StepWise USMLE
Microbiology · Bacteriology

Bacterial structure · pathogenesis · pharmacology

Cell envelope, antibiotic targets, spores, growth, culture, virulence strategies and toxins

🧱 Bacterial Cell Envelope

Layers common to all bacteria

  • Cytoplasmic (inner) membrane
    • Carries cell-wall-building enzymes
    • Transpeptidases and carboxypeptidases = penicillin-binding proteins (PBPs)
    • PBPs are the targets of β-lactam drugs
  • Peptidoglycan
    • Rigid mesh that resists osmotic lysis
    • Thickness explains Gram stain behavior

Gram-positive envelope

  • Very thick peptidoglycan
  • Retains crystal violet, stains purple
  • No outer membrane
  • Teichoic acids (in wall)
  • Lipoteichoic acid (anchored in membrane)
    • Both help adhesion to host surfaces
  • Surface proteins vary by genus
    • Example: M protein of Streptococcus pyogenes
  • Large peptidoglycan–teichoic acid fragments
    • Act as structural toxins
    • Attract neutrophils

Gram-negative envelope

  • Thin peptidoglycan in the periplasmic space
  • Loses crystal violet, counterstains pink
  • Outer membrane covers the wall
    • Porins let small molecules through
    • Lipoprotein anchors outer membrane to peptidoglycan
  • Outer leaflet is lipopolysaccharide (LPS)
    • Lipid A: embedded, the toxic part
    • Core polysaccharide
    • O antigen: variable chain, used for serotyping
  • Periplasm holds enzymes such as β-lactamases

What LPS does in the host

Lipid A binds TLR-4→Macrophage activation→TNF-α, IL-1, IL-6 release→Fever, vasodilation, shock risk
  • Also activates B cells without T-cell help (T-independent)

Comparison

FeatureGram-positiveGram-negative
PeptidoglycanThickThin
Outer membraneAbsentPresent (LPS)
EndotoxinNoYes (lipid A)
Teichoic acidsPresentAbsent
Periplasmic spaceMinimalProminent
Main adhesion toolTeichoic acidsPili (fimbriae)
Concept
  • Carriers harbor a potential pathogen without overt disease.

🛠️ Peptidoglycan Synthesis and Antibiotic Targets

Polymer structure

  • Backbone: alternating NAG (N-acetylglucosamine) and NAM (N-acetylmuramic acid)
  • Linked by β-1,4 bonds; lysozyme cuts this bond
  • Peptide stem hangs from each NAM
    • Begins as a pentapeptide
    • Ends in D-Ala–D-Ala
    • Third residue carries a free amine (L-lysine or diaminopimelic acid)
  • Cross-link formed between
    • Amine on residue 3 of one stem
    • D-Ala at position 4 of a neighboring stem
  • Terminal D-Ala is released during cross-linking

Synthesis steps

1 Cytoplasm: make NAG, NAM, add pentapeptide
2 Membrane: unit attached to bactoprenol carrier
3 NAG added to complete the disaccharide-peptide unit
4 Flipped across the membrane
5 Transglycosylase adds unit to growing chain
6 Transpeptidase cross-links stems (PBP)
7 Carrier recycled: pyrophosphate-carrier to phosphate-carrier
  • Carboxypeptidases trim terminal D-Ala residues
Why it is a good drug target
  • Peptidoglycan exists only in bacteria, so human cells are largely unaffected.

Cell-wall inhibitors

DrugStep blockedResistance
BacitracinCarrier recycling (dephosphorylation of bactoprenol pyrophosphate)Mostly used topically
β-lactamsBind PBPs, block transpeptidation
  • Altered PBP with lower affinity
  • β-lactamase destroys the drug
  • Reduced drug entry through porins (gram-negatives)
VancomycinBinds D-Ala–D-Ala, blocks transglycosylation and transpeptidationVanA and VanB change the terminus (D-Ala–D-Lac), so drug cannot bind

Antibiotic targets across the cell

TargetDrugs
Cell wall synthesisβ-lactams, vancomycin, bacitracin, polymyxin, daptomycin, isoniazid, ethambutol, ethionamide, cycloserine
30S ribosomeAminoglycosides, tetracyclines, tigecycline
50S ribosomeMacrolides, clindamycin, chloramphenicol, linezolid, quinupristin-dalfopristin, telithromycin
DNA replication or integrityFluoroquinolones, metronidazole, clofazimine
RNA synthesisRifampin, rifabutin
Folate pathway (antimetabolites)Sulfonamides, trimethoprim, dapsone, para-aminosalicylic acid
Exam trap
  • Tetracyclines bind the 30S subunit, whereas macrolides bind the 50S subunit.
  • Antibiotics work best on rapidly dividing cells in log phase.

📈 Endospores, Growth and Culture

Endospores

  • Formed by Bacillus and Clostridium (gram-positive rods)
  • Purpose: survival, not reproduction
    • One cell makes one spore
  • Trigger: nutrient depletion
  • Germination: warmth, moisture, nutrients
  • Withstand heat, drying, radiation, freezing, chemicals
  • Resistance features
    • Dehydrated core rich in calcium dipicolinate
    • Protective keratin-like protein coat
    • Thick, less cross-linked cortex of peptidoglycan
    • Heat-resistant enzymes
  • Core holds DNA, ribosomes, glycolytic enzymes
  • Autoclaving (steam under pressure) is needed to kill them
Vegetative cell→Starvation→Spore→Favorable conditions→Vegetative cell
  • Fungal spores differ: they serve reproduction

Growth curve phases

PhaseWhat happens
Lag
  • Cells adapt, no net increase
  • Induce enzymes for the medium
  • Cell number at start equals end
Log (exponential)
  • Binary fission, doubling at fixed intervals
  • Generation time measured here
  • Most susceptible to antibiotics
Stationary
  • Nutrients exhausted, wastes accumulate
  • New cells equal dying cells
  • Toxins and spores often produced
Death
  • Deaths exceed new cells
  • Log-scale decline
  • Binary fission: 1 → 2 → 4 → 8 cells

Culture principles

  • Obligate intracellular organisms cannot grow on artificial media
    • Viruses, rickettsiae, chlamydiae
    • Need cell culture, eggs or animals
  • Some organisms cannot be cultured at all, e.g., syphilis spirochete
  • Selective medium: favors certain organisms with nutrients or antibiotics
  • Differential medium: separates organisms by colony color or appearance

Special media

OrganismMediumType
AnaerobesThioglycolate brothOxygen-depleting
CorynebacteriumLöffler serum; tellurite agarSelective; differential
Enteric gram-negativesMacConkey; eosin methylene blueSelective and differential; differential
Enteric pathogens (Salmonella, Shigella)Hektoen; xylose-lysine-deoxycholateDifferential
Vibrio choleraeTCBS (alkaline)Selective
LegionellaCharcoal yeast extract with cysteineSelective
MycobacteriumLöwenstein-JensenSelective
Neisseria (sterile sites), HaemophilusChocolate agarEnriched
Neisseria (nonsterile sites)Thayer-Martin: chocolate agar plus vancomycin, colistin, nystatinSelective

Special growth requirements

RequirementOrganism
Cholesterol, purines, pyrimidinesMycoplasma
CysteineFrancisella, Brucella, Legionella, Pasteurella
X factor (hemin) and V factor (NAD)Haemophilus influenzae and H. aegyptius need both

🎯 Virulence Strategies

1. Colonization

  • Adhesion to host surfaces
    • Pili or fimbriae: main tool in gram-negatives
    • Teichoic acids: main tool in gram-positives
    • Adhesins: colonization factors, pertussis toxin, hemagglutinins
  • Biofilm formation on inert surfaces
    • Staphylococcus epidermidis (catheters, prostheses)
    • Streptococcus mutans (teeth)
    • Pseudomonas aeruginosa (lungs, catheters)

2. Avoiding phagocytosis and mucosal defense

  • Polysaccharide capsules
    • Streptococcus pneumoniae
    • Klebsiella pneumoniae
    • Haemophilus influenzae type b
    • Pseudomonas aeruginosa
    • Neisseria meningitidis
    • Cryptococcus neoformans (yeast)
  • Surface proteins that block uptake
    • S. pyogenes: M protein
    • S. aureus: protein A (binds Fc of IgG)
    • N. gonorrhoeae: pili
  • IgA protease cleaves secretory IgA
    • Neisseria, Haemophilus, S. pneumoniae

3. Acquiring nutrients

  • Siderophores chelate and import host iron

4. Antigenic variation

  • N. gonorrhoeae: pili and outer membrane proteins
  • Trypanosoma brucei: surface antigen switching
  • Enterobacteriaceae: capsular and flagellar antigens switched on or off
  • Influenza and HIV: mutation-driven change in surface proteins

5. Surviving inside cells

  • Block phagosome–lysosome fusion
    • Mycobacterium tuberculosis
  • Escape the phagosome into cytoplasm
    • Listeria monocytogenes
  • Invasins drive entry into non-phagocytic cells
    • Best characterized in Yersinia pseudotuberculosis
  • Viral injury comes from intracellular replication
    • Cell death, transformation or silent latency

6. Type III secretion

  • Needle-like channel injects effector proteins into host cells
  • Seen in E. coli, Salmonella, Yersinia, Pseudomonas aeruginosa, Chlamydia

7. Immune-mediated injury

MechanismExample
Antibody cross-reactivity with host tissueRheumatic fever after group A strep
Immune complex depositionPost-streptococcal glomerulonephritis
Delayed hypersensitivity and granulomas
  • Nerve damage in leprosy
  • Cavitation in tuberculosis
  • Tubal scarring and infertility after Chlamydia pelvic infection
Wall fragments driving inflammationGram-positive peptidoglycan–teichoic acid

8. Physical damage

  • Swelling in a closed space
    • Meningitis, neurocysticercosis
  • Bulk of the organism
    • Ascaris blocking the bile duct
High-yield
  • Intracellular pathogens trigger different immune responses, need different antibiotics and different culture methods.

☠️ Endotoxin and Exotoxin Principles

Endotoxin (LPS)

  • Structural component of the gram-negative outer membrane
  • Toxic part: lipid A
  • Released mainly when bacteria die or lyse
    • Neisseria meningitidis sheds outer membrane blebs while alive
  • Heat stable
  • Weakly immunogenic, so cannot be made into a toxoid
  • Main driver of gram-negative sepsis

Pathway to septic shock

LPS→TLR-4 on macrophages→TNF-α, IL-1, IL-6→Endothelial injury, vasodilation→Hypotension, shock
  • IL-1 is the major fever mediator
  • Bradykinin-mediated vasodilation worsens shock
  • Hageman factor (XII) activation triggers clotting
    • Leads to DIC

Exotoxins

  • Secreted protein toxins
  • Produced by both gram-positive and gram-negative species
  • Highly potent in tiny amounts
  • Can be inactivated by heat or chemicals into a toxoid
    • Keeps immunogenicity, loses toxicity
    • Basis of tetanus and diphtheria vaccines
  • Types by action
    • Enterotoxins: act on gut
    • Neurotoxins: act on nerves
    • Cytotoxins: kill cells

A–B toxin design

  • B subunit: binds a host receptor and delivers A into the cell
  • A subunit: enzymatic, often ADP-ribosyltransferase

Cytolysins

  • Damage membranes directly from outside
    • Clostridium perfringens alpha toxin: lecithinase (phospholipase C)
    • Staphylococcus aureus alpha toxin: forms membrane pores

Superantigens

  • Bind MHC II and T-cell receptor outside the antigen groove
  • Massive polyclonal T-cell activation with cytokine surge
Exam trap
  • Endotoxin cannot be toxoided, but exotoxins can.

💉 Major Exotoxins by Mechanism

Protein synthesis inhibitors

OrganismToxinMechanismEffect
Corynebacterium diphtheriae (gram +)Diphtheria toxinADP-ribosylates elongation factor 2 (eEF-2)Epithelial necrosis with pseudomembrane; cardiac and nerve injury
Pseudomonas aeruginosa (gram −)Exotoxin AADP-ribosylates eEF-2Hepatocyte injury, tissue necrosis
Shigella dysenteriae (gram −)Shiga toxinInactivates 60S ribosome (rRNA cleavage)Enterotoxic, cytotoxic and neurotoxic; bloody diarrhea
Enterohemorrhagic E. coli (gram −)Shiga-like toxin (verotoxin)Inactivates 60S ribosomeHemorrhagic colitis; hemolytic uremic syndrome

Neurotoxins

OrganismToxinMechanismEffect
Clostridium tetani (+)TetanospasminBlocks release of inhibitory glycine and GABASpastic paralysis, lockjaw
Clostridium botulinum (+)Botulinum toxinBlocks acetylcholine release at cholinergic junctionsFlaccid paralysis

Superantigens

OrganismToxinEffect
Staphylococcus aureus (+)TSST-1Fever, rash, capillary leak, shock; heightened sensitivity to LPS
Streptococcus pyogenes (+)Exotoxin A (pyrogenic or erythrogenic)Similar illness, scarlet fever rash, cardiotoxicity

Toxins that raise cAMP

OrganismToxinMechanismEffect
Enterotoxigenic E. coli (−)Heat-labile toxin (LT)ADP-ribosylates Gs, locking adenylate cyclase onSecretory diarrhea (heat-stable toxin ST acts through cGMP)
Vibrio cholerae (−)Cholera toxinSame as LTProfuse watery diarrhea
Bacillus anthracis (+)Anthrax toxins (three proteins)
  • Protective antigen = B component for both
  • Edema factor = adenylate cyclase
  • Lethal factor = protease
Edema, impaired phagocytosis, cell death
Bordetella pertussis (−)Pertussis toxinADP-ribosylates Gi, removing inhibition of adenylate cyclaseLymphocytosis, increased histamine sensitivity, insulin release

Cytolysins

OrganismToxinMechanismEffect
Clostridium perfringens (+)Alpha toxinLecithinaseMembrane breakdown, myonecrosis (gas gangrene)
Staphylococcus aureus (+)Alpha toxinInserts to form poresLeaky, lysed cells
High-yield
  • Diphtheria toxin and Pseudomonas exotoxin A share the same target, eEF-2.
  • Cholera, ETEC LT and pertussis toxin all alter G-protein signaling to change cAMP.
  • Tetanus blocks inhibitory transmitters, so muscles contract; botulism blocks excitatory transmitter, so muscles relax.