Polymerization of microtubules is ubiquitous in biological cells and under certain conditions it becomes oscillatory in time. Here simple reaction models are analyzed that capture such oscillations as well as the length distribution of microtubules. We assume reaction conditions that are stationary over many oscillation periods, and it is a Hopf bifurcation that leads to a persistent oscillatory microtubule polymerization in these models. Analytical expressions are derived for the threshold of the bifurcation and the oscillation frequency in terms of reaction rates as well as typical trends of their parameter dependence are presented. Both, a catastrophe rate that depends on the density of guanosine triphosphate (GTP) liganded tubulin dimers and a delay reaction, such as the depolymerization of shrinking microtubules or the decay of oligomers, support oscillations. For a tubulin dimer concentration below the threshold oscillatory microtubule polymerization occurs transiently on the route to a stationary state, as shown by numerical solutions of the model equations. Close to threshold a so-called amplitude equation is derived and it is shown that the bifurcation to microtubule oscillations is supercritical.
We investigate the response of two-dimensional pattern-forming systems with a broken up-down symmetry, such as chemical reactions, to spatially resonant forcing and propose related experiments. The nonlinear behavior immediately above threshold is analyzed in terms of amplitude equations suggested for a 1:2 and 1:1 ratio between the wavelength of the spatial periodic forcing and the wavelength of the pattern of the respective system. Both sets of coupled amplitude equations are derived by a perturbative method from the Lengyel-Epstein model describing a chemical reaction showing Turing patterns, which gives us the opportunity to relate the generic response scenarios to a specific pattern-forming system. The nonlinear competition between stripe patterns and distorted hexagons is explored and their range of existence, stability, and coexistence is determined. Whereas without modulations hexagonal patterns are always preferred near onset of pattern formation, single-mode solutions (stripes) are favored close to threshold for modulation amplitudes beyond some critical value. Hence distorted hexagons only occur in a finite range of the control parameter and their interval of existence shrinks to zero with increasing values of the modulation amplitude. Furthermore, depending on the modulation amplitude, the transition between stripes and distorted hexagons is either subcritical or supercritical.
The effects of a spatially periodic forcing on an oscillating chemical reaction as described by the Lengyel-Epstein model are investigated. We find a surprising competition between two oscillating patterns, where one is harmonic and the other subharmonic with respect to the spatially periodic forcing. The occurrence of a subharmonic pattern is remarkable as well as its preference up to rather large values of the modulation amplitude. For small modulation amplitudes we derive from the model system a generic equation for the envelope of the oscillating reaction that includes an additional forcing contribution, compared to the amplitude equations known from previous studies in other systems. The analysis of this amplitude equation allows the derivation of analytical expressions even for the forcing corrections to the threshold and to the oscillation frequency, which are in a wide range of parameters in good agreement with the numerical analysis of the complete reaction equations. In the nonlinear regime beyond threshold, the subharmonic solutions exist in a finite range of the control parameter that has been determined by solving the reaction equations numerically for various sets of parameters.
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