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Brock Biology of Microorganisms

Twelfth Edition

Madigan / Martinko Dunlap / Clark

Chapter 20
Metabolic Diversity: Phototrophy, Autotrophy, Chemolithotrophy, and Nitrogen Fixation
Lectures by Buchan & LeCleir
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I. The Phototrophic Way of Life

20.1 Photosynthesis 20.2 Chlorophylls and Bacteriochlorophylls 20.3 Carotenoids and Phycobilins

20.4 Anoxygenic Photosynthesis


20.5 Oxygenic Photosynthesis

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20.1 Photosynthesis
Photosynthesis is the most important biological process
on Earth
Phototrophs are organisms that carry out photosynthesis

Most phototrophs are also autotrophs

Photosynthesis requires light-sensitive pigments called chlorophyll

Photoautotrophy requires ATP production and CO2


reduction Oxidation of H2O produces O2 (oxygenic photosynthesis) Oxygen not produced (anoxygenic photosynsthesis)
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Patterns of Photosynthesis

Figure 20.2

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Patterns of Photosynthesis

Figure 20.2

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20.2 Chlorophylls and Bacteriochlorophylls


Organisms must produce some form of chlorophyll (or bacteriochlorophyll) to be photosynthetic Chlorophyll is a porphyrin

Number of different types of chlorophyll exist


Different chlorophylls have different absorption spectra

Chlorophyll pigments are located within special

membranes
In eukaryotes, called thylakoids In prokaryotes, pigments are integrated into cytoplasmic

membrane
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Structure and Spectra of Chloro- and Bacteriochlorophyll

Figure 20.3a

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Structure and Spectra of Chloro- and Bacteriochlorophyll

Absorption Spectrum
Figure 20.3b

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Structure of All Known Bacteriochlorophylls

Figure 20.4

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Structure of All Known Bacteriochlorophylls

Figure 20.4

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Structure of All Known Bacteriochlorophylls

Figure 20.4

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Photomicrograph of Algal Cell Showing Chloroplasts

Figure 20.5a

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Chloroplast Structure

Figure 20.5b

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20.2 Chlorophylls and Bacteriochlorophylls


Reaction centers participate directly in the conversion

of light energy to ATP


Antenna pigments funnel light energy to reaction centers Chlorosomes function as massive antenna complexes
Found in green sulfur bacteria

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The Chlorosome of Green Sulfur and Nonsulfur Bacteria

Electron Micrograph of Cell of Green Sulfur Bacterium

Figure 20.7

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The Chlorosome of Green Sulfur and Nonsulfur Bacteria

Model of Chromosome Structure


Figure 20.7

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20.3 Carotenoids and Phycobilins


Phototrophic organisms have accessory pigments in
addition to chlorophyll, including carotenoids and phycobiliproteins Carotenoids
Always found in phototrophic organisms Typically yellow, red, brown, or green Energy absorbed by carotenoids can be transferred to a reaction center

Prevent photo-oxidative damage to cells


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Structures of Some Common Carotenoids

Figure 20.9

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Absorption Spectrum with an Accessory Pigment

Figure 20.11

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20.4 Anoxygenic Photosynthesis


Anoxygenic photosynthesis is found in at least four phyla of Bacteria Electron transport reactions occur in the reaction

center of anoxygenic phototrophs


Reducing power for CO2 fixation comes from reductants present in the environment (i.e., H2S, Fe2+, or NO2-)
Requires reverse electron transport for NADH production in purple phototrophs
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Example of Electron Flow in Anoxygenic Photosynthesis

Figure 20.14

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Electron Flow in Purple, Green, Sulfur and Heliobacteria

Figure 20.18

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20.5 Oxygenic Photosynthesis


Oxygenic phototrophs use light to generate ATP and NADPH The two light reactions are called photosystem I and

photosystem II
Z scheme of photosynthesis
Photosystem II transfers energy to photosystem I

ATP can also be produced by cyclic photophosphorylation

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The Z scheme in Oxygenic Photosynthesis

Figure 20.19

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II. Autotrophy

20.6 The Calvin Cycle 20.7 Other Autotrophic Pathways in Phototrophs

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20.6 The Calvin Cycle


The Calvin Cycle
Named for its discoverer Melvin Calvin Fixes CO2 into cellular material for autotrophic growth

Requires NADPH, ATP, ribulose bisphophate


carboxylase (RubisCO), and phosphoribulokinase 6 molecules of CO2 are required to make 1 molecule of glucose

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The Calvin Cycle

Figure 20.22

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20.7 Other Autotrophic Pathways in Phototrophs


Green sulfur bacteria use the reverse citric acid cycle

to fix CO2
Green nonsulfur bacteria use the hydroxyproprionate pathway to fix CO2

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Autotrophic Pathways in Phototrophic Green Bacteria

Figure 20.24a

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Autotrophic Pathways in Phototrophic Green Bacteria

Figure 20.24b

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III. Chemolithotrophy

20.8 20.9

The Energetics of Chemolithotrophy Hydrogen Oxidation

20.10 Oxidation of Reduced Sulfur Compounds

20.11 Iron Oxidation


20.12 Nitrification

20.13 Anammox

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20.8 The Energetics of Chemolithotrophy


Chemolithotrophs are organisms that obtain energy

from the oxidation of inorganic compounds


Many sources of reduced molecules exist in the environment

The oxidation of different reduced compounds yields


varying amounts of energy

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Energy Yields from Oxidation of Inorganic Electron Donors

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20.9 Hydrogen Oxidation


Anaerobic H2 oxidizing Bacteria and Archaea are

known
Catalyzed by hydrogenase In the presence of organic compounds such as glucose, synthesis of Calvin cycle and hydrogenase enzymes is repressed

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Two Hydrogenases of Aerobic H2 Bacteria

Figure 20.25

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20.10 Oxidation of Reduced Sulfur Compounds


Many reduced sulfur compounds are used as electron donors
Discovered by Sergei Winogradsky H2S, S0, S2O3- are commonly used One product of sulfur oxidation is H+, which results in a lowering of the pH of its surroundings Sox system oxidizes reduced sulfur compounds directly to sulfate Usually aerobic, but some organisms can use nitrate as an

electron acceptor

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Sulfur Bacteria

Figure 20.26

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Oxidation of Reduced Sulfur Compounds

Steps in the Oxidation of Different Compounds

Figure 20.27a

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20.11 Iron Oxidation


Ferrous iron (Fe2+) oxidized to ferric iron (Fe3+)
Ferric hydroxide precipitates in water in the presence of O2 at neutral pH Many Fe oxidizers can grow at pH <1
Often associated with acidic pollution from coal mining activities ** Ferrous iron (Fe2+) is only available in anaerobic conditions or aerobically at acid pH

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20.11 Iron Oxidation


Some anoxygenic phototrophs can oxidize Fe2+
anaerobically using Fe2+ as an electron donor for CO2 reduction

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Iron-Oxidizing Bacteria

Acid Mine Drainage

Figure 20.28

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20.12 Nitrification
NH3 and NO2- are oxidized by nitrifying bacteria during the
process of nitrification Two groups of bacteria work in concert to fully oxidize ammonia to nitrate
NH3 > NO2- by Nitrosomonas sp. NO2- > NO3 by Nitrobacter sp.

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20.12 Nitrification
Key enzymes are ammonia monooxygenase,
hydroxylamine oxidoreductase, and nitrite oxidoreductase

Only small energy yields from this reaction


Growth of nitrifying bacteria is very slow

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20.13 Anammox
Anammox: anoxic ammonia oxidation

Performed by unusual group of obligate aerobes


Anammoxosome is compartment where anammox reactions occur
Protects cell from reactions occuring during anammox Hydrazine is an intermediate of anammox

Anammox is very beneficial in the treatment of sewage


and wastewater

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IV. Nitrogen Fixation

20.14 Nitrogenase and Nitrogen Fixation 20.15 Genetics and Regulation of Nitrogen Fixation

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Biogeochemical Cycling and Energy Flow

Nitrogen fixation N2

NH3

Process in which nitrogen gas is reduced to form ammonia Ammonia can be incorporated into cellular material Animals rely on prokaryotes to convert nitrogen gas to a form that can be assimilated to create biomass Nitrogenase mediates nitrogen fixation Enzyme complex is readily inactivated by oxygen Nitrogen fixing microbes called diazotrophs must have mechanism to protect nitrogenase complex from oxygen exposure

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20.14 Nitrogenase and Nitrogen Fixation


Only certain prokaryotes can fix nitrogen
Reaction is catalyzed by nitrogenase
Sensitive to the presence of oxygen

A wide variety of nitrogenases use different metal


cofactors Nitrogenase activity can be assayed using the acetylene reduction assay

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N2 Fixing Bacteria
May be free living or form symbiotic associations with higher

organisms particularly plants

May be aerobic or anaerobic,

May be autotrophic or heterotrophic

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N2 Fixation in Aerobic Environments


Cyanobacteria form heterocysts

Some aerobic bacteria produce capsule

Other aerobic bacteria have a high metabolic rate that consumes O2 near the cytoplasmic membrane

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FeMo-co, the Iron-Molybdenum Cofactor from Nitrogenase

Figure 20.35

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Nitrogenase Function

Figure 20.36

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Induction of Slime Formation by O2 in Nitrogen-Fixing Cells

Figure 20.37

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The Acetylene Reduction Assay for Nitrogenase Activity

Figure 20.39

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20.15 Genetics and Regulation of Nitrogen Fixation


Highly regulated process because it is such an energy-

demanding process
nif regulon coordinates regulation of genes essential to nitrogen fixation Oxygen and ammonia are the two main regulatory effectors

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