Department of Physiology, University of Ulsan College of Medicine, Seoul, Korea.
J Physiol. 2012 Sep 15;590(18):4447-63. doi: 10.1113/jphysiol.2012.229203. Epub 2012 Jun 6.
L-type Ca(2+) channels (ICaLs) are inactivated by an increase in intracellular [Ca(2+)], known as Ca(2+)-dependent inactivation (CDI). CDI is also induced by Ca(2+) released from the sarcoplasmic reticulum (SR), known as release-dependent inhibition (RDI). As both CDI and RDI occur in the junctional subsarcolemmal nanospace (JSS), we investigated which factors are involved within the JSS using isolated cardiac myocytes from the main pulmonary vein of the rabbit. Using the whole-cell patch clamp technique, RDI was readily observed with the application of a pre-pulse followed by a test pulse, during which the ICaLs exhibited a decrease in peak current amplitude and a slower inactivation. A fast acting Ca(2+) chelator, 1,2-bis(o-aminophenoxy)ethane-N,N,N',N'-tetraacetic acid (BAPTA), abolished this effect. As the time interval between the pre-pulse and test pulse increased, the ICaLs exhibited greater recovery and the RDI was relieved. Inhibition of the ryanodine receptor (RyR) or the SR Ca(2+)-ATPase (SERCA) greatly attenuated RDI and facilitated ICaL recovery. Removal of extracellular Na(+),which inhibits the Na(+)-Ca(2+) exchange (Incx), greatly enhanced RDI and slowed ICaL recovery, suggesting that Incx critically controls the [Ca(2+)] in the JSS. We incorporated the Ca(2+)-binding kinetics of the ICaL into a previously published computational model. By assuming two Ca(2+)-binding sites in the ICaL, of which one is of low-affinity with fast kinetics and the other is of high-affinity with slower kinetics, the new model was able to successfully reproduce RDI and its regulation by Incx. The model suggests that Incx accelerates Ca(2+) removal from the JSS to downregulate CDI and attenuates SR Ca(2+) refilling. The model may be useful to elucidate complex mechanisms involved in excitation–contraction coupling in myocytes.
L 型钙通道 (ICaLs) 可被细胞内 [Ca2+] 的增加失活,这种失活称为 Ca2+依赖性失活 (CDI)。CDI 也可被肌浆网 (SR) 释放的 Ca2+诱导,这种失活称为释放依赖性抑制 (RDI)。由于 CDI 和 RDI 均发生在连接部的亚肌小节纳空间 (JSS) 内,我们使用来自兔主肺动脉的分离心肌细胞,研究了 JSS 内涉及的因素。使用全细胞膜片钳技术,应用预脉冲后施加测试脉冲,即可轻易观察到 RDI,在此期间 ICaLs 的峰值电流幅度减小,失活速度变慢。快速作用的 Ca2+螯合剂 1,2-双(邻氨基苯氧基)乙烷-N,N,N',N'-四乙酸 (BAPTA) 可消除这种作用。随着预脉冲和测试脉冲之间的时间间隔增加,ICaLs 表现出更大的恢复,RDI 得到缓解。抑制兰尼碱受体 (RyR) 或 SR Ca2+-ATP 酶 (SERCA) 可显著减弱 RDI 并促进 ICaL 恢复。去除细胞外 Na+,抑制 Na+-Ca2+交换 (Incx),可显著增强 RDI 并减缓 ICaL 恢复,表明 Incx 可严格控制 JSS 中的 [Ca2+]。我们将 ICaL 的 Ca2+结合动力学纳入先前发表的计算模型中。通过假设 ICaL 中有两个 Ca2+结合位点,其中一个具有低亲和力和快速动力学,另一个具有高亲和力和较慢动力学,新模型能够成功再现 RDI 及其由 Incx 调节。该模型表明,Incx 可加速从 JSS 中去除 Ca2+,下调 CDI,并减弱 SR Ca2+的再填充。该模型可有助于阐明心肌细胞兴奋-收缩耦联中涉及的复杂机制。