Development of Complex Curricula for Molecular Bionics and Infobionics Programs within a consortial* framework**

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1 PETER PAZMANY UNIVERSITY SEMMELWEIS UNIVERSITY Development of Complex Curricula for Molecular Bionics and Infobionics Programs within a consortial* framework** Consortium leader PETER PAZMANY UNIVERSITY Consortium members SEMMELWEIS UNIVERSITY, DIALOG CAMPUS PUBLISHER The Project has been realised with the support of the European Union and has been co-financed by the European Social Fund *** **Molekuláris bionika és Infobionika Szakok tananyagának komplex fejlesztése konzorciumi keretben ***A projekt az Európai Unió támogatásával, az Európai Szociális Alap társfinanszírozásával valósul meg TÁMOP /2/A/KMR

2 Semmelweis University BIOCHEMISTRY BIOKÉMIA ANABOLISM OF CARBOHYDRATES A SZÉNHIDRÁTOK SZINTÉZISE TRETTER LÁSZLÓ TÁMOP /2/A/KMR

3 BIOCHEMISTRY: ANABOLISM OF CARBOHYDRATES Table of contents: Gluconeogenesis and its regulation Glycogen synthesis and its regulation Lactose synthesis TÁMOP /2/A/KMR

4 Gluconeogenesis Learning objectives At the end of the presentation students will be able: To demonstrate the difference between the steps of gluconeogenesis and glycolysis. To demonstrate the reactions, specific for gluconeogenesis. To understand the concept of gluconeogenesis i.e. that the higher multicellular organisms use many noncarbohydrate precursors for the biosynthesis of glucose, because the maintenance of blood sugar level is highly important. To understand the principles of metabolic regulation by the comparison the regulation of gluconeogenesis and glycolysis TÁMOP /2/A/KMR

5 Gluconeogenesis Gluconeogenesis DEF : biosynthesis of glucose from simpler, non-carbohydrate precursors The most important precursors of gluconeogenesis: lactate, pyruvate glycerol (glucogenic) amino acids fatty acids with odd number of carbon atoms Glucogenic amino acids DEF : amino acids with carbon skeleton that can be used in glucose synthesis during gluconeogenesis (e.g. alanine, aspartate) Antonym: Ketogenic amino acids DEF : amino acids with carbon skeleton that can be used in ketone body synthesis during starvation (e.g. leucine, lysine) TÁMOP /2/A/KMR

6 Gluconeogenesis The principles of gluconeogenesis I. Gluconeogenesis requires reactions from glycolysis, citric acid cycle and a few special reactions The nonequilibrium reactions during glucose catabolism obstruct the simple reversal of glycolysis The nonequilibrium reactions: 1. phosphoenol pyruvate pyruvate 2. fructose 6-P fructose 1,6-P 2 3. glucose glucose 6-P TÁMOP /2/A/KMR

7 Gluconeogenesis The principles of gluconeogenesis II. The circumvention of glycolytic reactions 1. Glycolysis: phosphoenol pyruvate pyruvate Pyruvate kinase ADP ATP Gluconeogenesis:pyruvate oxaloacetate phosphoenol pyruvate Pyruvate carboxylase Phosphoenol yruvate carboxykinase CO 2 ATP ADP+Pi GTP CO 2 GDP TÁMOP /2/A/KMR

8 Gluconeogenesis The principles of gluconeogenesis III. The circumvention of glycolytic reactions However: mitochondria are impermeable for oxaloacetate but oxaloacetate should be present in the cytosol in order to become a substrate for phosphoenol pyruvate carboxykinase (PEPCK) Problem solved by - reversible oxidoreduction and transamination - reversible transports OA Malate dehydrogenase (MDH) MAL Malate dehydrogenase (MDH) MAL OA OA NADH NAD + transaminase ASP ASP NAD + NAD H transaminase OA Glut α KG transaminases DEF : enzymes catalyzing the reversible transfer of amino group from an amino acid to an oxo acid resulting a new amino acid and a new oxo acid α KG Glut TÁMOP /2/A/KMR

9 Gluconeogenesis The principles of gluconeogenesis IV. The circumvention of glycolytic reactions 2. Glycolysis: fructose 6-P fructose 1,6-P2 Phosphofructokinase I ATP ADP Gluconeogenesis: Fructose 1,6-P 2 Fructose 6-P Fructose 1,6-P 2 ase H 2 O TÁMOP /2/A/KMR

10 Gluconeogenesis The principles of gluconeogenesis V. The circumvention of glycolytic reactions 3. Glycolysis: glucose glucose 6-P Hexokinase Glucokinase ATP ADP Gluconeogenesis: glucose 6-P glucose Gl 6-P Gl 6-Pase G6T Gl 6-P H 2 O Gl Pi Pi Gl ER Glucose 6-P is transported by glucose 6P transporter (G6T) then hydrolized in the ER by glucose 6Pase TÁMOP /2/A/KMR

11 Gluconeogenesis Gluconeogenesis in the liver from lactate and pyruvate Gl Gl Gl6Pase Gl 6-P ER Pi Fr 6-P Fr 1,6P2ase Fr 1,6-P PYR TRIOSE PHOSPHATES PEP PYR LACTATE LACTATE PEPCK OA ASP OA ASP PYR Ac-CoA PC OA CELL MEMBRANE MAL MALATE TCA CYCLE MITO CELL MEMBRANE TÁMOP /2/A/KMR

12 Gluconeogenesis Entry of glycerol into the gluconeogenesis (Liver) Gl Gl Gl6Pase Gl 6-P ADIPOSE TISSUE BLOOD ER Pi Fr 6-P Fr 1,6P2ase Glycerol Glycerol Aldolase Aldolase A B Fr 1,6-P Dihydroxyacetone-P NADH Glycerol kinase NAD Glycerol 3-P dehydrogenase ATP ADP Glycerol 3-P CELL MEMBRANE Glyceraldehyde 3-P CELL MEMBRANE TÁMOP /2/A/KMR

13 Gluconeogenesis Entry of fructose into the gluconeogenesis (Liver) Gl Gl Gl6Pase Gl 6-P Diet ER Pi Fr 6-P Fr 1,6P2ase Fructose Fructose Fr 1,6-P Fructokinase Fr 1-P Aldolase A Aldolase B Dihydroxyacetone-P Aldolase B CELL MEMBRANE Glyceraldehyde 3-P ATP Triokinase Glyceraldehyde CELL MEMBRANE TÁMOP /2/A/KMR

14 BIOCHEMISTRY: ANABOLISM OF CARBOHYDRATES Gluconeogenesis Entry of amino acids into the gluconeogenic pathway Gl Gl Gl6Pase Gl 6-P ER Pi Fr 6-P Fr 1,6P2ase Fr 1,6-P ALA AMINO ACIDS PYR Serine TRIOSE PHOSPHATES PEP PYR LACTATE CELL MEMBRANE PEPCK GOT OA MDH MAL ASP OA ASP MDH MALATE PYR Ac-CoA PC OA Fumarate TCA CYCLE α-kg Succ-CoA AMINO ACIDS MITO CELL MEMBRANE TÁMOP /2/A/KMR

15 BIOCHEMISTRY: ANABOLISM OF CARBOHYDRATES Entry of glycerol into the gluconeogenesis (Liver) Gl Gl Gl6Pase Gl 6-P ADIPOSE TISSUE BLOOD ER Pi Fr 6-P Fr 1,6P2ase Glycerol Glycerol Fr 1,6-P Glycerol kinase ATP ADP Aldolase A Aldolase B Dihydroxyacetone-P NADH NAD Glycerol 3-P dehydrogenase Glycerol 3-P CELL MEMBRANE Glyceraldehyde 3-P CELL MEMBRANE TÁMOP /2/A/KMR

16 Gluconeogenesis Entry of propionyl-coa into the gluconeogenesis (Liver) Fatty acids with odd number of carbon atoms Valine Isoleucine Methionine Cholesterol side chain Glucose Propionyl-CoA PEP PEPCK OA Malate Succinyl-CoA Fumarate Succinate TÁMOP /2/A/KMR

17 Gluconeogenesis Regulation of gluconeogenesis Gene expression Covalent modification Allosteric Enzyme Inducer repressor Pyruvate carboxylase Phosphoenol pyruvate carboxykinase (PEPCK) Fr 1,6-P 2 ase Pyruvate kinase Glucocort. Glucagon epinephrine (camp) Glucocort. Glucagon epinephrine (camp) Glucocort. Glucagon epinephrine (camp) Glucocort. Glucagon epinephrine (camp) phosphorylat ion dephosphory lation activator inhibitor insulin insulin insulin insulin Fructose 2,6-P 2 AMP - - Fructose 1,6-P 2 ATP, alanine TÁMOP /2/A/KMR

18 Gluconeogenesis Regulation at the PEPCK level The promoter region of PEPCK gene PEPCK (phosphoenol pyruvate carboxykinase) is the rate limiting step of gluconeogenezis All of the important hormones in the metabolism can regulate the PEPCK expression TÁMOP /2/A/KMR

19 G L Y C O L Y XS I S BIOCHEMISTRY: ANABOLISM OF CARBOHYDRATES Gluconeogenesis Regulation at the fr 2,6-P 2 ase level (liver) Fr 6-P X Fr 1,6-P 2 Glucagon Receptor [camp] PKA PFK2 Phosphatase activity - + [fr2,6-p 2 ] PFK2 P Kinase activity + The elevation of [camp] increases the rate of gluconeogensis and decreases the rate of glycolysis in the liver Cell membrane Glucose Fr 6-P Fr 1,6-P 2 G L U C O N E O G E N E S I S TÁMOP /2/A/KMR

20 Gluconeogenesis Regulation of gluconeogenesis I Regulated steps: pyruvate carboxylase PEPCK fr 2,6-P 2 ase Gl 6-Pase Pyruvate carboxylase is regulated step, it catalyses an irreversible reaction, it is at the beginning of the pathway, but it is not entirely comitted to the gluconeogenesis. Pyruvate carboxylase is also an anaplerotic enzyme of the citric acid cycle. Anaplerotic reactions can not be entirely shut off. Anaplerotic DEF : a reaction which can replenish the supply of intermediates in a metabolic pathway, e.g. in the citric acid cycle. PEPCK is a rate limiting, irreversible, heavily regulated step at the initial part of the pathway. It is important to note that neither allosteric nor phosphorylation type of regulation has not been detected on the enzyme. It is regulated exclusively on the gene expression level. Rate limiting step DEF : The slowest step in a metabolic pathway. Usually it is heavily regulated. E.g. The rate limiting step of glycolysis is the fr 6-P fr1,6-p 2 transformation TÁMOP /2/A/KMR

21 Gluconeogenesis Regulation of gluconeogenezis II. Fr 1,6-P 2 ase is also heavily regulated. This enzyme however, not the main regulatory site of the gluconeogenesis. It is far from the beginning of the pathway. It is particularly interesting that the regulation of PFK1 and the fr 1,6-P2ase is very much coordinated. Those conditions and effectors which stimulate Fr 1,6-P2ase activity inhibit PFK1 activity and vice versa. It is very tempting to say that because PFK1 is the rate-limiting enzyme of the glycolysis, Fr 1,6-P2ase should be the rate-limiting one of the gluconeogenesis, however it is not true. Gl6-Pase is also regulated, but not the rate limiting step either. The reasons for that 1. It is at the very end of the pathway, 2. It is not entirely committed to the gluconeogenesis but playing a role in the glycogenolysis as well. Glycogenolysis is activated earlier than gluconeogenesis, so the enzyme is not suitable for the fine tuning of the gluconeogenesis TÁMOP /2/A/KMR

22 Gluconeogenesis Summary Because of thermodinamic reasons gluconeogenesis is not a simple reversal of glycolysis The irreversible steps of glycolysis should be bypassed The most important gluconeogenic precursors join to the pathway at different levels The regulation of glucogenesis occurs at the irreversible reactions. The most important regulatory site of gluconeogenesis is the PEPCK enzyme TÁMOP /2/A/KMR

23 Major anabolic and catabolic pathways in glucose metabolism C A T A B O L I C A N A B O L I C TÁMOP /2/A/KMR

24 Glycogen synthesis Learning objectives At the end of the presentation students will be able: To demonstrate the difference between the steps of glycogenesis and glycogenolysis. To demonstrate the reactions, specific for glycogenesis. To understand the principles of simultaneous and opposite regulation of glycogenolysis and glycogenesis TÁMOP /2/A/KMR

25 Glycogen synthesis Overview of glycogen synthesis Glucokinase Gl 1-P Hexokinase phosphoglucomutase uridyltransferase Glucose Gl 6-P Gl 1-P UDP-glucose ATP ADP UTP PPi Glucose (n) H 2 O Glycogen synthase Pi+Pi UDP Glucose (n+1) Cost of glycogen synthesis: gluco-, hexokinase uridyltransferase Total cost 1 ATP 1 UTP 2 ATP equivalent TÁMOP /2/A/KMR

26 Glycogen synthesis Initiation of glycogen synthesis, The role of glycogenin Glycogenin Tyr-OH Self-glucosylating Primed glycogenin Glycogen synthase and Branching enzyme Tyr-O-(glucose)8 Glycogenin-Glycogen complex Tyr-O-(glycogen 8 UDP-gluc 8 UDP n UDP-gluc n UDP Glycogen synthase cannot initiate synthesis without having a primer TÁMOP /2/A/KMR

27 Glycogen synthesis The role of branching enzyme in the synthesis 4 Glycogen synthase 4 Branching enzyme UDP-gl 7UDP TÁMOP /2/A/KMR

28 Glycogen synthesis Regulation of glycogen synthesis I Main regulator: reversible phosphorylation/dephosphorylation camp Ca Protein kinases Glycogen synthase a active ATP Pi H 2 O Phosphoprotein phosphatase + - Insulin ADP camp Glycogen synthase b inactive + Gl 6-P P TÁMOP /2/A/KMR

29 Glycogen synthesis Cyclic AMP Secondary messenger DEF : An effector molecule synthesized within the cell in response to an external signal (first messenger) such as a hormone Cyclic AMP (camp) is formed from ATP by the adenylate cycle enzyme. First messengers, which could elevate camp level are glucagon, adrenalin on beta 1,2 receptors, ACTH etc. camp can activate protein kinase-a which can initiate a cascade of phosphorylation TÁMOP /2/A/KMR

30 Glycogen synthesis Regulation of glycogen synthesis II Elevation of [camp], or [Ca 2+ ] in the liver indicates the presence of glucagon and adrenaline. Meaning of glucagon: hypoglycemia, glycogen should be mobilized Glycogenesis should be inactivated. Meaning of adrenaline fight or fly stress, glycogen should be mobilized, glycogenesis should be inactivated camp, Ca2+, activate protein kinases which phosphorylate glycogen synthase Phosphorylated glycogen synthase is inactive synthesis stopped Phoshatases could remove the phosphorylation (inhibition) of the enzyme. camp inhibits the phosphatase, maintains phosphorylation of glycogen synthase (GS), maintains inhibition of the synthesis Insulin stimulates the phosphatase, relieves GS from inhibition, stimulates synthesis Gl 6-P precursor of the glycogen synthesis stimulates the inactive GS TÁMOP /2/A/KMR

31 Glycogen synthesis Glucose stimulates glycogen synthesis in the liver Increased blood glucose stimulates glycogen synthesis in the liver in an insulin independent manner as well. Glucose binds to phosphorylase a, and glucose-bound phosphorylase is a better substrate for phosphoprotein phosphatase. Phosphorylase acts as a glucose receptor. Glucose promotes inactivation of phosphorylase thus inhibits glycogenolysis. However glycogen synthesis should also be stimulated.active phosphorylase inhibits dephosphorylation of GS by protein phosphatase. But only phosphorylase a can inhibit this process.conversion of phosphorylase a to b relieves the inhibition, GS will be dephosphorylated, active, glycogen synthesis could start TÁMOP /2/A/KMR

32 Glycogen synthesis Glucose stimulates glycogen synthesis in the liver PP=phosphoprotein phosphatase TÁMOP /2/A/KMR

33 Glycogen synthesis Summary As gluconeogenesis is not the reversal of glycolysis, glycogen synthesis is also not the reversal of glycogenolysis. Glycogen synthesis needs energy, 2 high energy phosphate required/glucose incorporated into glycogen. Glycogen synthase is unable to prime de novo glycogen synthesis, In order to create a new glycogen molecule, a primer is needed, This primer is the self-catalytic glycogenin protein. The glycogen synthesis is regulated both by reversible phosphorylation/dephosphorylation, both by allosteric effectors. Among the allosteric effectors in the liver, the glucose is the most important. Glucose is able to stimulate glycogen synthesis in the liver in the absence of insulin as well TÁMOP /2/A/KMR

34 Lactose synthesis Lactose DEF: The disaccharide of the milk, containing glucose and galactose Lactose synthesis: in lactating mammary gland The enzyme:galactosyltransferase catalyzed reaction UDP-galactose + N-acetyl-glucosamine Dgalactosyl-N-acetyl-glucosamine protein attached This enzyme has a role in the glycoprotein synthesis The enzyme does not accept glucose as a galactose acceptor. However, after delivery the specificity of the enzyme changes UDP-galactose + glucose lactose + UDP lactose synthase Reason for change in the specificity: lactalbumin: produced by the mammary gland upon hormonal influence. Lactalbumin changes substrate specificity of galactosyltransferase. Lactalbumin-galactosyltransferase complex = lactose synthase Glycoprotein DEF : Proteins containing carbohydrate groups TÁMOP /2/A/KMR

35 Biochemistry: Anabolism of carbohydrates Recommended literature Orvosi Biokémia (Ed. Ádám Veronika) Textbook of Biochemistry Ed. Thomas Devlin 5 th -7th edition TÁMOP /2/A/KMR

36 Questions: 1. Which allosteric regulators can regulate both the glycolysis and the gluconeogenesis? 2. How would influence the elevation of camp the gluconeogenesis 3. Which reactions require ATP in gluconeogenesis starting from lactate 4. Which enzymes are active in phosphorylated form in the glycogen metabolism. 5. What is the effect of insulin on the gluconeogenesis? TÁMOP /2/A/KMR

37 Questions: Which statements are true for the synthesis of glycogen? 1. Glycogen synthase reaction connects free glucose molecules 2. Glycogen synthase requires ATP for the catalysis 3. During synthesis inorganic phosphate will be incorporated into the glycogen 4. The source of incorporated glucose is UDP-glucose 5. Branches are formed as postsynthetic modifications A:1,2,3, B:2,4 C:1,5, D:4,5 E:4 Which statements are true for the regulation of glycogen metabolism? 1. Phosphorylation stimulates glycogen phosphorylase activity 2. Phosphorylation decreases the activity of glycogen synthase 3. Phosphorylation increases the activity of glycogen synthase 4. Calcium increases the activity of phosphorylase kinase 5. Glucose decreases the activity of glycogen synthase in liver A:1,2,5, B:1,2,4, C:2,3,5, D:3,4, TÁMOP /2/A/KMR

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