Beta-oxidation of fatty acids takes place in the cytosol

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A long-chain fatty acid is dehydrogenated to create a trans double bond between C2 and C3. This is catalyzed by acyl CoA dehydrogenase to produce trans-delta 2-enoyl CoA. This enzyme uses NAD as an electron acceptor. Thiolysis occurs between C2 and C3 alpha and beta carbons of 3-ketoacyl CoA. Thiolase enzyme catalyzes the reaction when a new molecule of coenzyme A breaks the bond by nucleophilic attack on C3.

This releases the first two carbon units, as acetyl CoA, and a fatty acyl CoA minus two carbons. The process continues until all of the carbons in the fatty acid are turned into acetyl CoA. Fatty acids are oxidized by most of the tissues in the body. However, some tissues such as the red blood cells of mammals which do not contain mitochondria , [5] and cells of the central nervous system do not use fatty acids for their energy requirements, [6] but instead use carbohydrates red blood cells and neurons or ketone bodies neurons only.

The enzyme catalyzes the formation of a double bond between the C-2 and C The reaction is stereospecific , forming only the L isomer. This converts the hydroxyl group into a keto group. The thiol is inserted between C-2 and C For every cycle, the Acyl CoA unit is shortened by two carbon atoms. Odd-numbered saturated fatty acids[ edit ] In general, fatty acids with an odd number of carbons are found in the lipids of plants and some marine organisms.

Many ruminant animals form a large amount of 3-carbon propionate during the fermentation of carbohydrates in the rumen. The bicarbonate ion's carbon is added to the middle carbon of propionyl-CoA, forming a D-methylmalonyl-CoA. However, the D conformation is enzymatically converted into the L conformation by methylmalonyl-CoA epimerase , then it undergoes intramolecular rearrangement, which is catalyzed by methylmalonyl-CoA mutase requiring B12 as a coenzyme to form succinyl-CoA.

The succinyl-CoA formed can then enter the citric acid cycle. However, whereas acetyl-CoA enters the citric acid cycle by condensing with an existing molecule of oxaloacetate, succinyl-CoA enters the cycle as a principal in its own right.

Thus the succinate just adds to the population of circulating molecules in the cycle and undergoes no net metabolization while in it. When this infusion of citric acid cycle intermediates exceeds cataplerotic demand such as for aspartate or glutamate synthesis , some of them can be extracted to the gluconeogenesis pathway, in the liver and kidneys, through phosphoenolpyruvate carboxykinase , and converted to free glucose. Complete beta oxidation of linoleic acid an unsaturated fatty acid.

As in the above case, this compound is converted into a suitable intermediate by 3,2-Enoyl CoA isomerase. To summarize: Odd-numbered double bonds are handled by the isomerase. Even-numbered double bonds by the reductase which creates an odd-numbered double bond Peroxisomal beta-oxidation[ edit ] Fatty acid oxidation also occurs in peroxisomes when the fatty acid chains are too long to be handled by the mitochondria.

The same enzymes are used in peroxisomes as in the mitochondrial matrix, and acetyl-CoA is generated. It is believed that very long chain greater than C fatty acids, branched fatty acids, [11] some prostaglandins and leukotrienes [12] undergo initial oxidation in peroxisomes until octanoyl-CoA is formed, at which point it undergoes mitochondrial oxidation.

Through three enzymatic reactions that need cofactors of biotin and vitamin B12, propionyl CoA may be transformed into succinyl CoA. Succinyl CoA can subsequently join the citric acid cycle. Activation of fatty acids Long-chain fatty acids are activated in the cytoplasm of the cell by ATP and coenzyme A, resulting in the formation of fatty acyl-CoA.

Mitochondria activate short-chain fatty acids. Fatty acid activation uses the equivalent of two ATP molecules, since two high-energy phosphate bonds are broken. Transport of fatty acyl-CoA from the cytosol into mitochondria In the outer mitochondrial membrane, carnitine and fatty acyl-CoA from the cytosol combine to produce fatty acylcarnitine. When fatty acylcarnitine reaches the inner membrane, it transforms back into fatty acyl-CoA and moves into the matrix. Malonyl-CoA, a by-product of the production of fatty acids, inhibits the enzyme carnitine acyltransferase I, which catalyses the transfer of acyl groups from coenzyme A to carnitine.

Malonyl-CoA hence limits the transfer of fatty acids into mitochondria during their cytosolic production, preventing an unnecessary cycle synthesis followed by immediate degradation. The beta-oxidation of the fatty acid acyl-CoA takes place within the mitochondrion.

Enoyl-CoA is created when a double bond forms between the — and -carbons. Lhydroxyacyl-CoA dehydrogenase is an enzyme that only reacts with the L-isomer of hydroxyacyl-CoA. A thiolase that needs coenzyme A breaks the connection between the alpha and beta carbons of the ketoacyl-CoA. The two carbons at the carboxyl end of the original fatty acyl-CoA are converted to acetyl-CoA, and a fatty acyl-CoA that is two carbons smaller than the original is made using the remaining carbons.

Seven times are performed on the palmitoyl-CoA, which has 16 carbons. For a combined output of roughly Each of the seven NADH produces roughly 2. Different quantities of ATP will be produced by the oxidation of various fatty acids.

Oxidation of odd-chain and unsaturated fatty acids Odd-chain fatty acids. Acetyl-CoA and propionyl-CoA are produced by odd-chain fatty acids. The four stages of the oxidation spiral are repeated by these fatty acids, creating acetyl-CoA up to the last cleavage.

At this moment, propionyl-CoA is formed from the three remaining carbons. Acetyl-CoA cannot be converted to glucose, only protonated-CoA can. Unsaturated fatty acids: Which account for approximately half of the fatty acid remains in human lipids, need additional enzymes in relation to the four enzymes that catalyse the repeated phases of the oxidation spiral.