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Pollen tube growth is crucial for successful fertilization.In Arabidopsis thaliana,the ANXUR(ANX)/BUPS receptor kinase complex controls and maintains pollen tube growth in response to autocrine rapid alkalin-ization factor 4/19(RALF4/19)signaling; however,the molecular and cellular mechanisms underlying the ANX/BUPS-mediated regulation of pollen tube growth remain unclear.In this study,we found that pollen-specific LORELEI-like GPI-anchored proteins 2 and 3(LLG2/3)promote pollen tube growth in vitro and in vivo.LLG2/3 interacte with ANX/BUPS in a RALF4-concentration-dependent manner,suggesting that ANX/BUPS and LLG2/3 might form a receptor-coreceptor complex for perceiving RALF peptide signals.Disruption of the ANX/BUPS-LLG2/3 interaction led to the cytoplasmic retention of ANX1/2,in either llg2/3 knockdown mutants or in anx1/2 mutants lacking the J region,which mediates the ANX/BUPS-LLG2/3 interaction.Moreover,we found that RALF4 induced the production of reactive oxygen species(ROS),which stimulate pollen tube growth and reduce pollen burst rate.ROS levels are reduced in the pollen tubes of LLG2/3 RNAi lines,and application of exogenous H2O2 could partially rescue the defective pollen tube growth of LLG2/3 RNAi lines.Taken together,our study identifies LLG2/3 as novel regulatory components of pollen tube growth,and shows that they chaperone ANX/BUPS for secretion to the apical plasma membrane of pollen tube and act as coreceptors of ANX/BUPS in the acti-vation of ROS production for promoting pollen tube growth.