Xu H, Dixon J E, Barry D M, Trimmer J S, Merlie J P, McKinnon D, Nerbonne J M
Department of Molecular Biology and Pharmacology, Washington University, School of Medicine, St. Louis, Missouri 63110, USA.
J Gen Physiol. 1996 Nov;108(5):405-19. doi: 10.1085/jgp.108.5.405.
In the experiments here, the developmental expression of the functional Ca(2+)-independent, depolarization-activated K+ channel currents, Ito and IK, and of the voltage-gated K+ channel (Kv) alpha subunits, Kv1.2, Kv1.4, Kv1.5, Kv2.1, and Kv4.2 in rat ventricular myocytes were examined quantitatively. Using the whole-cell patch clamp recording method, the properties and the densities of Ito and IK in ventricular myocytes isolated from postnatal day 5 (P5), 10 (P10), 15 (P15), 20 (P20), 25 (P25), 30 (P30), and adult (8-12 wk) rats were characterized and compared. These experiments revealed that mean Ito densities increase fourfold between birth and P30, whereas IK densities vary only slightly. Neither the time- nor the voltage-dependent properties of the currents vary measurably, suggesting that the subunits underlying functional Ito and IK channels are the same throughout postnatal development. In parallel experiments, the developmental expression of each of the voltage-gated K+ channel alpha subunits, Kv1.2, Kv1.4, Kv1.5, Kv2.1, and Kv4.2, was examined quantitatively at the mRNA and protein levels using subunit-specific probes. RNase protection assays revealed that Kv1.4 message levels are high at birth, increase between P0 and P10, and subsequently decrease to very low levels in adult rat ventricles. The decrease in message is accompanied by a marked reduction in Kv1.4 protein, consistent with our previous suggestion that Kv1.4 does not contribute to the formation of functional K+ channels in adult rat ventricular myocytes. In contrast to Kv1.4, the mRNA levels of Kv1.2, Kv1.5, Kv2.1, and Kv4.2 increase (three- to five-fold) between birth and adult. Western analyses, however, revealed that the expression patterns of these subunits proteins vary in distinct ways: Kv1.2 and Kv4.2, for example, increase between P5 and adult, whereas Kv1.5 remains constant and Kv2.1 decreases. Throughout development, therefore, there is a mismatch between the numbers of Kv alpha subunits expressed and the functional voltage-gated K+ channel currents distinguished electrophysiologically in rat ventricular myocytes. Alternative experimental approaches will be required to define directly the Kv alpha subunits that underlie functional voltage-gated K+ channels in these (and other) cells. In addition, the finding that Kv alpha subunit protein expression levels do not necessarily mirror mRNA levels suggests that caution should be exercised in attempting functional interpretations of observed changes in mRNA levels alone.
在本实验中,对大鼠心室肌细胞中功能性钙(2+)非依赖性、去极化激活钾离子通道电流(Ito和IK)以及电压门控钾离子通道(Kv)α亚基Kv1.2、Kv1.4、Kv1.5、Kv2.1和Kv4.2的发育表达进行了定量检测。采用全细胞膜片钳记录方法,对出生后第5天(P5)、10天(P10)、15天(P15)、20天(P20)、25天(P25)、30天(P30)和成年(8 - 12周)大鼠分离的心室肌细胞中Ito和IK的特性及密度进行了表征和比较。这些实验表明,出生至P30期间,Ito的平均密度增加了四倍,而IK密度仅略有变化。电流的时间依赖性和电压依赖性特性均无明显变化,这表明在出生后的整个发育过程中,构成功能性Ito和IK通道的亚基是相同的。在平行实验中,使用亚基特异性探针在mRNA和蛋白质水平上对每个电压门控钾离子通道α亚基Kv1.2、Kv1.4、Kv1.5、Kv2.1和Kv4.2的发育表达进行了定量检测。核糖核酸酶保护分析显示,Kv1.4的信使核糖核酸水平在出生时较高,在P0至P10期间升高,随后在成年大鼠心室中降至极低水平。信使核糖核酸水平的下降伴随着Kv1.4蛋白的显著减少,这与我们之前的观点一致,即Kv1.4对成年大鼠心室肌细胞中功能性钾离子通道的形成没有贡献。与Kv1.4相反,Kv1.2、Kv1.5、Kv2.1和Kv4.2的mRNA水平在出生至成年期间增加了三至五倍。然而,蛋白质免疫印迹分析显示,这些亚基蛋白的表达模式以不同方式变化:例如,Kv1.2和Kv4.2在P5至成年期间增加,而Kv1.5保持不变,Kv2.1则减少。因此,在整个发育过程中,大鼠心室肌细胞中表达的Kvα亚基数与通过电生理学区分的功能性电压门控钾离子通道电流之间存在不匹配。需要采用其他实验方法来直接确定这些(以及其他)细胞中构成功能性电压门控钾离子通道的Kvα亚基。此外,Kvα亚基蛋白表达水平不一定反映mRNA水平这一发现表明,在仅根据观察到的mRNA水平变化进行功能解释时应谨慎行事。