The BEACH Domain-Containing Protein SPIRRIG Modulates Actin-Dependent Root Hair Development in Coordination with the WAVE/SCAR and ARP2/3 Complexes
AbstractRoot hairs are single cell protrusions that enable roots to optimize nutrient and water acquisition. They attain their tubular shapes by confining growth to the cell apex, a process called tip growth. The actin cytoskeleton and endomembrane system are essential for tip growth; however, little is known about how these cellular components coordinate their activities during this process. Here, we show that SPIRRIG (SPI), a BEACH domain-containing protein involved in membrane trafficking, and BRK1 and SCAR2, subunits of the WAVE/SCAR (W/SCR) and actin related protein (ARP)2/3 activation complexes, display polarized localizations to root hairs at distinct developmental stages. SPI accumulates at the root hair apex via post-Golgi vesicles and positively regulates tip growth by maintaining tip-focused vesicle secretion and filamentous-actin integrity. BRK1 and SCAR2 on the other hand, mark the root hair initiation domain to specify the position of root hair emergence. Live cell microscopy revealed that BRK1 depletion coincided with SPI accumulation as root hairs transitioned from initiation to tip growth. Furthermore, double mutant studies showed that SPI genetically interacts with BRK1 and ARP2/3. Taken together, our work uncovers a role for SPI in facilitating actin-dependent root hair development through pathways that intersect with the W/SCR and ARP2/3 complexes.