The joint regression of Finlay and Wilkinson (1963) and Eberhart and Russell (1966) and the stability variance of Shukla (1972) are the most used measures of adaptability and stability. Their classical estimates come with tests, not with the uncertainty a breeder needs to choose. bayes_stability() fits the three models by Gibbs sampling, so every slope, deviation variance and stability variance has a posterior distribution. This vignette gives the model, the sampler, an analysis of Maiz, a comparison of the three models and the safety-first index of Eskridge (1990).
2 Model
For genotype \(i\), environment \(j\) and plot \(l\), \[y_{ijl} = \mu + g_i + (1 + b_i)h_j + d_{ij} + e_{ijl}, \qquad d_{ij} \sim N(0, \sigma^2_{d,i}), \quad e_{ijl}
\sim N(0, \sigma^2),\] with \(\sum_i g_i = \sum_j h_j = \sum_i b_i = 0\). The environmental index \(h_j\) is a parameter, estimated jointly with the others, as in the Bayesian Finlay-Wilkinson regression of Lian and de los Campos (2016). The classical analysis plugs in the environment means instead and so ignores their error. The slope of genotype \(i\) is \(1 + b_i\), with average 1.
.model
Slopes
Deviation variance
Classical counterpart
"ER"
yes
\(\sigma^2_{d,i}\) for every genotype
Eberhart and Russell (1966); Bayesian version of Oliveira et al. (2018)
"FW"
yes
one \(\sigma^2_d\)
Finlay and Wilkinson (1963)
"Shukla"
no (\(b_i = 0\))
\(\sigma^2_{d,i}\) for every genotype: the stability variance
Shukla (1972); Edwards and Jannink (2006)
The replicates separate the deviations \(d_{ij}\) from the plot error \(e_{ijl}\), so the data must have replicates.
2.1 Priors
bam_stab_priors() sets normal priors on \(\mu\) and on the contrasts \(\mK_G\mathbf{g}\), \(\mK_E\mathbf{h}\) and \(\mK_G\mathbf{b}\) (so the sums are zero), and scaled inverse chi-squared priors on \(\sigma^2\) and \(\sigma^2_{d,i}\). The defaults are vague and in the unit of the trait: \(10^4\) times the variance of the response for the effects, and the least-squares variances as scales with one degree of freedom.
Priors of the Bayesian stability models
mu ~ N(0, 1e4 x var(y))
K_G g ~ N(0, 1e4 x var(y) I)
K_E h ~ N(0, 1e4 x var(y) I)
K_G b ~ N(0, 100 I) (slopes 1 + b_i)
sigma^2 ~ Inv-chi^2(1, within-cell mean square)
sigma_d^2 ~ Inv-chi^2(1, least-squares deviation variance)
2.2 The sampler
Every conditional distribution is normal or scaled inverse chi-squared. Drawing the deviations \(d_{ij}\) one at a time next to the genotype effects would mix slowly, because the mean of the deviations of a genotype and its effect \(g_i\) trade off. The sampler therefore draws \(\mu\), \(\mathbf{g}\), \(\mathbf{h}\) and \(\mathbf{b}\) with the deviations integrated out: each observed cell mean then has variance \(\sigma^2_{d,i} + \sigma^2/n_{ij}\). The deviations come next, then the variances. This is a partially collapsed Gibbs sampler: each sweep is still an exact draw from the posterior. Unbalanced data need nothing more, as the weights use the number of plots of every cell, and an empty cell has weight zero. The sampler is compiled; vignette 13 checks it against a plain R version.
The slopes of genotypes 4 and 5 lie above 1 with 95% posterior probability: they respond more than the average genotype to good environments. The slope of 8 lies below 1. The posterior probability that a slope exceeds 1 is in the column prob_slope_above_1. A breeder can use it directly, which a test of \(b_i = 1\) does not allow.
plot(x = fs, which ="slopes")
plot(x = fs, which ="lines")
plot(x = fs, which ="deviations")
The deviation variances are poorly determined from 20 environments, so their HPD intervals are wide and overlap. The Bayesian analysis shows this directly; a classical F test of \(\sigma^2_{d,i} = 0\) would hide it.
The posterior means of the slopes are close to the least-squares slopes (correlation 0.997). The deviation variances differ more. The posterior of a variance estimated from 20 environments is skewed to the right, so its mean lies above its mode. The prior, with its scale at the pooled least-squares value, also pulls the small variances up. The order of the genotypes changes less (rank correlation 0.73).
4 The safety-first index
Eskridge (1990) ranks genotypes by a lower quantile of their yield across environments, \(\mu + g_i - z_{1-\alpha}\sqrt{V_i}\), where \(V_i\) is the variance of genotype \(i\) across environments under the model:
Model
\(V_i\)
Eberhart-Russell
\(b_i^2 s^2_h (1 - 1/E) + \sigma^2_{d,i}\)
Finlay-Wilkinson
\(b_i^2 s^2_h (1 - 1/E)\)
Shukla
\(s^2_h + \sigma^2_{d,i}\)
Here \(s^2_h\) is the variance of the environmental index. The index is computed on every draw, so it has an HPD interval and every genotype has a probability of being the best on it. Here 6 is most probably the best (0.84).
s[order(-safety_first), list( Gen , mean , safety_first , safety_first_lower , safety_first_upper , prob_best_safety )]
Comparison of Bayesian models (smaller is better)
model terms variances DIC pD WAIC p_waic
<char> <int> <char> <num> <num> <num> <num>
1: ER NA deviation per genotype 11893.02 104.1660 11897.91 97.2984
2: FW NA one deviation variance 11896.06 101.2211 11901.92 95.4625
3: Shukla NA interaction per genotype 11897.47 108.9504 11903.37 101.7497
se lambda_last_lower lambda_last_mean dWAIC dWAIC_se
<num> <num> <num> <num> <num>
1: 46.1831 NA NA 0.0000 0.0000
2: 45.9799 NA NA 4.0128 5.0903
3: 45.4214 NA NA 5.4558 7.7438
Sources: Spiegelhalter et al. (2002); Watanabe (2010); Vehtari et al. (2017).
Full references: bam_cite(x).
WAIC prefers the ER model; the next, FW, is 4 above it (0.8 standard errors of the difference). The standard errors do not separate the three models here; the choice can follow the question, the slopes for adaptability and the variances for stability. The models are fitted to the same plots, so they can also be compared with the AMMI and GGE fits of bayes_ammi() (vignette 9).
6 The stability fit and the decision tools
The draws of the cell means \(\mu + g_i + (1 + b_i)h_j + d_{ij}\) feed the decision tools of vignette 9:
bam_interpret(x = fs , higher_better =TRUE , top =5 )
Bayesian Eberhart-Russell stability model
Best by the safety-first index (alpha = 0.05): 6, 5, 9, 3 and 4; most
probably the best: 6 (0.84).
Findings:
- The chains have converged (largest potential scale reduction factor 1.017;
smallest effective sample size of the main parameters 574).
- Slopes whose HPD interval lies above 1 (respond more than average to good
environments): 4 and 5; below 1 (respond less, relatively better in poor
environments): 8.
- Smallest deviation variance: 6 (77,600); largest: 1 (483,000).
- Share of each genotype's variation explained by the regression (R2): 1
0.73, 2 0.87, 3 0.92, 4 0.94, 5 0.95, 6 0.96, 7 0.88, 8 0.57, 9 0.94.
Recommendations:
- For a wide recommendation choose genotypes with a high mean, a slope near 1
and a small deviation variance; for favourable environments a slope above 1
with a high mean.
- The safety-first index combines the two: 6, 5 and 9 lead.
Cautions:
- The environmental index is estimated from the same trial; with few
environments the slopes are uncertain (see the HPD intervals).
Sources:
- Eberhart, S. A. and Russell, W. A. (1966). Stability parameters for
comparing varieties. Crop Science, 6(1), 36-40.
doi:10.2135/cropsci1966.0011183X000600010011x
- Oliveira, T. R. A., Carvalho, H. W. L., Nascimento, M., Costa, E. F. N.,
Amaral Junior, A. T., Gravina, G. A. and Carvalho Filho, J. L. S. (2018).
The Eberhart and Russell's Bayesian method used as an instrument to select
maize hybrids. Euphytica, 214, 64. doi:10.1007/s10681-018-2146-y
- Lian, L. and de los Campos, G. (2016). FW: an R package for
Finlay-Wilkinson regression that incorporates genomic/pedigree information
and covariance structures between environments. G3: Genes, Genomes,
Genetics, 6(3), 589-597. doi:10.1534/g3.115.026328
- Eskridge, K. M. (1990). Selection of stable cultivars using a safety-first
rule. Crop Science, 30, 369-374.
- Spiegelhalter, D. J., Best, N. G., Carlin, B. P. and van der Linde, A.
(2002). Bayesian measures of model complexity and fit. Journal of the Royal
Statistical Society B, 64(4), 583-639. doi:10.1111/1467-9868.00353
- Watanabe, S. (2010). Asymptotic equivalence of Bayes cross validation and
widely applicable information criterion in singular learning theory.
Journal of Machine Learning Research, 11, 3571-3594.
bam_cite(x = fs , style =c("text", "bibtex")[1] )
Eberhart, S. A. and Russell, W. A. (1966). Stability parameters for comparing varieties. Crop Science, 6(1), 36-40. doi:10.2135/cropsci1966.0011183X000600010011x
Oliveira, T. R. A., Carvalho, H. W. L., Nascimento, M., Costa, E. F. N., Amaral Junior, A. T., Gravina, G. A. and Carvalho Filho, J. L. S. (2018). The Eberhart and Russell's Bayesian method used as an instrument to select maize hybrids. Euphytica, 214, 64. doi:10.1007/s10681-018-2146-y
Lian, L. and de los Campos, G. (2016). FW: an R package for Finlay-Wilkinson regression that incorporates genomic/pedigree information and covariance structures between environments. G3: Genes, Genomes, Genetics, 6(3), 589-597. doi:10.1534/g3.115.026328
Eskridge, K. M. (1990). Selection of stable cultivars using a safety-first rule. Crop Science, 30, 369-374.
Spiegelhalter, D. J., Best, N. G., Carlin, B. P. and van der Linde, A. (2002). Bayesian measures of model complexity and fit. Journal of the Royal Statistical Society B, 64(4), 583-639. doi:10.1111/1467-9868.00353
Watanabe, S. (2010). Asymptotic equivalence of Bayes cross validation and widely applicable information criterion in singular learning theory. Journal of Machine Learning Research, 11, 3571-3594.
8 References
Finlay, K. W. and Wilkinson, G. N. (1963). The analysis of adaptation in a plant-breeding programme. Australian Journal of Agricultural Research, 14(6), 742–754. doi:10.1071/AR9630742
Eberhart, S. A. and Russell, W. A. (1966). Stability parameters for comparing varieties. Crop Science, 6(1), 36–40. doi:10.2135/cropsci1966.0011183X000600010011x
Shukla, G. K. (1972). Some statistical aspects of partitioning genotype-environmental components of variability. Heredity, 29(2), 237–245.
Eskridge, K. M. (1990). Selection of stable cultivars using a safety-first rule. Crop Science, 30, 369–374.
Lian, L. and de los Campos, G. (2016). FW: an R package for Finlay-Wilkinson regression that incorporates genomic/pedigree information and covariance structures between environments. G3: Genes, Genomes, Genetics, 6(3), 589–597. doi:10.1534/g3.115.026328
Oliveira, T. R. A., Carvalho, H. W. L., Nascimento, M., Costa, E. F. N., Amaral Junior, A. T., Gravina, G. A. and Carvalho Filho, J. L. S. (2018). The Eberhart and Russell’s Bayesian method used as an instrument to select maize hybrids. Euphytica, 214, 64. doi:10.1007/s10681-018-2146-y
Edwards, J. W. and Jannink, J.-L. (2006). Bayesian modeling of heterogeneous error and genotype x environment interaction variances. Crop Science, 46(2), 820. doi:10.2135/cropsci2005.0164