1. Chapter 7 Respiration

2. 2 Organisms can be classified based on how they obtain energy: –Autotrophs Able to produce their own organic molecules through photosynthesis –Heterotrophs Live on organic compounds produced by other organisms All organisms use cellular respiration to extract energy from organic molecules

3. 3 Cellular respiration Cellular respiration is a series of reactions Redox reactions Oxidized – loss of electrons Reduced – gain of electron Dehydrogenation – lost electrons are accompanied by protons –A hydrogen atom is lost (1 electron, 1 proton)

4. 4 Redox During redox reactions, electrons carry energy from one molecule to another Nicotinamide adenosine dinucleotide (NAD +) –An electron carrier –NAD + accepts 2 electrons and 1 proton to become NADH –Reaction is reversible

5. 5 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Product Enzyme NAD + NAD 2e – H+H+ +H + Energy-rich molecule NAD + 1. Enzymes that use NAD + as a cofactor for oxidation reactions bind NAD + and the substrate. 2. In an oxidation–reduction reaction, 2 electrons and a proton are transferred to NAD +, forming NADH. A second proton is donated to the solution. 3. NADH diffuses away and can then donate electrons to other molecules. Reduction Oxidation H H H H H H H H

6. In overall cellular energy harvest –Dozens of redox reactions take place –Number of electron acceptors including NAD + In the end, high-energy electrons from initial chemical bonds have lost much of their energy Transferred to a final electron acceptor 6

7. 7 Aerobic respiration –Final electron receptor is oxygen (O 2 ) Anaerobic respiration –Final electron acceptor is an inorganic molecule (not O 2 ), such as lactic acid Fermentation –Final electron acceptor is an organic molecule, such as ethanol

8. 8 Aerobic respiration C 6 H 12 O 6 + 6O 2 6CO 2 + 6H 2 O Free energy = – 686 kcal/mol of glucose Free energy can be even higher than this in a cell This large amount of energy must be released in small steps rather than all at once. –C 6 H 12 O 6 (general form for 6 carbon sugar such as glucose)

9. 9 2e – Electrons from food High energy Low energy 2H + 1 / 2 O2O2 Energy released for ATP synthesis H2OH2O Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

10. Electron carriers Many types of carriers used –Soluble, membrane-bound, move within membrane All carriers can be easily oxidized and reduced Some carry just electrons, some electrons and protons NAD+ acquires 2 electrons and a proton to become NADH 10

11. 11 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. HH O NH 2 + 2H CH 2 O O O H H NH 2 H H N N N CH 2 O O O H H NH 2 O–O– H H N N N HH O N N HH H C C OP OPO–O– OP O–O– OP N N OH CH 2 NADH: Reduced form of nicotinamide O O O O NAD + : Oxidized form of nicotinamide Reduction Oxidation Adenine OH NH 2 + H + CH 2 O–O–

12. 12 ATP Cells use ATP to drive endergonic reactions –ΔG (free energy) = –7.3 kcal/mol 2 mechanisms for synthesis 1. Substrate-level phosphorylation Transfer phosphate group directly to ADP During glycolysis 2. Oxidative phosphorylation ATP synthase uses energy from a proton gradient During electron transport chain

13. During Glycolysis

14. 14 Oxidation of Glucose The complete oxidation of glucose proceeds in stages: 1. Glycolysis 2. Pyruvate oxidation 3. Krebs cycle 4. Electron transport chain & chemiosmosis

16. 16 Glycolysis Converts 1 glucose (6 carbons) to 2 pyruvate (3 carbons) 10-step biochemical pathway Occurs in the cytoplasm Net production of 2 ATP molecules by substrate-level phosphorylation 2 NADH produced by the reduction of NAD + –Only process that occurs in red blood cells since they do not have mitochondria!

18. This shows all 10 reactions of Glycolysis Pay more attention to the previous slide

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20. 20 NADH must be recycled For glycolysis to continue, NADH must be recycled to NAD + by either: 1. Aerobic respiration –NADH goes and “drops off” electrons at electron transport chain, oxidized back to NAD+ –Oxygen is available as the final electron acceptor –Produces significant amount of ATP 2. Fermentation –Occurs when oxygen is not available –Organic molecule is the final electron acceptor so that NADH can be oxidized back to NAD+

21. 21 Fate of pyruvate Depends on oxygen availability. –When oxygen is present, pyruvate is oxidized to acetyl-CoA which enters the Krebs cycle Aerobic respiration –Without oxygen, pyruvate is reduced in order to oxidize NADH back to NAD + Fermentation

23. 23 Pyruvate Oxidation In the presence of oxygen, pyruvate is oxidized –Occurs in the mitochondria in eukaryotes multienzyme complex called pyruvate dehydrogenase catalyzes the reaction –Occurs at the plasma membrane in prokaryotes

24. 24 For each 3-carbon pyruvate molecule: –1 CO 2 Decarboxylation by pyruvate dehydrogenase –1 NADH –1 acetyl-CoA which consists of 2 carbons from pyruvate attached to coenzyme A Acetyl-CoA proceeds to the Krebs cycle Products of pyruvate oxidation

26. 26 Krebs Cycle Oxidizes the acetyl group from pyruvate Occurs in the matrix of the mitochondria Biochemical pathway of 9 steps in three segments 1.Acetyl-CoA + oxaloacetate → citrate 2.Citrate rearrangement and decarboxylation 3.Regeneration of oxaloacetate

28. 28 Krebs Cycle For each Acetyl-CoA entering (remember there are 2 for each glucose): –Release 2 molecules of CO 2 –Reduce 3 NAD + to 3 NADH –Reduce 1 FAD (electron carrier) to FADH 2 –Produce 1 ATP –Regenerate oxaloacetate

29. Don’t worry – you don’t have to memorize all of this Just wait until Biochem

30. 30 At this point Glucose has been oxidized to: –6 CO 2 –4 ATP –10 NADH –2 FADH 2 Electron transfer has released 53 kcal/mol of energy by gradual energy extraction Energy will be put to use to manufacture ATP These electron carriers proceed to the electron transport chain

31. Now onto Oxidative Phosphorylation Oxidative Phosphorylation Electron carriers are oxidized ADP is phosphorylated to become ATP Happens through Electron Transport Chain and Chemiosmosis 31

32. 32 Electron Transport Chain (ETC) ETC is a series of membrane-bound electron carriers Embedded in the inner mitochondrial membrane Electrons from NADH and FADH 2 are transferred to complexes of the ETC Each complex –A proton pump creating proton gradient –Transfers electrons to next carrier

34. 34 Chemiosmosis Accumulation of protons in the intermembrane space drives protons into the matrix via diffusion Membrane relatively impermeable to ions Most protons can only reenter matrix through ATP synthase –Uses energy of gradient to make ATP from ADP + P i

35. 35 ADP + P i Catalytic head Stalk Rotor H+H+ H+H+ Mitochondrial matrix Intermembrane space H+H+ H+H+ H+H+ H+H+ H+H+ ATP Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

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38. 38 Energy Yield of Respiration Theoretical energy yield (these values vary from book to book) –38 ATP per glucose for bacteria –36 ATP per glucose for eukaryotes Actual energy yield –30 ATP per glucose for eukaryotes –Reduced yield is due to “Leaky” inner membrane Use of the proton gradient for purposes other than ATP synthesis

40. 40 Regulation of Respiration Example of feedback inhibition (negative feedback) 2 key control points 1.In glycolysis Phosphofructokinase is allosterically inhibited by ATP and/or citrate 2.In pyruvate oxidation Pyruvate dehydrogenase inhibited by high levels of NADH Citrate synthetase inhibited by high levels of ATP

42. 42 Oxidation Without O 2 1.Anaerobic respiration –Use of inorganic molecules (other than O 2 ) as final electron acceptor –Many prokaryotes use sulfur, nitrate, carbon dioxide or even inorganic metals 2.Fermentation –Use of organic molecules as final electron acceptor

43. 43 Anaerobic respiration Methanogens –CO 2 is reduced to CH 4 (methane) –These bacteria found in diverse organisms including cows Sulfur bacteria –Inorganic sulphate (SO 4 ) is reduced to hydrogen sulfide (H 2 S) –Early sulfate reducers set the stage for evolution of photosynthesis

44. 44 a: © Wolfgang Baumeister/Photo Researchers, Inc.; b: NPS Photo Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. b.a. 0.625 µm

45. 45 Fermentation Reduces organic molecules in order to regenerate NAD + 1. Ethanol fermentation occurs in yeast –CO 2, ethanol, and NAD + are produced 2. Lactic acid fermentation –Occurs in animal cells (especially muscles, such as during sprinting) –Electrons are transferred from NADH to pyruvate to produce lactic acid

47. 47 Catabolism of Protein Amino acids undergo deamination to remove the amino group Remainder of the amino acid is converted to a molecule that enters glycolysis or the Krebs cycle –Alanine is converted to pyruvate –Aspartate is converted to oxaloacetate

48. 48 C HOO O C C O O C CO CHH CHH CHH C HH CH2NH2NH NH 3 HO Urea Glutamate α-Ketoglutarate Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

49. 49 Catabolism of Fat Fats are broken down to fatty acids and glycerol –Fatty acids are converted to acetyl groups by  -oxidation –Oxygen-dependent process The respiration of a 6-carbon fatty acid yields 20% more energy than 6-carbon glucose.

50. 50 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. CoA Fatty acid OO — C — C — C — ═ H H — ═ — ATP NADH H2OH2O Krebs Cycle PP i AMP + FADH 2 Fatty acid 2C shorter CoA Fatty acid OH H — C — C — C — — — H H — ═ — OH O H H — C — C — C ═ — — — H H — — CoA FAD Fatty acid OH — C ═ C — C — — H — ═ CoA Fatty acid O HO H — C — C — C — — — H H — ═ — CoA NAD + Acetyl-CoA

52. 52 Evolution of Metabolism Hypothetical timeline 1.Ability to store chemical energy in ATP 2.Evolution of glycolysis Pathway found in all living organisms 3.Anaerobic photosynthesis (using H 2 S) 4.Use of H 2 O in photosynthesis (not H 2 S) Begins permanent change in Earth’s atmosphere 5.Evolution of nitrogen fixation 6.Aerobic respiration evolved most recently