Selection Procedures for Poultry Flocks with Many Hatches
HANS ABPLANALP
Department of Poultry Husbandry, University of California, Davis
(Received for publication May 31, 1956)
INTRODUCTION
T HE main purpose of the present
study was to determine the effect of
differences in duration of hatching season
upon variation in production traits of
chickens. The accuracy of estimating ge-
netic worth of breeding animals and,
hence, the efficiency of selection decisions
depends among other factors on family
size, which in turn depends on the number
of hatches and thus on the duration of the
hatching season. It is conceivable that an
extended hatching season might, because
of purely environmental causes or inter-
action between genotypes and environ-
ment, lead to an increase in variation
among full sisters. Such added sources of
nongenetic variation would tend to coun-
teract the anticipated advantages of large
families.
Some of the problems pertaining to
family size have been discussed by Lerner
(1950) and by Lerner and Dempster
(1951). In a more recent study King and
Henderson (1954) have given quantitative
estimates of variation introduced by hatch
effects. They attempted to estimate the
bearing of hatch effects on the relative
efficiency of mass selection. However, the
assumptions underlying their study did
not involve a dependence of environ-
mental variability on family size as pre-
sumed by Lerner (1950). Furthermore, it
remains to be seen what effect hatch-to-
hatch differences may have when joint
family and individual performance is con-
sidered. A possible model that would lead
to increases in nongenetic variability with
extension of hatching season, can be
postulated by assuming some trend in
environmental conditions to which indi-
1285
viduals of successive hatches would be
exposed. Sexual maturity is an example of
a trait that depends on hatching date
(among others see Osborne, 1954). Thus,
if the end of a laying test is fixed for a
given date, individuals from successive
hatches will have decreasing periods of
time in which to complete the test. Under
those circumstances all hatch effects can
be excluded from comparisons of indi-
viduals, and therefore need not be con-
sidered as uncontrollable environmental
effects.
The problem of appropriate analyses of
such complex breeding experiments has
been worked out theoretically (see for
instance Henderson, 1953), but few ap-
plications to poultry have been published.
Since the amount of work required for an
appropriate statistical analysis of breed-
ing records may often be too great for
efficient operation, it is desirable to de-
termine by how much the accuracy of se-
lection decisions is reduced, when short
cuts are used. Thus it might be asked in
the present case whether it is necessary to
adjust individual production records for
date of hatching. It is also of interest to
know whether families represented by off-
spring of only early or late hatches are
given differential advantages and how im-
portant such biased family averages might
be for the over-all accuracy of selection
decisions.
Besides the main effects due to hatch
environment one may postulate interac-
tions between hatch environment and
genotypes. Their existence would lead to
changes in the relative ranking of families
under varied environmental conditions.
Environment-genotype interaction may
1286 H. ABPLANALP

be visualized by postulating the response dams that were represented by female off-
of a genotype to a particular environment spring in at least half of the hatches of a
as the sum of a general reaction that will given year.
rank the genotypes identically under all Data for the present study consisted of
conditions, plus a specific response elicited egg records of four populations of pullets
by the particular environment. If such hatched in the years 1947 to 1950. The
interactions exist selection will produce birds hatched in 1949 were offspring of the
permanent gains only on the basis of the 1947 population (since only two-year-old
general genotype. Adaptations of geno- dams were used in this analysis). The
types to specific conditions would, on the parents used for the 1950 hatches included
other hand, have to be considered as un- hens from the 1947, the 1948 and the 1949
controllable obstacles to accurate selec- populations.
tion for general performance. Gowe and Selection and propagation of the breed-
Wakely (1954) have published results that ing stock were subject to the following re-
tend to de-emphasize the importance of strictions:
genotype-environment interaction in pro- 1. All parents were selected when they
duction traits of chickens. Similar con- were almost two years old on the basis of
clusions may be drawn from an experi- their egg records. Other criteria, such as
ment in which the resistance of chickens egg weight, mortality of relatives, and
to leukosis was tested under conditions of production of relatives, were used as addi-
different experiment stations (King et al., tional evidence of relative merit.
1952). So far, the problem of interaction 2. The numbers of parents were rela-
in successive hatches of the same popula- tively low. Each male was mated to about
tion has not been studied, and most work- 2-8 females answering the requirement of
ers in the field have implicitly assumed representation in at least half of each
that they are nonexistent. year's hatches. Table 1 shows the numbers
of parents and offspring included in this
MATERIAL analysis.
The data used in this study were ob- 3. In some years a few of the male par-
tained from Kimber Farms, Inc. of Niles, ents were full brothers; likewise, there
California, representing production rec- were several full sisters among female
ords of a strain of SCWL chickens propa- parents. None of the latter were mated to
gated as a closed flock with emphasis on the same sire.
extremely large families. With minor ex- 4. The females were assigned as mates
ceptions, a period of about 20 weeks was to each sire for the entire period of hatch-
allowed in this line for the hatching of off- ing. Hence, no female was mated to more
spring from parental matings. than one male. The resulting offspring
Under these conditions the number of samples were thus divided into subgroups
female offspring produced by a hen in any according to a nested mode of classifica-
one week would vary according to her re- tion.
productive fitness and the chance dis- 5. All eggs were set at weekly intervals,
tribution of sex among the progeny. Some and individually pedigreed offspring used
dams represented only during a small in this study were hatched over periods of
segment of the hatching season could be 11 to 17 weeks (see Table 1).
expected to produce but little evidence 6. Female offspring were reared and
pertinent to this study. Consequently, the tested in floor pens; in the groups analyzed
analysis included only progeny from such none were eliminated during the 72 weeks
 SELECTION PROCEDURES 1287

TABLE 1.—Numbers of hens representing the four populations

Number of Number of 9
Year of
hatch Hatches Sires Dams Offspring dam per hatch

1947 15 7 17 427 1.67

1948 16 9 26 624 1.50
1949 17 8 31 827 1.57
1950 11 16 63 769 - 1.11

Combined populations 40 137 2,647 1.46

of each test, except by natural death or
accidents.
7. Each hatch of pullets was reared in a
separate group of pens. Four measure-
ments of production characteristics were
used in this study, namely:
(a) Average weight of eggs laid in March of the
first laying year, based on trap-nest records
4 days a week.
(b) Age at first egg.
(c) Winter production of eggs as determined by
trap-nest records 4 days a week, during a
period of 18 weeks preceding February 1 of
the first laying year.
(d) Production of eggs through 72 weeks of age,
measured by trapping 4 days a week.
Mortality and accidents eliminated
some birds before or during their produc-
tive life, leaving a number of egg records
incomplete. Egg-weight records were miss-
ing for the same reasons or because certain
pullets failed to lay eggs during the month
of March. For this study all birds with
missing egg records were excluded from
the analysis. Hence, only pullets that sur-
vived to March, laid eggs during that
time, and also had a record of age at first
egg were considered. Mortality after
March was not cause for excluding birds
from the analysis.
PROPERTIES OF THE POPULATIONS
Average performance of the four popu-
lations is shown in Table 2, for each trait
and each year separately. It may be noted
that egg-production averages increased
from 1947 to 1949 (presumably a result of

TABLE 2.—Coefficients of linear regression of egg weight and production traits on date of hatch

Population Mean Coefficient of Variance due Error

Production trait regression, b to regression
(Year) variance

Egg weight 1947 24.91 - .025 3.98 1.84

(oz. per doz.) 1948 24.63 - .080* 74.59 1.73
1949 24.76 - .079* 110.73 1.83
1950 24.90 .099* 52.14 1.80

Winter egg production 1947 35.7 - .326* 695.45 167.80

(4 days per week) 1948 37.4 -1.004* 11,616.52 149.44
1949 41.0 -2.110* 79,249.94 91.01
1950 40.8 .366* 708.73 166.35

Production to 72 weeks 1947 112.3 -1.127* 8,333.31 587.34

(4 days per week) 1948 114.3 -1.161* 15,552.84 827.97
1949 124.7 -1.134* 22,876.11 729.01
1950 117.3 - .024 2.97 616.16

Age at first egg 1947 26.69 .553* 2,003.28 12.42

(weeks) 1948 25.91 .323* 1,200.95 16.38
1949 25.44 .241* 1,030.80 6.72
1950 25.52 .217* 247.67 5.64

* Significant at the one percent level.

1288 H. ABPLANALP

selection), but dropped for the 1950 popu- variation other than hatch effects; they
lation. Figure 1 shows the dependence of are given in Table 2. All details of the
production traits on the date of hatching. analysis are discussed in the following sec-
Unbiased estimates of linear regressions tion. Most of the postulated linear regres-
were made by excluding all sources of sions can be considered significantly dif-
ferent from zero at the one percent level
PHENOTYPE AVERAGES AS DEPENDENT ON DATE OF NATCH except egg weight for 1947 and 72-week
production for 1950.
From the preliminary analysis of hatch
\ effects it appears that at least two types of
\
effects must be assumed to influence aver-
age performance of hatches:
1. Systematic effects, presumably due
II 13 IS
WEEK OF YEAR to progressive change in the date of hatch-
1947 ing. Such trends may result from the de-
1948
creasing length of testing periods when
egg records are terminated at a set time of
the year; winter production and March
egg weight undoubtedly were subject to
such effects. On the other hand, there
must exist some environmental factors
that tend to lengthen the time pullets need
Aga ol first igg
to reach sexual maturity. The assumption
7 9 II 13
WEEK OF YEAR
17 19 of linear trends in such a case is certainly
not generally possible, but may be reason-
PHENOTYPE AVERAGES AS DEPENDENT ON DATE OF HATCH able over a short range of seasonal condi-
tions such as exist for hatches of the early
spring months. Conclusions about them
are necessarily restricted to the conditions
of this study.
2. Nonsystematic environmental effects
that tend to be common to a group of
simultaneously hatched pullets, but differ
from hatch to hatch. Possibly these may
be traced to such factors as sporadic dis-
ease outbreaks, different housing condi-
tions, differential response of birds of dif-
ferent age to climatic variation, different
attendants, and many other ones. The
nature of these factors makes a prediction
of their effects impossible. Consequently
they are considered as random variables.
9 II 13 IS 17 19
WEEK OF YEAR
ANALYSIS OF VARIANCE
FIG. 1. The dependence of production traits on
date of hatching. Each point on the curves repre- The relative effect of various genetic
sents an average of three consecutive hatches. and environmental factors on observed
 SELECTION PROCEDURES 1289

differences between individuals was ap-
praised by means of the usual analysis of
variance techniques, particularly by means
of variance components. Statistical pro-
cedures for obtaining unbiased estimates
of variance components, adapted to ex-
periments with unequal class numbers of
corresponding subgroups of observations,
have been discussed by Henderson (1953).
In the present state of the theory and for
the case where all assumed effects of the
mathematical model are uncorrelated
random variables with zero means, the
following procedure seems feasible:
1. Choice of a mathematical model for
which the available data can furnish esti-
mates of variance components.
2. Selection of mean squares or sums of
squares appropriate to a different mathe-
matical model in which the same effects
are treated as fixed constants, and where
experimental classes are equal or propor-
tionate (orthogonal experiment).
3. Calculation of the expectations of the
mean squares or sums of squares in (2).
These will be linear functions of variance
components of the assumed random
effects.
TABLE 3.—Notation in the analysis of variance
4. Solution of the linear equations for
variance components.obtained by equat-
ing the above mean squares or sums of
squares in (2) with their expected value
in (3).
The suggested approach to the problem
does not necessarily provide estimates
with the smallest possible sampling vari-
ance. However, for an experiment of
complex design and with many assumed
effects, it is liable to be the only computa-
tionally feasible method.
The following mathematical model was
chosen for this analysis (see Table 3):
Xhijk= m+a h H +ai S -|-aij D +ahi H S
+ a h i i H D + a h i j k I + b • ZMjk,
where Xhijk stands for a production meas-
urement of the k th hen from the j t h dam
and the i th sire, hatched in the h th weekly
period. The effects postulated in the model
stand, successively, for a constant over-all
mean (m), random effects contributed by
hatch environment (ahH), sire genotype
(ai s ), dam genotype (a;j D ), hatch x sire
interaction (ahiHS), hatch x dam interac-
tion (ahijHD), and effects particular to in-
dividual offspring (ahijk1)- Finally, we

Numbers Variance
Source of Abbrevia- Subscript Range of of obser- Sum of Mean com-
variation tion (Level) Effect vations ponent
subscript squares square
affected estimate

Population (Year) P m 1 ... 4 m N (SS)P

H
Hatches H h 1 • • -h ah Nh (SS)H (MS)H H
Sires S i 1 •• • s ais Ni (SS)S (MS)S S

Dams D J 1 • • • di a„» N„ (SS)D (MS)D D

HatchXSire HS ahiHS Nu (SS)HS (MS)HS HS
Interaction

Hatch X Dam HD a. ..HD Nhij (SS)HD (MS)HD HD

<*hij
Interaction

Individuals I k 1 • • • ihijk ahijk 1 (SS)I (MS)I I

Linear Regression b m 1...4 b • Zhijk N ...
on Date of Hatch
 o

TABLE 4.—Sums of squares for estimating variance components and regression coefficients

Source of Uncorrected sums of squares Sums of squares due to regression Sums of crossproducts for regression estimate
variation Symbol Formula Symbol Formula Symbol Formula

X2 Z2
Population (US)P (RS)P b2 - ^ (US^JP
N N

Xh
Hatches (US)H V -2 (RS)H (RS)I (US)XZH
h Nh
2
^ X.i..2 2 7 • >
Sires (US)S (RS)S b E ^ (US)XZS w
T N; i Ni
>
X.ii. 2 7 .. 2 X.ijkZ.ijk >
Dams (US)D II (RS)D b ' H (US)XJD II 1J
Ni, i i N,j Nii

XbiJ
H X S Interaction (US)HS (RS)HS (RS)I (US) IZ HS

H X D Interaction (US)HD (RS)HD (RS)I (US)X2HD

Individuals (US)I x
1 1 1 1 wjk2 (RS)I W i l l i ; Zhiik2 (US) X J I 1 1 I XhijkZhijk
h i j k h i j k
h i j k
 SELECTION PROCEDURES 1291

shall assume that there exists a systematic
linear hatch effect, ( b z h i j k ) due to the
hatching date Zhijk- No assumptions are
made about the distributions of the postu-
lated random variables. It is assumed,
however, that:
1. All random effects are independent of
each other.
2. They have zero means.
3. All ahH have equal variance, H, and
that similarly the variance of all ai s is S,
etc., as shown in Table 3.
This model has been applied to each of
the four populations. I t is assumed that
the true values of the variance com-
ponents estimated are the same for all four
populations; hence the estimates will be
pooled for all final results.
The analysis of variance used for this
experiment follows Henderson's (1953)
procedure, except for differences in nota-
tion and added complexity due to assumed
regression and interactions. The sums of
squares uncorrected for mean and regres-
sion (US) are defined in Table 4. The
usual dot notation is used here to indicate
that sums have been taken over all sub-
scripts of Xhijk which are replaced by a
dot. Thus X.... represents the over-all sum
of all observations on a population. The
corresponding arithmetic average of the
whole population will be designated
as x.....
The expectations of uncorrected sums
of squares in terms of variance compo-
nents are given in Table 5. Each of the
coefficients (K's) given in Table 5 corre-
sponds to the variance component that is
given as a column heading.
Aside from variance components, each
uncorrected sum of squares contains vari-
ance due to the population mean; this
contribution is constant for all sums of
squares and equals Nm 2 .
Another portion of variability con-
sidered in the model is attributable to the
linear regression, b, of performance on
date of hatches. Consequently, each sum
of squares contains some portion of varia-
bility that would be eliminated if regres-
sion effects were nonexistent or if they
were removed by appropriate analysis of
the data. The contributions to total varia-
bility are considered as sums of squares
due to regression and are designated as
(RS). Table 4 gives the expected reduc-
tions in sums of squares. It will be noted
that each (RS) contains the parameter b 2 .
In order to obtain numerical values for the
various (RS), b 2 was replaced by its esti-
mate b 2 , and
.-_ (US)xJ-(US)x.D
~ (US) ZZ I-(US) ZZ HD
(US)XzI stands for the sum of crossprod-
ucts between x and z for individual obser-
vations, (US)XZD represents a similar sum
of crossproducts at the level of dam totals,

TABLE 5.—Expectation of uncorrected sums of squares (US) in terms of the general model

Variance components Constants due to

Sum of
squares H S D HS HD I m2 b2
(US)P* Kt K2 K3 K4 K5 1 N (RS)P
(US)H N K8 K, K„ K7 h N (RS)I
(US)S K8 N K9 K8 Kio s N (RS)S
(US1D Ku N N Ku Ku K13 N (RS)D
(US)HS N N Kl2 N Kl2
uta N (RS)I
(US)HD
(US)I
N
N
N
N
N
N
N
N
N
N vNhd N
N
(RS)I
(RS)I

* Each row in the above table stands for a linear equation: for instance: (US)P = HKi+SK 2 +DK
+HSK4+HDK6H-I+m2N-|-b2(RS)P.
1292 H. ABPLANALP

and (US)zZI and (US)ZZHD are uncor-
rected sums of squares of the independent
variable, (z). Appropriate formulas are
shown in Table 4.
The procedure outlined for excluding
variance due to regression from the subse-
quent analysis does not use all informa-
tion available for estimating regression.*
As the present analysis of covariance is
based on large numbers of degrees of free-
dom, a relatively small loss of information
is expected. Numerical values of b are
* An adjustment method for regression compa-
rable to the one used here has been given by Cochran
and Cox (1950, p. 81) for a factorial experiment.
They show that the approximate method gives cor-
rections closely comparable to those obtained by the
exact method. The approximation is recommended
there as a labor-saving device where a large number
of adjustments is needed.
given in Table 2 which also shows which of
the b's could reasonably be considered to
be different from zero. The appropriate
mean square for error is (MS)I, the mean
square due to individual deviations from
Xhij. subtotals. Corresponding numerical
values are given in Table 2. The derivation
of (MS)I will be given later (see Table 7).
The computation of numerical values
for all (US), (RS), and (K's) permits a
solution of the equations of Table 5. The
computation of mean squares leads to the
most convenient solution. For combined
populations the K's of variance compo-
nents in uncorrected sums of squares are
simply the sums of four corresponding
K-values from individual populations as
shown in Table 6.
Estimates of variance components were
derived according to two possible assump-

TABLE 6.—Formtdas for coefficients of variance components

Ki=———=203.13 K2
Lw
-=346.48
N N

E E Nii2 E E Nhi2
K 4 = — --j- =28.36
N N
EEENhii2 E NM 2
K 6 = - ^ - ^ =9.70 K 6 = E —T-T =395.21
N i> Ni,
2
EEN hii INM'
h
K7= E ^ =137.71 K8=£ =238.57
h JNh Ni

£N„« E E Nhu2
K0= E —T ; — = 8 1 7 . 9 2 Kio= 2-i _ h j =94.21
Ni

E Nh„« E Nhij2
K„= E E -=315.47 Ki2= E E -——=1076.34
l. i Nhi
K 13 =Edi=137

Uh,=Number of SireXHatch subclasses=515.

Vhd=Number of Dam X Hatch subclasses =1384.
h = S9.
s=40.
N=2647.
 SELECTION PROCEDURES 1293

TABLE 7.—Analysis of variance for combined populations, with adjustment for linear
regression of measurements on hatching date

Source of variation Symbol Sum of squares formula Degrees of freedom

Random Hatch Effects (SS)H (US)H-(US)P+(RS)I-(RS)P E (hm*)-4

m

Sires (SS)S ( U S ) S - ( U S ) P - (RS)S+(RS)P £(sm)-4

m
Dams (SS)D (US)D-(US)S-(RS)D+(RS)S
E E (dmi —Sm)
H X S Interaction (SS)HS (US)HS-(US)S-(US)H+(US)P
-(RS)P+(RS)S E (Um-Sm-hm)+4

H X D Interaction (SS)HD (US)HD-(US)D-(US)HS+(US)S

-(RS)S+(RS)D E E (Vm-Um-dmi+sm)
m i

Individuals (SS)I (US)I-(US)HD E (Nm-Vm)

:
Subscript m is used here to denote populations.

tions. In the first instance it was assumed
that linear trends in production could be
removed from observations in actual
breeding experiments. For such conditons
all hatch effects due to regression would be
absent, and the method of analysis shown
in Table 7 would be appropriate. Alterna-
tively, it was assumed that systematic
trends due to hatching date (or season)
could not be evaluated effectively under
conditions of actual breeding programs. It
might thus be more realistic to consider
the combined hatch effects as a random
variable. In this case no adjustments for
regression were made {i.e., in Table 7 (RS)
sums of squares were not subtracted).
Variance component estimates obtained
are shown in Table 8, with adjustment for
regression, and in Table 9 for. the case
where all hatch effects were considered as
a random variable.
The interpretation of results is facili-
tated by expressing expected variability
from different effects (variance compo-
nents) as percentage of total variability
(represented by the sum of all variance
components in the model). Tables 10 and
11 give these relative variance estimates
for adjusted and unadjusted analyses re-
spectively.
The variance estimates in Tables 10 and
11 provide direct information about the

TABLE 8.—Estimates of variance components when TABLE 9.—Estimates of variance components when all
ejects of seasonal trends are removed hatch effects are assumed to be a random variable

Source of M W Age at
^Ch W
^er Production A e at
l r fa5t
= Source of M a
, £h ^ " Production first
variation egg egg to 72 weeks variation wefht production to 72 weeks
weight production egg
H 0.0047 17.0308 17.9008 0.4181 H 0.1105 56.002 37.580 2.3125
S 0.296S 15.6720 52.2024 0.2820 S 0.2967 14.175 51.558 0.2972
D 0.3702 8.6951 43.6475 0.5569 D 0.3723 12.130 45.534 0.5445
HS 0.0122 3.2522 21.3959 0.5295 HS 0.0119 2.939 28.707 0.5562
HD 0.1084 2.6350 21.7350 -0.3049* HD 0.1066 3.431 2.191 -0.2925*
I 1.8000 138.1900 697.8060 9.6550 I 1.8002 138.190 697.806 9.6550

Total 2.5920 185.4751 854.6876 11.4415 Total 2.6982 226.867 863.376 13.3654

Negative components are assumed to be zero. * Negative components are assumed to be zero.
1294 H. ABPLANALP

TABLE 10.—Contributions of random effects to total production and are practically absent in
variability (expressed in percentage) after
exclusion of linear trends of hatches egg weight.
4. Genotype-environment interactions
Winter
Source of March
egg egg Produc-
tion to
Age
at first
contributed relatively little to total varia-
variation produc-
weight tion 72 weeks egg bility and may be of negligible magnitude.
H .18 9.18 2.09 3.65 5. Environmental effects peculiar to in-
S 11.44 8.45 6.11 2.46
D 14.28 4.69 5.11 4.87 dividuals and genetic effects due to ran-
HS .47 1.75 2.50 4.63
HD 4.18 1.42 2.55 .00 dom distribution of alleles heterozygous
I 69.45 74.51 81.64 83.49
in parental genotypes as given by I consti-
Total 100.00 100.00 100.00 100.00
Total interaction 4.65 3.17 5.05 4.63 tute the majority of observed variation for
2(D+S)
hi*= all traits considered. Egg weight shows
100
lowest individual variability, followed by
winter production, production to 72
relative effect of different sources of varia- weeks, and age at first egg with increas-
tion on possible gains from mass selection. ingly higher environmental variance.
They suggest the following conclusions: 6. These conclusions remain essentially
1. Sires and dams contributed about unchanged, whether or not adjustments
equally to the genetic variability of off- are made for linear regression of traits on
spring. hatching date.
2. The percent heritable variation ap-
APPLICATION
pear to be highest for March egg weight,
followed by winter production, egg pro- The derived estimates of variance com-
duction to 72 weeks of age, and age at first ponents were used in comparing the ex-
egg. pected results of various selection pro-
3. Environmental effects attributable to cedures. If selection pressure is assumed
hatch environment were most pronounced to be constant and if no repeat matings
in winter egg production and age at first are to be made the problem reduces to one
egg. They appear negligible for 72-week of determining which of the different esti-

TABLE 11.—Contributions of random effects to total variability (expressed in percentage) if all

hatch effects are considered random variables

Source of variation March egg Winter egg Production to Age at

(Component) weight production 72 weeks first egg

H' 4.10 24.68 4.35 17.30

S' 11.00 6.25 5.97 2.22
D' 13.80 5.35 5.27 4.07
HS' .44 1.32 3.33 4.16
HD' 3.95 1.51 .25 .00
I' 66.71 60.89 80.83 72.25

Total 100.00 100.00 100.00 100.00

Total Interaction 4.40 2.83 3.55 4.16

2(D'+S')
h2* = - .50 .23 .22 .13
100
2(D'+S')
.52 .31 .24 .15
S'+D'+HS'+HD'+I'
 SELECTION PROCEDURES 1295

mations of breeding worth is most accu-
rate.
Tables 10, 11, and 12 give heritability
estimates appropriate for four possible
types of mass selection.
The first type of heritability estimate,
2(D + S)
W likely to be met in practice, since unequal
H+S + D+HS+HD+I
is given in Table 10. Only random hatch
effects are assumed to exist. In reality, one
might expect such a situation if mass se-
lection were practiced in a breeding ex-
periment with only a few consecutive
hatches. It might also occur if each ob-
servation were adjusted in some manner
for linear regression on hatching date.
The second type of heritability estimate,
h 2 = 2(D'+S')
H'+S'+D'+HS'+HD'+I'
given in Table 11, predicts gains from se-
lection without adjustments for hatch
effects. Variance components derived from
the analysis of variance without adjust-
ment for linear regression of observations
on hatching data are denoted by a prime.
The results would apply were mass selec-
tion practiced in a population of several
hatches with some uncontrolled trends
existing from one hatch to the next; a
situation likely to hold true under most
practical circumstances.
A third estimate of heritability,
2(D'+S')
h 3 2 =-
S'+D'+HS'+HD'+r
pertains to a situation where hatch effects
are present but can be excluded com-
servations for all hatch effects. As shown
in Table 11, winter egg production and,
perhaps, age at first egg would show the
highest increase. The other two traits
show little rise in heritability by the ex-
clusion of hatch effects.
Exclusion of all hatch effects is not
representation of families leads to biased
estimates of hatch effects. Instead, one
might contemplate simple adjustment of
individual observations for hatch means.
In general, this leads to slightly biased
corrections since each hatch mean de-
pends on the precise numbers of offspring
by which families are represented. An
analysis of this abbreviated type was per-
formed with the available data, and vari-
ance components were derived by assum-
ing the model:
xijk" = m " + a i s " + a i i D " + a i j k I "
Double primes are used in this analysis to
indicate that each observation Xhuk was
first adjusted for the appropriate hatch
mean Xh...; thus, Xijk" = Xhijk—Xh.... The
usual heritability estimates were then de-
rived on the basis of variance components
for sires and dams. They are shown as h42
in Table 12. Since lu2 is affected by exist-
ing genotype-environment interactions, it
is comparable to li32 of the preceding
analysis, except for the incompleteness of
exclusion of hatch effects; this would tend
to reduce h42 below h32.
T A B L E 12.-—Analysis of variance and estimates of
heritability with observations adjusted for hatch means
March Winter Produc- Age
Statistic egg egg tion to
72 weeks at first
weight produc-
tion of age 6S

pletely by appropriate techniques of anal- (MS)S"« 26.852 1262.8 4930.8 47.76

(MS)D" 8.351 344.2 1551.2 20.44
ysis. In practice this will not be possible. (MS)I"=I" 1.969 164.7 811.7 13.03
D" .353 9.934 40.94 .410
By comparing h22 and h32 it is possible S" .293 14.721 54.06 .435
2(S"+D")
to appraise the potential increase in the h,'= .50 .26 .21 .12
S"+D"+I"
effectiveness of mass selection attainable
from proper adjustment of individual ob- * A double prime is used to distinguish statistics of this
analysis from those previously used.
1296 H. ABPXANALP

The relative efficiency of mass selection
under the four methods discussed above is
shown later (Table 16).
Information about the genotypes of in-
dividuals can be augmented from the per-
formance of related groups of individuals
by constructing selection indexes (Hazel,
1943; Lush, 1947; Robertson, 1955). Here,
only two types of genetic relationship are
to be considered: that due to the sire (half
sisters) and that due to the dam (full
sisters). Family averages and individual
performance can be combined into a selec-
tion index, which weighs them in such a
way as to give optimal selection decisions.
Some possible definitions for component
traits of individual breeding worth are
given in Table 13. They are chosen for
application to particular situations.
Index A is designed to represent a
breeding experiment in which decisions
about individual genotypes are desired
before all hatches have been tested. Each
hatch is treated as a separate population
from which animals may be selected as
soon as all its members have finished their
test. Individual performance is measured
as deviation from the average perform-
ance of full sisters; effects of dams, com-
mon to full sisters of the same hatch, are

TABLE 13.—Definitions of component traits in selection indexes and the corresponding mean squares

Source of genetic information

Index Individual phenotype Average dam contribution Average sire contribution

to offspring to offspring
(A)I (A)D (A)S

Xhijk Xhij. Xhij. — Xhi.. Xhi.. —Xh...

(US)I-(US)HD (US)HD-(US)HS (US)HS-(US)H
1259 869 456

Xhijk X.ij. x.i..—x....

(US)I-(US)D (US)D-(US)S (US)S-(US)P
2510 97 36

Xhijk Xhij,
(US)I-(US)HD (US)D-(US)S (US)S-(US)P
1259 3(5

D Xhijk— X..,.*
(US)I-(US)D (US)D-(US)P
2510 133

H, 53 (xhijk—Xh...) — 2~3 23 S (xhijk —Xh...)

E Xhijk—Xh.. h k h j k

Zi 2 (Xhljk—Xh...) 23 23 2~3 (xhijk—Xh...) 23 2323 23 (xhijk—Xh...)

h k h j k k
. (MS)I . (MS)D " ' ' . (MS)S
Nii

* Xhijk—x.... can be rewritten as (xhijk—x.ij.)+(x.ij.—x....). Thes2cond expression (x.ij.—x....) can

then be included in the evaluation of the dam component of the index. (See Lush, 1947 for a similar example).
 SELECTION PROCEDURES
estimated as deviations of full-sister
families from the average performance of
all offspring of the same sire and hatch,
and sire effects are estimated as deviations
of sire progenies of the same hatch about
the hatch mean.
Index B represents a case where the en-
tire population is to be evaluated simul-
taneously and where main effects of
hatches are disregarded. Expected dam
and sire contributions to offspring families
are estimated on the basis of offspring
raised over the entire hatching period.
Effects peculiar to individuals are esti-
mated from deviations of individual per-
formance from the family average of all
full sisters.
Index C is a combination of Index A
and Index B in that it evaluates individ-
uals on the basis of deviations of their per-
formance from the average of full sisters
raised in the same hatch, while parental
contributions are estimated without re-
gard to hatches.
Method D uses a selection index pro-
posed by Dempster and Lerner (1947). It
makes use of information from full-sister
families and that of individual perform-
ance but disregards half-sister groups with
a common sire. Individual phenotypes as
well as family performance are estimated
from deviations about the population
mean. This form of selection is more con-
venient for practical application than the
types of estimates used under methods A,
B, and C.
Index E, finally, represents a situation
where every observation is first corrected
for the average of the hatch group to
which the particular individual belongs.
Family averages are then computed on
the basis of these corrected observations,
including the entire range of hatches. Indi-
vidual performance is estimated from
deviations of corrected observations from
the appropriate dam progeny. Such pro-
1297
cedure might conceivably be advocated if
hatch effects were pronounced and syste-
matic. An evaluation of the selection
efficiency of Index E requires the analysis
of variance shown in Table 12.
Following Hazel's (1943) method of
constructing selection indexes, the com-
ponent measurements of genetic merit are
combined into a multiple regression equa-
tion of the form:
C=b 1 (A)I+b 2 (A)D+b 5 (A)S,
where C is the index value of a given indi-
vidual and b's are regression coefficients,
and where (A) I, (A)D and (A)S stand for
the defined effects attributable to indi-
vidual, dam, and sire, respectively.
If it is assumed that the three compo-
nent measurements can be defined in such
a way that they are statistically inde-
pendent, then the regression coefficients
(bi, b2, b3) are identical with the herita-
bilities of the corresponding component
traits. All indexes used in this study can
actually be expressed in terms of inde-
pendent components.
The accuracy of evaluation of a com-
ponent trait can be defined as the ratio of
its genetic variance to its phenotypic
variance (comparable to a heritability
estimate). Thus:
(G)I
bi = for individual deviations
(P)I
(G)D
b2= for dam effects
(P)D
(G)S
b3= for sire effects
(P)S
The phenotypic variability (P) con-
tributed by a defined component trait is
represented by the corresponding mean
square in the analysis of variance as
shown in Table 13. In order to estimate
genetic variability (G) attributable to de-
fined parental effects or due to individual
1298 H. ABPLANALP

T A B L E 14.—Mean squares and their composition

Computed mean squares

rw™™ ™ u nr„i„ T>-„J ~ A ~„* Average value oi mean squares in terms
Mean Degrees March Winter Produc- Age at
squares f w i a n c e com ponents
ot egg egg pro- tion to first
freedom weight tion 72 weeks egg
(MS)H 55 7.607 2677.0 2647.3 116.16 44.432H+ .886S+ .821D+6.670HS+2.328HD+1.000I
(MS)S 36 28.689 1368.9 5119.8 44.45 . 982H+63.903S+20.152D+5.839HS+2.348HD +1.0001
(MS)D 97 9.161 421.5 1614.0 21.53 .793H+ .000S+18.857D+ .793HS+2.281HD+1.000I
(MS)HS 420 2.215 159.4 858.5 11.61 - . 0 8 4 H - .116S+ .508D+4.861HS+1.987HD + 1.000I
(MS)HD 772 1.850 134.3 715.7 8.68 - . 1 0 0 H - .000S- .335D+ .1O0HS+1.748HD+ 1.0001
(MS)I 1259 1.800 138.2 697.8 9.66 .OO0H+ .O00S+ .O0OD+ .00OHS+ .OOOHD+1.000I
Genetic portions of mean squares (G) Average genetic value of mean squares
(MS)S 36 27.132 1176.6 4309.4 30.81 64.903S+21.152D
(MS)D 97 7.690 255.0 955.7 11.11 1.OO0S+19.857D
(MS)I 97.1 .84 1.000S+ 1.000D
1259 .669 26.3

deviations, the corresponding mean

squares have to be interpreted according
to their genetic and nongenetic compo-
sition. If estimates of all variance com-
ponents are available, the expected ge-
netic variability contained in each mean
square can be assessed, as shown in
Table 14.
In case of deviations of individual
phenotypes from their full-sister average
there is no experimental method for sepa-
rating genetic from nongenetic variabil-
ity. On the basis of Mendelian inheritance
within a randomly mating population it
shall be assumed that I, the variance com-
ponent due to individual deviations, con-
tains one half of the additive genetic
variability, G. For purposes of application
G may be estimated from 2(D + S). I,
therefore, contains genetic variance equiv-
alent to D + S.
The relative efficiency of different selec-
tion indexes will be judged by the genetic
gains expected from their application in
selection. As a basis for comparison, we
shall choose the genetic gains expected
from I122, that is, mass selection in a popu-
lation where no corrections for hatch
effects are made. Since both index and
mass selection exploit the same genetic
variability, (G), the relative efficiency of
index selection can be estimated by the
co-efficient of multiple correlation,
b 1 (G)I+b 2 (G)D/n d +b 3 (G)S/fi 8
Re >=j/
where fid and n8 are the average numbers
of offspring retained from dams and sires
respectively, under the assumed method
of evaluation.
For specific selection decisions on indi-
vidual pullets the number of full sisters is
known, and it is possible to replace n„ and
fia in each selection by the exact number
of relatives. As before, the required
weighting factors (regression coefficients
(bi, b2, b3)) can be obtained as the fraction
of the expected genetic mean square over
the expected phenotypic mean square,
using estimates of variance components
and known family sizes. For simplicity it
shall be assumed here that every parent is
represented equally by offspring in all k
hatches. Thus, each dam is supposed to
have had n offspring per hatch and each of
5 sires is assumed to have been mated to a
constant number of d dams. With these
assumptions the total population size be-
comes hnds, and the number of offspring
available from each hatch is rids.
Table 15 gives weighting factors for
family means and individual performance
on the basis of the above assumptions.
For equal class numbers, the factors in
Index E can be assumed to be equivalent
to those of Index C.
 SELECTION PROCEDURES 1299

TABLE 15.—Weighting factors for family and for individual deviations for various selection methods

Selection Estimates of
method effects Expected genetic variance Expected phenotypic variance
(Index)

Individual G+GH*
(Xhijk—Xhij.) 2 +E

Dam G(n+1) G(n+1) , GH(n+l) , ^

(Xhij. — Xhi..) \ 1 r~—HE

Sire G(nd+n+l) G(nd+n+l) GH(nd+n+n

(Xhi..-X h ...)

Individual CI G+GH
(Xhijk—X.ij.) ~2
Dam G(nh+1) G(nh+1) GH(n+l)
(x.u.-x.i..) 1 + 4 +E
G(ndh+nh+l) G(ndh+nh+l) 1 GH(nd+n+l) [_E
Sire

/ hnds—hnd\ / hnds—hnd\
Dam and Sire G
(X.ij.-X....)
(2+hn+-^T-) G 2+hn+
( -7s-— ) |
4
GH ( nds— nd \
-+E

* GH stands for the component of genotype-hatch interaction.

The formulas given in Table 15 can also
be used to evaluate the results for the hy-
pothetical case of selection with equal
family sizes. A comparison of the efficiency
of index selection in such a balanced popu-
lation with the expected outcome of selec-
tion in the actual experiment may indi-
cate how seriously unequal representation
of families affects the presumed efficiency
of selection. The situation may be ex-
plored by means of Index F, which corre-
sponds to Index B of the actual experi-
ment. For the balanced experiment it is
not possible to express family sizes as
whole numbers. For pooled populations
they assume the following values: fl=1.3
d = 3.43 §=10.00 h = 14.75. The results
for Index F are given in Table 16, to-
gether with those of the other selection
methods discussed. The tabulated values
of relative selection efficiency are
RC.G
Comparison of the relative efficiencies
of the selection methods in Table 16 sug-
gests the following conclusions:
1. Selection under exclusion of linear
trends in the performance of successive
hatches (hi2) shows some improvement in
efficiency over straightforward mass se-
lection (h22) for winter egg production and
age at first egg. Selection for March egg
weight and production to 72 weeks, on the
other hand, is not more effective when
linear trends are excluded.
1300 H. ABPLANALP

TABLE 16.—Relative efficiency of selection methods with respect to mass selection and heritabilily hi

Selection efficiency relative to hj

Selection method
March egg Winter egg Production to Age at
weight production 72 weeks first egg
Mass selection without adjustments for hatch
effects. (h2) 1.000 1.000 1.000 1.000
Mass selection under exclusion of linear trends
of hatches, (hi) _ 1.015 1.065 1.000 1.116
Mass selection under complete exclusion of all
hatch effects, (hs) 1.018 1.152 1.022 1.099
Mass selection with corrections of observations
for hatch means, (hi) .996 1.060 .965 .984
Index A; combination selection on the basis of
individual, dam average and sire average
within hatches. 1.124 1.277 1.135 1.179
Index B; combination selection on the basis of
individual, dam average and sire average dis-
regarding existing hatch effects. 1.190 1.482 1.485 1.789
Index C; combination selection, disregarding
hatch effects for sire and dam averages, but
not for individuals 1.206 1.523 1.495 1.816
Index D ; combination selection on the basis of
individual and dam average disregarding
existing hatch effects. 1.111 1.372 1.373 1.630
Index E; combination selection on the basis of
individual, dam average and sire average, with
adjustment of observations for hatch means
(biased). 1.190 1.506 1.467 1.749
Index F ; combination selection for the hypo-
thetical case of equal family representation in
all hatches. 1.220 1.542 1.517 1.883

2. Exclusion of all hatch effects from
comparisons of individuals (h32) has some
advantage over exclusion of linear trends.
3. Actual correction of observations for
hatch means (h42) results in slightly
biased comparisons of individuals and ac-
cordingly shows for three of the traits a
decreased selection efficiency as compared
with the hypothetical case of completely
excluded hatch effects (h32). In the present
experiment the corrections for hatch
means by this method resulted in a loss of
selection efficiency relative to mass selec-
tion (I122) for production to 72 weeks and
for age at first egg.
4. All index selection is more efficient
(as expected) than any of the other selec-
tion procedures.
(a) Selection on the basis of a combination of
parental effects and individual deviations of
offspring from family averages is relatively
inefficient if practiced within separate hatches
(Index A),
(b) Combination selection within the entire
population, where family performance is esti-
mated from all available offspring, shows
high efficiency, even when hatch effects are
disregarded (Index B). The improvement of
selection efficiency by this means is highest
for age at first egg (79 percent) and lowest
for egg weight (19 percent). Thus, the trait
with lowest heritability shows highest'possi-
ble improvements in selection precision when
family averages are used.
5. Refinements of selection procedures
such as represented by Index C do not
produce marked added improvements
over Index B.
6. Selection Index D, in which sire and
dam effects on offspring performance are
confounded, is almost as efficient as
Index B, where sire effects on offspring
are considered separately from dam ef-
fects. The loss in selection efficiency rela-
 SELECTION PROCEDURES
tive to Index B is 8 percent for egg weight,
11 percent for both winter production and
production to 72 weeks, and 16 percent
for age at first egg. Thus, it may be ad-
visable to use Index B for traits with very
low heritability.
7. Adjustments of individual perform-
ance for hatch means and subsequent ap-
plication of combination selection (Index
E) do not prove more efficient than com-
bination selection without hatch-adjusted
observations (Index B). This result may
be due to an overestimate of efficiency of
index B.
8. Biases of family means and losses in
selection precision from unequal family
sizes account for only slight losses in se-
lection efficiency (Index E compared to
Index F). Age at first egg, which has
lowest heritability of the four traits in-
vestigated, shows highest biases in family
averages.
Thus, the over-all results suggest that
corrections of individual observations for
hatch means do not improve selection
efficiency greatly, at least not under the
conditions of this experiment. Family se-
lection, as compared to mass selection on
the other hand, proves highly effective,
except for egg weight, which, having high
heritability, shows little potential im-
provement for either hatch corrections or
family selection.
DISCUSSION
Practical recommendations on the basis
of a study of limited experimental.ma-
terial in spite of an element of speculation
may apply to a rather wide range of con-
ditions and hence can serve as a useful
guide for breeding practice.
Hatching date was found to exert sig-
nificant effects upon all production traits
studied, particularly upon sexual maturity
and winter egg production. Similarly,
there were demonstrable random effects of
1301
hatch environment, with most pronounced
responses in winter production. These
findings generally agree with published
work of similar experiments by Garber
and Godbey (1952) and King and Hender-
son (1954). Thus, it appears safe to as-
sume that under many practical condi-
tions there are environmental factors
common to pullets that are hatched and
reared together. Most likely these hatch
effects vary considerably in magnitude be-
tween different generations of the same
population and between different flocks
(e.g., Bohren et al., 1952).
The study further indicates that the
magnitude of genotype-environment in-
teractions due to extended hatching sea-
sons is small for most practical purposes.
Combined genotype-environment inter-
actions account for only about 4 percent
variability among individuals; this is
somewhat less than variability due to
hatch effects.
As yet there have been few published
cases of strong genotype-environment
interactions in production traits of chick-
ens, although they exist under special con-
ditions (e.g., with respect to disease ex-
posure and viability, DeOme, 1943; Hutt
and Cole, 1947).
Although the estimates of hatch-geno-
type interaction presented in this study
amount to only about four percent of
total variability, it is possible that similar,
but more pronounced, interactions might
exist between successive generations of
the same population and the environ-
ments of successive years. Such interac-
tions might in part explain the fact that
selection within closed populations does
not produce as high gains as predicted
from heritability studies. If it is assumed
that genotype-environment interactions
generally exist, then it might be desirable
to adopt selection procedures that would
provide as wide a range of environmental
1302 H. ABPLANALP
conditions as possible within one genera-
tion. Consequently, one would expose
separate hatches to widely differing en-
vironments.
The magnitude of heritability estimates
derived here on the assumption of existing
genotype-environment interactions agrees
well with other published work.
King and Henderson (1954) give the
following average heritabilities from previ-
ously published studies: 0.57 for egg
weight, 0.31 for full year's production, and
0.27 for age at first egg. On general con-
siderations, and in agreement with practi-
cal recommendations advanced by others
(Lerner and Cruden, 1951), it appears
possible to improve egg weight by mass
selection alone, even in the presence of
hatch effects. The results presented agree
with this assumption and show that im-
provements in selection efficiency of 20
percent, at best, could be attained if se-
lection were based on combined family
and individual performance, with or
without adjustments for hatch means.
Practical considerations based on the
present findings suggest that family selec-
tion permits one to exclude most hatch
effects from selection decisions. However,
such methods may not be feasible or
practical unless the necessary calculations
can be carried out within a short time. In
this light the most useful method is un-
doubtedly family selection on the basis of
full sibs (Method D). Thus, if families are
well represented in all hatches, their aver-
ages will contain only a fraction of the
seasonal variation.
Under conditions of pronounced sea-
sonal trends in hatch effects, it would be
necessary to consider possible bias in com-
parisons either of individuals from differ-
ent hatches or of families hatched over
different ranges of possible hatching dates.
Under such conditions, the use of Index E
would seem appropriate. However, under
the conditions of this experiment an ad-
justment of observations for hatch means
did not appear to produce improved selec-
tion decisions. This may partly be due to
the fact that relatively few parents were
used within generations. Hence it is quite
possible that the sampling errors in the
derived estimates of variance components
lead to overestimates of heritability of
unadjusted observations (h22). Adjust-
ment of observations for hatch means
may be more appropriate in mass selection
than it is for combination selection. Also
in cases where production records are
terminated at a set date, rather than a
certain age of pullets, the effort of hatch
adjustment may become worthwhile.
If the labor of adjusting individual ob-
servations to hatch means exceeds the
available means, better representation of
parental matings might be insured ex-
perimentally by:
1. Excluding poorly represented families
before testing (as in the present study).
2. Allowing for longer reproduction pe-
riods for each hatch of offspring (saving
eggs for two weeks instead of one, as sug-
gested by King and Henderson (1954)).
3. Relying more strongly on family se-
lection than is indicated by considerations
of optimal selection for additive gene
effects.
Adjusting observations for hatch effects
by means of some presumed linear regres-
sion of performance of hens on date of
hatch does not appear a reasonable pro-
cedure because the magnitude of such
effects seems to be rather unpredictable.
Only age at first egg shows reasonably
constant linear regression on date of
hatch. For the same reason it does not
seem possible to make a general prediction
of the influence of such trends on expected
accuracy of mass selection.
 SELECTION PROCEDURES
The actual labor of adjusting observa-
tions for hatch means can be reduced con-
siderably if family representation within
hatches is good. Under these circum-
stances, one might construct a selection
index of type B or D without adjusting
individual performance for hatch means.
Selection could then be made on the basis
of this index, but parents would be chosen
in numbers proportionate to the size of
the hatch to which they belonged. This
procedure would, in effect, prevent large
hatch main effects from causing selective
discrimination against the members of a
particular hatching group.
The conclusions of this study regarding
the importance of hatch effects for selec-
tion efficiency are in disagreement with
recommendations made by Bohren et al.
(1952) and more recently by King and
Henderson (1954). The latter state: "It is
our conclusion that the hatch effect is of
sufficient importance to necessitate the
adjustment of data for this effect." The
discrepancy in results seems to be due
partly to the statistical methods em-
ployed in evaluating expected gains from
selection, and does not necessarily reflect
drastic differences in the magnitude of
hatch effects. King and Henderson (p.
162) attempted to evaluate the impor-
tance of hatch effects by alternately as-
suming that they existed, or that they
were not present at all, and by computing
heritability estimates on that basis. If
existing hatch effects are ignored this pro-
cedure results in a slight overestimate of
variance due to individuals and tends to
underestimate dam effects considerably.
I t appears that corrections for hatch ef-
fects, if performed without bias, improve
selection efficiency little, even when the
latter account for 25 percent of total
variability (winter egg production). These
potential improvements can perhaps also
1303
be realized in practice if selection in-
tensity of parents is kept constant within
hatches; if observations are merely ad-
justed for hatch means, the actual selec-
tion precision (lu2) will be somewhat less
than given by h32 because of biases in
hatch means or perhaps because of round-
ing errors in the calculation of adjusted
performance.
SUMMARY
From the results presented here, some
inferences can be drawn regarding the
effects of an extended hatching season on
the efficiency of artificial selection among
the resulting offspring. For the popula-
tions investigated, the analysis shows
that:
1. The effects of date of hatching for
laying flocks, hatched over 11 to 17 weeks
during spring months on production traits
were pronounced for age at first egg and
winter egg production; they were but
negligible for March egg weight and egg
production to 72 weeks of age. The same
relations hold for nonsystematic environ-
mental effects between hatches but to a
lesser extent.
2. Interaction between genotypes and
environment was generally low for all
traits.
3. Combined selection on the basis of
average family performance and indi-
vidual performance appeared to be effec-
tive in providing high accuracy of selec-
tion decisions, with or without adjustment
of observations for hatch means.
4. Slight additional improvements in se-
lection accuracy would be expected when
adjusting individual observations for the
corresponding hatch mean before selection
on combined family and individual per-
formance. However, such advantages
could not be demonstrated conclusively in
this study, possibly due to sampling errors
1304 H. ABPLANALP
in the derived variance components and
heritability estimates.
5. Unequal representation of families in
consecutive hatches did not appear to be a
serious obstacle to accurate selection.
ACKNOWLEDGEMENTS
The author is indebted to Kimber
Farms Inc. and their director of research
Dr. W. F. Lamoreux, for making avail-
able the data for this study. It is a pleas-
ure to acknowledge the help of Professor
I. Michael Lerner, who suggested the
problem and who made constructive sug-
gestions throughout the course of this
work.
REFERENCES
Bohren, B. B., G. D. Rapp and R. B. Arvidson,
1952. Increase in pullet egg size as a factor in
selection. Purdue Univ. Agr. Expt. Sta. Bull.
574.
Dempster, E. R., and I. M. Lerner, 1947. The
optimum structure of breeding flocks. I I .
Methods of determination. Genetics, 32: 576—
579.
DeOme, K. B., 1943. Intraperitoneal injection of
lymphomatous nerve tissue into resistant or
susceptible chickens. Poultry Sci. 22: 381-394.
Garber, M. J., and C. B. Godbey, 1952. The in-
fluence of sire, dam, and hatching date on
specific rate of growth of Single Comb White
Leghorn pullets from hatching to twelve weeks
of age. Poultry Sci. 3 1 : 945-955.
Gowe, R. S., and W. J. Wakely, 1954. Environ-
ment and poultry breeding problems. 1. The
NEWS AND NOTES
(Continued from page 1284)
SOUTH DAKOTA NOTES
W. Kohlmeyer, Head of the Department of
Poultry Husbandry, South Dakota State College,
returned to the College on September 1st, after
spending a year's leave in graduate study at Pur-
due University.
L. U. Rubida (B.S., South Dakota State, 1956)
has been appointed Graduate Teaching Assistant.
H. Woerpel (B.S., South Dakota State, 1956)
has been appointed Graduate Research Assistant.
(Continued on page 1348)
influence of several environments on the egg
production and viability of different genotypes.
Poultry Sci. 33 : 692-703.
Hazel, L. N., 1943. The genetic basis for construct-
ing selection indexes. Genetics, 28: 476-490.
Henderson, C. R., 1953. Estimation of variance and
covariance components. Biometrics, 9: 226-252.
Hutt, F. B., and R. K. Cole, 1947. Genetic control
of lymphomatosis in the fowl. Sci. 106: 379-
384.
King, D. F., P. K. Cole, F. B. Hutt and G. J.
Cottier, 1952. Tests in different environments
of fowls genetically resistant to leukosis. Poultry
Sci. 3 1 : 1027-1029.
King, S. C , and C. R. Henderson, 1954a. Variance
components analysis in heritability studies.
Poultry Sci. 3 3 : 147-154.
King, S. C , and C. R. Henderson, 1954b. Herita-
bility studies of egg production in the domestic
fowl. Poultry Sci. 3 3 : 155-169.
Lerner, I. M., 1950. Population Genetics and Animal
Improvement, xviii + 342 pp. Cambridge Uni-
versity Press.
Lerner, I. M., and D. Cruden, 1951. The heritabil-
ity of egg weight: The advantages of mass
selection and of early measurements. Poultry
Sci. 30:34-41.
Lerner, I. M., and E. R. Dempster, 1951. The
theory of multiple shifts. Poultry Sci. 30: 717-
722.
Lush, J. L., 1947. Family merit and individual
merit as bases for selection. Part I. Amer. Nat.
8 1 : 241-261.
Osborne, R., 1954. Sexual maturity in Brown
Leghorns and the relationship of genetic vari-
ance to differences in environment. Proc. Roy.
Soc. Edinburgh, Section B, 65: 285-298.
Robertson, A., 1955. Prediction equations in
quantitative genetics. Biometrics, 1 1 : 95-98.
MONTANA NOTES
Dr. E. R. Halbrook resigned effective October 1,
as Head of the Poultry Department, Montana State
College, to go to India as a Poultry Husbandry
Advisor, an assignment with the International Co-
operation Administration.
GENERAL MILLS NOTES
D. S. Carver has been appointed Broiler Spe-
cialist at General Mills Larro Research Farm, In-