ttERITABILITY

ESTIMATES OF GESTATION LENGTH AND BIRTH

WEIGHT IN HOLSTEIN-FRIESIAN
CATTLE AND THEIR
U S E I N S E L E C T I O N : I N D E X E S 1, 2
MOGENS PLUM, tIENRY ANDERSEN, '~ AND L. A. SWIGER 4
Departments of Dairy Science and Animal Science, ~Jniversity of Nebraska, Lincoln
ABSTRACT

I:[eritabilities of gestation length and birth weight of 958 Holstein calves

were 0.60 0.12 and 0.34 ~ 0.09, respectively, after the data were adjusted
for the effects of herd, sex, paYity, year, and season. The least-squares means
for birth weight and gestation length were 93.5 lb and 277.4 days, respectively.
The correlations between birth weight (BW) and gestation length (GL) were:
phenotypic, 0.43; genotypic, 0.37; and environmental, 0.52. The index for
selection for shorter gestation length is I = --GL + 0.10 BW and the index
for selection for lower birth weight is I = - - B W -- 0.14 GL.

A recent review of literature about gestation

length and birth weight in cattle and buffaloes
lists 21 references dealing with studies of gestation length in Holstein-Friesian cattle (1). I n
these 21 studies, the average gestation lengths
varied from 276 to 282 days. Since it is generally recommended not to rebreed cows until
at least 60 days post-partum, and since many
cows require more than one breeding for conception, the calving interval is often more than
12 months. From an economic standpoint, it
would be an advantage to have cows calve at
intervals of 12 months. One method to decrease
the length of the calving interval would be to
shorten the length of the gestation period. At
the same time, considerable interest is being
shown in obtaining smaller calves and a great
many dai~5, heifers are presently being bred to
beef bulls, to achieve this.
Since calf weight and gestation length are
correlated, it is important to know what effect
the shortening of the gestation length would
have on birth weight and vice versa.
Heritability estimates by several authors

range frmn zero to 0.71 for gestation length,

with the majority falling between 0.25 and 0.50
and from 0.22 to 0.51 for birth weight, with
most of the values falling in the 0.25 to 0.35
range (1). Thus, previous investigations indicate that the heritabilities are high enough to
make phenotypic selection for these characters
effective.
SOURCE OF DATA
The data came from the herds of Holstein
cattle maintained at the University of Nebraska's
North Platte and Scotts Bluff Experiment Stations. They consist of 958 gestation periods
and birth weights of single calves born alive
from 1947 to 1964. These calves were sired by
60 different bulls. The length of the gestation
period was the number of days from last service
to and including the day of calving but not
including the date of last selwice. Newborn
calves were weighed as soon after birth as possible and weights were recorded to the nearest
pound.
A N A L Y S I S OF DATA AND R E S U L T S

Adjustment o.f data. The data were first ad1%ceived for publication June 11, 1965.
:Published with the approval of the Director as
paper no. 1755, Journal Series, Nebraska Agricultural Experiment Station. This is a contribution
from Regional Project NC-2 Improvement of
Dairy Cattle Through :Breeding.
s Part of the data taken from a thesis presented
to the Graduate College by Henry Andersen in
partial fulfillment of the requirements for the
degree, Master of Science.
SPresent address: Institute for Artificial Insemination, Royal Veteri~xary mxd Agrizultural
College, Copenhagen, Demnark.
Present address: Department of Animal Science, The Ohio State University, Columbus, Ohio.

justed for the effects of herd, season, sex,

parity, and year. To separate these effects, the
data were analyzed by the least-squares method
described by Henderson (3) as Method 2 for a
mixed model.
Constants were fitted for the fixed main
effects: sex, year, season, parity, and herd,
with the random sire effects considered simultaneously. All interactions were ignored in
fitting these constants. Calves in Herd 1 had
0.96 day shorter gestation period and weighed
6.2 lb less at birth than calves in Herd 2. Nonsignificant effects of year ranged from --4.30
to +3.46 lb for birth weight and from --2.61

167~

ttERITABILITY ESTISIATE S
to + 2 . 5 6 days f o r gestation length. Effects of
parity, month of birth, and sex are given in
Table 1. The data were adjusted f o r the fixed
effects: herd, year, sex, parity, and season.
TABLE 1
Least-squares constants giving the effects of parity,
month of calving, and sex on gestation length
(t~ = 277.4 days) and birth weight
(~ = 93.5 lb)
Effect
Gestation
length

Birth
weight

(days)
+1

(lb )
--3

--1
+1
+1
+1
+3
--6
--1
+1
--1
+1
--2
0
--2
0
+1
0
+2
+1
+0.5
--0.5

+1
+1
0
0
+7
--6
+2
+4
+6
+1
--1
--2
--3
--3
--1
--1
--2
0
+4
--4

1
Parity

Month

Sex

2
3
4
5
6
7
January
February
March
April
May
June
July
August
September
October
November
December
Male
Female

The mean adjusted birth weight was 93.5

0.34 lb and the mean adjusted gestation length
was 277.4 0.22 days.

1673

ucts of the adjusted data to their expectations

according to Henderson (3). Table 2 gives the
mean squares and products, with their expectations. Table 3 lists the sire and half-sib components. The genetic component is computed
as the sire component divided by the average
relationship between paternal half-sibs. U n d e r
random mating this relationship would be 0.25,
but a sample of some sib relationships indicated that the relationship between paternal
half-sibs was closer to 0.29. Consequently, the
genetic component was computed as (Sire component)/0.29. This method of computation also
assumes that there are no epistatic effects, no
environmental correlations among a sire's progeny, and no covarianee between genotype and
environment. Only a little of the epistatic variance would be in the sire component, even i f
important. The design of the experiment was
such that the last two assumptions should be
met. The genetic component subtracted from
the total or phenotypic component leaves a remainder classified as the environmental component. These components are given in Table 3.
The heritability estimates of gestation length
and birth weight considered as characteristics
of the calf were computed as follows : h ~ = (Genetic c o m p o n e n t ) / ( P h e n o t y p i e
component).
These heritabilities are 0.60 0.12 for gestation length and 0.34 0.09 for birth weight.
The standard errors of the heritabilities were
computed according to the method given by
Swiger et al. (5) as S-(vo~) where
~t~ (1-- t)[l+
(k--1)t][2(N--1)]~/

~[ (~

s)(s -

1)]~

and N ---- total number of animals

t : intrasire correlation
lc = coefficient of sire component in sire
mean square
s = number of paternal half-sib groups.

Her#ability estimates of gestation length and

birth weight and the correlatio~ between gestatio~ length and birth weight. The components
of variance for sires and half-sibs were computed by equating the mean squares and prod-

TABLE 2
Analysis of variance and covariance for gestation length (GL) and birth weight (BW)
Mean squares and products
Source

d/f

GL

BW

GL BW

Sire
tta]f-sibs

59
898

166.38
38.02

391.83
142.13

98.99
32.76

Expectations
1.03z~ + 15.76~,,
ah~

TABLE 3
I-Ieritabilities and components of variance and covarlance for gestation length (GL) and
birth weight (BW)
Components
Trait
GL
BW
GL BW

Sire
8.07
15.57
4.14

I-Ialf-sib
38.02
142.13
32.76

Phenotypic
46.09
157.70
36.90

Genetic

Environmental

Heritability

27.83
53.69
14.28

18.26
104.01
22.62

0.60
0.34

1674

~ . P L U M , ]~1. A N D E R S E N , A N D L. A. S W I G E R

The phenotypic, genetic, and environmental correlations between gestation length and birth
weight were computed from the appropriate
components of variance and covariance given
in Table 3. The genetic correlation, for example, is 14.28/[ (53.69) (27.83) ] ~i. The values
for the correlations were rpp ~ 0.43, r~g ----0.37,
and r~, ----0.52.
APPLICATION

The principle of a selection index has been

described by Hazel (2). The purpose of a
selection index may be either to combine several
traits which all have some value into a composite net merit, or to use several correlated
traits to select for only one character. The latter
situation is found when the goal is to shorten
the length of the gestation period, and birth
weight is used as a secondary but correlated
character.
For a selection index where net merit is a
function of several traits, the weights for the
various traits are determined by the solution of
a set of simultaneous equations. The weights
are those that minimize the sum of the squares
of the differences between the indexes and the genetic values for net meri~ (6). I f P , P~,..., P~
refer to the phenotypes of the m characters and
G~M is the genotype of net merit, the b values
or weights to be given to characters 1 to m
are found by the solution of the following
equations :
blo'2pl +

b~a'.ole 2 -~ . . .

bto-vlP2 -~-

b:~r2e2 _1_

+ b.,~r~.~z.~ :

O'e~O~M

- _[_ bm-~'2Pm --- O'P2eNM

it is assumed that environmental effects

are independent of genetic effects, the covarianee between the phenotype of one character
and the genotype of another becomes the covariance between the genotypes of the two
characters. In the present case, net merit ( N M )
is the same as gestation length (Character 1)
and birth weight is a secondalT character (Character 2), which contains information about gestation length. Then (rP~oNMis equal to (r%~ and
~P2~M is equal to (r~G~. I f selection is for gestation length and only this one character is considered, there is one equation b~o:-P~ = cr-'~mand
the b value becomes the heritability o-~l/(r~ of
gestation length, or 0.60.
I f birth weight is included in the index, it is
necessary to solve two equations:
46.09 b~ + 36.90 b~ ----27.83
36.90 b~ + 157.70 b.~ ~-- 14.28
When

The solution of these equations gives an index

for shortening the gestation length
IaL : --0.6538 G L + 0 . 0 6 2 4 B W , or
I ~ , : - - G L + 0.10 BW.

Presently, many Holstein heifers are bred to

beef bulls to obtain smaller calves and have
easier parturition. I f it were the purpose to
select for lower birth weight, then birth weight
would be the primary character and gestation
length might be used as a secondary character.
I n this case, the b values are found from the
following equations :
157.70 b~ + 36.90 b~ ~ 53.69
36.90 b~ + 46.09 b~ =- 14.28,
giving an index for selection for lower birth
weight
I , , . ---- --0.3297 B W -- 0.0459, or
IBw =- - - B W - - 0.14 GL.
EXPECTED G E N E T I C C H A N G E S

The expected genetic change in a character j

per unit change in an index I(A Gffunit I) is
given by Morley (4) to be
Xsbj (rc~/(r~1~ where
Using the values from the index for gestation
length, the change in gestation length per unit
change in the index is
(0.6538) (27.83) -- (0.0624) (14.28)/(r~x
and the change in birth weight becomes
(0.6538) (14.28) -- (0.0624) (53.69)/(r-~t.
Multiplying these figures by ,r~, the following
values are obtained in standard units:
Change in gestation length----4.16(i) days
Change in birth weight
-= 1.44(i) pounds,
where i is the standardized selection differential
or selection intensity.
I f selection is carried out for gestation length
without considering the birth weight, the following values are obtained:
Change in gestation length =
4.10(i) days per generation
Change in birth weight ---2.10(i) pounds per generation.
Including birth weight adds very little to the
effectiveness of selection for gestation length,
but it does make the change in birth weight
smaller.
Table 4 gives the expected changes in gestation length and birth weight under different
intensities of selection, using the indexes for
either shorter gestation length or lower birth
weight. The selection index for gestation length
differs from the selection index for birth weight,
in that the first index has opposite signs for
the two characters, whereas the second index
has the same signs for the two characters.
The explanation for the coefficients of th-

HEI~ITABILITY ESTIMATES

1675

TABLE
4
Expected changes per generation in gestation ]ength and birth weight when selecting for
1) shorter gestation length an(] 2) ]ower birth weight using the appropriate index
Proportion selected
for breeding
Males

Females

Intensity
of
selection

90
70
50
90
70
50
90
70
50

(i)
0.80
0.95
1.10
0.98
1.13
1.28
1.13
1.28
1.43

- - ( % )
20
20
20
10
10
10
5
5
5

Reduction where goal is

Short GL
GL
BW

(days)
3.3
4.0
4.6
4.1
4.7
5.3
4.7
5.3
5.9

secondary t r a i t s being opposite in sign in the

two indexes is as follows: L e t I r e f e r to the
p r i m a r y t r a i t a n d 2 to t h e s e c o n d a r y trait.
Then
b~ : -- (o-z~p2o-~)/i) + (o-~'p~o-~)//D,
where D is the d e t e r m i n a n t , or

(lb)
1.2
1.4
1.6
1.4
1.6
1.8
1.6
1.8
2.0

(days)
1.1
1.3
1.5
1.4
1.6
1.8
1.6
1.8
2.0

REFERENCES

(2)

(3)

O-~G1 )
O"P1

(4)

or

T h a t is, b~ will be n e g a t i v e w h e n the h e r i t a b i l i t y

of T r a i t 1 is l a r g e r t h a n the coheritability a n d
positive when the h e r i t a b i ] i t y is smaller t h a n
the e o h e r i t a b i l i t y a n d when b o t h eovariances
are positive.
W h e r e the genetic a n d p h e n o t y p i e covariances
are b o t h negative, the o p p o s i t e is true. W h e n
the genetic covariance is negative a n d the p h e n o t y p i c eovariance positive, b_~ will be n e g a t i v e ;
a n d when the genetic covarianee is positive a n d

(lb)
3.4
4.1
4.7
4.2
4.8
5.5
4.8
5.5
6.1

(1) ANDEI~SEN, HENRY, AND FLU~I, MOGENS. 1965.

2
O'PIP2O- G1 ~ O"P10-G1G2

O-G1G2
O'PIP2

GL

the p h e n o t y p i e eovariance negative, b_~ will be

positive.

. % ~ % - (~1~)~.
W h e n b o t h genetic a n d p h e n o t y p i c covarianees are positive, b2 will be n e g a t i v e w h e n

Low BW
BW

(5)

(6)

Gestation Length and Birth Weight in

Cattle and Buffaloes: A Review. J. Dairy
Sci., 48: ]224.
HAZEL, L. iN. 1943. The Genetic Basis for
Constructing Selection Indexes. Genetics,
28 : 476.
HENDERSON, 1~. C. 1953. Estimation of Variance and Covariance Components. Biometrics, 9: 226.
~vioRr~, F. H. W. 1950. Selection for Economic Characters in Merino Sheep. Unpublished Ph.D. thesis, Iowa State University
Library, Ames.
S~VIGER, L. A., ttAaVEY, W. R., EVEaSON,
D. O., AND GaEOORY, K. E. 1964. The
Variance of Intraelass Correlation Involving Groups with One Observation. Biometrics, 20: 818.
Swmna, L. A., AND I-IAZEL, L. N. 1961. Optimum Length of Feeding Period in Selecting
for Gain of Beef Cattle. J. Animal Sci.,
20 : 189.