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Cells can withstand hostile environmental conditions manifest as largemechanical forces such as pressure gradients and/or shear stresses bydynamically changing their shape. Such conditions are realized in the Schlemm'scanal of the eye where endothelial cells that cover the inner vessel wall aresubjected to the hydrodynamic pressure gradients exerted by the aqueous humoroutflow. These cells form fluid-filled dynamic outpouchings of their basalmembrane called \textit{giant vacuoles}. The inverse of giant vacuoles arereminiscent of cellular blebs, extracellular cytoplasmic protrusions triggeredby local temporary disruption of the contractile actomyosin cortex. Inverseblebbing has been first observed experimentally during sprouting angiogenesis,but its underlying physical mechanisms are poorly understood. Here, we identifygiant vacuole formation as inverse blebbing and formulate a biophysical modelof this process. Our model elucidates how cell membrane mechanical propertiesaffect the morphology and dynamics of giant vacuoles and predicts coarseningakin to Ostwald ripening between multiple invaginating vacuoles. Our resultsare in qualitative agreement with observations from the formation of giantvacuoles during perfusion experiments. Our model not only elucidates thebiophysical mechanisms driving inverse blebbing and giant vacuole dynamics, butalso identifies universal features of the cellular response to pressure loadsthat are relevant to many experimental contexts.

Model of inverse bleb growth explains giant vacuole dynamics during cell mechanoadaptation