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Tomato ( Solanum lycopersicum ) is a model for climacteric fleshy fruit ripening studies. Tomato ripening is regulated by multiple transcription factors together with the plant hormone ethylene and their downstream effector genes. Transcription Factors APETALA2a (AP2a), NON-RIPENING (NOR) and FRUITFULL (FUL1/TDR4 and FUL2/MBP7) were reported as master regulators controlling tomato fruit ripening. Their proposed functions were derived from studies of the phenotype of spontaneous mutants or RNAi knock-down lines rather than, as it appears now, actual  null  mutants. To study TF function in tomato fruit ripening in more detail, we used CRISPR/Cas9-mediated mutagenesis to knock out the encoding genes, and phenotypes of these mutants are reported for the first time. While the earlier ripening, orange-ripe phenotype of  ap2a  mutants was confirmed, the  nor null  mutant exhibited a much milder phenotype than the spontaneous  nor  mutant. Additional analyses revealed that the severe phenotype in the spontaneous mutant is caused by a dominant-negative allele. Our approach also provides new insight into the independent and overlapping functions of  FUL1  and  FUL2 . Single and combined  null  alleles of  FUL1  and  FUL2  illustrate that these two genes have partially redundant functions in fruit ripening, but also unveil an additional role for  FUL2  in early fruit development.

Re-evaluation of transcription factor function in tomato fruit development and ripening with CRISPR/Cas9-mutagenesis