Secondary growth in the tomato stem is suppressed by the flowering regulators FUL2 and STM3
Herbaceous eudicot crops such as tomato accumulate substantial amounts of secondary xylem in their stems, particularly in the basal part. As plants progress from the vegetative phase to flowering and fruit development, secondary growth continues to provide mechanical support and facilitate transport. To uncover how secondary growth is regulated in the tomato stem, we used laser capture microdissection RNA-seq to establish tissue-specific transcriptomic profiles. This revealed that key regulators associated with wood formation in poplar are also acting in tomato. Furthermore, we found that the flowering regulators FUL2 and STM3 are substantially expressed in xylem tissue. Subsequent functional analysis identified FUL2 and STM3 as key repressors of secondary growth, with ful2 or stm3 knock-out mutants exhibiting strongly increased secondary growth in older tomato stems. Notably, a higher order mutant in which both ful2 and stm3 were knocked out displays a similar non-additive phenotype, indicating, together with yeast two-hybrid data showing physical interaction, that FUL2 and STM3 act together in a complex. The expression levels of both genes remain low until after the floral transition but strongly rise during subsequent stem development. Transcriptomic analyses in ful2, stm3, and wild-type stems at stages before and after detectable anatomical differences, showed that the FUL2-STM3 complex represses distinct sets of NAC and MYB candidate regulators associated with xylem development. Interestingly, FUL2 and STM3 control these regulators in a stage- and position-dependent way, with candidates showing upregulation only at later stages in mid-height stems being already upregulated at earlier stages in basal stem regions. Our findings provide a tissue-resolved transcriptomic resource for tomato stems and reveal a role for FUL2 and STM3 in limiting secondary xylem accumulation as stem development progresses.