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Essential knowledge 2.A.1:
All living systems require constant input of free energy a. Life requires a highly ordered system. b. Living systems do not violate the second law of thermodynamics, which states that entropy increases over time. c. Energy-related pathways in biological systems are sequential and may be entered at multiple points in the pathway. d. Organisms use free energy to maintain organization, grow and reproduce. e. Changes in free energy availability can result in changes in population size. f. Changes in free energy availability can result in disruptions to an ecosystem.
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Essential knowledge 2.A.2:
Organisms capture and store free energy for use in biological processes. a. Autotrophs capture free energy from physical sources in the environment b. Heterotrophs capture free energy present in carbon compounds produced by other organisms. c. Different energy-capturing processes use different types of electron acceptors. d. The light-dependent reactions of photosynthesis in eukaryotes involve a series of coordinated reaction pathways that capture free energy present in light to yield ATP and NADPH, which power the production of organic molecules.
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Essential knowledge 2.A.2:
Organisms capture and store free energy for use in biological processes. e. Photosynthesis first evolved in prokaryotic organisms; scientific evidence supports that prokaryotic (bacterial) photosynthesis was responsible for the production of an oxygenated atmosphere; prokaryotic photosynthetic pathways were the foundation of eukaryotic photosynthesis. f. Cellular respiration in eukaryotes involves a series of coordinateenzyme-catalyzed reactions that harvest free energy from simple carbohydrates. g. The electron transport chain captures free energy from electrons in a series of coupled reactions that establish an electrochemical gradient across membranes. h. Free energy becomes available for metabolism by the conversion of ATP→ADP, which is coupled to many steps in metabolic pathways.
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Cellular Respiration Usually involves breakdown of high energy glucose to low energy carbon dioxide and water (exergonic process) Energy extracted from glucose molecule: Released step-wise Allows ATP to be produced efficiently Oxidation-reduction enzymes include NAD+ and FAD as coenzymes
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Oxidation of Organic Fuel Molecules During Cellular Respiration
Glucose is oxidized and oxygen is reduced (Removes hydrogen) C6H12O6 + 6O CO2 + 6H2O + Energy becomes oxidized becomes reduced (Adds hydrogen) *oxygen is more electronegative than carbon, that means transfer of electrons releases energy to form more stable compounds with less free energy
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Cells mostly use glucose but also “burn” other organic molecules
Respiration releases the chemical bond energy of glucose, which is stored by the cell in the chemical bonds of ATP
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Cells typically store ~39-40% of glucose energy in ATP molecules - the rest is lost as heat (aerobic resp) Anaerobic resp. harvests only ~2% of energy in glucose
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NAD+ and FAD NAD+ (nicotinamide adenine dinucleotide)
Called a coenzyme of oxidation-reduction it can Oxidize a metabolite by accepting electrons Reduce a metabolite by giving up electrons Each NAD+ molecule used over and over again FAD (flavin adenine dinucleotide) Also a coenzyme of oxidation-reduction Sometimes used instead of NAD+
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NAD+ (nicotinamide adenine dinucleotide) is a coenzyme that cells make from vitamin niacin
NAD+ is used to shuttle electrons NAD+ removes 2 H+ and 2 e-, it keeps 1 H+ and 2 e- to become NADH or becomes reduced to become NADre
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Cellular Respiration: Overview of 4 Phases
1.Glycolysis: Occurs in cytoplasm Glucose broken down to two molecules of pyruvate ATP, NADH is formed 2.Preparatory reaction: Both pyruvates are oxidized to acetyl CoA Electron energy is stored in NADH Two carbons are released as CO2 3.Citric acid or Krebs cycle: Occurs in mitochondria Electron energy is stored in NADH and FADH2 ATP is formed
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4.Electron transport chain and chemiosmosis:
occurs on inner membrane of mitochondria Extracts energy from NADH & FADH2 * Both glycolysis and the citric acid cycle are exergonic and make small amounts of ATP (2 +2) by substrate level phosphorylation, the main function of glycolysis and Krebs is to supply last stage with electrons
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Glycolysis harvests energy by oxidizing glucose to pyruvate Glycolysis
Means “splitting of sugar”
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Glucose Breakdown: Glycolysis
Occurs in cytoplasm outside mitochondria Energy Investment Steps: Two ATP are used to activate glucose Glucose splits into two G3P molecules Energy Harvesting (payoff) Steps: Two electrons (as hydrogen atoms) are picked up by two NAD+ and converted to NADH Four ATP produced by substrate-level phosphorylation Net gain of two ATP Both G3Ps converted to pyruvates
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Energy investment phase
2 ATP used
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Energy payoff phase 4 ATP produced
2 used previously resulting in a net gain of 2 ATP molecules 2 NADH also produced
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Before the citric acid cycle can begin
Preparatory stage Before the citric acid cycle can begin Pyruvate must first be converted to acetyl CoA, which links the cycle to glycolysis
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Conversion of (2) pyruvate to (2) acetyl CoA
Preparatory stage Conversion of (2) pyruvate to (2) acetyl CoA Produces (2) NADH Releases (2) CO2
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Glucose Breakdown: The Preparatory Reaction
End product of glycolysis, pyruvate, enters the mitochondrial matrix Pyruvate converted to 2-carbon acetyl group Attached to Coenzyme A to form acetyl-CoA Electron picked up (as hydrogen atom) by NAD+
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Citric Acid cycle The citric acid cycle completes the energy-yielding oxidation of organic molecules The citric acid cycle
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Acetyl CoA joins with oxaloacetic acid to form citric acid**
For 2 Acetyl CoA (from 1 glucose) 4 CO2 released 2 ATP produced (slpp) 2 FADH2 produced (flavin adenine dinucleotide) another electron carrier molecule, similar to NADH
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Total for 1 glucose from beginning of glycolysis to end of Krebs cycle :
4 ATP 10 NADH 2 FADH2 6 CO2 released
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During oxidative phosphorylation, chemiosmosis couples electron transport to ATP synthesis
NADH and FADH2
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Stepwise Energy Harvest via NAD+ and the Electron Transport Chain
Cellular respiration Oxidizes glucose in a series of steps
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Electrons from organic compounds
Are usually first transferred to NAD+, a coenzyme
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NADH, the reduced form of NAD+, carries electrons to electron carrier molecules, which passes the electrons to the electron transport chain Electrons travel “down” the electron transport chain, where each electron carrier molecule in the chain is successively “lower” in energy
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Most of these carrier molecules are proteins which are embedded in the mitochondria membrane
The final and ultimate electron acceptor of electron transport chains is oxygen Cells use energy released in small amounts by redox rxns to produce ATP
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The electron transport chain
Passes electrons in a series of steps instead of in one explosive reaction
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Both glycolysis and the citric acid cycle
Can generate ATP by substrate-level phosphorylation
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The Pathway of Electron Transport
In the electron transport chain Electrons from NADH and FADH2 lose energy in several steps NADH and FADH2 capture electrons and transport them to “top” of ETC ETC uses “downhill” flow of electrons from NADH and FADH2 to pump H+ ions across membrane Each molecule of ETC will have successively lower energy levels Enzymes of ETC and ATP synthase are built into the inner membrane of mitochondria
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At the end of the chain Electrons are passed to oxygen, forming water
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Chemiosmosis: The Energy-Coupling Mechanism
ATP synthase Is the enzyme that actually makes ATP
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At certain steps along the electron transport chain
Electron transfer causes protein complexes to pump H+ from the mitochondrial matrix to the intermembrane space
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The resulting H+ gradient
Stores energy Drives chemiosmosis in ATP synthase
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Chemiosmosis Is an energy-coupling mechanism that uses energy in the form of a H+ gradient across a membrane to drive cellular work
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Electron Transport Chain and Chemiosmosis
H+ ions pumped across inner membrane (into intermembrane space) create an electrostatic gradient Because inner membrane pumps positive ions across membrane, inner surface becomes more negatively charged Increase of H+ ions creates osmotic concentration gradient
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Combined effects of osmotic concentration gradient, and electrostatic gradient create a powerful chemiosmotic gradient Because of strong chemiosmotic gradient, H+ ions move strongly back across membrane into matrix through special protein channel called ATP synthase
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NADH starts at the “top” of the electron transport chain, it’s electrons pass through three H+ ion pump/protein complexes FADH2 starts lower on the ETC, and its electrons only pass through two H+ ion pump/protein complexes
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2 H+ ions passing back across ATP synthase produces 1 ATP molecule
Since NADH transports across 6 H+ ions, it generates enough energy to produce ~ 3ATP/NADH
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Glycolysis + Krebs produce :
10 NADH X ~ 3ATP = ATP (COPP) 2 FADH2 X ~2ATP = 4 ATP (COPP) 32-34 ATP + 2 ATP – glycolysis (SLPP) + 2 ATP – Krebs (SLPP) 36-38 ATP
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An Accounting of ATP Production by Cellular Respiration
During respiration, most energy flows in this sequence Glucose to NADH to electron transport chain to proton-gradient to ATP
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About 39% of the energy in a glucose molecule
Is transferred to ATP during cellular respiration, making approximately ATP
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A Comparison of Chemiosmosis in Chloroplasts and Mitochondria
Generate ATP by the same basic mechanism: chemiosmosis
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Oxidative phosphorylation - oxidation of compounds provides energy to phosphorylate ATP - results in net loss of energy by organism (ETC of respiration) Photophosphorylation (PPP) - light provides energy for phosphorylation - organism increase in energy (ETC of photosynthesis)
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The spatial organization of chemiosmosis
Differs in chloroplasts and mitochondria
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In both organelles ATP synthase
Redox reactions of electron transport chains generate a H+ gradient across a membrane ATP synthase
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In the absence of oxygen
Fermentation enables some cells to produce ATP without the use of oxygen Cellular respiration Relies on oxygen to produce ATP In the absence of oxygen
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Glycolysis Can produce ATP with or without oxygen, in aerobic or anaerobic conditions Couples with fermentation to produce ATP
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Fermentation consists of
Types of Fermentation Fermentation consists of Glycolysis plus reactions that regenerate NAD+, which can be reused by glycolysis
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In alcohol fermentation
Pyruvate is converted to ethanol in two steps, one of which releases CO2
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During lactic acid fermentation
Pyruvate is reduced directly to NADH to form lactate (lactic acid) as a waste product Lactic acid causes muscle ache during exercise Used to make cheese and yogurt
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Fermentation and Cellular Respiration Compared
Both fermentation and cellular respiration Use glycolysis to oxidize glucose and other organic fuels to pyruvate
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Cellular respiration Produces more ATP = 36-38 compared to 2
18-19X more efficient
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Pyruvate is a key juncture in catabolism
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The Evolutionary Significance of Glycolysis
Probably evolved in ancient prokaryotes before there was oxygen in the atmosphere
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Glycolysis and the citric acid cycle connect to many other metabolic pathways
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The Versatility of Catabolism
Catabolic pathways Funnel electrons from many kinds of organic molecules into cellular respiration
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Biosynthesis (Anabolic Pathways)
The body Uses small molecules to build other substances These small molecules May come directly from food or through glycolysis or the citric acid cycle
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Metabolic Pool: Catabolism (1)
Foods: Sources of energy rich molecules Carbohydrates, fats, and proteins Catabolism (breakdown side of metabolism) Breakdown products enter into respiratory pathways as intermediates Carbohydrates Processed as above
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Metabolic Pool: Catabolism (2)
Breakdown products enter into respiratory pathways as intermediates (cont.) Proteins Broken into amino acids (AAs) Some AAs used to make other proteins Excess AAs deaminated (NH2 removed) in liver Results in poisonous ammonia (NH3) Quickly converted to urea Different R-groups from AAs processed differently
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Metabolic Pool: Anabolism (1)
All metabolic reactions part of metabolic pool Intermediates from respiratory pathways can be used for anabolism Anabolism (build-up side of metabolism): Carbs: Start with acetyl-CoA Basically reverses glycolysis (but different pathway) Fats G3P converted to glycerol Acetyls connected in pairs to form fatty acids
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Metabolic Pool: Anabolism (2)
Anabolism (cont.): Proteins: Made up of combinations of 20 different amino acids Some amino acids (11) can be synthesized from respiratory intermediates organic acids in citric acid cycle can make amino acids Add NH2 – transamination However, other amino acids (9) cannot be synthesized by humans Essential amino acids
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