Air spreading through wetted cellulose membranes: Implications for the safety function of hydraulic valves in plants

Plants transport water against the risk of cavitation inside xylem vessels, called ``embolism.'' As one of their hydraulic strategies, pit membranes composed of cellulose fibers have been known as safety valves that prevent the spreading of embolism towards adjacent xylem vessels. However, detailed observation of embolism spreading through a pit membrane is still lacking. Here, we hypothesized that the pit membranes normally remain to be wetted in xylem vessels and noticed in particular the hydraulic role of water film on air spreading that has been overlooked previously. For the hydrodynamic study of the embolism spreading through a wetted pit membrane, we investigated the penetration and spreading dynamics of air plugs through the wetted cellulose membrane in a channel flow. Air spreading exhibits two types of dynamics: continuous and discrete spreading. We elucidated the correlation of dynamic characteristics of air flow and pressure variations according to membrane thickness. Our study speculates that the thickness of pit membranes affects the behaviors of water film captured by cellulose fibers, and it is a crucial criterion for the reversible gating of further spreading of embolism throughout xylem networks.