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Unassisted ion transport through lipid membranes plays a crucial role in manycell functions without which life would not be possible, yet the precisemechanism behind the process remains unknown due to its molecular complexity.Here, we demonstrate a direct link between membrane potential fluctuations anddivalent ion transport. High-throughput wide-field second harmonic (SH)microscopy shows that membrane potential fluctuations are universally found inlipid bilayer systems. Molecular dynamics simulations reveal that suchvariations in membrane potential reduce the free energy cost of transient poreformation and increase the ion flux across an open pore. These transient porescan act as conduits for ion transport, which we SH image for a series ofdivalent cations (Cu$^{2+}$, Ca$^{2+}$, Ba$^{2+}$, Mg$^{2+}$) passing throughGUV membranes. Combining the experimental and computational results, we showthat permeation through pores formed via an ion-induced electrostatic field isa viable mechanism for unassisted ion transport.

Ion-induced transient potential fluctuations facilitate pore formation and cation transport through lipid membranes