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Nucleocytoplasmic transport is essential for cellular function, presenting a canonical example of rapid molecular sorting inside cells. It consists of a coordinated interplay between import/export of molecules in/out the cell nucleus. Here, we investigate the role of spatiotemporal dynamics of the nucleocytoplasmic transport and its regulation. We develop a biophysical model that captures the main features of the nucleocytoplasmic transport, in particular, its regulation through the Ran cycle. Our model yields steady-state profiles for the molecular components of the Ran cycle, their relaxation times, as well as the nuclear-to-cytoplasmic molecule ratio. We show that these quantities are affected by their spatial dynamics and heterogeneity within the nucleus. Specifically, we find that the spatial nonuniformity of Ran guanine exchange factor (RanGEF)—particularly its proximity to the nuclear envelope—increases the Ran content in the nucleus. We further show that RanGEF's accumulation near the nuclear envelope results from its intrinsic dynamics as a nuclear cargo, transported by the Ran cycle itself. Overall, our work highlights the critical role of molecular spatial dynamics in cellular processes and proposes new avenues for theoretical and experimental inquiries into the nucleocytoplasmic transport. Published by the American Physical Society2024more » « lessFree, publicly-accessible full text available October 1, 2025
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Vorticity, a measure of the local rate of rotation of a fluid element, is the driver of incompressible flow. In viscous fluids, powering bulk flows requires the continuous injection of vorticity from boundaries to counteract the diffusive effects of viscosity. Here we power a flow from within by suspending approximately cylindrical particles and magnetically driving them to rotate at Reynolds numbers in the intermediate range. We find that a single particle generates a localized three-dimensional region of vorticity around it—which we call a vortlet—that drives a number of remarkable behaviours. Slight asymmetries in the particle shape can deform the vortlet and cause the particle to self-propel. Interactions between vortlets are similarly rich, generating bound dynamical states. When a large number of vortlets interact, they spontaneously form collectively moving flocks. These flocks remain coherent while propelling, splitting and merging. If enough particles are added so as to saturate the flow chamber, a homogeneous three-dimensional active chiral fluid of vortlets is formed, which can be manipulated with gravity or flow chamber boundaries, leading to lively collective dynamics. Our findings demonstrate an inertial regime for synthetic active matter, provide a controlled physical system for the quantitative study of three-dimensional flocking in non-sentient systems and establish a platform for the study of three-dimensional active chiral fluids.more » « lessFree, publicly-accessible full text available October 8, 2025
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We re-examine the model proposed by Alexanderet al.(Phys. Fluids, vol. 18, 2006, 062103) for the closing of a circular hole in a molecularly thin incompressible Langmuir film situated on a Stokesian subfluid. For simplicity their model assumes that the surface phase is inviscid which leads to the result that the cavity area decreases at a constant rate determined by the ratio of edge tension to subfluid viscosity. We reformulate the problem, allowing for a regularising monolayer viscosity. The viscosity-dependent corrections to the hole dynamics are analysed and found to be non-trivial, even when the monolayer viscosity is small; these corrections may explain the departure of experimental data from the theoretical prediction when the hole radius becomes comparable to the Saffman–Delbrück length. Through fitting, under these relaxed assumptions, we find the edge tension could be as much as six times larger ($$\sim$$4.0 pN) than reported previously.more » « less
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Inspired by the recent realization of a two-dimensional (2-D) chiral fluid as an active monolayer droplet moving atop a 3-D Stokesian fluid, we formulate mathematically its free-boundary dynamics. The surface droplet is described as a general 2-D linear, incompressible and isotropic fluid, having a viscous shear stress, an active chiral driving stress and a Hall stress allowed by the lack of time-reversal symmetry. The droplet interacts with itself through its driven internal mechanics and by driving flows in the underlying 3-D Stokes phase. We pose the dynamics as the solution to a singular integral–differential equation, over the droplet surface, using the mapping from surface stress to surface velocity for the 3-D Stokes equations. Specializing to the case of axisymmetric droplets, exact representations for the chiral surface flow are given in terms of solutions to a singular integral equation, solved using both analytical and numerical techniques. For a disc-shaped monolayer, we additionally employ a semi-analytical solution that hinges on an orthogonal basis of Bessel functions and allows for efficient computation of the monolayer velocity field, which ranges from a nearly solid-body rotation to a unidirectional edge current, depending on the subphase depth and the Saffman–Delbrück length. Except in the near-wall limit, these solutions have divergent surface shear stresses at droplet boundaries, a signature of systems with codimension-one domains embedded in a 3-D medium. We further investigate the effect of a Hall viscosity, which couples radial and transverse surface velocity components, on the dynamics of a closing cavity. Hall stresses are seen to drive inward radial motion, even in the absence of edge tension.more » « less