Hollingworth Stephen, Chandler W Knox, Baylor Stephen M
Department of Physiology, University of Pennsylvania School of Medicine, Philadelphia, 19104, USA.
J Gen Physiol. 2006 Mar;127(3):291-307. doi: 10.1085/jgp.200509477.
The properties of Ca(2+) sparks in frog intact skeletal muscle fibers depolarized with 13 mM [K(+)] Ringer's are well described by a computational model with a Ca(2+) source flux of amplitude 2.5 pA (units of current) and duration 4.6 ms (18 degrees C; Model 2 of Baylor et al., 2002). This result, in combination with the values of single-channel Ca(2+) current reported for ryanodine receptors (RyRs) in bilayers under physiological ion conditions, 0.5 pA (Kettlun et al., 2003) to 2 pA (Tinker et al., 1993), suggests that 1-5 RyR Ca(2+) release channels open during a voltage-activated Ca(2+) spark in an intact fiber. To distinguish between one and greater than one channel per spark, sparks were measured in 8 mM [K(+)] Ringer's in the absence and presence of tetracaine, an inhibitor of RyR channel openings in bilayers. The most prominent effect of 75-100 microM tetracaine was an approximately sixfold reduction in spark frequency. The remaining sparks showed significant reductions in the mean values of peak amplitude, decay time constant, full duration at half maximum (FDHM), full width at half maximum (FWHM), and mass, but not in the mean value of rise time. Spark properties in tetracaine were simulated with an updated spark model that differed in minor ways from our previous model. The simulations show that (a) the properties of sparks in tetracaine are those expected if tetracaine reduces the number of active RyR Ca(2+) channels per spark, and (b) the single-channel Ca(2+) current of an RyR channel is <or=1.2 pA under physiological conditions. The results support the conclusion that some normal voltage-activated sparks (i.e., in the absence of tetracaine) are produced by two or more active RyR Ca(2+) channels. The question of how the activation of multiple RyRs is coordinated is discussed.
用含13 mM [K⁺]的林格氏液使青蛙完整骨骼肌纤维去极化后,其Ca²⁺火花的特性可以用一个计算模型很好地描述,该模型中Ca²⁺源通量的幅度为2.5 pA(电流单位),持续时间为4.6 ms(18℃;贝勒等人2002年的模型2)。这一结果,结合在生理离子条件下双层膜中报道的ryanodine受体(RyRs)单通道Ca²⁺电流值,0.5 pA(凯特伦等人,2003年)至2 pA(廷克等人,1993年),表明在完整纤维中电压激活的Ca²⁺火花期间,有1 - 5个RyR Ca²⁺释放通道打开。为了区分每个火花是一个还是多个通道,在含8 mM [K⁺]的林格氏液中,在不存在和存在丁卡因(双层膜中RyR通道开放的抑制剂)的情况下测量火花。75 - 100 μM丁卡因最显著的作用是使火花频率降低约六倍。其余的火花在峰值幅度、衰减时间常数、半高全宽(FDHM)、半高全宽(FWHM)和质量的平均值上有显著降低,但上升时间的平均值没有降低。用一个与我们之前的模型有微小差异的更新的火花模型模拟了丁卡因存在时的火花特性。模拟结果表明:(a)如果丁卡因减少每个火花中活性RyR Ca²⁺通道的数量,那么丁卡因存在时火花的特性是预期的;(b)在生理条件下,RyR通道的单通道Ca²⁺电流≤1.2 pA。结果支持这样的结论:一些正常的电压激活火花(即在不存在丁卡因的情况下)是由两个或更多活性RyR Ca²⁺通道产生的。文中还讨论了多个RyRs的激活是如何协调的问题。