Control of plant germline proliferation by SCFFBL17 degradation of cell cycle inhibitors

Flowering plants possess a unique reproductive strategy, involving double fertilization by twin sperm cells(1). Unlike animal germ lines, the male germ cell lineage in plants only forms after meiosis and involves asymmetric division of haploid microspores, to produce a large, non- germline vegetative cell and a germ cell that undergoes one further division to produce the twin sperm cells(2). Although this switch in cell cycle control is critical for sperm cell production and delivery, the underlying molecular mechanisms are unknown. Here we identify a novel F- box protein of Arabidopsis thaliana, designated FBL17 ( F- box- like 17), that enables this switch by targeting the degradation of cyclin- dependent kinase A; 1 inhibitors specifically in male germ cells. We show that FBL17 is transiently expressed in the male germ line after asymmetric division and forms an SKP1 -Cullin1 - F- box protein ( SCF) E3 ubiquitin ligase complex (SCFFBL17) that targets the cyclin- dependent kinase inhibitors KRP6 and KRP7 for proteasome- dependent degradation. Accordingly, the loss of FBL17 function leads to the stabilization of KRP6 and inhibition of germ cell cycle progression. Our results identify SCFFBL17 as an essential male germ cell proliferation complex that promotes twin sperm cell production and double fertilization in flowering plants.