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Echinocyte Shapes: Bending, Stretching and Shear Determine Spicule Shape and Spacing

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 Added by Ranjan Mukhopadhyay
 Publication date 2001
  fields Physics
and research's language is English




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We study the shapes of human red blood cells using continuum mechanics. In particular, we model the crenated, echinocytic shapes and show how they may arise from a competition between the bending energy of the plasma membrane and the stretching/shear elastic energies of the membrane skeleton. In contrast to earlier work, we calculate spicule shapes exactly by solving the equations of continuum mechanics subject to appropriate boundary conditions. A simple scaling analysis of this competition reveals an elastic length which sets the length scale for the spicules and is, thus, related to the number of spicules experimentally observed on the fully developed echinocyte.



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One of the most widely used methods for determination of the bending elasticity modulus of model lipid membranes is the analysis of the shape fluctuations of nearly spherical lipid vesicles. The theoretical basis of this analysis is given by Milner and Safran. In their theory the stretching effects are not considered. In the present study we generalized their approach including the stretching effects deduced after an application of statistical mechanics of vesicles.
A thin-walled tube, e.g., a drinking straw, manifests an instability when bent by localizing the curvature change in a small region. This instability has been extensively studied since the seminal work of Brazier nearly a century ago. However, the scenario of pressurized tubes has received much less attention. Motivated by rod-shaped bacteria such as E. coli, whose cell walls are much thinner than their radius and are subject to a substantial internal pressure, we study, theoretically, how this instability is affected by this internal pressure. In the parameter range relevant to the bacteria, we find that the internal pressure significantly postpones the onset of the instability, while the bending stiffness of the cell wall has almost no influence. This study suggests a new method to infer turgor pressure in rod-shaped bacteria from bending experiments.
Theoretical studies of nearly spherical vesicles and microemulsion droplets, that present typical examples for thermally-excited systems that are subject to constraints, are reviewed. We consider the shape fluctuations of such systems constrained by fixed area $A$ and fixed volume $V$, whose geometry is presented in terms of scalar spherical harmonics. These constraints can be incorporated in the theory in different ways. After an introductory review of the two approaches: with an exactly fixed by delta-function membrane area $A$ [Seifert, Z. Phys. B, 97, 299, (1995)] or approximatively by means of a Lagrange multiplier $sigma$ conjugated to $A$ [Milner and Safran, Phys. Rev. A, 36, 4371 (1987)], we discuss the determined role of the stretching effects, that has been announced in the framework of a model containing stretching energy term, expressed via the membrane vesicle tension [Bivas and Tonchev, Phys.Rev.E, 100, 022416 (2019)]. Since the fluctuation spectrum for the used Hamiltonian is not exactly solvable an approximating method based on the Bogoliubov inequalities for the free energy has been developed. The area constraint in the last approach appears as a self-consistent equation for the membrane tension. In the general case this equation is intractable analytically. However, much insight into the physics behind can be obtained either imposing some restrictions on the values of the model parameters, or studying limiting cases, in which the self-consistent equation is solved. Implications for the equivalence of ensembles have been discussed as well.
We report a theoretical study of DNA flexibility and quantitatively predict the ring closure probability as a function of DNA contour length. Recent experimental studies show that the flexibility of short DNA fragments (as compared to the persistence length of DNA l_P~150 base pairs) cannot be described by the traditional worm-like chain (WLC) model, e.g., the observed ring closure probability is much higher than predicted. To explain these observations, DNA flexibility is investigated with explicit considerations of a new length scale l_D~10 base pairs, over which DNA local bend angles are correlated. In this correlated worm-like chain (C-WLC) model, a finite length correction term is analytically derived and the persistence length is found to be contour length dependent. While our model reduces to the traditional worm-like chain model when treating long DNA at length scales much larger than l_P, it predicts that DNA becomes much more flexible at shorter sizes, which helps explain recent cyclization measurements of short DNA fragments around 100 base pairs.
Soft bodies flowing in a channel often exhibit parachute-like shapes usually attributed to an increase of hydrodynamic constraint (viscous stress and/or confinement). We show that the presence of a fluid membrane leads to the reverse phenomenon and build a phase diagram of shapes --- which are classified as bullet, croissant and parachute --- in channels of varying aspect ratio. Unexpectedly, shapes are relatively wider in the narrowest direction of the channel. We highlight the role of flow patterns on the membrane in this response to the asymmetry of stress distribution.
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