Citric Acid Cycle
Introduction to the Citric Acid Cycle
The citric acid cycle (CAC) occurs in the mitochondria following the entry of acetyl CoA, as well as fatty acid CoA or fatty acyl CoA.
Major products produced from this pathway are carbon dioxide (CO₂) and electron carriers NADH and FADH₂.
Overview of Initial Substrate and Product Formation
Initial Reaction:
Combines acetyl CoA with oxaloacetate (OAA).
This reaction is a condensation reaction and the catalyst for this process.
Product: Citrate (also known as citric acid).
Difference between citrate and citric acid:
Presence of hydronium on the carboxylic acids (at biological pH, citrate is deprotonated).
Notable Structure: Contains three carboxylic acid functional groups.
Alternative Terminology
Citric acid cycle is often referred to as TCA (tricarboxylic or tricarboxylate cycle).
Step-by-Step Mechanism of the Cycle
Reaction 1: Isomerization
Converts citrate to isocitrate through an isomerization process.
Enzyme Involved: Isomerase
Hydroxyl group moves from carbon 3 to carbon 2.
Significance: Facilitates the decarboxylation process (removal of CO₂) and enables the coupling of NADH production.
Water is used as an intermediate; it's removed as hydroxyl and added back in a different position.
Reaction 2: Oxidative Decarboxylation
Enzyme Involved: Isocitrate dehydrogenase
Process:
This reaction is identified as oxidative decarboxylation.
Products: NADH and CO₂.
Isocitrate is oxidized while NAD⁺ is reduced.
Final product is alpha-ketoglutarate.
Note: Two NADH produced for each glucose molecule; equivalently one NADH per acetyl CoA.
Reaction 3: Formation of Succinyl CoA
Conversion of alpha-ketoglutarate to succinyl CoA.
Enzyme Involved: Alpha-ketoglutarate dehydrogenase complex
Similar to the pyruvate dehydrogenase complex, coupling decarboxylation with NADH formation.
Additional product produced is CO₂.
This step is recognized as the second energy-generating reaction of CAC.
Total NADH produced thus far: Two per each acetyl CoA.
Reaction 4: Conversion to Succinate
Behavior: Removal of coenzyme A from succinyl CoA.
Product Formed: GTP through substrate-level phosphorylation (direct energy production).
GTP considered equivalent to ATP.
Enzyme Involved: Succinyl CoA synthetase.
Structural similarity noted between ATP and GTP; both are purines and share energy equivalence.
Reaction 5: Reduction of Fumarate
This reaction involves producing FADH₂.
Produced through the formation of a carbon-carbon double bond.
CO-Factor: FAD (Flavin adenine dinucleotide) is essential when producing carbon-carbon double bonds, in contrast to NADH, which is used to form carbonyl compounds.
Reaction 6: Addition of Water
Reaction: Water is added to incorporate an alcohol group, producing malate.
Malate is noted to have an L isomer, which is atypical among biologically active molecules.
Reaction 7: Oxidation of Malate
Enzyme Involved: Malate dehydrogenase
Process:
Oxidation of malate occurs here, removing two hydrogens.
Final product being oxaloacetate, thus completing the cycle.
Total products from one acetyl CoA:
Three NADH, one FADH₂, and one GTP.
Conclusion of Energy Yields
The outputs from the citric acid cycle will be analyzed to determine ATP equivalents produced from both glucose and fatty acids as the discussion moves forward.
Highlighting the cyclic nature of the citric acid cycle, oxaloacetate condenses with acetyl CoA again at the cycle's initiation, symbolizing the completion and renewal of the cycle.