Genomic selection in French dairy sheep: main results and design to implement genomic breeding schemes

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Genomic selection in French dairy sheep: main results and design to implement genomic breeding schemes F. Barillet *, J.M. Astruc, G. Baloche, D. Buisson, G. lagriffoul et al. * * INRA - Toulouse, France francis.barillet@toulouse.inra.fr Session S2 Dairy Sheep and Goats, Berlin, Germany, 23 May 2014 39 th ICAR Session, Berlin, Germany, 2014

Sheep dairying in France Western Pyrenean Manech (red and black faced) and Basco-Béarnais breeds 432,000 ewes Roquefort area Lacaune breed 890,000 ewes Corsica island Corsican breed 83,000 ewes

French dairy sheep breeding schemes Breeding objectives and # AI rams progeny tested per breed and per year Lacaune FY,PY,F%,P% + SCC + UDDER 440 % AI in nucleus 85% Red Manech FY,PY,F%,P% 150 50% Black Manech FY,PY,F%,P% 30 45% Basco- Béarnaise FY,PY,F%,P% 50 50% Corsican MY 20 30% 0 100 200 300 400 500 AI progeny-tested rams per year

Size of the reference populations (end of 2013) AI rams phenotyped and genotyped with the Illumina Ovine SNP50 beadchip fundings : Roquefort in, Genomia and Degeram projects Breed # AI genotyped rams Years of birth # SNP available for GEBV estimations Lacaune 4,841 1999 to 2013 42,039 Basco-Béarnaise 509 2000 to 2012 Manech black faced 331 1999 to 2009 38,287 Manech red faced 1,424 1999 to 2009 4

Improvement of GEBV in French dairy sheep (from 2010 to 2013) GBLUP Evaluation in 2 steps Evaluation in 1 step Pseudo-ss-GBLUP (using all rams and daughter-yield-deviation) GBLUP Bayes Cπ PLS spls Duchemin et al, JDS 2012 ss-gblup (using all phenotypes and pedigrees as in animal model) Test of different GEBV methods including unknown parent groups Heterogeneity of variance within herd (in progress)

Accuracy of GEBV using GBLUP or other methods in Lacaune breed (1,806 in training population, and 681 born in 2007-2008 in validation population) Accuracy of EBV / GEBV Methods Milk Fat SCS EBV BLUP (parent average) 0.37 0.46 0.39 GEBV GBLUP 2 steps 0.42 0.56 0.44 Bayes Cπ 0.44 0.57 0.45 PLS 0.41 0.56 0.43 spls 0.42 0.56 0.43 GEBV (genomic) always better than EBV (parent average) Nearly no difference between GEBV methods Duchemin et al, JDS 2012, 95

Improvement of GEBV in French dairy sheep (from 2010 to 2013) GBLUP Evaluation in 2 steps Evaluation in 1 step Pseudo-ss-GBLUP (using all rams and daughter-yield-deviation) GBLUP Bayes Cπ PLS spls Duchemin et al, JDS 2012 ss-gblup (using all phenotypes and pedigrees as in animal model) including unknown parent groups Heterogeneity of variance within herd (in progress)

Accuracy of EBV / GEBV in dairy sheep : comparison between BLUP and GBLUP estimates Lacaune breed 2,900 progeny tested rams : - 1,593 in training population (born between 1999 & 2005) - 707 excluded (born in 2006 & 2007) - 592 (born in 2008 or 2009) in validation EBV (polygenic) based on BLUP : pseudo-blup GEBV (genomic) estimates using pseudo-ss-gblup (2 steps) or ss-gblup (1 step) Baloche et al, JDS 2014, 97

Use of reduced (2007) and full data sets (2011) to assess accuracy (according to Interbull recommendations) 450 400 350 300 250 200 150 100 50 0 1998 1999 2000 2001 2002 2003 2004 2005 2006 2007 2008 2009 Baloche et al, JDS 2014, 97 training Validation not used Birth year of the rams

Accuracy gain in GBLUP (GEBV) over BLUP (EBV) in dairy sheep Trait Accuracy (reliability) BLUP (PA) GBLUP 2 steps GBLUP 1 step Accuracy gain in GEBV over EBV (PA) Milk yield 0.32 0.43 0.47 0.15 Fat content 0.58 0.65 0.71 0.13 Protein content 0.54 0.62 0.70 0.16 SCS 0.49 0.59 0.59 0.10 Teat angle 0.47 0.58 0.66 0.19 Genomic (GEBV) always better than pedigree (EBV) But accuracy gain lower than in dairy cattle (large pop) Baloche et al, JDS 2014, 97

Summary of accuracy gain in (French) dairy cattle and sheep breeds Trait MILK YIELD Cattle (France) Sheep (France) Holstein Montbeliarde Lacaune Manech red faced Accuracy Parent average (EBV) Genomic (GEBV) Accuracy gain 0.33 0.60 0.33 0.30 0.47 0.27 0.32 0.47 0.15 0.29 0.43 0.14 Reference population (training) Effective size of the Population (breed) 4,000 1,200 1,900 1,000 45 125 250 170

From conventional to genomic breeding scheme in French dairy sheep Lacaune breed : 1 breed and 2 breeding schemes (companies) Pyrenean breeds : 1 company and 3 breeding schemes / 3 breeds (Basco-Bearnaise, Manech black faced, Manech red faced) Following presentation based on 1 Lacaune breeding scheme (1 company) Objective (defined by the managers of the breeding scheme) : is it possible to get at least a similar genetic gain without extra cost?

From conventional to genomic breeding scheme in French dairy sheep precision = accuracy g per year = precision x selection rate x σg generation interval Precision : comparable for conventional and genomic scheme Generation interval : quite similar for conventional and genomic scheme Selection rate : Objective to be reached comes mostly from possible selection rate in genomic versus conventional situation (breeding scheme), given the constraint of no-extra costs.

Classical AI scheme Genomic AI scheme (present) (near futur) Sampling rams(240 rams) Lay-off (230 rams) 115 Progeny-tested rams 70 45 20 Selection rate (r) at 2.5-year-old: 50% rams age 8-months 1.5 year 2.5 years 3.5 years 4.5 years 5.5 years Candidate genotyped rams Young rams Young rams Progeny-tested rams (genomic rams) (low) Selection rate (r2) at 2.5-year-old Genomic selection rate (r1) at 3-4-month-old Number of alive AI rams in the AI center Classical AI scheme Genomic AI scheme can be reduced thanks to 700 suppression of lay-off 14

Objective : is it possible to get at least a similar genetic gain without extra costs? Annual genetic gain ( g / year) in genomic versus classical scheme depending on possible genomic selection rate (r1) given the constraint of no extra-costs NEW COSTS Breeding and genotyping of a number of young candidate rams (1 to 4 month-old) suitable for genomic selection rate (r1) COST DECREASE Which reduction of the number of alive AI rams in the AI center (thanks to suppression of lay-off )?

Fine modelling Genomic AI scheme of physiological (near futur) constraints in the framework of extensive use of AI in fresh semen with highly seasoned period 8-months 1.5 year 2.5 years 3.5 years 4.5 years Candidate genotyped rams Young rams Young rams Progeny-tested rams (genomic rams) (low) Selection rate (r2) at 2.5-year-old Genomic selection rate (r1) at 3-4-month-old allows us to define a range of total alive AI rams (given the age structure in genomic situation) # alive AI rams in the AI center Genomic AI scheme 400 versus to 450 700-40 % 16 in classical scheme

Modeling a genomic scheme in dairy sheep (illustration with 1 Lacaune breeding company) Candidate genotyped rams 130 Young rams 110 Young rams 80 Progeny tested rams 60 40 Total : 420 AI in the AI center (genomic rams) 8-months-old 1.5-yr-old 2.5-yr-old 3.5-yr-old 4.5-yr-old Genomic selection rate (r1) at 3-4 month old : {1/3;1/4;1/5;1/6;1/7} Selection rate (r2) at 2.5-yr-old after progeny-test results {1;0.9;0.8;0.7}

Annual genetic gain (in genetic standard deviation ) according to r1 (genomic selection rate) and r2 (selection after progeny test) for one Lacaune breeding scheme impact of r1 much higher than impact of r2 annual genetic gain (nearly) always higher with genomic selection extra genetic gain much lower than in dairy cattle Buisson et al EAAP 2013, p.369

Co-evolution of annual genetic gain and costs according to the genomic selection rate (r1) scénario 2 scénario 1 G : + 10 to + 15 % Which decision? 1. Scenario 1 with r1=1/3 (current genotyping cost (115 ) 2. Scenario 2 with r1=1/4 (if genotyping cost (85 )

To take the decision to move to genomic selection in French dairy sheep Efficient current (classical) French dairy sheep schemes: close to their optimum. Are we confident in our modelling of French dairy sheep genomic schemes and expected annual genetic gain? We performed a genomic experiment to check / validate it.

Experimental design : AI rams born in 2011 from 46 sires 30 ram lamb per sire : 46 families et 928 genotyped candidates Genomic rams (GR) : chosen on their GBLUP at 3 month-old Genomic rams (GR) Classical rams (CR) 18 ram lambs per sire ±1/3 6 ram lambs per sire 3 ram lambs to be progeny-tested per sire genomic selection rate 1/2 standard & 1/2 developpement 12 ram lambs per sire 6 ram lambs to be progeny-tested per sire 21

Distribution of total merit index (TMI) ssgblup for CR and GR - at 2.5-year-old - TMI mean : - GR : + 201 points - CR : + 79 points Superiority of GR : +122 points (0.50 TMI std) Consequences : % rams above TMI 100 points (culling rate at 2.5-yr-old) -GR : 24 % -CR : 48 %

Conclusion Significant reduction (by 30 % to 40 %) of the number of alive AI rams in the case of genomic selection (GS) versus classical selection. More flexible GS breeding scheme allowing, at the same cost, an annual genetic gain increased by 10%-15 %, with a genomic selection rate (r1) at 3-month-old between 1/3 and ¼. The genomic selection experiment performed for AI rams born in 2011 confirms the relevance of selection rate (r1) and (r2) equal respectively to 0.3 and 0.80 in this GS situation experiment. Genomic selection will be implemented in 2015 in the French Lacaune breed and in a near futur in Pyrenean breeds (Basco- Béarnaise and Manech). 39 th ICAR Session, Berlin, Germay ny, 2014

Collaborations and fundings INRA Livestock Breeding Institute F. Barillet J.M. Astruc G. Baloche G. Lagriffoul D. Buisson H. Larroque A. Legarra Dairy sheep breeding companies C. Soulas, X. Aguerre, F. Fidele B. Giral-Viala, P. Boulenc P. Panis, P. Guibert G. Frégeat