TwoComponent Regulatory System TwoComponent Regulatory SystemTCS twocomponent signal

第六章 细菌双组分调节系统 Two-Component Regulatory System

• 细菌双组分调节系统又叫细菌双组分系统（ Two-Component Regulatory System，TCS ）双组分信号传 导系统（two-component signal transduction systems，TCSTS）

细胞内的双组分体系调节的反应 nutrient acquisition onitrogen ophosphorus ocarbon energy metabolism oelectron transport systems ouptake and catabolic machinery virulence oplasmid transfer odegradative secretions otoxin production oadherence factors adaptation to physical or chemical aspects of the environment op. H oosmolarity olight quality complex developmental pathways osporulation ofruiting body development oswarmer cell production

A single cell may have many two-component systems E. coli 178 信号蛋白中, 62 参与了双组分系统 26 histidine kinases (PSI-BLAST search of E. coli genome for signal transmitters) 36 response regulators (PSI-BLAST search of E. coli genome for signal receivers) in 10 sub-families (families & functions) 整个信号转导过程只由两个组分(HPK和RR)即可完成，是一典型 的、最简单的TCSTS。需要强调的是，随着研究的广泛深入，发 现TCSTS并非想象中的那么简单，在一些条件下，可能不止一个 HPK或RR参与了一种信号的转导；而在一个菌细胞中也可能有多 种二元系统存在，来感应或转导不同的信息。

受体复合体主要由细胞膜上的4个可甲基化趋化蛋白（methyl-accepting chemotaxis proteins or MCPs）, 组氨酸激酶Che. A、连接蛋白Che. W组成. 当诱导物结合时促进从顺时针（ clockwise signalling [CWS] ）向逆时针转变（counter-clockwise signalling [CCWS]）

Figure 1. Example of a basic two-component system. The E. coli osmolarity-response system consists of an HPK osmosensor (Env. Z) and an RR transcription factor (Omp. R) (Pratt and Silhavy, 1995). Env. Z autophosphorylates using ATP as the phosphate donor. The phosphate from the transmitter module of Env. Z is then transferred to an Asp residue in the receiver module of Omp. R, thereby affecting the promoter interactions of the Omp. R DNAbinding module, which regulates the transcription of two porin genes, omp. F and omp. C. Changes in osmolarity are perceived by the amino-terminal module of Env. Z. In response to such changes, Env. Z changes the level of phosphorylated Omp. R. The dotted lines depict intra-protein regulatory interactions. The dashed line depicts phosphorylation/dephosphorylation events. P, Phosphoryl group; H, His; D, Asp.

Regulation of Translation by Antisense RNA

Fig. B 3

The initiation of sporulation is governed in part by the activities of two spatially separated sigma factors

磷酸盐吸收 The activation of sigma F is accomplished through a phosphorelay system

The adenylylation/deadenylylation cycle that controls the activity of glutamine synthetase. 腺苷脱腺苷控制谷氨酰胺合成酶（GS）活性。氮信号被腺（尿）苷酰转移酶（ATase或 UTase）识别。低氮激活UTase，尿苷酰化的PII协助ATase对GS-AMP 去腺苷酰化。