Lockhart with a twist: Modelling cellulose microfibril deposition and reorientation reveals twisting plant cell growth mechanisms.

Plant morphology emerges from cellular growth and structure. The turgor-driven diffuse growth of a cell can be highly anisotropic: significant longitudinally and negligible radially. Such anisotropy is ensured by cellulose microfibrils (CMF) reinforcing the cell wall in the hoop direction. To maintain the cell's integrity during growth, new wall material including CMF must be continually deposited. We develop a mathematical model representing the cell as a cylindrical pressure vessel and the cell wall as a fibre-reinforced viscous sheet, explicitly including the mechano-sensitive angle of CMF deposition. The model incorporates interactions between turgor, external forces, CMF reorientation during wall extension, and matrix stiffening. Using the model, we reinterpret some recent experimental findings, and reexamine the popular hypothesis of CMF/microtubule alignment. We explore how the handedness of twisting cell growth depends on external torque and intrinsic wall properties, and find that cells twist left-handedly 'by default' in some suitable sense. Overall, this study provides a unified mechanical framework for understanding left- and right-handed twist-growth as seen in many plants.