BPK 312 Nutrition for Fitness & Sport. Lecture 3
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1 BPK 312 Nutrition for Fitness & Sport Lecture 3 Nutrient Roles in Bioenergetics 1. Learning Objectives for Lecture 3 2. Bioenergetics/Conservation of Energy 3. Redox Reactions 4. ATP/Phosphocreatine 5. Cellular Oxidation/Electron Transport/Oxidative Phosphorylation 6. Energy Release from Macronutrients 7. The Metabolic Mill 1
2 1. Lecture 3 Learning Objectives (LO3) LO3-1: Define bioenergetics and give a description of the nutritionenergy interaction, energy transfer from macronutrients and cellular respiration. LO3-2: Define and explain oxidation-reduction reactions as well as how they are important in energy transfer from macronutrients to ATP. LO3-3: Explain how phosphocreatine and high energy phosphate bonds allow short term, explosive activity. LO3-4: Explain the details of how electron transport in the respiratory chain and oxidative phosphorylation allow the transfer of energy from macronutrients to ATP. LO3-5: Explain and describe the steps of energy transfer from: (i) glycogen and glucose to ATP in glycolysis, (ii) from coenzymes to high energy phosphate bonds in the citric acid cycle and (iii) from lactate to glycogen in the Cori Cycle LO3-6: Name the sources of fat for catabolism and describe the steps in transfer of energy from triacylglycerols in beta-oxidation to the electron transport chain for ATP production. LO3-7:Explain deamination, transamination and how the carbon skeletons from amino acids are catabolized to give gluconeogenic as well as ketogenic intermediates for energy transfer. 2
3 2. Bioenergetics & Conservation of Energy Bioenergetics refers to flow of energy within a living system. Aerobic chemical reactions do & anaerobic chemical reactions do not require oxygen. Energy is transferred from the sun to plants by photosynthesis using chlorophyll, H 2 O & CO 2 to produce carbohydrates (CHO) including glucose. Overall equation for photosynthesis: Cellular respiration in animals allows recovery of food chemical energy stored in plants Herbivores, carnivores and omnivores transfer energy transfer from different food sources Image Source: 3
4 2. Bioenergetics & Conservation of Energy On engraisse pas les cochons à l'eau claire Jeanne Beauregard, né Archambault, Calixa-Lavallée, Qc, Energy and Laws of Thermodynamics First law Energy is neither created nor destroyed, but instead, transforms from one state to another without being used up. There are six forms of interchangeable energy states: Chemical, Light, Electric, Mechanical, Heat, Nuclear Biologic Work Takes one of three forms: Mechanical work of muscle contraction Chemical work for synthesizing molecules Transport work that concentrates diverse substances in body fluids 4
5 2. Bioenergetics & Conservation of Energy Recall Potential Energy and Kinetic Energy Potential energy (PE) refers to energy associated with a substance s structure or position. Kinetic Energy (KE) refers to energy of motion. PE and KE constitute the total energy of any system. Releasing PE transforms it into KE of motion. Energy transformation in the human body depend on: (i) Oxidation-reduction (redox) reactions & (ii) Chemical reactions that conserve & liberate energy in Adenosine Triphosphate (ATP) 5
6 3. Redox Reactions Oxidation reduction reactions couple: Oxidation = a substance loses H +, electrons or oxygen giving a valence Reduction = a substance gains electrons giving a valence Redox reactions power the body s energy transfer processes. LIGHT MODERATE LDH STRENUOUS MAXIMAL LDH Fig 4-5: Example of a redox reaction during intense exercise - the reduction of pyruvate to give lactate & subsequent oxidation of lactate to give pyruvate during recovery cf slide 22 (Lactate Dehydrogenase = LDH) 6
7 4. Adenosine Triphosphate (ATP) & Phosphocreatine (PCr) ATP is the body s primary energy carrier molecule that captures free energy in high energy phosphate bonds Examples of work carried out in the body using ATP Digestion Circulation 2 outermost phosphate bonds are high energy phosphate bonds. Splits rapidly without O 2 Only g of ATP stored in body there is a continual resynthesis of ATP Nerve Conduction Muscle Contraction Glandular Tissue Synthesis Fig 4-8: Adenosine Triphosphate (ATP), the body s energy currency that powers all biological work 7
8 4. Adenosine Triphosphate (ATP) & Phosphocreatine (PCr) Potential energy (PE) is extracted from macronutrients in food & conserved within phosphate bonds within ATP. Chemical PE in ATP powers all biologic work. Adenosine TriPhosphatase (ATPase) for ATP degradation & energy release for rapid anaerobic energy supply ATP + H 2 O ATP Synthesis ATPase ADP + P i kcal/mol Fig 4-7: Simplified ATP image ATP Synthase ADP + P i ATP 8
9 4. Adenosine Triphosphate (ATP) & Phosphocreatine (PCr) Phosphocreatine (PCr): The Energy Reservoir In addition to ATP, PCr is another high-energy phosphate compound. PCr quickly releases large amounts of energy when bonds between creatine & phosphate are broken. Cells store 4 6 x more PCr than ATP Is a reservoir of high-energy phosphate bonds, for shortterm 8-10 s explosive, all out muscular exercise Phosphorylation gives energy transfer in phosphate bonds ATPase Creatine Phosphokinase Fig 4-9: ATP & PCr sources of anaerobic phosphate bond energy. Energy released from splitting PCr helps resynthesize ATP from ADP & Pi; Adenosine triphosphatase 9 (ATPase)
10 5. Cellular Oxidation, Electron Transport Chain (ETC) & Oxidative Phosphorylation ACTIVATION ENERGY ACTIVATION ENERGY sudden release of all chemical energy slow step-wise release of chemical energy BURNING OF GLUCOSE CELLULAR OXIDATON OF GLUCOSE Fig 4-6: Burning glucose in a fire vs. cellular oxidation of glucose 10
11 5. Cellular Oxidation, Electron Transport Chain (ETC) & Oxidative Phosphorylation Most energy for ATP phosphorylation is from oxidation of hydrogen (H) from macronutrients, CHO, lipids & protein Constitutes the mechanism for aerobic energy metabolism Involves the transfer of hydrogen atoms & electrons Loss of hydrogen= oxidation & gain of hydrogen=reduction Highly specific dehydrogenase co-enzymes are reduced with H from macronutrients Nicotinamide Adenine Dinucleotide (NAD + ) from niacin (Vit B 3 ) Flavin Adenine Dinucleotide (FAD) from riboflavin (Vit B 2 ) NADH & FADH 2 are 2 high energy molecules carrying H & their electrons Mitochondria contain cytochrome carrier molecules on their inner membrane that remove electrons from H & pass them to O 2 Electron transport by cytochromes is the respiratory chain Chemical Reactions in Mitochondria Animation Button nb change create to transfer of energy in this animation 11
12 5. Cellular Oxidation, Electron Transport Chain (ETC) & Oxidative Phosphorylation Oxidative Phosphorylation Refers to energy transfer through phosphate bonds Oxidative phosphorylation synthesizes ATP by transferring H & electrons from NADH and FADH 2 to oxygen. >90% of body s ATP synthesis Fig 4-10: Schematic diagram for oxidation of hydrogen from NADH & FADH 2 for subsequent electron transport for the reduction of O 2. 12
13 5. Cellular Oxidation, Electron Transport Chain (ETC) & Oxidative Phosphorylation Electron Transport & Oxidative Phosphorylation 1 Cytochrome 2 e- Cytochrome 2e- Cytochrome 2e- Cytochrome 2e- Cytochrome 2e- 2 3 Electron Transport Animation Button Fig 4-11: In the body chemical energy is liberated with each of 3 hydrogen/electron pairs from NADH & FADH 2 are shuttled by 5 mitochondrial cytochromes; cytochromes are Fe 13 containing proteins. This energy is conserved in ATP in high energy phosphate bonds.
14 5. Cellular Oxidation, Electron Transport & Oxidative Phosphorylation Electron Transport Chain (ETC) & Oxidative Phosphorylation Theoretical value for aerobic ATP production from oxidation of H & subsequent phosphorylation is: NADH + H ADP + 3 P i + ½ O 2 NAD + + H 2 O + 3 ATP ATP needs to be transported out of the mitochondria at the cost of some ATP On average the net yield is 2.5 ATP synthesized per NADH, when FADH 2 donates H this gives on average a net yield of 1.5 ATP synthesized from each hydrogen pair 14
15 5. Cellular Oxidation, Electron Transport Chain (ETC) & Oxidative Phosphorylation Efficiency of Electron Transport Chain (ETC) & Oxidative Phosphorylation Formation of each mole of ATP conserves ~ 7 kcal of energy Since 2.5 moles ATP is produced per mole of NADH then 2.5 x 7 kcal = ~18 kcal is conserved as chemical energy The relative efficiency is ~34% for transferring chemical energy by ETC-oxidative phosphorylation since 1 mole of NADH liberates 52 kcal, i.e. ~18 kcal/52 kcal x 100 = ~34%. Remaining 66% of this energy is dissipated as heat 15
16 5. Cellular Oxidation, Electron Transport Chain (ETC) & Oxidative Phosphorylation Role of Oxygen in Energy Metabolism 3 conditions for ATP re-synthesis using energy from macronutrients Cond. 1: Availability of reduced NADH & FADH 2 in tissues Cond. 2: Presence of oxidizing agent O 2 in the tissues Cond. 3: Sufficient concentration of the enzymes & mitochondria in the tissues to ensure energy transfer reactions proceed at their appropriate rate Oxygen is the final electron acceptor in the respiratory chain & combines with hydrogen to form water. Strenuous Exercise In Cond. 2 if there is inadequate O 2 in the tissues or in Cond 3 if the rate of delivery of O 2 is inadequate these give an imbalance between H release & acceptance by O 2, i.e. its reduction. Electron flow down ETC backs up, H accumulates & lactate forms as give in Fig 4-15 on a following slide in this lecture. 16
17 6. Energy Release from Macronutrients Sources for ATP formation include: i. Glucose derived from liver glycogen ii. Triacylglycerol & glycogen molecules stored within skeletal muscle cells/fibers iii. Free fatty acids (FFA) derived from triacylglycerol in liver and adipocytes that enter the bloodstream for delivery to active muscle iv. Intramuscular & liverderived carbon skeletons of amino acids Fig
18 6. Energy Release from Macronutrients Intramuscular Energy Stores Fig 4-13: Macronutrient Fuel Sources Mitochondrion Glycogen Glucose aa FFA TAG Glucose Deaminated aa Liver produces rich sources of amino acids (aa) & glucose (glycogen) Adipocytes give large amounts of free fatty acids (FFA) These compounds are released into blood & are carried to skeletal m. Most energy transfer takes place in mitochondria within skeletal m. Intramuscular energy sources include ATP, PCr, Triacylglycerol (TAG), 18 glycogen & carbon skeletons from aa s FFA Citric Acid Cycle ATP
19 6. Energy Release from Macronutrients Energy Release from Carbohydrates C 6 H 12 O O 2 6CO 2 + 6H 2 O kcal/mol 1 function of CHO is to supply energy for cellular work. in a bomb calorimeter the complete breakdown of 1 mol of glucose of 180 g liberates 686 kcal of energy Synthesis of 1 mol ATP needs 7.3 kcal of energy All energy in glucose oxidation could give 94 mol of ATP In muscle phosphate bonds conserve only 34%, i.e. 34% of 686 kcal/mol = 233 kcal/mol of energy in ATP bonds with the remainder dissipated as heat. 1 mol of glucose breakdown gives 233 kcal/7.3 kcal x mol -1 = 32 mol of ATP 19
20 6. Energy Release from Macronutrients Glucose Degradation Occurs in two stages: 1. Anaerobic: Glucose breaks down relatively rapidly to 2 molecules of pyruvate in the reactions of glycolysis 2. Aerobic: Pyruvate degrades further to carbon dioxide and water in the reactions of the citric acid cycle 20
21 6. Energy Release from Macronutrients Glycolysis In cytosol & anaerobic cond. Glycolysis gives 5-10% of total ATP from a glucose molecule Substrate-level phosphorylation in glycolysis gives net gain of 2 ATP Hydrogen release in glycolysis gives 2 NADH for max exercise <90 s Glycogenolysis gives net gain 3 ATP b/c 1 st step bypassed Lactate formation Glycogen phosphorylase glycolysis.html Fig 4-13 ENZYMES 1. Hexokinase 2. Glucose 6- Phosphate isomerase 3. Phosphofructokinase 4. Aldolase 5. Triosephosphate isomerase 6. Glyceraldehyde 3- phosphate dehydrogenase 7. Phosphoglycerate kinase 8.Phosphoglycerom utase 9. Enolase 10. Pyruvate kinase 21
22 6. Energy Release from Macronutrients Lactate Formation & Use In heavy exercise when energy demand exceeds O 2 supply, ETC can t process all NADH Depends on reaction 6 in glycolysis & for NAD + availability to oxidize 3-phosphoglyceraldehyde Lactate Dehydrogenase = LDH dramatically slows glycolytic rate & lactic acid production results Lactate is a valuable source of chemical energy in the Cori Cycle nb at physiological ph, lactic acid dissociates to lactate & H + Fig 4-15: Lactic Acid Formation when excess H + from NADH temporarily combines with pyruvate. This frees NAD + to accept more H + from glycolysis, cf slide 6 22
23 6. Energy Release from Macronutrients Cori Cycle Lactate is a valuable source of chemical energy during exercise 3 2 Fig Lactic acid from skeletal muscle enters venous circulation & dissociates to lactate & H + 2. Lactate enters liver where it is converted to pyruvate & then via gluconeogenesis, there is a resynthesis of glucose. 3. Blood glucose as well as muscle & liver glycogen can subsequently be maintained. 4. Glucose is released from liver to arterial blood to active skeletal muscle. Cori Cycle Animation Button Glucose animation_library/hp-25-cori_cycle/cori_cycle.html 23
24 6. Energy Release from Macronutrients Citric Acid Cycle (CAC) 2 nd stage of CHO breakdown is the CAC. Irreversible joining of pyruvate with CoA, a Vit. B derivative, from Vit B 6 or pantothenic acid, to acetyl-coa This releases 2 H atoms to reduce both NAD + & FAD Fig 4-18 fumarate oxaloacetate malate citrate Citric Acid Cycle Animation Button isocitrate e.g. Pyruvate + NAD + + CoA acetyl-coa + CO 2 + NADH + H + The acetyl portion of acetyl-coa joins with oxaloacetate to form citrate from citric acid. Each acetyl-coa gives 2 CO 2 & 4 pairs of hydrogen atoms, plus 1 high energy Guanosine-5'- triphosphate (GTP) Succinate Succinyl-CoA oxalosuccinate α-ketoglutarate wolterskluwer_vitalstream_ com/animation_library/ HP-16-citric_acid/ citric_acid_cycle.html 24
25 6. Energy Release from Macronutrients Fig Schematic Diagram of hydrogen formation & subsequent oxidation during aerobic energy metabolism. Phase 1: CAC generates H atoms during breakdown of acetyl CoA Phase 2: ATP is reformed when these H s are oxidized via aerobic electron transport - oxidative phosphorylation 25
26 6. Energy Release from Macronutrients Fig 4-19: Net Yield of 32 ATP molecules during complete oxidation of 1 glucose molecule through glycolysis, the CAC & electron transport chain 26
27 6. Energy Release from Macronutrients Energy Release from Fat Lipolysis Stored fat represents the body s biggest source of PE. Energy sources for fat catabolism include: i. Triacylglycerol stored directly in skeletal m. fiber ii. Circulating triacylglycerol (TAG) in lipoprotein complexes iii. Circulating free fatty acids Lipolysis Animation download.lww.com/ wolterskluwer_vitalstr eam_com/ animation_library/ HP-26-triacylglycerol/ triacylglycerol.html Hormone Sensitive Lipase 3,5 cyclic monophosphate (camp) TAG + 3 H 2 O glycerol + 3 fatty acids 3 steps in lipoysis, steps 1&2 with HSL, Step 3 with HSL & monoglyceride lipase camp Activation: stimulated by epinephrine, norepinephrine (e.g. exercise), glucagon, growth hormone + inhibited by lactate, insulin & ketones these circulating factors don t enter cell but activate camp & Hormone Sensitive Lipase 27
28 6. Energy Release from Macronutrients Adipocytes TAG fat droplets take up to 95% of adipocyte volume & is major FFA source Lipase stimulates glycerol & FFA release from adipocytes FFA bind to albumin in the plasma Long chain fatty acids enter muscle fibers by diffusion or by protein mediated transport &: (i) form muscle intracellular TAG Fig 4-20: Fat storage & mobilization or lipolysis Lipase (ii) bind to CoA & then to carnitine by actions of carnitineacyl-coa transferase I & II fatty acids enter mitochondria (iii) Carnitine + fatty acyl-coa à acylcarnitine + CoA (iv) end product is Acetyl-CoA à CAC & ETC to give ATP (iv) [Acetyl-CoA]/[CoA] ratio FA transfer to mitoch. Fat Mobilzation Animation download.lww.com/ wolterskluwer_vitalstre am_com/ animation_library/ HP-17-fat_mobilization/ fat_mobilization.html Short & medium chain FA diffuse freely into mitochon., cf Lec #7, slides 283-5
29 6. Energy Release from Macronutrients Breakdown of Glycerol and Fatty Acids Glycerol Provides carbon skeletons for glucose synthesis, enters glycolytic pathway as 3-phosphoglyceraldehyde to give ATP by substrate-level phosphorylation Fatty acids Beta (ß)-oxidation for fatty acid oxidation converts a free fatty acid to multiple acetyl-coa molecules. H + released during fatty acid catabolism is oxidized through the respiratory chain. Note CAC rate depends on concentration of its intermediates, including oxaloacetate & malate, that are derived from CHO. A low CHO diet can limit fatty acid oxidation, due a slow rate of the citric acid cycle. C 15 H 32 O O 2 16CO H 2 O kcal 29
30 6. Energy Release from Macronutrients Breakdown of Glycerol and Fatty Acid Fragments Electron transport chain accepts pairs of hydrogen from glycolysis, citric acid cycle and ß-oxidation *** Fat burns in a carbohydrate flame*** Fig 4-21: General scheme of glycerol & fatty acid fragment breakdown 30
31 6. Energy Release from Macronutrients Energy Release from Protein Protein plays a role as an energy substrate during endurance activities and heavy trainings. Deamination: Nitrogen is removed from the amino acid by the liver Transamination: when an amino group is passed to another compound remaining carbon skeletons enter metabolic pathways to produce ATP. especially evident for the branched chained amino acids leucine, isoleucine, valine, glutamine & aspartate Excessive intake of protein is converted to body fat. 31
32 6. Energy Release from Macronutrients A. Alanine Structure B. Transamination The nitrogen containing amine group is transferred to other compounds Allows availability of the carbon skeleton to enter into energy metabolism e.g. the compound enters into the citric acid cycle wolterskluwer_vitalstream_com/ animation_library/hp-23-transamination/ transamination.html Transamination Animation Glutamate Glutamine transaminase α-ketoglutaric acid Pyruvate Alanine Fig A: Chemical structure of aa alanine B: Transamination 32
33 6. Energy Release from Macronutrients Glucogenic & Ketogenic Amino Acids Carbon skeletons of amino acids that form pyruvate or directly enter the citric acid cycle are glucogenic because they can form glucose Carbon skeletons of amino acids that form acetyl-coa are ketogenic because they can t form glucose molecules but rather synthesize fat Fig 4-21: Glucogenic and ketogenic amino acids. 33
34 6. Energy Release from Macronutrients Deamination Gluconeogenesis Alanine Transaminase Glucose Alanine Cycle Animation Button In prolonged exercise this cycle accounts for 10-15% of total exercise energy requirement after 4 h of continuous light exercise alaninederived glucose accounts for 45% of the livers total glucose release Fig. 1-20: Glucose-Alanine Cycle HP-09-alanine_glucose/alanineglucose.html 34
35 7. The Metabolic Mill The citric acid cycle is a vital link between food energy and the chemical energy of ATP. The citric acid cycle also provides intermediates that cross the mitochondrial membrane into the cytosol to synthesize bio-nutrients. 35
36 BPK 312 Nutrition for Fitness & Sport Lecture 3 Summary Slide Nutrient Roles in Bioenergetics 1. Learning Objectives for Lecture 3 2. Bioenergetics/Conservation of Energy 3. Redox Reactions 4. ATP/Phosphocreatine 5. Cellular Oxidation/Electron Transport/Oxidative Phosphorylation 6. Energy Release from Macronutrients 7. Metabolic Mill 36
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