PowerPoint ® Lecture Presentations prepared by John Zamora Middle Tennessee State University C H A P T E R © 2015 Pearson Education, Inc. Metabolic Diversity.

© 2015 Pearson Education, Inc. I. Phototrophy 13.1 Photosynthesis and Chlorophylls 13.2 Carotenoids and Phycobilins 13.3 Anoxygenic Photosynthesis 13.4 Oxygenic Photosynthesis 13.5 Autotrophic Pathways

© 2015 Pearson Education, Inc. 13.1 Photosynthesis and Chlorophylls Photosynthesis is the conversion of light energy to chemical energy Phototrophs carry out photosynthesis Most phototrophs are also autotrophs Photosynthesis requires light-sensitive pigments called chlorophylls Photoautotrophy requires ATP production and CO 2 reduction

© 2015 Pearson Education, Inc. 13.1 Photosynthesis and Chlorophylls Photoautotrophy Oxidation of H 2 O produces O 2 (oxygenic photosynthesis; Figure 13.1) Oxygen not produced (anoxygenic photosynsthesis; Figure 13.1)

© 2015 Pearson Education, Inc. Prokaryotic phototrophs Purple and green bacteriaCyanobacteria Anoxygenic Light ADP ATP Reducing powerCarbonEnergy Oxygenic Reducing powerCarbonEnergy electrons ADP Light ATP Figure 13.1

© 2015 Pearson Education, Inc. 13.1 Photosynthesis and Chlorophylls Organisms must produce some form of chlorophyll (or bacteriochlorophyll) to be photosynthetic (Figure 13.2a) Chlorophyll is related to porphyrins Number of different types of chlorophyll exist Different chlorophylls have different absorption spectra (Figure 13.2b)

© 2015 Pearson Education, Inc. 13.1 Photosynthesis and Chlorophylls Cyanobacteria produce chlorophyll a (Figure 13.3) Prochlorophytes produce chlorophyll a and b Anoxygenic phototrophs produce bacteriochlorophylls

© 2015 Pearson Education, Inc. 13.1 Photosynthesis and Chlorophylls Reaction centers participate directly in the conversion of light energy to ATP Antenna pigments funnel light energy to reaction centers (Figure 13.4)

© 2015 Pearson Education, Inc. 13.1 Photosynthesis and Chlorophylls Chlorophyll pigments are located within special membranes In eukaryotes, they are called thylakoids (Figure 13.5) In prokaryotes, pigments are integrated into cytoplasmic membrane (Figure 13.6) Chlorosomes function as massive antenna complexes (Figure 13.7) Found in green sulfur bacteria and green nonsulfur bacteria

© 2015 Pearson Education, Inc. 13.2 Carotenoids and Phycobilins Phototrophic organisms have accessory pigments in addition to chlorophyll, including carotenoids and phycobiliproteins Carotenoids (Figure 13.8) Always found in phototrophic organisms Typically yellow, red, brown, or green (Figure 13.9) Energy absorbed by carotenoids can be transferred to a reaction center Prevent photooxidative damage to cells

© 2015 Pearson Education, Inc. 13.2 Carotenoids and Phycobilins Phycobiliproteins are main antenna pigments of cyanobacteria and red algae Form into aggregates within the cell called phycobilisomes (Figure 13.10) Allow cell to capture more light energy than chlorophyll alone

© 2015 Pearson Education, Inc. 13.3 Anoxygenic Photosynthesis Anoxygenic photosynthesis is found in four phyla of Bacteria Photosynthesis apparatus is embedded in membranes (Figure 13.11) Electron transport reactions occur in the reaction center of anoxygenic phototrophs (Figure 13.11)

© 2015 Pearson Education, Inc. 13.3 Anoxygenic Photosynthesis Reducing power for CO 2 fixation comes from reductants present in the environment (i.e., H 2 S, Fe 2+, or NO 2– ) Requires reverse electron transport for NADH production in purple phototrophs (Figure 13.12a) Electrons are transported in the membrane through a series of proteins and cytochromes (Figure 13.12b)

© 2015 Pearson Education, Inc. Photosynthetic membrane In (cytoplasm) Out (periplasm) Arrangement of protein complexes in the purple bacterium reaction center Light ADP + P i Q Quinone pool RC Fe-S bc 1 e–e– ATP Q Q Q Q Q Q Q e–e– e–e– e–e– e–e– ATPase P870 P870* Bph Q Figure 13.12b

© 2015 Pearson Education, Inc. 13.3 Anoxygenic Photosynthesis For a purple bacterium to grow autotrophically, the formation of ATP is not enough Reducing power (NADH) is also necessary Reduced substances such as H 2 S are oxidized, and the electrons eventually end up in the "quinone pool" of the photosynthetic membrane (Figure 13.13)

© 2015 Pearson Education, Inc. Purple bacteriaGreen sulfur bacteriaHeliobacteria P870* Bchl BPh Q Cyt bc 1 Cyt c 2 P870 Light Reverse electron flow NADH P840* Chl a FeS Fd Cyt bc 1 Q Cyt c 553 P840 Light P798* Chl a–OH FeS Fd Cyt bc 1 Q Cyt c 553 Light P798 Figure 13.13

© 2015 Pearson Education, Inc. 13.4 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 (Figure 13.14) Photosystem II transfers energy to photosystem I ATP can also be produced by cyclic photophosphorylation

© 2015 Pearson Education, Inc. 13.5 Autotrophic Pathways The Calvin cycle (Figure 13.16) Named for its discoverer, Melvin Calvin Fixes CO 2 into cellular material for autotrophic growth Requires NADPH, ATP, ribulose bisphophate carboxylase (RubisCO), and phosphoribulokinase 6 molecules of CO 2 are required to make 1 molecule of glucose (Figure 13.17)

© 2015 Pearson Education, Inc. 13.5 Autotrophic Pathways Green sulfur bacteria use the reverse citric acid cycle to fix CO 2 (Figure 13.19a) Green nonsulfur bacteria use the hydroxypropionate pathway to fix CO 2 (Figure 13.19b)

© 2015 Pearson Education, Inc. II. Chemolithotrophy 13.6 Inorganic Compounds as Electron Donors 13.7 Hydrogen (H 2 ) Oxidation 13.8 Oxidation of Reduced Sulfur Compounds 13.9 Iron (Fe 2+ ) Oxidation 13.10 Nitrification and Anammox

© 2015 Pearson Education, Inc. 13.6 Inorganic Compounds as Electron Donors Chemolithotrophs are organisms that obtain energy from the oxidation of inorganic compounds Mixotrophs are chemolithotrophs that require organic carbon as a carbon source Many sources of reduced molecules exist in the environment The oxidation of different reduced compounds yields varying amounts of energy

© 2015 Pearson Education, Inc. 13.7 Hydrogen (H 2 ) Oxidation Anaerobic H 2 -oxidizing Bacteria and Archaea are known Catalyzed by hydrogenase Calvin cycle and hydrogenase enzymes allow chemolithotrophic growth (Figure 13.20)

© 2015 Pearson Education, Inc. Cytoplasmic hydrogenase Cell material ADP ATP Electron transport generates a proton motive force. Membrane-integrated hydrogenase Out In 2e – Q e–e– cyt bc 1 cyt ccyt aa 3 NADH Figure 13.20

© 2015 Pearson Education, Inc. 13.8 Oxidation of Reduced Sulfur Compounds Many reduced sulfur compounds are used as electron donors (Figure 13.22a) Discovered by Sergei Winogradsky H 2 S, S 0, S 2 O 3 – are commonly used One product of sulfur oxidation is H +, which lowers 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

© 2015 Pearson Education, Inc. Electron transport AMP APS reductase Adenosine phosphosulfate (APS) Substrate-level phosphorylation Sulfite oxidase Electron transport 8 e – 2 e – Sulfite oxidase (Sox) ADP e–e– 2 e – PiPi Figure 13.22a

© 2015 Pearson Education, Inc. 13.8 Oxidation of Reduced Sulfur Compounds Electrons from reduced sulfur compounds reach the electron transport system (Figure 13.22b) Transported through the chain to O 2 Generates a proton motive force that leads to ATP synthesis by ATPase

© 2015 Pearson Education, Inc. 13.9 Iron (Fe 2+ ) Oxidation Ferrous iron (Fe 2+ ) is oxidized to ferric iron (Fe 3+ ) Ferric hydroxide precipitates in water Many Fe oxidizers can grow at pH < 1 Often associated with acidic pollution from coal mining activities (Figure 13.23) Some anoxygenic phototrophs can oxidize Fe 2+ anaerobically using Fe 2+ as an electron donor for CO 2 reduction

© 2015 Pearson Education, Inc. 13.9 Iron (Fe 2+ ) Oxidation Ferrous iron oxidation begins in the periplasm, where rusticyanin oxidizes Fe 2+ to Fe 3+ (Figure 13.24) Rusticyanin then reduces cytochrome c, and this subsequently reduces cytochrome a Cytochrome a interacts with O 2 to form H 2 O ATP is synthesized from ATPases in the membrane Autotrophy in Acidithiobacillus ferrooxidans is driven by the Calvin cycle

© 2015 Pearson Education, Inc. Electron transport generates proton motive force. Outer membrane cyt c Rusticyanin Periplasm Reverse e – flow In NAD + Q cyt bc 1 cyt c cyt aa 3 ATP NADH ADP Cell material (pH 6) Out (pH 2) e–e– e–e– + ATP Figure 13.24

© 2015 Pearson Education, Inc. 13.10 Nitrification and Anammox NH 3 and NO 2 – are oxidized by nitrifying bacteria during the process of nitrification Two groups of bacteria work in concert to fully oxidize ammonia to nitrate Key enzymes are ammonia monooxygenase, hydroxylamine oxidoreductase, and nitrite oxidoreductase Only small energy yields from this reaction Growth of nitrifying bacteria is very slow

© 2015 Pearson Education, Inc. 13.10 Nitrification and Anammox Ammonia-oxidizing bacteria (Figure 13.26) NH3 is oxidized by ammonia monooxygenase, producing NH 2 OH and H 2 O Hydroxylamine oxidoreductase then oxidizes NH 2 OH to NO 2 – Electrons and protons are used to generate ATP

© 2015 Pearson Education, Inc. Electron transport generates a proton motive force. Oxidation of hydroxylamine Oxidation of ammonia Reduction of oxygen 4 e – 2 e – HAO AMO Reverse e – flow to make NADH Cyt aa 3 Out ADP + P i Q ATP Cyt c Figure 13.26

© 2015 Pearson Education, Inc. 13.10 Nitrification and Anammox Nitrite-oxidizing bacteria (Figure 13.27) Nitrite (NO 2 – ) is oxidized by the enzyme nitrite oxidoreductase to nitrate (NO 3 – ) Electrons and protons are used to generate ATP

© 2015 Pearson Education, Inc. 13.10 Nitrification and Anammox Anammox: anoxic ammonia oxidation Performed by unusual group of obligate aerobes (Figure 13.28a and b) Anammoxosome is a compartment where anammox reactions occur (Figure 13.28c) Protects cell from reactions occurring during anammox Hydrazine is an intermediate of anammox Anammox is very beneficial in the treatment of sewage and wastewater

© 2015 Pearson Education, Inc. III. Fermentations 13.11 Energetic and Redox Considerations 13.12 Lactic and Mixed-Acid Fermentations 13.13 Clostridial and Propionate Fermentations 13.14 Fermentations Without Substrate-Level Phosphorylation 13.15 Syntrophy

© 2015 Pearson Education, Inc. 13.11 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

© 2015 Pearson Education, Inc. 13.11 Energetic and Redox Considerations In the absence of external electron acceptors, compounds can be fermented (Figure 13.29) ATP is usually synthesized by substrate-level phosphorylation Energy-rich phosphate bonds from phosphorylated organic intermediates transferred to ADP Redox balance is achieved by production of fermentation products

© 2015 Pearson Education, Inc. 13.11 Energetic and Redox Considerations Most fermentations require that organic intermediates contain an energy-rich phosphate bond or a molecule of coenzyme A In many fermentations, redox balance is maintained by the production of molecular hydrogen (H 2 ; Figure 13.30) H 2 production involves transfer of electrons from ferredoxin to H + by a hydrogenase

© 2015 Pearson Education, Inc. 13.12 Lactic and Mixed-Acid Fermentations Fermentations are classified by either the substrate fermented or the products formed Wide variety of organic compounds can be fermented Lactic acid bacteria produce lactic acid Lactic acid fermentation can occur by homofermentative (Figure 13.31a) and heterofermentative (Figure 13.31b) pathways

© 2015 Pearson Education, Inc. Figure 13.31b ATP Glucose NADH Entner– Doudoroff pathway Pyruvate 6- Phosphogluconic acid Glucose 6-phosphate ADP NADH ATP Ribulose 5-phosphate + CO 2 Xylulose 5-phosphate PiPi NAD + Phospho- ketolase ADP PiPi 1,3-Bisphospho- glyceric acid NAD + Pyruvate – Lactate Ethanol ATP ADP NADH G-3-P NAD + Acetaldehyde Acetyl phosphate Glycer- aldehyde 3-P Heterofermentative

© 2015 Pearson Education, Inc. 13.12 Lactic and Mixed-Acid Fermentations The Entner–Doudoroff Pathway A variant of the glycolytic pathway Glucose 6-phosphate is converted to pyruvate and glyceraldehyde 3-phosphate Yields half the ATP of glycolysis A widespread pathway for sugar catabolism in bacteria

© 2015 Pearson Education, Inc. 13.12 Lactic and Mixed-Acid Fermentations Mixed-acid fermentations (Figure 13.32) Generate acids Acetic, lactic, and succinic Sometimes also generate neutral products Example: butanediol Characteristic of enteric bacteria

© 2015 Pearson Education, Inc. Glucose Glycolysis 2 Pyruvate Acetate ADP NADH 2 PiPi Phosphoroclastic reaction Acetaldehyde Ethanol Acetyl-CoA Acetoacetyl-CoA Acetoacetate β-Hydroxybutyryl-CoA Crotonyl-CoA Butyryl-CoA Butyraldehyde Butanol Butyrate Isopropanol Acetone ADP ATP NADH– consuming reactions–– butyrate, 2 NADH; butanol, 4 NADH ATP + Figure 13.33

© 2015 Pearson Education, Inc. 13.13 Clostridial and Propionate Fermentations Some Clostridium species ferment amino acids using a complex biochemical pathway known as the Stickland reaction (Figure 13.34)

© 2015 Pearson Education, Inc. Oxidation stepsReduction steps Alanine2 Glycine NAD + NADH Pyruvate, CoA Acetyl-CoA CoA PiPi Acetyl~P ADP NADH NAD + Acetate 2 ADP 2 Acetyl~P ATP 2 Substrate-level phosphorylation 2 Acetate ATP Figure 13.34

© 2015 Pearson Education, Inc. 13.13 Clostridial and Propionate Fermentations Secondary fermentation Fermentation of fermentation products Fermentation of ethanol plus acetate by Clostridium kluyveri Propionic acid fermentation (Figure 13.35) Propionibacterium and related prokaryotes produce propionic acid as a major fermentation product

© 2015 Pearson Education, Inc. 13.14 Fermentations Without Substrate-Level Phosphorylation Fermentations of certain compounds do not yield sufficient energy to synthesize ATP Catabolism of the compound can then be linked to ion pumps that establish a proton or sodium motive force Propionigenium modestum catabolizes succinate under strictly anoxic conditions (Figure 13.36a) Establishes a sodium motive force Sodium motive force forms ATP

© 2015 Pearson Education, Inc. 13.14 Fermentations Without Substrate-Level Phosphorylation Oxalobacter formigenes catabolizes oxalate and produces formate (Figure 13.36b) Formate is excreted from the cell Export of formate from the cell establishes a proton motive force Proton motive force forms ATP

© 2015 Pearson Education, Inc. 13.15 Syntrophy Syntrophy Process whereby two or more microbes degrade a substance neither can degrade alone Most syntrophic reactions are secondary fermentations Most reactions are based on interspecies hydrogen transfer H 2 production by one partner is linked to H 2 consumption by the other (Figure 13.37) Syntrophic reactions are important for the anoxic portion of the carbon cycle

© 2015 Pearson Education, Inc. 14.5 Syntrophy Syntrophy (cont'd) H 2 consumption affects the energetics of the reaction carried out by the H 2 producer, allowing the reaction to be exothermic (Figure 13.38)

© 2015 Pearson Education, Inc. IV. Anaerobic Respiration 13.16 Principles of Anaerobic Respiration 13.17 Nitrate Reduction and Denitrification 13.18 Sulfate and Sulfur Reduction 13.19 Acetogenesis 13.20 Methanogenesis 13.21 Other Electron Acceptors

© 2015 Pearson Education, Inc. 13.16 Principles of Anaerobic Respiration In anaerobic respiration, electron acceptors other than O 2 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 (Figure 13.39)

© 2015 Pearson Education, Inc. 13.16 Principles of Anaerobic Respiration In the assimilative metabolism of an inorganic compound (e.g., NO 3 –, SO 4 2–, CO 2 ), the reduced compounds are used in biosynthesis During anaerobic respiration, the reduction of inorganic compounds is called dissimilative metabolism because the reduced products are excreted

© 2015 Pearson Education, Inc. 13.17 Nitrate Reduction and Denitrification Inorganic nitrogen compounds are the most common electron acceptors in anaerobic respiration All products of nitrate reduction (denitrification) are gaseous (Figure 13.40) Denitrification is the main biological source of gaseous N 2

© 2015 Pearson Education, Inc. Figure 13.40 Nitrate reduction (Escherichia coli) Denitrification (Pseudomonas stutzeri) Nitrate reductase Nitrite reductase Nitric oxide reductase Nitrous oxide reductase Gases

© 2015 Pearson Education, Inc. 13.17 Nitrate Reduction and Denitrification Biochemical pathway for dissimilative nitrate reduction has been well studied (Figure 13.41) Enzymes of the pathway are repressed by oxygen

© 2015 Pearson Education, Inc. 13.18 Sulfate and Sulfur Reduction Inorganic sulfur compounds can be used as electron acceptors in anaerobic respiration Reduction of SO 4 2– to H 2 S proceeds through several intermediates and requires activation of sulfate by ATP (Figure 13.42)

© 2015 Pearson Education, Inc. 13.18 Sulfate and Sulfur Reduction During sulfate reduction, electron transport reactions lead to proton motive force formation (Figure 13.43) This drives ATP synthesis by ATPase Many different compounds can serve as electron donors in sulfate reduction Examples: H 2, organic compounds, phosphite Some sulfur-reducing bacteria can gain additional energy through disproportionation of sulfur compounds

© 2015 Pearson Education, Inc. 13.19 Acetogenesis Acetogens and methanogens use CO 2 as an electron acceptor in anaerobic respiration H 2 is the major electron donor for both groups of organisms (Figure 13.44)

© 2015 Pearson Education, Inc. Reduction of CO 2 to methyl group Formyl tetrahydrofolate Methyl tetrahydrofolate Methyl B 12 ATP Reduction of CO 2 to carbonyl group CO dehydrogenase Energy- conserving steps ATP Acetate ADP Acetyl-CoA ATP Na + motive force CoA Figure 13.45

© 2015 Pearson Education, Inc. 13.20 Methanogenesis H 2 is the major electron donor for methanogenesis (Figure 13.49) Additional electron donors exist Examples: formate, CO, organic compounds (Figure 13.50)

© 2015 Pearson Education, Inc. 13.20 Methanogenesis Autotrophy in methanogens occurs via the acetyl-CoA pathway Energy conservation in methanogenesis is linked to both proton and sodium motive forces (Figure 13.51)

© 2015 Pearson Education, Inc. Figure 13.51 Site of reduction in MPH red MPH ox Electron transport generates a proton motive force. Out In Hetero- disulfide reductase F 420-ox MPH red cyt b red F 420-red MPH ox cyt b ox 2e – ADP ATP

© 2015 Pearson Education, Inc. 13.21 Other Electron Acceptors Fe 3+, Mn 4+, ClO 3 –, and various organic compounds can serve as electron acceptors for bacteria (Figure 13.52) Fe 3+ is abundant in nature, and its reduction is a major form of anaerobic respiration

© 2015 Pearson Education, Inc. 13.21 Other Electron Acceptors Reduction of arsenate has been employed for cleanup of toxic wastes and groundwater (Figure 13.53) Halogenated compounds can also serve as electron acceptors via a process called reductive dechlorination (dehalorespiration)

© 2015 Pearson Education, Inc. 13.21 Other Electron Acceptors Organic electron acceptors Fumarate reduced to succinate Trimethyl amine oxide (TMAO) Dimethyl sulfoxide (DMSO) Halogenated organic compounds

© 2015 Pearson Education, Inc. 13.21 Other Electron Acceptors Proton reduction Pyrococcus furiosus Member of the Archaea Grows optimally at 100°C on sugars; small peptides as electron donors May have the simplest anaerobic respiration mechanism (Figure 13.54) Organism uses modified glycolysis, and protons in anaerobic respiration are linked to ATPase activity

© 2015 Pearson Education, Inc. 13.22 Aerobic Hydrocarbon Metabolism Oxygen is used as a direct reactant in certain biochemical reactions Oxygenases Enzymes that catalyze the incorporation of atoms of oxygen from O 2 into organic compounds Two classes: Monooxygenases: incorporate one oxygen atom (Figure 13.55a) Dioxygenases: incorporate both oxygen atoms

© 2015 Pearson Education, Inc. 13.22 Aerobic Hydrocarbon Metabolism Many microbes can use aliphatic and aromatic hydrocarbons as electron donors when growing aerobically Oxygenases are central enzymes in these biochemical reactions Aerobic degradation of aromatic compounds involves ring oxidation (Figure 13.56)

© 2015 Pearson Education, Inc. 13.23 Aerobic Methanotrophy Methylotrophs use compounds that lack C–C bonds as electron donors and carbon sources Methanotrophs are methylotrophs that use CH 4 The initial step in methanotrophy requires methane monooxygenase (MMO; Figure 13.57) In the MMO reaction, CH 4 is converted to CH 3 OH and H 2 O Other oxidation steps convert CH 3 OH to CO 2

© 2015 Pearson Education, Inc. 13.23 Aerobic Methanotrophy Methanotrophs are classified into two physiological groups that differ in the pathways invoked for assimilation of carbon into cell material Type I: ribulose monophosphate pathway (Figure 13.58b) Type II: serine pathway (Figure 13.58a) Both pathways assimilate carbon at the oxidation level of formaldehyde

© 2015 Pearson Education, Inc. 13.24 Anoxic Hydrocarbon Metabolism Aliphatic and aromatic hydrocarbons and organic compounds containing only carbon and hydrogen can be oxidized anaerobically (Figure 13.59) First step in degradation is the addition of oxygen to the molecule through the incorporation of fumarate Hydrocarbons are oxidized to intermediates that can be catabolized via the citric acid cycle

© 2015 Pearson Education, Inc. Hexane (C 6 ) Addition of fumarate 1-Methylpentylsuccinate (C 10 ) Removal of CO 2 and CoA transfer to form succinate Beta-oxidation (see Figure 13.55b) To NO 3 – or SO 4 2– reduction, generating a proton motive force Succinate (C 4 ) Fumarate( C 4 ) Acetyl-CoA Hexane catabolism Toluene catabolism Anaerobic benzoyl-CoA pathway (see Figure 13.60) To NO 3 – or SO 4 2– reduction, generating a proton motive force To anaerobic respiration Succinyl-CoA Activation with CoA CoA transfer Benzylsuccinate Fumarate addition Propionyl-CoA Figure 13.59

© 2015 Pearson Education, Inc. 13.24 Anoxic Hydrocarbon Metabolism Methane The simplest hydrocarbon Can be oxidized under anoxic conditions by a consortium containing sulfate-reducing bacteria and methanotrophic archaea (Figure 13.61)

PowerPoint ® Lecture Presentations prepared by John Zamora Middle Tennessee State University C H A P T E R © 2015 Pearson Education, Inc. Metabolic Diversity.

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PowerPoint ® Lecture Presentations prepared by John Zamora Middle Tennessee State University C H A P T E R © 2015 Pearson Education, Inc. Metabolic Diversity.

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