Nutrition, Culture, and Metabolism of Microorganisms, Part II 01-27-15
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Nutrition, Culture, and
Metabolism of Microorganisms,
Part II
01-27-15
7/17/2019 NutCultCatab II
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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
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3.8 Glycolysis
•Two reaction series are linked to energyconservation 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
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Energy-rich
intermediates ADP
BA B P P DC~ ~
P i
Substrate-level phosphorylation
Oxidative phosphorylation
ADP P i
Energizedmembrane
Less energized
membrane
Dissipation of proton
motive force coupled
to ATP synthesis
+
ATP
ATP
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3.8 Glycolysis
• Fermented substance is both an electrondonor and an electron acceptor
• Glycolysis (Embden-
Meyerhof pathway): acommon pathway for catabolism of glucose
– Anaerobic process
– Three stages
• Overview
–Glucose consumed
– Two ATPs produced
– Fermentation products generated• Some harnessed by humans for consumption
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3.9 Fermentative Diversity and the
Respiratory Option
• Fermentations classified by products
formed
– Ethanol
– Lactic acid
– Propionic acid
– Mixed acids
– Butyric acid
– Butanol
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3.9 Fermentative Diversity and the
Respiratory Option
• Fermentations classified by substratefermented – 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
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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
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3.10 Respiration: Electron Carriers
• Electron Transport Systems – Membrane associated
– Mediate transfer of electrons
–
Conserve some of the energy released duringtransfer and use it to synthesize ATP
– Many oxidation –reduction enzymes areinvolved in electron transport•
NADH dehydrogenases• flavoproteins
• iron –sulfur proteins
• cytochromes
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3.10 Respiration: Electron Carriers
Isoalloxazine ring
Ribitol
Oxidized (FMN)
Reduced (FMNH2)
E 0′ of FMN/FMNH2 (or FAD/FADH2) = – 0.22 V
• NADH dehydrogenases
– proteins bound to inside surface of cytoplasmicmembrane
– active site binds NADH and accepts 2 electrons
and 2 protons that are passed to flavoproteins• Flavoproteins
– contains flavin prostheticgroup (e.g., FMN, FAD)
that accepts 2 electronsand 2 protons butonly donates theelectrons to the next
protein in the chain
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Porphyrinring
Pyrrole
Heme (a porphyrin)
Histidine-N
Cysteine-S
Amino acid Amino acid
S-Cysteine
N-Histidine
Protein
Cytochrome
3.10 Respiration: Electron Carriers
•
Cytochromes – Proteins that contain
heme prosthetic
groups
– Accept and donate a
single electron via
the iron atom
in heme
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Cysteine
Cysteine
Cysteine
Cysteine
Cysteine
Cysteine
Cysteine
Cysteine
3.10 Respiration: Electron Carriers
•
Iron –
Sulfur Proteins – Contain clusters of iron
and sulfur
• Example: ferredoxin
– Reduction potentials
vary depending on
number and position of
Fe and S atoms – Carry electrons
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Oxidized
Reduced
3.10 Respiration: Electron Carriers
• Quinones
– Hydrophobic non-protein-containing
molecules that participate in electron
transport
– Accept electrons andprotons but pass
along electrons
only
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3.11 Respiration: The Proton Motive Force
• Electron transport system oriented incytoplasmic membrane so that electronsare separated from protons
• Electron carriers arranged in membrane inorder of their reduction potential
•
The final carrier in the chain donates theelectrons and protons to the terminalelectron acceptor
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3.11 The Proton Motive Force
• During electron transfer, several protonsare released on outside of the membrane
– Protons originate from NADH and thedissociation of water
• Results in generation of pH gradient and anelectrochemical potential across themembrane (the proton motive force)
– The inside becomes electrically negative andalkaline
– The outside becomes electrically positive andacidic
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CYTOPLASM E N V I R O N M E N
T
Complex II
Succinate
Fumarate
Qcycle
0.22
0.0
0.1
0.36
0.39
E 0(V)
E 0(V)
3.11 The Proton Motive Force• Complex I ( NADH:quinone
oxidoreductase) – NADH donates e to FMN
– FMNH2 donates e to quinone
• Complex II (succinate dehydrogenasecomplex)
– Bypasses Complex I – Feeds e and H+ from FADH directly to
quinone pool
• Complex III (cytochrome bc 1 com plex) – Transfers e from quinones to
cytochrome c – Cytochrome c shuttles e to
cytochromes a and a3
• Complex IV (cytochromes a and a3 ) – Terminal oxidase; reduces O2 to H2O
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MembraneOut
In
F1
Foc12
b2
a
3.11 The Proton Motive Force
• ATP synthase (ATPase):complex that convertsproton motive force into ATP
• Two components – F 1: multiprotein
extramembrane complex,faces cytoplasm
– Fo: proton-conductingintramembrane channel
• Reversible; dissipatesproton motive force
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3.12 The Citric Acid Cycle
•Citric acid cycle (CAC): pathway throughwhich pyruvate is completely oxidized toCO2
– 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
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3.12 The Citric Acid Cycle
• The citric acid cycle generates manycompounds 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 tetrapyrrolecompounds
– Acetyl-CoA: necessary for fatty acid biosynthesis
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3.13 Catabolic Diversity
• Microorganisms demonstrate a wide rangeof mechanisms for generating energy
– Fermentation
– Aerobic respiration – Anaerobic respiration
– Chemolithotrophy
–
Phototrophy
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Electron donor
(organic compound)
Organic e –
acceptors
Electron
acceptors
Electron
acceptors
Light
Fermentation
C h e m o t r o p h s
Aerobic
respiration
Anaerobic respiration
Chemoorganotrophy
Anaerobic respiration
Aerobic respiration
Chemolithotrophy
P h o t o t r o p h s
Electron
transport
Generation of pmf
Organic
compound
Cell materialCell material
Photoheterotrophy Photoautotrophy
Phototrophy
Electron transport/
generation of pmf
Electron transport/
generation of pmf
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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 toaerobic respiration
– Dependent on electron transport,generation of a proton motive force, and
ATPase activity
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3.13 Catabolic Diversity
• Chemolithotrophy – Uses inorganic chemicals as electron donors
• Examples include hydrogen sulfide (H2S), hydrogengas (H2), ferrous iron (Fe2+), ammonia (NH3)
– Typically aerobic – Begins with oxidation of inorganic electron
donor
– Uses electron transport chain and protonmotive force
– Autotrophic; uses CO2 as carbon source
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3.13 Catabolic Diversity
• Phototrophy : uses light as energysource
– Photophosphorylation: light-mediated ATP
synthesis – Photoautotrophs: use ATP for assimilation
of CO2 for biosynthesis
– Photoheterotrophs: use ATP for
assimilation of organic carbon for
biosynthesis