E. P. Cunningham and C. R.

Henderson
Traits in Beef Cattle
Estimation of Genetic and Phenotypic Parameters of Weaning
1965, 24:182-187. J ANIM SCI
http://www.journalofanimalscience.org/content/24/1/182
services, is located on the World Wide Web at:
The online version of this article, along with updated information and
www.asas.org
by guest on July 14, 2014 www.journalofanimalscience.org Downloaded from by guest on July 14, 2014 www.journalofanimalscience.org Downloaded from
E S T I MAT I ON OF GE NE T I C AND P HE NOT YP I C P ARAME T E RS
OF WE ANI NG T RAI T S I N BE E F CAT T L E 1
E . P . C U N N I N G H A I V s ~ A N D C . R . H E N D E R S O N
Cornell Uni versi t y, It haca, Ne w Yor k
p
REWEANI NG performance is recognized
as one of the most i mport ant factors in
determining the economic ret urns from beef
cattle, and in genetic i mprovement of the
beef cattle popul at i on for overall product i vi t y.
Genetic response to selection in such a popula-
tion depends on the i nt ensi t y of selection, the
genetic vari abi l i t y of the traits concerned,
t he relation among them, and the accuracy
with which we can i dent i fy the individuals
in the popul at i on with their genotypes. Efforts
to maximize this response for preweaning per-
formance require a knowledge of the genetic
and phenot ypi c paramet ers of the components
of this trait. A number of estimates of these
paramet ers have been derived from records in
experiment station herds. The present paper
reports on the estimation of environmental
effects, variance components, heritabilities and
correlations for weaning traits in a large popu-
lation of commercial herds.
Ma t e r i a l s and Me t h o d s
The dat a available for this st udy were col-
lected on 83 farms in New York over the
years 1946 to 1960, and comprised 7,971 re-
cords of the Angus, Hereford and Short horn
breeds. The breeding pat t er n of these herds
in N'ew York is largely seasonal, i.e., the cows
caIv~ in March, April and May and the calves
are weaned in October, November and De-
cember. They are usually sold at weaning
as feeder calves. Each record consisted of a
weight and t ype score on a weanling calf with
i nformat i on on the i dent i t y of the calf, sire,
dam and herd, and on the breed, sex, year
and date of birth, date of weighing and creep-
feed status of t he calf. Because of deficiencies
of Various kinds in the dat a and the restric-
tions imposed for the purposes of t he analysis,
3,133, or 39% of these records were discarded.
Thi s reduction eliminated dat a for the first
x Based in part on a thesis submitted by the senior author
to the Graduate School, Cornell University, in partial fulfillment
of the requirements for the Ph.D. degree.
Present address: Animal Breeding and Genetics Department,
The Agricultural Institute, Castleknock, Co. Dublin, Ireland.
3 years of the program from t he Short horn
breed and from 28 herds, most of which had
ver y small numbers of cows.
Trai t s Studied. Preweani ng performance
may be eval uat ed by means of records on
weight at weaning ( WW) , average daily gain
from bi rt h to weaning ( ADG) and, to a lesser
extent, by the t ype score or grade of the calf
at weaning ( TS) . ADG is a function of WW,
days of age ( DA) and bi rt h weight ( BW) .
I f growth is linear with age of calf, t hen ADG
WW- BW
may be expressed as DA . In the pre-
sent dat a bi rt h weight was not available, and
was t aken as a const ant for each breed (60 lb.
for Angus and 70 lb. for Her ef or d) . I n this
case WW standardized for age of calf may be
considered the same t rai t as ADG, differing
from it onl y because of a scaling factor. Thi s
is confirmed by the values of 0.98 and 0.93
obtained for t he genetic correlation of ADG
and WW by Koch and Cl ark (1955a) and
Lehmann et al. (1961), al t hough in bot h of
these studies actual bi rt h weights were used.
The assumption of l i neari t y of growt h with
age was examined by dividing t he age of calf
into 10-day intervals from 120 to 250 days
and plotting t he unadj ust ed means of t he
records for each interval. Thi s was done
separat el y for each breed, and the response
appeared to be closely linear. I n addition,
tests of the significance of linear regression
were computed, using unadj ust ed weaning
weights and single-day age intervals from
120 to 250 days. These are present ed in table
1. The values of the regression coefficient were
1.67 for the Angus and 1.49 for t he Herefords.
All WW records were convert ed to ADG
figures, and the traits ADG and TS were
t hen t aken as a measure of preweaning pre-
formance. TS was based on t he st andard
feeder calf grades, with each grade being
divided into plus, average and minus. Nu-
merical values were t hen assigned t o t he
various grades, ranging from 4 (Common) to
17 ( Fancy + ) . Almost all of t he calves in
t he program were graded by one man.
182
by guest on July 14, 2014 www.journalofanimalscience.org Downloaded from
ESTI MATI NG WEANI NG TRAI TS I N CATTLE i 33
T AB L E 1. ANAL YS I S OF VAR I ANC E ] ? OR L I NE AR R E GR E S S I ON OF
WE ANI NG WE I GHT ON DAYS OF AGE ~
An g u s He r e f o r d
So u r c e d. f . MS f d. f . MS f
L i n e a r r e g r e s s i o n 1 7 7 3 6 2 7 6 . 0 1 8 6 1 . 8 * * 1 2 9 8 2 5 7 2 . 0 4 8 0 . 5 * *
De v i a t i o n s f r o m
l i n e a r r e g r e s s i o n 128 4 1 5 5 . 3 1 . 3 124 6 2 0 7 . 3 1 . 2
E r r o r 3279 3 1 6 2 . 8 9 1732 5 1 1 6 . 2 . .
a Includes 427 records which were discarded in the l at er analyses for lack of age-of-dam information.
~ p~. 01.
General i nformat i on relating to the dat a is
summarized in table 2.
Method of Analysis. Each ADG and TS
record was classified by breed, sex, age of dam,
year, sire, herd, creep-feed status and mont h
of birth. Separate analyses were carried out
for t he two breeds. Mean values for bot h
t rai t s varied little with mont h of bi rt h, and
this classification was ignored. Sires were
nested within herds and herds within creep-
feeding classes. Classification by dams was
not considered, and the dam cont ri but i on to
variation was t aken to be a random variable.
The following model was constructed from
the five classifications, sex, age of dam, year,
sire, herd, and their interactions, for purposes
of locating and testing t he principal sources
of variation:
Xijklmq=/~+ SI--~Aj+Yk-~- SAIj-~- SYlk+Plm
- ~- ei j kl mq
where
Xijklmq is the single observed ADG or TS
is the popul at i on mean
Si is the effect of t he ith sex
Aj is the effect of the j t h age of dam
Yk is the effect of the kth year
Plm is the effect of the lmth sire-herd
class
eijklmq iS a random error specific to this
observation.
Ther e were t hree sexes: bull, heifer and steer.
Age of dam was grouped into seven classes
as follows: 2, 3, 4, 5, 6, 7 to 10 and over 10
years of age. These groupings were set up
from the observed distribution of the mean
T AB L E 2. ME ANS AND S I R E AND HE R D
NUMB E R S AND NUMB E R OF C AL VE S
F OR E AC H B R E E D
N0. No. No. Mean Mean Mean age
of of of ADG, type of calf
Breed calves herds sires l b. / day scorea days
Angus 3190 39 121 1.829 11.626 195.86
Hereford 1648 16 55 1.812 11.534 199.37
a Faney--[-~-17, Fa nc yml 6, Fancy--~---15, etc.
ADG for different ages of the dam from 2
to 20 years.
Various approxi mat e tests of two- and
t hree-way interactions have been proposed
(Snedecor and Cox, 1935; Swiger, 1961). As
a prel i mi nary step, the two- and t hree-way
interactions among t he sex, age of dam and
year classes were pl ot t ed and examined. They
indicated no large interaction effects. An an-
alysis of variance was t hen comput ed for t he
model given. The nonorthogonal nat ure of
the dat a required the solution of t he least
squares equations, which amount ed to a set of
199 equations in the Angus breed. The size of
t he comput er, therefore, set a limit to the size
of the model which could be dealt with. These
equations were solved by part i t i oni ng and ab-
sorption. Analysis of variance for bot h breeds
for ADG is given in table 3.
All four classes of main effects were highly
significant sources of variation. The sex by
age-of-dam i nt eract i on was nonsignificant,
and t he sex by year interaction bordered on
significance at the 5% level. Bot h of these
i nt eract i on classes were t herefore dropped
from the model. The "r emai nder " sum of
squares, which was obtained as a difference
between the sum of squares due to the ul t i mat e
sex by age-of-dam by year by herd-sire sub-
classes and t hat due to the full model, was
used to test all interactions not specified in
t he model. Thi s proved significant, which is
not surprising, considering the number of
degrees of freedom involved. However, t he
small size of t he "r emai nder " mean squares
suggested t hat the model was a reasonable
one, and the analysis was pursued with the
model given, with the i nt eract i on classes omit-
t ed and t he herd-sire classification broken
down i nt o herds and sires-within-herds.
The met hod of analysis used was based on
Met hod I I of Henderson (1953). Thi s is a
two-stage procedure for the estimation of
variance component s in mixed model situa-
tions, in which t he first stage is the least
by guest on July 14, 2014 www.journalofanimalscience.org Downloaded from
184 CUNNI NGHAM AND HENDERS ON
T AB L E 3. ANALYS I S OF VAR I ANC E F OR AVE R AGE DAI L Y GAI N
An g u s
Sum of Mean
Sour ce d. f. s qua r e s s qua r e
He r e f o r d
S u m of Me a n
d. f. s qua r e s s qua r e
To t a l 3190 10946. 53
Sex 2 22. 97 11. 48" *
Age of d a m 6 12. 60 2. 10" *
Ye a r 11 1. 87 0 . 1 7 " *
Sex x age of d a m 12 0. 67 0 . 0 5
Sex x ye a r 22 1. 60 0 . 0 7 *
He r d- s i r e 120 89. 33 0. 74**
Ful l mo d e l 174 10807. 94 . .
Ul t i ma t e s ubcl as s es 1605 10882. 52
Re ma i n d e r 1431 74. 58 0. ' 05"*
Ec r or 1585 6 4 . 0 0 0 . 0 4
1648 5605. 97
2 11. 57 5 . 7 ; * *
6 4 . 8 7 0 . 8 1 " *
11 4. 53 0 . 4 1 " *
12 0 . 6 9 0 . 0 5
22 2. 63 0 . 1 1 "
54 59. 50 1. 10" *
108 5500. 78 . .
607 5538. 68
499 37. 89 0. 07" *
1041 67. 28 0 . 0 6
P < . 0 5 .
* * P < . O I .
squares est i mat i on of t he fixed effects and the
correction of t he dat a by subt ract i ng from
each observat i on an appr opr i at e linear func-
tion of these est i mat es.
Under cert ai n conditions the l east squares
est i mat es of the fixed effects can be i mproved.
Thi s possibility arises when the fixed classes
are unequal l y spread over the r andom classes,
since the difference between the class t ot al s
for the l at t er will t hen contain i nformat i on
on the former. I n the present case, for instance,
the different years are unequal l y represent ed
in all herds, and, therefore, the herd differences
have par t i al l y confounded with t hem i nforma-
tion on the year effects. Thi s i nformat i on can
be recovered and combi ned with the regul ar
least squares est i mat es in a j oi nt weighted
set of est i mat es, which will be an i mpr ovement
on those derived by regular least squares
met hods, provi ded t hat the weights are known
or can be est i mat ed wi t h sufficient accur acy
and precision. Thi s recovery is rel at i vel y
simple in bal anced dat a. I n unbal anced cases,
such as the present one, it is usual l y i mpos-
sible if conventional anal yses are used. Thi s
difficulty can be overcome by the appl i cat i on
of a comput at i onal procedure for the deri va-
tion of maxi mum likelihood estimates, which
applies in mixed model situations under very
general conditions (Henderson, 1952, 1963;
Henderson e t a l . , 1959). These est i mat es are,
in fact, aut omat i cal l y weighted l east squares
estimates. The weights used are vari ance
component s.
Fr om a combi nat i on of these techniques
an i t erat i ve procedure was developed in which
weighted l east squares est i mat i on of the fixed
effects in the model al t ernat es with vari ance
component est i mat i on. Thi s procedure is de-
scribed in detail by Cunni ngham (1962).
For the est i mat i on of component s of covari -
ance between the t rai t s ADG and TS, the
regul ar covari ance anal ogue of Met hod I I
(Henderson, 1953) was used. Est i mat es of
the heri t abi l i t y of each t rai t were const ruct ed
using the vari ance component est i mat es, and
the genetic, phenot ypi c and envi ronment al
correlations between the two t rai t s were com-
put ed from the est i mat es of the vari ance and
covari ance component s.
Re s ul t s and Di s c us s i o n
F i x e d E f f e c t s . The envi ronment al effects for
which est i mat es were obt ai ned were the t hree
sexes, seven ages of dam and 12 years. I n fact ,
these par amet er s were not i ndi vi dual l y esti-
mabl e, and t he quant i t i es est i mat ed were the
differences between par amet er s wi t hi n each
classification. These quant i t i es were est i mat ed
by regular l east squares, weighted l east
squares, and by the i t erat i ve est i mat i on proc-
ess. The i t erat i ve pr ocedur e was carri ed
t hrough four i t erat i ons in t he case of the
Angus dat a, and seven in the Her ef or d dat a,
at which poi nt the est i mat es appear ed to have
stabilized in each case.
The rel at i ve effectiveness of the different
set s of est i mat es as correction fact ors was
measur ed by using t hem t o adj ust t he dat a
and t hen compari ng t he equal i t y of means
wi t hi n sex, age of dam and year classes. Ther e
was little difference bet ween t he est i mat es ob-
t ai ned by the weighted l east squares and i t era-
tive procedures as measured by t he differences
between t he means wi t hi n classes. The regul ar
by guest on July 14, 2014 www.journalofanimalscience.org Downloaded from
E S T I MAT I NG WE ANI NG T R AI T S I N C AT T L E 185
l e a s t s quar es e s t i ma t e s wer e l ess ef f ect i ve as
c or r e c t i on f act or s. Of t he t hr ee me t hods of
e s t i ma t i on, t he i t e r a t i ve pr oc e dur e a ppe a r s
t o gi ve t he be s t e s t i ma t e s f or use as cor r ect i on
f act or s . Th e r e was evi dence t ha t t he va r i a nc e
of t he i t e r a t i ve e s t i ma t e s was l ess t ha n t h a t
f or t he r e gul a r a nd we i ght e d l e a s t s quar es
e s t i ma t e s .
Th e e s t i ma t e s obt a i ne d b y t he i t e r a t i ve pr o-
cedur e ar e pr e s e nt e d i n t a bl e 4. Th e sex effect s
a r e gi ven as de vi a t i ons f r om t he st eer effect ,
t he a ge - of - da m effect s as de vi a t i ons f r om t he
7- t o 10- year age gr oup effect a nd t he y e a r
effect s as de vi a t i ons f r om t he 1955 effect.
Thus , t he val ues f or sex a nd age of d a m ma y
be used, wi t h si gns r ever s ed, as cor r ect i on
f act or s t o a d j u s t ADG a nd TS val ues t o a
s t a n d a r d val ue r e pr e s e nt e d b y t he cal f f r om
a cow aged 7 t o 10 year s .
Th e p a t t e r n of t hese e s t i ma t e s c onf or ms
wi t h t h a t t o be f ound i n t he ext ens i ve l i t e r a -
t ur e. Th e pa r a l l e l a na l ys e s f or t he t wo
br e e ds agr ee wel l i n t he sex a nd a ge - of - da m
es t i mat es . Bul l cal ves gr ow f as t er t ha n st eer s,
a nd st eer s f as t er t ha n hei f er s. Bul l cal ves al s o
gr a de b e t t e r t ha n s t eer s or hei f er s, b u t hei f er s
gr a de c ons i de r a bl y b e t t e r t ha n st eer s. Th e ob-
TABLE 4. ESTIMATES OF ENVIRONMENTAL
PARAMETERS OBTAINED BY ITERATIVE
ESTIMATION PROCEDURE
Angus Hereford
Item ADG TS ADG TS
Sex
Bull 0.108 0. 634 0.113 0. 499
Heifer - - . 142 0.532 - - . 107 0.307
Steer 0.000 0.000 0. 000 0. 000
Age of dam
2 --. 261 --. 497 - - . 264 - - . 474
3 --. 159 --. 063 --. 192 - - . 288
4 --. 069 0.106 --. 081 0.146
5 - - . 036 0.056 --. 027 0.212
6 --. 025 0.059 - - . 010 0..127
7-10 0.000 0.000 0.000 0.000
~10 - - . 029 --. 565 0.022 - - . 090
Year
1949 0.085 1.350 0.046 0.491
1950 0.067 0.587 0.020 0.391
1951 0.082 0.674 0.003 0.910
1952 0.055 0.271 0.024 0.254
1953 0.005 0.408 - - . 009 0.556
1954 0.013 --. 205 0.113 0.673
1955 0.000 0.000 0.000 0.000
1956 - - . 010 - - . 030 0.031 0.728
1957 --. 088 --. 244 - - . 270 --. 628
1958 - - . 078 --. 291 --. 082 0.360
1959 --. 051 - - . 408 --. 157 --1. 270
1960 --. 001 --. 192 --. 057 --. 445
s er ved di f f er ences be t we e n bul l s a nd s t eer s a r e
i n p a r t a t t r i b u t a b l e t o t he effect s of sel ect i on,
i n t h a t br e e de r s t end t o s ave t he heavi er ,
hi gher g r a d i n g cal ves as bul l s a nd t o c a s t r a t e
t he r e ma i nde r . Th e we i ght e d me a n of t he mal e
cal ves f or ADG was 0. 158 ( Angus ) a nd 0. 160
( He r e f o r d ) hi gher t h a n t he me a n for f emal es ,
a nd f or TS was 0. 454 ( Angus ) a nd 0. 153
( He r e f o r d ) bel ow t he f emal e mean. ADG i m-
pr oves wi t h age of d a m i n a c ur vi l i ne a r f as hi on
t o a ma x i mu m a t 7 t o 10 ye a r s or over . Th e
s ame p a t t e r n hol ds t r ue f or TS, except t h a t
t he val ues t end t o r e a c h a ma x i mu m a t ear l i er
ages ( 4 t o 5 y e a r s ) a nd t o decl i ne a f t e r t h a t
age. Th e effect of s el ect i on a mong cows was
i gnor ed i n t he c o mp u t a t i o n of t hese age- of -
d a m effect s, whi ch ma y be bi a s e d u p wa r d as
a r es ul t ( Lus h a nd Shr ode, 1950) . Th e si gni f i -
cance of t hes e pos s i bl e bi as es i n beef c a t t l e
t r a i t s is di s cus s ed b y Ko c h a nd Cl a r k ( 1 9 5 5 a ) .
A s l i ght br e e d di f f er ence does show up i n
t he e s t i ma t e s of t he y e a r effect s. Th e Angus
d a t a a p p e a r t o be l ess va r i a bl e t h a n t he
He r e f or d. Th i s i s p r o b a b l y a r ef l ect i on of
di f f er ences i n t he a ve r a ge he r d e nvi r onme nt
for t he t wo br eeds . Th e decl i ne i n ADG over
t he pe r i od s t udi e d, whi ch is i ndi c a t e d b y t he
e s t i ma t e s of t he y e a r effect s, ma y be due t o
t he g r a d u a l i ncl us i on i n t he p r o g r a m i n t he
l a t e r ye a r s of her ds wi t h poor er e nvi r onme nt
or genet i c ma t e r i a l .
Components o] Variance and Covariance.
Th e e s t i ma t i on of c ompone nt s of va r i a nc e f or
bot h t r a i t s was c a r r i e d out on a wi t hi n- br e e d
basi s, usi ng modi f i c a t i ons of Me t h o d I I ( He n -
der s on, 1953) . Th e d a t a wer e f i r st c or r e c t e d
for sex, a ge - of - da m a nd y e a r effect s. Me a n
s quar es wer e t hen c o mp u t e d f or her ds , si r es,
her ds b y ye a r s a nd er r or . Fi n a l l y , t hes e me a n
s quar es wer e e qua t e d t o t hei r expect ed val ues
a nd t he r es ul t i ng e qua t i ons wer e s ol ved f or
t he r e qui r e d va r i a nc e c ompone nt s . Th e pr o-
por t i on of t he va r i a t i on i n t he a d j u s t e d r ecor ds
whi ch was due t o each of t hese sour ces i s i n-
di c a t e d i n t a bl e 5. Th e b r e a k d o wn of t he va r i -
a t i on is s i mi l ar for b o t h t r a i t s a nd bot h br eeds .
Th e f act t h a t a gr e a t e r p r o p o r t i o n of t he
va r i a nc e f or TS is i n r es i dual a nd l ess i n he r ds
ma y r ef l ect t he s ubj e c t i ve n a t u r e of t he t y p e
s cor i ng pr oc e dur e , or pos s i bl y t h a t he r d en-
vi r onme nt has a gr e a t e r effect on ADG t ha n
on TS.
Th e a c t u a l e s t i ma t e s of t he va r i a nc e a nd
c ova r i a nc e Component s ne e de d i n t he c ompu-
t a t i on of he r i t a bi l i t i e s a nd c or r e l a t i ons ar e
by guest on July 14, 2014 www.journalofanimalscience.org Downloaded from
186
TABLE 5. PERCENT OF VARIATION IN SEX,
AGE OF DAM AND YEAR ADJUSTED
RECORDS WHICH IS DUE TO
VARIOUS SOURCES
Angus
Source ADG TS
CUNNI NGHAM AND HENDERSON
herd heritability requires, in addition, the as-
sumption that all herd-to-herd variation is
nongenetic. The values for both traits in each
breed are given in table 7.
Hereford These values fall within the range of those
ADG TS reported elsewhere (Warwick, 1958), but
are considerably larger than most of the esti-
15 3 mates in the literature, which indicate a value
10 11 close to 0.3 for ADG and 0.2 for TS. It seems
3 5
72 81 probable that the present values are overesti-
mates resulting from biases in the variance
component estimates. This is borne out by
the values of 0.48 and 0.28 obtained for the
repeatability of these traits in the same data
(Cunningham and Henderson, 1965).
Several factors may contribute to the infla-
tion of the heritability estimates. The denom-
inator may be reduced due to underestimation
of ~r2e, resulting from correlated errors. Since
a2e is the within-sire-herd-subclass variance,
TABLE 7. ESTIMATES. OF THE HERITABILITY
OF ADG AND TS
Herds 18 9
Sires 11 11
Herds x years 9 6
Residual 62 74
given in table 6. The necessary components
are:
a2n=her d component of variance
~2s/~i=sire-within-herd component of vari-
ance
a2e=-error component of variance
(ri2s/R= sire-within-herd component of co-
variance
alze=error component of covariance
The values obtained separately for the two
breeds show good agreement except that the
herd component of variance is considerably
larger for the Angus than for the Hereford
data. The greater inherent variability of the
Hereford records is reflected in the larger error
components of variance and covariance for
that breed.
Heritabilities. The heritability of each trait
was computed from the variance component
estimates on both an intra- and inter-herd
basis, using the formulas
4 0 " 2 8 / H 4 O ' 2 S / H
h 2 i n t r a - a n d h 2 1 n t e r ~ '
~ - b ~ ' ~ 9 ~ -t- ~
T h e s e formulas require the following as-
sumptions: (1) that identifiable environmental
variation has been corrected for, (2) that
e p i s t a t i c gene action may be neglected, (3)
that there is random mating with respect to
the genes involved in this trait, (4) that the
frequencies of these g e n e s a r e stable, and
(5) that the effects of these genes, as devia-
tions from the mean, do not change from one
generation to the next. The formula for inter-
TABLE 6. ESTIMATES OF COMPONENTS OF
VARIANCE AND COVARIANCE FOR
ADG AND TS
B r e e d T r a i t ~2H a ~ s / H ~r2e a12 S/ H ~12 e
A D G 0 . 0 1 2 0 0 . 0 0 7 0 0 . 0 4 0 4
A n g u s T S 0 . 2 9 4 9 0 . 3 6 9 5 2 . 3 9 6 4 0 . 0 4 6 7 0 . 0 6 2 4
A D G 0 . 0 1 3 0 0 . 0 0 9 2 0 . 0 6 4 6
H e r e f o r d T S 0 0 9 1 8 0 . 3 5 2 7 2 . 5 5 7 1 0 . 0 3 0 4 0 ' . 1 4 7 0
Heritability estimate
Breed Trait Intra-herd Inter-herd
Angus ADG 0. 590 0. 471
TS 0. 534 0.483
Hereford # 1-)G 0.400 0.425
TS 0.485 0.410
the fact that observations within such a sub-
class may be repeated records on the same
dam, or records on closely related dams would
reduce the estimate. It could also be reduced,
and the estimate of sire variance simultan-
eously increased by any differential or assorta-
tive mating, i.e., by mating the better cows
with one sire, the poorer with another. Though
the records were corrected for age of dam,
differential mating of dam might have a similar
effect. Furthermore, the fact that sires were
partially compounded with years would have
an inflationary effect on the sire variance if
an interaction of sires with years existed and
was ignored in the analysis. All these factors
tend to bias the ratio
O- 2~ ,TT
O ' 2 S / H + 0 ~ e
upward. This ratio is multiplied by four to
give heritability, and hence even a moderate
bias in the variance components would result
in a large bias in the heritability.
An alternative interpretation of these high
heritability estimates is that there exists con-
by guest on July 14, 2014 www.journalofanimalscience.org Downloaded from
ESTI MATI NG WEANI NG TRAI TS I N CATTLE 187
TABLE 8. ESTIMATES OF CORRELATIONS
BETWEEN ADG AND TS
Correlation Angus Hereford
Genetic 0.61 0.36
Phenotypic 0.28 0.36
Environmental --. 95 0.37
si derabl y more additive genetic vari at i on for
these t rai t s in New York commercial beef
herds. There may be some t rut h in this, be-
cause all the reports from elsewhere are based
on few and large herds, often at experi ment
stations, i n which selection may have reduced
the genetic vari abi l i t y.
Correlations. The genetic, phenot ypi c and
envi ronment al correlations bet ween the two
t rai t s were comput ed from the vari ance and
covariance component estimates of table 7.
The values obt ai ned are given i n t abl e 8.
The estimates agree wi t h some of those
reported in other studies (Koch and Clark,
1955b; Lehmann et al . , 1961). However,
the large discrepancy between the values for
the two breeds reduces the val ue of these
estimates. Furt hermore the sampl i ng vari ance
of the estimates is probabl y quite large, and
the possible biases affecting the heri t abi l i t y
estimates may also influence these correlations.
There appears to be a positive and fairly large
genetic relationship between these two traits.
Summa r y
The genetic and phenot ypi c paramet ers of
average dai l y gai n from bi rt h to weani ng
( ADG) and t ype score at weani ng (TS) have
been eval uat ed from 4,838 records on beef
cattle i n New York. The records were drawn
from 39 herds wi t h 121 sires of the Angus
breed and from 16 herds with 55 sires of
the Hereford breed for the years 1949 to 1960
inclusive.
The dat a were tested for the presence of
i nt eract i on effects. Est i mat es were obt ai ned
by an iterative procedure of the effects of sex,
age-of-dam and year differences on the two
traits. The records were adj ust ed for these
fixed effects and component s of vari ance and
covariance were est i mat ed for herds, sires,
herds by years, and error. These component s
were used to comput e estimates of the herit-
abi l i t y of each t rai t and of the genetic, pheno-
typic and envi ronment al correlations bet ween
them.
Li t e r a t ur e Ci t ed
Cunningham, E. P. 1962. Estimation of genetic and
phenotypic parameters in a beef cattle population.
Ph.D. Thesis. Cornell University, Ithaca, New
York.
Cunningham, E. P. and C. R. Henderson. 1965.
Repeatability of weaning traits in beef cattle. J.
Animal Sci. 24:188.
Henderson, C. R. 1952. Specific and general combin-
ing ability. In Heterosis (J. W. Gowen, Editor).
Iowa State College Press, Ames, Iowa.
Henderson, C. R. 1953. Estimation of variance and
covariance components. Biometrics 9:226.
Henderson, C. R. 1963. Selection index and expected
genetic advance. Statistical genetics and plant
breeding. National Academy of Sciences--National
Research Council publication No. 982.
Henderson, C. R., O. Kempthorne, S. R. Searle and
C. M. von Krosigk. 1959. The estimation of envi-
ronmental and genetic trends from records subject
to culling. Biometrics 15:192.
Koch, R. M. and R. T. Clark. 1955a. Influence of
sex, season of birth and age of dam on economic
traits in range beef cattle. J. Animal Sci. 14:386.
Koch, R. M. and R. T. Clark. 1955b. Genetic and
environmental relationships among economic char-
acters in beef cattle. I. Correlation among paternal
and maternal half-sibs. J. Animal Sci. 14:775.
Lehmann, R. P., J. A. Gaines, R. C. Carter, K. P.
Bovard and C. M. Kincaid. 1961. Selection indexes
for weanling traits in beef calves. J. Animal Sci.
20:53.
Lush, Jay L. and Robert R. Shrode. 1950. Changes
in milk production with age and milking fre-
quency. J. Dairy Sci. 33:338.
Snedecor, G. W. and Gertude M. Cox. 1935. Dis-
proportionate subclass numbers in tables of mul-
tiple classification. Iowa Agr. Exp. Sta. Res. Bul.
180.
Swiger, L. A. 1961. Genetic and environmental in-
fluences on gain of beef cattle during various
periods of life. J. Animal Sci. 20:183.
Warwick, E. J. 1958. Fifty years of progress in
breeding beef cattle. J. Animal Sci. 17:922.
by guest on July 14, 2014 www.journalofanimalscience.org Downloaded from