Bacterial cell wall

Membrane Lipid Diversity

 Archaea have the most extreme variations in
phospholipid side-chain structures.

- Ether links
between
glycerol and
fatty acids
- Hydrocarbon
chains are
branched
terpenoids.
Prokaryotic cell
 Cell Wall Functions
• Maintains shape of the bacterium
– almost all bacteria have one
• Helps protect cell from osmotic lysis
• Helps protect from toxic materials
• May contribute to pathogenicity
 Bacterial Cell Wall
• Composed of peptidoglycan
• Peptidoglycan (murein, sacculus)
– rigid structure that lies just outside the cell
plasma membrane
• Scientists describe two basic types of
bacterial cell walls:
– Gram-positive and Gram-negative based on
Gram staining
• Gram-positive: stain purple; thick peptidoglycan
• Gram-negative: stain pink or red; thin peptidoglycan
and outer membrane
 Bacterial Cell Walls: Peptidoglycan

• Structure of Peptidoglycan
• rigid layer that provides strength
• typically composed of
• alternating modified glucose (N-acetylglucosamine
and N-acetylmuramic acid) in β-1,4 linkages
• amino acids L-alanine, D-alanine, D-glutamic acid,
and either L-lysine or diaminopimelic acid (DAP)
• cross-linked differently in gram-negative bacteria and
gram-positive bacteria (often “interbridges”)
• can be destroyed by lysozyme (enzymes that cleave
glycosidic bond between sugars)
• found in human secretions, major defense against
bacterial infection
• 100+ distinct peptidoglycans have been described
 The Bacterial Cell Wall and Outer Layers

The cell wall is a single, interlinked

molecule that encloses the entire cell.
Comparison of the structures of glucose, NAG, and NAM.
 Peptidoglycan
• Alternating NAM and NAG molecules are linked in
rows of 10 to 65 sugars to form a carbohydrate
“backbone” (the glycan portion of
peptidoglycan) connected by β1→4 linkage.

• Peptidoglycan is unique to bacteria, although

archaea build analogous structures whose overall
physical nature is similar.

• The amount and exact composition of

peptidoglycan vary among the major bacterial
groups.
Possible structure of peptidoglycan.
 Linking of peptidoglycan
• Adjacent rows are linked by polypeptides (the peptide portion
of peptidoglycan).

• Although the structure of the polypeptide link varies, it always

includes tetrapeptide side chains, which consist of four amino
acids attached to NAMs in the backbone.

• The amino acids occur in an alternating pattern of D and

L forms. This is unique because the amino acids found in
other proteins are L forms.

• Parallel tetrapeptide side chains may be directly bonded to

each other or linked by a peptide cross-bridge, consisting of a
short chain of amino acids

• The NAM-NAG backbone is the same in all bacterial

species; the tetrapeptide side chains and the peptide
cross-bridges vary from species to species
 Peptides in peptidoglycan
• Several different amino acids are present in Cell
wall
• Three of these amino acids are not found in
proteins:
- D-glutamic acid
- D-alanine
- mesodiaminopimelic acid

• The presence of D-amino acids protects against

degradation by most peptidases, which recognize
only the L-isomers of amino acid residues
 Cell Envelope of Bacteria
 Most bacteria have additional envelope layers
that provide structural support and protection.
 Envelope composition defines:
- Gram-positive bacteria (thick cell wall)
- Example: the phylum Firmicutes

- Gram-negative bacteria (thin cell wall)

- Example: the phylum Proteobacteria

- Mycobacteria (complex, multilayered cell wall)

- Example: the phylum Actinomycetes
Cell Envelope: Gram-positive and Gram-negative
 Gram-Positive Cell
Walls
• Composed primarily of
peptidoglycan
• May also contain teichoic
acids (negatively charged)
– help maintain cell envelope
– protect from environmental
substances
– may bind to host cells
• some gram-positive bacteria
have layer of proteins on
surface
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of peptidoglycan
21
 Gram-Positive Cell Envelope
S-layer
- An additional protective layer commonly found in
free-living bacteria and archaea
- Crystalline layer of thick subunits consisting of
protein or glycoprotein
- May contribute to cell shape and help protect the cell
from osmotic stress
 Periplasm
• Depending on the organism, the periplasm can contain
several different classes of proteins.

• Hydrolytic enzymes, which function in the initial

degradation of food molecules

• Binding proteins, which begin the process of

transporting substrates

• Chemoreceptors, which are proteins that govern the

chemotaxis response.

• Most of these proteins reach the periplasm by way of a

protein-exporting system present in the cytoplasmic
membrane
 Periplasmic Space of
Gram-positive Bacteria
• Lies between plasma membrane and
cell wall and is smaller than that of
Gram-negative bacteria
• Periplasm has relatively few proteins
• Enzymes secreted by Gram-positive
bacteria are called exoenzymes
– aid in degradation of large nutrients
 Gram-Negative
Cell Walls
• More complex than Gram-
positive
• Consist of a thin layer of
peptidoglycan surrounded
by an outer membrane
• Outer membrane
composed of lipids,
lipoproteins, and
lipopolysaccharide (LPS)
• No teichoic acids
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 Gram-Negative Cell Walls
• Peptidoglycan is ~5-10% of cell wall
weight
• Periplasmic space differs from that in
Gram-positive cells
– may constitute 20–40% of cell volume
– many enzymes present in periplasm
• hydrolytic enzymes, transport proteins and
other proteins

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 Gram-Negative Cell
Walls

• outer membrane lies outside the thin

peptidoglycan layer 28
The Bacterial Cell Wall and Outer Layers

Lipoprotein and
Lipopolysaccharide
(LPS)
Murein lipoprotein: in
the inner leaflet of the
outer membrane that
forms a peptide bond with
peptidoglycan
LPS: in the outward-facing
leaflet of the outer
membrane
 Lipopolysaccharides
• Lipopolysaccharide (LPS), also called
Endotoxin, is an important part of the outer
membrane and can be used to identify
Gram-negative bacteria.

• It is an integral part of the cell wall and is

not released until the cell walls of dead
bacteria are broken down.
Lipopolysaccharide (LPS)
• Consists of three
parts
– lipid A
– core polysaccharide
– O side chain (O
antigen)
• Lipid A embedded in
outer membrane
• Core polysaccharide,
O side chain extend
out from the cell
 Structure of the lipopolysaccharide
of gram-negative Bacteria
• The chemistry of lipid A and the polysaccharide
components varies among species of gram-negative
Bacteria.
• Major components (lipid A–KDO–core–O-specific) are
typically the same. The O-specific polysaccharide is
highly variable among species.
• KDO, ketodeoxyoctonate; Hep, heptose; Glu, glucose;
Gal, galactose; GluNac, N-acetylglucosamine; GlcN,
glucosamine; P, phosphate. Glucosamine and the lipid A
fatty acids are linked through the amine groups.
 Lipid A
• The lipid A portion is responsible
for the toxic properties that make
any Gram-negative infection a
potentially serious medical
problem.

• It causes fever and dilates blood

vessels, so the blood pressure
drops precipitously.

• Because bacteria release

endotoxin mainly when they are
dying, killing them may increase
the concentration of this very toxic
substance.

• Thus, antibiotics given late in an

infection may cause a worsening
of symptoms, or even death of the
patient.
 Importance of LPS
• contributes to negative charge on cell
surface
• helps stabilize outer membrane structure
• may contribute to attachment to surfaces
and biofilm formation
• creates a permeability barrier
• protection from host defenses (O antigen)
• can act as an endotoxin (lipid A)

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