Ginsburg K S, Weber C R, Bers D M
Department of Physiology, Loyola University Chicago, Stritch School of Medicine, Maywood, Illinois 60153, USA.
J Gen Physiol. 1998 Apr;111(4):491-504. doi: 10.1085/jgp.111.4.491.
In steady state, the Ca content of the sarcoplasmic reticulum (SR) of cardiac myocytes is determined by a balance among influx and efflux pathways. The SR Ca content may be limited mainly by the ATP-supplied chemical potential that is inherent in the gradient between SR and cytosol. That is, forward Ca pumping from cytosol to SR may be opposed by energetically conservative reverse pumping dependent on intra-SR free [Ca]. On the other hand, SR Ca loading may be limited by dissipative pathways (pump slippage and/or pump-independent leak). To assess how SR Ca content is limited, we loaded voltage-clamped ferret ventricular myocytes cumulatively with known amounts of Ca via L-type Ca channels (ICa), using Na-free solutions to prevent Na/Ca exchange. We then measured the maximal resulting caffeine-released SR Ca content under control conditions, as well as when SR Ca pumping was accelerated by isoproterenol (1 micro M) or slowed by thapsigargin (0.2-0.4 micro M). Under control conditions, SR Ca content reached a limit of 137 micro mol.liter cytosol-1 (nonmitochondrial volume) when measured by integrating caffeine-induced Na/Ca exchange currents lintegraINaCaXdt) and of 119 micro mol.liter cytosol-1 when measured using fluorescence signals dependent on changes in cytosolic free Ca ([Ca]i). When Ca-ATPase pumping rate was slowed 39% by thapsigargin, the maximal SR Ca content decreased by 5 (integralINaCaXdt method) or 23% (fluorescence method); when pumping rate was increased 74% by isoproterenol, SR Ca content increased by 10% (fluorescence method) or 20% (integralINaCaXdt method). The relative stability of the SR Ca load suggests that dissipative losses have only a minor influence in setting the SR Ca content. Indeed, it appears that the SR Ca pump in intact cells can generate a [Ca] gradient approaching the thermodynamic limit.
在稳态下,心肌细胞肌浆网(SR)的钙含量由流入和流出途径之间的平衡决定。SR钙含量可能主要受SR与胞质溶胶之间梯度所固有的ATP供应化学势的限制。也就是说,从胞质溶胶向SR的正向钙泵浦可能会受到依赖于SR内游离[Ca]的能量保守反向泵浦的对抗。另一方面,SR钙负载可能受耗散途径(泵浦滑动和/或非泵浦依赖性泄漏)的限制。为了评估SR钙含量是如何受到限制的,我们使用无钠溶液通过L型钙通道(ICa)向电压钳制的雪貂心室肌细胞累积加载已知量的钙,以防止钠/钙交换。然后,我们测量了在对照条件下以及当SR钙泵浦被异丙肾上腺素(1微摩尔)加速或被毒胡萝卜素(0.2 - 0.4微摩尔)减慢时咖啡因释放的SR钙含量的最大值。在对照条件下,通过积分咖啡因诱导的钠/钙交换电流(积分INaCaXdt)测量时,SR钙含量达到137微摩尔·升胞质溶胶 -1(非线粒体体积)的极限,而使用依赖于胞质游离钙([Ca]i)变化的荧光信号测量时为119微摩尔·升胞质溶胶 -1。当毒胡萝卜素使钙 - ATP酶泵浦速率减慢39%时,最大SR钙含量下降了5%(积分INaCaXdt方法)或23%(荧光方法);当异丙肾上腺素使泵浦速率增加74%时,SR钙含量增加了10%(荧光方法)或20%(积分INaCaXdt方法)。SR钙负载的相对稳定性表明耗散损失对设定SR钙含量的影响较小。实际上,完整细胞中的SR钙泵似乎能够产生接近热力学极限的[Ca]梯度。