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Glycolysis Goal: use 6 carbon compounds to produce _____________________ __________________________________________________ Remember 1st law of thermodynamics: Matter (energy) can be ___________________________________ Energy in C-C bonds of glucose is transferred ___________________ Specifically: The break down of _________________ into _________________ Oxidative process accompanied by the reduction of NAD+ Glycolysis Energy Production: - __________ ATP - __________ ATP - __________ ATP - 10 separate enzymatic reactions to produce pyruvate from glucose - we will study each reaction in detail. - you must know: - each enzyme, name & order - order and structure of each metabolite What is fate of Pyruvate? 1 Glycolysis Fate of pyruvate: Aerobic (presence of O2) - citric acid cycle breaks down pyruvate ________________ - reduces ________________ _______________________ - next lecture series Anaerobic (lack of O2) 1. Fermentation: loss of CO2 and _______________ 2. Anaerobic glycolysis: reduction of pyruvate to lactate - this lecture series 2 Glycolysis History Louis Pasteur 1822 - 1895 - French Scientist - Inventor of pasteurization: heating of liquids to kill harmful organisms - sponsored by French wine industry to study fermentation - determined in 1860 that glucose supplied more energy in presence of O2 3 Glycolysis History a.k.a.: Embden - Meyerhof Pathway Otto Fritz Meyerhof Gustav Georg Embden 1884 - 1951 1874 - 1933 German born biochemist - German born biochemist - University of Kiel - between 1912 and 1918 determined the intermediates of glycolysis - awarded Nobel prize in physiology or medicine, 1922 - Worked out precise steps involved in the breakdown of glucose - Not included on Nobel prize 4 Glycolysis Two phases or parts to glycolysis (10 steps between both phases) Phase 1: - 2 “ ” of phosphorylating glucose twice. - forms fructose 1,6,-bisphosphate - split into two molecules of glyceraldehyde-3-phosphate - ______________ Phase 2: - conversion of glyceraldehyde-3-phosphate to _________________ - coupled formation _________ - _________________ 5 Glycolysis Structure of Sugars, Chapter 16 a.k.a. carbohydrates, saccharides, sugars: simple organic compounds that are ________________________________________________ - Can range in carbon number from 3 (simplest) to 4, 5, 6, and 7 (most complex) Simplest 3 carbon sugars - Exist as an aldehyde or ketone 6 Glycolysis Structure of Sugars, Chapter 16 Sugars have ____________________________ ___________________ used in glycolysis 7 Glycolysis Structure of Sugars, Chapter 16 Numbering of carbons in sugars is based on _____________________________. - the most oxidized _________ _______________________ In ketone sugars the _______________ __________________ 8 Glycolysis Structure of Sugars, Chapter 16 5 or 6 carbon sugars normally exist ___________________ The free electrons on OH of C-5 bond with ________________ forming cyclic structure Cyclic structure is similar to pyran: a six member ring with 5 carbons and one oxygen Can produce α and β forms called anomers 9 Glycolysis Structure of Sugars, Chapter 16 Different representations of sugar structures Fischer projections - _______________________ stereochemistry of 3-D molecule Haworth projections - a perspective representation of the _______ ____________________________ Fischer linear cyclized 10 Glycolysis Structure of Sugars, Chapter 16 Hermann Emil Fischer Walter Haworth 1852 - 1919 1883 - 1950 - German organic chemist - British organic chemist - worked at University of Berlin - worked at St. Andrews University - determined the structure and stereochemistry of sugars between 1884 and 1894 - worked on structure of polysaccharides starting in 1915 - Nobel Prize in Chemistry, 1902 - Nobel Prize in Chemistry, 1937 11 Glycolysis Phase 1 Step 1 - phosphorylation of glucose producing _________________ ____________________ - reaction carried out by the enzyme - requires - phosphorylation of glucose with out ATP has ΔG = _______________ - coupled to ATP hydrolysis to give reaction ΔG = _______________ 12 Glycolysis Phase 1 Step 1 - Glucose-6-phosphate (G6P) formation Hexokinase Remember : kinase is an enzyme that transfers a phosphate from ATP to substrate Hexokinase undergoes large _______________________________ _____________________________ Glucose becomes almost __________________________________________ Glucose not bound Glucose bound This is an example of what kind of enzymesubstrate interaction? _____________ 13 Glycolysis Phase 1 Step 2 - G6P is isomerized to - reaction carried out by the enzyme - ring structure is broken producing aldehyde at C-1 which is then _________________________ - C-2 OH is oxidized to give ketone, ring is closed connecting ______ _________________ - no net oxidation or reduction 14 Glycolysis Phase 1 Step 3 - Fructose-6-phosphate (F6P) is phosphorylated to give ________ ________________________ - reaction carried out by the enzyme - requires ______________ - phosphorylation of F6P with out ATP has ΔG > 0 - coupled to ATP hydrolysis to give reaction ΔG = -3.4 kcal/mole 15 Glycolysis Phase 1 Step 3 - Fructose-1,6-bisphosphate formation Phosphofructokinase (PFK) - Both G6P and F6P can be utilized by Do not have to . - Once PFK produces fructose-1,6-bisphosphate it is . It must go - reaction carried out by PFK is not reversible - PFK is key regulatory enzyme in glycolysis pathway 16 Glycolysis Phase 1 Step 3 - Fructose-1,6-bisphosphate formation Phosphofructokinase (PFK) ATP is an allosteric regulator of PFK In the presence of low [ATP] In the presence of high [ATP] Reasoning? High cellular [ATP] - no need for __________________ - PFK inactivated, _______________ ___________ Low cellular [ATP] - requirement for _______________ - PFK activated, ________________ 17 Glycolysis Phase 1 Step 4 - Fructose-1,6-bisphosphate is split into two 3-carbon molecules: ________________________________ and _____________________________ - reaction carried out by the enzyme - intermediates no - DHAP and GA3P 18 Glycolysis Phase 1 Step 5 - DHAP is converted to - reaction carried out by the enzyme - reaction produces a second - now 6 carbon glucose has been 19 Glycolysis Phase 2 - two molecules of each intermediate take part in each reaction Step 6 - GA3P is oxidized to - reaction carried out by the enzyme - involves oxidation of GA3P, NAD+ as e- acceptor, and phosphorylation at C-1 - Phosphate comes not from ATP but from inorganic phosphate: 20 Glycolysis Phase 2 Step 6 - GA3P is oxidized to 1,3-bisphosphoglycerate Takes place in a two step reaction 1. GA3P is oxidized to 3-phosphoglycerate - aldehyde is oxidized 21 Glycolysis Phase 2 Step 6 - GA3P is oxidized to 1,3-bisphosphoglycerate Takes place in a two step reaction 2. 3-phosphoglycerate is phosphorylated by Pi This reaction has ΔG , but overall ΔG 22 Glycolysis Phase 2 Step 7 - 1,3-bisphosphoglycerate is dephosphorylated to produce ATP - reaction carried out by the enzyme - two ATP are produced per - thus two ATP used in Phase 1 - substrate level phosphorylationATP production by the direct transfer of phosphate from intermediate ___________________ _____________________________23 Glycolysis Phase 2 Step 8 - 3-phosphoglycerate is converted to - reaction carried out by the enzyme - this reaction takes place to set up next reaction 24 Glycolysis Phase 2 Step 9 - 2-phosphoglycerate loses H2O to produce - reaction carried out by the enzyme - e- transfer is not involved - requires Mg2+ which binds H2O for elimination 25 Glycolysis Phase 2 Step 10 - phosphoenolpyruvate is dephosphorylated to form ATP from ADP, producing - reaction carried out by the enzyme - two ATP are produced per glucose molecule, substrate level phosphor. - pyruvate kinase is allosterically regulated by - inhibited by high levels 26 Glycolysis Regulation points of glycolysis 1. ___________________ Glucose-6-phosphate production 2. ____________________ Fructose 1,6-bisphosphate production inhibited by ATP 3. _____________ pyruvate produced inhibited by ATP 27 Glycolysis What is the energy production of glycolysis? 2 ATP used in phase 1 - ________________ - ________________ 4 ATP produce in phase 2 - __________________ - __________________ Overall ATP production: __________ 28 Glycolysis What is the fate of pyruvate? Depends on the ___________________ __________ conditions: - pyruvate enters citric acid cycle (TCA cycle), ______________ to generate reducing agents for ATP production (next lecture series) _____________ conditions: 1. Anaerobic glycolysis, the production of ____________________. Occurs in muscles 2. Alcohol fermentation, production of ethanol from pyruvate 29 Glycolysis Anaerobic Glycolysis - Pyruvate is reduced to lactate - occurs in - only form of glycolysis - catalyzed by enzyme - oxidizes NADH to NAD+ reduction 30 Glycolysis Anaerobic Glycolysis Why carry out anaerobic glycolysis? - citric acid cycle (TCA cycle) requires . - but during anaerobic conditions - lactate dehydrogenase produces - this NAD+ can then be used by _________________________ _________________________ - only place in glycolysis NAD+ required, thus glycolysis can continue and produce __________ ____________________________ 31 Glycolysis Alcoholic Fermentation 2 steps in the production of ethanol from pyruvate 1. The enzyme produce acetaldehyde removes CO2 from pyruvate to 32 Glycolysis Alcoholic Fermentation 2 steps in the production of ethanol from pyruvate 2. The enzyme ethanol reduces acetaldehyde to Produces NAD+ that can then be used by glyeraldehyde-3-phosphate dehydrogenase reduction 2H+ + 2e- NAD+ NADH 33 Glycolysis Alcoholic Fermentation Beer making Extract sugars from malted barley in water yeast Aerobic glycolysis Anaerobic fermentation CO2 CO2 Place in air tight bottle, yeast produced CO2 carbonates beer Use of O2 EtOH EtOH EtOH EtOH EtOH Air tight container with exhaust for gas 34