Bacterial cells possess multiple cytoskeletal proteins involved in a wide range of cellular processes. These cytoskeletal proteins are dynamic, but the driving forces and cellular functions of these dynamics remain poorly understood. Eukaryotic cytoskeletal dynamics are often driven by motor proteins, but in bacteria no motors that drive cytoskeletal motion have been identified to date. Here, we quantitatively study the dynamics of the Escherichia coli actin homolog MreB, which is essential for the maintenance of rod-like cell shape in bacteria. We find that MreB rotates around the long axis of the cell in a persistent manner. Whereas previous studies have suggested that MreB dynamics are driven by its own polymerization, we show that MreB rotation does not depend on its own polymerization but rather requires the assembly of the peptidoglycan cell wall. The cell-wall synthesis machinery thus either constitutes a novel type of extracellular motor that exerts force on cytoplasmic MreB, or is indirectly required for an as-yetunidentified motor. Biophysical simulations suggest that one function of MreB rotation is to ensure a uniform distribution of new peptidoglycan insertion sites, a necessary condition to maintain rod shape during growth. These findings both broaden the view of cytoskeletal motors and deepen our understanding of the physical basis of bacterial morphogenesis.bacterial cytoskeletal dynamics | cell-wall organization | peptidoglycan synthesis | cell growth C ytoskeletal proteins play an important role in bacterial morphogenesis (1). Of the bacterial cytoskeletal proteins, the widely conserved actin homolog MreB is particularly important for bacterial cells to elongate and maintain a rod-like shape. MreB forms polymers that are associated with the cell membrane and distributed along the length of the cell in many rod-shaped bacteria (2). These polymeric MreB structures are essential for the maintenance of rod-like cell shape, as their disruption leads to cell rounding. Although MreB is essential for proper morphogenesis, bacterial cell shape is ultimately determined by the shape of the peptidoglycan cell-wall sacculus, which in turn is controlled by the cell-wall synthesis machinery. The cell wall, which is composed of stiff glycan strands cross-linked by flexible peptide linkers, forms a load-bearing structure that can counteract the intracellular turgor pressure. Cell-wall assembly requires peptidoglycan subunits to be synthesized, polymerized into glycan strands by transglycosylase enzymes, and cross-linked into the existing cell-wall network by transpeptidase enzymes. MreB directly or indirectly associates with a number of proteins that have been implicated in cell-wall assembly, such that MreB is believed to act upstream of the cell-wall assembly machinery to direct the synthesis enzymes to the sites of cell-wall insertion.Previous studies have demonstrated that MreB structures are dynamic in Bacillus subtilis and Caulobacter crescentus (3-7). To date, no motor proteins have been shown to either ...
Overdamped Brownian motion of a self-propelled particle is studied by solving the Langevin equation analytically. On top of translational and rotational diffusion, in the context of the presented model, the 'active' particle is driven along its internal orientation axis. We calculate the first four moments of the probability distribution function for displacements as a function of time for a spherical particle with isotropic translational diffusion, as well as for an anisotropic ellipsoidal particle. In both cases the translational and rotational motion is either unconfined or confined to one or two dimensions. A significant non-Gaussian behaviour at finite times t is signalled by a non-vanishing kurtosis γ(t). To delimit the super-diffusive regime, which occurs at intermediate times, two timescales are identified. For certain model situations a characteristic t(3) behaviour of the mean-square displacement is observed. Comparing the dynamics of real and artificial microswimmers, like bacteria or catalytically driven Janus particles, to our analytical expressions reveals whether their motion is Brownian or not.
Self-propelled particles move along circles rather than along a straight line when their driving force does not coincide with their propagation direction. Examples include confined bacteria and spermatozoa, catalytically driven nanorods, active, anisotropic colloidal particles and vibrated granulates. Using a non-Hamiltonian rate theory and computer simulations, we study the motion of a Brownian "circle swimmer" in a confining channel. A sliding mode close to the wall leads to a huge acceleration as compared to the bulk motion, which can further be enhanced by an optimal effective torque-to-force ratio.
The phase-field-crystal model is by now widely used in order to predict crystal nucleation and growth. For colloidal solidification with completely overdamped individual particle motion, we show that the phase-field-crystal dynamics can be derived from the microscopic Smoluchowski equation via dynamical density-functional theory. The different underlying approximations are discussed. In particular, a variant of the phase-field-crystal model is proposed which involves less approximations than the standard phase-field-crystal model. We finally test the validity of these phase-field-crystal models against dynamical density-functional theory. In particular, the velocities of a linear crystal front from the undercooled melt are compared as a function of the undercooling for a two-dimensional colloidal suspension of parallel dipoles. Good agreement is only obtained by a drastic scaling of the free energies in the phase-field-crystal model in order to match the bulk freezing transition point.
Many bacterial species colonize surfaces and form dense 3D structures, known as biofilms, which are highly tolerant to antibiotics and constitute one of the major forms of bacterial biomass on Earth. Bacterial biofilms display remarkable changes during their development from initial attachment to maturity, yet the cellular architecture that gives rise to collective biofilm morphology during growth is largely unknown. Here, we use high-resolution optical microscopy to image all individual cells in Vibrio cholerae biofilms at different stages of development, including colonies that range in size from 2 to 4,500 cells. From these data, we extracted the precise 3D cellular arrangements, cell shapes, sizes, and global morphological features during biofilm growth on submerged glass substrates under flow. We discovered several critical transitions of the internal and external biofilm architectures that separate the major phases of V. cholerae biofilm growth. Optical imaging of biofilms with single-cell resolution provides a new window into biofilm formation that will prove invaluable to understanding the mechanics underlying biofilm development.biofilm | community | self-organization | emergent order | nematic order
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