Ca(2+)-regulated actin-binding protein. Involved in actin filaments bundling. Caps the barbed end of actin filaments and is able to sever them in a calcium-dependent manner. Required for the construction of actin collars in pollen tubes. Acts redundantly with VLN5 (AC Q9LVC6) to generate thick actin filament bundles and to regulate polarized pollen tube growth (PubMed:23715472). Acts redundantly with VLN3 (AC O81645) to regulate directional organ growth and in sclerenchyma development (PubMed:22209875, PubMed:22563899, respectively). Present in the apical and subapical regions of pollen tubes. Expressed in all tissues examined. Mainly detected in the root epidermis and vasculature. Expressed in the root cap. No visible phenotype and no visible effect on pollen tube growth. Decreased severing frequency of actin filaments. Vln2 and vln5 double mutants have pollen tubes curled and wider at some regions along the tube. They accumulate actin filaments at the tips of pollen tubes (PubMed:23715472). Vln2 and vln3 double mutants show anomaly in the growth direction of organs (PubMed:22209875) and defects in sclerenchyma development, but no alterations in the secondary cell-wall machinery (PubMed:22563899). Belongs to the villin/gelsolin family.