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Faculty: Dr. Alvin Fox

Suggested reading: Murray, Third edition Chapter 3

KEY WORDS

KEY

WORDS

Cell envelope

Cell wall

Cell membrane

Outer membrane

Peptidoglycan

Braun lipoprotein

Porins

Lipopolysaccharide (endotoxin)

Teichoic acid

Teichuronic acid

Lipoteichoic acid

Mycolic acid

Undecaprenol (bactoprenol)

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

Cell

Envelope

The cell envelope may be defined defined as the

cell cell membrane membrane and and cell cell wall wall plus plus anan outer outer

membrane membrane ifif one one isis present present.

The cell

cell wall

wall consists of the peptidoglycan

peptidoglycan

layer and

layer

and attached

attached structures

structures.

Bacterial cell

Bacterial

cell envelopes

envelopes fall

fall

into

into two

two major

major categories:

categories:

Gram positive

Gram

positive and Gram

Gram negative

negative

Gram staining is based on characteristics that reflect major structural differences between the two groups.

Peptidoglycan

Peptidoglycan

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

Peptidoglycan consists

Peptidoglycan

consists of:

of:

  • 1. Glycan

Glycan (polysaccharide)

(polysaccharide) backbone

backbone consisting of

  • A. N-acetyl muramic acid (Mur)

  • B. N-acetyl glucosamine (Gln)

  • 2. Peptide

Peptide side

side chains

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

Gram Positive

Positive Cell

Cell Envelope

Envelope

Lipoteichoic

Gram Gram Positive Positive Cell Cell Envelope Envelope Lipoteichoic acid Peptidoglycan-teichoic acid r r r r

acid

Peptidoglycan-teichoic acid

r r r r r r r r r r
r
r
r
r
r
r
r
r
r
r

Cytoplasmic

membrane

Cytoplasm

Cytoplasm

Gram Positive

Gram

Positive Cell

Cell Envelope

Envelope

Covalently bound to the thick peptidoglycan are:

  • a. Teichoic Teichoic acid acid their backbones are usually phosphorus- containing polymers of ribitol or glycerol or

  • b. Teichuronic Teichuronic acid acid which are glucuronic acid- containing polysaccharides.

These negatively charged molecules are believed to be involved in concentrating metal ions from the surroundings.

Teichoic Teichoic acids acids can also direct direct autolytic autolytic enzymes enzymes to sites of peptidoglycan digestion (autolysis), one of the steps in cell wall biosynthesis.

Gram Negative

Gram

Negative Cell

Cell Envelope

Envelope

Outer Membrane
Outer Membrane
Gram Negative Gram Negative Cell Cell Envelope Envelope Outer Membrane Porin Lipopolysaccharide Braun lipoprotein Peptidoglycan r

Porin

Lipopolysaccharide

Braun lipoprotein
Braun lipoprotein
Gram Negative Gram Negative Cell Cell Envelope Envelope Outer Membrane Porin Lipopolysaccharide Braun lipoprotein Peptidoglycan r

Peptidoglycan

Gram Negative Gram Negative Cell Cell Envelope Envelope Outer Membrane Porin Lipopolysaccharide Braun lipoprotein Peptidoglycan r
Gram Negative Gram Negative Cell Cell Envelope Envelope Outer Membrane Porin Lipopolysaccharide Braun lipoprotein Peptidoglycan r
r
r
Gram Negative Gram Negative Cell Cell Envelope Envelope Outer Membrane Porin Lipopolysaccharide Braun lipoprotein Peptidoglycan r
Gram Negative Gram Negative Cell Cell Envelope Envelope Outer Membrane Porin Lipopolysaccharide Braun lipoprotein Peptidoglycan r
Gram Negative Gram Negative Cell Cell Envelope Envelope Outer Membrane Porin Lipopolysaccharide Braun lipoprotein Peptidoglycan r

Inner (cytoplasmic) membrane

Gram Negative Gram Negative Cell Cell Envelope Envelope Outer Membrane Porin Lipopolysaccharide Braun lipoprotein Peptidoglycan r
Gram Negative Gram Negative Cell Cell Envelope Envelope Outer Membrane Porin Lipopolysaccharide Braun lipoprotein Peptidoglycan r

Cytoplasm

Cytoplasm

Gram Negative

Gram

Negative Cell

Cell Envelope

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 ß 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 ofof Lipopolysaccharide

Structure

Lipopolysaccharide

of of Lipopolysaccharide Structure

Acid

Acid fast

fast and

and related

related bacteria

bacteria

(mycobacteria, nocardia

(mycobacteria,

nocardia and

and corynebacteria)

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

Synthesis ofof cell

cell envelope

envelope

macromolecules

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

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

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.