Cellulose Synthase Complex and Remorin Nanodomains Mediate Stress Resilience Through Cell Wall-Plasma Membrane Attachments
HighlightsO_LIThe plasma membrane forms attachments to the plant cell wall that are revealed by hyperosmotic shock and correlate with tolerance to stress. C_LIO_LICellulose Synthase Complex (CSC) clusters and REMORIN (REM) nanodomains localize to cell wall-plasma membrane attachment sites. C_LIO_LICSC density at the plasma membrane determines the extent of cell wall-plasma membrane attachment under hyperosmotic stress. C_LIO_LIREMs rapidly form nanodomains under hyperosmotic stress and are associated with SHOU4/4L, CSC exocytosis inhibitors that limit CSC density at the plasma membrane. C_LI The outer cell surface of an organism is the frontline for detecting and responding to environmental stimuli. In plants, this interface consists of the plasma membrane that lies beneath the cell wall and remains associated with it through attachment sites. These wall-membrane attachments become evident upon hyperosmotic shock, when severe water loss causes the membrane to retract from the wall. Despite their long-standing observation, the molecular identity and function of these attachments remain poorly understood. Here, we identified two mechanisms governing wall-membrane attachments: one dependent on the Cellulose Synthase Complex (CSC), whose density at the plasma membrane positively correlates with resistance to hyperosmotic stress, and the other on REMORIN (REM), which acts antagonistically to the CSC mechanism. Using proximity-labeling proteomics, we identified SHOU4/4L as REM-associated proteins that mediate this antagonism. Together, our findings reveal how wall-membrane attachments are patterned to mediate plant cell resilience under water stress.