What are the true values of the bending modulus of simple lipid bilayers?

• Large uncertainties exist for values of the bending modulus of membranes.
• The effect of sugar is considered, including new X-ray data.
• Effect of tilt on data analysis is evaluated for different experimental methods.
• Estimates of values of the tilt modulus are discussed.

Values of the bending modulus KC are reviewed, and possible causes for the considerable differences are discussed. One possible cause is the use of glucose and sucrose in the classical micromechanical manipulation and shape analysis methods. New data, using the more recent low angle X-ray method, are presented that do not support an effect of glucose or sucrose on KC. Another possible cause is using an incomplete theory to interpret the data. Adding a tilt term to the theory clearly does not affect the value obtained from the shape analysis method. It is shown that a tilt term, using a value of the modulus Kθ indicated by simulations, theory, and estimated from order parameters obtained from NMR and from the wide angle X-ray method, should also not affect the value obtained using the micromechanical manipulation method, although it does require a small correction when determining the value of the area compressibility modulus KA. It is still being studied whether including a tilt term will significantly affect the values of KC obtained using low angle X-ray data. It remains unclear what causes the differences in the experimental values of KC for simple lipid bilayers.