Previous mutagenesis studies with bacteriorhodopsin have shown that reprotonation of the Schiff's base is the rate-limiting step in the photocycle of the D96N mutant, whereas retinal re-isomerization and return of the protein to the initial state constitute the rate-limiting events in the photocycle of the L93A mutant. Thus, in the D96N mutant, decay of the M intermediate is slowed down by more than 100-fold at pH 7. In the L93A mutant, decay of the O intermediate is slowed down by 250-fold. We report here that in the L93A, D96N double mutant, decay of the M intermediate, as well as the formation and decay of the O intermediate, are slowed down dramatically. The photocycle is completed by the decay of a long-lived O intermediate, as in the L93A mutant. The decay of the M and O intermediates in the double mutant parallels the behavior seen in the single mutants over a wide temperature and pH range, arguing that the observed independence is an intrinsic property of the mutant. The slow decay of the M and O intermediates can be selectively and independently reversed under conditions identical to those used for the corresponding intermediates in the D96N and L93A single mutants. Because the effects of the two individual mutations are preserved in the double mutant and can be independently reversed, we conclude that residues Asp 96 and Leu 93 act independently and at different stages of the bacteriorhodopsin photocycle. These results also show that formation of the O intermediate only requires protonation of the Schiff's base and is independent of the protonation of Asp 96 from the aqueous medium.

译文

先前对细菌视紫红质的诱变研究表明,席夫碱的再质子化是D96N突变体光循环中的限速步骤,而视网膜再异构化和蛋白质返回到初始状态则构成了光循环中的限速事件。L93A突变体。因此,在D96N突变体中,M中间体的衰变在ph7下减慢了100倍以上。在L93A突变体中,O中间体的衰变减慢250倍。我们在这里报告说,在L93A,D96N双突变体中,M中间体的衰变以及O中间体的形成和衰变显着减慢。像L93A突变体一样,通过长寿命O中间体的衰变来完成光循环。双突变体中M和O中间体的衰变与在宽温度和pH范围内单个突变体中看到的行为相似,认为观察到的独立性是突变体的内在特性。M和O中间体的缓慢衰变可以在与D96N和L93A单个突变体中相应中间体所使用的条件相同的条件下选择性地和独立地逆转。由于两个个体突变的作用保留在双突变体中并且可以独立逆转,因此我们得出结论,残基Asp 96和Leu 93独立且在细菌视紫红质光循环的不同阶段起作用。这些结果还表明,O中间体的形成仅需要席夫碱的质子化,并且与水性介质中Asp 96的质子化无关。

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