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Chapter 12 : Cell-Interactive Sensing of the Environment

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Abstract:

can respond to nutrient limitation in either of two ways: (i) it can enter a stationary phase, in which cells grow very slowly, or (ii) it can stop growth and initiate the development of fruiting bodies with the eventual differentiation of spores. In and , stationary phase triggers differentiation into environmentally resistant rod cells, but this resistance is not manifest in despite the presence of a stationary phase and a stationary-phase sigma factor, encoded by . The time-ordered sequence of morphological and biochemical changes is coordinated by several signals passed between cells. One of these signals, A-factor, plays an important role in evaluating and responding to starvation. Purification of A-factor was based on an A-signal-dependent promoter. To identify A-factor, substances that would allow an A-signal-deficient mutant strain to express the Tn5 lacZ transcriptional fusion Ω4521 were sought from wild-type cells. Importantly, (p)ppGpp directly assesses the cell's capacity to synthesize any protein for which it has mRNA. There is as yet no evidence that other starvation-sensing pathways play a role in initiating development, even though the level of cyclic AMP rises twofold after starvation. The (p)ppGpp induces A-factor production, which requires . A-factor, being soluble, creates a pool outside the cells. Each cell then responds to the total amount of A-factor produced by all cells. The choice of response to depletion of nutrients is critical for cell survival.

Citation: Kaiser D. 2000. Cell-Interactive Sensing of the Environment, p 263-275. In Brun Y, Shimkets L (ed), Prokaryotic Development. ASM Press, Washington, DC. doi: 10.1128/9781555818166.ch12
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Figures

Image of FIGURE 1
FIGURE 1

Alternatives. Depending on the nutritional regime, the cells may respond to starvation by growing slowly (A) or they may develop a fruiting body (B).

Citation: Kaiser D. 2000. Cell-Interactive Sensing of the Environment, p 263-275. In Brun Y, Shimkets L (ed), Prokaryotic Development. ASM Press, Washington, DC. doi: 10.1128/9781555818166.ch12
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Image of FIGURE 2
FIGURE 2

Fruiting body development in M. xanthus. Development was initiated at 0 h by replacing nutrient medium with a buffer devoid of a usable carbon or nitrogen source. The lower right frame shows a fruiting body which has split open, revealing spores inside. This frame is three times the magnification of the others. (Scanning electron microscopy by J. Kuner. Reproduced from Kaiser et al., 1985.).

Citation: Kaiser D. 2000. Cell-Interactive Sensing of the Environment, p 263-275. In Brun Y, Shimkets L (ed), Prokaryotic Development. ASM Press, Washington, DC. doi: 10.1128/9781555818166.ch12
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Image of FIGURE 3
FIGURE 3

A-signal-defective developmental mutants of M, xanthus. The vertical arrows point to the times at which each of the lacZ fusions to a developmentally regulated promoter begins to be expressed. These fusions are reporters of normal development. The gray horizontal bar indicates the period during which there is normal expression of the reporter fusions in asg mutants. To the right of the bar, almost all development is defective (no, or gready reduced, reporter expression) because these reporters are asg dependent. An exception is Ω14469, which is expressed at 4 h but is asg independent. The columns to the right indicate the morphological phenotype with respect to aggregation and sporulation of the mutants.

Citation: Kaiser D. 2000. Cell-Interactive Sensing of the Environment, p 263-275. In Brun Y, Shimkets L (ed), Prokaryotic Development. ASM Press, Washington, DC. doi: 10.1128/9781555818166.ch12
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Image of FIGURE 4
FIGURE 4

Biosynthesis of pppGpp and ppGpp on ribosomes depends on the relA protein. Evidence for this scheme is summarized in Cashel et al., 1996.

Citation: Kaiser D. 2000. Cell-Interactive Sensing of the Environment, p 263-275. In Brun Y, Shimkets L (ed), Prokaryotic Development. ASM Press, Washington, DC. doi: 10.1128/9781555818166.ch12
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Image of FIGURE 5
FIGURE 5

Time course of (p)ppGpp and GTP levels in M. xanthus after the initiation of starvation by a shift from a nutrient medium rich in amino acids and peptides to a buffered salts medium devoid of any carbon or nitrogen source. (p)ppGpp increases rapidly after starvation.

Citation: Kaiser D. 2000. Cell-Interactive Sensing of the Environment, p 263-275. In Brun Y, Shimkets L (ed), Prokaryotic Development. ASM Press, Washington, DC. doi: 10.1128/9781555818166.ch12
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Image of FIGURE 6
FIGURE 6

The promoter for Ω4521 and the effects on transcription of changing individual bases. Those effects show that the promoter is recognized by sigma-54, encoded by rpoN. Boxes enclose the -24 hexamer and -12 pentamer, which are identical in the Ω4521 and mbhA promoters. Bases identical to the general sigma-54 consensus are shown in bold type. Arrows indicate the base changed in each mutant. The vertical position of the mutant form of the base indicates the percent wild type β-galactosidase activity in the mutant.

Citation: Kaiser D. 2000. Cell-Interactive Sensing of the Environment, p 263-275. In Brun Y, Shimkets L (ed), Prokaryotic Development. ASM Press, Washington, DC. doi: 10.1128/9781555818166.ch12
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Image of FIGURE 7
FIGURE 7

Ribosomes, sensing starvation, produce (p)ppGpp. (p)ppGpp initiates a stringent response, the production of A-factor, and expression of early developmentally regulated genes. A stringent response involves the arrest of the synthesis of ribosomes and the major polymers of the cell (Singer and Kaiser, 1995).

Citation: Kaiser D. 2000. Cell-Interactive Sensing of the Environment, p 263-275. In Brun Y, Shimkets L (ed), Prokaryotic Development. ASM Press, Washington, DC. doi: 10.1128/9781555818166.ch12
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FIGURE 8

A second battery of genes, exemplified by Ω4521, depends on starvation and extracellular A-factor. The first battery consists of starvation-dependent, but A-factor-independent, genes, such as the preaggregation genes shown in Fig. 7 . Sigma-54 promoters are receptive to at least two regulatory inputs, one via the sigma factor-polymerase holoenzyme and another via the required sigma-54 activator protein.

Citation: Kaiser D. 2000. Cell-Interactive Sensing of the Environment, p 263-275. In Brun Y, Shimkets L (ed), Prokaryotic Development. ASM Press, Washington, DC. doi: 10.1128/9781555818166.ch12
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Tables

Generic image for table
TABLE 1

A-factor activity of amino acids

Citation: Kaiser D. 2000. Cell-Interactive Sensing of the Environment, p 263-275. In Brun Y, Shimkets L (ed), Prokaryotic Development. ASM Press, Washington, DC. doi: 10.1128/9781555818166.ch12
Generic image for table
TABLE 2

A-factor production

Citation: Kaiser D. 2000. Cell-Interactive Sensing of the Environment, p 263-275. In Brun Y, Shimkets L (ed), Prokaryotic Development. ASM Press, Washington, DC. doi: 10.1128/9781555818166.ch12

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