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Tension plays a vital role in pore formation in biomembranes, but the mechanism of pore formation remains unclear. We investigated the temperature dependence of the rate constant of constant tension (σ)–induced pore formation in giant unilamellar vesicles of lipid membranes using an experimental method we developed. By analyzing this result, we determined the activation energy ( ) of tension-induced pore formation as a function of tension. A constant ( ) that does not depend on tension was found to contribute significantly to . Analysis of the activation energy clearly indicated that the dependence of on in the classical theory is correct, but that the classical theory of pore formation is not entirely correct due to the presence of . We can reasonably consider that is a nucleation free energy to form a hydrophilic pre-pore from a hydrophobic pre-pore or a region with lower lateral lipid density. After obtaining , the evolution of a pre-pore follows a classical theory. Our data provide valuable information that help explain the mechanism of tension-induced pore formation in biomembranes and lipid membranes.


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