arXiv subjects

Raymond E. Goldstein

Publications and source records attributed to Raymond E. Goldstein.

At least 19 recordsLinked to original sources

arXiv2026

Theory of Cell Body Lensing and Phototaxis Sign Reversal in "Eyeless" Mutants of $Chlamydomonas$

Phototaxis of many species of green algae relies upon directional sensitivity of their membrane-bound photoreceptors, which arises from the presence of a pigmented "eyespot" behind them that blocks light passing through the cell body from reaching the photoreceptor. A decade ago it was discovered that the spherical cell body of the alga $Chlamydomonas~reinhardtii$ acts as a lens to concentrate incoming light, and that in "eyeless" mutants of $Chlamydomonas$ the consequence of that focused light reaching the photoreceptor from behind is a reversal in the sign of phototaxis relative to the wild type behavior. We present a quantitative theory of this sign reversal by completing a recent simplified analysis of lensing [Yang, et al., Phys. Rev. E 113, 022401 (2026)] and incorporating it into an adaptive model for $Chlamydomonas$ phototaxis. This model shows that phototactic dynamics in the presence of lensing is subtle because of the existence of internal light caustics when the cellular index of refraction exceeds that of water. During each period of cellular rotation about its body-fixed axis, the photoreceptor receives two competing signals: a relatively long, slowly-varying signal from the direct illumination, and a stronger, shorter, rapidly-varying lensed signal. The reversal of the sign of phototaxis is then a consequence of the dominance of the flagellar photoresponse to the signal with the higher time derivative. These features lead to a quantitative understanding of phototaxis sign reversal, including bistability in the direction choice, a prediction that can be tested in single-cell tracking studies of mutant phototaxis.

cond-mat.soft↗

arXiv2025

Phototactic Decision-Making by Micro-Algae

We study how simple eukaryotic organisms make decisions in response to competing stimuli in the context of phototaxis by the unicellular alga $Chlamydomonas~reinhardtii$. While negatively phototactic cells swim directly away from a collimated light beam, when presented with two beams of adjustable intersection angle and intensities, we find that cells swim in a direction given by an intensity-weighted average of the two light propagation vectors. This geometrical law is a fixed point of an adaptive model of phototaxis and minimizes the average light intensity falling on the anterior pole of the cell. At large angular separations, subpopulations of cells swim away from one source or the other, or along the direction of the geometrical law, with some cells stochastically switching between the three directions. This behavior is shown to arise from a population-level distribution of photoreceptor locations that breaks front-back symmetry of photoreception.

cond-mat.soft↗

arXiv2025

Algal Optics

Nearly a decade ago it was discovered that the spherical cell body of the alga $Chlamydomonas~reinhardtii$ can act as a lens to concentrate incoming light onto the cell's membrane-bound photoreceptor and thereby affect phototaxis. Since many nearly transparent cells in marine environments have complex, often non-axisymmetric shapes, this observation raises fundamental, yet little-explored questions in biological optics about light refraction by the bodies of microorganisms. There are two distinct contexts for such questions: the $absorption$ problem for $incoming$ light, typified by photosynthetic activity taking place in the chloroplasts of green algae, and the $emission$ problem for $outgoing$ light, where the paradigm is bioluminescence emitted from scintillons within dinoflagellates. Here we examine both of these aspects of ``algal optics" in the special case where the absorption or emission is localized in structures that are small relative to the overall organism size, taking into account both refraction and reflections at the cell-water boundary. Analytical and numerical results are developed for the distribution of light intensities inside and outside the body, and we establish certain duality relationships that connect the incoming and outgoing problems. For strongly non-spherical shapes we find lensing effects that may have implications for photosynthetic activity and for the angular distribution of light emitted during bioluminescent flashes.

cond-mat.soft↗

arXiv2025

Photometric Decision-Making During the Dawn Choruses of Cicadas

We report the first quantitative study of the onset of dawn choruses of cicadas in several natural habitats. A time-frequency analysis of the acoustical signals is used to define an order parameter for the development of collective singing. The ensemble of recordings reveals that the chorus onset times accurately track the changing sunrise times over the course of many weeks, occurring within civil twilight at a solar elevation of -$3.8^\circ \pm 0.2^\circ$. Despite day-to-day variations in the amplitude of fully developed choruses, the order parameter data collapse to a common sigmoidal curve when scaled by those amplitudes and shifted by the onset time, revealing a characteristic rise time of ~60 s for a chorus to reach saturation amplitude. The results are used to obtain the cumulative distribution function of singing as a function of ground illumination, from which is obtained a generalized susceptibility which exhibits a narrow peak with a half-width of $\sim\! 12\%$. The variance of the order parameter exhibits a similar peak, suggesting that a generalized fluctuation-dissipation theorem holds for this system. A model of decision-making under ramps of a control parameter is developed and can achieve a quantitative match to the data. It suggest that sharpness of the susceptibility peak reflects cooperative decision-making arising from acoustic communication.

cond-mat.soft↗

arXiv2024

Spatiotemporal distribution of the glycoprotein pherophorin II reveals stochastic geometry of the growing ECM of $Volvox~carteri$

The evolution of multicellularity involved the transformation of a simple cell wall of unicellular ancestors into a complex, multifunctional extracellular matrix (ECM). A suitable model organism to study the formation and expansion of an ECM during ontogenesis is the multicellular green alga $Volvox~carteri$, which, along with the related volvocine algae, produces a complex, self-organized ECM composed of multiple substructures. These self-assembled ECMs primarily consist of hydroxyproline-rich glycoproteins, a major component of which is pherophorins. To investigate the geometry of the growing ECM, we fused the $yfp$ gene with the gene for pherophorin II (PhII) in $V.~carteri$. Confocal microscopy reveals PhII:YFP localization at key structures within the ECM, including the boundaries of compartments surrounding each somatic cell and the outer surface of the organism. Image analysis during the life cycle allows the stochastic geometry of those growing compartments to be quantified. We find that their areas and aspect ratios exhibit robust gamma distributions and exhibit a transition from a tight polygonal to a looser acircular packing geometry with stable eccentricity over time, evoking parallels and distinctions with the behavior of hydrated foams. These results provide a quantitative benchmark for addressing a general, open question in biology: How do cells produce structures external to themselves in a robust and accurate manner?

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arXiv2024

The 2024 Motile Active Matter Roadmap

Activity and autonomous motion are fundamental aspects of many living and engineering systems. Here, the scale of biological agents covers a wide range, from nanomotors, cytoskeleton, and cells, to insects, fish, birds, and people. Inspired by biological active systems, various types of autonomous synthetic nano- and micromachines have been designed, which provide the basis for multifunctional, highly responsive, intelligent active materials. A major challenge for understanding and designing active matter is their inherent non-equilibrium nature due to persistent energy consumption, which invalidates equilibrium concepts such as free energy, detailed balance, and time-reversal symmetry. Furthermore, interactions in ensembles of active agents are often non-additive and non-reciprocal. An important aspect of biological agents is their ability to sense the environment, process this information, and adjust their motion accordingly. It is an important goal for the engineering of micro-robotic systems to achieve similar functionality. With many fundamental properties of motile active matter now reasonably well understood and under control, the ground is prepared for the study of physical aspects and mechanisms of motion in complex environments, of the behavior of systems with new physical features like chirality, of the development of novel micromachines and microbots, of the emergent collective behavior and swarming of intelligent self-propelled particles, and of particular features of microbial systems. The vast complexity of phenomena and mechanisms involved in the self-organization and dynamics of motile active matter poses major challenges, which can only be addressed by a truly interdisciplinary effort involving scientists from biology, chemistry, ecology, engineering, mathematics, and physics.

cond-mat.soft↗

arXiv2024

Feeders and Expellers, Two Types of Animalcules With Outboard Cilia, Have Distinct Surface Interactions

Within biological fluid dynamics, it is conventional to distinguish between "puller" and "pusher" microswimmers on the basis of the forward or aft location of the flagella relative to the cell body: typically, bacteria are pushers and algae are pullers. Here we note that since many pullers have "outboard" cilia or flagella displaced laterally from the cell centerline on both sides of the organism, there are two important subclasses whose far-field is that of a stresslet, but whose near field is qualitatively more complex. The ciliary beat creates not only a propulsive force but also swirling flows that can be represented by paired rotlets with two possible senses of rotation, either "feeders" that sweep fluid toward the cell apex, or "expellers" that push fluid away. Experimental studies of the rotifer $Brachionus~plicatilis$ in combination with earlier work on the green algae $Chlamydomonas~reinhardtii$ show that the two classes have markedly different interactions with surfaces. When swimming near a surface, expellers such as $C.~reinhardtii$ scatter from the wall, whereas a feeder like $B.~plicatilis$ stably attaches. This results in a stochastic "run-and-stick" locomotion, with periods of ballistic motion parallel to the surface interrupted by trapping at the surface.

cond-mat.soft↗

arXiv2024

Stochastic Voronoi Tessellations as Models for Cellular Neighborhoods in Simple Multicellular Organisms

Recent work on distinct multicellular organisms has revealed a hitherto unknown type of biological noise; rather than a regular arrangement, cellular neighborhood volumes, obtained by Voronoi tessellations of the cell locations, are broadly distributed and consistent with gamma distributions. We propose an explanation for those observations in the case of the alga $Volvox$, whose somatic cells are embedded in an extracellular matrix (ECM) they export. Both a solvable one-dimensional model of ECM growth derived from bursty transcriptional activity and a two-dimensional "Voronoi liquid" model are shown to provide one-parameter families that smoothly interpolate between the empirically-observed near-maximum-entropy gamma distributions and the crystalline limit of Gaussian distributions governed by the central limit theorem. These results highlight a universal consequence of intrinsic biological noise on the architecture of certain tissues.

cond-mat.soft↗

arXiv2024

The Architecture of Sponge Choanocyte Chambers Maximizes Mechanical Pumping Efficiency

Sponges, the basalmost members of the animal kingdom, exhibit a range of complex architectures in which microfluidic channels connect multitudes of spherical chambers lined with choanocytes, flagellated filter-feeding cells. Choanocyte chambers can possess scores or even hundreds of such cells, which drive complex flows entering through porous walls and exiting into the sponge channels. One of the mysteries of the choanocyte chamber is its spherical shape, as it seems inappropriate for inducing directional transport since many choanocyte flagella beat in opposition to such a flow. Here we combine direct imaging of choanocyte chambers in living sponges with computational studies of many-flagella models to understand the connection between chamber architecture and directional flow. We find that those flagella that beat against the flow play a key role in raising the pressure inside the choanocyte chamber, with the result that the mechanical pumping efficiency, calculated from the pressure rise and flow rate, reaches a maximum at a small outlet opening angle. Comparison between experimental observations and the results of numerical simulations reveal that the chamber diameter, flagellar wave number and the outlet opening angle of the freshwater sponge $E. muelleri$, as well as several other species, are related in a manner that maximizes the mechanical pumping efficiency. These results indicate the subtle balances at play during morphogenesis of choanocyte chambers, and give insights into the physiology and body design of sponges.

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arXiv2024

Biophysical Fluid Dynamics in a Petri Dish

The humble Petri dish is perhaps the simplest setting in which to examine the locomotion of swimming organisms, particularly those whose body size is tens of microns to millimetres. The fluid layer in such a container has a bottom no-slip surface and a stress-free upper boundary. It is of fundamental interest to understand the flow fields produced by the elementary and composite singularities of Stokes flow in this geometry. Building on the few particular cases that have previously been considered in the literature, we study here the image systems for the primary singularities of Stokes flow subject to such boundary conditions - the stokeslet, rotlet, source, rotlet dipole, source dipole and stresslet - paying particular attention to the far-field behavior. In several key situations, the depth-averaged fluid flow is accurately captured by the solution of an associated Brinkman equation whose screening length is proportional to the depth of the fluid layer. The case of hydrodynamic bound states formed by spinning microswimmers near a no-slip surface, discovered first using the alga $Volvox$, is reconsidered in the geometry of a Petri dish, where the power-law attractive interaction between microswimmers acquires unusual exponentially screened oscillations.

cond-mat.soft↗

arXiv2023

Anti-Diffusion in an Algae-Bacteria Microcosm: Photosynthesis, Chemotaxis, and Expulsion

In Nature there are significant relationships known between microorganisms from two kingdoms of life, as in the supply of vitamin B$_{12}$ by bacteria to algae. Such interactions motivate general investigations into the spatio-temporal dynamics of metabolite exchanges. Here we study by experiment and theory a model system: a coculture of the bacterium $B. subtilis$, an obligate aerobe that is chemotactic to oxygen, and a nonmotile mutant of the alga $C. reinhardtii$, which photosynthetically produces oxygen when illuminated. Strikingly, when a shaft of light illuminates a thin, initially uniform suspension of the two, the chemotactic influx of bacteria to the photosynthetically active region leads to expulsion of the algae from that area. This effect arises from algal transport due to spatially-varying collective behavior of bacteria, and is mathematically related to the ``turbulent diamagnetism" associated with magnetic flux expulsion in stars.

cond-mat.soft↗

arXiv2023

How do Cicadas Emerge Together? Thermophysical Aspects of Their Collective Decision-Making

Certain periodical cicadas exhibit life cycles with durations of 13 or 17 years, and it is now generally accepted that such large prime numbers arise evolutionarily to avoid synchrony with predators. Less well explored is the question of how, in the face of intrinsic biological and environmental noise, insects within a brood emerge together in large successive swarms from underground during springtime warming. Here we consider the decision-making process of underground cicada nymphs experiencing random but spatially-correlated thermal microclimates like those in nature. Introducing short-range communication between insects leads to a model of consensus building that maps on to the statistical physics of an Ising model with a quenched, spatially correlated random magnetic field and annealed site dilution, which displays the kinds of collective swarms seen in nature.

cond-mat.soft↗

arXiv2023

Swimming, Feeding and Inversion of Multicellular Choanoflagellate Sheets

The recent discovery of the striking sheet-like multicellular choanoflagellate species $Choanoeca~flexa$ that dynamically interconverts between two hemispherical forms of opposite orientation raises fundamental questions in cell and evolutionary biology, as choanoflagellates are the closest living relatives of animals. It similarly motivates questions in fluid and solid mechanics concerning the differential swimming speeds in the two states and the mechanism of curvature inversion triggered by changes in the geometry of microvilli emanating from each cell. Here we develop fluid dynamical and mechanical models to address these observations and show that they capture the main features of the swimming, feeding, and inversion of $C.~flexa$ colonies.

cond-mat.soft↗

arXiv2022

Stabilization of Microbial Communities by Responsive Phenotypic Switching

Clonal microbes can switch between different phenotypes and recent theoretical work has shown that stochastic switching between these subpopulations can stabilize microbial communities. This phenotypic switching need not be stochastic, however, but could also be in response to environmental factors, both biotic and abiotic. Here, motivated by the bacterial persistence phenotype, we explore the ecological effects of such responsive switching by analyzing phenotypic switching in response to competing species. We show that the stability of microbial communities with responsive switching differs generically from that of communities with stochastic switching only. To understand the mechanisms by which responsive switching stabilizes coexistence, we go on to analyze simple two-species models. Combining exact results and numerical simulations, we extend the classical stability results for the competition of two species without phenotypic variation to the case in which one species switches, stochastically and responsively, between two phenotypes. In particular, we show that responsive switching can stabilize coexistence even when stochastic switching on its own does not affect the stability of the community.

q-bio.PE↗

arXiv2022

Collapse of a hemicatenoid bounded by a solid wall: instability and dynamics driven by surface Plateau border friction

The collapse of a catenoidal soap film when the rings supporting it are moved beyond a critical separation is a classic problem in interface motion in which there is a balance between surface tension and the inertia of the surrounding air, with film viscosity playing only a minor role. Recently [Goldstein, et al., Phys. Rev. E 104, 035105 (2021)], we introduced a variant of this problem in which the catenoid is bisected by a glass plate located in a plane of symmetry perpendicular to the rings, producing two identical hemicatenoids, each with a surface Plateau border (SPB) on the glass plate. Beyond the critical ring separation, the hemicatenoids collapse in a manner qualitatively similar to the bulk problem, but their motion is governed by the frictional forces arising from viscous dissipation in the SPBs. Here we present numerical studies of a model that includes classical friction laws for SPB motion on wet surfaces and show consistency with our experimental measurements of the temporal evolution of this process. This study can help explain the fragmentation of bubbles inside very confined geometries such as porous materials or microfluidic devices.

cond-mat.soft↗

arXiv2021

Biofilm Growth Under Elastic Confinement

Bacteria often form surface-bound communities, embedded in a self-produced extracellular matrix, called biofilms. Quantitative studies of their growth have typically focused on unconfined expansion above solid or semi-solid surfaces, leading to exponential radial growth. This geometry does not accurately reflect the natural or biomedical contexts in which biofilms grow in confined spaces. Here we consider one of the simplest confined geometries: a biofilm growing laterally in the space between a solid surface and an overlying elastic sheet. A poroelastic framework is utilised to derive the radial growth rate of the biofilm; it reveals an additional self-similar expansion regime, governed by the stiffness of the matrix, leading to a finite maximum radius, consistent with our experimental observations of growing $Bacillus~subtilis$ biofilms confined by PDMS.

cond-mat.soft↗

arXiv2021

Cellular organization in lab-evolved and extant multicellular species obeys a maximum entropy law

The prevalence of multicellular organisms is due in part to their ability to form complex structures. How cells pack in these structures is a fundamental biophysical issue, underlying their functional properties. However, much remains unknown about how cell packing geometries arise, and how they are affected by random noise during growth - especially absent developmental programs. Here, we quantify the statistics of cellular neighborhoods of two different multicellular eukaryotes: lab-evolved "snowflake" yeast and the green alga $Volvox~carteri$. We find that despite large differences in cellular organization, the free space associated with individual cells in both organisms closely fits a modified gamma distribution, consistent with maximum entropy predictions originally developed for granular materials. This 'entropic' cellular packing ensures a degree of predictability despite noise, facilitating parent-offspring fidelity even in the absence of developmental regulation. Together with simulations of diverse growth morphologies, these results suggest that gamma-distributed cell neighborhood sizes are a general feature of multicellularity, arising from conserved statistics of cellular packing.

cond-mat.soft↗

arXiv2021

Fluid mechanics of mosaic ciliated tissues

In tissues as diverse as amphibian skin and the human airway, the cilia that propel fluid are grouped in sparsely distributed multiciliated cells (MCCs). We investigate fluid transport in this "mosaic" architecture, with emphasis on the trade-offs that may have been responsible for its evolutionary selection. Live imaging of MCCs in embryos of the frog $Xenopus~laevis$ shows that cilia bundles behave as active vortices that produce a flow field accurately represented by a local force applied to the fluid. A coarse-grained model that self-consistently couples bundles to the ambient flow reveals that hydrodynamic interactions between MCCs limit their rate of work so that when the system size is large compared to a single MCC, they best shear the tissue at low area coverage, a result that mirrors findings for other sparse distributions such as cell receptors and leaf stomata.

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