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• In cells, electrons are the most important source of chemical potential energy.
• The amount of potential energy in an electron is based on its position relative to positive and negative charges.
– Electrons closer to negative charges (from other electrons) and farther from positive charges (in nuclei of nearby atoms), have higher potential energy.
• In general, a molecule’s potential energy is a function of its electrons’ configuration and position.
• During a redox reaction, electrons can be transferred completely from one atom to another, or they can simply shift their position in covalent bonds.
• The carbon atoms of glucose are oxidized to form carbon dioxide, and the oxygen atoms in oxygen are reduced to form water:
C6H12O6 + 6 O2 6 CO2 + 6 H2O + energy
glucose oxygen carbon water dioxide
In cells, glucose is oxidized through a long series of carefully controlled redox reactions. The resulting change in free energy is used to synthesize ATP from ADP and Pi. Together, these reactions comprise cellular respiration.
• All organisms use glucose to build fats, carbohydrates, and other compounds; cells recover glucose by breaking down these molecules.
– Glucose is used to make ATP through either cellular respiration or fermentation.
Cellular respiration produces ATP from a molecule with high potential energy – usually glucose. Each of the four steps of cellular respiration consists of a series of chemical reactions, and a distinctive starting molecule and characteristic set of products.
• Glycolysis, a series of 10 chemical reactions, is the first step in glucose oxidation.
• All of the enzymes needed for glycolysis are found in the cytosol.
• In glycolysis, glucose is broken down into two 3-carbon molecules of pyruvate, and the potential energy released is used to phosphorylate ADP to form ATP.
• Substrate-level phosphorylation occurs when ATP is produced by the enzyme-catalyzed transfer of a phosphate group from an intermediate substrate to ADP.
– This is how ATP is produced in glycolysis and the citric acid cycle.
• In an electron transport chain a proton gradient provides energy for ATP production; the membrane protein ATP synthase uses this energy to phosphorylate ADP to form ATP. This process is called oxidative phosphorylation.
• During glycolysis, high levels of ATP inhibit the enzyme phosphofructokinase, which catalyzes one of the early reactions.
• Phosphofructokinase has two binding sites for ATP:
1. The active site, where ATP phosphorylates fructose-6-phosphate, resulting in the synthesis of fructose-1,6-bisphosphate
2. A regulatory site
High ATP concentrations cause ATP to bind at the regulatory site, changing the enzyme’s shape and dramatically decreasing the reaction rate at the active site.
Pyruvate processing is under both positive and negative control. Abundant ATP reserves inhibit the enzyme complex; large supplies of reactants, such as acetyl CoA and NADH, and low supplies of products, such as ATP, stimulate it.
• During the third step of glucose oxidation, the acetyl CoA produced by pyruvate processing enters the citric acid cycle, located in the mitochondrial matrix.
– Each acetyl CoA is oxidized to two molecules of CO2.
• Some of the potential energy released is used to
1. Reduce NAD+ to NADH.
2. Reduce flavin adenine dinucleotide (FAD) to FADH2 (another electron carrier).
3. Phosphorylate GDP to form GTP (later converted to ATP).
• A series of carboxylic acids is oxidized and recycled in the citric acid cycle.
• Citrate (the first molecule in the cycle) is formed from pyruvate and oxaloacetate (the last molecule in the cycle).
• The citric acid cycle completes glucose oxidation. The energy released by the oxidation of one acetyl CoA molecule is used to produce 3 NADH, 1 FADH2, and 1 GTP, which is then converted to ATP.
The citric acid cycle can be turned off at multiple points, via several different mechanisms of feedback inhibition.
To summarize, the citric acid cycle starts with acetyl CoA and ends with CO2. The potential energy that is released is used to produce NADH, FADH2, and ATP. When energy supplies are high, the cycle slows down.
• For each glucose molecule that is oxidized to 6 CO2, the cell reduces 10 molecules of NAD+ to NADH and 2 molecules of FAD to FADH2, and produces 4 molecules of ATP by substrate-level phosphorylation.
• The ATP can be used directly for cellular work.
• However, most of glucose’s original energy is contained in the electrons transferred to NADH and FADH2, which then carry them to oxygen, the final electron acceptor.
• During the fourth step in cellular respiration, the high potential energy of the electrons carried by NADH and FADH2 is gradually decreased as they move through a series of redox reactions.
• The proteins involved in these reactions make up what is called an electron transport chain (ETC).
• O2 is the final electron acceptor. The transfer of electrons along with protons to oxygen forms water.
• The energy released as electrons move through the ETC is used to pump protons across the plasma membrane into the intermembrane space, forming a strong electrochemical gradient.
• The protons then move through the enzyme ATP synthase, driving the production of ATP from ADP and Pi.
• Because this mode of ATP production links the phosphorylation of ADP with NADH and FADH2 oxidation, it is called oxidative phosphorylation.
• Most of the ETC molecules are proteins containing chemical groups that facilitate redox reactions. All but one of these proteins are embedded in the inner mitochondrial membrane.
– In contrast, the lipid-soluble ubiquinone (Q) can move throughout the membrane.
• During electron transport, NADH donates electrons to a flavin-containing protein at the top of the chain, but FADH2 donates electrons to an iron-sulfur protein that passes electrons directly to Q.
• The ETC pumps protons from the mitochondrial matrix to the intermembrane space. The proton-motive force from this electrochemical gradient can be used to make ATP in a process known as chemiosmosis.
• ETC proteins are organized into four large multiprotein complexes (called complex I–IV) and cofactors. Protons are pumped into the intermembrane space from the mitochondrial matrix by complexes I and IV.
• Q and the protein cytochrome c transfer electrons between complexes.
• The vast majority of the “payoff” from glucose oxidation occurs via oxidative phosphorylation; ATP synthase produces 25 of the 29 ATP molecules produced per glucose molecule during cell respiration.
• Oxygen is the most effective electron acceptor because it is highly electronegative. There is a large difference between the potential energy of NADH and O2 electrons which allows the generation of a large proton-motive force for ATP production.
• Cells that do not use oxygen as an electron acceptor cannot generate such a large potential energy difference. Thus, they make less ATP than cells that use aerobic respiration.
• In most organisms, cellular respiration cannot occur without oxygen. Fermentation, a metabolic pathway that regenerates NAD+ from stockpiles of NADH, allows glycolysis to continue producing ATP in the absence of oxygen.
• Fermentation occurs when pyruvate or a molecule derived from pyruvate accepts electrons from NADH.
• This transfer of electrons oxidizes NADH to NAD+.
– With NAD+ present, glycolysis can continue to produce ATP via substrate-level phosphorylation.
• In lactic acid fermentation, pyruvate produced by glycolysis accepts electrons from NADH. Lactate and NAD+ are produced.
– Lactic acid fermentation occurs in muscle cells.
• In alcohol fermentation, pyruvate is enzymatically converted to acetaldehyde and CO2. Acetaldehyde accepts electrons from NADH. Ethanol and NAD+ are produced.
• Fermentation is extremely inefficient compared with cellular respiration.
– Fermentation produces just two ATP molecules per glucose molecule, compared with about 29 ATP molecules per glucose molecule in cellular respiration.
– Consequently, organisms never use fermentation if an appropriate electron acceptor is available for cellular respiration.
Cellular Respiration Interacts with Metabolic Pathways
• Energy and carbon are cells’ two fundamental requirements.
– They need high-energy electrons for generating chemical energy in the form of ATP, and a source of carbon-containing molecules for synthesizing macromolecules.
• Metabolism includes thousands of different chemical reactions.
– Catabolic pathways involve the breakdown of molecules and the production of ATP.
– Anabolic pathways result in the synthesis of larger molecules from smaller components.
• Molecules found in carbohydrate metabolism are used to synthesize macromolecules such as RNA, DNA, glycogen or starch, amino acids, fatty acids, and other cell components.