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Chapter 9 : Motility and Chemotaxis

Author: Linda L. McCarter1
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Affiliations: 1: Microbiology Department, The University of Iowa, Iowa City, IA 52242
Content Type: Monograph
Publication Year: 2006

Category: Environmental Microbiology

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Abstract:

To possess a functioning flagellar motility system, three things are requisite: a propeller, a motor, and a system of navigation. In this chapter, the swimming and swarming systems are considered separately. There is a central processing system that is shared by both flagellar systems, i.e., chemotaxis, and it allows the bacteria to detect signals in their environment and respond by modifying their movement. Various types of polar flagellar genes and polar flagellar and linked chemotaxis genes have been provided in the chapter. Chemotaxis integrates environmental signaling to modulate behavior by biasing movement toward more favorable conditions or away from unfavorable environments. Chemotaxis integrates environmental signaling to modulate behavior by biasing movement toward more favorable conditions or away from unfavorable environments. One consequence is that not just motility, but also chemotaxis is important for survival and colonization. In , a number of studies show differential expression of various motility and chemotaxis genes under in vivo and in vitro conditions. Bacterial chemotaxis has been most extensively studied in organisms with a few (~6 to 10) peritrichously arranged flagella, e.g., , serovar Typhimurium, and . In swimmer cells, the methyl-accepting chemotaxis proteins (MCPs) localize to both poles; in the swarmer cells, MCPs are found at the poles and at intervals along the cell body. The result is that polar flagellar function influences expression of cell surface polysaccharide, which has important consequences for biofilm formation and host colonization.

Citation: McCarter L. 2006. Motility and Chemotaxis, p 115-132. In Thompson F, Austin B, Swings J (ed), The Biology of Vibrios. ASM Press, Washington, DC. doi: 10.1128/9781555815714.ch9
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Motility and Chemotaxis
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Citation: McCarter L. 2006. Motility and Chemotaxis, p 115-132. In Thompson F, Austin B, Swings J (ed), The Biology of Vibrios. ASM Press, Washington, DC. doi: 10.1128/9781555815714.ch9
/content/10.1128/9781555815714.ch09.ch09fig01
FIGURE 1
Flagellation patterns of liquid- and plategrown The swimmer cells (left panel) possess single, sheathed polar flagella. This electron micrograph is of planktonically grown cells, stained with uranyl acetate. Cell proportions are ~1 × 2 μm. The swarmer cells (right panels) are elongated and display numerous lateral flagella in addition to the single, sheathed polar flagellum. swarmer cells were harvested from a plate and stained with phosphotungstic acid. The arrows point to the partially dissolved sheath of polar flagellum. The diameter of the sheathed polar flagellum is ~30 nm, and the diameter of the unsheathed lateral flagellum is ~15 nm.
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10.1128/9781555815714/f0117-01.gif
Image of FIGURE 1
FIGURE 1

Flagellation patterns of liquid- and plategrown The swimmer cells (left panel) possess single, sheathed polar flagella. This electron micrograph is of planktonically grown cells, stained with uranyl acetate. Cell proportions are ~1 × 2 μm. The swarmer cells (right panels) are elongated and display numerous lateral flagella in addition to the single, sheathed polar flagellum. swarmer cells were harvested from a plate and stained with phosphotungstic acid. The arrows point to the partially dissolved sheath of polar flagellum. The diameter of the sheathed polar flagellum is ~30 nm, and the diameter of the unsheathed lateral flagellum is ~15 nm.

Citation: McCarter L. 2006. Motility and Chemotaxis, p 115-132. In Thompson F, Austin B, Swings J (ed), The Biology of Vibrios. ASM Press, Washington, DC. doi: 10.1128/9781555815714.ch9
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FIGURE 2
Regulatory hierarchy and morphogenetic pathway for polar flagellar biogenesis. Flagellar assembly is an ordered process, initiating with integral membrane proteins forming a scaffold for the export and assembly of the remainder of the organelle and culminating with polymerization of the filament ( ). Gene expression is strictly regulated and tied into morphogenesis of the organelle (reviewed by Aldridge and Hughes, 2002). Early polar gene expression requires σ and late gene expression σ. The scheme of control seems conserved for the sp. in which it has been studied, and this figure, which depicts a general pathway for gene expression and organelle biogenesis, represents a summary of what is currently known (Kim and McCarter, 2000; Prouty et al., 2001; Millikan and Ruby, 2003, 2004). The master flagellar regulator FlrA (a.k.a. FlaK) directs transcription of class 2 genes, which encode the export and assembly apparatus and other regulators that act sequentially. FlhF and FlhG regulate flagellar number and placement by an unknown mechanism, but one that also influences transcription ( ). Expression of class 3 genes is controlled by the activity of a two-component regulator, FlrC (a.k.a. FlaM) and results in the assembly of the hook and basal body. Completion of the hook allows export of the anti-sigma factor FlgM, which frees the σ to direct transcription of class 4 genes (Correa et al., 2004).
10.1128/9781555815714/f0122-01_thmb.gif
10.1128/9781555815714/f0122-01.gif
Image of FIGURE 2
FIGURE 2

Regulatory hierarchy and morphogenetic pathway for polar flagellar biogenesis. Flagellar assembly is an ordered process, initiating with integral membrane proteins forming a scaffold for the export and assembly of the remainder of the organelle and culminating with polymerization of the filament ( ). Gene expression is strictly regulated and tied into morphogenesis of the organelle (reviewed by Aldridge and Hughes, 2002). Early polar gene expression requires σ and late gene expression σ. The scheme of control seems conserved for the sp. in which it has been studied, and this figure, which depicts a general pathway for gene expression and organelle biogenesis, represents a summary of what is currently known (Kim and McCarter, 2000; Prouty et al., 2001; Millikan and Ruby, 2003, 2004). The master flagellar regulator FlrA (a.k.a. FlaK) directs transcription of class 2 genes, which encode the export and assembly apparatus and other regulators that act sequentially. FlhF and FlhG regulate flagellar number and placement by an unknown mechanism, but one that also influences transcription ( ). Expression of class 3 genes is controlled by the activity of a two-component regulator, FlrC (a.k.a. FlaM) and results in the assembly of the hook and basal body. Completion of the hook allows export of the anti-sigma factor FlgM, which frees the σ to direct transcription of class 4 genes (Correa et al., 2004).

Citation: McCarter L. 2006. Motility and Chemotaxis, p 115-132. In Thompson F, Austin B, Swings J (ed), The Biology of Vibrios. ASM Press, Washington, DC. doi: 10.1128/9781555815714.ch9
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FIGURE 3
Regulatory hierarchy and morphogenetic pathway for lateral flagellar biogenesis. Although it is a distinct flagellar system, with its own complete set of flagellar genes, the lateral flagellar hierarchy of resembles the polar hierarchy in that a σ−dependent transcriptional regulator directs expression of intermediate genes, and σ controls late flagellar gene expression (Stewart and McCarter, 2003). The master lateral regulator LafK has also been shown to affect polar gene transcription (Kim and McCarter, 2004).
10.1128/9781555815714/f0125-01_thmb.gif
10.1128/9781555815714/f0125-01.gif
Image of FIGURE 3
FIGURE 3

Regulatory hierarchy and morphogenetic pathway for lateral flagellar biogenesis. Although it is a distinct flagellar system, with its own complete set of flagellar genes, the lateral flagellar hierarchy of resembles the polar hierarchy in that a σ−dependent transcriptional regulator directs expression of intermediate genes, and σ controls late flagellar gene expression (Stewart and McCarter, 2003). The master lateral regulator LafK has also been shown to affect polar gene transcription (Kim and McCarter, 2004).

Citation: McCarter L. 2006. Motility and Chemotaxis, p 115-132. In Thompson F, Austin B, Swings J (ed), The Biology of Vibrios. ASM Press, Washington, DC. doi: 10.1128/9781555815714.ch9
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Tables

Motility and Chemotaxis
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Citation: McCarter L. 2006. Motility and Chemotaxis, p 115-132. In Thompson F, Austin B, Swings J (ed), The Biology of Vibrios. ASM Press, Washington, DC. doi: 10.1128/9781555815714.ch9
/content/10.1128/9781555815714.ch09.ch09tab01
TABLE 1
Flagellation patterns of species
Generic image for table
TABLE 1

Flagellation patterns of species

Citation: McCarter L. 2006. Motility and Chemotaxis, p 115-132. In Thompson F, Austin B, Swings J (ed), The Biology of Vibrios. ASM Press, Washington, DC. doi: 10.1128/9781555815714.ch9
/content/10.1128/9781555815714.ch09.ch09tab02
TABLE 2
Polar flagellar and linked chemotaxis genes
Generic image for table
TABLE 2

Polar flagellar and linked chemotaxis genes

Citation: McCarter L. 2006. Motility and Chemotaxis, p 115-132. In Thompson F, Austin B, Swings J (ed), The Biology of Vibrios. ASM Press, Washington, DC. doi: 10.1128/9781555815714.ch9
/content/10.1128/9781555815714.ch09.ch09tab03
TABLE 3
lateral flagellar genes
Generic image for table
TABLE 3

lateral flagellar genes

Citation: McCarter L. 2006. Motility and Chemotaxis, p 115-132. In Thompson F, Austin B, Swings J (ed), The Biology of Vibrios. ASM Press, Washington, DC. doi: 10.1128/9781555815714.ch9
/content/10.1128/9781555815714.ch09.ch09tab04
TABLE 4
Multiple copies of potential chemotaxis genes are encoded in genomes
Generic image for table
TABLE 4

Multiple copies of potential chemotaxis genes are encoded in genomes

Citation: McCarter L. 2006. Motility and Chemotaxis, p 115-132. In Thompson F, Austin B, Swings J (ed), The Biology of Vibrios. ASM Press, Washington, DC. doi: 10.1128/9781555815714.ch9

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