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三、 肽链合成的终止

 瑛娜 2017-11-28

三、 肽链合成的终止。肽链合成的终止

包括两个阶段:肽链合成的终止反应

(termination reaction),即新合成的

肽链从最后一个tRNA的释放。终止后反应

(post-termination reaction)包括tRNA

的逐出和核糖体与mRNA的分离以及核糖

体大小亚基的分离。在已经测序的基因

中,代表C-末端氨基酸的密码子后面,总

有一个终止密码子。每一个基因中只要有

一个终止密码子就可以实现肽链合成的

终止。没有一种tRNA能识别终止密码子。

终止密码子是由蛋白质因子来识别,这与

其他密码子由tRNA来识别不同。大肠杆菌

三种终止密码子UAGUAAUGA。细菌

UAA使用频率最高,其次是UGAUAG

低。大肠杆菌有两种蛋白质因子催化终止

反应,称为释放因子(releasing factor

RF)。RF1识别UAAUAGRF2识别UGA

UAARF3可刺激RF1RF2的活性。RF作用

于核糖体的A位点,但必须在P位有肽酰

-tRNA时,才能识别A位点的终止密码子。

RF在细胞内的浓度远比起始因子和延伸

因子低,每个细胞内有600个左右,大约

相当于每10个核糖体有一个释放因子。

RF1 and RF2 recognize the termination

codons and activate the ribosome to

hydrolyze the peptidyl tRNA. Cleavage

of polypeptide from tRNA takes place

by a reaction analogous to the usual peptidyl

transfer, except that the acceptor is H2O instead of aminoacyl-tRNA.Then RF1 or RF2 is released from the ribosome by RF-3, which is a GTP-binding

protein related to EF-G . RF3 resembles the

GTP-binding domains of EF-Tu and EF-G, and RF1/2 resemble the C-terminal of EF-G, which mimics tRNA. This suggests that the action of RF3 on RF1/2

utilize the same site that is used by the

elongation factors. The termination reaction

involves release of the completed polypeptide, but leaves a deacylated tRNA and the mRNA still associated with the ribosome. The post-termination reaction involves the dissocation of the remaining components (tRNA, mRNA, 30S and 50S subunits) requires the factor RRF, ribosome recycling factor. Like the other factors involved in release, RRF has a structure that mimics tRNA, except that it lacks an equivalent

for the 3amino acid-binding region . IF-3 is also required in this reaction. RRF acts on the 50S subunit, and IF-3 acts to remove deacaylated

tRNA from the 30S subunit. Once the subunits have separated, IF-3 remains

necessary, of course, to prevent their reassociation.

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