COMPARISION OF STATISTICAL
MODELS
FOR DAYS TO 114 kg AND BACKFAT DEPTH
M.T. See
Summary
Five models were studied for the
genetic evaluation of backfat depth and days to 114 kg in Duroc swine. Models
were compared using likelihood ratio tests on a chi-square distribution. The
results clearly indicated the importance of including the litter effect. Results also indicated that for
days to 114 kg the model providing the best fit (p < .001) included all
random effects, while for backfat (p < .005) pe was not included. However,
permanent environmental effects account for only 1% of the variation for days
and can probably be safely ignored. In
addition, the complete model resulted in markedly larger heritability estimates
due in part to the large negative genetic correlations observed between direct
and maternal effects. The data structure associated with the estimation of
this correlation should be considered when selecting a model for genetic
evaluations.
Introduction
The objective of this project was to evaluate the
suitability of differing animal models for the genetic evaluation of growth
rate and backfat depth in swine. Most evaluations of backfat depth and days to
114 kg from swine utilize a model that includes direct genetic and litter as random
effects. Applying the model with the best fit should result in the most accurate genetic
evaluation. Therefore more complete models should be considered. However, inaccuracies due to data structure
and computational complexity should be considered.
Material and Methods
Data were provided by the United
Duroc Swine Registry. The analysis dataset consisted of 44,223 records, 1677
contemporary groups and 220 levels for sex(herd). Five single trait animal models (Table 1) were compared for days
and backfat depth. The fixed part of
the model (contemporary group and sex(herd) was constant across all five models
(Table 1).
(Co)variance components were
estimated using the MTDFREML computer programs. This set of programs determines the best set of parameter
estimates by minimizing -2 times the restricted log likelihood function (L), that
is, -2L =
constant + log|R| + log|C| + y'Py where C is the
full-rank coefficient matrix for the mixed model equations and y'Py is
the weighted sums of squares for the residuals. A likelihood ratio test was used to compare the six models. This statistical procedure consists of
subtracting the value of -2L from the
model with more parameters from -2L corresponding to the model with fewer parameters. The difference was compared to a chi-square
distribution with degrees of freedom equal to the difference in the number of
parameters estimated for the two models.
The restricted log likelihood function, order and number of iterations
(Table 2) were obtained at convergence of the MTDFREML iterative process.
Table 1.
Models
|
|
Model1
|
s2g
|
s2m
|
sg,m
|
s2c
|
s2pe
|
s2e
|
|
1
|
g+e
|
x
|
|
|
|
|
x
|
|
2
|
g+c+e
|
x
|
|
|
x
|
|
x
|
|
3
|
g+m+c+e
|
x
|
x
|
|
x
|
|
x
|
|
4
|
g+m+r+c+e
|
x
|
x
|
x
|
x
|
|
x
|
|
5
|
g+m+r+c+pe+e
|
x
|
x
|
x
|
x
|
x
|
x
|
1 g = random genetic effect, m = random maternal
genetic effect, c = random litter effect, pe = random effect of permanent
maternal environment, and e= random residual effect. |
Table 2. Model information and
likelihood estimates.
|
|
|
Iterations
|
|
-2 log
likelihood
|
|
|
|
Order
|
BF
|
Days
|
BF
|
Days
|
Parameters
|
|
1
|
52338
|
26
|
31
|
142717.0
|
271586
|
2
|
|
2
|
64627
|
49
|
43
|
142420.6
|
268962
|
3
|
|
3
|
115068
|
73
|
79
|
142420.3
|
268960
|
4
|
|
4
|
115068
|
140
|
198
|
142403.1
|
268869
|
5
|
|
5
|
121983
|
278
|
410
|
142403.1
|
268770
|
6
|
|
|
|
|
|
|
|
|
Results and Discussion
Likelihood ratio tests (Table 3) indicate that for days to
114 kg the model providing the best fit (p < .001) included all random
effects, while for backfat (p < .005) pe was not included. However,
permanent environmental effects account for only 1% of the variation for days
and can probably be safely ignored. Genetic parameters estimated from all models
are shown in Table 4 for days and Table 5 for backfat depth. Heritability
estimates from the best fitting models were .45 for days and .43 for backfat
depth. Heritability for maternal genetic effects was .10 and .03 for days and
backfat depth. Heritability estimates from the Model 2 that balances fit with
computational ease and data structure were .29 for days and .35 for backfat.
Table 3. Likelihood ratio test (LRT) between models for
days
(upper off-diagonals) and backfat depth (lower off-diagonals) models.
|
|
Model
|
1
|
2
|
3
|
4
|
5
|
|
1
|
g+e
|
-
|
**
|
**
|
**
|
**
|
|
2
|
g+c+e
|
**
|
-
|
NS
|
**
|
**
|
|
3
|
g+m+c+e
|
**
|
NS
|
-
|
**
|
**
|
|
4
|
g+m+r+c+e
|
**
|
*
|
**
|
-
|
**
|
|
5
|
g+m+r+c+pe+e
|
**
|
*
|
*
|
NS
|
-
|
|
NS p > .1
* p < .005
** p < .001 |
Table 4.
Genetic parameters for days to 114 kg in Duroc swine.
|
|
Model
|
h2
|
m2
|
c2
|
pe2
|
rg,m
|
|
1
|
g+e
|
.55
|
|
|
|
|
|
2
|
g+c+e
|
.29
|
|
.23
|
|
|
|
3
|
g+m+c+e
|
.29
|
.01
|
.23
|
|
|
|
4
|
g+m+r+c+e
|
.45
|
.11
|
.23
|
|
-.76
|
|
5
|
g+m+r+c+pe+e
|
.45
|
.10
|
.23
|
.01
|
-.78
|
Table 5.
Genetic parameters for adjusted backfat depth in Duroc swine.
|
|
Model
|
h2
|
m2
|
c2
|
pe2
|
rg,m
|
|
1
|
g+e
|
.46
|
|
|
|
|
|
2
|
g+c+e
|
.35
|
|
.08
|
|
|
|
3
|
g+m+c+e
|
.34
|
.003
|
.07
|
|
|
|
4
|
g+m+r+c+e
|
.43
|
.03
|
.07
|
|
-.53
|
|
5
|
g+m+r+c+pe+e
|
.43
|
.03
|
.07
|
.00
|
-.53
|
When maternal effects are included in the model the
heritability is greatly increased, 55% for days and 23% for backfat. This
increase in heritability estimate would be expected to create a wider
distribution among EPDs, resulting in more desirable numbers for trait
leaders.
The results clearly indicated the importance of including
the litter effect (Model 2). However, to
accurately estimate litter effects sows should have more than 1 litter. In this data 60% of the sows were
represented by 1 litter. Consideration needs to be given to data structure and
multiple litters per nucleus female. Model 3 was fit only to describe the observed increase in
heritability from adding maternal genetic effects and the result do not
significantly differ from model 2. The
results indicate that the large negative genetic correlation between direct and
maternal effects causes the increase in both direct and maternal heritabilites.
To accurately estimate maternal genetic effects and correlation between direct
and maternal effects sires should have multiple daughters with measured progeny
and daughters should have multiple litters with measured offspring. This also
raises concerns over contemporary group structure and numbers of records per
female and sire.
It should also be considered that Model 2 is more
conservative by not taking into account
maternal and permanent environmental effects that are prone to errors due to
data structure. In addition, Model 2 is
easier computationally. Therefore careful consideration should be given to both
Model 2 and Model 4 when selecting a model for genetic evaluations of
post-weaning traits in swine.
Implications
Applying the model with the best
fit should result in the most accurate genetic evaluation. However,
inaccuracies due to data structure and computational complexity should be
considered. When maternal effects are included in the model the
heritability is greatly increased, 55% for days and 23% for backfat. This
result is due in part to a large negative genetic correlation between direct
and maternal effects. The accuracy of the estimation of this correlation should
be considered when selecting a model