gene experimentally. Lastly, the summary-based TWAS cannot account for rare variants that are poorly captured by the LD reference panel, or optimally capture non-linear relationships between SNPs and expression. Additional sources of information could potentially be incorporated to improve the prediction, including significant trans-associations11,28; allele-specific expression47,48; splice-QTLs effecting individual exons10; haplotype effects; and SNP-specific functional priors20,49–51.
