Bacterial Structure in Relationship to Pathogenicity (page 2)
(This chapter has 2 pages)
© 2008 Kenneth Todar, PhD
S-layers
are proteins in the
outermost cell envelope of a
broad range of bacteria. S-layers are composed of a
single
protein or glycoprotein species (mw 40-200 kDa) and exhibit either
oblique, square or hexagonal lattice symmetry with unit cell dimensions
in the range of 3 to 30 nm. S-layers are generally 5 to 10 nm thick and
show pores of identical size (diameter, 2 - 8 nm) and morphology.

Many
Gram-negative and Gram-positive bacteria, as well a many archaea,
possess a regularly structured layer called an S-layer attached to the
outermost portion of their cell wall. It is composed of protein or
glycoprotein and in electron micrographs, has a pattern resembling a
tiled surface. Transmission electron micrograph of a freeze-etched,
metal shadowed preparation of a bacterial cell with an S-layer with
hexagonal lattice
symmetry. Bar = 100nm.
http://www.foresight.org/conference/MNT7/Papers/Pum/index.html
S-layers have been associated with a number of possible functions that
relate to pathogenicity. S-layers can function as adhesins, enabling
the bacterium
to adhere to host cell membranes and tissue surfaces in order to
colonize. Many of the cell-associated protein adhesins used by
pathogens are components of the S-layer. The
S-layer may protect bacteria from harmful enzymes or changes in pH.
Like many other surface components, S-layers contribute to virulence by
protecting the
bacterium against complement and attack by phagocytes.
The cell wall of a bacterium is
an essential structure that protects the delicate cell protoplast from
osmotic lysis. The cell wall of Bacteria consists of a polymer of
disaccharides
cross-linked by short chains of amino acids (peptides). This molecule
is
a type of peptidoglycan called murein. Murein is unique
to
the Domain, Bacteria. In the Gram-positive bacteria,
the cell wall is thick
(15-80 nanometers), consisting of several layers of peptidoglycan
complexed
with molecules called teichoic acids. In the Gram-negative
bacteria,
the cell wall is relatively thin (10 nanometers) and is composed of a
single
layer of peptidoglycan surrounded by a membranous structure called the
outer
membrane.

The structure of the muramic
acid subunit in the peptidoglycan Escherichia. coli. The
molecule
consists of N-acetyl glucosamine (NAG) attached (via a beta 1,4 link)
to N-acetyl-muramic acid (NAM). Attached to the NAM is a peptide chain,
which (in the case of E. coli, as illustrated) consists of
L-alanine,
D-glutamate, diaminopimelic acid and D-alanine. Some antibiotics,
including
bacitracin, act by blocking the synthesis of the muramic acid subunit.
Penicillin and related antibiotics (beta lactams), as well as
vancomycin,
block the assembly of the muropeptide subunits into the peptidoglycan
polymer.
The cell wall, more properly the cell
envelope, is a complicated structure, fundamentally different in
Gram-positive and Gram-negative bacteria. Cell wall components are
major
determinants of virulence in both groups of bacteria. Endotoxin,
inherent to all Gram-negative bacteria, is toxic to animals in a
variety
of ways. Peptidoglycan and LPS, as well as some teichoic acids, induce
the alternate complement pathway leading to inflammation. Teichoic
acids
and O-specific polysaccharides may be used as adhesins by
Gram-positive
and Gram-negative bacteria, respectively. Some cell wall components
protect
against phagocytic engulfment or digestion. Variations in the
macromolecular
structure of cell wall components may be at the basis of antigenic
variation
as well as specific host resistance to pathogens.

E. coli 0157.
Transmission
electron micrograph (CDC). O157 refers to the antigenic type of E.
coli
which, in this case, is based on the precise molecular structure of the
O-specific polysaccharide in the cell wall LPS.
The essential outer membrane of Gram-negative bacteria is the target
for attack by complement, hydrophobic agents and certain antibiotics.
Murein
(peptidoglycan) is dismantled by a host enzyme, lysozyme, found in most
body fluids. Several antibiotics, mainly the beta lactams, exert their
antimicrobial effect by blocking the synthesis and assembly of
peptidoglycan.

Schematic drawing the outer
membrane of a Gram-negative bacterium.
The membranes of bacteria are
structurally
similar to the cell membranes of eucaryotes, except that bacterial
membranes
consist of saturated or monounsaturated fatty acids (rarely
polyunsaturated
fatty acids) and do not normally contain sterols. The plasma membrane
is
an exceptionally dynamic structure in bacteria which mediates
permeability,
transport, secretion and energy generation. In terms of pathogenesis of
a bacterium, it is often dependent upon the integrity and function
of its plasma membrane. The membrane might be responsible for secretion
of toxins, resistance to antimicrobial agents, tactic responses or
sensing
other environmental signals to turn on genes for virulence.
Endospores are bacterial
structures
(resting cells) formed by a few groups of bacteria as intracellular
structures,
but ultimately they are released as free endospores. Biologically,
endospores
are a fascinating type of cell. Endospores exhibit no signs of life,
being
described as cryptobiotic. They are highly resistant to environmental
stresses
such as high temperature (some endospores can be boiled for hours and
retain
their viability), irradiation, strong acids, disinfectants, etc. They
are thought to be the most durable cell produced in nature. Although
cryptobiotic,
they retain viability indefinitely, such that under appropriate
environmental
conditions, they germinate back into vegetative cells.
Endospores are formed mainly by two genera of Gram-positive
bacteria: Bacillus,
the aerobic sporeformers, and Clostridium, the anaerobic
sporeformers.
Both genera contain pathogens, and the endospores produced by these
bacteria
invariably play some role in the toxicity, transmission or survival of
the pathogen.

Spore stain of a Bacillus
species. (CDC). Mature spores stain green whether free or still
inside
the vegetative sporangium. Vegetative cells and sporangia stain red.
The
Schaeffer-Fulton stain technique was applied. The primary stain,
malachite
green, is forced into the spores by heating the prepared slide to
boiling
for 4-5 minutes. After washing, the vegetative cells are counterstained
with safranine.
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