Lipid Metabolism

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Slide 1 :

1 Lipid Metabolism Hanley N. Abramson, Ph.D. Professor, Department of Pharmaceutical Sciences Wayne State University October 2008

Slide 2 :

2 Fatty Acids Ester Thioester

Slide 3 :

3 Fatty Acids as Stored Energy Fatty acids are the body’s principal form of stored energy Carbon almost completely reduced as CH2 Very closely packed in storage tissues - not hydrated as sugars are

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4 Dietary Fatty Acids Comprise 30-60% of caloric intake in average American diet Triacylglycerols, phospholipids, sterol esters Principal sources: dairy products, meats

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5 Digestion of Dietary Triacylglycerols Occurs in duodenum Facilitated by Bile salts (emulsification) Alkaline medium (pancreatic juice) Pancreatic lipases OH OH TAG MAG Intestinal lipases Glycerol + Fatty Acids Blocked by Orlistat (“Fat Blocker”) - Xenical/Alli

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6 Epithelial Cell (Intestinal Wall) Intestinal lumen MAG Glycerol Fatty Acids TAG Lipoprotein Chylomicrons Lymphatics Blood (bound to albumin) Adipose Tissue And Muscle

Slide 7 :

7 Adipocytes

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8 Fat Storage Mainly as triacylglycerols (triglycerides) in adipose cells Constitute 84% of stored energy Protein - 15% Carbohydrate (glucose or glycogen) - <1%

Slide 9 :

9 Processing of Lipid Reserves: Overview 1. Lipid Mobilization: In adipose tissue TAGs hydrolyzed to fatty acids plus glycerol 2. Transport of Fatty Acids in Blood To Tissues Activation of Fatty Acids as CoA Ester Transport into Mitochondria 5. Metabolism to Acetyl CoA

Slide 10 :

Release of Fatty Acids from Triacylglycerols

Slide 11 :

11 Adipose Cell Hormone (Adrenalin, Glucagon, ACTH) Receptor (7TM) ATP c-AMP Adenylyl Cyclase Activates Activates lipase Triacylglycerols Glycerol + Fatty acids Blood Lipolysis Insulin blocks this step

Slide 12 :

12 ATP c-AMP AMP Inactive Kinase Activated Kinase Inactive Lipase Activated Lipase P Triacyl- glycerol Glycerol + Fatty Acids Phosphatase (Hormone-sensitive Lipase) Insulin favors formation of the inactive lipase Adenylyl cyclase Phosphodiesterase Enhanced by insulin Enhanced by glucagon

Slide 13 :

13 Acylglycerol Lipases Triacylglycerol Lipase Diacylglycerol Lipase OH OH OH Monoacylglycerol Lipase OH OH OH Triacylglycerol (TAG) Diacylglycerol (DAG) Monoacylglycerol (MAG) Glycerol

Slide 14 :

14 Fate of Glycerol OH OH OH Glycerol In Liver: Dihydroxyacetone Phosphate Pyruvate Glucose Glycolysis Gluconeogenesis

Slide 15 :

15 Beta Oxidation Cleavage of fatty acids to acetate in tissues Occurs in mitochondria

Slide 16 :

16 Steps in Beta Oxidation Fatty Acid Activation by Esterification with CoASH Membrane Transport of Fatty Acyl CoA Esters Carbon Backbone Reaction Sequence Dehydrogenation Hydration Dehydrogenation Carbon-Carbon Cleavage (Thiolase Reaction)

Slide 17 :

17 Fatty Acid Activation by Esterification with CoASH CoASH + RCO2H + ATP RCOSCoA + AMP + PPi AcylCoA Synthetase 2 Pi Pyrophos- phatase Occurs in outer mitochondrial membrane for long chain fatty acids ATP AMP + PPi -32.3 CoASH + RCO2H RCOSCoA +31.5 PPi 2 Pi -33.6 ?G0’(KJ/mole) -34.4

Slide 18 :

18 Membrane Transport of Fatty Acyl CoA Esters Transported across inner mitochondrial membrane by translocase

Slide 19 :

19 Source: http://cellbio.utmb.edu/cellbio/mitochondria_1.htm Carnitine acyltransferase I Carnitine acyltransferase II Translocase

Slide 20 :

20 Beta Oxidation Reaction Sequence Occurs in Mitochondria Repeat Sequence (?-ketothiolase)

Slide 21 :

21 Complete Beta Oxidation of Palmitoyl CoA 7 Cycles 8 CH3COSCoA + 7 FADH2 + 7 NADH + 7 H+

Slide 22 :

22 Energetics of Complete Oxidation of Fatty Acids Palmitic Acid Palmitoyl CoA -2 CH3COSCoA CO2 + H2O 108 High Energy Phosphate Bonds Generated Net 106 TCA Cycle 106 High Energy Phosphate Bonds ?G0’ = 3,233 KJ/Mole For Palmitic Acid CO2: ?G0’ = - 9,790 KJ/Mole Efficiency of ?-Oxidation = 33%

Slide 23 :

23 Complete Oxidation Fatty Acids: 9 kcal/g Carbohydrates: 4 kcal/g Protein: 4 kcal/g

Slide 24 :

24 American Golden Plover

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25 Arctic Tern

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26 Camel

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27 Beta Oxidation of Odd Carbon Fatty Acids 5 Cycles 5 CH3COSCoA + CH3CH2COSCoA Propionyl CoA D-Methylmalonyl CoA L-Methylmalonyl CoA Succinyl CoA TCA Cycle Propionyl CoA Carboxylase ATP/CO2 Epimerase Mutase Vit. B12

Slide 28 :

28 Beta Oxidation of Unsaturated Fatty Acids Oleoyl CoA Beta Oxidation (3 Cycles) cis-?3 Isomerase trans-?2 Continuation of Beta Oxidation

Slide 29 :

29 Ketogenesis: Formation of Ketone Bodies 2 CH3COSCoA CH3COCH2COSCoA Thiolase CH3COSCoA Acetoacetyl CoA HO2C-CH2-C-CH2COSCoA OH CH3 ?-Hydroxy-?-methylglutaryl CoA (HMG CoA) HMG CoA Synthase Cholesterol (in cytosol) Several steps Ketogenesis (in liver: mitochon- drial matrix) See Slide 78

Slide 30 :

30 Ketogenesis: Formation of Ketone Bodies (Cont’d.) HO2C-CH2-C-CH2COSCoA OH CH3 HMG CoA Acetoacetate HMG CoA lyase - CH3COSCoA - CO2 CH3COCH3 Acetone (volatile) CH3CHCH2CO2 OH ?-Hydroxybutyrate NADH + H+ NAD+ Dehydrogenase Ketone bodies are important sources of energy, especially in starvation CH3COCH2CO2

Slide 31 :

31 ?-Hydroxybutyrate Acetoacetate Succinyl CoA Succinate Acetoacetyl CoA ?-Ketoacyl CoA transferase 2 Acetyl CoA Thiolase TCA Cycle Ketone Bodies As Energy Sources In liver Acetoacetate is major energy source in cardiac muscle and renal cortex; also in brain in starvation and diabetes Not found in liver Combines with oxaloacetate

Slide 32 :

32 Ketones in Diabetes Mellitus In presence of insulin: Enhanced glucose uptake by tissues Decreased mobilization of lipids by adipocytes In absence of insulin: Decreased glucose uptake by tissues Increased mobilization of lipids by adipocytes

Slide 33 :

33 Ketones in Diabetes Mellitus Biochemical consequences of decreased insulin production: Glucose not taken up by liver Decreased oxaloacetate to combine with acetyl CoA to enter TCA Adipocytes release fatty acids into blood Increased production of ketone bodies in liver

Slide 34 :

34 CH3COCH2CO2H pKa = 3.6 Acetoacetic Acid CH3CHCH2CO2H pKa = 4.7 ?-Hydroxybutyric acid OH Concentration of acetoacetic acid can result in metabolic acidosis (pH 7.1) affinity of Hb for O2. Metabolic Acidosis in Untreated Diabetes Mellitus

Slide 35 :

35 Fatty Acid Biosynthesis

Slide 36 :

36 Fatty Acid Synthesis vs. Degradation Intermediates Site Enzymes Redox Coenzymes Synthesis Degradation Linked to SH in Linked to CoASH Proteins (Acyl Carrier Proteins) Cytosol Mitochondria Components of Separate Polypeptides Single Peptide NADP+ / NADPH NAD+ / NADH

Slide 37 :

37 Fatty Acid Biosynthesis Occurs in cytosol Starts with acetyl CoA Problem: Most acetyl CoA produced in mitochondria Acetyl CoA unable to traverse mitochondrial membrane

Slide 38 :

38 Mitochondrial membrane Cytosol Mitochondria Glucose Pyruvate Pyruvate Acetyl CoA Oxalo- acetate Citrate Citrate Acetyl CoA Pyruvate Dehydrogenase ATP-Citrate Lyase Malate Oxaloacetate Malic enzyme Malate dehydrogenase Note: Acetyl CoA cannot be converted to glucose Citrate As Carrier of Acetate Groups

Slide 39 :

39 Fatty Acid Biosynthesis: Formation of Malonyl CoA CH3COSCoA + ATP + HCO3- -O2CCH2COSCoA Acetyl CoA Carboxylase + ADP + Pi + H+ Malonyl CoA Committed step in fatty acid synthesis Reaction is irreversible Regulation of acetyl CoA carboxylase activity: by palmitoyl CoA by citrate by insulin by epinephrine and glucagon Malonyl CoA inhibits carnitine acyl transferase I Blocks beta oxidation

Slide 40 :

40 Fatty Acid Biosynthesis: Role of Acyl Carrier Proteins CH3COSCoA CH3CO-S-ACP -O2CCH2COSCoA -O2CCH2CO-S-ACP Acetyl Transferase Malonyl Transferase Acetyl ACP Malonyl ACP ACP = Acyl carrier protein

Slide 41 :

41 Fatty Acid Biosynthesis: Formation of Acetoacetyl ACP CH3CO-S-ACP + -O2CCH2CO-S-ACP CH3COCH2CO-S-ACP + CO2 Acetoacetyl ACP ?-Ketoacyl ACP Synthetase

Slide 42 :

42 Fatty Acid Biosynthesis: Formation of Butyryl ACP CH3COCH2CO-S-ACP CH3CCH2CO-S-ACP OH H Acetoacetyl ACP ?-D-Hydroxybutyryl ACP ?-Ketoacyl ACP reductase NADPH + H+ NADP+ CH3C=C-CO-S-ACP H H ?-Hydroxyacyl ACP dehydratase - H2O Crotonyl ACP CH3CH2CH2CO-S-ACP Butyryl ACP 2,3-trans- Enoyl ACP reductase NADPH + H+ NADP+

Slide 43 :

43 Fatty Acid Biosynthesis: Sources of NADPH Pentose Phosphate Pathway: CHO OH OH OH OP HO CO2- OH OH OH OP HO NADP+ NADPH + H+ NADP+ NADPH + H+ CO2 OH OH OH OP O Ribulose-5- phosphate 6-Phospho- gluconate Glucose-6- phosphate Malic Enzyme: HO-CH-CO2- CH2CO2- Malate CO2 NADP+ NADPH + H+ Malic Enzyme CH3CCO2- O Pyruvate

Slide 44 :

44 Fatty Acid Biosynthesis: Chain Elongation CH3CH2CH2CO-S-ACP -O2CCH2CO-S-ACP + CH3CH2CH2COCH2CO-S-ACP CH2CH2CH2CHCH2CO-S-ACP CH3CH2CH2C=CCO-S-ACP H H OH

Slide 45 :

45 Fatty Acid Biosynthesis: Chain Elongation (Cont’d) CH3(CH2)3CH2CO-S-ACP CH3CH2CH2C=CCO-S-ACP H H NADPH + H+ NADP+ CH3(CH2)13CH2CO-S-ACP 5 Cycles Palmitoyl ACP CH3(CH2)13CH2CO2- Palmitate Thioesterase

Slide 46 :

46 Fatty Acid Biosynthesis: Fatty Acid Synthase in Animals Consists of a single polypeptide containing three distinct domains Conducts all steps in fatty acid synthesis except function of acyl CoA carboxylase

Slide 47 :

47 Orlistat: A Fatty Acid Synthase (FAS) Inhibitor Anti-obesity (Inhibits pancreatic lipase in git) Inhibits thioesterase domain of FAS Anti-cancer (experimental): FAS overexpressed in several tumor types; inhibition induces apoptosis

Slide 48 :

48 The Crystal Structure of a Mammalian Fatty Acid Synthase Timm Maier, Marc Leibundgut, Nenad Ban* Sept. 5, 2008

Slide 49 :

49 Further Processing of Fatty Acids: Elongation CH3(CH2)13CH2COSCoA Palmitoyl CoA CH3(CH2)13CH2COCH2COSCoA CH3(CH2)13CH2CCH2COSCoA OH H NADH + H+ NAD+ Thiolase Dehydrogenase L-? Configuration CH3COSCoA In mitochondria and at surface of endoplasmic reticulum

Slide 50 :

50 Further Processing of Fatty Acids: Elongation (Cont’d) CH3(CH2)13CH2CCH2COSCoA OH H CH3(CH2)13CH2C=CCOSCoA H H - H2O Hydratase CH3(CH2)13CH2CH2CH2COSCoA Stearoyl CoA NADPH + H+ NADP+ Dehydrogenase

Slide 51 :

51 Further Processing of Fatty Acids: Unsaturation CH3(CH2)13CH2CH2CH2COSCoA CH3(CH2)7C=C(CH2)7COSCoA + H2O H H Stearoyl CoA Oleoyl CoA This reaction occurs in eukaryotes Endoplasmic reticulum membrane Stearoyl CoA Desaturase O2

Slide 52 :

52 Further Processing of Fatty Acids: Polyunsaturation CH3(CH2)7C=C(CH2)7CO2H H H Oleic acid Plants: Further unsaturation occurs primarily in this region Animals: Further unsaturation occurs primarily in this region (18:1?9) 9 Linoleic acid (18:2?9, 12) 12 9 Linolenic acid (18:3?9, 12, 15) 15 12 9 Essential dietary fatty acids in mammals

Slide 53 :

53 Formation of Arachidonate in Mammals Linoleic acid 14 11 8 5 Arachidonic acid (20:4?5, 8, 11, 14) (Eicosa-5,-8,11,14-tetraenoic acid) As CoA ester: 1) Elongation 2) Desaturation x 2 Prostaglandins

Slide 54 :

54 Omega-3 Fatty Acids ?-3 double bond Eicosapentaenoic acid (20:5?5, 8, 11, 14, 17) Docahexaenoic acid (22:6?4, 7, 10, 13, 16, 19) Found in fish oils, esp. cold water fish Important in: Growth regulation Modulation of inflammation Platelet activation Lipoprotein metabolism

Slide 55 :

55 Metabolite Regulation of Fatty Acid Synthesis and Breakdown Pyruvate Acetyl CoA Malonyl CoA Palmitoyl CoA Citrate Inhibits Stimulates Beta Oxidation Blocks Glucose

Slide 56 :

56 Hormonal Regulation of Fatty Acid Synthesis and Breakdown ATP cAMP AMP Adenylyl cyclase Glucagon and epinephrine Stimulates Phosphodiesterase Insulin Stimulates Activates Protein Kinase Inactivates ACC by phosphorylation Inhibition of fatty acid synthesis Activates triacyl- glycerollipase Inactivates lipase

Slide 57 :

57 Synthesis of Phosphatidate Dihydroxyacetone Phosphate (from glycolysis) Glycerol Phosphatidate (formed in endoplasmic reticulum) Diacylglycerol (important in cell signaling) R3COSCoA Diacylglycerol acyltransferase (liver) Triacylglycerol (transported to adipocytes and muscle)

Slide 58 :

58 Synthesis of Glycerophospholipids R=H; CDP ethanolamine R=CH3; CDP choline CDP = cytidine diphosphate Diacylglycerol + Transferase R3=NH3; Phosphatidylethanolamine R3=N(CH3)3; Phosphatidylcholine Serine Ethanolamine + + Phosphatidylserine

Slide 59 :

59 Respiratory Distress Syndrome Most frequently seen in premature infants Also called hyaline membrane disease Failure to produce sufficient dipalmitoyl phosphatidylcholine, which normally is found in the extracellular fluid surrounding alveoli; decreases surface tension of fluid to prevent lung collapse Treatment in infants born before 30 weeks includes administration of artificial lung surfactant (e.g., Exosurf or Pumactant)

Slide 60 :

60 Synthesis of Glycero-phospholipids (Cont’d) Phosphatidate Cytidine diphosphate (CDP) diacylglycerol Phosphatidyl- inositol + Diacylglycerol (DAG) Phospholipase C (plasma membrane) Both IP3 and DAG are important second messengers in cell signaling pathways Inositol-1,4,5- triphosphate (IP3) Phosphorylation of 4 & 5 OH groups

Slide 61 :

61 Synthesis of Glycero-phospholipids (Cont’d) Cytidine diphosphate (CDP) diacylglycerol Cardiolipin: formed in inner mitochondrial membrane; plays role in oxidative phosphorylation

Slide 62 :

62 Synthesis of Glycero-phospholipids (Cont’d) Dihydroxyacetone Phosphate (from glycolysis) Plasmalogens (Abundant in cardiac tissue and CNS)

Slide 63 :

63 Synthesis of Sphingolipids CH3(CH2)14COSCoA + HCO3-2 CoASH 3-Ketosphingosine synthase CH3(CH2)14CO-CHCH2OH NH3+ 2S,3-Ketosphinganine 3 Steps CH3(CH2)12CH=CH-CH-CH-CH2OH OH Ceramide Palmitoyl CoA Serine trans CH3(CH2)nCONH

Slide 64 :

64 Synthesis of Sphingolipids (Cont’d) CH3(CH2)12CH=CH-CH-CH-CH2OH CH3(CH2)nCONH OH Ceramide Phosphatidylcholine Diacylglycerol CH3(CH2)12CH=CH-CH-CH-CH2O-P-OCH2CH2N(CH3)3 CH3(CH2)nCONH OH O O- + Sphingomyelin Cerebrosides Gangliosides trans trans

Slide 65 :

65 Synthesis of Gangliosides CH3(CH2)12CH=CH-CH-CH-CH2OH CH3(CH2)nCONH OH Ceramide CH3(CH2)12CH=CH-CH-CH-CH2O-Sugar CH3(CH2)nCONH OH Cerebroside Ganglioside trans trans Glucose or galactose Ceramide - Sugar - Sugar - GalNAc - Gal NAN NAN = N-acetylneuraminate GalNAc = N-acetylgalactose

Slide 66 :

66 Lipid Storage Diseases (Gangliosidoses)

Slide 67 :

67 Tay-Sachs Disease Ceramide - O - Glucose - Galactose - N-Acetylgalactose Hexoseaminidase A catalyzes cleavage of this glycoside linkage GM2 (a ganglioside): Autosomal recessive disorder characterized by deficiency of hexoseaminidase A; accumulation of gangliosides in brain Most prevalent in Jews from Eastern Europe For further information see: http://www.marchofdimes.com/professionals/681_1227.asp

Slide 68 :

68 Other Gangliosidoses Gaucher’s disease: Fabry’s disease: Nieman-Pick disease: Ceramide - O - Glucose Ceramide - O - Glucose - O - Galactose - O - Galactose Ceramide - Phosphate - Choline ?-glucosidase ?-galactosidase sphingomyelinase

Slide 69 :

69 Synthesis of Eicosanoids R’= H or CH3 In cell membrane Hydrolysis of sn-2 ester bond by phospholipase A2 (PLA2) Arachidonate

Slide 70 :

70 Synthesis of Eicosanoids: PLA2 Activation Various stimuli: Activation of Hormones, autacoids, etc. Membrane-bound Receptors PLA2 Activity Ca+2 Arachidonate release and eicosanoid synthesis are important mediators of tissue injury and inflammation

Slide 71 :

71 Synthesis of Eicosanoids: Prostaglandin Synthesis Cyclic endoperoxide Hydroperoxide Prostaglandin endoperoxide synthetase (Cyclooxygenase) Cyclooxygenase Hydroperoxidase Prostaglandin endoperoxide synthetase (also called cyclooxygenase) possesses both cyclooxygenase and hydroperoxidase activity Two forms of cyclooxygenase: COX -1 - constitutively expressed COX -2 - inducible PGH2 PGG2

Slide 72 :

72 Cyclooxygenase (COX) Inhibitors Nonsteroidal antiinflammatory drugs: Acetylsalicylic acid (aspirin) Ser-530 Irreversible inhibition of COX by acetylation of the active site Actions of Aspirin: Antiinflammatory (COX-2 inhibition) GI injury (COX-1 inhibition)

Slide 73 :

73 COX-2 Selective Inhibitors Rofecoxib (Vioxx) Celecoxib (Celebrex) Glucocorticoids block COX-2 expression

Slide 74 :

74 Prostaglandins PGH2 PGE2 PGD2 PGF2a Prostaglandins exhibit a variety of actions on different tissues

Slide 75 :

75 Prostacyclin and Thromboxanes PGH2 Prostacyclin (PGI2): Blocks platelet aggregation Prostacyclin synthase Thromboxane synthase Thromboxane A2 (TxA2): Promotes platelet aggregation (t1/2 = 30 sec.) Non-Enzymatic Thromboxane B2 (TxB2): inactive

Slide 76 :

76 Leukotriene Biosynthesis Arachidonic acid 5-Hydroperoxyeicosa- 6,8,11,14-tetraenoic acid (5-HPETE) 5-Lipoxygenase Leukotriene A4 (LTA4) 5-Lipoxygenase Glutathione LTC4 synthase Leukotriene C4 (LTC4) Leukotriene E4 (LTE4) - Glu - Gly LTA Hydrolase Leukotriene B4 (LTB4) Leukotrienes are important mediators of inflammation Cysteinyl leukotrienes

Slide 77 :

77 Leukotriene Biosynthesis (Cont’d) Arachidonic acid 12-Lipoxygenase 12-Hydroperoxyeicosa- 5,8,10,14-tetraenoic acid (12-HPETE) 12-Hydroxyeicosa- 5,8,10,14-tetraenoic acid (12-HETE)

Slide 78 :

78 Leukotriene Biosynthesis Inhibition Zileuton (Zyflo) An inhibitor of 5-lipoxygenase Used in the treatment of asthma

Slide 79 :

79 Cholesterol Biosynthesis: Formation of Mevalonate 2 CH3COSCoA CH3COCH2COSCoA Thiolase CH3COSCoA Acetoacetyl CoA HO2C-CH2-C-CH2COSCoA OH CH3 ?-Hydroxy-?-methyl- glutaryl CoA (HMG CoA) HMG CoA Synthase HO2C-CH2-C-CH2CH2OH OH CH3 3R-Mevalonic acid HMGCoA reductase CoASH Key control step in cholesterol biosynthesis Liver is primary site of cholesterol biosynthesis

Slide 80 :

80 Cholesterol Biosynthesis: Processing of Mevalonate -O2C-CH2-C-CH2CH2OH OH CH3 Mevalonate -O2C-CH2-C-CH2CH2OPOP CH3 OH 2 Steps ATP 5-Pyrophospho- mevalonate CH2=C-CH2CH2OPOP CH3 - CO2 - H2O Isopentenyl pyrophosphate CH3-C=CH2CH2OPOP CH3 Dimethylallyl pyrophosphate Isomerase

Slide 81 :

81 Cholesterol Biosynthesis: Isoprenoid Condensation Head Tail Head Tail Isopentenyl Pyrophosphate (IPP) Dimethylallyl pyrophosphate Head to tail Condensation Geranyl Pyrophosphate (GPP) Farnesyl Pyrophosphate (FPP) Head to tail condensation of IPP and GPP Tail to tail condensation of 2 FPPs Squalene Head Tail Head Tail Isoprenes Geranyl transferase Geranyl transferase Squalene synthase

Slide 82 :

82 Isoprenoids Widely distributed in nature Generally contain multiple of 5 carbons: Monoterpene; 10 carbons Sesquiterpene: 15 carbons Diterpene: 20 carbons Menthol: a monoterpene Lycopene: a tetraterpene

Slide 83 :

83 Conversion of Squalene to Cholesterol Squalene Squalene monooxygenase 2,3-Oxidosqualene cyclase Lanosterol 20 Steps Cholesterol Acyl-CoA: cholesterol acyltransferase Cholesterol esters (principal transport form in blood) O2 Squalene- 2,3-epoxide

Slide 84 :

84 Inhibition of Cholesterol Biosynthesis Atorvastatin (Lipitor): resembles intermediate HMG CoA Mevalonate Intermediate HMGCoA reductase

Slide 85 :

85 Transformations of Cholesterol: Bile Salts Cholesterol Cholic acid R = CH2SO3- Taurocholate R = CO2- Glycocholate Detergents

Slide 86 :

86 Transformations of Cholesterol: Steroid Hormones Cholesterol Estradiol Progesterone Cortisol Testosterone Vitamin D

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Lipid Mobilization:. In adipose tissue TAGs hydrolyzed to fatty acids plus .... Enhanced glucose up more Lipid Mobilization:. In adipose tissue TAGs hydrolyzed to fatty acids plus .... Enhanced glucose uptake by tissues; Decreased mobilization of lipids by … less
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