Person: Amir, Ariel
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Amir
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Ariel
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Amir, Ariel
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Publication Stochastic tunneling across fitness valleys can give rise to a logarithmic long-term fitness trajectory(American Association for the Advancement of Science (AAAS), 2019-07-05) Guo, Yipei; Vucelja, Marija; Amir, ArielPublication Mechanical strain-sensing implicated in cell shape recovery in Escherichia coli(2017) Wong, Felix; Renner, Lars D.; Özbaykal, Gizem; Paulose, Jayson; Weibel, Douglas B.; van Teeffelen, Sven; Amir, ArielThe shapes of most bacteria are imparted by the structures of their peptidoglycan cell walls, which are determined by many dynamic processes that can be described on various length-scales ranging from short-range glycan insertions to cellular-scale elasticity.1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11 Understanding the mechanisms that maintain stable, rod-like morphologies in certain bacteria has proved to be challenging due to an incomplete understanding of the feedback between growth and the elastic and geometric properties of the cell wall.3, 4, 12, 13, 14 Here we probe the effects of mechanical strain on cell shape by modeling the mechanical strains caused by bending and differential growth of the cell wall. We show that the spatial coupling of growth to regions of high mechanical strain can explain the plastic response of cells to bending4 and quantitatively predict the rate at which bent cells straighten. By growing filamentous E. coli cells in donut-shaped microchambers, we find that the cells recovered their straight, native rod-shaped morphologies when released from captivity at a rate consistent with the theoretical prediction. We then measure the localization of MreB, an actin homolog crucial to cell wall synthesis, inside confinement and during the straightening process and find that it cannot explain the plastic response to bending or the observed straightening rate. Our results implicate mechanical strain-sensing, implemented by components of the elongasome yet to be fully characterized, as an important component of robust shape regulation in E. coli.Publication Theory of Interacting Dislocations on Cylinders(American Physical Society, 2013) Amir, Ariel; Paulose, Jayson; Nelson, DavidWe study the mechanics and statistical physics of dislocations interacting on cylinders, motivated by the elongation of rod-shaped bacterial cell walls and cylindrical assemblies of colloidal particles subject to external stresses. The interaction energy and forces between dislocations are solved analytically, and analyzed asymptotically. The results of continuum elastic theory agree well with numerical simulations on finite lattices even for relatively small systems. Isolated dislocations on a cylinder act like grain boundaries. With colloidal crystals in mind, we show that saddle points are created by a Peach-Koehler force on the dislocations in the circumferential direction, causing dislocation pairs to unbind. The thermal nucleation rate of dislocation unbinding is calculated, for an arbitrary mobility tensor and external stress, including the case of a twist-induced Peach-Koehler force along the cylinder axis. Surprisingly rich phenomena arise for dislocations on cylinders, despite their vanishing Gaussian curvature.Publication Simultaneous regulation of cell size and chromosome replication in bacteria(Frontiers Media S.A., 2015) Ho, Po-Yi; Amir, ArielBacteria are able to maintain a narrow distribution of cell sizes by regulating the timing of cell divisions. In rich nutrient conditions, cells divide much faster than their chromosomes replicate. This implies that cells maintain multiple rounds of chromosome replication per cell division by regulating the timing of chromosome replications. Here, we show that both cell size and chromosome replication may be simultaneously regulated by the long-standing initiator accumulation strategy. The strategy proposes that initiators are produced in proportion to the volume increase and is accumulated at each origin of replication, and chromosome replication is initiated when a critical amount per origin has accumulated. We show that this model maps to the incremental model of size control, which was previously shown to reproduce experimentally observed correlations between various events in the cell cycle and explains the exponential dependence of cell size on the growth rate of the cell. Furthermore, we show that this model also leads to the efficient regulation of the timing of initiation and the number of origins consistent with existing experimental results.Publication Bending forces plastically deform growing bacterial cell walls(Proceedings of the National Academy of Sciences, 2014) Amir, Ariel; Babaeipour, F.; McIntosh, D. B.; Nelson, David; Jun, SuckjoonCell walls define a cell shape in bacteria. They are rigid to resist large internal pressures, but remarkably plastic to adapt to a wide range of external forces and geometric constraints. Currently, it is unknown how bacteria maintain their shape. In this work, we develop experimental and theoretical approaches and show that mechanical stresses regulate bacterial cell-wall growth. By applying a precisely controllable hydrodynamic force to growing rod-shaped Escherichia coli and Bacillus subtilis cells, we demonstrate that the cells can exhibit two fundamentally different modes of deformation. The cells behave like elastic rods when subjected to transient forces, but deform plastically when significant cell wall synthesis occurs while the force is applied. The deformed cells always recover their shape. The experimental results are in quantitative agreement with the predictions of the theory of dislocation-mediated growth. In particular, we find that a single dimensionless parameter, which depends on a combination of independently measured physical properties of the cell, can describe the cell's responses under various experimental conditions. These findings provide insight into how living cells robustly maintain their shape under varying physical environments.Publication Getting into shape: How do rod-like bacteria control their geometry?(Springer Science + Business Media, 2014) Amir, Ariel; van Teeffelen, SvenRod-like bacteria maintain their cylindrical shapes with remarkable precision during growth. However, they are also capable to adapt their shapes to external forces and constraints, for example by growing into narrow or curved confinements. Despite being one of the simplest morphologies, we are still far from a full understanding of how shape is robustly regulated, and how bacteria obtain their near-perfect cylindrical shapes with excellent precision. However, recent experimental and theoretical findings suggest that cell-wall geometry and mechanical stress play important roles in regulating cell shape in rod-like bacteria. We review our current understanding of the cell wall architecture and the growth dynamics, and discuss possible candidates for regulatory cues of shape regulation in the absence or presence of external constraints. Finally, we suggest further future experimental and theoretical directions, which may help to shed light on this fundamental problem.Publication Cell Size Regulation in Bacteria(American Physical Society (APS), 2014) Amir, ArielVarious bacteria such as the canonical gram negative Escherichia coli or the well-studied gram positive Bacillus subtilis divide symmetrically after they approximately double their volume. Their size at division is not constant, but is typically distributed over a narrow range. Here, we propose an analytically tractable model for cell size control, and calculate the cell size and interdivision time distributions, as well as the correlations between these variables. We suggest ways of extracting the model parameters from experimental data, and show that existing data for E. coli supports partial size control, and a particular explanation: a cell attempts to add a constant volume from the time of initiation of DNA replication to the next initiation event. This hypothesis accounts for the experimentally observed correlations between mother and daughter cells as well as the exponential dependence of size on growth rate.Publication Details Matter: Noise and Model Structure Set the Relationship between Cell Size and Cell Cycle Timing(Frontiers Media S.A., 2017) Barber, Felix; Ho, Po-Yi; Murray, Andrew; Amir, ArielOrganisms across all domains of life regulate the size of their cells. However, the means by which this is done is poorly understood. We study two abstracted “molecular” models for size regulation: inhibitor dilution and initiator accumulation. We apply the models to two settings: bacteria like Escherichia coli, that grow fully before they set a division plane and divide into two equally sized cells, and cells that form a bud early in the cell division cycle, confine new growth to that bud, and divide at the connection between that bud and the mother cell, like the budding yeast Saccharomyces cerevisiae. In budding cells, delaying cell division until buds reach the same size as their mother leads to very weak size control, with average cell size and standard deviation of cell size increasing over time and saturating up to 100-fold higher than those values for cells that divide when the bud is still substantially smaller than its mother. In budding yeast, both inhibitor dilution or initiator accumulation models are consistent with the observation that the daughters of diploid cells add a constant volume before they divide. This “adder” behavior has also been observed in bacteria. We find that in bacteria an inhibitor dilution model produces adder correlations that are not robust to noise in the timing of DNA replication initiation or in the timing from initiation of DNA replication to cell division (the C+D period). In contrast, in bacteria an initiator accumulation model yields robust adder correlations in the regime where noise in the timing of DNA replication initiation is much greater than noise in the C + D period, as reported previously (Ho and Amir, 2015). In bacteria, division into two equally sized cells does not broaden the size distribution.Publication Is cell size a spandrel?(eLife Sciences Publications, Ltd, 2017) Amir, ArielAll organisms control the size of their cells. We focus here on the question of size regulation in bacteria, and suggest that the quantitative laws governing cell size and its dependence on growth rate may arise as byproducts of a regulatory mechanism which evolved to support multiple DNA replication forks. In particular, we show that the increase of bacterial cell size during Lenski’s long-term evolution experiments is a natural outcome of this proposal. This suggests that, in the context of evolution, cell size may be a 'spandrel' DOI: http://dx.doi.org/10.7554/eLife.22186.001Publication Surprises in Numerical Expressions of Physical Constants(Mathematical Association of America, 2016) Amir, Ariel; Lemeshko, Mikhail; Tokieda, TadashiIn science, as in life, ‘surprises’ can be adequately appreciated only in the presence of a null model, what we expect a priori. In physics, theories sometimes express the values of dimensionless physical constants as combinations of mathematical constants like π or e. The inverse problem also arises, whereby the measured value of a physical constant admits a ‘surprisingly’ simple approximation in terms of well-known mathematical constants. Can we estimate the probability for this to be a mere coincidence, rather than an inkling of some theory? We answer the question in the most naive form.