Abstract
Microtubules (MTs) are an important part of the cytoskeleton. In the living cell, MTs are non-equilibrium polymers with complex chemical and mechanical properties. During polymerization, tubulin dimers are believed to form an intermediate open sheet structure, which eventually closes into a tube. This implies flexibility in the lateral bonds between protofilaments. Previous research from Wang and Nogales (2005, Nature, 435:911) has shown a bistability in these bonds under non-physiological conditions, leading to the pairing of protofilaments with alternating binding angles. We have here used atomic force microscopy (AFM) to image the fine structure of taxol-stabilized MTs that were flattened inside-out against the substrate surface. We were able to reach single-monomer resolution, and observed a highly reproducible lateral zigzag pattern of protofilaments, demonstrating that a structural bistability also exists in regular microtubules. To estimate the angular orientation of protofilaments in the flattened MTs, we used the tubulin atomic structure to reconstruct the AFM images.
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