Two-Dimensional Brain Microtubule Structures Behave as Memristive Devices

Microtubules (MTs) are cytoskeletal structures that play a central role in a variety of cell functions including cell division and cargo transfer. MTs are also nonlinear electrical transmission lines that produce and conduct electrical oscillations elicited by changes in either electric field and/or ionic gradients. The oscillatory behavior of MTs requires a voltage-sensitive gating mechanism to enable the electrodiffusional ionic movement through the MT wall. Here we explored the electrical response of non-oscillating rat brain MT sheets to square voltage steps. To ascertain the nature of the possible gating mechanism, the electrical response of non-oscillating rat brain MT sheets (2D arrays of MTs) to square pulses was analyzed under voltage-clamping conditions. A complex voltage-dependent nonlinear charge movement was observed, which represented the summation of two events. The first contribution was a small, saturating, voltage-dependent capacitance with a maximum charge displacement in the range of 4 fC/mu m(2). A second, major contribution was a non-saturating voltage-dependent charge transfer, consistent with the properties of a multistep memristive device. The memristive capabilities of MTs could drive oscillatory behavior, and enable voltage-driven neuromorphic circuits and architectures within neurons.