Body Weight and Egg Weight Dynamics in Layers

R. J. DI MASSO,*,†,‡ A. M. DOTTAVIO,*,‡ Z. E. CANET,* and M. T. FONT,†,‡,1

*Cátedra de Genética y Biometrı́a, Facultad de Ciencias Veterinarias, Ovidio Lagos y Ruta 33, CC 166, 2170 Casilda,
Argentina, †Instituto de Genética Experimental, Facultad de Ciencias Médicas, Santa Fe 3100, 2000 Rosario, Argentina,
and ‡Consejo de Investigaciones, Universidad Nacional de Rosario, Santa Fe 3100, 2000 Rosario, Argentina

ABSTRACT The association between body weight-age at the lowest maturing rate (MEW = 0.922), the inverse
and egg weight-age patterns was studied in a segregat- being true for birds in Group 3 (AEW = 55.7 g and MEW
ing population of laying hens belonging to the F3 = 0.737). Birds belonging to Groups 2 and 4 were

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generation of a Rhode Island Red × White Leghorn
reciprocal cross. Body weight and egg weight were
expressed as a function of time using the model
developed by Weatherup and Foster. Each hen was
characterized in terms of its asymptotic body weight
(ABW), maturing rate for body weight (MBW), asymp-
totic egg weight (AEW), and maturing rate for egg
weight (MEW) values. Four groups of hens were
distinguished by means of a principal component
analysis. Birds belonging to Groups 1 and 3 were
discriminated for their egg weight-age pattern. Group 1
included hens laying the heaviest eggs (AEW = 66.1 g)
(Key words: body weight, egg weight, Weatherup and Foster model, principal component analysis, layer)
distinguished for their body weight-age pattern. Hens in
Group 2 showed the lowest ABW (1,893 g) and MBW
(0.764) whereas the heaviest (ABW = 2,802 g) and less
mature (MBW = 0.929) birds were found in Group 4.
The results confirm the partial pleiotropic basis of the
body weight-egg weight correlation, evincing the feasi-
bility of applying selective pressure not only on each
character separately but also on maturing rate indepen-
dently of asymptotic weight within each trait. This
strategy could be implemented using a biological
selection index based on principal component analysis
equations.
1998 Poultry Science 77:791–796

INTRODUCTION low asymptotic weight and a high maturing rate

Growth is a complex biological process resulting from
genetic factors and environmental circumstances, which
is usually represented, both in domestic and experimen-
tal animals, as an age-dependent S-shaped change in
body weight. The dynamic of this time-related change
can be described by means of different mathematical
models such as the four-parameters Richards model
(Richards, 1959) and its variants, the logistic, Gompertz,
and von Bertalanffy equations (Krause et al., 1967;
Knizetova et al., 1983; Grossman and Bohren, 1985;
Grossman et al., 1985; Ricklefs, 1985; Zelenka et al., 1986;
Kachman et al., 1988; Barbato, 1991).
From an economics point of view, certain associations
between the two most relevant parameters that charac-
terize such models (average size at maturity and
maturing rate) are more desirable than others for animal
production purposes. In layers, for example, an advanta-
geous growth pattern would be the combination of a
(Grossman and Bohren, 1985). This advantage is so in
laying birds because, in contrast to meat-type poultry,
growth is not important per se but it is obviously
necessary for hens to attain the productive phase of their
cycle. Therefore, an early maturing light bird would
express two propitious features in comparison with
other growth patterns, a short nonproductive phase
prior to sexual maturity and low maintenance costs.
With regard to egg weight, it might be advantageous to
have laying hens producing, quickly and regularly, eggs
within a limited weight range (Poggenpoel and Duckitt,
1988).
In addition, body weight and egg weight show a
positive genetic correlation (Siegel, 1962; Festing and
Nordskog, 1967) and, as the egg represents the marketa-
ble product, a compromise exists between reducing
body weight while keeping commercially acceptable egg
weights. This relationship was considered by Nordskog
and Briggs (1968) when discussed the body weight-egg
production paradox.

Received for publication May 15, 1997.

Accepted for publication February 17, 1998. Abbreviation Key: ABW = asymptotic body weight; AEW = asymp-
1To whom correspondence should be addressed: totic egg weight; MBW = maturing rate for body weight; and MEW =
dimasso@cidoc.edu.ar maturing rate for egg weight.

791
792 DI MASSO ET AL.
In poultry, genetic variation for body weight has been
determined and exploited in different experiments of
body growth selection (Siegel, 1962; Festing and Nord-
skog, 1967; Krause et al., 1967; Marks, 1978; Dunnington
and Siegel, 1985). Similarly, egg weight, alone or
combined with other traits, has been extensively used as
a selective criterion (Festing and Nordskog, 1967;
Kolstad, 1980; Liljedahl and Weyde, 1980; Sorensen et al.,
1980). Both variables are usually measured at specific
ages or during a specific period of time along the laying
cycle, reducing an intrinsically dynamic process to a
static determination. This approach reduces the potential
information that experimental data could yield about the
phenomenon under study (Cock, 1966).
This paper discusses the association between body
weight-age and egg weight-age patterns in a segregating
population of laying hens, using a longitudinal analysis
in accordance with the dynamic nature of both
processes. This approach could be applied in poultry to
define selective criteria in genetic improvement plans.
MATERIALS AND METHODS
A random sample of 50 hens belonging to the F3
generation of a Rhode Island Red × White Leghorn
reciprocal cross was used. Parental lines belong to the
stocks maintained at the Facultad de Ciencias Veterinar-
ias, Universidad Nacional de Rosario, Casilda, República
Argentina. Birds hatched in spring were collectively
floor reared under natural photoperiod until 18 wk of
age, and then individually caged under a minimum of
14 h lighting. Hens were maintained under identical
managemental conditions.
Individual body weights were registered weekly to
the nearest 10 g from 18 to 48 wk of age. All the eggs
laid in the same period were identified and weighed the
evening of the same day to the nearest 0.1 g.
Body weights and egg weights for each individual
were expressed as a function of time using the equation
(Y = A – B rt) of Weatherup and Foster (1980), where A
= mature body weight or mature egg weight; B = the
range in body weight or egg weight from t = 0 to the
asymptote; and r = the rate at which the respective
mature weight is approached (r < 1). The same model
was used for fitting both response variables because, on
the one hand it is a reparameterization of Brody’s
exponential growth equation (Brody, 1945), a special
case of Richards function (Richards, 1959) and, on the
other hand, it adequately describes how egg weight
increases with age in various commercial avian species
(Shalev and Pasternak, 1993). Estimation was by non-
linear least squares (Draper and Smith, 1981), using an
iterative procedure via Marquardt’s (1963) algorithm.
Goodness of fit was evaluated by the coefficient of
determination (R2). Each hen was described by the
following traits: asymptotic body weight (ABW), matur-
ing rate for body weight (MBW), asymptotic egg weight
(AEW), and maturing rate for egg weight (MEW).
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FIGURE 1. Scatterplot of the two first principal components, with
the ordination plane partitioned using Lefkovitch’s method. Group 1:
First quadrant (⁄), Group 2: Second quadrant (+), Group 3: Third
quadrant (◊), and Group 4: Fourth quadrant (π).
Four groups of layers were defined by means of a
principal component analysis (Tatsuoka, 1971). Firstly, a
two-dimensional ordination was obtained in which
every point (hen) had as its coordinates two scores
obtained from two different linear combinations of the
aforementioned four traits (ABW, MBW, AEW, and
MEW). Secondly, a divisive classification was performed
by partitioning the ordination plane using Lefkovitch’s
(l976) method. Two divisions were made by breaking
the x-axis and the y-axis of the Cartesian coordinate
system at their respective centroids. As a consequence,
four quadrants were defined, numbered from one to
four in counterclockwise order. Hens in each quadrant
were identified and their individual body weight-age
and egg weight-age curves were fitted. In both cases,
estimations of A and r were considered as new
variables. Differences among groups (quadrants) were
evaluated by one-way analysis of variance (Sokal and
Rohlf, 1969) and when statistically significant, the means
were separated by Duncan’s multiple range test
(Kramer, 1956). A discriminant analysis (Kleinbaum and
Kupper, 1978) was performed to corroborate differences
among groups in their jointly considered body weight
and egg weight patterns.
RESULTS
The proportion of the total variance accounted for by
the two first principal components was 79% (Figure 1).
The first principal component (Y1) showed a positive
and significant correlation with the four traits included
in the analysis: ABW, MBW, AEW, and MEW (Table 1).
Accordingly, the highest Y1 values corresponded to hens
with the highest asymptotic body weight and the lowest
 BODY WEIGHT AND EGG WEIGHT IN LAYERS 793
TABLE 1. First (Y1) and second (Y2) principal component their body weight-age and egg weight-age patterns
equations and Pearson’s correlation coefficients between
body weight-age and egg weight-age curves parameters when they were simultaneously considered, as a high
and principal component values1 proportion of the observations (more than 92%) were
correctly classified.
Pearson’s correlation Parameter estimates for growth curves and egg
coefficients (r)
weight patterns are presented in Tables 4 and 5,
Variable Y1 Y2 respectively. The R2 values indicate an excellent agree-
Asymptotic body weight (ABW) 0.781 –0.424 ment between the curve estimated by the proposed
Maturing rate for body weight (MBW) 0.643 –0.622 mathematical model and the experimental observations.
Asymptotic egg weight (AEW) 0.763 0.439
Maturing rate for egg weight (MEW) 0.602 0.658 Figures 2 and 3 illustrate theoretical body weights and
11st principal component Y = 0.5570 ABW + 0.4583 MBW + 0.5436
egg weights, respectively, for each group using the
1
AEW + 0.4293 MEW. 2nd principal component Y2 = –0.3886 ABW – Weatherup and Foster equation (1980) plotted against
hen age.

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0.5694 MBW + 0.4025 AEW + 0.6023 MEW.

maturing rate for body weight (> r) and the highest
asymptotic egg weight and the lowest maturing rate for
egg weight. The second principal component (Y2)
showed a negative correlation with both body weight
traits, whereas a positive correlation was evident for the
same characters for egg weight (Table 1). In conse-
quence, hens with the highest Y2 values showed the
lowest asymptotic body weight and the highest matur-
ing rate for this variable jointly with the highest
asymptotic egg weight and the lowest maturing rate for
egg weight.
The partitioning of the scatterplot for the first two
principal components (Y1: x-axis and Y2: y-axis) allowed
the identification of four groups of hens (Figure 1 and
Table 2). Birds belonging to Groups 1 and 3 (first and
third quadrant, respectively) were discriminated for
their egg weight-age pattern. Group 1 included hens
laying the heaviest eggs (AEW = 66.1 ± 2.18 g) at the
lowest maturing rate (MEW = 0.922 ± 0.009), the inverse
being true for birds in Group 3 (AEW: 55.7 ± 0.98 g;
MEW: 0.737 ± 0.025). Birds belonging to Groups 2 and 4
(second and fourth quadrants respectively) were distin-
guished for their body weight-age pattern. Hens in
Group 2 showed the lowest ABW (1,893 ± 46 g) and the
highest MBW (0.764 ± 0.029) whereas the heaviest (2,802
± 116 g) and less mature (0.929 ± 0.008) birds were found
in Group 4.
Table 3 shows the results of a discriminate analysis
performed with data distributed in these four groups.
This technique confirmed differences between groups in
DISCUSSION
Body weight and egg weight are two relevant
productive traits in poultry. Body weight has shown to
be highly responsive to selection in chickens and genetic
improvement for growth has been basically performed
selecting for body weight at fixed ages. Genetic
modification of body weight in meat-type chickens
increased feed intake, body fat, and age at sexual
maturity and decreased reproductive performance as
undesirable correlated responses to artificial selection
(Barbato, 1991), whereas, in laying hens, a correlated
modification of egg weight and a decline in egg
production and other components of reproductive
fitness have been described by Festing and Nordskog
(1967) as a consequence of selection for body weight. On
the other hand, egg weight is also highly heritable in
chickens and its economic implications are obvious. In
layers, each 1-g increment in average egg weight may
improve income by about 4% whereas in meat-type
poultry, the same increase may enhance marketing
weight by 2 to 13 g (Shalev and Pasternak, 1993).
The genetic antagonism based on the positive genetic
correlation between body weight and egg weight
indicates the necessity to develop particular strategies
for combining in the same bird a desirable weight-age
pattern for both productive traits, implying a simultane-
ous change in both weight-age trajectories. Among the
different possible approaches available to change the
shape of these curves are 1) to apply selection pressure
at discrete points on the curve, 2) to construct a selection
index to exert selective pressure simultaneously at all

TABLE 2. Nonlinear least squares estimates (mean ± standard error) of Weatherup and Foster equation parameters for individual
body weight-age and egg weight-age data of hens grouped by means of a principal component analysis

Group 1 Group 2 Group 3 Group 4

Quadrant 1 Quadrant 2 Quadrant 3 Quadrant 4
Variable (n = 11) (n = 12) (n = 12) (n = 15)
Asymptotic body weight, g 2,344.17 ± 103.09a 1,892.85 ± 46.46b 2,194.98 ± 105.58a 2,801.72 ± 116.01c
Maturing rate for body weight 0.8704 ± 0.0161a 0.7636 ± 0.0293b 0.8715 ± 0.0153a 0.9294 ± 0.0081c
Asymptotic egg weight, g 66.11 ± 2.18a 57.07 ± 0.98b 55.72 ± 0.98b 60.86 ± 1.12c
Maturing rate for egg weight 0.9217 ± 0.0093a 0.8650 ± 0.0171b 0.7370 ± 0.0254c 0.8611 ± 0.0115b
a–cValues with different superscript differ significantly (P < 0.05).
794 DI MASSO ET AL.
TABLE 3. Discriminant analysis—Results of between groups classification analysis

Contrast
between Discriminant
groups False (–) Sensibility False (+) Specificity rate
(%)
1 vs 2 8.3 91.7 0 100 95.7
1 vs 3 8.3 91.7 0 100 95.7
1 vs 4 9.1 90.9 6.7 93.3 92.3
2 vs 3 0 100 0 100 100
2 vs 4 0 100 0 100 100
3 vs 4 0 100 6.3 93.7 96.3

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points of the curve, or 3) to determine a mathematical
function and then to develop an optimum curve by
selecting for desired values of parameters of this
function (Grossman and Bohren, 1985). Festing and
Nordskog (1967) studied the response to selection for
body weight and egg weight in White Leghorns and
concluded that these characters are evidently controlled
by both independent and pleiotropic genes.
A dynamic study of the same traits in a segregating
population of layers derived from a reciprocal cross
between two genotypes with different asymptotic body
weight showed that egg weight and body weight
increase monotonically with age approaching an asymp-
tote. As both sets of longitudinal data showed a similar
behavior, they were fitted with the same mathematical
model. This pattern is typical of the curve relating egg
weight and age of hen (Cowen et al., 1964; Weatherup
and Foster, 1980; Shalev and Pasternak, 1993; Minvielle
et al., 1994) whereas for body weight-age, data results
from using postinflexion growth values (Brody, 1945).
This dynamic approach allowed us to discriminate
four growth patterns divided in two subgroups (Groups
1 and 4 vs Groups 2 and 3) with very similar
20-wk body weights within subgroup but different
between them, and with different asymptotic weight
within subgroup. With regard to maturing rate, there
were three categories ranking from the earliest maturing
hens with the lowest r value (Group 2) to the late
maturing birds belonging to Group 4 (with the highest r
value). Hens in Quadrants 1 and 3 showed similar
maturing rates for body weight, a condition made
evident in the parallelism of their growth curves.
Groups 1 and 4, with the highest ABW, showed
different patterns of egg size increase with age. The
birds belonging to the first quadrant had a higher initial
egg weight and a lower maturing rate than those
allocated in Quadrant 4. As a consequence, average egg
weight of layers in Group 1 still increased at 50 wk of
age whereas the same trait was nearly stabilized for
hens in Group 4. Hens localized in the other two
quadrants (Groups 2 and 3) displayed a differential
behavior related to maturing rate but not to AEW.
In accordance with the positive genetic correlation
between body weight and egg weight, those birds with
the highest MBW (Groups 1 and 4) showed also the
highest AEW but hens in Group 1 were lighter and laid
heavier eggs than hens in Group 4. The remaining
groups (2 and 3), with similar average MEW, exhibited
significantly different MBW. These differences among
groups in their jointly considered body weight and egg
weight patterns were corroborated by the discriminant
analysis.

TABLE 4. Nonlinear least squares estimates of Weatherup and

Foster equation parameters for mean body weight-age data of
hens grouped by means of a principal component analysis1

Variable
Location A B r R2
(g)
1st quadrant 2,306.54 13,192.89 0.8625 0.962
2nd quadrant 1,876.74 124,671.75 0.7600 0.950
3rd quadrant 2,137.73 11,585.42 0.8718 0.983
4th quadrant 2,711.77 5,002.37 0.9258 0.991
FIGURE 2. Theoretical mean body weight-age curves of laying
1A = asymptotic body weight; B = range in body weight from t = 0 hens grouped by means of a principal component analysis. Group 1:
to A; r = maturing rate for body weight; and R2 = coefficient of First quadrant (⁄), Group 2: Second quadrant (+), Group 3: Third
determination. quadrant (◊), and Group 4: Fourth quadrant (π).
 BODY WEIGHT AND EGG WEIGHT IN LAYERS
TABLE 5. Nonlinear least squares estimates of Weatherup and
Foster equation parameters for mean egg weight-age data of
hens grouped by means of a principal component analysis1
Variable
Location A B r R2
(g)
1st quadrant 66.58 139.38 0.9212
2nd quadrant 56.52 436.39 0.8714
3rd quadrant 55.66 14,496.08 0.7629
4th quadrant 60.66 1,076.14 0.8426
1A = asymptotic egg weight; B = range in egg weight from t = 0 to
A; r = maturing rate for egg weight; and R2 = coefficient of
determination.
The results confirm the partial pleiotropic basis of the
genetic correlation between body weight and egg
weight, evincing the feasibility of applying an indepen-
dent selective pressure on each character; also, within
each trait, applying independent selection pressure on
maturing rate separately from asymptotic weight,
modifying both weight-age patterns at the same time.
Perhaps this type of strategy could be accomplished by
means of a biological selection index based on principal
component analysis equations as it has been proposed
by Godshalk and Timothy (1988). Although the heritable
basis of the various combinations of traits in the four
groups of hens herein described has not yet been
demonstrated, such a procedure would potentially
eliminate those problems associated with conventional
selection indexes (Hazel, 1943), such as errors in genetic
parameter estimates and difficulties in assigning relative
economic weight to traits.
FIGURE 3. Theoretical mean egg weight-age curves of laying hens
grouped by means of a principal component analysis. Group 1: First
quadrant (⁄), Group 2: Second quadrant (+), Group 3: Third quadrant
(◊), and Group 4: Fourth quadrant (π).
795
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