Use of structured antedependence models for the genetic

analysis of growth curves1

F. Jaffrezic2, E. Venot, D. Laloe, A. Vinet, and G. Renand

INRA Quantitative and Applied Genetics, 78352 Jouy-en-Josas Cedex, France

ABSTRACT: Growth curve analysis is an important
issue for many agricultural and laboratory species, for
both phenotypic and genetic studies. The aim of this
paper is to present the use of a novel statistical ap-
proach, namely the structured antedependence (SAD)
models, to deal with this issue. The basic idea of these
models is that an observation at time t can be explained
by the previous observations. These models are espe-
cially appropriate to deal with cumulative traits such
as growth, as BW at age t clearly depends on BW mea-
sures at ages (t 1), (t 2), etc. These models were
applied on an INRA experimental Charolais herd data
set. The data comprised BW records for 560 cows born
over an 11-yr period (from 1988 to 1998) from 60 sires
and 369 dams. The proposed SAD models were com-
pared with the well-known random regression (RR)
models that are already widely used in various areas
of longitudinal data analysis. It was found that the
SAD models fit the growth process better with far fewer
parameters than the RR models (9 instead of 16 covari-
ance parameters for the phenotypic analysis, and 14
instead of 21 for the genetic analysis). Despite this
smaller number of covariance parameters, the likeli-
hood value was found to be much higher with the SAD
vs. the RR models, with a difference of 262.9 for the
phenotypic analysis with a quartic polynomial for the
RR and 751.5 for the genetic analysis with a cubic poly-
nomial for both the genetic and environmental parts of
the RR model. The SAD models also proved to be better
able to interpolate missing values. Heritability, genetic,
and environmental correlation coefficients were esti-
mated for weights from birth to adulthood. The struc-
tured antedependence models proved, in this study, to
be very appropriate to model growth data in a parsimo-
nious and flexible way.

Key Words: Beef Cattle, Genetic Analysis, Growth Curves, Random Regression Models,
Structured Antedependence Models

2004 American Society of Animal Science. All rights reserved. J. Anim. Sci. 2004. 82:34653473

Introduction chain Monte Carlo methods to estimate the parameters,

Growth curve analysis is an important issue for many
agricultural species such as beef cattle, pigs, chicken,
and rabbit. In the case of genetic studies, a few method-
ologies have been proposed. Blasco et al. (2003) pro-
posed using parametric growth curves, such as the
Gompertz curve, that have interpretable parameters in
terms of growth rate and adult BW. Each parameter
of the curve is then decomposed into genetic and envi-
ronmental components, as in classical quantitative ge-
netics theory, and selection can then be made on these
parameters. This approach, however, leads to nonlinear
mixed models, and therefore requires the use of Markov
1
Thanks to W. G. Hill for very interesting comments and ideas.
Data were provided by the INRA experimental center located near
Bourges, France.
2 man, 2000). These models are especially appropriate
Correspondencee-mail: jaffrezic@dga2.jouy.inra.fr.
Received June 17, 2004.
Accepted September 7, 2004.
which can be quite time consuming, especially for large
data files.
On the other hand, Meyer (2002, 2004) proposed us-
ing random regression (RR) models (Diggle et al., 1994),
as in the classical longitudinal data analyses. This ap-
proach has already been extensively investigated in the
context of test-day records in dairy cattle and proved
to have a few drawbacks. In particular, polynomial
functions tend to show border effects that lead to the
prediction of unreasonable values at the extreme points
of the data. A possible way to overcome this drawback
would be to use spline functions instead of polynomials;
they do, however, still require a quite large number
of parameters.
Other approaches have been proposed in the context
of longitudinal data analyses, such as structured ante-
dependence (SAD) models (Nunez-Anton and Zimmer-
for cumulative traits, as the observation at time t is
defined as a function of the previous observations.

3465
3466
The aim of this paper was to investigate the use of
these SAD models for growth curve analysis and apply
them to the genetic analysis of an experimental beef
cattle data set.
Materials and Methods
Structured Antedependence Models
Let Yi(tj) be the observed body weight at age tj for
animal i. It is assumed that it can be decomposed into
genetic and environmental components, as follows:
Yi(tj) = (tj) + gi(tj) + ei(tj)
where (tj) corresponds to the fixed effects and includes
the mean curve in the population. Functions gi(tj) and
ei(tj) are Gaussian processes, which are independent of
one another, have an expected value of 0 at each age,
and covariance functions G(tj,tk) and E(tj,tk), respec-
tively, for two ages tj and tk. They represent the age-
dependent genetic and environmental deviations, re-
spectively, for animal i. It is assumed that J measure-
ment times are available with t1 < t2 < ... < tJ. These
ages are assumed to be the same for all the individuals;
however, the model is able to deal with missing values.
The idea of antedependence models, as originally pro-
posed by Gabriel (1962), is that an observation at time
tj can be explained by the previous observations. An
antedependence structure of order r is defined by the
fact that the kth observation (k > r) given the r preceding
ones is independent of all further observations (Gabriel,
1962). Generalizing this concept to genetic analysis, a
second-order structured antedependence model for the
genetic part gi(tj), for animal i, can be written as:
gi(t0) = gi(t0)
gi(t1) = 11 gi(t0) + gi(t1)
gi(tj) = 1j gi(tj 1)) + 2j gi(tj 2) + gi(tj)
for j 2.
Parameters 1j and 2j are termed antedependence
parameters. As originally proposed by Gabriel (1962),
these parameters were estimated for each time j. There-
fore, if J measurement times were considered, (J 1)
parameters 1j and (J 2) parameters 2j had to be
estimated. If the number of measurement times is large
and the order of antedependence high, that will lead to
a very large number of parameters to be estimated.
Therefore, Nunez-Anton and Zimmerman (2000) pro-
posed in their SAD models to decrease this number of
parameters by fitting a parametric function of the lag
time for these antedependence terms. For example,
when using an exponential function, for j = 1, ..., J:
1j (tj, tj1) = exp(1 [tj tj1]) and 2j (tj, tj2) = exp(2
[tj tj2])
Jaffrezic et al.
A further advantage of modeling the antedependence
parameters with a continuous function of the lag time
is that it allows dealing with unequally spaced data,
as well as unbalanced data. It is therefore not required
that the animals be weighed at regular time intervals
or that the times of measurement are the same for all
the animals. To be able to find an appropriate paramet-
ric curve, the time spacing should not be completely
random. The model can, for example, easily deal with
smaller time intervals at earlier rather than late ages.
Although antedependence models are based on an
idea similar to autoregressive models, due to the initial
condition gi(t0) = gi(t0), there is no constraint on the
[1] antedependence parameters 1j and 2j, whereas they
have to be between 1 and 1 for autoregressive models.
Therefore, any parametric function can be used to
model these parameters, depending on the biological
process studied.
Parameters gi(tj) are assumed normally distributed,
with mean 0 and variance g2(tj) that are termed inno-
vation variances. Gabriel (1962) originally suggested
estimating one innovation variance for each time of
measurement. This again leads to a quite large number
of parameters when many different times of measure-
ment are analyzed. In their structured antedependence
models, Nunez-Anton and Zimmerman (2000) again
suggested using a continuous function of time to model
these innovation variances, such as polynomials. For a
quadratic polynomial, for instance:
ln g2(tj) = a + b tj + c tj2 [5]
In this case, therefore, only three parameters will
have to be estimated, regardless of the number of times
of measurement J. This parametric modeling of the
error variances can also be related to the structural
[2] model proposed by Foulley and Quaas (1995). The use
of a polynomial function to model the variances might,
[3] as with the RR models, show some border effects. These
border effects will, however, be far less important for
[4] the SAD models than for the RR case as the estimated
covariance function of the process is not a simple func-
tion of these polynomials but rather a complex combina-
tion of the antedependence parameters and innovation
variances. In practice, simple parametric functions are
sufficient to adequately model these variances. When
possible, it is best to fit an unstructured antedepen-
dence model (UAD) as originally proposed by Gabriel
(1962), which will give a nonparametric estimation of
the antedependence parameters and innovation vari-
ances and will allow one to choose the most appropriate
parametric functions of time.
To relate this modeling to the genetic covariance func-
tion G(tj,tk), for two ages tj and tk, Pourahmadi (1999)
showed that a Cholesky decomposition of the inverse
of the covariance matrix can be easily calculated using
the antedependence parameters and innovation vari-
ances. If there are J measurement times, let G be the
 Genetic analysis of growth curves 3467
genetic covariance matrix, of dimension J J. It can Table 1. Likelihood values (Log L) and Bayesian informa-
be shown that: tion criterion (BIC) for the phenotypic analysis using dif-
ferent structured antedependence (SAD) models, with
G1 = LD1L [6] antedependence up to order 3, and with the antedepen-
dence parameters modeled as constant (const), linear (lin),
where L is a lower triangular matrix with 1 on the quadratic (quad), or exponential (exp) functions of time,
diagonal and negatives of the antedependence coeffi- different random regression (RR) models based on poly-
cients rj (r = 1, ..., R for SAD(R), j = 1, ..., J) below nomials up to order 4 (quartic), and a completely unstruc-
the diagonal entries, and D is a diagonal matrix with tured model (US) with a 10 10 covariance matrix
innovation variances j2(j = 1, ..., J) as components. An Modela Parametersb Log L BIC
interesting computational property is that the inverse
G1 is sparse. Indeed, for a second-order antedepen- US 55 2,003.4 1,766.9
SAD3 quad-lin-const 9 1,615.7 1,577.0
dence model, for instance, only the first two subdiago- SAD3 quad-const-const 8 1,608.3 1,573.9
nals are nonzero. Antedependence and innovation vari- SAD3 quad-lin-lin 10 1,616.9 1,573.9
ance parameters can be estimated by restricted maxi- SAD2 quad-lin 8 1,606.8 1,572.4
mum likelihood procedures. In the following example, SAD2 quad-const 7 1,598.2 1,568.1
SAD1 exp 5 1,553.1 1,531.6
these parameters were estimated using the OWN func-
SAD1 quad 6 1,551.8 1,526.0
tion of ASREML (Gilmour et al., 2002), which requires SAD2 const 5 1,527.0 1,505.5
specification of the covariance matrix and derivatives SAD3 const 6 1,529.5 1,503.7
with respect to each of the parameters. As presently SAD1 lin 5 1,524.6 1,503.1
implemented, it is necessary to build the J J covari- SAD1 const 4 1,503.0 1,485.8
RR quartic 16 1,352.8 1,284.0
ance matrix, which will be the same for all the animals.
RR cubic 11 1,063.9 1,016.6
The J times of measurement are therefore assumed to RR quad 7 763.7 733.6
be the same for all the individuals, and should not be RR linear 4 422.8 405.6
too large (20 at most). It is not required, however, that a
Numbers following SAD indicate the order of antedependence.
these ages should be equally spaced or that the animals b
Parameters = number of covariance parameters in the model.
have measurements at all times.

Presentation of the Data
Data analyzed in this study came from an INRA ex-
perimental Charolais herd (Mialon et al., 2001). The
data set comprised BW records for 560 cows born over
an 11-yr period (from 1988 to 1998) from 60 sires and
369 dams. Data were collected monthly from 1998 to
2003, and only measurements at 10 different ages were
considered here for each animal, at approximately 0,
112, 224, 364, 540, 720, 900, 1,260, 1,620 and 1,980 d.
Although the same ages were considered for each ani-
mal, they were unequally spaced and some records were
missing. The fixed effects were the year of birth of the
animal, twinning effect and the dam age at calving. To
obtain the most accurate fit, the mean curve in the
population was modeled nonparametrically by fitting
one mean at each age by including in the fixed effects
(tj) of Eq. [1] the variable age as a factor. This was
possible because only 10 measurements were consid-
ered for each animal and the data had very few missing
values. In other practical cases, however, it could be
better to use parametric or semiparametric functions
of age, such as spline functions.
and the antedependence parameters were either con-
sidered as constant, linear, quadratic, or exponential
functions of age. Random regression models based on
polynomials up to order 4 (quartic) were used.
Models were compared using the likelihood values
and the Bayesian information criterion (BIC; Schwarz,
1978): BIC = ln L 0.5 nc ln(N p), where ln L is the
restricted maximum likelihood value, nc is the number
of covariance parameters in the model, p is the number
of fixed effects (also equal to rank (X)), and N is the total
number of observations. The number of fixed effects in
the model was equal to 25 and the total number of
observations was 5,455.
This analysis was followed by a genetic study. An
animal model was used and the pedigree file comprised
807 animals. Several SAD and RR models were com-
pared for both the genetic and permanent environmen-
tal parts. As before, models were compared using the
likelihood values and the BIC criterion. The estimated
genetic parameters obtained at different ages for the
chosen models are presented in the next section.
Results

Analysis Phenotypic Analysis

First, a phenotypic analysis of the data was per-
formed to compare various different SAD and RR mod-
els. Here, models with antedependence up to order 3
were investigated. The innovation variances were as-
sumed to be changing as a quadratic function of age,
As shown in Table 1, it was found that even the
simplest first-order SAD model (with only four parame-
ters for the covariance structure) provided a better like-
lihood value than the quartic RR model (with 16 pa-
rameters).
3468
Figure 1. Estimated first-order antedependence param-
eter obtained nonparametrically with the unstructured
antedependence model (UAD(1)) and with a quadratic
polynomial function with the structured antedependence
model (SAD(1)).
The SAD model chosen in this phenotypic study,
based on the BIC criterion, is a third-order antedepen-
dence model that can be written for any animal i at age
t as follows:
yit = 1tyi(t 1) + 2tyi(t 2) + 3yi(t 3) + eit
where the antedependence parameters and innovation
variances were modeled as follows:
1t = 1.72 0.32 t + 0.02 t2, 2t =
0.07 + 0.04 t, 3 = 0.11 and
Var(eit) = exp(3.60 + 0.85 t 0.05 t2)
The 10 different ages considered here were coded as t =
1, ..., 10.
The choice of these polynomial functions to model the
antedependence parameters was based on a prelimi-
nary analysis using a first-order unstructured antede-
pendence model. Figure 1 presents the nonparametric
estimates for the 1t antedependence parameter using
the first-order unstructured antedependence model and
estimates obtained with a first-order SAD model using a
quadratic polynomial. It can be seen that the quadratic
function is appropriate to model this first antedepen-
dence parameter.
As pointed out in the Materials and Methods section,
it might also be possible in other practical examples to
use nonlinear functions, such as exponential functions
that have asymptotes. Estimation of these nonlinear
Jaffrezic et al.
terms can easily be implemented in ASREML (Gilmour
et al., 2002) using the OWN function; it simply requires
specifying the derivatives of the chosen function with
respect to each of the parameters.
The antedependence model presented in Eq. [7] re-
quired only nine parameters for the covariance struc-
ture and gave a likelihood value of 1,615.7, whereas
the most complicated RR model required 16 parame-
ters, with a likelihood value equal to 1,352.8.
Figure 2 gives the correlation functions estimated
with the chosen SAD and RR models. To evaluate the
goodness-of-fit of these functions, an unstructured co-
variance matrix (10 10) was also fitted to the data.
The estimated correlation function obtained with this
saturated model is also presented in Figure 2. One of the
main drawbacks of the RR models based on polynomial
functions can clearly be seen here, namely border effects
(Druet et al., 2003). In fact, although the global shape
of the correlation function seems to be close to the un-
structured one, a strongly negative correlation was esti-
mated at early ages, which seems to have no biological
meaning and is not observed with either the SAD or
unstructured model.
Conversely, Figure 3 gives the estimated variance
functions obtained with the different models, and they
all seem quite similar. As expected, the variances were
found to quite strongly increase with age (and BW).
It was found that both the chosen SAD and RR models
were able to fit individual phenotypic curves well when
all the observations were available, even for irregular
patterns. The main difficulty of the RR model seemed
to be the prediction of missing observations at early
ages (due to the poor estimation of the correlation func-
[7] tion at these ages). Figure 4 shows, for example, the
interpolations obtained with the different models for
an animal missing values at ages 112 and 224 d. Both
the SAD and the unstructured model predicted the
same values, whereas the RR model predicted lower
BW values (the difference was approximately equal to
20 kg at age 112 d). The accuracy of the covariance
structure is therefore particularly important for dealing
with missing values.
To check the interpolation abilities of both models at
early ages, observations at age 112 d were deleted from
the data for all the animals born since 1995, which
represented 220 animals and approximately 40% of the
population analyzed. To compare the predictions ob-
tained with the SAD and RR models to the actual BW
values at age 112 d for these animals, the Vonesh con-
cordance coefficient (rc) was used (Vonesh et al., 1996)
with the following formula:
i=1 (yi yi)2
M
rc = 1 [8]
i=1 (yi y)2 + i=1 (yi y)2 + M(y y)2
M M
where M is the number of animals with deleted values
(here M = 220), yi represents the actual observation for
 Genetic analysis of growth curves 3469

Figure 2. Estimated phenotypic correlation functions obtained with the unstructured (US) model and the structured
antedependent (SAD) and random regression (RR) models chosen according to the Bayesian information criterion
(BIC) values given in Table 1.

animal i at age 112 d, yi is the predicted value obtained
either with the SAD or RR model, and y and y are the
averages of the observed and predicted values, respec-
tively. This rc coefficient has values between 1 and 1,
with a perfect fit at 1 and a lack of fit for negative values.
When calculating this coefficient for the interpolation
of the 40% missing values at age 112 d, it was found
that the prediction ability was much better with the
SAD than with the RR model. In fact, the concordance
coefficient was found to be equal to 0.83 for the SAD
model for the predicted vs. actual observed BW values,
and was 0.44 for the RR model.
The differences in the interpolation abilities were less
important for other ages; for example, rc = 0.69 at 224
d of age for the SAD model compared with 0.58 for the
RR model, or rc = 0.74 at 540 d of age for the SAD model
compared with 0.70 for the RR model.
Finally, the rc coefficient was equal to 0.74 with the
SAD model compared with 0.69 with the RR model
for the latest age (1,980 d), showing a slightly better
extrapolation ability of the SAD model. Figure 5 illus-
trates this prediction ability for an animal with a clas-
sical growth curve. In this particular example, the phe-
notypic prediction obtained with the RR model was de-
creasing, whereas the BW for this animal would be
expected to remain stable, as predicted by the SAD
model, or to keep increasing slightly, as actually ob-
served.
Genetic Analysis
Model comparison results for the genetic analysis are
presented in Table 2. As for the phenotypic analysis, it
was found that the simplest first-order SAD model, with
a constant antedependence parameter, had a higher
likelihood value than a cubic RR model, while requiring
far fewer parameters for the covariance structure (eight
instead of 21 for the RR model). The best-fitting SAD
model regarding the likelihood value and the BIC crite-
rion was a second-order antedependence, with a qua-
dratic first-order antedependence parameter, a con-
stant second-order antedependence parameter, and
quadratic innovation variances for both the genetic and
environmental parts.
3470 Jaffrezic et al.

Figure 3. Estimated phenotypic variance functions ob-

tained with the unstructured (US) model and the struc-
tured antedependent (SAD) and random regression (RR)
models chosen according to the Bayesian information cri-
terion (BIC) values given in Table 1.
Figure 5. Observed and extrapolated phenotypic
growth curves obtained with the chosen structured ante-
dependent (SAD) and random regression (RR) models
for an animal with observations for age greater than 1,620
d deleted.

For the genetic part, the chosen model was as follows with parameter estimates equal to 1t = 1.29 0.11 t +
(for any animal i, at age t): 0.01 t2, 2 = 0.34, and Var(et) = 15.8 + 33.8 t 1.1 t2.
For the environmental part, the chosen model was
yit = 1t yi(t 1) + 2 yi(t 2) + eit [9] as follows:

yit = 1t yi(t 1) + 2 yi(t 2) + eit [10]

where 1t = 1.50 0.24 t + 0.01 t2, 2 = 0.05, and

Var(eit) = 157.4 + 174.4 t 3.5 t2.
Figure 6 shows the estimated genetic correlations
obtained with this SAD model and the cubic RR model.

Table 2. Likelihood values (Log L) and Bayesian informa-

tion criterion (BIC) for the genetic analysis
Modela Parametersb Log L BIC

SAD2 quad-constc 14 1,871.9 1,811.7

SAD2 const 10 1,635.7 1,592.7
SAD12 const 9 1,595.6 1,556.9
SAD1 const 8 1,588.7 1,554.3
SAD13 const 10 1,596.0 1,553.0
RR cubicd 21 1,120.4 1,030.0
RR quad 13 815.0 759.1
RR linear 7 470.2 440.1
a
The same model was used for both the genetic and environmental
parts.
b
Parameters = number of covariance parameters in the model.
Figure 4. Phenotypic interpolated growth curves ob- c
SAD2 quad-const corresponds to a second-order structured antede-
tained with the unstructured (US) model and the struc- pendence model with quadratic (quad) first-order antedependence
tured antedependent (SAD) and random regression (RR) parameter and constant (const) second order.
d
RR cubic corresponds to random regression model based on polyno-
models chosen as in Figures 2 and 3, for an animal with mial of order three (cubic) for both the genetic and environmental
missing observations at ages 112 and 224 d. parts.
 Genetic analysis of growth curves
Figure 6. Estimated genetic correlation functions ob-
tained with the structured antedependent (SAD) and ran-
dom regression (RR) models chosen according to the
Bayesian information criterion (BIC) values given in Ta-
ble 2.
The patterns were quite similar, although a more im-
portant drop was observed for the RR model at early
ages. These genetic correlations were equal to more
than 0.95 at late ages and approximately 0.4 at early
ages. On the other hand, for the environmental correla-
tions the same drawback was observed for the RR model
as in the phenotypic analysis. In fact, as shown in Fig-
ure 7, the correlations even became negative at very
early ages, although there seems to be no biological
reason for this. As expected, the environmental correla-
tions obtained with both methods were found to be
much lower than the genetic ones. They ranged from
about 0.1 for most ages, to around 0.6 for a few adjacent
3471
Figure 7. Estimated environmental correlation func-
tions obtained with the structured antedependent (SAD)
and random regression (RR) models chosen according to
the Bayesian information criterion (BIC) values given in
Table 2.
ages. Due to the very large number of parameters to
be estimated, the completely unstructured model (with
110 parameters) did not properly reach convergence in
this genetic study, and therefore no reference model
is available.
Figure 8 gives the estimation of the heritability over
age for the chosen SAD model. The pattern is character-
istic of heritability estimates found in the literature.
In a review of univariate estimates of heritability coef-
ficients, Koots et al. (1994) computed average values of
0.35, 0.27, 0.35, and 0.50 for birth weight, weaning
weight, yearling weight, and adult cow weight, respec-
3472 Jaffrezic et al.
Figure 8. Estimated heritability functions over age ob-
tained with the chosen structured antedependent (SAD)
and random regression (RR) models, as presented in Ta-
ble 2.
tively. A similar pattern was obtained by Bullock et al.
(1993) with field data records of Polled Hereford cows:
0.49, 0.24, 0.30, and 0.52, respectively. In two studies
of growth curves up to 2 yr of age, Meyer (2002, 2004)
also obtained a similar pattern for heritability coeffi-
cient estimates, with RR models. In the current study,
heritability of birth and adult cow weights (0.54 and
0.70) were among the highest estimates found in the
literature. Figure 8 also presents the heritability esti-
mates obtained with the cubic RR model; it was also
found to be increasing with age, although the heritabil-
ity estimated for the adult body weight was slightly
lower than with the SAD model (equal to 0.6). The main
difference in the pattern of these two heritability curves
is for the birth weight. The RR model estimated a very
low heritability at birth, equal to 0.05, whereas it is
expected to be much higher as previously found in the
literature (Bullock et al., 1993; Koots et al., 1994).
Discussion
Use of longitudinal models over the simple multitrait
model for growth curve analysis has many advantages.
In particular, they are able to capture the continuous
nature of the process, and therefore allow prediction of
the genetic values at any time, even when no measure-
ment is available. They also allow the analysis of unbal-
anced data and irregularly spaced measurements.
Among them, the SAD models in this study proved
to be appropriate to model growth data. They, in fact,
offer a parsimonious and flexible way of modeling this
process, and were found to be able to better fit the
covariance structure of the data than polynomial-based
random regression models. It was found in the pheno-
typic study that even the simplest first-order SAD
model (with only four parameters) provided a better fit
(higher likelihood value) than a fourth-order RR model
(with 16 parameters). Similar results were obtained in
the genetic study where a simple first-order SAD model,
for both the genetic and environmental parts (with
eight parameters), provided a better fit than a cubic
RR model (with 21 parameters).
The chosen SAD and RR models proved able to fit
individual phenotypic curves very well, when all the
observations were available. Differences were noted be-
tween the two methodologies for interpolating the miss-
ing values. Due to the poor estimation of the correlation
structure obtained with the RR models, they generally
predicted the missing values less accurately than the
SAD models.
To further improve the modeling of the covariance
structure, it would also be possible to model variance
heterogeneity due to other environmental factors, as
proposed by Foulley and Quaas (1995), and Robert-
Granie et al. (2002) for simple RR models. This is
straightforward in the SAD models, as a structural
model is already used to model the changes of the inno-
vation variances over time. Extension of this heteroge-
neous model can also be applied to the antedependence
parameters, to correct, for example, for preferential
treatments influencing the growth rate of some
animals.
A multivariate extension of the SAD models has been
presented by Jaffrezic et al. (2003), which would allow
one to study, for example, the dependence between BW
and feed intake, or any other time-dependent trait of
economic interest.
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