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1 number 23 Done by A. Rawajbeh Corrected by Doctor F. Al-Khateeb

2 Ketone bodies Ketone bodies are used by the peripheral tissues like the skeletal and cardiac muscles, where they are the preferred source of energy, and by brain during prolonged fasting. So what is the advantage of using ketone bodies by the brain during prolonged fasting? Before fasting, glucose concentration is about mmol/l, and after 2-3 days, blood glucose is still high at about 4mmol/L (70% of its normal level). This maintenance of glucose level is achieved by gluconeogenesis. Triacylglycerols: cannot be used to produce glucose because only glycerol (which represents a small part of the TAG molecule) can be converted to DHAP (an intermediate in gluconeogenesis), TAGs are not considered as resources for glucose carbons.

3 Proteins: are used to nourish the intermediates of gluconeogenesis after they are degraded to smaller amino acids [except leucine and lysine]. BUT Proteins aren t stored to be used as a source of energy, they have their own functions in our bodies. Thus, using proteins as fuels is not good during fasting. Ketone bodies: they are found in the blood with increasing levels during fasting as the insulin/glucagon ratio decreases and catecholamines are secreted (they activate degradation of TAGs by lipases in white adipose tissues releasing free fatty acids that would be converted to ketone bodies by the liver). Ketones bodies level increases in the blood, so the brain starts to use it as a source of energy, that means that the brain starts depending on fatty acids as source of energy instead of proteins. So, proteins are preserved to do their functions. After 40 days of continuous fasting, the brain has decreased its consumption of glucose by 40-60% The numbers in the figure are not for memorization.

4 Adaptation of the brain and the muscles to use ketone bodies as a source of energy would preserve proteins so this is an advantage. **************************************************************************** Fatty acid synthesis **FA synthesis occurs mainly in the liver **FA synthesis requires carbon source (acetyl-coa), reducing power )NADPH) and energy input (ATP). -From where are acetyl CoA provided for fatty acid synthesis? >>from glucose metabolism Why is NADPH needed? >> Degradation of fatty acids to acetyl-coa is oxidative so logically their synthesis is reductive, so we need reducing power for this biosynthetic pathway, this is provided by NADPH.

5 Why energy? >>because we build larger molecules from smaller ones. Just like we get energy from fatty acids oxidation, fatty acid synthesis needs energy. Fatty acid synthesis from acetyl CoA molecules (condensation reaction between acyl units) has a high positive free energy change, we can use ATP molecules to provide energy making the synthesis favorable. {this occurs by carboxylation>decarboxylation mechanism as you see in gluconeogenesis [pyruvate>oaa>pep>>>glucose]} **The general reactions in FA synthesis and FA oxidation are just opposite to each other. Steps of FA synthesis: IT OCCURS IN THE CYTOSOL 1-Acetyl CoA is converted to malonyl CoA to activate synthesis. **By acetyl CoA carboxylase **Malonyl CoA is a three-carbon molecule containing two carboxyl groups [same as succinyl which is a 4-carbon molecule]. **Carboxylation needs CO2, biotin, and ATP.

6 2-FATTY ACYL SYNTHASE (FAS) **large molecule, has 7 active sites(domains) so it is a multifunctional enzyme. ** homo-dimeric, composed of two polypeptides. [Each one alone is inactive. When they form the dimer structure, they become active.] **catalyzes the reactions more efficiently, as the intermediates are transferred from an active site to the next one in the same protein. Two important domains: -One domain is known as the condensing enzyme, it contains an SH group and thus can form a thioester bond with the acyl molecules -One domain is called the Acyl Carrier Protein ACP: *has phosphopantetheine, a derivative of pantothenic acid (B5) with reactive SH group, that forms thioester bonds with acyl molecules. *The difference between the two domains is that the first one catalyzes the condensation reaction and the second carries acyl units and presents them to the other catalytic domains. *Phosphpantotheine is similar to CoA. BUT instead of a phosphorylated ADP, it is linked to the ACP protein (only one of the two phosphates is removed). SO it likes a very large CoA

7 #FAS enzyme is not found in bacteria, instead many enzymes catalyze the FA synthesis. A)-Acetyl CoA binds to ACP SH group then it is transferred to the condensing enzyme SH group. Malonyl-CoA molecule binds to ACP SH group. The condensation reaction occurs between acetyl CoA and malonyl CoA releasing a CO2 (the CO2 we used in carboxylation why? to produce a high energy level in malonyl to prepare it for condensation, the break of this bond (the CH₂- COO bond) releases the needed energy for this buildup reaction to proceed). Another energy source for this condensation reaction is the cleavage of the thioester bond. So, the total reaction has negative free energy and it is spontaneous(favorable). >> Ketoacyl ACP (carried by ACP) is produced B)-Then, it is reduced by NADPH, converting the keto group into a hydroxyl group. C)-Then, dehydration occurs forming a double bond. D)-Then, another reduction reaction breaks the double bond consuming another NADPH molecule.

8 Now we have a 4-carbon acyl (butyric acid) attached to ACP>>butyryl- ACP The cycle is repeated by transferring the 4-carbon acyl to the condensing domain SH, then, binding of new malonyl CoA to ACP-SH, then, THE condensation and the subsequent reactions occur again, adding 2 carbons in each cycle, so we only synthesis fatty acids with even number of carbons. The cycle is repeated until we have a 16-carbon acyl (palmitoyl ACP). Palmitoyl thioesterase domain cleaves the thioester bond between palmitoyl and the S of ACP. Now the most important questions: -How many cycles are required to produce palmitate(16-carbons)? -How many malonyl CoA molecules are required? -How many acetyl CoA molecules are required? -How many acetyl CoA is required totally? -How many NADPH is required? >>> <<< NOTE THAT: 8 Acetyl CoA molecules are needed totally: 7 are converted to malonyl-coa and one is used as such as acetyl CoA. Production of cytosolic acetyl CoA

9 **FA synthesis occurs in the cytosol while oxidation occurs in the mitochondria, so they are separated. **The inner mitochondrial membrane (IMM) is impermeable to acetyl CoA. **Acetyl CoA is produced in the mitochondria from pyruvate and fatty acid oxidation. **Acetyl-CoA and oxaloacetate condensate into Citrate [IMM is permeable to citrate] by citrate synthase. **Citrate completes the cycle if energy is required and if not it accumulates then it is released to the cytosol [NADH/NAD+ ratio determines that]. **In the cytosol, citrate is cleaved into acetyl CoA and OAA by ATP citrate lyase [We need energy to produce the high energy thioester bond (from CoA), so ATP is needed]. **OAA cannot cross the IMM but it is converted to malate by cytosolic malate dehydrogenase. **Then malate can either enter into the mitochondria or undergo an oxidative decarboxylation (by NADP-dependent malate dehydrogenase) and get converted to pyruvate reducing one NADP into NADPH, this is what usually happens because the enzyme quickly binds to malate before it enters the mitochondria. **These NADPH molecules are used in FA synthesis. Note: OAA is converted to Malate by Oxidizing NADH and then Malate is converted to Pyruvate by Reducing NADP+.

10 OAA s structure in the diagram is wrong. OAA is OOC-CO-CH₂-COO

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