The Motility of Mollicutes

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The Motility of Mollicutes Charles W. Wolgemuth, Oleg Igoshin, George Oster  Biophysical Journal  Volume 85, Issue 2, Pages 828-842 (August 2003) DOI: 10.1016/S0006-3495(03)74523-8 Copyright © 2003 The Biophysical Society Terms and Conditions

Figure 1 Application of the active helical filament model to the swimming of Spiroplasma. (a) The graph plots the displacement of the center of mass in the direction of the helix axis as a function of time, and shows the average propulsive velocity is positive and roughly constant. The insert shows frames from the movie of the flexing helix (for movie, see the Supplemental Material). (b) Plot of speed versus specific viscosity for Spiroplasma. The dots are data taken from Gilad et al. (2003), and the dashed line is 0.98(η/ηH2O)1/4. See Appendix C for details. Biophysical Journal 2003 85, 828-842DOI: (10.1016/S0006-3495(03)74523-8) Copyright © 2003 The Biophysical Society Terms and Conditions

Figure 2 Gliding motility of Mycoplasma mobile. (a) Schematic representation of propulsive and attachment organelle in Mycoplasma containing filament bundle. The filament bundle can elongate and shorten by bending or by longitudinal shortening, e.g., due to filament unbundling and entropic contraction. In either case, the projected length alternates between lengths L1 and L2, producing a step size of δ=L2−L1. (b) Schematic of the proposed mechanochemical cycle for the cell. The filament bundle attaches at the front and contracts, stretching the cell body. The elastically coupled cell body slides forward, consolidating the forward step. Finally, the nose detaches and extends, completing the cycle. The mathematical model given in Appendix E follows the sequence of events shown in the figure. Triggered by the periodic membrane potential, the mechanochemical deformation of the filament bundle can occur in several ways. (c–d) Load velocity curves for bending (c) and contraction (d) deformations for the no-slip (solid line) and back-slip (dashed line) models. The data points are from Miyata et al. (2002) for two temperatures. In both cases, the model extrapolates to the same temperature-independent stall force of ∼25–27pN. The details of the simulation and parameters used are given in Appendix E. Biophysical Journal 2003 85, 828-842DOI: (10.1016/S0006-3495(03)74523-8) Copyright © 2003 The Biophysical Society Terms and Conditions

Figure 3 Application to the swimming of Synechoccocus. (a) Cartoon showing how the internal bending wave is translated by the spicules into an external compression/amplitude wave. (b) Computed bending wave moving from left to right. The plot shows the conformation of the active filament at three time steps with chronological order (solid line, dotted line, and dashed line; for movie, see the Supplemental Material). Biophysical Journal 2003 85, 828-842DOI: (10.1016/S0006-3495(03)74523-8) Copyright © 2003 The Biophysical Society Terms and Conditions

Figure 4 Cartoon of possible mechanism of gliding A-motility of Myxococcus. A proposed propulsive cycle built on the ability of helically wound filaments to constrict and elongate the cell body ratcheted by slime. Slime extruded from posterior pores prevent the rear from moving backward, so that filament contraction pushes the front forward. Relaxation of the filaments permits the turgor pressure to restore the cell's original length and diameter. During this phase of the cycle, slime extrusion from the posterior pores pushes the rear forward rather than pulling the front end backward (Wolgemuth et al., 2002). Thus the role of slime extrusion is to prevent backward movement, and to consolidate leading end protrusion into a forward step. For the passive mechanism driven by turgor pressure oscillations, the filament angle must be greater than 54°; otherwise raising the turgor pressure will cause the cylinder to shorten and fatten. Biophysical Journal 2003 85, 828-842DOI: (10.1016/S0006-3495(03)74523-8) Copyright © 2003 The Biophysical Society Terms and Conditions

Figure 5 Minimizing elastic strain energy causes the elastic filament in Spiroplasma to assume a geodesic locus along the cell surface (figure redrawn from Trachtenberg, 1998). Biophysical Journal 2003 85, 828-842DOI: (10.1016/S0006-3495(03)74523-8) Copyright © 2003 The Biophysical Society Terms and Conditions

Figure 6 (a) Diagram of the experimental system. A bead is attached to the cell body and subject to a load force, Fload, from a fluid flow or laser trap. (b) The mechanical cycle of the model motor. (c) The phases of the mechanochemical cycle showing the relationship between the contracted/extended and detached/attached states. Biophysical Journal 2003 85, 828-842DOI: (10.1016/S0006-3495(03)74523-8) Copyright © 2003 The Biophysical Society Terms and Conditions

Figure 7 (a) Variables for the bending formula. (b) Projected length, Lp, versus dimensionless force, 2F˜/(Bκ2). Biophysical Journal 2003 85, 828-842DOI: (10.1016/S0006-3495(03)74523-8) Copyright © 2003 The Biophysical Society Terms and Conditions

Figure 8 A mechanochemical oscillator. (a) The (L, c) phase plane showing the shape of the nullclines and the limit cycle. (b) Enlargement of the limit cycle in the phase plane and (c) time plot. Biophysical Journal 2003 85, 828-842DOI: (10.1016/S0006-3495(03)74523-8) Copyright © 2003 The Biophysical Society Terms and Conditions