How to Track Protists in Three Dimensions

We present an apparatus optimized for tracking swimming microorganisms in the size range 10-1000 microns, in three dimensions (3D), far from surfaces, and with negligible background convective fluid motion. CCD cameras attached to two long working distance microscopes synchronously image the sample from two perpendicular directions, with narrowband dark-field or bright-field illumination chosen to avoid triggering a phototactic response. The images from the two cameras can be combined to yield 3D tracks of the organism. Using additional, highly directional broad-spectrum illumination with millisecond timing control the phototactic trajectories in 3D of organisms ranging from Chlamydomonas to Volvox can be studied in detail. Surface-mediated hydrodynamic interactions can also be investigated without convective interference. Minimal modifications to the apparatus allow for studies of chemotaxis and other taxes.Comment: 8 pages, 7 figure

Fluid dynamics of bacterial turbulence

Self-sustained turbulent structures have been observed in a wide range of living fluids, yet no quantitative theory exists to explain their properties. We report experiments on active turbulence in highly concentrated 3D suspensions of Bacillus subtilis and compare them with a minimal fourth-order vector-field theory for incompressible bacterial dynamics. Velocimetry of bacteria and surrounding fluid, determined by imaging cells and tracking colloidal tracers, yields consistent results for velocity statistics and correlations over two orders of magnitude in kinetic energy, revealing a decrease of fluid memory with increasing swimming activity and linear scaling between energy and enstrophy. The best-fit model parameters allow for quantitative agreement with experimental data.Comment: 5 pages, 4 figure

Oscillatory Flows Induced by Microorganisms Swimming in Two-dimensions

We present the first time-resolved measurements of the oscillatory velocity field induced by swimming unicellular microorganisms. Confinement of the green alga C. reinhardtii in stabilized thin liquid films allows simultaneous tracking of cells and tracer particles. The measured velocity field reveals complex time-dependent flow structures, and scales inversely with distance. The instantaneous mechanical power generated by the cells is measured from the velocity fields and peaks at 15 fW. The dissipation per cycle is more than four times what steady swimming would require.Comment: 4 pages, 4 figure

Meso-scale turbulence in living fluids

Turbulence is ubiquitous, from oceanic currents to small-scale biological and quantum systems. Self-sustained turbulent motion in microbial suspensions presents an intriguing example of collective dynamical behavior amongst the simplest forms of life, and is important for fluid mixing and molecular transport on the microscale. The mathematical characterization of turbulence phenomena in active non-equilibrium fluids proves even more difficult than for conventional liquids or gases. It is not known which features of turbulent phases in living matter are universal or system-specific, or which generalizations of the Navier-Stokes equations are able to describe them adequately. Here, we combine experiments, particle simulations, and continuum theory to identify the statistical properties of self-sustained meso-scale turbulence in active systems. To study how dimensionality and boundary conditions affect collective bacterial dynamics, we measured energy spectra and structure functions in dense Bacillus subtilis suspensions in quasi-2D and 3D geometries. Our experimental results for the bacterial flow statistics agree well with predictions from a minimal model for self-propelled rods, suggesting that at high concentrations the collective motion of the bacteria is dominated by short-range interactions. To provide a basis for future theoretical studies, we propose a minimal continuum model for incompressible bacterial flow. A detailed numerical analysis of the 2D case shows that this theory can reproduce many of the experimentally observed features of self-sustained active turbulence.Comment: accepted PNAS version, 6 pages, click doi for Supplementary Informatio

Direct measurement of the flow field around swimming microorganisms

Swimming microorganisms create flows that influence their mutual interactions and modify the rheology of their suspensions. While extensively studied theoretically, these flows have not been measured in detail around any freely-swimming microorganism. We report such measurements for the microphytes Volvox carteri and Chlamydomonas reinhardtii. The minute ~0.3% density excess of V. carteri over water leads to a strongly dominant Stokeslet contribution, with the widely-assumed stresslet flow only a correction to the subleading source dipole term. This implies that suspensions of V. carteri have features similar to suspensions of sedimenting particles. The flow in the region around C. reinhardtii where significant hydrodynamic interaction is likely to occur differs qualitatively from a "puller" stresslet, and can be described by a simple three-Stokeslet model.Comment: 4 pages, 4 figures, accepted for publication in PR

Ultrasonic Methods for Characterizing the Interface in Composites

Micromechanical modeling studies of composite materials have highlighted the need for better characterization of the interface zone in composite materials. Bulk behavior in composites has been predicted to be strongly influenced by the local elastic properties, residual stresses, and adhesion of the interface. Techniques to nondestructively measure these newly perceived quantities of importance do not exist. Thus it is not possible experimentally to (i) confirm the micromechanical model predictions, (ii) explore the relationships between interface properties and processing variables, and (iii) ensure acceptable interface properties in commercially fabricated composites. We report here the current status of a SDIO/ONR funded research program directed at developing experimental techniques for characterizing the interface zone in composites through the use of ultrasonic interface waves [1]

Cronin Effect and High Transver Momentum Suppression in D+Au Collisions

Great interest has attached to recent D+Au, s^(1/2) = 200 A GeV data at RHIC, obtained with the BRAHMS detector. Between pseudorapidity eta=0 and eta=3.2 the appropriately defined ratio R[DAu/PP], comparing transverse momentum spectra of D+Au to P+P exhibits a steady decrease with eta. This diminuition is examined within a two-stage simulation, the last stage being a purely hadronic, reduced energy cascade. The result is an adequate description of the data including the so-called Cronin effect. Additionally there is clear evidence for suppression, in the second stage, of relatively high transverse momentum, eta=0, leading mesons, i.e. the Cronin effect, only near mid-rapidity, is appreciably muted by final state interactions.Comment: 13 pages, 6 figure

Fluid dynamics and noise in bacterial cell-cell and cell-surface scattering

Bacterial processes ranging from gene expression to motility and biofilm formation are constantly challenged by internal and external noise. While the importance of stochastic fluctuations has been appreciated for chemotaxis, it is currently believed that deterministic long-range fluid dynamical effects govern cell-cell and cell-surface scattering - the elementary events that lead to swarming and collective swimming in active suspensions and to the formation of biofilms. Here, we report the first direct measurements of the bacterial flow field generated by individual swimming Escherichia coli both far from and near to a solid surface. These experiments allowed us to examine the relative importance of fluid dynamics and rotational diffusion for bacteria. For cell-cell interactions it is shown that thermal and intrinsic stochasticity drown the effects of long-range fluid dynamics, implying that physical interactions between bacteria are determined by steric collisions and near-field lubrication forces. This dominance of short-range forces closely links collective motion in bacterial suspensions to self-organization in driven granular systems, assemblages of biofilaments, and animal flocks. For the scattering of bacteria with surfaces, long-range fluid dynamical interactions are also shown to be negligible before collisions; however, once the bacterium swims along the surface within a few microns after an aligning collision, hydrodynamic effects can contribute to the experimentally observed, long residence times. As these results are based on purely mechanical properties, they apply to a wide range of microorganisms.Comment: 9 pages, 2 figures, http://www.pnas.org/content/108/27/1094

Strangeness Suppression in Proton-Proton Collisions

We analyse strangeness production in proton-proton (pp) collisions at SPS and RHIC energies, using the recently advanced NeXus approach. After having verified that the model reproduces well the existing data, we interpret the results: strangeness is suppressed in proton-proton collisions at SPS energy as compared to electron-positron (e+e-) annihilation due to the limited masses of the strings produced in the reaction, whereas high energy pp and e+e- collisions agree quantitatively . Thus strangeness suppression at SPS energies is a consequence of the limited phase-space available in string fragmentation.Comment: 7 Figures, 4 Page

Consistent Treatment of Soft and Hard Processes in Hadronic Interactions

The QCD improved parton model is a very successful concept to treat processes in hadronic interactions, whenever large partonic transverse momenta are involved. However, cross sections diverge in the limit p_T -> 0, and the usual treatment is the definition of a lower cutoff p_T_min, such that processes with a smaller p_T -- so-called soft processes -- are simply ignored, which is certainly not correct for example at RHIC energies. A more consistent procedure amounts to introduce a technical parameter Q_0^2, referred to as soft virtuality scale, which is nothing but an artificial borderline between soft and hard physics. We will discuss such a formalism, which coincides with the improved parton model for high p_T processes and with the phenomenological treatment of soft scattering, when only small virtualities are involved. The most important aspect of our approach is that it allows to obtain a smooth transition between soft and hard scattering, and therefore no artificial dependence on a cutoff parameter should appear.Comment: 19 pages, 19 figure