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Marco Polin

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Published work

13 published item(s)

preprint2026arXiv

Kin-ematic Exclusion in Active Matter: Modelling Mutual Inhibition in \textit{Pseudomonas aeruginosa} Sibling Colonies

The striking variety of macroscopic morphologies displayed by bacterial colonies depends on microscopic environmental and behavioural details in a manner that is currently not well understood. A surprising example is sibling inhibition, whereby isogenic bacterial colonies spreading in soft agar hydrogels tend to avoid each other and form sharp demarcation lines when growing nearby. Here we investigate this effect with the common pathogen \textit{Pseudomonas aeruginosa}, by combining quantitative density measurements with a minimal biophysical model. Our results show that the phenomenon does not depend on gel compression, lethal inhibition or quorum sensing-dependent cell communication. Instead, colony separation is driven by localised nutrient depletion through a dynamic feedback between growth and motility. The model, which is calibrated using experimental data, captures key observations including the dependence of inhibition strength on the initial nutrient concentration. This work establishes nutrient availability and non-lethal motility inhibition as central factors underlying sibling inhibition, providing a generalisable framework for microbial spatial dynamics with implications for understanding bacterial interactions in tissues, soils and engineered microbiomes.

preprint2022arXiv

Steering self-organisation through confinement

Self-organisation is the spontaneous emergence of spatio-temporal structures and patterns from the interaction of smaller individual units. Examples are found across many scales in very different systems and scientific disciplines, from physics, materials science and robotics to biology, geophysics and astronomy. Recent research has highlighted how self-organisation can be both mediated and controlled by confinement. Confinement occurs through interactions with boundaries, and can function as either a catalyst or inhibitor of self-organisation. It can then become a means to actively steer the emergence or suppression of collective phenomena in space and time. Here, to provide a common framework for future research, we examine the role of confinement in self-organisation and identify overarching scientific challenges across disciplines that need to be addressed to harness its full scientific and technological potential. This framework will not only accelerate the generation of a common deeper understanding of self-organisation but also trigger the development of innovative strategies to steer it through confinement, with impact, e.g., on the design of smarter materials, tissue engineering for biomedicine and crowd management.

preprint2020arXiv

A Python based automated tracking routine for myosin II filaments

The study of motor protein dynamics within cytoskeletal networks is of high interest to physicists and biologists to understand how the dynamics and properties of individual motors lead to cooperative effects and control of overall network behaviour. Here, we report a method to detect and track muscular myosin II filaments within an actin network tethered to supported lipid bilayers. Based on the characteristic shape of myosin II filaments, this automated tracking routine allowed us to follow the position and orientation of myosin II filaments over time, and to reliably classify their dynamics into segments of diffusive and processive motion based on the analysis of displacements and angular changes between time steps. This automated, high throughput method will allow scientists to efficiently analyse motor dynamics in different conditions, and will grant access to more detailed information than provided by common tracking methods, without any need for time consuming manual tracking or generation of kymographs.

preprint2016arXiv

A brief introduction to the model microswimmer {\it Chlamydomonas reinhardtii}

The unicellular biflagellate green alga {\it Chlamydomonas reinhardtii} has been an important model system in biology for decades, and in recent years it has started to attract growing attention also within the biophysics community. Here we provide a concise review of some of the aspects of {\it Chlamydomonas} biology and biophysics most immediately relevant to physicists that might be interested in starting to work with this versatile microorganism.

preprint2016arXiv

Entrainment dominates the interaction of microalgae with micron-sized objects

The incessant activity of swimming microorganisms has a direct physical effect on surrounding microscopic objects, leading to enhanced diffusion far beyond the level of Brownian motion with possible influences on the spatial distribution of non-motile planktonic species and particulate drifters. Here we study in detail the effect of eukaryotic flagellates, represented by the green microalga Chlamydomonas reinhardtii, on microparticles. Macro- and micro-scopic experiments reveal that microorganism-colloid interactions are dominated by rare close encounters leading to large displacements through direct entrainment. Simulations and theoretical modelling show that the ensuing particle dynamics can be understood in terms of a simple jump-diffusion process, combining standard diffusion with Poisson-distributed jumps. This heterogeneous dynamics is likely to depend on generic features of the near-field of swimming microorganisms with front-mounted flagella.

preprint2016arXiv

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.

preprint2016arXiv

Phototaxis beyond turning: persistent accumulation and response acclimation of the microalga Chlamydomonas reinhardtii

Phototaxis is an important reaction to light displayed by a wide range of motile microorganisms. Flagellated eukaryotic microalgae in particular, like the model organism Chlamydomonas reinhardtii, steer either towards or away from light by a rapid and precisely timed modulation of their flagellar activity. Cell steering, however, is only the beginning of a much longer process which ultimately allows cells to determine their light exposure history. This process is not well understood. Here we present a first quantitative study of the long timescale phototactic motility of Chlamydomonas at both single cell and population levels. Our results reveal that the phototactic strategy adopted by these microorganisms leads to an efficient exposure to light, and that the phototactic response is modulated over typical timescales of tens of seconds. The adaptation dynamics for phototaxis and chlorophyll fluorescence show a striking quantitative agreement, suggesting that photosynthesis controls quantitatively how cells navigate a light field.

preprint2015arXiv

Metachronal waves in the flagellar beating of $Volvox$ and their hydrodynamic origin

Groups of eukaryotic cilia and flagella are capable of coordinating their beating over large scales, routinely exhibiting collective dynamics in the form of metachronal waves. The origin of this behaviour -- possibly influenced by both mechanical interactions and direct biological regulation -- is poorly understood, in large part due to lack of quantitative experimental studies. Here we characterise in detail flagellar coordination on the surface of the multicellular alga $Volvox~carteri$, an emerging model organism for flagellar dynamics. Our studies reveal for the first time that the average metachronal coordination observed is punctuated by periodic phase defects during which synchrony is partial and limited to specific groups of cells. A minimal model of hydrodynamically coupled oscillators can reproduce semi-quantitatively the characteristics of the average metachronal dynamics, and the emergence of defects. We systematically study the model's behaviour by assessing the effect of changing intrinsic rotor characteristics, including oscillator stiffness and the nature of their internal driving force, as well as their geometric properties and spatial arrangement. Our results suggest that metachronal coordination follows from deformations in the oscillators' limit cycles induced by hydrodynamic stresses, and that defects result from sufficiently steep local biases in the oscillators' intrinsic frequencies. Additionally, we find that random variations in the intrinsic rotor frequencies increase the robustness of the average properties of the emergent metachronal waves.

preprint2014arXiv

Flagellar Synchronization Through Direct Hydrodynamic Interactions

Microscale fluid flows generated by ensembles of beating eukaryotic flagella are crucial to fundamental processes such as development, motility and sensing. Despite significant experimental and theoretical progress, the underlying physical mechanisms behind this striking coordination remain unclear. Here, we present a novel series of experiments in which the flagellar dynamics of two micropipette-held somatic cells of Volvox carteri, with measurably different intrinsic beating frequencies, are studied by high-speed imaging as a function of their mutual separation and orientation. From analysis of beating time series we find that the interflagellar coupling, which is constrained by the lack of chemical and mechanical connections between the cells to be purely hydrodynamical, exhibits a spatial dependence that is consistent with theoretical predictions. At close spacings it produces robust synchrony which can prevail for thousands of flagellar beats, while at increasing separations this synchrony is systematically degraded by stochastic processes. Manipulation of the relative flagellar orientation reveals the existence of both in-phase and antiphase synchronized states, which is consistent with dynamical theories. Through dynamic flagellar tracking with exquisite spatio-temporal precision, we quantify changes in beating waveforms that result from altered coupling configuration and distance of separation. The experimental results of this study prove unequivocally that flagella coupled solely through a fluid medium can achieve robust synchrony despite significant differences in their intrinsic properties.

preprint2013arXiv

Antiphase Synchronization in a Flagellar-Dominance Mutant of Chlamydomonas

Groups of beating flagella or cilia often synchronize so that neighboring filaments have identical frequencies and phases. A prime example is provided by the unicellular biflagellate Chlamydomonas reinhardtii, which typically displays synchronous in-phase beating in a low-Reynolds number version of breaststroke swimming. We report here the discovery that ptx1, a flagellar dominance mutant of C. reinhardtii, can exhibit synchronization in precise antiphase, as in the freestyle swimming stroke. Long-duration high-speed imaging shows that ptx1 flagella switch stochastically between in-phase and antiphase states, and that the latter has a distinct waveform and significantly higher frequency, both of which are strikingly similar to those found during phase slips that stochastically interrupt in-phase beating of the wildtype. Possible mechanisms underlying these observations are discussed.

preprint2013arXiv

Ciliary contact interactions dominate surface scattering of swimming eukaryotes

Interactions between swimming cells and surfaces are essential to many microbiological processes, from bacterial biofilm formation to human fertilization. However, in spite of their fundamental importance, relatively little is known about the physical mechanisms that govern the scattering of flagellated or ciliated cells from solid surfaces. A more detailed understanding of these interactions promises not only new biological insights into structure and dynamics of flagella and cilia, but may also lead to new microfluidic techniques for controlling cell motility and microbial locomotion, with potential applications ranging from diagnostic tools to therapeutic protein synthesis and photosynthetic biofuel production. Due to fundamental differences in physiology and swimming strategies, it is an open question whether microfluidic transport and rectification schemes that have recently been demonstrated for pusher-type microswimmers such as bacteria and sperm cells, can be transferred to puller-type algae and other motile eukaryotes, as it is not known whether long-range hydrodynamic or short-range mechanical forces dominate the surface interactions of these microorganisms. Here, using high-speed microscopic imaging, we present direct experimental evidence that the surface scattering of both mammalian sperm cells and unicellular green algae is primarily governed by direct ciliary contact interactions. Building on this insight, we predict and verify experimentally the existence of optimal microfluidic ratchets that maximize rectification of initially uniform Chlamydomonas reinhardtii suspensions. Since mechano-elastic properties of cilia are conserved across eukaryotic species, we expect that our results apply to a wide range of swimming microorganisms.

preprint2011arXiv

Dynamic Critical Behavior of the Chayes-Machta Algorithm for the Random-Cluster Model. I. Two Dimensions

We study, via Monte Carlo simulation, the dynamic critical behavior of the Chayes-Machta dynamics for the Fortuin-Kasteleyn random-cluster model, which generalizes the Swendsen-Wang dynamics for the q-state Potts ferromagnet to non-integer q \ge 1. We consider spatial dimension d=2 and 1.25 \le q \le 4 in steps of 0.25, on lattices up to 1024^2, and obtain estimates for the dynamic critical exponent z_{CM}. We present evidence that when 1 \le q \lesssim 1.95 the Ossola-Sokal conjecture z_{CM} \ge β/νis violated, though we also present plausible fits compatible with this conjecture. We show that the Li-Sokal bound z_{CM} \ge α/νis close to being sharp over the entire range 1 \le q \le 4, but is probably non-sharp by a power. As a byproduct of our work, we also obtain evidence concerning the corrections to scaling in static observables.

preprint2010arXiv

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.