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Brock Biology of Microorganisms

Twelfth Edition

Madigan / Martinko Dunlap / Clark

Chapter 21
Metabolic Diversity: Catabolism of Organic Compounds
Lectures by Buchan & LeCleir
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I. Fermentations
21.1 Fermentations: Energetic and Redox Considerations 21.2 Fermentative Diversity: Lactic and Mixed-Acid Fermentations 21.3 Fermentative Diversity: Clostridial and Propionic Acid Fermentations 21.4 Fermentations without Substrate-Level

Phosphorylation
21.5 Syntrophy

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21.1 Fermentations: Energetic and Redox Considerations


Two mechanisms for catabolism of organic compounds
Respiration
Exogenous electron acceptors are present to accept
electrons generated from the oxidation of electron donors

Fermentation
Electron donor and acceptor are the same compound Relatively little energy yield

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21.1 Fermentations: Energetic and Redox Considerations


In the absence of external electron acceptors, compounds
can be catabolized anaerobically by fermentation ATP is usually synthesized by substrate-level phosphorylation
Energy-rich phosphate bonds from phosphorylated organic intermediates transferred directly to ADP

Redox balance is achieved by production and secretion of fermentation products

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The Essentials of Fermentation

Figure 21.1

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21.1 Fermentations: Energetic and Redox Considerations

A requirement for most fermentations is that organic intermediates can be generated that contain an energy-rich phosphate bond or a molecule of

coenzyme-A

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Energy-Rich Compounds Involved in SLP

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Fermentation
End products of fermentation include
Lactic acid Ethanol Butyric acid

Propionic acid
2,3-Butanediol Mixed acids

All are produced in a series of reactions to produce appropriate terminal electron acceptors
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Anaerobic Breakdown of Major Fermentable Substrates

Figure 21.2

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21.2 Lactic and Mixed-Acid Fermentations


Fermentations are classified by either the substrate

fermented or the productions formed


A wide variety of organic compounds can be fermented

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Common Bacterial Fermentations

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21.2 Lactic and Mixed-Acid Fermentations

Lactic acid fermentation can occur by homofermentative and heterofermentative pathways

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Glucose Fermentation by Homofermentations

Figure 21.4

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21.2 Lactic and Mixed-Acid Fermentations


The Entner-Doudoroff Pathway
A variant of the glycolytic pathway A widespread pathway for sugar catabolism in bacteria

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21.2 Lactic and Mixed-Acid Fermentations


Mixed-Acid Fermentations
Generate acids
Acetic, lactic, and succinic Sometimes also generate neutral products
E.g., butanediol

Characteristic of enteric bacteria

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II. Anaerobic Respiration


21.6 Anaerobic Respiration: General Principles 21.7 Nitrate Reduction and Denitrification 21.8 Sulfate and Sulfur Reduction 21.9 Acetogenesis

21.10 Methanogenesis
21.11 Proton Reduction 21.12 Other Electron Acceptors 21.13 Anoxic Hydrocarbon Oxidation Linked to Anaerobic Respiration
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21.6 Anaerobic Respiration: General Principles


In anaerobic respiration electron acceptors other than O2
are used Anaerobic and aerobic respiratory systems are similar
But anaerobic respiration yields less energy than aerobic respiration

Energy released from redox reactions can be determined by comparing reduction potentials of each electron acceptor
Animation: Electon Transport: Aerobic & Anaerobic Conditions

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Major Forms of Anaerobic Respiration

Figure 21.12

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21.7 Nitrate Reduction and Denitrification

Inorganic nitrogen compounds are the most common electron acceptors in anaerobic respiration

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Oxidation States of Key Nitrogen Compounds

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21.7 Nitrate Reduction and Denitrification

All products of nitrate reduction (denitrification) are


gaseous Denitrification is the main biological source of gaseous N2

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21.7 Nitrate Reduction and Denitrification

The biochemical pathway for dissimilative nitrate reduction has been well-studied Enzymes of the pathway are repressed by oxygen

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Respiration and Anaerobic Respiration

Figure 21.14c

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21.8 Sulfate and Sulfur Reduction

Several inorganic sulfur compounds can be used as electron acceptors in anaerobic respiration

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Sulfur Compounds and Electron Donors for SO42- Reduction

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21.9 Acetogenesis
Acetogens and methanogens use CO2 as an
electron acceptor in anaerobic respiration
H2 is the major electron donor for both groups of organisms

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The Processes of Methanogenesis and Acetogenesis

Figure 21.17

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21.10 Methanogenesis

Methanogenesis
Involves a complex series of biochemical reactions that use novel coenzymes

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Fluorescence Due to the Methanogenic Coenzyme F420

Autofluourescence in Cells of the Methanogen Methanosarcina barkeri

F420 fluorescence in Cells of the Methanogen Methanobacterium formicicum

Figure 21.20

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21.10 Methanogenesis

H2 is the major electron donor for methanogenesis

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Methanogenesis from CO2 plus H2

Figure 21.21

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21.12 Other Electron Acceptors

Fe3+, Mn4+, ClO3-, and various organic compounds can serve as electron acceptors for bacteria Fe3+ is abundant in nature and its reduction is a major form of anaerobic respiration

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Alternative Electron Acceptors for Anaerobic Respirations

Figure 21.25

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21.12 Other Electron Acceptors

The reduction of arsenate has been employed for clean-up of toxic wastes and groundwater

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21.13 Anoxic Hydrocarbon Oxidation

Methane
The simplest hydrocarbon Can be oxidized under anoxic conditions by a consortia containing sulfate-reducing bacteria and

methanotrophic archaea

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21.15 Aerobic Hydrocarbon Oxidation


Many bacteria and eukaryotic microbes can use aliphatic and aromatic hydrocarbons as electron

donors when growing aerobically


Oxygenases are central enzymes in these biochemical reactions Aerobic aromatic compound degradation involves ring oxidation

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21.16 Methylotrophy and Methanotrophy


Methylotrophs use compounds that lack C-C bonds as electron donors and carbon sources Methanotrophs are methylotrophs that use CH4
The initial step in methanotrophy requires methane monooxygenase (MMO)

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21.17 Hexose, Pentose, and Polysaccharide Metabolism

Sugars and polysaccharides are common

substrates for chemoorganotrophs


Polysaccharides such as cellulose and starch are

common in nature
Their breakdown yields hexoses and pentoses that are

readily catabolized by microbes

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Naturally Occurring Polysaccharides Yielding Sugars

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