SCFKMD controls cytokinin signaling by regulating the degradation of type-B response regulators

Cytokinins are plant hormones that play critical roles in growth and development. In Arabidopsis, the transcriptional response to cytokinin is regulated by action of type-B Arabidopsis response regulators (ARRs). Although central elements in the cytokinin signal transduction pathway have been identified, mechanisms controlling output remain to be elucidated. Here we demonstrate that a family of F-box proteins, called the KISS ME DEADLY (KMD) family, targets type-B ARR proteins for degradation. KMD proteins form an S-PHASE KINASE-ASSOCIATED PROTEIN1 (SKP1)/Cullin/F-box protein (SCF) E3 ubiquitin ligase complex and directly interact with type-B ARR proteins. Loss-of-function KMD mutants stabilize type-B ARRs and exhibit an enhanced cytokinin response. In contrast, plants with elevated KMD expression destabilize type-B ARR proteins leading to cytokinin insensitivity. Our results support a model in which an SCFKMD complex negatively regulates cytokinin responses by controlling levels of a key family of transcription factors.