Polarized tip growth is a fundamental cellular process in many eukaryotes.In this study,we examined the dynamic restructuring of the actin cytoskeleton and its relationship to vesicle transport during pollen tip growth in Arabidopsis.We found that actin filaments originating from the apical membrane form a specialized structure consisting of longitudinally aligned actin bundles at the cortex and inner cytoplasmic filaments with a distinct distribution.Using actin-based pharmacological treatments and genetic mutants in combination with FRAP (fluorescence recovery after photobleaching) technology to visualize the transport of vesicles within the growth domain of pollen tubes,we demonstrated that cortical actin filaments facilitate tip-ward vesicle transport.We also discovered that the inner apical actin filaments prevent backward movement of vesicles,thus ensuring that sufficient vesicles accumulate at the pollen tube tip to support the rapid growth of the pollen tube.The combinatorial effect of cortical and inteal apical actin filaments perfectly explains the generation of the inverted "V" cone-shaped vesicle distribution patte at the pollen tube tip.When pollen tubes tu,apical actin filaments at the facing side undergo depolymerization and repolymerization to reorient the apical actin structure toward the new growth direction.This actin restructuring precedes vesicle accumulation and changes in tube morphology.Thus,our study provides new insights into the functional relationship between actin dynamics and vesicle transport during rapid and directional pollen tube growth.