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FA + GLYCEROL • • • GLYCEROL o Gluconeogenesis o FA synthesis CARNITINE ACYLTRANSFERASE II – located in the INNER MITOCHONDRIAL MEMBRANE BETA OXIDATION • FA -> Acetyl CoA • 4 steps: FA o o Oxidation o Storage o Free w/ albumin OXIDATION (1ST) Oxidation of FA by Acyl CoA Dehydrogenase Forms a double bond between C2 & C3 Produces FADH2 from FAD LIPOPROTEIN LIPASE – enzyme FREE FATTY ACIDS – virtually all cells can use Fatty Energy except Brain, RBC & Medulla BETA OXIDATION • Major pathway for catabolism of FA • 2 carbon fragments are cleaved producing Acetyl CoA (enters Kreb’s Cycle) o HYDRATION o OXIDATION (2ND) STEPS: Hydroxyacyl CoA oxidized by NAD+ Became 3-Ketoacyl CoA Forms NADH 1. TRANSPORT OF FA INTO MITOCHONDRIA • After FA are taken up by a cell, it is converted to a CoA derivative (fatty acylCoA) o • Fatty aclyCoA synthetase/thiokinase • Transport to mitochondria THIOLYTIC CLEAVAGE (THIOLYSIS) *Mitochondrial membrane is impermeable Cleavage of 3-Ketoacyl CoA by thiol group of another CoA Produces shorter Acyl CoA and Acetyl CoA CARNITINE SHUTTLE • Acyl group must first be transferred Cytosolic CoA to Carnitine • Catalyzed by Carnitine Acyltransferase I • Forming Fatty Acyl Carnitine I & II • Inhibited by: MALONYL CoA CARNITINE ACYLTRANSFERASE I – located in the OUTER MITOCHONDRIAL MEMBRANE Add H2O ENERGY YIELD FROM BETA OXIDATION • • Each time an acetyl group (2 carbons) is cleaved from acyl chain, it produces: o 1 NADH o 1 Acetyl CoA o 1 FADH2 Therefore: *Carnitine is not needed inside the mitochondria because it is only a shuttle o PALMITATE (16:0) 16/2 = 8 – 1 = 7cycles * Fats yield high energy • O.H.O.T. FORMULA = (n/2) – 1 PALMITATE – 7 CYCLES o 7 NADH x3ATP = 21 7 FADH2 x2ATP = 14 RELEASE OF FA FROM TAG 8Acetyl CoA x 12ATP = 96 131 Energy investment FAcyl CoA synthase _-2 Total ATP 129 • Mobilization of STORED FAT • HORMONE – SENSITIVE LIPASE o Breaks stored fat o Present in ADIPOSE TISSUES 18 carbon = 8cycles o 8 NADH x3ATP = 24 8 FADH2 x2ATP = 16 TAG FA + GLYCEROL 9 Acetyl CoA x12ATP = 108 HORMONE – SENSITIVE LIPASE 148 – 2 = 146 • OXIDATION OF ODD NUMBERED FA • Same process until last 3 is reached • PROPIONYL CoA instead of Acetyl CoA • Propionyl CoA -> Methyl Malonyl CoA by enzyme Propionyl CoA Carboxylase • Methylmalonyl CoA -> Succinyl CoA by enzyme Methylmalonyl CoA Mutase • Activators: o Epinephrine o Low level glucose Reactivators: o Insulin o High level glucose FATE OF GLYCEROL • Succinyl CoA – can enter Kreb’s Cycle • Glycerol cannot be metabolized by adipocytes • COMPUTATION: • Lacks GLYCEROL KINASE transport to liver • Phosphorylated or converted to DHAP • Can participate in glycolysis/gluconeogenesis o 17 carbons – 3= 14/2= 7-1 = 6cycles 6 NADH x3ATP = 18 6 FADH2 x2ATP = 7 Acetyl CoA x12ATP = 84 12 114 FA SYNTHESIS – can be created from Carbohydrates & proteins • Location: primarily in the LIVER; kidney, lactating mammary glands, adipose tissues • Required energy: -2 112 Kreb’s Cycle (Succinyl CoA) 118 • o +6 6 from Kreb’s cycle is constant for ODD • NADPH, Acetyl CoA, ATP MAJOR SOURCES OF NADPH o HMP shunt o • NADP dependent MALATE HYDROGENASE Byproduct: NADPH Convert Malate to Pyruvate o Occurs in cytosol: FA synthesis o Acetyl CoA inside mitochondria o CITRATE SHUTTLE OAA + Acetyl CoA to produce CITRATE Enzyme: ATP CITRATE LYASE – removes citrate 2. CARBOXYLATION OF ACETYL CoA TO MALONYL CoA o Outside the cytosol o Acetyl CoA Carboxylase o Requires ATP o Coenzyme = BIOTIN o Regulated step / IRREVERSIBLE 2 WAYS OF REGULATION OF ENZYME • • REDUCTION DEHYDRATION REDUCTION o Fatty acid synthase enzyme (7 activity) 2 domains o TOTAL RESULT OF 7 STEPS – production of 4 Carbon compound STEPS: 1. PRODUCTION OF CYSTOLIC ACETYL CoA FURTHER ELONGATION AND DESATURATION OF FA CHAINS • Palmitate can be further elongated or desaturated • In humans, up to 9C double bonds ESSENTIAL FA – have double bonds beyond C9 • LINOLEIC ACID 18:2 (9,12) • LINOLENIC ACID 18:3 (9,12,15) RELATIONSHIP OF GLUCOSE METABOLISM & PAMITATE SYNTHESIS: FA = Acetyl CoA + ATP + NADPH 1. Glycolysis produce pyruvate 2. OAA is produced by gluconeogenesis SHORT TERM o Undergoes activation by Citrate 3. Acetyl CoA – mitochondria o Inactivated by Malonyl CoA 4. Citrate 5. Carbons of Cytosolic Acetyl CoA LONG TERM o o Prolonged consumption of high carb or fat free diets increases enzyme synth (PRO) High fat diet/fasting reduces FA synth (ANTI) 3. FA SYNTHASE Opposite of oxidation o CONDENSATION STORAGE: TAG= 3FA + GLYCEROL FA FATTY ACYL CoA TAG (Storage Fat) = glycerol phosphate + 3Fattyacyl CoA INSULIN – storage/anabolic GLUCAGON – degradation/catabolic - Rosette Go 100208