The search for the genetic determinism of prolificacy variability in sheep has evidenced several major mutations in genes playing a crucial role in the control of ovulation rate. In the Noire du Velay (NV) sheep population, a recent genetic study has evidenced the segregation of such a mutation named FecL L . However, based on litter size (LS) records of FecL L non-carrier ewes, the segregation of a second prolificacy major mutation was suspected in this population. In order to identify this mutation, we have combined a case/control genome-wide association study with ovine 50k SNP chip genotyping, whole genome sequencing, and functional analyses. A new single nucleotide polymorphism (OARX:50977717T > A, NC_019484) located on the X chromosome upstream of the BMP15 gene was evidenced to be highly associated with the prolificacy variability (P = 1.93E-11). The variant allele was called FecX N and shown to segregate also in the Blanche du Massif Central (BMC) sheep population. In both NV and BMC, the FecX N allele frequency was estimated close to 0.10, and its effect on LS was estimated at +0.20 lamb per lambing at the heterozygous state. Homozygous FecX N carrier ewes were fertile with increased prolificacy in contrast to numerous mutations affecting BMP15. At the molecular level, FecX N was shown to decrease BMP15 promoter activity and supposed to impact BMP15 expression in the oocyte. This regulatory action was proposed as the causal mechanism for the FecX N mutation to control ovulation rate and prolificacy in sheep.
11For many decades, prolificacy has been selected in meat sheep breeds as a polygenic 12 trait but with limited genetic gain. However, the discovery of major genes affecting 13 prolificacy has changed the way of selection for some ovine breeds implementing 14 gene-assisted selection as in the French Lacaune and Grivette meat breeds, or in the 15 Spanish Rasa Aragonesa breed. Based on statistical analysis of litter size parameters 16 from 34 French meat sheep populations, we suspected the segregation of a mutation 17 in a major gene affecting prolificacy in the Noire du Velay and in the Mouton Vendéen 18 breeds exhibiting a very high variability of the litter size. After the genotyping of 19 mutations known to be present in French sheep breeds, we discovered the segregation 20 of the FecL L mutation at the B4GALNT2 locus and the FecX Gr mutation at the BMP15 21 locus in Noire du Velay and Mouton Vendéen, respectively. The frequency of ewes 22 carrying FecL L in the Noire du Velay population was estimated at 21.2% and the 23 Mouton Vendéen ewes carrying FecX Gr at 10.3%. The estimated mutated allele effect 24 of FecL L and FecX Gr on litter size at +0.4 and +0.3 lamb per lambing in Noire du Velay 25 and Mouton Vendéen, respectively. Due to the fairly high frequency and the rather 26 strong effect of the FecL L and FecX Gr prolific alleles, specific management programmes 27including genotyping should be implemented for a breeding objective of prolificacy 28 adapted to each of these breeds. 30In ovine breeds raised for meat purposes, numerical productivity represents an 32 important technical and economic lever. The objective is to reach an optimum for the 33 economic profitability of breeding. Improvement of this numerical productivity is 34 achieved by increasing the number of lambs born per ewe at each lambing, i.e. the 35 prolificacy, associated with the improvement of lamb viability as well as the maternal 36 quality. This leads to increased post-natal survival and growth rate of the lambs. For 37 decades, genetic selection efforts have been made particularly on improving prolificacy 38 of sheep breeds. However, prolificacy is a weakly heritable polygenic trait (h 2 = 0.05 -39 0.2) (see the review by Bradford (Bradford, 1985)), allowing limited genetic gain. 40 Nevertheless in some breeds, a very large effect on ovulation rate (OR) and litter size 41 (LS) due to single mutation in fecundity major genes (called Fec genes, reviewed in 42 (Fabre et al., 2006)) has been demonstrated. The first evidence of the segregation of 43 a prolificacy major gene was established in the early 1980's in Australian Booroola 44 Merino. This was implicated by the observation of a large variability of LS and OR in 45 this population and the presence of extremely prolific ewes in this low prolific breed 46 ( Piper and Bindon, 1982; Davis et al., 1982). The causal mutation named FecB B was 47 France particularly, two genetic programmes were implemented to discover and to 57 manage mutations with major effect in order to improve the pro...
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