An ancient alkalinization factor informs Arabidopsis root development

The power of hydrogen (pH) regulates virtually all cellular activities. In both plants and animals, cell-to-cell variations in pH correlate with key developmental transitions^1–5, yet the underlying regulators and associated functions remain elusive. Here, we report that members of the REMORIN (REM) protein family function as inhibitors of the H⁺-ATPases thereby promoting extracellular pH (pHe) alkalinization. This, in turn, regulates various cell surface processes, including steroid hormone signaling, and coordinates developmental transitions in the Arabidopsis thaliana root. Inhibition of H⁺-ATPases by REMs represents an evolutionary innovation that predates the origin of the root system itself. This study thus uncovers an ancient alkalinization mechanism co-opted by the root developmental program and infers that pHe patterning may have shaped morphogenesis evolution.