Cytokinins are involved in regulation of tomato pericarp thickness and fruit size

Although cytokinins (CKs) regulate fruit development, no direct genetic evidence supports the role of endogenous CKs in pericarp growth or development or fruit size. Here, we report that the reduction in levels of endogenous active CKs via overexpression of the CK-inactivating enzyme gene AtCKX2 specifically in fruit tissues resulted in reduced pericarp thickness and smaller fruit size compared with wild-type control fruits. Pericarp thickness and single fruit weight in transgenic plants were significantly reduced. Analysis of paraffin sections showed that the reduced pericarp thickness was due largely to a decreased number of cells, and thus decreased cell division. Transcriptome profiling showed that the expression of cell division- and expansion-related genes was reduced in AtCKX2-overexpressing fruits. In addition, the expression of auxin-signaling and gibberellin-biosynthetic genes was repressed, whereas that of gibberellin-inactivating genes was enhanced, in AtCKX2-overexpressing fruits. These results demonstrate that endogenous CKs regulate pericarp cell division and subsequently fruit size. They also suggest that CKs interact with auxin and gibberellins in regulating tomato pericarp thickness and fruit size.