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Many bacterial cells secrete some extracellular material in the form of a capsule or a slime layer. A slime layer is loosely associated with the bacterium and can be easily washed off, whereas a capsule is attached tightly to the bacterium and has definite boundaries. Capsules can be seen under a light microscope by placing the cells in a suspension of India ink. The capsules exclude the ink and appear as clear halos surrounding the bacterial cells. Capsules are usually polymers of simple sugars (polysaccharides), although the capsule of Bacillus anthracis is made of polyglutamic acid. Most capsules are hydrophilic (“water-loving”) and may help the bacterium avoid desiccation (dehydration) by preventing water loss. Capsules can protect a bacterial cell from ingestion and destruction by white blood cells (phagocytosis). While the exact mechanism for escaping phagocytosis is unclear, it may occur because capsules make bacterial surface components more slippery, helping the bacterium to escape engulfment by phagocytic cells. The presence of a capsule in Streptococcus pneumoniae is the most important factor in its ability to cause pneumonia. Mutant strains of S. pneumoniae that have lost the ability to form a capsule are readily taken up by white blood cells and do not cause disease. The association of virulence and capsule formation is also found in many other species of bacteria.
A capsular layer of extracellular polysaccharide material can enclose many bacteria into a biofilm and serves many functions. Streptococcus mutans, which causes dental caries, splits the sucrose in food and uses one of the sugars to build its capsule, which sticks tightly to the tooth. The bacteria that are trapped in the capsule use the other sugar to fuel their metabolism and produce a strong acid (lactic acid) that attacks the tooth enamel. When Pseudomonas aeruginosa colonizes the lungs of persons with cystic fibrosis, it produces a thick capsular polymer of alginic acid that contributes to the difficulty of eradicating the bacterium. Bacteria of the genus Zoogloea secrete fibers of cellulose that enmesh the bacteria into a floc that floats on the surface of liquid and keeps the bacteria exposed to air, a requirement for the metabolism of this genus. A few rod-shaped bacteria, such as Sphaerotilus, secrete long chemically complex tubular sheaths that enclose substantial numbers of the bacteria. The sheaths of these and many other environmental bacteria can become encrusted with iron or manganese oxides.
Flagella, fimbriae, and pili
Many bacteria are motile, able to swim through a liquid medium or glide or swarm across a solid surface. Swimming and swarming bacteria possess flagella, which are the extracellular appendages needed for motility. Flagella are long, helical filaments made of a single type of protein and located either at the ends of rod-shaped cells, as in Vibrio cholerae or Pseudomonas aeruginosa, or all over the cell surface, as in Escherichia coli. Flagella can be found on both gram-positive and gram-negative rods but are rare on cocci and are trapped in the axial filament in the spirochetes. The flagellum is attached at its base to a basal body in the cell membrane. The protomotive force generated at the membrane is used to turn the flagellar filament, in the manner of a turbine driven by the flow of hydrogen ions through the basal body into the cell. When the flagella are rotating in a counterclockwise direction, the bacterial cell swims in a straight line; clockwise rotation results in swimming in the opposite direction or, if there is more than one flagellum per cell, in random tumbling. Chemotaxis allows a bacterium to adjust its swimming behavior so that it can sense and migrate toward increasing levels of an attractant chemical or away from a repellent one.
Not only are bacteria able to swim or glide toward more favorable environments, but they also have appendages that allow them to adhere to surfaces and keep from being washed away by flowing fluids. Some bacteria, such as E. coli and Neisseria gonorrhoeae, produce straight, rigid, spikelike projections called fimbriae (Latin for “threads” or “fibers”) or pili (Latin for “hairs”), which extend from the surface of the bacterium and attach to specific sugars on other cells—for these strains, intestinal or urinary-tract epithelial cells, respectively. Fimbriae are present only in gram-negative bacteria. Certain pili (called sex pili) are used to allow one bacterium to recognize and adhere to another in a process of sexual mating called conjugation (see below Bacterial reproduction). Many aquatic bacteria produce an acidic mucopolysaccharide holdfast, which allows them to adhere tightly to rocks or other surfaces.
The cytoplasm
Although bacteria differ substantially in their surface structures, their interior contents are quite similar and display relatively few structural features.
