RESEARCH ARTICLE o

Genotype imputation is a key step in the analysis of genome-wide association studies (GWAS) and genomic prediction [1]. Since obtaining whole genome sequences until recently was very costly, genomic studies have mostly relied on single nucleotide polymorphism (SNP) marker arrays, where only a small fraction of polymorphisms are preselected to be highly polymorphic and to cover the whole genome [2]. Furthermore, it has been suggested that many quantitative trait nucleotide (QTN) are rare variants [3]. This means that the analyses have typically relied on linkage disequilibrium (LD) between markers on the SNP panel and the causal polymorphism, since QTN that affect a certain trait are not expected to be among the com-monly available SNPs from marker panels. Genotype im-putation allows the analysis of variants not represented in * Correspondence