LORELEI-LIKE GPI-ANCHORED PROTEINS 2/3 Regulate Pollen Tube Growth as Chaperones and Coreceptors for ANXUR/BUPS Receptor Kinases in Arabidopsis.

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 alkalinization 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 which stimulate pollen tube growth and reduce pollen burst rate. levels are reduced in the pollen tubes of LLG2/3 lines, and application of exogenous H₂O₂ could partially rescue the defective pollen tube growth of LLG2/3 duanyu1615 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 activation of duanyu1670 production for promoting pollen tube growth.

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