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Fruit flavor is defined as the perception of the food by the olfactory and gustatory systems, and is one of the main determinants of fruit quality. Tomato flavor is largely determined by the balance of sugars, acids and volatile compounds. Several genes controlling the levels of these metabolites in tomato fruit have been cloned, including LIN5 , ALMT9 , AAT1 , CXE1 , and LoxC . The aim of this study was to identify any association of these genes with trait variation and to describe the genetic diversity at these loci in the red-fruited tomato clade comprised of the wild ancestor Solanum pimpinellifolium , the semi-domesticated species Solanum lycopersicum cerasiforme and early domesticated Solanum lycopersicum . High genetic diversity was observed at these five loci, including novel haplotypes that could be incorporated into breeding programs to improve fruit quality of modern tomatoes. Using newly available high-quality genome assemblies, we assayed each gene for potential functional causative polymorphisms and resolved a duplication at the LoxC locus found in several wild and semi-domesticated accessions which caused lower accumulation of lipid derived volatiles. In addition, we explored gene expression of the five genes in nine phylogenetically diverse tomato accessions. In general, the expression patterns of these genes increased during fruit ripening but diverged between accessions without clear relationship between expression and metabolite levels. 

Summary The mechanisms that regulate the vast diversity of plant organ shapes such as the fruit remain to be fully elucidated. TONNEAU1 Recruiting Motif proteins (TRMs) have been implicated in the control of organ shapes in a number of plant species, including tomato. However, the role of many of them is unknown. TRMs interact with Ovate Family Proteins (OFPs) via the M8 domain. However, thein plantafunction of the TRM‐OFP interaction in regulating shape is unknown.We used CRISPR/Cas9 to generate knockout mutants in TRM proteins from different subclades and in‐frame mutants within the M8 domain to investigate their roles in organ shape and interactions with OFPs.Our findings indicate that TRMs impact organ shape along both the mediolateral and proximo‐distal axes of growth. Mutations inSltrm3/4andSltrm5act additively to rescue the elongated fruit phenotype ofovate/Slofp20(o/s) to a round shape. Contrary, mutations inSltrm19andSltrm17/20aresult in fruit elongation and further enhance the obovoid phenotype in theo/smutant.This study supports a combinatorial role of the TRM‐OFP regulon where OFPs and TRMs expressed throughout development have both redundant and opposing roles in regulating organ shape.