
of
have
a
tw in n in g
a b i l i t y , i n te rm s o f l i f e t i m e e f f i c i e n c y , co u ld be a s s o c i a t e d w ith th e
c a p a b i l i t y f o r overcom ing s t r e s s .
P ric e (1985) s t a t e s t h a t th e r e e x i s t s a n e v o lu tio n a r y , a s w e ll a s
o n to g e n ic ,
d e te rm in a n t i n
th e re s p o n s e t o
s tre s s .
In
e v o lu tio n a r y
te rm s, n a tu r a l s e l e c t i o n w ould te n d t o re d u c e th e s t r e s s f u l e f f e c t s o f
c e rta in
c o n d itio n
g e n e r a tio n .
th a t
a ffe c ts
a
g iv e n p o p u la tio n
g e n e ra tio n
a fte r
In th e case o f sh eep u n d er ra n g e c o n d itio n s o f M ontana,
th e o n ly a l t e r n a t i v e f o r t h e i r s u r v i v a l was t o have g e s t a t i o n d u rin g
w in te r .
T h is c a p a b i l i t y
a lre a d y
e x is t e d i n
in tr o d u c e d t o U n ite d S t a t e s from E u ro p e.
needs
of
sheep
fo r
a l t e r n a t i v e s w ith o u t
c o n d itio n s .
s u rv iv a l,
it
s u b s ta n tia l
w ill
d o m e s tic a te d sh e e p when
I f we c o n s id e r th e b io lo g ic a l
be d i f f i c u l t
a lte ra tio n
of
th e
to
fin d
o th e r
e n v iro n m e n ta l
U n d e sira b le e f f e c t s o f h ig h te m p e ra tu re s d u rin g g e s t a t i o n
i n ewes ( T h w a ite s, 1985) a p p a r e n tly h a s an e v o lu tio n a r y e f f e c t .
P e rh a p s
th e
overcom ing s t r e s s
same g e n e ti c
c h a ra c te ris tic
a ls o e x i s t s i n
th e
e x is tin g
group o f s i n g l e s .
in
tw in s f o r
One way t o
V
o b se rv e i t would be th e e v a lu a tio n o f t h e i r r e l a t i v e grow th r a t e from
75
w eaning to 12 mo o f a g e .
Moberg (1985) s t a t e s t h a t I n each e v e n t i n
which s t r e s s f u l c o n d itio n s ta k e p la c e h o m e o sta s is i s com prom ised.
The
a c t i v e f u n c tio n o f th e c e n t r a l n erv o u s sy stem (CNS) d i c t a t e s w hich o f
th e p h y s io lo g ic a l sy ste m s m ust re sp o n d t o m a in ta in h o m e o s ta s is .
th e a b i l i t y
e ffic ie n t
o f c e r t a i n a n im a ls f o r
fu n c tio n in g
of
a fa st
CNS u n d er
Maybe
and a d e q u a te re s p o n s e
s tre s s
or
g iv e them a c u m u la tiv e
s u p e r i o r i t y i n t h e i r l i f e t i m e p ro d u c tiv e p e rfo rm a n c e s.
T h is ap p ro ach
g iv e s an a p p a re n t e x p la n a tio n t o th e a s s o c i a t i o n betw een
th e "in d e x o f
C e p h a liz a tio n » and lo n g e v ity s u g g e s te d by Com fort (1 9 6 1 ).
G e n e tic a d a p t a b i l i t y r e l i e s i n l a r g e p a r t on g e n e ti c e f f e c t s o th e r
th a n a d d i t i v e .
Dobzhansky (1 9 5 1 )
in d ic a te d
t h a t h e te r o z y g o te s have
a d a p tiv e s u p e r i o r i t y o v e r hom ozygotes, which i s th e e s s e n t i a l c o n d itio n
f o r th e e s ta b lis h m e n t o f b a la n c e d polym orphism s.
r e f e r s t o th e h ig h e r p l a s t i c i t y o f h e te r o z y g o te s .
A lso , L e rn e r ( I 95U)
I h i s means t h a t i f
th e r e e x i s t s a d d itiv e g e n e tic v a r i a b i l i t y f o r d e a lin g w ith s t r e s s ,
would be a t a v e ry low l e v e l .
th e
b reed h a v in g
th e
h ig h e s t
The low a d d i t i v e v a r i a b i l i t y
p e rfo rm a n c e s
a p p a re n t s u p p o rt to t h i s s p e c u la tio n .
(T a rg h e e s)
it
found i n
w o u ld
g iv e
76
SUMMARY AND CONCLUSIONS
Under th e ra n g e
e ffic ie n t
ewes.
in
c o n d itio n s o f M ontana,
life tim e
p ro d u c tio n
(E I)
T arghee e v e s w ere m o re
th a n R a m b o u ille t and Colum bia
Colum bias w ere th e p o o re s t i n p ro d u c tiv e e f f i c i e n c y , b u t had th e
h ig h e s t g re a se wool p ro d u c tio n .
Ewes re m a in in g i n th e f l o c k lo n g e r (6
y r ) showed th e h ig h e s t a v e ra g e y e a r ly p e rfo rm an c es f o r a l l
tra its .
p r o d u c tiv e
A p p a re n tly , t h i s ad v an tag e was due to s e l e c t i o n .
Ages a t m a tu r ity w ere a p p ro x im a te ly 39 mo, 38 mo and 41 mo f o r
R a m b o u ille ts,
th e
T arghees and C olum bias, r e s p e c t i v e l y .
s m a lle s t m ature
M a tu re
s iz e s
w e re
s iz e
and C olum bias th e
7 0 .0 4 ,
7 2 .4 2
Targhee and Columbia b re e d s ,
m a tu rin g r a t e s
and 73.11
re s p e c tiv e ly .
(k ) fo llo w e d by R a m b o u ille ts.
R a m b o u ille ts had
s lo w e s t m a t u r i n g
kg f o r
ra te .
th e R a m b o u ille t,
T arg b ees had
th e f a s t e r
Colum bia ewes w ere th e
s lo w e s t grow ing group.
Ewes re m a in in g i n th e f l o c k , u n t i l th ey had 4 and 5 r e p r o d u c tiv e
re c o rd s,
w e re
s ig n ific a n t.
s m a lle r
at
M atu rin g r a t e
s i g n i f i c a n t l y s lo w e r.
m a tu rity ,
but
d iffe re n c e s
w e re
not
(k ) i n th e f i v e - r e c o r d group o f ewes was
Phe no t y p i c a l l y ,
th e s lo w e r m a tu rin g r a t e
c o u ld '
be e x p la in e d by th e h ig h e r r a t e o f p ro d u c tio n i n th e f i v e - r e c o r d g ro u p .
There w ere
b re e d
d iffe re n c e s
in
g ro w th
ra te s
d u rin g
a ll
th e
i n t e r v a l s p re v io u s t o 18 mo o f age ( b ir th - w e a n in g , w eaning -12 mo, 12-18
mo, b i r t h -12 mo and b ir t h - 1 8 m o).
th e
in te rv a l
w e a n in g -12
R a m b o u ille ts h ad f a s t e r grow th i n
mo i n t e r v a l ,
e x c e lle d i n th e 12 t o 18 mo i n t e r v a l .
but
T a rg h e e s and C olum bias
77
H e rita b ility
e s tim a te s from
th e
p o o led d a ta
s e t w ere .4 3 ,
.3 3 ,
. 11 , . 68 , .4 2 , .56 and .15 f o r th e av erag e number o f lam bs b o rn , number
o f lam bs w eaned,
w e ig h t o f lamb w eaned,
m a tu rin g r a t e and e f f i c i e n c y in d e x .
grow th s t a t i s t i c s
f le e c e w e ig h t,
m atu re s i z e ,
The h ig h e s t h e r ! t a b i l i t i e s f o r th e
(AGR, RGR and AMR) were o b se rv e d i n w eaning t o 12 mo
in te rv a l.
G e n e tic c o r r e l a t i o n s betw een p ro d u c tiv e t r a i t s and B rody’ s grow th
p a ra m e te rs A and k d e riv e d from th e p o oled d a ta s e t i n d i c a t e d t h a t ewes
w ith h ig h e r a d d i t i v e g e n e ti c p o t e n t i a l f o r weaned lam b p r o d u c tio n
w ill
a ls o have g e n e tic p o t e n t i a l f o r l a r g e r m atu re
ewes
s iz e s .
However,
s e le c te d f o r h ig h a d d i t i v e g e n e ti c p ro d u c tio n p o t e n t i a l f o r lam bs a t
w eaning would show no c o r r e l a t e d e f f e c t f o r m a tu rin g r a t e .
e f f i c i e n c y ( E l ) , th e o p p o s ite o c c u r s .
l a r g e r m atu re s i z e s
I n te rm s o f
Ewes w ith g e n e ti c p o t e n t i a l f o r
(A) te n d e d t o h av e lo w e r e f f i c i e n c y ,
but f a s te r
m a tu rin g r a t e s ( k ) .
G e n e tic c o r r e l a t i o n s o f ATFP w ith A and k w ere low i n d i c a t i n g t h a t
in c r e a s e d m ature
ra te
(k ) w i l l
w e a n in g
to
s i z e w i l l n o t a f f e c t ATFP and t h a t f a s t e r m a tu rin g
te n d t o
th e
w e ll a s r e p r o d u c tiv e
to
red u ce ATFP.
12 mo o f a g e i n t e r v a l
r e l a t i o n s h i p s w ith
w e a n in g
s lig h tly
su b se q u e n t
e ffic ie n c y .
12 mo i n t e r v a l ,
show ed
Growth s t a t i s t i c s
th e
h ig h e s t
p r o d u c tiv e t r a i t s
a p p e a rs to
th e
g e n e tic
ew es,
as
in
th e
be th e m ost e f f i c i e n t
and
The RGR grow th
in
fo r
s ta tis tic ,
p r a c t i c a l f o r s e l e c t i v e p u rp o s e s .
In d e x e s o f s e l e c t i o n f o r ATWW w hich in c lu d e RGR2 g av e th e h ig h e s t
r e l a t i v e re s p o n se to
s e le c tio n
compared t o m ass s e l e c t i o n f o r
ATWW.
The adv an tag e was 380% g r e a t e r s e l e c t i o n re s p o n s e p e r g e n e r a tio n .
When
78
EI was th e
tra it
of in te r e s t ,
grow th
p a ra m e te rs A and k showed no
p r a c t i c a l adv an tag e i n th e in d e x e s , however AGR2 and RGR2, e s p e c i a l l y
BGR2,
produced a
re sp o n se o f 117%.
fo r e ffic ie n c y .
s i g n i f i c a n t im provem ent o n th e
e x p e c te d
s e le c tio n
T h is i n d i c a t e s a p o s s ib le p r a c t i c a l use i n s e l e c t i o n
ATFP showed a sm a ll p o s i t i v e e f f e c t when in c lu d e d i n
th e in d e x e s f o r im p ro v in g t h e a c c u ra c y f o r s e l e c t i o n f o r e i t h e r ATW o r
El.
Twins r a i s e d a s tw in s te n d e d t o grow slo w e r up t o w eaning th a n d id
s in g le s r a i s e d a s s i n g l e s and tw in r a i s e d a s s i n g l e s ,
fa s te r
in
th e w eaning t o
12 mo i n t e r v a l .
b u t th e y grew
Twins h ad s m a lle r m atu re
s i z e s th an s in g le s and slo w e r m a tu rin g r a t e s .
Ewes b o rn tw in s produced
2.2 kg more lamb a t w eaning each y e a r , on th e a v e ra g e , th a n ewes b o rn
s in g le s ,
and th e y had
th e h i g h e s t
p ro d u c tiv e
e ffic ie n c y
in d e x e s .
However, th e y produced .1 kg l e s s g re a s e f l e e c e w e ig h t y e a r ly th a n ewes
born s i n g l e s .
The g e n e tic r e l a t i o n s h i p s betw een grow th d u rin g t h e i n t e r v a l from
w eaning t o 12 mo and th e p ro d u c tiv e t r a i t s and e f f i c i e n c y s u g g e s t t h a t
p ro d u c tiv e
life tim e
e ffic ie n c y
co u ld
be im proved by s e l e c t i n g f o r
grow th d u rin g t h a t p e rio d i n th e a n im a l's l i f e .
Some c o n c e p tu a l id e a s w ere p u t f o r t h r e g a r d in g p o s s i b l e b io l o g i c a l
b a s i s , i n a d d i t i o n t o lin k a g e and p le io tr o p h y w hich may c o n tr ib u te t o
t h e g e n e tic c o r r e l a t i o n s o b ta in e d betw een grow th i n t h e w eaning t o 12
mo i n t e r v a l
and l i f e t i m e
p r o d u c tiv e
e ffic ie n c y .
T h is in v o lv e d
th e
e f f e c t o f s e a s o n a l s t r e s s d u rin g th e w in te r p e r io d f o llo w in g w eaning t o
w hich each group o f young ewes w ere s u b je c te d .
I t i s in
t h i s p e r io d
when th e young ewes b e g in t o depend on t h e i r own a b i l i t y to fo ra g e f o r
79
food and i t i s th e p e rio d when p u b e rty n o rm ally o c c u r s .
Thus, i t i s a n
im p o rta n t
to a d a p t t o
dev elo p m en tal
b io lo g ic a lly
s tre s s fu l
p e rio d ,
s itu a tio n .
r e q u i r i n g th e a b i l i t y
The c a p a c ity
to
grow u n d er
c o n d itio n s was c o n s id e re d a good m easure o f a d a p t a b i l i t y .
ewes may undergo a n even g r e a t e r s t r e s s th a n s i n g l e s .
a
th e s e
Twin born
The g e n o ty p es
a s s o c ia te d w ith t h i s a d a p tiv e a b i l i t y e x p re s s e d a s grow th re s p o n s e , may
a ls o be th e g e n o ty p es t h a t a f f e c t r e p r o d u c tiv e f i t n e s s .
The a n im a ls
t h a t a r e a b le to re sp o n d f a v o r a b ly to th e e a r ly s t r e s s o f th e w eaning
t o 12 mo p e rio d a p p a r e n tly a r e a ls o a b le to p ro d u ce w e ll l a t e r i n l i f e .
I
80
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APPENDIX
TABLE 22. AVERAGES, VARIATION AND RANGES OF LIFETIhE AVERAGE OF TOTAL
NUMBER OF LAMBS BORN (ATLB) FOR EWE GROUPS WITHIN BREEDS
B reed
Number o f
re c o rd s
R a n b o u ille t
Targhee
Columbia
N
Average
S ta n d a rd C oef. o f
d e v ia t io n v a r i a t i o n
3
4
5
74
64
164
1.28
1 .3 9
1.46
.284
.299
3
4
5
74
47
217
1.38
.388
1.34
1.52
.356
.312
3
4
5
54
1.23
21
100
1.20
.398
.444
1.47
.302
.366
Low est
H ig h e st
2 8 .6 5
2 0 .4 0
2 0 .4 8
.33
.75
.60
2.00
2.00
2.20
2 8 .1 2
26.43
2 0 .5 0
.67
.25
.80
2.00
2.20
3 2 .3 9
3 6 .9 7
2 0 .4 8
.67
.50
.80
2 .2 5
2 .4 0
2 .3 3
2.00
TABLE 23. AVERAGES, VARIATION AND RANGES OF LIFETIME AVERAGE OF THE
TOTAL NUMBER OF LAMBS WEANED (ATLW) FOR EWE GROUPS WITHIN BREEDS
B reed
Number o f
re c o rd s
R am b o u llle t
T argbee
Columbia
N
Average
3
4
5
74
64
164
1.01
.407
1.16
1.25
.322
.300
40.14
2 7 .7 8
2 4 .0 0
3
4
5
74
47
217
1.12
1 .1 9
1.33
.350
.332
.312
3 1 .3 2
27.96
2 3 .4 5
.60
2.00
2.00
2.00
3
4
5
21
100
.90
.93
1.27
.409
.364
.271
. 4 5 .6 7
3 9 .1 5
21.24
.00
2.00
.50
.80
1 .7 5
54
S ta n d a rd C oef. o f
d e v ia tio n v a r i a t i o n
Low est
H ig h e s t
.00
2.00
.50
.40
2.00
.33
.25
1 .7 5
1.80
86
TABLE 24.
AVERAGES, VARIATION AND RANGES OF LIFETIME AVERAGE OF THE
TOTAL OF KILOGRAMS OF LAMBS WEANED (ATWW) FOR EWE GROUPS WITHIN BREEDS
B reed
Number of
re c o rd s
R am b o u ille t
T arghee
Columbia
N
Average
S ta n d a rd C oef. o f
d e v ia t io n v a r i a t i o n
Low est
H ig h e st
7 1.40
6 8 .1 8
8 0 .4 8
3
4
5
74
64
164
3 8 .4 0
44.07
47.90
14.84
11.82
3 8 .7 4
.00
26.82
11.20
2 3 .3 9
1 6.25
17.54
3
4
5
74
47
217
45.44
4 8 .3 2
54.17
12.87
13.37
11.90
2 8 .3 3
2 7 .6 8
2 1 .9 8
3
4
5
54
21
100
3 6 .6 5
3 7 .5 8
50.72
16.63
13.21
11.05
4 5 .3 7
3 5 .1 4
2 1 .7 9
14.83
9.10
2 4.82
.00
1 8 .0 8
26.62
7 9 .4 0
6 8 .9 8
8 4 .1 2
7 8 .1 7
7 0 .3 5
7 2 .9 2
TABLE 25.
AVERAGES, VARIATION AND RANGES ' OF LIFETIhE AVERAGE OF
KILOGRAMS OF GREASE FLEECE PRODUCED (ATFP) FOR EWE GROUPS WITHIN BREEDS
B reed
Number o f
R am b o u ille t
Targhee
Columbia
N
A verage
S ta n d a rd
C oef. o f
Low est
H ig h e s t
11.17
11.43
10.64
3 .3 2
3 .2 0
5 .5 8
5 .9 8
6 .4 0
3
4
5
64
164
4 .3 2 ,
4 .5 2
4 .5 4
.4 8
.52
.4 8
3
4
5
74
47
217
4 .6 5
4 .8 2
4 .8 6
.56
.61,
.55
12.73
1 1 .2 8
3 .4 5
3 .3 2
3.51
3
4
5
54
21
100
4 .8 2
4 .9 7
5 .1 9
.6 4
.85
.60
13.35
17.09
11.55
2 .8 4
3 .7 0
•
12.12
3.62
3.00
5 .8 0
6.10
6 .3 8
6 .0 8
6 .2 4
6 .5 5
87
TABLE 26. AVERAGES, VARIATION AND RANGES OF MATURE SIZE (A) FOR EWE
GROUPS WITHIN BREEDS
B reed
Number o f
re c o rd s
R am b o u llle t
Targhee
Columbia
N
Average
S ta n d a rd C oef. o f
d e v ia tio n v a r i a t i o n
Low est
H ig h e st
53.62
84.81
8 9 .3 5
8 5.79
3
4
5
74
64
164
6 9 .5 5
7 1 .3 6
7 1.5 6
6 .6 4
7.1
5 .4 9
9 .5 5
9.9 9
7 .6 7
56.88
3
4
5
74
47
217
7 2 .9 5
7 2 .9 6
7 4 .2 5
8.3 5
7 .3 0
1 1 .4 5
56.11
10.01
6.78
9 .1 4
5 8 .8 5
5 8 .5 7
3
4
5
54
21
100
7 4 .5 3
7 5 .7 2
7 3 .5 6
9.7 0
5 .3 8
6.31
13.01
7 .1 0
8 .5 8
5 4 .2 4
6 4 .6 1
5 9 .1 2
58.15
96.84
9 4 .4 5
9 3.70
98.76
86.06
93.38
TABLE 27. AVERAGES, VARIATION AND RANGES OF BRODY’ S CONSTANT OF
INTEGRATION (B) FOR EWE GROUPS WITHIN BREEDS
B reed
tber o f
lords
R am b o u lllet
3
4
5
74
64
164
6 3 .3 6
6 3 .9 0
5.8 4
5.97
4 .8 4
3
4
5
74
47
217
6 5 .5 4
6 5 .4 5
66.16 .
7 .2 8
6 .7 4
6 .1 4
3
4
5
54
6 6 .4 5
21
100
67.20
8 .5 0
4.51
5 .5 5
Targhee
Columbia
N
Average
62.52
6 5.4 6
Low est
H ig h e st
9 .3 5
9 .4 2
7 .5 8
4 7 .9 2
5 1 .4 5
5 3 .2 2
7 6 .5 0
7 8 .2 9
7 7 .8 7
11.11
1 0 .2 9
9 .2 8
49.61
52.71
5 4 .6 4
85.45
86.63
12.81
6 .7 2
8 .4 8
51.17
5 6 .6 5
5 2 .5 8
86.85
7 4.80
82.15
S ta n d a rd C oef. o f
d e v ia tio n v a r i a t i o n
85.82
88
TABLE 28. AVERAGES, VARIATION AND RANGES OF MATURING RATE (K) FOR EWE
GROUPS WITHIN BREEDS
B reed
Number o f
re c o rd s
R am b o u llle t
Targhee
Columbia
N
Average
S ta n d a rd C oef. o f
d e v ia t io n v a r i a t i o n
Low est
H ig h e s t
.0740
.0627
.0637
.1889
.1842
.2057
.2428
.1979
.2314
3
4
5
74
64
164
.1160
.1104
.026
.030
22.36
.1072
.024
22.22
3
4
5
74
47
217
.1216
.1154
.035
.029
.027
2 9 .1 9
2 4 .9 0
2 4 .2 2
.0612
3
4
5
54
.1060
21
100
.0947
.1042
.039
.034
.034
36.56
3 6 .2 5
32.31
.0494
.0443
.0556
.1101
27.56
.0446
.0625
.2178
.1904
.3260
TABLE 29. AVERAGES, VARIATION AND RANGES OF EWE EFFICIENCY INDEX (E I)
FOR EWE GROUPS WITHIN BREEDS
B reed
Number o f
re c o rd s
R am b o u ille t
T arghee
Columbia
N
Average
S ta n d a rd C oef. o f
d e v ia tio n v a r i a t i o n
3
4
5
74
64
164
.557
.623
.673
.218
.174
.162
3 9 .2 0
2 7 .9 6
23.99
3
4
5
74
47
217
.635
.671
.731
.200
.198
3 1 .5 8
2 9 .5 2
.152
20.80
54
.498
.501
.693
.230
.185
.151
46.15
3 7 .0 0
3
4
6
21
100
21.80
Low est
H ig h e st
.0000
.2062
1.007
.986
1 .109
.1531
.1157
.3381
1 .013
1 .1 7 2
.0000
1 .027
.2089
.2370
.4010
1.216
.916
1.062
89
TABLE 30. AVERAGES, VARIATION AND RANGES OF ABSOLUTE GROWTH RATE IN
THE INTERVAL BIRTH-WEANING (AGRI) FOR EWE GROUPS WITHIN BREEDS
Breed
Number o f
re c o rd s
R am b o u ille t
Targhee
Columbia
N Average
3
4
5
74
64
164
3
4
5
3
4
5
21
100
7 .7 3 6
S ta n d a rd C oef. o f
d e v ia t io n v a r i a t i o n
Lowest
H ig h e st
7 .8 9 6
1.114
1 .2 4 9
1.165
14.41
1 5.49
14.75
5.080
5.580
5 .1 5 0
74
47
217
8 .54 8
8.351
8.483
1.271
1.107
1.226
14.87
5.620
13.16
5 .6 2 0
5 .5 6 0
12.250
11.225
11.750
54
8.461
8.183
8 .25 7
1.115
1.341
1.329
13.55
1 6 .3 9
16.09
6.280
6.060
10.900
8.062
14.45
5 .1 4 0
9.925
10.200
10.825
11.050
11.450
TABLE 3 1 .
AVERAGES, VARIATION AND RANGES OF ABSOLUTE GROWTH RATE IN
THE INTERVAL WEANING-12 MONTHS (AGR2) FOR EWE GROUPS WITHIN BREEDS
B reed
Number o f
re c o rd s
R am b o u ille t
T arghee
Columbia
N
Average
S ta n d a rd C oef. o f
d e v ia tio n v a r i a t i o n
Low est
H ig h e s t
3
4
5
74
64
164
1.437
1 .42 5
1.504
694
470
623
4 8 .3 3
3 2 .9 8
41.46
3
4
5
74
47
217
1 .19 9
1.193
1.090
514
610
42.92
5 1 .1 4
49.57
.200
-.1 2 2
-.3 1 1
3 .0 3 3
2 .5 6 3
3
4
5
54
.978
1.040
1.044
7 3 .6 2
51.26
6 5 .1 9
- .5 1 4
.144
- .4 4 3
2 .4 3 8
2 .0 1 3
3 .4 0 0
21
100
540
720
534
681
.244
.256
.000
3 .4 1 3
2 .7 1 3
3.817
2.600
90
TABLE 32.
AVERAGES, VARIATION AND RANGES OF ABSOLUTE GRWTH RATE IN
THE INTERVAL 12-18 MONTHS (AGR3) FOR EWE GROUPS WITHIN BREEDS
B reed
Number o f
re c o rd s
R m b o u llle t
T arghee
Colmnbia
N
Average
S ta n d a rd C oef. o f
d e v ia tio n v a r i a t i o n
Low est
H ig h e st
4 3 .7 5
. 5 2 .0 3
5 4 .1 3
- .0 4 0
.150
5 .6 3 3
5 .6 3 3
7 .0 5 0
.800
.967
6 .9 5 0
6 .8 2 5
.000
8.700
.450
.850
.6 8 3
8 .3 2 5
7 .1 7 5
7 .0 6 7
3
4
5
74
64
164
2.677
2.451
2 .9 1 8
1.171
3
4
5
74
47
217
3.326
3 .3 4 9
3 .5 6 5
1.613
1.577
1.722
4 8 .4 9
47.07
48.31
3
4
5
54
3 .510
3 .3 8 8
3.396
1.955
1.857
1.552
5 5 .7 0
54.82
4 5 .6 8
21
100
1.276
1.579
.250
TABLE 3 3 .
AVERAGES, VARIATION AND RANGES OF ABSOLUTE GRWTH RATE IN
THE HTERVAL BIRTH-12 MONTHS (AGR4) FOR EWE GROUPS WITHIN BREEDS
B reed
Number o f
re c o rd s
R a m b o u ille t
T arghee
Columbia
N
A verage
S ta n d a rd C oef. o f
d e v ia tio n v a r i a t i o n
Low est
H ig h e s t
3
4
5
74
64
164
3 .71 5
3 .7 4 4
3.664
468
475
481
12.61
2 .7 8 3
12.71
13.13
2.686
2.700
4 .9 4 5
• 4.727
5 .0 5 5
3
4
74
47
3 .7 6 5
3.707
605
575
16.08
15.51
2 .131
2 .4 3 8
5 .7 5 5
4 .9 8 2
5
217
3.667
493 .
13.45
2.562
4 .9 6 4
3
4
5
54
3 .53 0
3 .49 2
3 .54 8
,462
,732
,502
1 3 .0 9
2 0 .9 7
14.14
2 .6 6 9
4 .5 5 5
5 .0 6 4
4 .4 0 0
21
100
2.238
2 .4 5 6
91
TABLE 34.
AVERAGES, VARIATION AND RANGES OF ABSOLUTE GROWTH RATE IN
THE INTERVAL BIRTH-18 MONTHS (AGR5) FOR EWE GROUPS WITHIN BREEDS
B reed
Number o f
re c o rd s
R a m b o u ille t
Targhee
Columbia
N
Average
S ta n d a rd C oef. o f
d e v ia tio n v a r i a t i o n
3
4
5
74
64
164
3 .33 5
3 .2 6 0
3 .3 5 8
.343
.356
.343
3
4
5
74
47
217
3 .5 4 4
3 .5 1 7
3.544
.400
.430
.434
3
4
5
54
21
100
3 .4 3 7
3 .34 7
3 .4 1 3
.480
.401
.366
1 0 .2 9
10.92
10.22
Low est
H ig h e st
2.612
4 .076
4.106
4 .1 6 3
2 .5 5 3
2 .6 7 6
1 1.30
2.661
12.22
2 .7 4 4
2 .6 4 7
4 ,6 6 5
4 .6 5 3
4.913
2 .6 7 6
2 .8 3 5
2 .4 5 6
4 .8 8 2
4.294
4 .400
12.24
13.96
11.98
10.76
TABLE 35.
AVERAGES, VARIATION AND RANGES OF RELATIVE GROWTH RATE IN
THE INTERVAL
BIR TH-WEANING (RGRI) FOR EWE GROUPS WITHIN BREEDS
B reed
Number o f
re c o rd s
R a m b o u llle t
T arghee
Columbia
N
A verage
S ta n d a rd C oef. o f
d e v ia t io n v a r i a t i o n
3
4
5
74
64
164
.2122
.018
.2165
.022
.2202
.025
3
4
5
74
47
217
.2181
.2125
.2125
.011
3
4
5
54
21
100
.2223
.2156
.2140
Low est
H ig h e s t
.1722
.1685
.2805
.2908
.2847
8.31
1 0 .2 8
11.45
.1706
.017
.017
4 .9 2
8 .3 7
7 .9 5
.1726
.1722
.2274
.2272
.2317
.014
.014
.016
6 .1 3
6 .3 2
7 .4 0
.1730
.1736
.1739
.2272
.1751
.2828
.2312
92
TABLE 36.
AVERAGES, VARIATION AND RANGES OF RELATIVE GROWTH RATE IN
THE INTERVAL WEANING-12 MONTHS (RGR2) FOR EME GROUPS WITHIN BREEDS
B reed
Number o f
re c o rd s
R a m b o u llle t
N
Average
S ta n d a rd C oef. o f
d e v ia t io n v a r i a t i o n
3
4
5
74
64
164
.0302
.012
.0298
.0314
.009
T arghee
3
4
5
74
47
217
C oltm bla
3
4
5
54
21
100
Low est
H ig h e s t
.0051
.0059
.011
4 1.08
3 1 .3 2
32.04
.0000
.0602
.0491
.0716
.0247
.0242
.0223
.010
.011
.010
3 9 .8 2
4 6.53
46.91
.0045
-.0 0 3 3
—.0081
.0549
.0602
.0478
.0206
.0221
.015
-.0 1 2 9
.0042
.0217
.013
7 1 .7 7
46.27
5 9 .7 3
.0488
.0429
.0609
.010
-.0100
TABLE 37.
AVERAGES, VARIATION AND RANGES OF RELATIVE GROWTH RATE IN
THE INTERAL
12 -18 MONTHS (RGR3 )
FOR EWE GROUPS WITHIN BREEDS
B re e d s
Number of
re c o rd s
N
Average
S ta n d a rd C oef. o f
d e v ia tio n v a r i a t i o n
Low est
H ig h e st
R a m b o u ille t
3
4
5
74
' 64
164
.0433
.0399
.0461
.017
.019
.023
40.13
4 8 .5 7
4 9 .5 0
.0007
.0049
.0026
.0925
.0925
.0972
T arghee
3
4
5
74
47
217
.0501
.0507
.0533
.022
.021
.022
4 4 .1 6
4 1 .2 7
4 2 .0 0
.0137
.0151
.0000
.0919
.0939
.1259
3
4
5
54
21
100
.0534
.0539
.0530
.025
.0 2 8
.025
4 6.69
5 2 .5 2
4 1.57
.0074
.0129
.0125
.1117
.1144
.0989
Colvm bia
-
93
TABLE 38.
AVERAGES, VARIATION AND RANGES OF RELATIVE GROWTH RATE IN
THE INTERVAL BIRTH-12' MONTHS (RGR4) FOR BfE GROUPS WITHIN BREEDS
B reed
Number o f
re c o rd s
R am b o u llle t
T arghee
Columbia
3
N
Average
S ta n d a rd C oef. o f
d e v ia tio n v a r i a t i o n
Low est
H ig h e st
7 .4 0
.0677
.0625
.06 35
.0850
.0842
.0844
.0664
.0651
.0641
.0913
.0916
.0895
.0629
.0635
.0638
.0847
.0841
.0850
4
5
74
64
164
.0797
.0783
.0774
.0047
.0055
.0057
3
4
5
74
47
217
.0781
.0769
.0760
.0068
.0067
8 .2 9
8 .8 5
8 .7 9
3
4
5
54
21
100
.0770
.0755
.0758
.0060
.0073
.0064
7 .7 2
9 .6 2
8.51
.0065
5 .8 5
7.12
TABLE 39.
AVERAGES, VARIATION AND RANGES OF RELATIVE GROWTH RATE IN
THE INTERVAL BIRTH-18 MONTHS (RGR5) FOR EWE GROUPS WITHIN BREEDS
B reed
Number o f
re c o rd s
R am b o u llle t
Targhee
Columbia
3
N
Average
S ta n d a rd C oef. o f
d e v ia tio n v a r i a t i o n
Low est
H ig h e st
;
4
5
74
64
164
.0545 ■
.0541
.0543
.0015
.0015
.0015
2 .8 5
2 .7 4
2 .8 4
.0501
.0499
.0585
.0585
.0585
3
4
5
74
47
217
.0544
.0542
.0543
.0012
.0016
2.31
2 .9 7
2 .4 8
.0508
.0504
.0503
.0592
.0554
.0583
3
4
5
54
.0543
.0539
.0540
.0016
.0014
.0015
3.00
.0482
.0509
.0505
.0582
.0556
.0590
21
100
.0013
2.51
2.76
.0506
94
TABLE 40.
AVERAGES, VARIATION AND RANGES OF ABSOLUTE MATURING RATE IN
THE INTERVAL BIRTH-WEANING (AMRI) FOR EWE GROUPS WITHIN BREEDS
B reeds
Number o f
re c o rd s
R am b o u lllet
N
Average
S ta n d a rd C oef. o f
d e v ia tio n v a r i a t i o n
Low est
H ig h e st
.0826
3
4
5
74
64
164
.1116
.1133
.1105
.014
.016
.015
12.89
13.87
13.89
‘ .0780
.0720
.1402
.1509
.1428
T arghee
3
4
5
74
47
217
.1177
.1149
.1145
.016
.013
.015
13.37
11.67
1 3 .0 8
.0886
.0809
.0813
.1617
.1403
.1476
Columbia
3
4
5
54
21
100
.1151
.1085
.1128
.017
.019
.019
17.45
17.21
16.84
.0772
.0822
.0756
.1557
.1448
.1755
TABLE 41.
AVERAGES, VARIATION AND RANGES OF ABSOLUTE MATURING RATE IN
THE INTERVAL WEANING-12 MONTHS (AMR2) FOR EWE GROUPS WITHIN BREEDS
B reed
Number o f
re c o rd s
R am b o u llle t
3
4
5
Targfree
3
4
5
Columbia
3
4
5
N
A verage
74
64
164
.0208
.0203
.0213
74
47
217
.0166
.0166
54
21
100
.0149
.0134
.0140
.0143
S ta n d a rd C oef. o f
d e v ia t io n v a r i a t i o n
010
Low est
H ig h e st
.0035
.0041
.0503
.0395
.0519
007
009
4 8 .8 4
3 6 .5 4
42.71
007
009
008
4 4 .9 8
5 4 .0 9
5 2 .3 6
-.0 0 1 8
-.0 0 4 5
O il
008
7 9 .2 7
-.0080
76.62
.0017
010
6 6 .9 4
-.0058
.0000
.0022
.0410
.0436
.0408
.0408
.0283
•.0 4 9 7
95
TABLE 42.
AVERAGES, VARIATION AND RANGES OF ABSOLUTE MATURING RATE IN
THE INTERVAL 12-18 MONTHS (AMR3) FOR EHE GROUPS WITHIN BREEDS
B reed
Number o f
re c o rd s
R am b o u llle t
Targhee
Columbia
3
N
Average
4
5
74
64
164
.0383
.0342
.0405
3
4
5
74
47
217
.0456
.0455
.0475
3
4
5
54
.0466
.0442
.0459
21
100
S ta n d a rd C oef. o f
d e v ia tio n v a r i a t i o n
.016
Low est
.0 1 8
41.35
5 1 .4 4
—.0006
.0035
.021
52.61
.0022
.021
.020
.022
47.00
4 4 .1 6
4 6 .4 2
.0098
.0144
.024
.023
51.78
52.57
4 4 .2 8
.0063
.0116
.020
.0000
.0110
H ig h e st
.0814
.0789
.0950
.0892
.0896
.1285
.1002
.0848
.0987
TABLE 43.
AVERAGES, VARIATION AND RANGES OF ABSOLUTE MATURING RATE IN
THE INTERVAL BIRTH-12 MONTHS (AMR4) FOR EHE GROUPS WITHIN BREEDS
B reeds
Number o f
re c o rd s
R am b o u ilIet
T arghee
Columbia
3
N
Average
S ta n d a rd C oef. o f
d e v ia t io n v a r i a t i o n
Lowest
H ig h e st
.0732
4
5
74
64
164
.0537
.0529
.0515
.007
.008
.008
13.24
1 4 .3 0
14.74
.0394
.0375
.0369
3
4
5
74
47
217
.0520
.0512
.0497
.008
.009
.007
15.99
17.01
14.67
.0322
.0716
.0349
.0758
3
4
5
54
.0482
.0465
.0486
.0 0 9
.011
18.41
2 3 .4 4
15.87
.0320
.0667
.0689
.0687
21
100
.008
.0357
.0265
.0326
.0710
.0725
.0705
96
TABLE 44.
AVERAGES, VARIATION AND RANGES OF ABSOLUTE MATURING RATE IN
THE INTERVAL BIRTH-18 MONTHS (AMR5) FOR EHE GROUPS WITHIN BREEDS
B reeds
Number o f
re c o rd s
R a m b o u llle t
T arghee
:
Columbia
3
N
Average
S ta n d a rd C oef. o f
d e v ia tio n v a r i a t i o n
Lowest
H ig h e st
.0401
.0393
.0370
.0579
.0570
.0552
.0568
.0556
4
5
74
64
164
.0481
.0459
.0470
.004
.005
.004
10.12
8.38
3
4
5
74
47
217
.0480
.0483
.0478
.005
.004
.004
9.1 4
8 .7 4
.0362
.0326
.0362
.0581
3
4
5
54
21
100
.0465
.0443
.0465
.006
.005
.004
13.90
11.08
9 .1 9
.0339
.0336
.0359
.0603
.0559
.0554
7.81
8.70
97
TABLE 45.
PREDICTED BODY WEIGHTS AND PERCENT OF MATURE WEIGHT (A) FOR
EACH BREED AND EWE GROUP USING BRODY'S GROWTH EQUATION DERIVED IN THIS
STUDY IN SIX MONTH INTERVALS
Age
B irth
6 mo kg
*a
12 mo kg
%
18 mo kg
%
24 mo kg
%
30 mo kg
%
36 mo kg
%
42 mo kg
%
48 mo kg
%
54 mo kg
*
60 mo kg
%
66 mo kg
%
72 mo kg
%
78 mo kg
%
84 mo kg
%
a
B reeds
R am b o u ille t Targhee Columbia
7 .3 5
38.76
5 5.34
5 4.44
7 7 .7 2
62.25
7 .4 7
40.80
5 6 .3 4
57.03
7 8 .7 5
6 4 .9 3
88.88
89.66
6 6 .1 5
94.44
68.77
94.96
7 0 .6 4
97.54
7 1 .5 5
68.10
97.23
69.07
98.61
98.80
69.56
99.31
99.97
7 0.03
99.98
7 0.04
7 1 .9 9
99.41
72.21
99.71
7 2 .3 2
99.86
7 2 .37
99.93
7 2 .3 9
99.96
7 2 .4 0
99.97
72.41
9 9 .9 8
7 2 .4 2
100.00
100.00
69.80
99.66
6 9 .9 2
99.82
6 9 .9 8
99.91
70.01
99.96
70.02
7 .6 4
3 9 .3 7
53.85
5 5 .7 2
Grouos
3 re c o rd s 4 re c o rd s 5 re c o rd s
7.20
,
76.21
6 4 .1 5
87.7*
6 8 .4 9
93.68
7 0 .7 3
96.74
7 1 .8 8
98.31
7 2 .4 7
99.12
7 2 .7 8
99.54
7 2 .9 4
99.76
7 3 .0 2
99.87
7 3 .0 6
99.93
7 3 .0 8
99.95
7 3 .0 9
99.97
7 3 .1 0
99.98
,
40.00
5 6 .1 5
56.00
78.61
63.80
89.56
67.60
94.90
6 9 .4 6
97.51
7 0 .3 6
98.77
70.81
7 .6 3
3 9 .6 4
54.83
5 5 .8 0
7 7 .1 8 !
6 3 .9 6
88.47
6 8 .0 8
94.17
70.16
97.05 •
71.21
98.50
7 1 .7 4
99.23
99.41
7 1 .0 2
9 9 .7 0
7 1 .1 3
99.85
99.61
7 2 .1 5
99.80
71.18
72.22
99.92
7 1 .2 0
99.95
71.21
99.97
7 1 .2 2
9 9 .9 8
7 1 .2 3
99.90
7 2 .2 5
99.94
7 2 .2 7
99.97
100.00
9 9.98
72.01
72.28
9 9 .9 8
72.28
P e rc e n ta g e s r e p r e s e n t th e d e g re e o f m a tu r ity a t each age,
7 .6 5
3 9 .3 2
54.56
5 5 .4 2
7 6.90
63.60
88.25
6 7 .7 6
94.03
6 9.87
96.96
7 0 .9 4
98.44
7 1 .4 9
99.20
7 1 .7 7
9 9.59
71.91
9 9.79
7 1 .9 8
99.88
72.02
99.94
72.04
99.97
7 2 .0 5
9 9 .9 8
7 2 .0 5
9 9 .9 8
98
TABLE 46.
LEAST SQUARES MEANS OF BIRTH-REARING EFFECTS FOR EACH BREED
(HARVEY ANALYSIS, MODEL B)
T ra it
ATLB
ATLW
ATWW
ATFP
A
k
BI
AGRI
AGR2
AGR3
AGR4
AGR5
RGRI
RGR2
RGR3
RGR4
RGR5
AMRI
AMR2
AMR3
AMR4
AMR5
R am b o u llle t
Targhee
Columbia
S/S
T/S
T/T
S/S
<■
T/S
T/T
S/S
T/S
T/T
1.33
1 .1 9
44.51
4 .5 0
7 1 .5 5
••1 1 .8 0
•62.50
8.46
• 1 .3 4
2.61
• • 3.81
• • 3 .3 5
•2 1 .7 0
• • 2 .7 0
4 .1 0
• • 7.80
• • 5.40
••1 1 .8 0
• • 1.90
3 .6 0
• • 5.40
• • 4 .7 0
1.47
1 .1 8
45.90
4.37
6 9.73
12.20
6 6 .1 0
8.56
1.44
2 .6 2
3.86
3 .3 9
22.60
2.80
4 .1 0
7.90
5.50
12.30
2 .1 0
3 .7 0
5.60
4.90
1 .4 5
1.21
44.44
4.44
6 9 .6 4
10.30
6 6 .9 0
7.40
1 .5 5
2 .6 8
3 .5 5
3.21
21.90
3 .3 0
4 .4 0
7 .8 0
5 .4 0
10.70
2 .2 0
3 .8 0
5 .1 0
4 .6 0
1.46
1.2 4
4 9 .4 3
• 4.84
• • 7 4 .9 2
• • 1 2 .3 0
•6 6 .2 0
• • 9 .1 0
• • 1.1 0
2 .9 9
• • 3 .9 4
• • 3 .5 7
• 21.40
• • 2 .1 0
4 .4 4
• • 7 .8 0
• • 5 .4 0
••1 2 .2 0
• • 1.5 0
• 3.9 0
5 .3 0
4.8 0
1.47
1 .3 2
5 2 .5 2
4 .7 2
6 8 .9 2
12.00
7 6 .0 0
8 .1 9
1 .20
2 .8 6
3.71
3 .3 8
21.70
2 .5 0
4 .6 0
7 .9 0
5 .5 0
11.90
1.7 0
4.2 0
5 .4 0
4.90
1.53
1.2 8
50.87
4.6 5
7 2 .3 9
11.20
70.60
7 .9 9
1 .3 9
3.1 3
3 .7 2
3.4 6
21.50
2.90
4 .8 0
7 .9 0
5 .4 0
11.10
1 .9 0
4 .3 0
5.20
4.8 0
1.33
1.0 8
4 2.74
5 .1 2
•7 5 .1 2
10.60
57.00
• • 8.85
• • .81
3 .2 7
3.61
3 .4 2
21.60
• • 1.60
5 .0 0
• • 7 .6 0
• • 5 .4 0
••1 1 .9 0
• • 1.10
4 .3 0
4 .8 0
4 .6 0
1 .4 8
1.2 0
4 6.19
5.1 6
7 1 .8 4
10.90
6 5 .0 0
8 .7 3
.93
3 .1 3
3 .5 9
3 .3 4
2 2 .3 0
1.90
5 .1 0
7 .7 0
5 .4 0
12.20
1.30
4 .3 0
5.0 0
4 .7 0
1.42
1.13
4 5 .4 2
4.97
7 2 .0 9
10.20
6 3 .0 0
7 .7 8
1.27
3 .1 9
3 .5 6
3 .3 6
2 1 .7 0
2 .7 0
5 .0 0
7 .8 0
5 .5 0
10.90
1.80
4 .3 0
5 .0 0
4 .7 0
U n its o f m e asu re : A, ATWW and ATFP a s k g ; k a s %/mo r a t e o f grow th
d e c li n e ; AGR a s kg/m o; RGR a s % o f w e ig h t g a in p e r month r e l a t i v e to
Y tg; AMR a s % o f w e ig h t g a in p e r month r e l a t i v e t o A; and E I a s % o f A.
« ( P < .0 5 ) , • ( P < .0 1 ) .
99
TABLE 4 7 .
ESTIMATES® OF HERITABILITIES, GENETIC AND PHENOTYPIC
CORRELATIONS FOR LIFETIME PRODUCTION AND GROWTH TRAITS OF RAMBOUILLET
EWES
T ra its
ATLB
ATLW
ATWW
ATFP
A
k
EI
AGRI
AGR2
AGR3
AGR4
AGR5
RGRI
RGR2
RGR3
RGR4
RGRh
AMRI
AMR2
AMR3
AMR4
AMR5
ATLB
.qR + .?7
.9 4 ± ,2 0
1 .8 0 ± 6 .4
- .1 4 ± ,4 9
•60± ,53
- .4 8 ± ,4 8
• 97±3.2
-.2 4 ± ,6 1
“ •26±>37
-.0 5 ± ,5 0
.06± ,44
• 3 8 ± J.3
- .0 7 ± ,40
.28±>38
-.0 5 ± ,4 3
- .1 9 ± ,3 5
- .2 0 ± ,3 5
-.6 1 ± ,5 5
.22± ,36
-.21±>45
-.2 8 ± ,4 1
-.45± > 48
ATLW
.73
.41+ .27
1 .9 7 ± 6 .7
“ »30±,50
.4 4 ± ,5 4
- .2 3 ± ,4 4
1 .4 2 ± 3 .0
- .4 8 ± ,6 4
•43±,37
.2 8 ± ,4 8
•23± ,42
•63±3.5
- .3 5 ± ,4 2
.4 5 ± ,3 8
- .3 8 ± ,4 2
- .1 6 ± ,3 4
- .4 0 ± ,3 7
-.S B ±,51
»41±,35
-.4 3 ± ,4 4
•01± ,39
- .2 0 ± ,4 5
ATUV
ATFP
.66
- .0 0 2
.96
—.06
.01+ .26
—.06
.66+ .27
-1 .2 8 ± 5 .8
.
1 1±,41
1 .8 6 ± 8 .3
-.8 6 ± 3 .6
-.0 5 ± ,4 0
-.0 3 ± 7 .0
-.9 1 ± 3 .5
.2 0 * , 4 8
-1 .8 4 ± 8 .1
1.68* 7 .1
-.2 0 * ,3 2
-.6 3 * 3 .0 ■ -.0 0 5 ± ,4 2
- .1 6 * ,3 6
.8 9 ± 3 .9
- . 60* 1.3
2 .8 6 * 3 3 .
- I .3 8 * 6 .0
- . 1 2±,34
1 .7 4 * 7 .4
- .2 7 * ,3 4
-1 .0 4 * 4 .6
•08±,37
- .6 2 * 2 .8
- .5 1 ± ,3 3
-1 .4 8 * 6 .4
-.6 5 ± ,3 7
-2 .5 9 * 1 1 .
•21±,41
- .2 4 ± ,3 2
1 .4 9 ± 6 .3
-1 .2 1 * 5 .1
.1 8 * ,4 0
- .1 4 * 1 .3
-.1 9 ± ,3 5
- .7 6 * 3 .2
-.1 0 * ,4 3
A
.025
-.1 1
- .1 1
.3 2
.66 + .2 7
- . 6 9±.» 6 8
.2 2 * 1 .9
• 1 1±j.50
.17±,31
-.0 1 *..42
•32± ,32
.47±,81
- .2 0 * ,3 4
.1 2 * . 32
-* 0 1 * ,3 6
.0 1 * ,2 9
-.3 0 ± ,3 1
-•6 3 ± ,5 9
•01*,31
-.1 6 * ,4 1
-.48± > 47
- .8 4 * ,7 5
k
.07
.17
.19
- .0 9
- .5 5
.6Q+.27
.2 0 * 1 .2
- .0 8 * ,4 9
.4 7 ± ,2 8
.5 5 * ,4 9
.4 1 * ,3 0
- .1 8 * . 96
.2 6 * .3 2
.4 7 ± ,3 0
-.4 4 ± ,4 2
.25*»27
.6 1 * .2 5
.43*>30
.5 8 ± ,2 4
- .3 5 * .3 8
•9 1 * .0 8
.7 2 * . 17
H e r i t a b i l i t y e s tim a te s a r e u n d e rlin e d on d ia g o n a l, p h e n o ty p ic
c o r r e l a t i o n s a r e above th e d ia g o n a l, and g e n e tic c o r r e l a t i o n s a r e below
th e d ia g o n a l.
100
TABLE 47.
(C o n tin u ed )
T ra it
EI
AGRI
ATLB
.60
- .01
ATLW
.92
—.03
ATWW
.96
—.01
ATFP
- .1 5
.22
A
—.38
.33
k
.3 3
.24
EI
.04*.?fi
-.1 1
AGRI -1 .7 5 ± 6 .1
.35+ .27
AGR2
1.2 4 ± 4 .0 -.35± > 47
AGR3
-.4 9 ± 2 .1 -.63±#61
AGR4
.3 8 ± J .8
• 11±,45
AGR5
1.59± 7.0 -1 .5 5 ± 2 .3
RGRI
-.9 8 ± 3 .4
.7 9 ± ,2 2
RGR2
1.39±.4.5 -•43±>56
RGR3
-.8 6 ± 2 .9 -.3 0 ± ,5 0
RGR4
-•35dkJ .5
•22± ,34
RGR5
-.7 9 ± 2 .7
•25± ,35
AMRI -1.39±_4.4
•70±,24
AMR2
1.31+4.1 -.3 7 ± ,4 8
AMR3
-.8 2 ± 2 .7
.41±j,55
AMR4
•5 8 ± J.8 - .0 3 ± ,4 2
AMR5
.44±) .4 - .5 9 ± ,5 2
AGR2
.07
.07
.07
-.0 0 4
.15
.20
.03
—.26
1 .05+.27
-.4 1 ± ,4 3
•86 12
1 .3 5 ± 2 .2
- .1 3 ± ,2 7
•99±,01
-.4 9 ± ,4 0
.4 6 ± ,1 8
.2 5 ± ,2 2
-.4 0 ± ,4 0
.98±,01
-.4 6 ± ,4 1
. 6 9±# 16
•30±,31
AGR3
.12
.06
.07
- .0 2
.11
- .1 3
.02
- .0 5
- .4 3
.5 3 + .2 7
- .7 4 ± ,5 9
.O ltl.1
-.2 8 ± ,3 9
-.3 3 t.4 2
1 .0 5 ± ,0 4
-•31±>36
• 07±,30
- .4 7 ± ,4 8
- . jM±,43
1 .0 1 ± ,0 3
-.6 6 ± ,5 6
• 13±,41
AGR4
.05
.05
.06
.19
.43
.37
- .0 5
.46
.69
- .3 9
.71 + .2 7
.79± ,93
.1 0 ± ,3 0
•81 ±j» 18
-.6 7 ± # 5 4
•59±..18
.41±,.24
-.1 5 ± > 3 6
.8 2 ± ,1 4
-»69±j.56
.6 7 ± ,1 7
-.0 0 3 ± ,3 7
AGR5
.13
.0 8
.10
.17
.53
.23
- .0 5
.45
.28
.44
.6 3
.08 + .2 6
—. 6 I^-I .3
I .4 1 ± 2 .5
.47±J.O
.4 7 ± ,9 3
1 .0 4 ± j.6
•-1 .6 5 * 3 .0
1 .2 7 ± 2 .2
.31±>92
.2 4 ± ,8 5
.1 1 ± 3 .0
101
TABLE 47.
(C o n tin u ed )
T ra its
RGRI
ATLB
ATLW
ATWW
ATFP
A
k
EI
AGRI
AGR2
AGRB
AGRU
AGR5
RGRI
RGR2
RGRB
RGRU
RGR5
AMRI
AMRid
AMRB
AMRU
AMR5
-.0 0 5
- .0 6
- .0 5
-.OU
.01
.16
-.0 5
.65
- .0 9
- .2 0
.20
.07
- • 17±,28
- .1 5 ± ,3 2
.U6±,19
.3U±,23
.78± ,17
—.09± ,26
—.19*>34
.2U±,27
.01±,3U
RGR2
.06
.07
.06
- .0 9
.03
.13
.06
-.UU
.96
- .3 9
.51
.13
- .1 3
.QQ+.27
-.U U ±,39
.5 0 ± ,1 8
.25±>22
-.U2±,UU
•99±#01
-.U ItfU O
.7 0 t# 18
.3 7 ± ,3 2
RGRB
.09
.OU
.OU
- .0 7
- .0 3
- .1 9
.03
- .1 7
- .5 3
.96
- .5 8
.21
- .2 2
- .4 6
.7F+ .27
- .2 7 t,3 1
.07t>26
- .2 U t,3 8
-.5 0 t.3 9
1 .0 0 t,0 2
-.6 2 tf U 8
•22t,3 U
RGRU
.03
.01
.01
- .1 0
.06
.23
.01
.24
.41
- .3 4
.53
.22
.44
.39
- .4 0
1 .2 8 + .Pfi
. 661^13
.14±>28
.5 0 t,1 7
-.3 0 t,3 3
• 5 7 t,1 7
.1 9 ± ,2 8
RGR5
.04
- .0 3
- .0 4
- .1 7
- .0 4
.2 5
- .3 0
.13
.11
.24
.22
.3 8
.16
.10
.21
.52
I .26+.2fi
•36t*28
•31±f21
.1 5 t,2 7
• 6 0 tf 2 1
• 7 6 t,2 2
AMRI
—.02
.03
.06
-.0 0 1
- .3 7
.63
.15
.75
—.36
- .1 3
.16
.06
.64
—.46
- .1 4
.20
.15
.8 7 + .2 7
-.2 9 t,3 5
-.2 3 ± ,3 9
.3 1 ± ,2 9
.1 1 t,4 1
102
TABLE 47.
(Continued)
T ra its
AMR2
ATLB
.07
ATLW
.1 0
ATWW
.10
ATFP
- .0 9
A
- .0 7
k
.3 3
EI
.12
AGRI
- .3 4
AGR2
.97
AGR3
- .4 5
AGR4
.60
AGR5
.16
RGRI
—.10
RGR2
.97
RGR3
- .5 3
RGR4
.42
RGR5
.13
AMRI
- .2 9
AMR2 I .15+ .27
AMR3 - •45± ,39
AMR4
.7 9 ± ,1 2
AMR5
.4 4 ± ,2 8
AMR3
.11
.09
.09
- .1 0
- .1 7
.02
.13
- .1 3
- .48
.95
-.5 1
.28
- .2 0
- .4 2
.97
- .3 7
.23
- .0 2
- .4 5
.64+ .2 7
-.5 3 ± * 4 4
•33±*33
AMR4
.04
.14
.15
—.08
- .4 1
.85
.27
.18
.56
—.48
.64
.18
.19
.48
- .5 7
.50
.26
.46
.66
—•38
.85+ .27
.62± ,21
AMR5
.11
.20
.21
- .1 7
- .5 4
.83
.3 5
.09
.10
.31
.16
.42
.07
.08
.25
.16
.42
.47
.22
.46
.62
.55 + .2 7
103
TABLE 4 8 .
ESTIMATES3 OF HERITABILITIESf GENETIC AND PHENOTYPIC
CORRELATIONS FOR LIFETIME PRODUCTICN AND GROWTH TRAITS OF TARGHEE EWES
T ra its
ATUB
ATLB
.24+.22
ATLW
ATWW
ATFP 1.24± 3.6
A
-.2 0 £ l.0
k
.4 4 ± ,7 3
EI
AGRI 1 .60± 4.3
AGR2 - .9 9 ± j.1
AGR3
.72± ,86
AGR4 -.005± #62
AGRS
•31±,52
RGRI
RGR2 - .1 3 ± J .6
RGR3 3 .4 5 ± 4 4 .
RGR4
.5 0 ± J.2
RGR5
AMRI 1 .0 1 * 3 .2
AMR2 -1 .0 2 ± J .2
AMR3
AMR4
.0 2 * ,5 4
AMR5
.9 2 ± ,9 9
ATLW
ATVW
.72
-.03+.22
.6 4
.92
-.20+.23
ATFP
A
k
.03
.005
.04
.04+.21
-4 .4 6 * 3 3 .
.8 7 * 3 .0
.01
- .1 3
—.08
.25
.12+.21
- .1 9 * 3 .3
.10
.15
.17
—.0 8
- .4 8
~2-3+.22
-6 .1 2 * 2 3 .
-.5 2 * 2 .1
.2 3 * 3 .9
-2 .1 4 * 5 .4
1 .7 2 * 4 .4
.24*2.1
-.2 9 * 3 .1
2 .4 8 * 2 .4
.07*^89
1.06*^57
.6 8 * 3 .9
1.14±>91
- I .2 4 * 3 .5
•90*,50
- .0 4 * ,5 4
—.1 6 * 2 .2
5 .2 9 * 7 0 .
- .4 7 * 2 .9
- .5 5 * 3 .5
I .3 0 * 3 .4
12.3 3 * 3 5 7 . -6 .5 0 * 8 4 .
-2 .5 3 * 3 .6
2 .3 0 * 2 .5
- .2 4 * 2 .4
.2 1 * 2 .0
- .9 7 * 2 .4
-.5 3 * 3 .4
.8 0 ± ,4 7
1 .1 6 * ,8 7
.1 5 * 3 .3
.8 6 * 3 .4
- .6 7 * 3 .3
1 .0 7 * 2 .5
1 .0 0 ± ,1 5
.28*^81
H e rita b ility
e s tim a te s
a r e u n d e rlin e d
on d ia g o n a l, p h e n o ty p ic
c o r r e l a t i o n s a re above th e d ia g o n a l, and g e n e tic c o r r e l a t i o n s a r e below
th e d ia g o n a l.
104
TABLE 48. » (Continued)
ATLB
ATLW
ATWW
ATFP
A
k
EI
AGRl
AGR2
AGRB
AGR4
AGR5
RGRI
RGR2
RGRB
RGR4
RGR5
AMRI
AMR2
AMRB
AMR4
AMR5
.59
.8 9
.90
- .0 6
- .4 2
.3 2
.OQ+.22
AGRI
AGR2
AGRB
AGR4
AGR5
.10
.12
.03
.10
.15
.69
.10
- .1 5
.49
.2 8
.54
.68
.48 + .2 1
- .5 5
- .0 5
.02
.07
.10
.19
-.0 1
- .2 3
.IQ + .21
- .6 4 ± J .3
1 .2 6 ± ,4 8
•51± ,52
.06
-.0 0 5
.35
—.26
—.06
—.01
- .2 8
.18+.21
.32*^81
1 .1 5 * ,4 8
.06
—.0 2
.07
.16
.47
.38
—.01
.58
.5 9
- .2 0
.1 1 + .2 2
.64±>26
.46± 3.4
1 .0 0 ± ,0 7
17.07±225. -3 .9 8 * 5 3 .
-2 .0 1 ± 5 .6
1 .7 6 ± J.8
-1 .1 7 * 3 .9
2 .5 7 * 2 9 .
—.8 8 * 3 .8
1 .3 5 t.9 6
1.24*3 8 .
.7 5 * ,7 t
.4 0 * ,7 0
4 .6 6 * 5 8 .
- .2 1 ± ,7 3
.16± 2.3
.24± 2.5
.6 9 t J .6
.9 8 * ,0 5
-.3 5 * 3 .4
1 .1 4 * 3 .8
.16*,81
1.19*^62
- .4 6 * , 84
.25± ,61
- .1 8 ± 3 .2
1 .4 6 ± 4 .3
1 .1 6 ± ,4 9
1 .5 9 * 3 .4
- .9 9 * 3 .2
- . 4 8 * , 92
.73±>27
1 .4 0 * 3 .0
- .0 9 * ,3 9
.97±>66
.11
- .0 0 2
.04
.12
.43
.26
- .1 1
.04+.21
.54± 2.8
3 .2 3 ± 8 .9
•3 1 ± J.1
.8 7 ± J .8
i
EI
O
Co
T ra its
105
TABLE 48 .
T ra its
(C o n tin u ed )
RGRI
ATLB
-.0 1
ATLW
- .0 5
ATWW
- .0 4
ATFP
- .0 6
A
.02
k
.1 4
EI
- .0 4
AGRI
.56
AGR2
- .0 9
AGR3
- .1 7
AGR4
.16
AGR5
.03
Rtihl —.104+.22
RGR2
RGR3
RGR4
RGR5
AMRI
AMR2
AMR3
AMR4
AMR5
RGR2
RGR3
RGR4
RGR5
AMRI
- .07
- .0 4
.004
.01
- .0 2
.1 2
.02
- .4 2
.97
—.26
.40
.13
- .1 3
.13+ .21
6 .8 3 ± 8 9 .
2 .1 9±2.5
.06
.07
.03
- .0 9
.11
- .3 3
—.01
- .2 2
- .4 4
.92
-.5 1
.22
-.2 1
-•3 6
.0 1 + .2 0
-4 .3 3 * 5 7 .
- .0 3
.00
.01
- .1 3
.04
.18
.003
.14
.28
.18
.42
.20
.23
.27
- .2 7
.11+ .21
.8 8 * 2 .3
1 .0 4 ± ,0 9
-.8 9 * 3 4 .
6 .4 1 * 8 5 .
.8 3 * 3 .5
2 .3 1 * 2 .5
.10
.10
.10
- .0 7
—•36
.69
.22
.6 8
- .3 2
- .3 1
.22
—.05
.57
- .4 2
- .3 3
.11
- .1 1
.12+.21
.7 7 * 3 .7
1 .4 0 ± ,9 4
1 .6 6 * 3 .7
4 .9 6 * 6 5 .
-4 .6 3 * 5 8 .
1 .8 4 * 3 .6
2 .5 1 * 2 .9
.61±,51
.1 1 * 3 .2
- .0 0 3
- .0 0 4
.01
- .1 3
.10
.04
—.02
- .0 3
.18
.24
.19
.35
-.0 1
.18
.20
.48
—.0 6 + .2 2
106
TABLE 48.
(Continued)
T ra its
AMR2
AMR3
AMR4
AMR5
ATLB
ATLW
ATWW
ATFP
A
k
EI
AGRI
AGR2
AGR3
AGR4
AGR5
RGRI
RGR2
RGR3
RGR4
RGR5
AMRI
AMR2
AMR3
AMR4
AMR5
—.06
—.02
.03
.0 0 2
- .1 2
.3 0
.08
- .3 3
.97
- .3 5
.47
.12
—.10
.97
- .4 6
.28
.17
- .1 5
.17+.21
.11
.14
.10
—*11
—.05
—.08
.11
- .1 9
- .3 5
.89
—*41
.26
- .1 8
—.28
.93
- .2 2
.20
- .1 7
- .3 4
-.12+.2%
.04
.08
.12
- .0 7
- .4 2
.85
.26
.20
.51
- .5 3
.59
.07
.15
.43
- .6 3
.42
.12
.56
.60
- .4 0
.4S+.21
.6 9 ± ,3 8
.13
.20
.21
- .1 5
- .4 6
.77
.36
.05
.21
.19
.23
.32
.03
.18
.12
.21
.32
.42
.30
.39
.64
.16+.21
1 .27 ± ,55
1.30£J .1
107
TABLE 4 9 .
ESTIMATES3 OF HERITABILITIES, GENETIC AND
PHENOTYPIC
CORRELATIONS FOR LIFETIME PRODUCTI CN AND GROWTH TRAITS OF COLUMBIA EWES
T ra its
ATLB
.07
.10
.16
I .T2+.T6
.5 8 4 ,3 1
- .3 6 4 ,3 2
- .2 1 4 ,5 0
- .9 9 1 ,5 9
.9 7 1 ,2 2
- .2 6 i,3 4
.7 0 4 ,3 5
.2 5 1 ,3 6
-.5 4 4 ,3 5
.9 4 4 ,2 3
- .5 7 1 ,4 5
.0 4 ± ,4 9
-.3 0 t,2 9
-1 .1 4 4 ,5 7
.9 3 1 ,2 5
- .6 0 i , 4 0
.0 5 4 ,3 9
.4 5 4 ,3 4
A
.06
- .0 4
.03
.29
.7 0 + .TB
- .3 8 4 ,8 2
-.7 7 1 ,7 1
.2 0 4 ,5 2
.3 1 4 ,3 5
.1 3 4 ,4 3
.5 8 4 ,4 3
.5 1 1 ,3 7
.2 9 4 ,4 0
.2 0 4 ,3 5
- .3 1 1 ,4 9
.8 4 4 ,8 8
-.0 1 i,3 5
- .5 9 4 , 71
.1 0 4 ,3 8
- .4 5 1 ,5 0
- .5 1 1 ,7 8
- .7 4 4 ,7 0
k
.21
.17
.11
- .1 1
0
.63
.96
.T5+.TQ
.1 4 i,4 8
.3 2 i,6 7
.1 5 1 ,5 3
.7 9 1 ,2 7
.0 1 t,7 5
.6 4 4 ,5 8
- .6 0 t,6 l
.8 3 1 ,7 5
.1 5 1 ,6 8
- .0 7 1 ,5 3
. 784,62
-.9 2 ± ,8 5
I . I 64J . I
.0 2 t,4 8
- .3 4 1 ,6 4
.6 9 1 ,6 0
- . 764,65
.4 8 4 ,7 0
-.1 8 i,5 3
ATFP
I*
.72
.4 8 + .TQ
.9 4 i,0 8
.1 9 1 ,4 2
.0 7 1 ,5 6
.3 5 1 ,4 4
.8 8 i,1 4
.0 3 1 ,6 5
.5 6 i,4 7
-.4 7 1 ,5 1
•69 i ,6 2
.1 6 1 ,5 9
- .1 4 1 ,4 7
.6 8 ± ,4 9
-.7 1 i,6 6
.7 0 1 ,7 6
.071,41
- . 1 4 i ,5 2
.6 4 t,5 0
- . 4 8 i ,5 0
.5 7 1 ,5 9
.0 9 1 ,4 8
ATWV
NO
ATLB
.6 5 + .TB
ATLW
.7 7 ± ,2 6
ATWW .6 1 ± ,4 2
ATFP
.5 2 ± ,3 8
A
.2 5 ± ,4 9
k
.4 0 ± ,3 7
EI
.5 3 ± ,4 6
AGRI
.44± ,54
AGR2
.40± ,36
AGR3
•29±,45
AGR4
.7 2 ± ,4 8
AGR5
.8 2 1 ,4 5
RGRI .-.0 3 ± ,3 9
RGR2
.3 8 1 ,3 6
RGR3 ■-.1 2 i,5 2
RGR4
.2 2 1 ,6 4
RGR5
.1 6 1 ,3 5
AMRI
.1 0 i,4 6
AMR2
.4 2 1 ,3 7
AMR]
.19± ,45
AMR4
.4 7 1 ,4 9
AMR5
.4 5 1 ,3 7
ATLW
I .OQ+.T7
.6 2 1 ,5 0
.0 6 ± ,4 5
- .0 7 4 ,3 0
.1 1 1 ,3 8
- .0 5 1 ,4 4
.0 4 i,4 2
-.1 6 1 ,3 1
. 01i ,3 0
.1 3 4 ,4 5
.4 6 1 ,4 7
.2 5 4 ,2 8
.7 0 i.1 9
.1 0 4 ,3 1
.4 5 4 ,3 7
.8 7 4 ,1 0
.9 2 4 ,0 7
H e r i t a b i l i t y e s tim a te s a r e u n d e rlin e d on d ia g o n a l, p h e n o ty p ic
c o r r e l a t i o n s a r e above th e d ia g o n a l, and g e n e tic c o r r e l a t i o n s a r e below
th e d ia g o n a l.
108
TABLE i»9.
(Continued)
T ra its
EI
ATLB
ATLW
ATWW
ATFP
A
k
EI
AGRI
AGR2
AGR3
AGR4
AGR5
RGRI
RGR2
HGRB
RGR4
RGR5
AMRI
AMR2
AMRB
AMR4
AMR5
.60
.9 3
.95
.0 5
—.26
.2 9
-35+.1Q
-.1 2 ± ,7 6
.4 3 ± ,5 4
.6 l± j,68
.4 3 ± ,7 4
-.1 9 ± ,6 9
-.13± >54
.6 3 ± ,5 8
-.7 0 ± ,7 9
.86± ,93
.10± ,47
.08± ,61
.61± ,57
-.5 0 ± ,5 8
«82±,67
.3 0 ± ,4 9
AGRI
.14
.10
.08
- .0 3
.25
.25
- .0 0 4
.5 1 + .3 9
-•73±#65
•55±,55
-.6 1 ± ,4 4
-.0 1 ± ,6 4
1 .0 4 ± ,3 4
- .7 2 ± ,7 0
.4 2 ± ,6 9
.5 9 ± ,7 0
-.1 4 ± ,4 0
.6 5 ± ,2 8
-.7 8 ± ,7 2
.4 4 ± ,5 7
-.63± »56
-.1 7 ± ,4 5
AGR2
.16
.0 5
.10
.33
.15
.18
.06
- .3 1
I.P R + .36
-.4 3 ± ,4 4
1.01± ,21
.3 2 ± ,3 6
- .5 1 ± ,3 5
•99±,01
- *6 9±j 6 5
.2 4 ± ,4 7
-.1 1 ± ,2 7
-.75± > 5 4
.98± ,01
-.5 9 ± ,5 1
.5 9 ± ,2 8
-.06±>31
AGR3
.09
.14
.16
.03
.30
- .1 9
.07
.03
- .3 4
.8 2 + .3 8
■.07±>51
.7 8 ± ,3 0
-.0 1 t» 3 6
-.47± > 44
.8 3 t» 1 2
• 97±,98
•79±>25
.28±>46
- .4 5 ± ,4 8
.8 3 1 ,1 1
- .1 4 i,5 7
.4 1 1 ,3 5
AGR4
.27
.17
.20
.31
.34
.38
.09
.45
.67
- .2 3
.5 5 + .3 8
.5 9 1 ,3 4
-.1 8 i,4 2
1 .0 2 1 ,2 8
-.5 2 ± ,8 9
.9 7 1 ,5 7
- .0 5 1 ,3 9
-.8 0 i,5 2
.9 6 i,2 4
- .3 7 1 ,6 5
.3 9 1 ,4 5
.1 0 1 ,4 4
AGR5
.31
.27
.3 0
.27
.5 4
.17
.13
.47
.2 9
.52
.70
.5 8 + .3 8
-.I 6 i,4 2
.3 1 1 ,3 7
.2 9 1 ,5 2
1 .4 9 1 ,9 4
.5 6 1 ,3 2
-.3 6 i,5 0
.2 7 1 ,3 9
.4 2 i,4 2
.1 2 t,5 5
.2 2 1 ,4 1
109
TABLE 49.
T ra its
ATLB
ATLW
ATWW
ATFP
A
k
EI
AGHI
AGR2
AGR3
AGR4
AGR5
RGRI
RGR2
RGR3
RGR4
RGR5
AMRI
AMR2
AMR3
AMR4
AMR5
(Continued)
RGRI
- .05
- .1 1
- .1 2
- .2 1
- .0 4
.0 8
- .1 0
.49
- .1 6
- .2 3
.05
- .0 5
I . 14+.17
-.55± > 35
.0 4 ± ,4 2
—.10±>51
- .1 5 ± ,2 8
.62± ,26
- .5 8 ± ,3 7
-.10± > 37
-.3 4 ± ,4 1
-.32± > 33
RGR2
.13
.05
.10
.28
.08
.13
.08
- .4 5
.97
- .3 4
.54
.18
—.18
I .1 1 + .ifi
-.6 9 ± # 6 2
•15±,47
- .0 6 ± ,2 7
-.6 7 ± ,5 3
.99±>01
-.57± > 49
.7 0 ± ,2 9
.04± ,31
RGR3
- .0 3
.07
.08
- .0 8
.08
- .2 8
.05
—.1 8
-.5 1
.90
—.56
.15
- .2 2
—.46
.6 1 + .IQ
•73±3.0
•96±>33
.4 7 ± ,5 9
- .6 5 ± ,6 7
.92±>07
-.18±>71
.4 7 ± ,4 4
RGR4
.18
.19
.16
—.0 8
—.06
.30
.18
.22
.23
—.18
.43
.28
.39
.21
—.28
.Ifi+ .IQ
.7 9 ± ,3 2
—.08±>60
• 19±>49
.69± ,85
.2 9 ± ,6 2
.5 0 ± ,4 8
RGR5
.15
.25
.23
- .1 2
- .0 9
.18
.25
.06
- .0 0 4
.36
.12
.35
.13
.01
.35
.60
1 .4 0 » .1 6
- .0 7 1 ,3 3
- .0 8 1 ,2 8
.8 7 1 ,2 3
.0 2 1 ,3 8
.5 0 1 ,2 3
AMRI
.07
.09
.01
—.26
- .5 2
.70
.16
.6 9
- .3 9
- .1 9
.14
.002
.47
—.46
- .2 2
.23
.10
.7Q+.18
—.6 3 1 ,5 0
.6 0 t,4 5
.0 1 i,5 4
.4 5 1 ,2 9
110
TABLE 49.
(Continued)
T ra its
AMR2
ATLB
.16
ATLW
.06
ATWW
.10
ATFP
.26
A
- .0 4
k
.30
EI
.11
AGRI
.37
AGR2
.97
- .4 0
AGR3
AGR4
.60
.1 8
AGR5
RGRI
- . 16
RGR2
.97
RGR3
- .5 3
RGR4
.24
.01
RGR5
AMRI
- .3 0
AMR2 I .1 6+ .17
AMR3 - •49±>50
AMR4
.7 5 ± ,2 4
AMR5
• 13±,32
AMRS
AMR4
AMR5
.07
.1 8
.17
—.10
- .1 1
.04
.19
—»06
- .4 4
.90
- .3 9
.30
-.2 1
- .4 0
.92
- .1 6
.43
.0 2
- .4 3
.8 1 + .IR
• 13±,53
.75± ,2 7
.20
.17
.13
.02
- .5 0
.87
.27
.20
.50
- .4 5
.63
.1 8
.08
.43
- .5 9
.43
.16
.55
.60
- .2 9
.SB+.IR
•75±>20
.26
.31
.26
- .0 7
- .5 4
.86
.41
.20
.13
.1 8
.32
.40
.005
.09
.05
.33
.43
.58
.22
.41
.73
.QQ+.17
Ill
TABLE 50.
AMONG SIRES COMPONENTS OF VARIANCE AND COVARIANCE FOR
LIFETIME PRODUCTION AND GROWTH TRAITS (POOLED DATA FROM RAMBOUILLET,
TARGHEE AND COLUMBIA EWES)a
T ra its
ATLB
ATLW
ATWW
ATFP
A
k
BI
AGRI
ATLB
ATLW
ATWW
ATFP
11.516
9.2949
8.9750
235.49
208.33
3769.8
4 .8 0 7 6 .
4.2046
117.27
49.654
aV alu es a r e m u ltip l ie d by IO^
C o v arian c e s a re on th e d ia g o n a l
A
k
EI
48.855 .12880 3 .0 2 6 5
32.487 .07598 2 .9 8 5 9
2004.8 .08170 49.497
17.659 -.4 0 9 4 9 .72560
4027.5 -1 2 .8 5 0 -1 9 .6 2 9
.08750 .14685
1.1682
AGRI
7.9 3 2 8
-4 .9 9 6 9
-2 0 6 .4 3
-3 5 .6 8 9
28.958
.11336
-3 .8 8 0 4
39.097
112
TABLE 50.
T ra its
ATLB
ATLW
ATWW
ATFP
A
k
EI
AGRI
AGR2
AGRB
AGR4
AGR5
RGRI
RGR2
RGRB
(Continued)
AGR2
2 .1 2 1 4
9 .1 9 1 5
3 6 0 .0 6
18.451
6 6 .4 7 1
.6 4 3 9 4
4 .6 7 3 5
- 1 9 .1 0 9
4 6 .4 4 8
AGRB
8 .2 7 4 5
- 3 .2 4 3 7
-7 7 .6 6 7
-5 .0 9 8 1
2 4 1 .2 1
—.8 6 8 6 9
- 3 .6 6 4 9
2 4 .5 3 4
- 2 4 .5 9 9
5 9 .9 3 7
AGR4
3.0 3 5 1
4.4791
181.31
I .4283
7 1 .3 4 4
.4 1 3 6 8
1 .7 9 4 5
- 4 .3 4 8 8
2 5 .5 5 8
-7 .1 7 4 2
1 5 .3 6 4
AGR5
4 .5 5 7 0
2 .5 1 4 8
1 0 3 .9 9
- 2 .3 1 7 9
9 6 .7 7 4
- .0 1 4 3 5
.38743
- .5 2 6 6 9
8 .1 3 5 4
1 5 .3 8 4
5 .9 5 7 6
7 .5 6 3 0
RGRI
.01419
-.1 0 9 1 1
- 3 .9 3 8 8
- .4 5 0 4 4
-2 .3 2 6 1
.0 1 2 9 2
- .0 3 2 7 8
.85 0 6 7
-.2 1 0 0 8
- 1 0 .7 0 9
.02102
-.0 8 2 0 3
.02 2 7 5
RGR2
RGRB
.03252
.18 8 9 2
7 .2 3 8 0
.33943
.55 0 0 4
.0 1 3 3 2
.10400
- .3 7 1 2 8
.84052
- .5 0 3 7 5
.4 5 9 7 9
.1 3 4 5 9
- .0 0 4 3 5
.01 5 4 5
.0 5 1 8 9
- . 0 8805
- 2 .7 2 5 7
- 1 .0 0 0 5
2 .4 8 8 4
- .0 1 2 9 8
- .0 6 4 4 4
.38217
- .4 6 9 1 2
.7 5 4 7 8
- .1 7 6 4 7
.15947
- .0 0 0 3 4
- .0 0 9 2 9
.01 0 6 2
113
TABLE 50.
T ra its
ATLB
ATLW
ATWW
ATFP
A
k
BI
AGRI
AGR2
AGR3
AGRn
AGR5
RGRI
RGR2
RGR3
RGRU
RGR5
AMRI
AMR2
AMR3
AMR4
AMR5
(Continued)
RGR4
.00039
.0 1 3 3 7
.683 4 2
- .0 3 4 6 6
- .2 3 0 1 4
.0 0 3 2 7
.01329
.01823
.061 8 7
- .0 1 3 3 7
.04550
.0 13 4 8
.00082
.00115
-.0 0 0 3 0
.000 3 0
RGR5
.00162
.00162
.07545
- .0 3 3 5 5
- .0 6 4 4 5
.00152
.00 2 1 9
.01753
.02 0 2 8
.01 2 6 4
.02183
.0 1 6 3 2
.00031
.00 0 3 4
.00015
.00 0 1 4
.00013
AMRI
AMR2
.01643
- .1 4 2 7 3
-6 .6 2 4 6
-.5 4 9 0 1
- 6 .2 7 1 3
.0 2 4 7 0
- .0 2 8 8 6
.50 9 4 2
- .3 6 4 0 0
—.1 0 6 0 0
- .1 6 5 1 3
- .1 7 6 2 5
.01610
- .0 0 5 9 7
.00050
.00 0 6 4
.00032
.0 1 7 9 9
.03070
.13161
4 .8 2 2 1
.23071
-.1 8 8 6 1
.01260
.07602
- .2 9 0 1 5
.63907
- .4 0 0 2 8
.3 3 9 4 8
.0 8 9 2 2
-.0 0 2 4 4
.0 1 1 7 9
- .0 0 7 1 8
.00101
.00032
- .0 0 3 5 8
.00 9 1 5
AMR3
.09036
- .0 6 2 2 5
-2 .1 2 9 1
-.1 0 5 2 1
1 .4 1 3 7
- .0 0 5 0 9
- .0 4 3 4 7
.4 0 1 1 5
- .3 6 7 3 2
.65433
- .1 1 0 2 2
.1 7 5 8 4
.00033
- .0 0 7 3 2
.0 0 8 6 5
-.0 0 0 0 3
.00025
.00256
- .0 0 5 4 8
.00781
AMR4
AMRS
.01137
.04476
1.2 8 4 6
- .0 1 9 6 9
- 2 .3 0 2 6
.0 1 6 7 8
.04580
-.1 0 6 5 6
.30396
- .2 7 9 5 0
.15435
.0 0 2 8 3
.00195
.00 6 0 3
- .0 0 4 3 0
.00085
.00036
.0 0 2 5 9
.00495
- .0 0 2 5 2
.00406
.03 5 2 8
.01756
.2 9 3 4 4
- .0 6 7 5 3
-1 .4 3 4 7
.0 0 9 3 5
.02146
- .0 1 6 4 5
.06976
.0 4 2 1 8
.0 4 0 3 3
.04 1 4 3
.00 0 5 0
.0 0 1 5 0
.0 0 0 4 5
.00 0 3 7
.0 0 0 2 9
.0 0 2 0 8
.00 1 3 9
.00 1 4 3
.00 1 7 3
.0 0 1 6 2
m
TABLE 51.
WITHIN SIRES COMPONENTS OF VARIANCE AND COVARIANCE FOR
LIFETIME PRODUCTION AND GROWTH TRAITS (POOLED DATA FROM RAMBOUILLET,
TARGHEE AND COLUMBIA EWES)3
T ra its
ATLB
ATLW
ATWW
ATFP
A
k
BI
ATLB
ATLW
ATWW
9 5 .2 5 2
6 8 .7 4 9
9 9 .7 5 8
2 2 9 7 .6
3 5 4 8 .4
138899
3V a lu e s a r e m u l t i p l i e d by IO^
C o v a ria n c e s a r e o n th e d ia g o n a l
ATFP
A
k
EI
AGRI
.86411 3 1 .1 8 2 17.730
- .5 2 1 5 7 1 .0 2 9 0
- 2 .4 2 3 2 - 2 4 8 .8 3
1.3061 4 9 .7 3 5 1 0 .5 0 5
1 4 9 .9 2 - 6 9 5 2 .8 4 9 .1 8 5 1 9 1 6 .3 6 5 4 .7 6
2 4 2 .1 2 905.31 - .8 3 7 9 9 - 6 .2 0 3 7 8 9 .3 7 3
3 4 5 1 8 . - 6 9 .4 6 4 - 3 7 9 .0 5 1 9 7 8 .8
.5 3 1 6 5 1.2521
5 .6 4 1 7
2 9 .1 2 2 - 8 .3 2 0 4
115
TABLE 51.
T ra its
ATLB
ATLW
ATWW
ATFP
A
k
EI
AGRI
AGR2
AGR3
AGRH
AGR5
RGRI
RGR2
RGR3
(Continued)
AGR2
3 .5 3 1 8
- 7 .0 1 3 9
- 7 0 .2 5 2
1 1 .0 5 9
3 0 8 .4 8
1.6711
-3 .2 5 7 9
-9 0 .4 7 2
1 8 5 .6 2
AGR3
1 8 .7 1 7
2 5 .8 4 7
892.41
3 .5 3 5 3
1 0 1 0 .9
- 2 .9 4 2 2
3 .4 2 5 6
- 2 6 .4 2 8
-1 0 2 .5 0
4 8 8 .5 3
AGR4
AGR5
1 1 .0 2 6
9 .1 8 3 9
7 .1 3 0 2
1 .2 7 2 8
3 9 6 .5 0
2 3 6 .1 9
3 7 .9 6 2
2 9 .2 9 8
8 5 3 .3 4
935.71
2 .9 9 0 3
1 .0 6 2 9
- 4 .0 1 4 5 - 2 .5 6 1 3
170.11
1 1 9 .4 0
8 6 .8 7 4
28.4 1 1
- 6 5 .6 2 4
8 6 .4 9 5
1 1 3 .3 8 • 5 9 .4 6 2
6 7 .4 0 7
RGRI
- .0 2 2 3 2
—.0 8 6 10
-2 .3 5 1 1
- .1 3 9 6 4
2 .7 6 0 4
.0 2 7 1 0
.05096
5 .3 6 5 7
-.2 9 6 4 1
-1 .4 6 8 0
.6 6 3 3 4
.2 4 0 6 2
.11 6 7 3
RGR2
.02427
- .1 4 6 1 2
- 2 .5 5 3 0
- .0 8 3 9 9
.41043
.01551
- .0 3 2 2 6
- 3 .0 8 5 9
3 .3 5 2 8
- 1 .7 4 6 4
1.0841
.2 0 6 7 6
- .0 0 9 1 7
.06544
RGR3
.13 9 8 9
.3 0 6 5 4
8 .5 6 4 2
- .3 6 4 3 6
1 .5 6 2 5
- .0 5 7 7 4
.10 3 8 3
-2 .0 9 3 1
- 2 .0 1 5 6
6 .3 1 8 3
- 1 .8 5 9 8
.3 9 4 5 4
- .0 2 4 8 7
-.0 3 0 6 1
.09 5 3 3
116
TABLE 51.
T ra its
ATLB
ATLW
ATWW
ATFP
A
k
EI
AGRI
AGR2
AGR3
AGRU
AGRS
RGRI
RGR2
RGR3
RGRU
RGR5
AMRI
AMR2
AMR3
AMRU
AMR5
(Continued)
RGR4
RGR5
AMRI
AMR2
AMR3
AMR4
AMR5
.01716
.0 1 2 3 2
.26U75
—«06708
.38715
.0 0 6 5 7
.00085
.269 1 0
.20951
-.2 9 U 9 8
.2U161
.09051
.00613
.0 03 7 0
- .0 0 5 5 9
.002 7 0
.01392
.01U82
.52503
- .0 3 2 2 9
.09341
.00157
.00593
.02U28
.027U2
.1 5 2 9 9
.03693
.06836
.00077
.0005U
.00186
.0 0 0 6 2
.00059
.23372
.50711
1 8 .5 4 2
- .1 6 0 2 2
- 2 5 .4 7 2
.18 9 2 9
.45617
7 .6 2 3 8
- 1 .7 4 7 4
- 1 .8 8 5 5
1 .0082
.20357
.07 0 4 9
- .0 4 3 9 3
.03112
.00307
.00015
.14 5 4 8
.04848
- .0 3 8 2 8
.75560
-.0 8 0 9 1
-3 .3 4 2 1
.03901
.04158
- 1 .7 2 2 5
2 .4 9 5 3
- 1 .6 3 2 9
.99914
.17 6 1 8
-.0 0 5 1 1
.04654
- .0 2 8 3 2
.00285
.00038
- .0 1 9 1 8
.03582
.26540
.5 1 1 6 7
1 6 .2 6 8
- .4 6 5 3 9
-7 .1 5 8 0
-.0 0 2 2 3
.28071
- 1 .5 8 6 0
-1 .6 4 3 9
5 .9 7 8 8
-1 .4 5 6 7
.5 6 3 3 4
- .0 2 1 5 8
- .0 2 5 1 4
.0 8 6 7 2
-.0 0 4 5 7
.00192
- .0 1 0 9 9
-.0 2 1 6 1
.08785
.11460
.1 6 7 6 2
7 .1 9 8 4
- .1 1 3 8 8
- 1 1 .3 3 4
.090 00
.19118
.99451
.98341
- 1 .5 8 1 4
.97303
.1 7 9 9 9
.00733
.0 1 4 6 5
- .0 2 7 0 6
.0 0 3 1 0
.00043
.03111
.01628
-.0 1 6 0 1
.02126
.14 5 3 8
.2 4 6 0 9
9 .3 0 8 6
- .1 8 4 7 9
- 9 .1 3 1 4
.0 5 9 1 8
.20197
.36 0 1 5
.17451
.51566
.2 5 0 5 3
.30 6 5 3
.0 0 1 7 2
.0 0 2 2 9
.0 0 4 4 9
.00093
.00083
.0 1 8 9 9
.00436
.01226
.00951
.00990
MONTANA STATt UNIVOtSITV UM ARKS
Ill
762 1001 3856 7