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8 Chemosensory Signal Transduction Systems in

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Abstract:

The majority of the analyses have focused on two-component signal transduction (TCST) systems, even though these are likely to compose only a fraction of the total systems that have evolved to translate information derived from the environment. The majority of all work on chemotaxis systems up until the last decade focused on the control of flagellum-based motility. is therefore an ideal organism for analysis of chemosensory signal transduction systems in bacteria. There are two primary reasons for our investigation into the role of the Che6 chemosensory system. First, the gene order is consistent with that found in other organisms known to utilize TFP-based motility. Second, a mutant allele known to suppress the mutant developmental defect was mapped to (suppressor of ), which we now know is cotranscribed as part of the operon. As does not possess phycobilisomes, Cpc7 is not likely to function as a phycocyanobilin lyase per se but might function in another light-dependent adaptation process. In summary, the chemosensory systems found in appear to have evolved in order to regulate functions that need temporal control mechanisms. In that regard may represent a case study encompassing both extremes where several systems regulate motility and several systems regulate alternative functions. Analysis of chemosensory signal transduction systems in each model organism is an excellent example of the modular nature of signal transduction and the evolution of bacterial genomes.

Citation: Kirby J, Berleman J, Müller S, Li D, Scott J, Wilson J. 2008. 8 Chemosensory Signal Transduction Systems in , p 135-147. In Whitworth D (ed), Myxobacteria. ASM Press, Washington, DC. doi: 10.1128/9781555815677.ch8

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Figures

Image of Figure 1
Figure 1

Domain topology of TCST systems. (A) A prototypical TCST system is shown. A variable input (sensor) domain is covalently bound to the histidine kinase domain. Phosphorylation occurs on a conserved histidine residue. The phosphoryl group is transferred to a conserved aspartate residue within the receiver domain (Rec) in the response regulator. The prototypical RR output is a DNA-binding domain capable of influencing gene expression. The vertical bar represents the cytoplasmic membrane. (B) The specialized TCST system that controls chemotaxis is shown. The MCP transducer is depicted as transmembrane and is coupled by CheW to the CheA kinase. Only two methyl groups are shown to represent methylation of the receptor by CheR. Phosphorylated CheB can remove these methyl groups. Methylation is a hallmark feature of the chemotaxis TCST systems. Phosphotransfer to the response regulator CheY influences its ability to bind the FliM switch component at the flagellar motor. (C) A chemosensory system such as the Che3 system found in is depicted. Chemosensory systems represent a composite of the prototypical TCST and the specialized chemotaxis TSCT systems.

Citation: Kirby J, Berleman J, Müller S, Li D, Scott J, Wilson J. 2008. 8 Chemosensory Signal Transduction Systems in , p 135-147. In Whitworth D (ed), Myxobacteria. ASM Press, Washington, DC. doi: 10.1128/9781555815677.ch8
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Image of Figure 2
Figure 2

Genetic organization of the eight chemosensory systems in . Each cluster or operon is defined by the existence of a gene encoding a homolog to CheA (black). Genes encoding homologs to CheW (dotted), MCPs (white), CheB (squares), CheR (circles), CheC (gray), and CheY-like response regulators (shaded) are shown. ORFs that do not display homology to any known chemotaxis gene are also shown (diagonal stripes).

Citation: Kirby J, Berleman J, Müller S, Li D, Scott J, Wilson J. 2008. 8 Chemosensory Signal Transduction Systems in , p 135-147. In Whitworth D (ed), Myxobacteria. ASM Press, Washington, DC. doi: 10.1128/9781555815677.ch8
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Image of Figure 3
Figure 3

The mutants display premature development on rich medium. Both the parent (DZ2) and mutant were grown at 32°C on rich medium (CYE) for 3 days and photographed (Kirby and Zusman, 2003); both colonies are approximately 2 cm in diameter. The mutant displays a rippling phenotype characteristic of development. Additionally, aggregates that resemble fruiting bodies are visible at the colony edge under higher magnification.

Citation: Kirby J, Berleman J, Müller S, Li D, Scott J, Wilson J. 2008. 8 Chemosensory Signal Transduction Systems in , p 135-147. In Whitworth D (ed), Myxobacteria. ASM Press, Washington, DC. doi: 10.1128/9781555815677.ch8
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Image of Figure 4
Figure 4

Model for Che3 signal transduction. The current model for Che3 chemosensory signal transduction is shown. CrdB is an outer membrane lipoprotein with a C-terminal OmpA-like peptidoglycan binding domain and is predicted to sense envelope stress. A conformational change in CrdB transmits a signal to the MCP receptor complex affecting CheA kinase levels in a manner similar to that observed during chemotaxis. CrdC is homologous to CheW and may affect coupling of the receptors to the CheA3 kinase. Results indicate that CrdA is autoregulatory and regulates expression of , , and other genes, thereby affecting development as described previously (Kirby and Zusman, 2003). The cytoplasmic and outer membranes (black lines) and the peptidoglycan layer (cross-hatching) demarcate the periplasm.

Citation: Kirby J, Berleman J, Müller S, Li D, Scott J, Wilson J. 2008. 8 Chemosensory Signal Transduction Systems in , p 135-147. In Whitworth D (ed), Myxobacteria. ASM Press, Washington, DC. doi: 10.1128/9781555815677.ch8
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Figure 5

The mutant displays defective motility and aggregation. The DZF1 parent and mutant are shown. Cells were grown in rich medium (CYE), washed in buffer (MMC), and plated on either 0.3% agar containing CYE or on 1.5% agar with low nutrients (CF). DZF1 cells swarm on rich medium (A) and develop on low-nutrient medium to produce fruiting bodies (B). The mutant cells display major defects in swarming (C) and development (D) relative to the parent.

Citation: Kirby J, Berleman J, Müller S, Li D, Scott J, Wilson J. 2008. 8 Chemosensory Signal Transduction Systems in , p 135-147. In Whitworth D (ed), Myxobacteria. ASM Press, Washington, DC. doi: 10.1128/9781555815677.ch8
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Tables

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Table 1

Chemosensory systems in

Citation: Kirby J, Berleman J, Müller S, Li D, Scott J, Wilson J. 2008. 8 Chemosensory Signal Transduction Systems in , p 135-147. In Whitworth D (ed), Myxobacteria. ASM Press, Washington, DC. doi: 10.1128/9781555815677.ch8

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