Viscoelastic mechanics of living cells

In cell mechanotransduction, cells respond to external forces or to perceive mechanical properties of their supporting substrates by remodeling themselves. This ability is endowed by modulating cells' viscoelastic properties, which dominates over various complex cellular processes. The viscoelasticity of living cells, a concept adapted from rheology, exhibits substantially spatial and temporal variability. This review aims not only to discuss the rheological properties of cells but also to clarify the complexity of cellular rheology, emphasizing its dependence on both the size scales and time scales of the measurements. Like typical viscoelastic materials, the storage and loss moduli of cells often exhibit robust power-law rheological characteristics with respect to loading frequency. This intrinsic feature is consistent across cell types and is attributed to internal structures, such as cytoskeleton, cortex, cytoplasm and nucleus, all of which contribute to the complexity of cellular rheology. Moreover, the rheological properties of cells are dynamic and play a crucial role in various cellular and tissue functions. In this review, we focus on elucidating time- and size-dependent aspects of cell rheology, the origins of intrinsic rheological properties and how these properties adapt to cellular functions, with the goal of interpretation of rheology into the language of cell biology.