Buckling-induced zebra stripe patterns in nematic F-actin

Rather than forming a simple and uniform nematic liquid crystal, concentrated solutions of semiflexible polymers, such as F-actin, have been observed to display a spatially periodic switching of the nematic director. When observed with polarization microscopy, these patterns appear as alternating light and dark bands, often referred to as zebra stripe patterns. Zebra stripe patterns, although not fully characterized, are due to periodic orientation distortions in the nematic order. We characterize such patterns by using a combination of two techniques. Using polarization microscopy, we quantify the periodic orientation distortions and show that the magnitude of the order parameter also varies periodically in the striped domains. When using fluorescently labeled filaments as markers, filaments spanning the striped domains are seen to undergo large angle bends. With fluorescence, clear density differences between adjacent stripes are also observed with domains of lesser density corresponding to strongly bent filaments. By directly comparing patterned areas with both polarization and fluorescence techniques, we show that periodic variation in the orientation, order parameter, filament bending, and density are correlated. We propose that these effects originate from the coupling of orientation and density that occurs for highly concentrated solutions of long semiflexible polymers subject to shear flows, as previously proposed [P. de Gennes, Mol. Cryst. Liq. Cryst. (Phila. Pa.) 34, 177 (1977)]. After cessation of shearing, strong interfilament interactions and high compressibility can lead to periodic buckling from the relaxation of filaments stretched during flows. The characterization of zebra stripe patterns presented here provides evidence that buckling in confined F-actin nematics produces strong periodic bending that is responsible for the observed features.