II. THE PROKARYOTIC CELL: BACTERIA
B. PROKARYOTIC CELL STRUCTURE
2a. The Gram-Positive Cell Wall
The overall purpose of this Learning Object is:
1) to learn the chemical makeup and functions associated with the cell wall of gram-positive bacteria;
2) to introduce the relationship between components of the gram-positive cell wall and the initiation of body defenses; and
3) to introduce the relationship between components of the gram-positive cell wall and bacterial pathogenicity.
LEARNING OBJECTIVES FOR THIS SECTION
In this section
on Prokaryotic Cell Structure we are looking at the various organelles or structures
that make up a bacterium. As mentioned in the introduction to this section,
a typical bacterium usually consists of:
We will now look at the gram-positive bacterial cell wall.
The Gram-Positive Cell Wall (def)
As mentioned in the previous section on peptidoglycan, gram-positive bacteria are those that retain the initial dye crystal violet during the Gram stain procedure and appear purple when observed through the microscope.
© Daniel Cavanaugh, Mark Keen, authors, Licensed for use, ASM MicrobeLibrary.Common gram-positive bacteria of medical importance include Streptococcus pyogenes, Streptococcus pneumoniae, Staphylococcus aureus, Enterococcus faecalis, and Clostridium species.
Highlighted Bacterium: Enterococcus speciesClick on this link, read the description of Enterococcus, and be able to match the bacterium with its description on an exam.
A. Structure and Composition of the Gram-Positive Cell Wall
1. In electron micrographs, the gram-positive cell wall appears as a broad, dense wall 20-80 nm thick and consisting of numerous interconnecting layers of peptidoglycan (see Figs. 1A and 1B). Chemically, 60 to 90% of the gram-positive cell wall is peptidoglycan. In gram-positive bacteria it is thought that the peptidoglycan is laid down in cables of several cross-linked glycan strands approximately 50 nm wide. These cables then themselves become cross-linked for further cell wall strength.
2. Interwoven in the cell wall of gram-positive are teichoic acids and lipoteichoic acids. Teichoic acids extend through and beyond the rest of the cell wall and are polyalcohols composed of polymers of glycerol, phosphates, and the sugar alcohol ribitol and are covalently bound to the peptidoglycan. Teichoic acids covalently bound to cytoplasmic membrane lipids are called lipoteichoic acids (see Fig. 1B).
3. The outer surface of the peptidoglycan is studded with surface proteins that differ with the strain and species of the bacterium (see Fig. 1B).
4. The periplasm is the gelatinous material between the peptidoglycan and the cytoplasmic membrane.
To view an electron micrograph of Streptococcus showing a gram-positive cell wall, see the Rockefeller University web page.
B. Functions of the Gram-Positive Cell Wall Components
1. The peptidoglycan in the gram-positive cell wall prevents osmotic lysis (def).
2. The teichoic acids probably help make the cell wall stronger (see Fig. 1B).
3. The surface proteins (see Fig. 1B) in the bacterial peptidoglycan, depending on the strain and species, carry out a variety of activities, including:
- Functioning as enzymes.
- Serving as adhesins (def). Adhesins enable the bacterium to adhere intimately to host calls and other surfaces in order to colonize and resist flushing (See Fig. 5 ).
- Occasionally functioning as invasins (def). Invasins enable some bacteria to penetrate host cells.
- Aiding certain bacteria in resisting phagocytic destruction.
The role of these cell wall surface proteins will be discussed in greater detail later in Unit 2 under Bacterial Pathogenicity.
4. The periplasm contains enzymes for nutrient breakdown.
C. Significance of Gram-Positive Cell Wall Components to the Initiation of Body Defenses
1. In order to protect against infection, one of the things the body must initially do is detect the presence of microorganisms. The body does this by recognizing molecules unique to microorganisms that are not associated with human cells. These unique molecules are called pathogen-associated molecular patterns or PAMPs (def). (Because all microbes, not just pathogenic microbes, possess PAMPs, pathogen-associated molecular patterns are sometimes referred to as microbe-associated molecular patterns or MAMPs.)
Fragments of peptidoglycan and teichoic acids are PAMPS associated with the cell wall of gram-positive bacteria. In addition, bacteria and other microorganisms also possess mannose-rich glycans (short carbohydrate chains with the sugar mannose or fructose as the terminal sugar) that function as PAMPs. These mannose-rich glycans are common in microbial glycoproteins and glycolipids but rare in those of humans (see Fig. 6).
These PAMPS bind to pattern-recognition receptors or PRRs (def) on a variety of defense cells of the body and trigger such innate immune defenses as inflammation (def), fever, and phagocytosis.
Inflammation (def) is the first response to infection and injury and is critical to body defense. Basically, the inflammatory response is an attempt by the body to restore and maintain homeostasis (def) after injury. Most of the body defense elements are located in the blood, and inflammation is the means by which body defense cells and body defense chemicals leave the blood and enter the tissue around an injured or infected site.
Body defense cells called macrophages (def), and dendritic cells (def) have pattern recognition receptors such as toll-like receptors on their surface that are specific for the peptidoglycan fragments and lipoteichoic acids in the gram-positive cell wall and/or to NODs in their cytoplasm that are specific for peptidoglycan fragments.
The binding of these cell wall components to their corresponding pattern recognition receptors triggers the macrophages to release various defense regulatory chemicals called cytokines (def), including IL-1, IL-6, IL-8, TNF-alpha, and PAF. The cytokines then bind to cytokine receptors on target cells and initiate inflammation and activate both the complement pathways (def) and the coagulation pathway (see Fig. 2).
2. The peptidoglycan and teichoic acids also activate the alternative complement pathway and the lectin pathway (def), defense pathways that play a variety of roles in body defense.
D. Significance of Gram-Positive Cell Wall Components to Bacterial Pathogenicity
During severe systemic infections with large numbers of bacteria present, however, high levels of gram-positive PAMPs are released resulting in excessive cytokine production by the macrophages and other cells and this, in turn, can harm the body (see Fig. 3).
E-Medicine article on infections associated with organisms mentioned in this Learning Object. Registration to access this website is free.
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