arXiv subjects
Raymond E. Goldstein
Publications and source records attributed to Raymond E. Goldstein.
Why clothes don't fall apart: tension transmission in staple yarns
The problem of how staple yarns transmit tension is addressed within abstract models in which the Amontons-Coulomb friction laws yield a linear programming (LP) problem for the tensions in the fiber elements. We find there is a percolation transition such that above the percolation threshold the transmitted tension is in principle unbounded, We determine that the mean slack in the LP constraints is a suitable order parameter to characterize this supercritical state. We argue the mechanism is generic, and in practical terms corresponds to a switch from a ductile to a brittle failure mode accompanied by a significant increase in mechanical strength.
Microbial mutualism at a distance: the role of geometry in diffusive exchanges
The exchange of diffusive metabolites is known to control the spatial patterns formed by microbial populations, as revealed by recent studies in the laboratory. However, the matrices used, such as agarose pads, lack the structured geometry of many natural microbial habitats, including in the soil or on the surfaces of plants or animals. Here we address the important question of how such geometry may control diffusive exchanges and microbial interaction. We model mathematically mutualistic interactions within a minimal unit of structure: two growing reservoirs linked by a diffusive channel through which metabolites are exchanged. The model is applied to study a synthetic mutualism, experimentally parameterised on a model algal-bacterial co-culture. Analytical and numerical solutions of the model predict conditions for the successful establishment of remote mutualisms, and how this depends, often counterintutively, on diffusion geometry. We connect our findings to understanding complex behaviour in synthetic and naturally occurring microbial communities.
A compact Eulerian representation of axisymmetric inviscid vortex sheet dynamics
A classical problem in fluid mechanics is the motion of an axisymmetric vortex sheet evolving under the action of surface tension, surrounded by an inviscid fluid. Lagrangian descriptions of these dynamics are well-known, involving complex nonlocal expressions for the radial and longitudinal velocities in terms of elliptic integrals. Here we use these prior results to arrive at a remarkably compact and exact Eulerian evolution equation for the sheet radius $r(z,t)$ in an explicit flux form associated with the conservation of enclosed volume. The flux appears as an integral involving the pairwise mutual induction formula for vortex loop pairs first derived by Helmholtz and Maxwell. We show how the well-known linear stability results for cylindrical vortex sheets in the presence of surface tension and streaming flows [A.M. Sterling and C.A. Sleicher, $J.~Fluid~Mech.$ ${\bf 68}$, 477 (1975)] can be obtained directly from this formulation. Furthermore, the inviscid limit of the empirical model of Eggers and Dupont [$J.~Fluid~Mech.$ $\textbf{262}$ 205 (1994); $SIAM~J.~Appl.~Math.$ ${\bf 60}$, 1997 (2000)], which has served as the basis for understanding singularity formation in droplet pinchoff, is derived within the present formalism as the leading order term in an asymptotic analysis for long slender axisymmetric vortex sheets, and should provide the starting point for a rigorous analysis of singularity formation.
The role of tumbling frequency and persistence in optimal run-and-tumble chemotaxis
One of simplest examples of navigation found in nature is run-and-tumble chemotaxis. Tumbles reorient cells randomly, and cells can drift toward attractants or away from repellents by biasing the frequency of these events. The post-tumble swimming directions are typically correlated with those prior, as measured by the variance of the reorientation angle distribution. This variance can range from large, in the case of bacteria, to so small that tumble events are imperceptible, as observed in choanoflagellates. This raises the question of optimality: why is such a range of persistence observed in nature? Here we study persistent run-and-tumble dynamics, focusing first on the optimisation of the linearised chemotactic response within the two-dimensional parameter space of tumble frequency and angular persistence. Although an optimal persistence does exist for a given tumble frequency, in the full parameter space there is a continuum of optimal solutions. Introducing finite tumble times that depend on the persistence can change this picture, illuminating one possible method for selecting tumble persistence based on species-specific reorientation dynamics. Moving beyond linear theory we find that optimal chemotactic strengths exist, and that these maximise reaction when swimming in a wrong direction, but have little or no reaction when swimming with even the slightest projection along the chemoattractant gradient.
Mechanics and Variability of Cell Sheet Folding in the Embryonic Inversion of $Volvox$
Many embryonic deformations during development are the global result of local cell shape changes and other local active cell sheet deformations. Morphogenesis does not only therefore rely on the ability of the tissue to produce these active deformations, but also on the ability to regulate them in such a way as to overcome the intrinsic variability of and geometric constraints on the tissue. Here, we explore the interplay of regulation and variability in the green alga $Volvox$, whose spherical embryos turn themselves inside out to enable motility. Through a combination of light sheet microscopy and theoretical analysis, we quantify the variability of this inversion and analyse its mechanics in detail to show how shape variability arises from a combination of geometry, mechanics, and active regulation.
Scattering of biflagellate microswimmers from surfaces
We use a three-bead-spring model to investigate the dynamics of bi-flagellate micro-swimmers near a surface. While the primary dynamics and scattering are governed by geometric-dependent direct contact, the fluid flows generated by the swimmer locomotion are important in orienting it toward or away from the surface. Flagellar noise and in particular cell spinning about the main axis help a surface-trapped swimmer escape, whereas the time a swimmer spends at the surface depends on the incident angle. The dynamics results from a nuanced interplay of direct collisions, hydrodynamics, noise and the swimmer geometry. We show that to correctly capture the dynamics of a biflagellate swimmer, minimal models need to resolve the shape asymmetry.
Spontaneous Oscillations of Elastic Filaments Induced by Molecular Motors
It is known from the wave-like motion of microtubules in motility assays that the piconewton forces that motors produce can be sufficient to bend the filaments. In cellular phenomena such as cytosplasmic streaming, molecular motors translocate along cytoskeletal filaments, carrying cargo which entrains fluid. When large numbers of such forced filaments interact through the surrounding fluid, as in particular stages of oocyte development in $Drosophila~melanogaster$, complex dynamics are observed, but the detailed mechanics underlying them has remained unclear. Motivated by these observations, we study here perhaps the simplest model for these phenomena: an elastic filament, pinned at one end, acted on by a molecular motor treated as a point force. Because the force acts tangential to the filament, no matter what its shape, this "follower-force" problem is intrinsically non-variational, and thereby differs fundamentally from Euler buckling, where the force has a fixed direction, and which, in the low Reynolds number regime, ultimately leads to a stationary, energy-minimizing shape. Through a combination of linear stability theory, analytical study of a solvable simplified "two-link" model, and numerical studies of the full elastohydrodynamic equations of motion we elucidate the Hopf bifurcation that occurs with increasing forcing of a filament, leading to flapping motion analogous to the high Reynolds number oscillations of a garden hose with a free end.
Run stop shock, run shock run: Spontaneous and stimulated gait-switching in a unicellular octoflagellate
In unicellular flagellates, growing evidence suggests control over a complex repertoire of swimming gaits is conferred intracellularly by ultrastructural components, resulting in motion that depends on flagella number and configuration. We report the discovery of a novel, tripartite motility in an octoflagellate alga, comprising a forward gait ($run$), a fast knee-jerk response with dramatic reversals in beat waveform ($shock$), and, remarkably, long quiescent periods ($stop$) within which the flagella quiver. In a reaction graph representation, transition probabilities show that gait switching is only weakly reversible. Shocks occur spontaneously but are also triggered by direct mechanical contact. In this primitive alga, the capability for a millisecond stop-start switch from rest to full speed implicates an early evolution of excitable signal transduction to and from peripheral appendages.
Evaporation-driven convective flows in suspensions of non-motile bacteria
We report a novel form of convection in suspensions of the bioluminiscent marine bacterium $Photobacterium~phosphoreum$. Suspensions of these bacteria placed in a chamber open to the air create persistent luminiscent plumes most easily visible when observed in the dark. These flows are strikingly similar to the classical bioconvection pattern of aerotactic swimming bacteria, which create an unstable stratification by swimming upwards to an air-water interface, but they are a puzzle since the strain of $P.~phosphoreum$ used does not express flagella and therefore cannot swim. Systematic experimentation with suspensions of microspheres reveals that these flow patterns are driven not by the bacteria but by the accumulation of salt at the air-water interface due to evaporation of the culture medium; even at room temperature and humidity, and physiologically relevant salt concentrations, the rate of water evaporation is sufficient to drive convection patterns. A mathematical model is developed to understand the mechanism of plume formation, and linear stability analysis as well as numerical simulations were carried out to support the conclusions. While evaporation-driven convection has not been discussed extensively in the context of biological systems, these results suggest that the phenomenon may be relevant in other systems, particularly those using microorganisms of limited motility.
Traces of surfactants can severely limit the drag reduction of superhydrophobic surfaces
Superhydrophobic surfaces (SHSs) have the potential to achieve large drag reduction for internal and external flow applications. However, experiments have shown inconsistent results, with many studies reporting significantly reduced performance. Recently, it has been proposed that surfactants, ubiquitous in flow applications, could be responsible, by creating adverse Marangoni stresses. Yet, testing this hypothesis is challenging. Careful experiments with purified water show large interfacial stresses and, paradoxically, adding surfactants yields barely measurable drag increases. This suggests that other physical processes, such as thermal Marangoni stresses or interface deflection, could explain the lower performance. To test the surfactant hypothesis, we perform the first numerical simulations of flows over a SHS inclusive of surfactant kinetics. These simulations reveal that surfactant-induced stresses are significant at extremely low concentrations, potentially yielding a no-slip boundary condition on the air--water interface (the "plastron") for surfactant amounts below typical environmental values. These stresses decrease as the streamwise distance between plastron stagnation points increases. We perform microchannel experiments with thermally-controlled SHSs consisting of streamwise parallel gratings, which confirm this numerical prediction. We introduce a new, unsteady test of surfactant effects. When we rapidly remove the driving pressure following a loading phase, a backflow develops at the plastron, which can only be explained by surfactant gradients formed in the loading phase. This demonstrates the significance of surfactants in deteriorating drag reduction, and thus the importance of including surfactant stresses in SHS models. Our time-dependent protocol can assess the impact of surfactants in SHS testing and guide future mitigating designs.
A biology journal that can teach physicists a lesson in peer review
This is a Commentary in $Physics~Today$ on the novel review process developed by the biology journal $eLife$, with the suggestion that it be adopted by physics journals.
Elastohydrodynamic Synchronization of Adjacent Beating Flagella
It is now well established that nearby beating pairs of eukaryotic flagella or cilia typically synchronize in phase. A substantial body of evidence supports the hypothesis that hydrodynamic coupling between the active filaments, combined with waveform compliance, provides a robust mechanism for synchrony. This elastohydrodynamic mechanism has been incorporated into `bead-spring' models in which flagella are represented by microspheres tethered by radial springs as internal forces drive them about orbits. While these low-dimensional models reproduce the phenomenon of synchrony, their parameters are not readily relatable to those of flagella. More realistic models which reflect the elasticity of the axonemes and active force generation take the form of fourth-order nonlinear PDEs. While computational studies have shown synchrony, the effects of hydrodynamic coupling between nearby filaments governed by such models have been theoretically examined only in the regime of interflagellar distances $d$ large compared to flagellar length $L$. Yet, in many biological situations $d/L \ll 1$. Here, we first present an asymptotic analysis of the hydrodynamic coupling between two filaments in the regime $d/L \ll 1$, and find that the form of the coupling is independent of the details of the internal forces that govern the motion of the filaments. The analysis is like the localized induction approximation for vortex filament motion, extended to the case of mutual induction. To understand how the coupling mechanism leads to synchrony of extended objects, we introduce a heuristic model of flagellar beating, a single fourth-order nonlinear PDE whose form is derived from symmetry considerations, the physics of elasticity, and the overdamped nature of the dynamics. Analytical and numerical studies of this model illustrate how synchrony between two filaments is achieved through the asymptotic coupling.
A Theory of Shape-Shifting Droplets
Recent studies of cooled oil emulsion droplets uncovered transformations into a host of flattened shapes with straight edges and sharp corners, driven by a partial phase transition of the bulk liquid phase. Here, we explore theoretically the simplest geometric competition between this phase transition and surface tension in planar polygons, and recover the observed sequence of shapes and their statistics in qualitative agreement with experiments. Extending the model to capture some of the three-dimensional structure of the droplets, we analyze the evolution of protrusions sprouting from the vertices of the platelets and the topological transition of a puncturing planar polygon.
Filter-feeding, near-field flows, and the morphologies of colonial choanoflagellates
Efficient uptake of nutrients from the environment is an important component in the fitness of all microorganisms, and its dependence on size may reveal clues to the origins of evolutionary transitions to multicellularity. Because potential benefits in uptake rates must be viewed in the context of other costs and benefits of size, such as varying predation rates and the increased metabolic costs associated with larger and more complex body plans, the uptake rate itself is not necessarily that which is optimized by evolution. Uptake rates can be strongly dependent on local organism geometry and its swimming speed, providing selective pressure for particular arrangements. Here we examine these issues for choanoflagellates, filter-feeding microorganisms that are the closest relatives of the animals. We explore the different morphological variations of the choanoflagellete $Salpingoeca~rosetta$, which can exist as a swimming cell, a sessile thecate cell, and as colonies of cells in various shapes. In the absence of other requirements and in a homogeneously nutritious environment, we find that the optimal strategy to maximize filter feeding by the collar of microvilli is to swim fast, which favours swimming unicells. In large external flows, the sessile thecate cell becomes advantageous. Effects of prey diffusion are discussed and also found to be to the advantage of the swimming unicell.
Aerotaxis in the Closest Relatives of Animals
As the closest unicellular relatives of animals, choanoflagellates serve as useful model organisms for understanding the evolution of animal multicellularity. An important factor in animal evolution was the increasing ocean oxygen levels in the Precambrian, which are thought to have influenced the emergence of complex multicellular life. As a first step in addressing these conditions, we study here the response of the colony-forming choanoflagellate $Salpingoeca~rosetta$ to oxygen gradients. Using a microfluidic device that allows spatio-temporal variations in oxygen concentrations, we report the discovery that $S.~rosetta$ display positive aerotaxis. Analysis of the spatial population distributions provides evidence for logarithmic sensing of oxygen, which enhances sensing in low oxygen neighborhoods. Analysis of search strategy models on the experimental colony trajectories finds that choanoflagellate aerotaxis is consistent with stochastic navigation, the statistics of which are captured using an effective continuous version based on classical run-and-tumble chemotaxis.
Long-range interactions and phase defects in chains of fluid-coupled oscillators
Eukaryotic cilia and flagella are chemo-mechanical oscillators capable of generating long-range coordinated motions known as metachronal waves. Pair synchronization is a fundamental requirement for these collective dynamics, but it is generally not sufficient for collective phase-locking, chiefly due to the effect of long-range interactions. Here we explore experimentally and numerically a minimal model for a ciliated surface; hydrodynamically coupled oscillators rotating above a no-slip plane. Increasing their distance from the wall profoundly effects the global dynamics, due to variations in hydrodynamic interaction range. The array undergoes a transition from a traveling wave to either a steady chevron pattern or one punctuated by periodic phase defects. Within the transition between these regimes the system displays behavior reminiscent of chimera states.
Coordinated Beating of Algal Flagella is Mediated by Basal Coupling
Cilia and flagella often exhibit synchronized behavior; this includes phase locking, as seen in {\it Chlamydomonas}, and metachronal wave formation in the respiratory cilia of higher organisms. Since the observations by Gray and Rothschild of phase synchrony of nearby swimming spermatozoa, it has been a working hypothesis that synchrony arises from hydrodynamic interactions between beating filaments. Recent work on the dynamics of physically separated pairs of flagella isolated from the multicellular alga {\it Volvox} has shown that hydrodynamic coupling alone is sufficient to produce synchrony. However, the situation is more complex in unicellular organisms bearing few flagella. We show that flagella of {\it Chlamydomonas} mutants deficient in filamentary connections between basal bodies display markedly different synchronization from the wild type. We perform micromanipulation on configurations of flagella and conclude that a mechanism, internal to the cell, must provide an additional flagellar coupling. In naturally occurring species with 4, 8, or even 16 flagella, we find diverse symmetries of basal body positioning and of the flagellar apparatus that are coincident with specific gaits of flagellar actuation, suggesting that it is a competition between intracellular coupling and hydrodynamic interactions that ultimately determines the precise form of flagellar coordination in unicellular algae.