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CHAPTER 27 PROKARYOTES AND THE ORIGINS OF METABOLIC DIVERSITY Section B1: The Structure, Function, and Reproduction of Prokaryotes 1. Nearly all prokaryotes have a cell wall external to the plasma membrane 2. Many prokaryotes are motile Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings Introduction • Most prokaryotes are unicellular. • Some species may aggregate transiently or form true colonies, even extending to division of labor between specialized cell types. • The most common shapes among prokaryotes are spheres (cocci), rods (bacilli), and helices. Fig. 27.3 Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings • Most prokaryotes have diameters in the range of 15 um, compared to 10-100 m for most eukaryotic cells. • However, the largest prokaryote discovered so far has a diameter of 0.75 mm. • It is a sulfur-metabolizing marine bacterium from coastal sediments off Namibia. Fig. 26.4 Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings 1. Nearly all prokaryotes have a cell wall external to the plasma membrane • In nearly all prokaryotes, a cell wall maintains the shape of the cell, affords physical protection, and prevents the cell from bursting in a hypotonic environment. • Most bacterial cell walls contain peptidoglycan, a polymer of modified sugars cross-linked by short polypeptides. • The walls of archaea lack peptidoglycan. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings • The Gram stain is a valuable tool for identifying specific bacteria, based on differences in their cell walls. • Gram-positive bacteria have simpler cell walls, with large amounts of peptidoglycans. Fig. 27.5a Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings • Gram-negative bacteria have more complex cell walls and less peptidoglycan. • An outer membrane on the cell wall contains lipopolysaccharides, carbohydrates bonded to lipids. Fig. 27.5b Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings • Among pathogenic bacteria, gram-negative species are generally more threatening than grampositive species. • The lipopolysaccharides on the walls are often toxic and the outer membrane protects the pathogens from the defenses of their hosts. • Gram-negative bacteria are commonly more resistant than gram-positive species to antibiotics because the outer membrane impedes entry of antibiotics. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings • Many antibiotics, including penicillins, inhibit the synthesis of cross-links in peptidoglycans, preventing the formation of a functional wall, particularly in gram-positive species. • These drugs are a very selective treatment because they cripple many species of bacteria without affecting humans and other eukaryotes, which do not synthesize peptidoglycans. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings • Many prokaryotes secrete another sticky protective layer, the capsule, outside the cell wall. • Capsules adhere the cells to their substratum. • They may increase resistance to host defenses. • They glue together the cells of those prokaryotes that live as colonies. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings • Another way for prokaryotes to adhere to one another or to the substratum is by surface appendages called pili. • Pili can fasten pathogenic bacteria to the mucous membranes of its host. • Some pili are specialized for holding two prokaryote cells together long enough to transfer DNA during conjugation. Fig. 27.6 Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings 2. Many prokaryotes are motile • About half of all prokaryotes are capable of directional movement. • The action of flagella, scattered over the entire surface or concentrated at one or both ends, is the most common method of movement. • The flagella of prokaryotes differ in structure and function from those of eukaryotes. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings • In a prokaryotic flagellum, chains of a globular protein wound in a tight spiral from a filament which is attached to another protein (the hook), and the basal apparatus. Fig. 27.7 • Rotation of the filament is driven by the diffusion of protons into the cell through the basal apparatus after the protons have been actively transported by proton pumps in the plasma membrane. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings • A second motility mechanism is found in spirochetes, helical bacteria. • Two or more helical filaments under the cell wall are attached to a basal motor attached to the cell. • When the filaments rotate, the cell moves like a corkscrew. • A third mechanism occurs in cells that secrete a jet of slimy threads that anchors the cells to the substratum. • The cell glides along at the growing end of threads. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings • In a relatively uniform environment, a flagellated cell may wander randomly. • In a heterogenous environment, many prokaryotes are capable of taxis, movement toward or away from a stimulus. • With chemotaxis, binding between receptor cells on the surface and specific substances results in movement toward the source (positive chemotaxis) or away (negative chemotaxis). • Other prokaryotes can detect the presence of light (phototaxis) or magnetic fields. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings