Freedman N J, Lefkowitz R J
Howard Hughes Medical Research Institute, Department of Medicine (Cardiology), Duke University Medical Center, Durham, North Carolina 27710, USA.
Recent Prog Horm Res. 1996;51:319-51; discussion 352-3.
Waning responsiveness to continuous or repeated stimulation constitutes the phenomenon of desensitization, which pervades biological systems. Over the last several years, molecular mechanisms for desensitization of cellular signaling through G protein-coupled receptors have been delineated, particularly at the level of the receptors themselves. This review focuses on those aspects of G protein-coupled receptor desensitization which occur within minutes of agonist exposure. Agonist-dependent desensitization of these receptors can reduce their signaling responsiveness to maximum stimulation by up to 70-80%; indeed, in some receptor systems, the process of receptor desensitization appears to effect the termination of the cellular signaling response. Agonist-induced desensitization involves phosphorylation of G protein-coupled receptors by two currently recognized classes of serine/threonine protein kinases. Second messenger-dependent kinases, phosphorylating a variety of proteins, mediate a generalized cellular hyporesponsiveness termed heterologous desensitization. G protein-coupled receptor kinases (GRKs) phosphorylate specifically agonist-occupied, or activated, receptors, and thereby initiate receptor-specific, or homologous, desensitization. GRK-mediated receptor phosphorylation facilitates the binding of an inhibitory arrestin protein to the phosphorylated receptor, an event which substantially impairs receptor signaling. The GRK family comprises six, and the arrestin family comprises four known members. Each GRK demonstrates the ability to phosphorylate a limited number of model receptor substrates, but the widespread expression of several GRKs and the two somatic arrestins suggest that GRK-initiated homologous desensitization is of general importance to a wide range of G protein-coupled receptors. Exploration of the substrate specificity for GRKs and the ligand specificity for arrestins remains in its early stages. Currently, GRKs can most readily be differentiated by the mechanisms each employs to associate with the plasma membrane. Accumulating data from human disease states and transgenic mice attest to the physiologic significance of GRK-initiated receptor desensitization.
对持续或重复刺激的反应性减弱构成了脱敏现象,这一现象遍及生物系统。在过去几年中,已阐明了通过G蛋白偶联受体使细胞信号脱敏的分子机制,尤其是在受体自身水平。本综述聚焦于激动剂暴露数分钟内发生的G蛋白偶联受体脱敏的那些方面。这些受体的激动剂依赖性脱敏可使其对最大刺激的信号反应性降低多达70 - 80%;实际上,在某些受体系统中,受体脱敏过程似乎会导致细胞信号反应的终止。激动剂诱导的脱敏涉及目前已确认的两类丝氨酸/苏氨酸蛋白激酶对G蛋白偶联受体的磷酸化。第二信使依赖性激酶可磷酸化多种蛋白质,介导一种称为异源脱敏的普遍细胞低反应性。G蛋白偶联受体激酶(GRKs)特异性地磷酸化激动剂占据的或活化的受体,从而引发受体特异性或同源脱敏。GRK介导的受体磷酸化促进抑制性抑制蛋白与磷酸化受体的结合,这一事件会严重损害受体信号传导。GRK家族有六个成员,抑制蛋白家族有四个已知成员。每个GRK都显示出磷酸化有限数量的模型受体底物的能力,但几种GRK和两种体细胞抑制蛋白的广泛表达表明,GRK引发的同源脱敏对广泛的G蛋白偶联受体具有普遍重要性。对GRKs的底物特异性和抑制蛋白的配体特异性的探索仍处于早期阶段。目前,GRKs最容易通过各自与质膜结合的机制来区分。来自人类疾病状态和转基因小鼠的越来越多的数据证明了GRK引发的受体脱敏的生理意义。