Possible transient liquid crystal phase during the laying out of connective tissues:: α-chitin and collagen as models

Morphogenesis of extracellular matrices can be considered from different perspectives. One is that of ontogenesis, i.e., an organism's development, which is mostly concerned with the spatiotemporal regulation of genes, cell differentiation and migration. Complementary to this purely biological point of view, a physico-chemical approach can help in understanding complex mechanisms by highlighting specific events that do not require direct cellular control. Because of a structural similarity between some biological systems and liquid crystals, it was supposed that similar mechanisms could be involved. In this respect, it is important to determine the intrinsic self-assembly properties driving the ordering of biological macromolecules. Here we review in vitro studies of the condensed state of major biological macromolecules from extracellular matrices and related theories describing a mesophase transition in suspensions of rodlike particles. Dilute suspensions of collagen or chitin are isotropic, i.e., the macromolecules can take on any orientation in the fluid. Beyond a critical concentration, an ordered nematic phase appears with a higher volume fraction. The two-phase coexistence can be seen between crossed polarizers since the nematic phase is strongly birefringent and appears bright, whereas the isotropic phase remains dark. A widespread property of these structural macromolecular scaffolds is their chirality. Although the origin of chirality in colloidal suspensions is still a subject of debate, the helical nature of the cholesteric phase can be quantified. Small angle x-ray scattering performed on shear-aligned samples can help demonstrate the cholesteric nature of the anisotropic phase, inferred from optical observations. Liquid-like positional local order is revealed by the presence of broad interference peaks at low angle. The azimuthal profiles of these patterns are fitted to determine the value of the nematic order parameter at the transition. A few physico-chemistry experiments can assess the nature of the transition, and in turn, applying theoretical models can prove useful in predicting and controlling the structure of assemblies of biological macromolecules.