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Free, publicly-accessible full text available April 11, 2026
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Abstract The cytoskeleton is an active composite of filamentous proteins that dictates diverse mechanical properties and processes in eukaryotic cells by generating forces and autonomously restructuring itself. Enzymatic motors that act on the comprising filaments play key roles in this activity, driving spatiotemporally heterogeneous mechanical responses that are critical to cellular multifunctionality, but also render mechanical characterization challenging. Here, we couple optical tweezers microrheology and fluorescence microscopy with simulations and mathematical modeling to robustly characterize the mechanics of active composites of actin filaments and microtubules restructured by kinesin motors. It is discovered that composites exhibit a rich ensemble of force response behaviors–elastic, yielding, and stiffening–with their propensity and properties tuned by motor concentration and strain rate. Moreover, intermediate kinesin concentrations elicit emergent mechanical stiffness and resistance while higher and lower concentrations exhibit softer, more viscous dissipation. It is further shown that composites transition from well‐mixed interpenetrating double‐networks of actin and microtubules to de‐mixed states of microtubule‐rich aggregates surrounded by relatively undisturbed actin phases. It is this de‐mixing that leads to the emergent mechanical response, offering an alternate route that composites can leverage to achieve enhanced stiffness through coupling of structure and mechanics.more » « lessFree, publicly-accessible full text available April 10, 2026
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Free, publicly-accessible full text available January 13, 2026
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Abstract Polyacrylamide hydrogels are widely used in biomedical applications due to their tunable mechanical properties and charge neutrality. Our recent tribological investigations of polyacrylamide gels have revealed tunable and pH-dependent friction behavior. To determine the origins of this pH-responsiveness, we prepared polyacrylamide hydrogels with two different initiating chemistries: a reduction–oxidation (redox)-initiated system using ammonium persulfate (APS) andN,N,N′N′-tetramethylethylenediamine (TEMED) and a UV-initiated system with 2-hydroxy-4′-(2-hydroxyethoxy)-2-methylpropiophenone (Irgacure 2959). Hydrogel swelling, mechanical properties, and tribological behavior were investigated in response to solution pH (ranging from ≈ 0.34 to 13.5). For polyacrylamide hydrogels in sliding contact with glass hemispherical probes, friction coefficients decreased fromµ = 0.07 ± 0.02 toµ = 0.002 ± 0.002 (redox-initiated) and fromµ = 0.05 ± 0.03 toµ = 0.003 ± 0.003 (UV-initiated) with increasing solution pH. With hemispherical polytetrafluoroethylene (PTFE) probes, friction coefficients of redox-initiated hydrogels similarly decreased fromµ = 0.06 ± 0.01 toµ = 0.002 ± 0.001 with increasing pH. Raman spectroscopy measurements demonstrated hydrolysis and the conversion of amide groups to carboxylic acid in basic conditions. We therefore propose that the mechanism for pH-responsive friction in polyacrylamide hydrogels may be credited to hydrolysis-driven swelling through the conversion of side chain amide groups into carboxylic groups and/or crosslinker degradation. Our results could assist in the rational design of hydrogel-based tribological pairs for biomedical applications from acidic to alkaline conditions. Graphical abstractmore » « less
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Here we present a new, compact magnetic tweezers design that enables precise application of a wide range of dynamic forces to soft materials without the need to raise or lower the magnet height above the sample. This is achieved through the controlled rotation of the permanent magnet array with respect to the fixed symmetry axis defined by a custom-built iron yoke. These design improvements increase the portability of the device and can be implemented within existing microscope setups without the need for extensive modification of the sample holders or light path. This device is particularly well-suited to active microrheology measurements using either creep analysis, in which a step force is applied to a micron-sized magnetic particle that is embedded in a complex fluid, or oscillatory microrheology, in which the particle is driven with a periodic waveform of controlled amplitude and frequency. In both cases, the motions of the particle are measured and analyzed to determine the local dynamic mechanical properties of the material.more » « less
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The ability of cells to reorganize in response to external stimuli is important in areas ranging from morphogenesis to tissue engineering. While nematic order is common in biological tissues, it typically only extends to small regions of cells interacting via steric repulsion. On isotropic substrates, elongated cells can co-align due to steric effects, forming ordered but randomly oriented finite-size domains. However, we have discovered that flat substrates with nematic order can induce global nematic alignment of dense, spindle-like cells, thereby influencing cell organization and collective motion and driving alignment on the scale of the entire tissue. Remarkably, single cells are not sensitive to the substrate’s anisotropy. Rather, the emergence of global nematic order is a collective phenomenon that requires both steric effects and molecular-scale anisotropy of the substrate. To quantify the rich set of behaviours afforded by this system, we analyse velocity, positional and orientational correlations for several thousand cells over days. The establishment of global order is facilitated by enhanced cell division along the substrate’s nematic axis, and associated extensile stresses that restructure the cells’ actomyosin networks. Our work provides a new understanding of the dynamics of cellular remodelling and organization among weakly interacting cells.more » « less
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