Up-regulation of LsKN1 promotes cytokinin and suppresses gibberellin biosynthesis to generate wavy leaves in lettuce.

Lettuce (Lactuca sativa) is one of the most popular vegetables worldwide, and diverse leaf shapes, including wavy leaves, are important commercial traits. In this study, we examined the genetics of wavy leaves using an F2 segregating population, and identified a major QTL controlling wavy leaves. The candidate region contained LsKN1, which has previously been shown to be indispensable for leafy heads in lettuce. Complementation tests and knockout experiments verified the function of LsKN1 in producing wavy leaves. The LsKN1∇ allele, which has the insertion of a transposon and has previously been shown to control leafy heads, promoted wavy leaves in our population. Transposition of the CACTA transposon from LsKN1 compromised its function for wavy leaves. High expression of LsKN1 up-regulated several key genes associated with cytokinin (CK) to increase the content in the leaves, whereas it down-regulated the expression of genes in the gibberellin (GA) biosynthesis pathway to decrease the content. Application of CK to leaves enhanced the wavy phenotype, while application of GA dramatically flattened the leaves. We conclude that the changes in CK and GA contents that result from high expression of LsKN1 switch determinate cells to indeterminate, and consequently leads to the development of wavy leaves.