From cytoskeletal assemblies to living materials

Many subcellular structures contain large numbers of cytoskeletal filaments. Such assemblies underlie much of cell division, motility, signaling, metabolism, and growth. Thus, understanding cell biology requires understanding the properties of networks of cytoskeletal filaments. While there are well established disciplines in biology dedicated to studying isolated proteins - their structure (Structural Biology) and behaviors (Biochemistry) - it is much less clear how to investigate, or even just describe, the structure and behaviors of collections of cytoskeletal filaments. One approach is to use methodologies from Mechanics and Soft Condensed Matter Physics, which have been phenomenally successful in the domains where they have been traditionally applied. From this perspective, collections of cytoskeletal filaments are viewed as materials, albeit very complex, 'active' materials, composed of molecules which use chemical energy to perform mechanical work. A major challenge is to relate these material level properties to the behaviors of the molecular constituents. Here we discuss this materials perspective and review recent work bridging molecular and network scale properties of the cytoskeleton, focusing on the organization of microtubules by dynein as an illustrative example.