Department of Physics and Chemical Physics Program, University of Colorado, Boulder, CO 80309.
Chirality Research Center, Hiroshima University, Higashi Hiroshima, Hiroshima 739-8526, Japan.
Proc Natl Acad Sci U S A. 2022 Jun 14;119(24):e2200930119. doi: 10.1073/pnas.2200930119. Epub 2022 Jun 7.
Biological functionality is often enabled by a fascinating variety of physical phenomena that emerge from orientational order of building blocks, a defining property of nematic liquid crystals that is also pervasive in nature. Out-of-equilibrium, "living" analogs of these technological materials are found in biological embodiments ranging from myelin sheath of neurons to extracellular matrices of bacterial biofilms and cuticles of beetles. However, physical underpinnings behind manifestations of orientational order in biological systems often remain unexplored. For example, while nematiclike birefringent domains of biofilms are found in many bacterial systems, the physics behind their formation is rarely known. Here, using cellulose-synthesizing bacteria, we reveal how biological activity leads to orientational ordering in fluid and gel analogs of these soft matter systems, both in water and on solid agar, with a topological defect found between the domains. Furthermore, the nutrient feeding direction plays a role like that of rubbing of confining surfaces in conventional liquid crystals, turning polydomain organization within the biofilms into a birefringent monocrystal-like order of both the extracellular matrix and the rod-like bacteria within it. We probe evolution of scalar orientational order parameters of cellulose nanofibers and bacteria associated with fluid-gel and isotropic-nematic transformations, showing how highly ordered active nematic fluids and gels evolve with time during biological-activity-driven, disorder-order transformation. With fluid and soft-gel nematics observed in a certain range of biological activity, this mesophase-exhibiting system is dubbed "biotropic," analogously to thermotropic nematics that exhibit solely orientational order within a temperature range, promising technological and fundamental-science applications.
生物功能通常是通过构建块的取向有序所产生的各种迷人物理现象来实现的,这是向列液晶的一个定义性质,在自然界中也普遍存在。在非平衡状态下,这些技术材料的“活”类似物存在于从神经元髓鞘到细菌生物膜的细胞外基质和甲虫的角质层等生物实体中。然而,生物系统中取向有序表现的物理基础往往仍未得到探索。例如,虽然许多细菌系统中都存在类似向列的双折射生物膜域,但它们形成的物理原理却鲜为人知。在这里,我们使用合成纤维素的细菌,揭示了生物活性如何导致这些软物质系统的流体和凝胶类似物中的取向有序,无论是在水中还是在固体琼脂上,都存在畴之间的拓扑缺陷。此外,营养物质的供给方向起着类似于传统向列液晶中约束表面摩擦的作用,将生物膜内的多畴组织转变为细胞外基质和其中棒状细菌的双折射单晶体状有序。我们探测了与流体-凝胶和各向同性-向列转变相关的纤维素纳米纤维和细菌的标量取向序参数的演化,展示了高度有序的活性向列流体和凝胶如何随着时间的推移在生物活性驱动的无序-有序转变过程中演变。在一定范围的生物活性下观察到流体和软凝胶向列相,这个具有中间相的系统被称为“生物向列”,类似于仅在温度范围内表现出取向有序的热致向列,有望在技术和基础科学应用中得到应用。