Lecture Sixteen: METABOLIC ENERGY: [Based on GENERATION Chapter 15

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1 Lecture Sixteen: METABOLIC ENERGY: [Based on GENERATION Chapter 15 AND STORAGE Berg, (Figures in red are for the 7th Edition) Tymoczko (Figures in Blue are for the 8th Edition) & Stryer] Two major questions integral to Biochemistry: How do cells extract energy and reducing power from their environment? How do cells synthesise the building blocks of their macromolecules? Answering here mainly the first question IMPORTANT NOTE: Although metabolic reactions may look complicated the number of kinds of reaction is small and metabolic pathways are regulated in very similar ways A Thermodynamically Unfavourable Reaction Can Be Driven By A Favourable One A reaction can occur spontaneously (the change in free energy) is ( - G ) However the overall free-energy change for a coupled series of reactions is the of the free-energy changes of the individual steps So a thermodynamically favourable reaction (- G) can drive a thermodynamically unfavourable reaction (+ G) coupled to it There are three ways in which the reactions can be coupled A shared chemical intermediate (Eg: and substrate level phosphorylation) An activated protein conformation (Eg: ) Ionic (electrochemical) gradients across membranes (Eg: oxidative phosphorylation) Figure 15-8, page 417 (15-8, page 451) (15-8, page 432) What Is Energy Required For? Active transport of solutes across membranes against their concentration and/or charge gradient Biosynthesis Signal Amplification What Is Energy Obtained From? Oxidation of foodstuffs (chemotrophs) Trapping light energy (phototrophs) Part of this obtained energy is changed into (stored as) a highly accessible form before it is used This store in most energy-requiring systems is Adenosine Triphosphate (ATP) Figure 15-3, page 412 (15-3, page 447) (15-3, page 427)

2 ATP is an molecule It is an energy carrier containing two PHOSPHOANHYDRIDE BONDS A large amount of energy (~ G -12kcal/mol) is released when ATP is hydrolysed to ADP and phosphate (P i ) And exactly the same is stored when ATP is synthesised So the cycling between ATP and ADP is the fundamental mode of energy exchange in biological systems the ENERGY CURRENCY ATP Is Continuously Formed And Consumed ATP is as a long term store A typical cell consumes ATP within ONE minute of its formation therefore turnover of ATP is very high A resting human consumes ~40 kg of ATP/24 hrs ATP is an example of an The group carried is the phosphoryl group Note: ATP does NOT have special bonds, other than that large amounts of energy are released when they are hydrolysed under cellular conditions NADH and FADH 2 are further examples of Activated Carriers These two molecules are the major electron carriers in the oxidation of fuel molecules Chemotrophs derive free energy from oxidation of fuel molecules, such as glucose and fatty acids In aerobic organisms the ultimate electron acceptor is oxygen, BUT The electrons from the fuel molecules are NOT transferred directly to oxygen, but are instead transferred to special electron carriers Pyridine nucleotides (i.e. ) Or flavins (FADH 2 ) These reduced electron carriers then transfer their electrons to oxygen Via an electron transport chain located in the inner mitochondrial membrane (IMM) The energy released is used to synthesise ATP (a process known as oxidative phosphorylation) Figure 15-13, page 420 (15-13, page 454) (15-13, page 435) Nicotinamide adenine dinucleotide (NAD + is a major electron acceptor in the oxidation of fuel molecules In the oxidation of a substrate NAD + accepts an H + and two e, (equivalent to a hydride ion, H ), to form NADH

3 NOTE: NADH is used primarily for the generation of ATP The reduced form of NADP + is NADPH Used almost exclusively for biosynthesis This needs reducing power in addition to ATP The other major electron carrier in the oxidation of fuel molecules is flavin adenine dinucleotide (FAD) Figure 15-14, page 421 (15-14, page 455) (15-14, page 436) The reactive part of FAD is the isoalloxazine ring This accepts two e and two H + (equivalent to two hydrogens) to form FADH 2 Figure 15-15, page 421 (15-15, page 455) (15-15, page 436) NADH, NADPH and FADH 2 are slowly to react with oxygen and ATP is only slowly hydrolysed Catalysts (enzymes) are required to speed these processes So these enzymes can control the flow of free energy and reductive power General: TABLE 15-2 page 422 (TABLE 15-2, page 456) (TABLE 15-2, page 437) The essential message here is: Most interchanges of activated groups in metabolism are accomplished by a rather small set of carriers Stages In The Extraction Of Energy From Foodstuffs Hans Krebs described three stages in the extraction of energy from foodstuffs Figure 15-12, page 419 (15-12, page 455) (15-12, page 434) STAGE 1 Large molecules of food broken into smaller units Proteins to amino acids Polysaccharides to simple sugars (eg ) Fats to glycerol and fatty acids NOTE: No useful energy is generated in this stage STAGE 2 STAGE 3 These numerous small molecules are degraded to a few simple units that play a central role in metabolism Most are converted to acetyl CoA NOTE: Some ATP is generated at this stage

4 Citric acid cycle and oxidative phosphorylation NOTE: 90% of ATP is generated in this last stage Metabolic Pathways Are Regulated In Three Principal Ways By controlling: The amounts of Amount depends on rate of synthesis and rate of breakdown The amount of most enzymes is regulated primarily by adjusting the rate of transcription of genes encoding them The catalytic activities of these Controlled in several ways a) Reversible allosteric control b) Reversible covalent modification The accessibility of For example by controlling their entry into the cell And finally: An important general principle of metabolism is: Biosynthetic and Degradative pathways are almost always distinct NOT THE SAME Many reactions in metabolism are controlled by the ENERGY STATUS of the cell i.e.: One measure of the energy status of the cell is the balance between concentrations of ATP and AMP within the cell, known as the ENERGY CHARGE energy charge = [ATP] + 1/2 [ADP] [ATP] + [ADP] + [AMP] Summary of Lecture Sixteen Coupling a favourable one with an unfavourable reaction can drive the latter Three ways The free energy donor and store in many systems is: ATP Continuously being consumed (high turnover) NADH and FADH 2 are major electron carriers Ultimate electron carrier is Oxygen

5 Activated Carriers NADH and FADH 2 electron transfer occurs in the mitochondrial membrane Interchanges in metabolism use a small select number of carriers for the process Energy from Foodstuffs Three stages Metabolic Pathway Regulation Principal is STAGE 3 => Produces 90% of ATP Three principal ways

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