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Faculty: Dr.

Alvin Fox
Suggested reading: Murray, Third edition
Chapter 3

KEY WORDS
Cell envelope

Lipopolysaccharide (endotoxin)

Cell wall

Teichoic acid

Cell membrane

Teichuronic acid

Outer membrane

Lipoteichoic acid

Peptidoglycan

Mycolic acid

Braun lipoprotein

Undecaprenol (bactoprenol)

Porins

Endospore

Topics for discussion:


1. The structure of the Gram negative, Gram
positive and acid fast cell envelopes.
2. The composition and function of unique
cell envelope macromolecules and their
biosynthesis.
3. Endospores, which are unusual in many
ways.

Cell Envelope
The cell envelope may be defined as the
cell membrane and cell wall plus an outer
membrane if one is present.
present
The cell wall consists of the peptidoglycan
layer and attached structures.
structures

Bacterial cell envelopes fall


into two major categories:
Gram positive and Gram negative
Gram staining is based on characteristics that reflect
major structural differences between the two groups.

Peptidoglycan
A single bag-shaped, highly crosslinked macromolecule that surrounds
the bacterial cell membrane and
provides rigidity.

Peptidoglycan consists of:


1. Glycan (polysaccharide) backbone consisting of
A. N-acetyl muramic acid (Mur)
B. N-acetyl glucosamine (Gln)
2. Peptide side chains containing
A. D- and L- amino acids
B. Diaminopimelic acid (in some cases)
C. The side chains are cross-linked by peptide
bridges which vary in
structure among bacterial species.

Muramic acid, D-amino acids and


diaminopimelic acid are not
synthesized by mammals.

PG is found in all eubacteria except


Chlamydia and Mycoplasma.

Gram Positive Cell Envelope


Lipoteichoic
acid
r

Peptidoglycan-teichoic acid
r

r
r

Cytoplasmic membrane

Cytoplasm

r
r

r
r

Gram Positive Cell Envelope


Covalently bound to the thick peptidoglycan are:
a. Teichoic acid their backbones are usually phosphoruscontaining polymers of ribitol or glycerol or
b. Teichuronic acid which are glucuronic acidcontaining polysaccharides.
These negatively charged molecules are believed to be
involved in concentrating metal ions from the surroundings.
Teichoic acids can also direct autolytic enzymes to sites of
peptidoglycan digestion (autolysis), one of the steps in cell
wall biosynthesis.

Outer Membrane

Gram Negative Cell Envelope


Porin

Braun lipoprotein
r

Inner (cytoplasmic)
membrane

Cytoplasm

Lipopolysaccharide

Peptidoglycan

Gram Negative Cell Envelope


Covalently linked to the thin peptidoglycan is
a. Braun lipoprotein which binds the outer membrane to the
cell wall.
b. Proteins and phospholipids
c. Lipopolysaccharide which consists of three regions:
a. an outer O antigen,
b. a middle core which contains several sugars (heptoses
and ketodeoxyoctonic acid), not found elsewhere in
nature
c. an inner lipid A region which contains hydroxy fatty
acids (uncommon in nature). The molecule displays
endotoxin activity.
Porins in the outer membrane help form channels to allow passage of
small hydrophilic nutrients (such as sugars) through the outer membrane.

Structure of Lipopolysaccharide

Acid fast and related bacteria


(mycobacteria, nocardia and corynebacteria)
The cell envelopes of these organisms are considerably more
complex than other bacteria.
Mycolic acid (long, branch chained fatty acids) is covalently
bound via a polysaccharide to peptidoglycan.
Additional mycolic acid-containing compounds and other
complex lipids form a thick waxy membranous layer
outside the peptidoglycan layer.

Synthesis of cell envelope


macromolecules
Peptidoglycan:
1. The precursor subunit (muramyl pentapeptide
attached to uridine diphosphate, UDP) is synthesized
in the cytoplasm and passed to the cell membrane.
2. The subunit is moved enzymatically from the
nucleotide to a lipid carrier (undecaprenol/bactoprenol)
and built into a completed subunit (disaccharide
pentapeptide with attached bridge peptide).
3. The completed subunits are then exported to the cell
wall.

4. The undecaprenol is recirculated in the cell membrane


and used again.
5. The glycan backbones of the existing cell wall is
enzymatically broken (by autolysins) to allow insertion
of the newly synthesized subunit.
6. Cross-linking of the peptide side-chain of the inserted
subunit to the existing chain then occurs enzymatically
(penicillin binding proteins).
7. Completed subunits of teichoic and teichuronic acids
are also synthesized in the cell membrane (on lipid
carriers) before transport and insertion into the existing
cell wall.

Lipopolysaccharide
1. Lipid A is assembled in the cell membrane and the
core sugars attached sequentially.
2. O-antigen subunits are independently synthesized (on
a lipid carrier as in peptidoglycan synthesis).
3. The fully synthesized O-antigen is then attached to the
lipid A-core (generating lipopolysaccharide) in the cell
membrane before passage/insertion into the outer
membrane.

Endospores
1. These modified Gram positive bacterial cells have an
unusual cell envelope that contains a cell membrane and
an outer membrane.
2. The peptidoglycan layer is less cross-linked than in
most bacterial cells and contains a dehydrated form of
muramic acid.
3. The spore peptidoglycan is referred to as a cortex and
is found between the two membranes.

4. A coat consisting of highly cross-linked keratin is found


around the outside of the cell.
5. The bacterial spore is highly resistant to chemical
agents because of this coat.
6. When sporulation occurs, cell division is unequal and
the larger so-called "mother cell" envelops the daughter
cell.
7. The cell membrane of the daughter cell constitutes the
inner membrane of the spore and the cell membrane of
the mother forms the outer membrane.

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