ROS and calcium oscillations are required for polarized root hair growth

Root hairs are filamentous extensions from epidermis of plant roots with growth limited to the apical dome. Cell expansion undergoes tightly regulated processes, including the coordination between cell wall loosening and cell wall crosslinking, to form the final shape and size. Tip-focused gradients and oscillations of reactive oxygen species (ROS) together with calcium ions (Ca²⁺) as indispensable regulated mechanisms control rapid and polarized elongation of root hair cells. ROS homeostasis mediated by plasma membrane-localized NADPH oxidases, known as respiratory burst oxidase homologues (RBOHs), and class III cell wall peroxidases (PRXs), modulates cell wall properties during cell expansion. The expression levels of RBOHC, an NADPH oxidase that produces ROS, and class III PRXs are directly upregulated by ROOT HAIR DEFECTIVE SIX-LIKE 4 (RSL4), encoding a basic-helix-loop-helix (bHLH) transcription factor, to modulate root hair elongation. Cyclic nucleotide-gated channels (CNGCs), as central regulators of Ca²⁺ oscillations, also regulate root hair extension. Here, we review how the gradients and oscillations of Ca²⁺ and ROS interact to promote the expansion of root hair cells.