AbstractDuring reproduction in flowering plants, pollen grains form a tube that grows in a polarized fashion through the female tissues to eventually fertilize the egg cell. These highly polarized pollen tubes have a rapid rate of growth that is supported by a tip-focused delivery of membrane and cell wall components. To gain a better understanding of how this growth is regulated, we investigated the function RABA4D, a member of the Arabidopsis thaliana RabA4 subfamily of Rab GTPase proteins. Here, we show that RABA4D was expressed in a pollen-specific manner and that enhanced yellow fluorescent protein (EYFP)-RabA4d-labeled membrane compartments localized to the tips of growing pollen tubes. Mutant pollen in which the RABA4D gene was disrupted displayed bulged pollen tubes with a reduced rate of growth in vitro and displayed altered deposition of some cell wall components. Expression of EYFP-RabA4d restored wild-type phenotypes to the raba4d mutant pollen tubes, while expression of EYFP-RabA4b did not rescue the raba4d phenotype. In vivo, disruption of RABA4D resulted in a male-specific transmission defect with mutant raba4d pollen tubes displaying aberrant growth in the ovary and reduced guidance at the micropyle. We propose that RabA4d plays an important role in the regulation of pollen tube tip growth.