No Arabic abstract
The eukaryotic flagellum beats with apparently unfailing periodicity, yet responds rapidly to stimuli. Like the human heartbeat, flagellar oscillations are now known to be noisy. Using the alga textit{C. reinhardtii}, we explore three aspects of nonuniform flagellar beating. We report the existence of rhythmicity, waveform noise peaking at transitions between power and recovery strokes, and fluctuations of interbeat intervals that are correlated and even recurrent, with memory extending to hundreds of beats. These features are altered qualitatively by physiological perturbations. Further, we quantify the recovery of periodic breastroke beating from transient hydrodynamic forcing. These results will help constrain microscopic theories on the origins and regulation of flagellar beating.
A quantitative description of the flagellar dynamics in the procyclic T. brucei is presented in terms of stationary oscillations and traveling waves. By using digital video microscopy to quantify the kinematics of trypanosome flagellar waveforms. A theoretical model is build starting from a Bernoulli-Euler flexural-torsional model of an elastic string with internal distribution of force and torque. The dynamics is internally driven by the action of the molecular motors along the string, which is proportional to the local shift and consequently to the local curvature. The model equation is a nonlinear partial differential wave equation of order four, containing nonlinear terms specific to the Korteweg-de Vries (KdV) equation and the modified-KdV equation. For different ranges of parameters we obtained kink-like solitons, breather solitons, and a new class of solutions constructed by smoothly piece-wise connected conic functions arcs (e.g. ellipse). The predicted amplitude and wavelengths are in good match with experiments. We also present the hypotheses for a step-wise kinematical model of swimming of procyclic African trypanosome.
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.
Cell migration in morphogenesis and cancer metastasis typically involves interplay between different cell types. We construct and study a minimal, one-dimensional model comprised of two different motile cells with each cell represented as an active elastic dimer. The interaction between the two cells via cadherins is modeled as a spring that can rupture beyond a threshold force as it undergoes dynamic loading via the attached motile cells. We obtain a phase diagram consisting of chase-and-run dynamics and clumping dynamics as a function of the stiffness of the interaction spring and the threshold force. We also find that while feedback between cadherins and cell-substrate interaction via integrins accentuates the chase-run behavior, feedback is not necessary for it.
We study the motion of oil drops propelled by actin polymerization in cell extracts. Drops deform and acquire a pear-like shape under the action of the elastic stresses exerted by the actin comet. We solve this free boundary problem and calculate the drop shape taking into account the elasticity of the actin gel and the variation of the polymerization velocity with normal stress. The pressure balance on the liquid drop imposes a zero propulsive force if gradients in surface tension or internal pressure are not taken into account. Quantitative parameters of actin polymerization are obtained by fitting theory to experiment.
Mechanical stress plays an intricate role in gene expression in individual cells and sculpting of developing tissues. However, systematic methods of studying how mechanical stress and feedback help to harmonize cellular activities within a tissue have yet to be developed. Motivated by our observation of the cellular constriction chains (CCCs) during the initial phase of ventral furrow formation in the Drosophila melanogaster embryo, we propose an active granular fluid (AGF) model that provides valuable insights into cellular coordination in the apical constriction process. In our model, cells are treated as circular particles connected by a predefined force network, and they undergo a random constriction process in which the particle constriction probability P is a function of the stress exerted on the particle by its neighbors. We find that when P favors tensile stress, constricted particles tend to form chain-like structures. In contrast, constricted particles tend to form compact clusters when P favors compression. A remarkable similarity of constricted-particle chains and CCCs observed in vivo provides indirect evidence that tensile-stress feedback coordinates the apical constriction activity. We expect that our particle-based AGF model will be useful in analyzing mechanical feedback effects in a wide variety of morphogenesis and organogenesis phenomena.