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The dynamics of stochastic attrition viewed as an absorption time on a terminating Markov chain.pdf

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The dynamics of stochastic attrition viewed as an absorption time on a terminating Markov chain.pdf

Cell Calcium 38 (2005) 73–86 The dynamics of stochastic attrition viewed as an absorption time on a terminating Markov chain Hilary DeRemigio, Gregory D. Smith ? Department of Applied Science, McGlothlin-Street Hall, Rm 305, The College of William and Mary, Williamsburg, VA 23187, USA Received 27 April 2005; received in revised form 31 May 2005; accepted 1 June 2005 Abstract Localized Ca2+ elevations known as Ca2+ puffs and sparks are cellular signals that arise from the cooperative activity of clusters of inositol 1,4,5-trisphosphate receptors and ryanodine receptors clustered at Ca2+ release sites on the surface of the endoplasmic reticulum or sarcoplasmic reticulum. When Markov chain models of these intracellular Ca2+-regulated Ca2+ channels are coupled via a mathematical representation of Ca2+ microdomain, simulated Ca2+ release sites may exhibit the phenomenon of “stochastic Ca2+ excitability” where the inositol 1,4,5-trisphosphate receptors (IP3Rs) or ryanodine receptors (RyRs) open and close in a concerted fashion. Interestingly, under some conditions simulated puffs and sparks can be observed even when the single-channel model used does not include slow Ca2+ inactivation or, indeed, any long-lived closed/refractory state [V. Nguyen, R. Mathias, G. Smith, Stochastic automata network descriptor for Markov chain models of instantaneously-coupled intracellular Ca2+ channels, Bull. Math. Biol. 67 (2005) 393–432]. In this case, termination of the localized Ca2+ elevation occurs when all of the intracellular channels at a release site simultaneously close through a process referred to as stochastic attrition [M. Stern, Theory of excitation-contraction coupling in cardiac muscle, Biophys. J. 63 (1992) 497–517]. In this paper, we investigate the statistical properties of stochastic attrition viewed as an absorption time on a terminating Markov chain that represents a Ca2+ release site composed of N two-state channels that are activated by Ca2+. Assuming that the loca