Nutrition, Culture, and

Metabolism of Microorganisms,
Part II
01-27-15

IV. Essentials of Catabolism

3.8 Glycolysis
3.9 Fermentative Diversity and the
Respiratory Option
3.10 Respiration: Electron Carriers
3.11 Respiration: The Proton Motive Force
3.12 Respiration: Citric Acid Cycle
3.13 Catabolic Diversity

3.8 Glycolysis
Two reaction series are linked to energy
conservation in chemoorganotrophs:
fermentation and respiration
Differ in mechanism of ATP synthesis
Fermentation: substrate-level
phosphorylation; ATP directly synthesized
from an energy-rich intermediate
Respiration: oxidative phosphorylation; ATP
produced from proton motive force formed by
transport of electrons

Energy-rich
P i intermediates
A

B~P

ADP

ATP

C ~P

Substrate-level phosphorylation

Energized
membrane
Dissipation of proton
motive force coupled
to ATP synthesis

ADP + P i

Less energized
membrane

Oxidative phosphorylation

ATP

3.8 Glycolysis
Fermented substance is both an electron
donor and an electron acceptor
Glycolysis (Embden-Meyerhof pathway): a
common pathway for catabolism of glucose
Anaerobic process
Three stages

Overview
Glucose consumed
Two ATPs produced
Fermentation products generated
Some harnessed by humans for consumption

3.9 Fermentative Diversity and the

Respiratory Option
Fermentations classified by products
formed
Ethanol
Lactic acid
Propionic acid
Mixed acids
Butyric acid
Butanol

3.9 Fermentative Diversity and the

Respiratory Option
Fermentations classified by substrate
fermented

Usually NOT glucose

Amino acids
Purines and pyrimidines
Aromatic compounds

Saccharomyces cerevisiae can carry out

fermentation or respiration
Carries out the one most beneficial
Respiration generates more ATP
Fermentation occurs when conditions are anoxic

3.10 Respiration: Electron Carriers

Aerobic Respiration
Oxidation using O2 as the terminal electron
acceptor
Higher ATP yield than fermentations
ATP produced at the expense of the proton motive
force, which is generated by electron transport

3.10 Respiration: Electron Carriers

Electron Transport Systems
Membrane associated
Mediate transfer of electrons
Conserve some of the energy released during
transfer and use it to synthesize ATP
Many oxidationreduction enzymes are
involved in electron transport

NADH dehydrogenases
flavoproteins
ironsulfur proteins
cytochromes

3.10 Respiration: Electron Carriers

NADH dehydrogenases
proteins bound to inside surface of cytoplasmic
membrane
active site binds NADH and accepts 2 electrons
and 2 protons that are passed to flavoproteins

Flavoproteins
contains flavin prosthetic
group (e.g., FMN, FAD)
that accepts 2 electrons
and 2 protons but
only donates the
Oxidized (FMN)
electrons to the next
Reduced (FMNH )
protein in the chain
E of FMN/FMNH (or FAD/FADH ) = 0.22 V
Isoalloxazine ring

Ribitol

3.10 Respiration: Electron Carriers

Porphyrin
ring

Cytochromes
Proteins that contain
heme prosthetic
Pyrrole
groups
Accept and donate a
single electron via Heme (a porphyrin)
the iron atom
Histidine-N
in heme

Protein

N-Histidine

Cysteine-S

S-Cysteine
Amino acid Amino acid

Cytochrome

3.10 Respiration: Electron Carriers

IronSulfur Proteins
Contain clusters of iron
and sulfur

Cysteine

Cysteine

Cysteine

Cysteine

Example: ferredoxin

Reduction potentials
vary depending on
number and position of
Fe and S atoms
Carry electrons

Cysteine
Cysteine
Cysteine
Cysteine

3.10 Respiration: Electron Carriers

Quinones
Hydrophobic non-protein-containing
molecules that participate in electron
transport
Accept electrons and
protons but pass
along electrons
only
Oxidized

Reduced

3.11 Respiration: The Proton Motive Force

Electron transport system oriented in
cytoplasmic membrane so that electrons
are separated from protons
Electron carriers arranged in membrane in
order of their reduction potential
The final carrier in the chain donates the
electrons and protons to the terminal
electron acceptor

3.11 The Proton Motive Force

During electron transfer, several protons
are released on outside of the membrane
Protons originate from NADH and the
dissociation of water

Results in generation of pH gradient and an

electrochemical potential across the
membrane (the proton motive force)
The inside becomes electrically negative and
alkaline
The outside becomes electrically positive and
acidic

3.11 The Proton Motive Force

E0(V)

Complex I (NADH:quinone
oxidoreductase)

0.22

NADH donates to FMN

FMNH2 donates e to quinone

Bypasses Complex I
Feeds e and H+ from FADH directly to
quinone pool

0.0
ENVIRONMENT

Complex II (succinate dehydrogenase

complex)

Q
cycle

Complex IV (cytochromes a and a3)

Succinate
Fumarate

0.1

CYTOPLASM

Complex III (cytochrome bc1 complex)

Transfers e from quinones to
cytochrome c
Cytochrome c shuttles e to
cytochromes a and a3

Complex II

0.36

0.39

Terminal oxidase; reduces O2 to H2O

E0(V)

3.11 The Proton Motive Force

F1: multiprotein
extramembrane complex,
faces cytoplasm
Fo: proton-conducting
intramembrane channel Membrane c12

Reversible; dissipates
proton motive force

F1

b2

In

ATP synthase (ATPase):

complex that converts
proton motive force into
ATP
Two components

a
Fo

Out

3.12 The Citric Acid Cycle

Citric acid cycle (CAC): pathway through
which pyruvate is completely oxidized to
CO2
Initial steps (glucose to pyruvate) same as
glycolysis
Per glucose molecule, 6 CO2 molecules
released and NADH and FADH generated
Plays a key role in catabolism and
biosynthesis

Energetics advantage to aerobic

respiration

3.12 The Citric Acid Cycle

The citric acid cycle generates many
compounds available for biosynthetic purposes
-Ketoglutarate and oxalacetate (OAA):
precursors of several amino acids; OAA also
converted to phosphoenolpyruvate, a precursor
of glucose
Succinyl-CoA: required for synthesis of
cytochromes, chlorophyll, and other tetrapyrrole
compounds
Acetyl-CoA: necessary for fatty acid biosynthesis

3.13 Catabolic Diversity

Microorganisms demonstrate a wide range
of mechanisms for generating energy

Fermentation
Aerobic respiration
Anaerobic respiration
Chemolithotrophy
Phototrophy

Fermentation

Electron donor

(organic compound)

Organic e
acceptors

Electron
acceptors

Aerobic
respiration

Anaerobic respiration
Chemoorganotrophy

Electron transport/
generation of pmf

Light
Organic
compound
Electron
transport

Aerobic respiration

Electron
acceptors
Anaerobic respiration
Chemolithotrophy

Photoautotrophy

Photoheterotrophy

Phototrophs

Chemotrophs

Electron transport/
generation of pmf

Generation of pmf
Cell material

Phototrophy

Cell material

3.13 Catabolic Diversity

Anaerobic Respiration
The use of electron acceptors other than
oxygen
Examples include nitrate (NO3), ferric iron
(Fe3+), sulfate (SO42), carbonate (CO32),
certain organic compounds

Less energy released compared to

aerobic respiration
Dependent on electron transport,
generation of a proton motive force, and
ATPase activity

3.13 Catabolic Diversity

Chemolithotrophy
Uses inorganic chemicals as electron donors
Examples include hydrogen sulfide (H2S), hydrogen
gas (H2), ferrous iron (Fe2+), ammonia (NH3)

Typically aerobic
Begins with oxidation of inorganic electron
donor
Uses electron transport chain and proton
motive force
Autotrophic; uses CO2 as carbon source

3.13 Catabolic Diversity

Phototrophy: uses light as energy
source
Photophosphorylation: light-mediated ATP
synthesis
Photoautotrophs: use ATP for assimilation
of CO2 for biosynthesis
Photoheterotrophs: use ATP for
assimilation of organic carbon for
biosynthesis