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. 2021 Dec 3;12:761728. doi: 10.3389/fgene.2021.761728

TABLE 1.

An overview of the 18 examples of alleles with a heterozygote advantage and a homozygote disadvantage in livestock species.

Species (breed) Trait (References) Gene(s) involved Type of variant Inheritance Heterozygote advantage Homozygote disadvantage
Pig (Large White) Fetal lethality Derks et al. (2018) BBS9, BMPER Frameshift (deletion) Autosomal recessive Increased feed intake and growth Fetal death
Pig (Large White) Leg weakness Matika et al. (2019) MSTN Stop-gained Autosomal recessive Increase in muscle depth and decrease in fat depth Leg weakness syndrome
Pig (Finnish Yorkshire) Immotile short-tail sperm Sironen et al. (2012) SPEF2 Frameshift (insertion) Autosomal recessive High female litter size Male infertility
Pig (Pietrain, Landrace) Malignant hyperthermia Vogeli et al. (1994) RYR1 Missense Autosomal recessive High lean meat content Pale soft exudative meat
Cattle (Belgian Blue) Crooked tail Fasquelle et al. (2009) MRC2 Frameshift and missense Autosomal recessive Enhanced muscular development Crooked tail syndrome
Cattle (Nordic Red) Embryonic lethality Kadri et al. (2014) RNASEH2B Frameshift (deletion) Autosomal recessive High milk yield Embryonic lethal
Cattle (Belgian Blue and Shorthorn) Roan coat Seitz et al. (1999) KITLG Missense Autosomal Co-dominant Roan phenotype White heifer disease
Sheep Mastitis Rupp et al. (2015) SOCS2 Missense Autosomal recessive Higher body weight and milk production (also in homozygotes) Mastitis
Sheep Fecundity Hanrahan et al. (2004) BMP15, GDF9 Stop-gained and missense BMP15: X-linked GDF9: Autosomal recessive Increase ovulation rate and litter size Female infertility
Sheep (Soay) Polledness Wiedemar and Drögemüller (2015) RXFP2 Frameshift (insertion) Autosomal recessive Higher overall fitness HoP/HoP males: lower reproductive success
Ho+/Ho+ males: lower survival
Sheep Chondrodysplasia Beever et al. (2006) FGFR3 Missense Autosomal recessive Larger animals Spider lamb syndrome
Sheep Callipygous phenotype Kim et al. (2004) DLK1 - PEG11 Intergenic Polar overdominance Muscular hypertrophy -
Chicken (Wyandotte) Rose comb Imsland et al. (2012) MNR2, CCDC108 Inversion Autosomal dominant Rose comb Male infertility
Chicken Creeper Jin et al. (2016) IHH Deletion autosomal dominant semi-lethal Short legs Lethal before hatch
Horse (Appaloosa, Knabstrupper) Leopard complex spotting Bellone et al. (2010) TRPM1 Frameshift (insertion) Autosomal incompletely dominant Leopard complex spotting Congenital night blindness
Horse (American Paint) Frame pattern Ayala-Valdovinos et al (2016) EDNRB Missense Autosomal dominant Frame overo Lethal white foal syndrome
Rabbit Dwarfism Carneiro et al. (2017) HMGA2 Deletion Autosomal recessive Dwarfism Lethal
Rabbit English Spotting Fontanesi et al. (2014) KIT Regulatory Autosomal recessive English spotting Dilated (“mega”) cecum and ascending colon

The table also includes one example of polar-overdominance, of which the phenotype depends on the parental origin of the variant.