Calcium signal transmission by axonemal microtubules as an optimized information pathway in cilia and flagella.

Calcium plays a key role in signal transduction in eukaryotic cells. Besides controlling local functions of cells calcium ions are responsible for the generation of global signals such as waves and spikes. Pulsatile increases of calcium concentrations are generally considered to have a much higher fidelity of information transfer than simple tonic changes, since they are much less prone to noisy fluctuations. In that respect, it was clearly revealed that Ca2+ has very crucial involvement in many signaling pathways in cilia and flagella. We earlier established a model in which axonemal microtubules exhibit the features of nonlinear polyelectrolitic electric transmissions lines for efficient transport of cations, primarily Ca2+. These microtubules guide accumulated "ionic clods" which serve as the pulsatile signals aimed to regulate pertaining motor proteins, dyneins and kinesis. We here consider such Ca2+ signals in axoneme in the context of Shannon's and Fisher's information theories. It appears that the fast drift of these "ionic clouds" represents the optimized calcium signaling for control of "flagellary beats" as well as intraflagellary transport of proteins essential for the construction, elongation and maintenance of eukaryotic cilia and flagella themselves.

Calcium signaling modulates the dynamics of cilia and flagella.

To adapt to changing environments cells must signal and signaling requires messengers whose concentration varies with time in space. We here consider the messenger role of calcium ions implicated in regulation of the wave-like bending dynamics of cilia and flagella. The emphasis is on microtubules as polyelectrolytes serving as transmission lines for the flow of Ca2+ signals in the axoneme. This signaling is superimposed with a geometric clutch mechanism for the regulation of flagella bending dynamics and our modeling produces results in agreement with experimental data.

Nonlinear ionic pulses along microtubules.

Microtubules are cylindrically shaped cytoskeletal biopolymers that are essential for cell motility, cell division and intracellular trafficking. Here, we investigate their polyelectrolyte character that plays a very important role in ionic transport throughout the intra-cellular environment. The model we propose demonstrates an essentially nonlinear behavior of ionic currents which are guided by microtubules. These features are primarily due to the dynamics of tubulin C-terminal tails which are extended out of the surface of the microtubule cylinder. We also demonstrate that the origin of nonlinearity stems from the nonlinear capacitance of each tubulin dimer. This brings about conditions required for the creation and propagation of solitonic ionic waves along the microtubule axis. We conclude that a microtubule plays the role of a biological nonlinear transmission line for ionic currents. These currents might be of particular significance in cell division and possibly also in cognitive processes taking place in nerve cells.