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Genomic selection on rice 
Early generation selection in a recurrent 
selection breeding program within a 
synthetic population 
Since 1967 / Science to cultivate change 
Cécile Grenier 
Tuong-Vi Cao
Genomic selection 
Since 1967 / Science to cultivate change 
• Decreased genotyping costs and 
new statistical methods enable 
simultaneous estimation of all 
marker effects! 
• GS – a new form of MAS that 
estimates all marker effects across 
the whole genome to calculate 
genome estimated breeding values 
(GEBVs ) 
• Markers are not tested for 
significance – all markers are used 
in selection
Genomic selection on Rice 
Can genomic selection can be applied on rice synthetic population (SP) 
managed through recurrent selection (RS)? 
Can GS be adapted to Recurrent Genomic Selection (RGS)? 
Since 1967 / Science to cultivate change 
Theme 2 – Varietal Development
The breeding scheme 
Since 1967 / Science to cultivate change 
Recombination 
Candidate 
units 
Evaluation 
Phenotype in target 
environments 
Synthetic Population 
3000 S0 plants 
Varieties 
Selected 
units 
Evaluations
The SP derived training and 
breeding population 
Fixation through 
SSD for ~350 lines 
Since 1967 / Science to cultivate change 
Synthetic Population 
3000 S0 plants 343 S2:4 and S3:5 families 
Extraction of 
400 S0 plants 
Recombination 
(35 plants) 
Training Population 
Breeding Population
Testing the Feasibility of Genomic Selection through Cross-Validations 
Phenotypes (Y) Genotypes (X) 
Since 1967 / Science to cultivate change 
343 families 
(from a SP with 10 cycles of recombination) 
Whole Genome Regression Model
Since 1967 / Science to cultivate change 
GBS technology 
6,874 SNP with MAF ≥ 2.5% (1 marker every ~ 57 kb) 
4,098 SNP with MAF ≥ 10.0% (1 marker every ~ 95 kb) 
LD decay curve for chromosome 1 and MAF ≥ 10% 
For ½ initial r², the average extent of LD is ~ 0.639 Mb, i.e. at least 610 
markers are required to cover the whole genome
Heatmap (G matrix of 343 individuals with 
Un-rooted Neighbor Joining 
(dissimilarity matrix among 343 individuals with 6874 SNP) 
6874 SNP) 
Since 1967 / Science to cultivate change 
The genetic material
Since 1967 / Science to cultivate change 
The genetic material 
Evaluation of the 343 families (301 S2:4 and 42 S3:5) under a Lattice Design with 2 repetitions 
Panicle weight (h2=0.19) Grain yield (h2=0.30) 
Flowering date (h2=0.86) 
Plant height (h2=0.61)
Testing the Feasibility of Genomic Selection through Cross-Validations 
Phenotypes (Y) Genotypes (X) 
Since 1967 / Science to cultivate change 
343 families 
(from a SP with 10 cycles of recombination) 
Whole Genome Regression Model 
k-folds cross-validation: 
 100 samplings of Training Population (TP) and Validation Population (VP) 
 100 cor(y, X) 
 Mean of correlations: ‘Predictive ability of genomic selection’
GS in Rice synthetic populations 
Regression models 
G-BLUP 
Ridged Regression 
Bayesian LASSO 
Bayesian RR 
Since 1967 / Science to cultivate change 
Limit for r² MAF (%) No. SNP 
r² <= 0.75 
2.5 1758 
5 1158 
10 678 
r² <= 0.90 
2.5 4314 
5 3268 
10 2152 
r² <= 1.00 
2.5 6874 
5 5605 
10 4098 
k No. ind. 
[tst] 
3 114 
6 57 
9 38 
Incidence matrix 
choice of SNP markers based on LD and MAF 
FD (Flowering date) {h2 = 0.86} 
PH (Plant height) {h2 = 0.61} 
PW (Panicle weight) {h2 = 0.19} 
GY (Grain yield) {h2 = 0.30} 
k-folds cross-validation 
fraction k of the population (n=343) 
used for validation 
Traits
Statistical models for GS 
Criteria rrBLUP B-RR B-LASSO 
Variable selection No No Yes 
Marker effects All markers with same 
2 σ2, λ2 
Since 1967 / Science to cultivate change 
Penalized regressions 
– Parametric linear regression models (frequentist and Bayesians) 
• Ridge Regression (RR), Best Linear Unbiased Predictors (BLUP), Least Absolute Shrinkage and 
Selection Operator (LASSO), G-BLUP, RR-BLUP, LASSO, Bayesian RR, Bayesian LASSO… 
effect 
– Non-parametric nonlinear models 
• RKHS, NN, RBFNN 
All marker have an effect Some markers have null 
effect 
Parameter shrinkage 
of estimates effects 
Same extend of 
shrinkage 
Same extend of 
shrinkage 
Marker-specific shrinkage 
Hyper-parameters No σβ 
Distribution of effects Gaussian Gaussian Double exponential 
Best for… Trait controlled by many 
loci w. small effects 
Trait controlled by few loci 
varying in effect size
Regression model and marker effects 
Bayesian LASSO (MAF2.5 - r2≤0.75) with 9-fold CV -- Grain yield 
 cor(ŷ[tst], y[tst]) = 0.25 
 cor(ŷ, y) = 0.84 
Since 1967 / Science to cultivate change 
Marker Effects
Accuracy is function of trait genetic architecture, 
heritability and, for FD, of choice of markers 
Bayesian LASSO (9 X matrices) with 9-fold CV 
Grain yield {h2 = 0.30} Panicle weight {h2 = 0.19} 
Since 1967 / Science to cultivate change 
0.600 
0.500 
0.400 
0.300 
0.200 
0.100 
0.600 
0.500 
0.400 
0.300 
0.200 
0.100 
0.000 
0 2000 4000 6000 8000 
0.600 
0.500 
0.400 
0.300 
0.200 
0.100 
0.000 
0 2000 4000 6000 8000 
0.600 
0.500 
0.400 
0.300 
0.200 
0.100 
0.000 
0 2000 4000 6000 8000 
0.000 
0 2000 4000 6000 8000 
Flowering date {h2 = 0.86} 
Plant height {h2 = 0.61} 
No. SNP 
Accuracy (cor(ŷ, y)) 
No. SNP 
Accuracy (cor(ŷ, y))
Selection of markers (predictors) based on LD 
improved the accuracy for oligogenic traits 
Flowering date (9 X matrices, 3-fold CV, Ridged Regression) 
2.5% 
5.0% 
Since 1967 / Science to cultivate change 
7.5% 
10% 
0.50 
0.40 
0.30 
0.20 
0.10 
0.00 
-0.10 
0 1000 2000 3000 4000 5000 6000 7000 
Accuracy = corr(Yobs, Yhat) 
Number of SNP 
No. SNP 
r² <= 0.75 r² <= 0.80 r² <= 0.90 r² <= 1.00 
Series5 Series6 Series7 
Accuracy (cor(ŷ, y))
Slight superiority of the Bayesian Statistics 
Since 1967 / Science to cultivate change 
Grain Yield (9 X matrices with 9-fold CV) 
0.350 
0.300 
0.250 
0.200 
0.150 
0.100 
0.050 
0.000 
0 2000 4000 6000 8000 
BL 
BRR 
GBLUP 
RR 
No. SNP 
Accuracy (cor(ŷ, y))
The RS breeding scheme 
Since 1967 / Science to cultivate change 
Recombination 
Candidate 
units 
Evaluation 
Phenotype in target 
environments 
Synthetic Population 
3000 S0 plants 
Varieties 
Selected 
units 
Evaluations
The RGS breeding scheme 
Since 1967 / Science to cultivate change 
Recombination 
Candidate 
units 
Whole 
Genome 
Genotyping 
Breeding Population 
3000 S0 plants 
Promising lines 
Selected 
units 
GEBVs 
Genomic 
prediction 
MET Evaluations 
GS models 
Evaluations 
Training Population New varieties
Since 1967 / Science to cultivate change 
Conclusions 
• Yes, GS on rice synthetic population is feasible! 
• Although not fantastic accuracies were achieved, it was 1 site, 1 year and a 
first promising result 
• Small accuracy may still be worth considering the cost of field evaluation, 
the gain in time to select during the off-season and the possibility to apply 
stronger selection intensity 
• Soon to come: 
 More data, more sites, and more adequate statistics (experimental 
design and multi-site evaluations accounted in the model) for 
nonparametric non linear models 
 GS on the breeding population using the entire training population 
to develop the genomic prediction model 
 Maximizing the benefit of GS on earlier generation of the RS 
scheme (S0 generation)

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Genomic selection on rice

  • 1. Genomic selection on rice Early generation selection in a recurrent selection breeding program within a synthetic population Since 1967 / Science to cultivate change Cécile Grenier Tuong-Vi Cao
  • 2. Genomic selection Since 1967 / Science to cultivate change • Decreased genotyping costs and new statistical methods enable simultaneous estimation of all marker effects! • GS – a new form of MAS that estimates all marker effects across the whole genome to calculate genome estimated breeding values (GEBVs ) • Markers are not tested for significance – all markers are used in selection
  • 3. Genomic selection on Rice Can genomic selection can be applied on rice synthetic population (SP) managed through recurrent selection (RS)? Can GS be adapted to Recurrent Genomic Selection (RGS)? Since 1967 / Science to cultivate change Theme 2 – Varietal Development
  • 4. The breeding scheme Since 1967 / Science to cultivate change Recombination Candidate units Evaluation Phenotype in target environments Synthetic Population 3000 S0 plants Varieties Selected units Evaluations
  • 5. The SP derived training and breeding population Fixation through SSD for ~350 lines Since 1967 / Science to cultivate change Synthetic Population 3000 S0 plants 343 S2:4 and S3:5 families Extraction of 400 S0 plants Recombination (35 plants) Training Population Breeding Population
  • 6. Testing the Feasibility of Genomic Selection through Cross-Validations Phenotypes (Y) Genotypes (X) Since 1967 / Science to cultivate change 343 families (from a SP with 10 cycles of recombination) Whole Genome Regression Model
  • 7. Since 1967 / Science to cultivate change GBS technology 6,874 SNP with MAF ≥ 2.5% (1 marker every ~ 57 kb) 4,098 SNP with MAF ≥ 10.0% (1 marker every ~ 95 kb) LD decay curve for chromosome 1 and MAF ≥ 10% For ½ initial r², the average extent of LD is ~ 0.639 Mb, i.e. at least 610 markers are required to cover the whole genome
  • 8. Heatmap (G matrix of 343 individuals with Un-rooted Neighbor Joining (dissimilarity matrix among 343 individuals with 6874 SNP) 6874 SNP) Since 1967 / Science to cultivate change The genetic material
  • 9. Since 1967 / Science to cultivate change The genetic material Evaluation of the 343 families (301 S2:4 and 42 S3:5) under a Lattice Design with 2 repetitions Panicle weight (h2=0.19) Grain yield (h2=0.30) Flowering date (h2=0.86) Plant height (h2=0.61)
  • 10. Testing the Feasibility of Genomic Selection through Cross-Validations Phenotypes (Y) Genotypes (X) Since 1967 / Science to cultivate change 343 families (from a SP with 10 cycles of recombination) Whole Genome Regression Model k-folds cross-validation:  100 samplings of Training Population (TP) and Validation Population (VP)  100 cor(y, X)  Mean of correlations: ‘Predictive ability of genomic selection’
  • 11. GS in Rice synthetic populations Regression models G-BLUP Ridged Regression Bayesian LASSO Bayesian RR Since 1967 / Science to cultivate change Limit for r² MAF (%) No. SNP r² <= 0.75 2.5 1758 5 1158 10 678 r² <= 0.90 2.5 4314 5 3268 10 2152 r² <= 1.00 2.5 6874 5 5605 10 4098 k No. ind. [tst] 3 114 6 57 9 38 Incidence matrix choice of SNP markers based on LD and MAF FD (Flowering date) {h2 = 0.86} PH (Plant height) {h2 = 0.61} PW (Panicle weight) {h2 = 0.19} GY (Grain yield) {h2 = 0.30} k-folds cross-validation fraction k of the population (n=343) used for validation Traits
  • 12. Statistical models for GS Criteria rrBLUP B-RR B-LASSO Variable selection No No Yes Marker effects All markers with same 2 σ2, λ2 Since 1967 / Science to cultivate change Penalized regressions – Parametric linear regression models (frequentist and Bayesians) • Ridge Regression (RR), Best Linear Unbiased Predictors (BLUP), Least Absolute Shrinkage and Selection Operator (LASSO), G-BLUP, RR-BLUP, LASSO, Bayesian RR, Bayesian LASSO… effect – Non-parametric nonlinear models • RKHS, NN, RBFNN All marker have an effect Some markers have null effect Parameter shrinkage of estimates effects Same extend of shrinkage Same extend of shrinkage Marker-specific shrinkage Hyper-parameters No σβ Distribution of effects Gaussian Gaussian Double exponential Best for… Trait controlled by many loci w. small effects Trait controlled by few loci varying in effect size
  • 13. Regression model and marker effects Bayesian LASSO (MAF2.5 - r2≤0.75) with 9-fold CV -- Grain yield  cor(ŷ[tst], y[tst]) = 0.25  cor(ŷ, y) = 0.84 Since 1967 / Science to cultivate change Marker Effects
  • 14. Accuracy is function of trait genetic architecture, heritability and, for FD, of choice of markers Bayesian LASSO (9 X matrices) with 9-fold CV Grain yield {h2 = 0.30} Panicle weight {h2 = 0.19} Since 1967 / Science to cultivate change 0.600 0.500 0.400 0.300 0.200 0.100 0.600 0.500 0.400 0.300 0.200 0.100 0.000 0 2000 4000 6000 8000 0.600 0.500 0.400 0.300 0.200 0.100 0.000 0 2000 4000 6000 8000 0.600 0.500 0.400 0.300 0.200 0.100 0.000 0 2000 4000 6000 8000 0.000 0 2000 4000 6000 8000 Flowering date {h2 = 0.86} Plant height {h2 = 0.61} No. SNP Accuracy (cor(ŷ, y)) No. SNP Accuracy (cor(ŷ, y))
  • 15. Selection of markers (predictors) based on LD improved the accuracy for oligogenic traits Flowering date (9 X matrices, 3-fold CV, Ridged Regression) 2.5% 5.0% Since 1967 / Science to cultivate change 7.5% 10% 0.50 0.40 0.30 0.20 0.10 0.00 -0.10 0 1000 2000 3000 4000 5000 6000 7000 Accuracy = corr(Yobs, Yhat) Number of SNP No. SNP r² <= 0.75 r² <= 0.80 r² <= 0.90 r² <= 1.00 Series5 Series6 Series7 Accuracy (cor(ŷ, y))
  • 16. Slight superiority of the Bayesian Statistics Since 1967 / Science to cultivate change Grain Yield (9 X matrices with 9-fold CV) 0.350 0.300 0.250 0.200 0.150 0.100 0.050 0.000 0 2000 4000 6000 8000 BL BRR GBLUP RR No. SNP Accuracy (cor(ŷ, y))
  • 17. The RS breeding scheme Since 1967 / Science to cultivate change Recombination Candidate units Evaluation Phenotype in target environments Synthetic Population 3000 S0 plants Varieties Selected units Evaluations
  • 18. The RGS breeding scheme Since 1967 / Science to cultivate change Recombination Candidate units Whole Genome Genotyping Breeding Population 3000 S0 plants Promising lines Selected units GEBVs Genomic prediction MET Evaluations GS models Evaluations Training Population New varieties
  • 19. Since 1967 / Science to cultivate change Conclusions • Yes, GS on rice synthetic population is feasible! • Although not fantastic accuracies were achieved, it was 1 site, 1 year and a first promising result • Small accuracy may still be worth considering the cost of field evaluation, the gain in time to select during the off-season and the possibility to apply stronger selection intensity • Soon to come:  More data, more sites, and more adequate statistics (experimental design and multi-site evaluations accounted in the model) for nonparametric non linear models  GS on the breeding population using the entire training population to develop the genomic prediction model  Maximizing the benefit of GS on earlier generation of the RS scheme (S0 generation)