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Physics of Microswimmers - Single Particle Motion and Collective Behavior
Locomotion and transport of microorganisms in fluids is an essential aspect
of life. Search for food, orientation toward light, spreading of off-spring,
and the formation of colonies are only possible due to locomotion. Swimming at
the microscale occurs at low Reynolds numbers, where fluid friction and
viscosity dominates over inertia. Here, evolution achieved propulsion
mechanisms, which overcome and even exploit drag. Prominent propulsion
mechanisms are rotating helical flagella, exploited by many bacteria, and
snake-like or whip-like motion of eukaryotic flagella, utilized by sperm and
algae. For artificial microswimmers, alternative concepts to convert chemical
energy or heat into directed motion can be employed, which are potentially more
efficient. The dynamics of microswimmers comprises many facets, which are all
required to achieve locomotion. In this article, we review the physics of
locomotion of biological and synthetic microswimmers, and the collective
behavior of their assemblies. Starting from individual microswimmers, we
describe the various propulsion mechanism of biological and synthetic systems
and address the hydrodynamic aspects of swimming. This comprises
synchronization and the concerted beating of flagella and cilia. In addition,
the swimming behavior next to surfaces is examined. Finally, collective and
cooperate phenomena of various types of isotropic and anisotropic swimmers with
and without hydrodynamic interactions are discussed.Comment: 54 pages, 59 figures, review article, Reports of Progress in Physics
(to appear
A practical review on the measurement tools for cellular adhesion force
Cell cell and cell matrix adhesions are fundamental in all multicellular
organisms. They play a key role in cellular growth, differentiation, pattern
formation and migration. Cell-cell adhesion is substantial in the immune
response, pathogen host interactions, and tumor development. The success of
tissue engineering and stem cell implantations strongly depends on the fine
control of live cell adhesion on the surface of natural or biomimetic
scaffolds. Therefore, the quantitative and precise measurement of the adhesion
strength of living cells is critical, not only in basic research but in modern
technologies, too. Several techniques have been developed or are under
development to quantify cell adhesion. All of them have their pros and cons,
which has to be carefully considered before the experiments and interpretation
of the recorded data. Current review provides a guide to choose the appropriate
technique to answer a specific biological question or to complete a biomedical
test by measuring cell adhesion
Possible origins of macroscopic left-right asymmetry in organisms
I consider the microscopic mechanisms by which a particular left-right (L/R)
asymmetry is generated at the organism level from the microscopic handedness of
cytoskeletal molecules. In light of a fundamental symmetry principle, the
typical pattern-formation mechanisms of diffusion plus regulation cannot
implement the "right-hand rule"; at the microscopic level, the cell's
cytoskeleton of chiral filaments seems always to be involved, usually in
collective states driven by polymerization forces or molecular motors. It seems
particularly easy for handedness to emerge in a shear or rotation in the
background of an effectively two-dimensional system, such as the cell membrane
or a layer of cells, as this requires no pre-existing axis apart from the layer
normal. I detail a scenario involving actin/myosin layers in snails and in C.
elegans, and also one about the microtubule layer in plant cells. I also survey
the other examples that I am aware of, such as the emergence of handedness such
as the emergence of handedness in neurons, in eukaryote cell motility, and in
non-flagellated bacteria.Comment: 42 pages, 6 figures, resubmitted to J. Stat. Phys. special issue.
Major rewrite, rearranged sections/subsections, new Fig 3 + 6, new physics in
Sec 2.4 and 3.4.1, added Sec 5 and subsections of Sec
Phase separation and rotor self-assembly in active particle suspensions
Adding a non-adsorbing polymer to passive colloids induces an attraction
between the particles via the `depletion' mechanism. High enough polymer
concentrations lead to phase separation. We combine experiments, theory and
simulations to demonstrate that using active colloids (such as motile bacteria)
dramatically changes the physics of such mixtures. First, significantly
stronger inter-particle attraction is needed to cause phase separation.
Secondly, the finite size aggregates formed at lower inter-particle attraction
show unidirectional rotation. These micro-rotors demonstrate the self assembly
of functional structures using active particles. The angular speed of the
rotating clusters scales approximately as the inverse of their size, which may
be understood theoretically by assuming that the torques exerted by the
outermost bacteria in a cluster add up randomly. Our simulations suggest that
both the suppression of phase separation and the self assembly of rotors are
generic features of aggregating swimmers, and should therefore occur in a
variety of biological and synthetic active particle systems.Comment: Main text: 6 pages, 5 figures. Supplementary information: 5 pages, 4
figures. Supplementary movies available from
httP://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1116334109/-/DCSupplementa
Universal entrainment mechanism governs contact times with motile cells
Contact between particles and motile cells underpins a wide variety of
biological processes, from nutrient capture and ligand binding, to grazing,
viral infection and cell-cell communication. The window of opportunity for
these interactions is ultimately determined by the physical mechanism that
enables proximity and governs the contact time. Jeanneret et al. (Nat. Comm. 7:
12518, 2016) reported recently that for the biflagellate microalga
Chlamydomonas reinhardtii contact with microparticles is controlled by events
in which the object is entrained by the swimmer over large distances. However,
neither the universality of this interaction mechanism nor its physical origins
are currently understood. Here we show that particle entrainment is indeed a
generic feature for microorganisms either pushed or pulled by flagella. By
combining experiments, simulations and analytical modelling we reveal that
entrainment length, and therefore contact time, can be understood within the
framework of Taylor dispersion as a competition between advection by the no
slip surface of the cell body and microparticle diffusion. The existence of an
optimal tracer size is predicted theoretically, and observed experimentally for
C. reinhardtii. Spatial organisation of flagella, swimming speed, swimmer and
tracer size influence entrainment features and provide different trade-offs
that may be tuned to optimise microbial interactions like predation and
infection.Comment: New analytical entrainment theory; includes Supplementary
informations as Appendix; Supplementary movies available upon reques
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