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Metabolism

QuestionAnswer
Metabolic Principles: All microbes follow core rules: they conserve energy (make ATP), use reducing power (electrons for redox reactions), and maintain redox balance by passing electrons to external acceptors.
Metabolic Diversity: Microbial diversity comes from mixing and modifying metabolic reactions—horizontal gene transfer and enzyme variations create many pathways (e.g., different electron donors/acceptors).
Conservation of Energy (in Cells): The principle that cells conserve energy by linking the movement of electrons to the production of ATP.
Redox Reactions: Chemical reactions where electrons are transferred from an electron donor to an electron acceptor; these reactions drive electron flow in cells.
Reduction Potential (E₀′): A measure of a molecule’s tendency to donate or accept electrons; higher (more positive) values indicate a stronger tendency to accept electrons.
Energy Yield and E₀′ Difference: The greater the difference in reduction potential between paired half-reactions, the more energy is released and available for cellular work.
Electron Donors and Acceptors: Cells can use many organic or inorganic substances as electron donors, pairing them with a wide range of possible electron acceptors.
ATP Synthesis Pathways: Electron flow powers ATP production through three mechanisms — substrate-level phosphorylation, oxidative phosphorylation, or photophosphorylation.
Fermentation: A chemotrophic metabolism that requires no external electron acceptor. ATP is produced by substrate-level phosphorylation, and electron balance is maintained by reducing intermediates that are excreted as fermentation products.
Phototrophs: Organisms that obtain their energy from light.
Chemotrophs: Organisms that obtain their energy from chemical compounds.
Organotrophs: Organisms that use organic molecules as their electron source.
Lithotrophs: Organisms that use inorganic molecules as their electron source.
Heterotrophs: Organisms that use organic carbon as their carbon source.
Autotrophs: Organisms that use CO₂ as their carbon source.
Aerobic Respiration: Metabolism using O₂ as the electron acceptor; ATP is produced via oxidative phosphorylation driven by a proton motive force; electron donors can be organic (chemoorganotrophs) or inorganic (chemolithotrophs).
Chemolithotrophs: Organisms that generate energy by oxidizing inorganic compounds (e.g., H₂, NH₃, Fe²⁺) as electron donors, often using O₂ as the electron acceptor; energy yield varies by compound.
Anaerobic Respiration: Metabolism using electron acceptors other than O₂; yields less energy than aerobic respiration; dominates in oxygen-poor environments.
Reducing Power: Low-potential electron carriers (e.g., NAD(P)H, reduced ferredoxin) used for biosynthesis; generated by oxidizing organic compounds in chemoorganotrophs. Chemolithotrophs/phototrophs often need energy-coupled reactions to reduce NAD(P)⁺ or ferredoxin.
Redox Balance: Maintenance of electron flow in cells by transferring electrons to external acceptors (respiration) or intermediates (fermentation); ensures proper metabolism and availability of reducing power.
Three methods of energy coupling to increase reducing power or achieve redox balance: ATP hydrolysis coupling, reverse electron transport, electron bifurcation.
ATP Hydrolysis Coupling: Uses energy from ATP breakdown to drive otherwise difficult reactions and increase reducing power.
Reverse Electron Transport: Uses proton motive force to push electrons uphill and reduce NAD(P)⁺ when electron donors are weak.
Electron Bifurcation: Splits electrons from one donor to reduce both high- and low-potential acceptors at once, powered by flavin enzymes.
Bifurcating Enzyme Mechanism: Flavin-containing enzymes split electrons, sending one to a high-potential acceptor and one to a low-potential acceptor. This generates strong reducing intermediates and boosts fermentative energy yield by oxidizing NADH to NAD⁺, often producing H₂.
Assimilative Processes: incorporate inorganic nutrients into cells. Consume energy (ATP and reducing power), only done to acquire nutrients for biosynthesis, variety of assimilative reduction reactions.
What’s the most important Assimilative Process? CO2 fixation.
Dissimilative Processes: conserve energy, electron acceptors must be reduced and excreted, dissimilative reductions are part of anaerobic respiration.
Autotrophs & CO2: autotrophs can assimilate CO2 into cell material. Include many microbes, including almost all phototrophs and chemolitotrophs. CO2 supplies carbon for biosynthesis. In phototrophs, CO2 is the ultimate photosynthetic electron acceptor.
Mixotrophs: Organisms that can use both inorganic and organic sources of energy or carbon, switching depending on availability.
The Calvin Cycle: most widespread, globally important pathway for CO2 fixation, used by all oxygenic phototrophs. The key enzyme is RubisCO (reduces O2 to G3P), requires NADPH and ATP to synthesize 1 fructose-6-phosphate from CO2).
Carboxysomes & The Calvin Cycle: carboxysomes are proteinaceous microcompartments containing RubisCO. Carboxysome also protects RubisCO from O2, which competes with CO2. Cycle originated before the Great Oxidation Event.
Photosynthesis: use of light energy to drive biosynthesis, like sunlight and infrared radiation in the dark sea floor.
What kind of organisms do photosynthesis? Photosynthetic organisms are autotrophs (reduce CO2 to organic compounds → photoautotrophy), not all phototrophs are autotrophs, photoheterotrophs are phototrophs that use organic carbon as a carbon source.
Light Reactions: Convert light energy into ATP and proton motive force.
Light-Independent Reactions (Calvin Cycle): Fix CO₂ into organic compounds using ATP and reducing power from light reactions.
Oxygenic Photosynthesis: Uses H₂O as an electron donor, produces O₂, performed by cyanobacteria, algae, and plants.
Anoxygenic Photosynthesis: Uses donors other than H₂O (e.g., H₂S), does not produce O₂; light drives CO₂ reduction and ATP production.
Respiratory Processes Defined by Electron Donor: Electron donor (oxidized; energy source): Organic compounds, H₂, sulfur compounds, nitrogen compounds (nitrification, anammox), Fe²⁺
Respiratory Processes Defined by Electron Acceptor: O₂ (aerobic respiration), NO₃⁻/NO₂⁻ (denitrification), SO₄²⁻ (sulfate reduction), CO₂ (methanogenesis), Fe³⁺ (iron reduction)
Nitrification: Aerobic, two-step bacterial process that oxidizes ammonia (NH₃) to nitrite (NO₂⁻), then nitrite to nitrate (NO₃⁻). Performed by specialized chemolithotrophic bacteria.
Nitrifying Bacteria: Chemolithotrophic microorganisms that catalyze nitrification. Most are specialists (Nitrosomonas, Nitrobacter) performing only one oxidation step; some Nitrospira can perform both steps completely.
Nitrification Carbon Metabolism: Aerobic nitrifying bacteria fix CO₂ via the Calvin cycle, generating NAD(P)H through reverse electron flow.
Metabolic Flexibility of Nitrifiers: Nitrite oxidizers can grow chemoorganotrophically on organics. Ammonia oxidizers range from obligate chemolithotrophs to autotrophs/mixotrophs; archaeal ammonia oxidizers use a modified 3-hydroxypropionate/4-hydroxybutyrate cycle.
Denitrification: Anaerobic respiration using nitrate (NO₃⁻) as the electron acceptor, reduced to N₂. Some microbes fully reduce to N₂, others stop at nitrite. Releases N₂O and causes soil nitrogen loss.
Denitrifying Microorganisms: Mostly Proteobacteria (e.g., Pseudomonas) that are facultative aerobes; some Archaea and one eukaryote (Globobulimina pseudospinescens) also denitrify. Many can use alternative electron acceptors (Fe³⁺, organics) anaerobically or ferment.
Ecological Impact of Denitrification: Detrimental to agriculture (removes fertilizer nitrate), produces N₂O greenhouse gas and NO (consumes ozone), NO₂⁻ contributes to acid rain. Beneficial in sewage treatment by removing fixed nitrogen that would otherwise trigger algal blooms.
Methanogenesis: Anaerobic respiration where CO₂ (or small C compounds) is the terminal electron acceptor, producing methane (CH₄)
Washington-Paine Experiment The first U.S. scientific experiment showed that “magical mud” containing microbes could produce methane from organic matter, demonstrating that microbial activity, not just chemistry, drives methane production in anaerobic environments.
Methane Hydrate-Flammable Ice: A crystalline solid where methane is trapped in a water-ice lattice. It forms under high pressure and low temperature (deep ocean sediments or permafrost) and releases flammable methane when heated or depressurized.
Comparison Between Methane and CO2: methane is 84x more potent than CO2 in the short term.
Methanogenesis from CO2 + H2: H2 is the major electron donor for methanogenesis. Uses reductive acetyl-CoA pathway to assimilate CO2. Has two types with/without cytochromes.
Methanogenesis Without Cytochromes: relies heavily on electron bifurcation to generate reducing power. These organisms use H₂ to reduce CO₂ step-by-step, producing methane while recycling CoM and CoB.
Methanogenesis With Cytochromes: Some methanogens use cytochromes in their electron transport chain, creating a stronger H⁺ or Na⁺ gradient across the membrane and producing more ATP than methanogens without cytochromes.
Methanogenesis with Methanol and Acetate: methanol or acetate replaces H₂ + CO₂. Methanol provides electrons while methyl groups are reduced to CH₄. Strong H⁺/Na⁺ gradients increase ATP yield and allow reverse steps for extra energy.
Limitations with Fermentation: defined by lack of an external electron acceptor. Two major challenges are that it conserves much less energy than respiratory organisms, difficult to achieve redox balance.
Diversity of Fermentation: there’s tremendous reaction diversity. Many only ferment when lacking external electron acceptors anoxically. Many are more exclusively fermentative.
Achieving Redox Balance: Total atoms and electrons in reactants/substrates must equal products. Achieved by excreting fermentation products (acids, alcohols) and through H₂ production mechanisms.
Fermentations of Obligate Anaerobes: Many fermenters are obligate anaerobes, growing in highly reducing conditions and unable to tolerate O₂, often producing H₂. Ex: Clostridium species
Secondary Fermentation: Fermentation where products of one organism's fermentation serve as substrates for another organism. Common in microbial communities living in close association. Important in food production and environmental nutrient cycling.
Propionic Acid Fermentation: Used by Propionibacterium, converting lactate into propionate, acetate, and CO₂, generating ATP via substrate-level phosphorylation. Responsible for Swiss cheese holes and part of the human skin microbiome.
Syntrophy: Obligate cooperation between two different microbes to perform a thermodynamically unfavorable reaction that neither can do alone. Most syntrophic reactions are secondary fermentations involving interspecies electron transfer.
Interspecies Electron Transfer (IET): One species donates electrons and another accepts them. Types: Direct (DIET) via cell-to-cell contact or nanowires, and Mediated (MIET) via diffusing products like H₂.
Direct Interspecies Electron Transfer (DIET): A syntrophic process in which microbes pass electrons directly between cells through physical contact or conductive structures like nanowires/pili, allowing electron exchange without diffusible intermediates.
Mediated Interspecies Electron Transfer (MIET): A syntrophic process where microbes transfer electrons indirectly using diffusible electron carriers such as H₂ or formate, which shuttle electrons between the donor and the partner organism.
Ecology of Syntrophs: Syntrophs drive anoxic carbon cycling, consuming fermentation products and H₂ for anaerobic respiration. They function when methanogenesis or acetogenesis is the terminal process and are unnecessary in oxic conditions or with abundant electron acceptors.
Created by: smurtab
 

 



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