! ! Answer 1:! You have asked a classic question in biology, and of course, a very important one.

How living things produce usable energy is important not only from the perspective of understanding life, but it could also help us to design more e"cient energy harvesting and producing products - if we could "mimic" how living cells deal with their energy balance, we might be able to vastly improve our technology. For example, a plant is a much better harvester of sunlight than even our best solar panel. And of course, if we understand energy use, it can also help us deal with human diseases such as diabetes.! ! Now, the answer to your question can be found in any basic biology text book, but sometimes, there is so much information packed into such a text book that it can be di"cult to extract the information you need or more often, to view all of that information in a larger context. Let's try to tackle your question in several parts.! ! First, we need to know what ATP really is - chemically, it is known as adenosine triphosphate. ATP is a usable form of energy for cells - the energy is "trapped" in a chemical bond that can be released and used to drive other reactions that require energy (endergonic reactions).! ! Photosynthetic organisms use energy from sunlight to synthesize their own fuels. They can convert harvested sunlight into chemical energy (including ATP) to then drive the synthesis of carbohydrates from carbon dioxide and water. When they synthesize the carbohydrates, oxygen gets released. Globally, more than 10 billion tons of carbon is "xed" by plants every year - this means that carbon molecules are converted from being part of a simple gas (carbon dioxide) into more complex, reduced molecules (carbohydrates), making carbon available as food for nonphotosynthesizers (and of course, providing oxygen). They use some of the carbohydrate for their own growth and reproduction. It is pretty remarkable when you think about it - have you been to Sequoia National Park or seen the redwoods along our northwest coast? Massive trees, right? Think about the fact that most of that mass is in the form of carbon that was pulled out of the air as carbon dioxide!!

! The process of photosynthesis is two-part. First, there are the light reactions, where light is converted into chemical energy (a reduced electron carrier and ATP). This occurs in the thylakoids (stacked membranes) of the chloroplasts. The ATP and electron carriers are then used in a second set of reactions, called the light-independent reactions. This also occurs in the chloroplasts, but in an area called the stroma. In this case, carbon dioxide gets used to produce sugars in a series of reactions called the Calvin Cycle, C4 photosynthesis, and crassulacean acid metabolism. You can look in any basic bio textbook to see how much "energy" or "sugar" is produced in each step of the process.! ! In non-photosynthesizers, the fuel has to be consumed. The most common chemical fuel is the sugar glucose (C6H12O6)... Other molecules, such as fats or proteins, can also supply energy, but (usually) they have to rst be converted to glucose or some intermediate that can be used in glucose metabolism.! ! Now this brings us to the next part - how do we go from glucose to ATP? This is achieved through the process of "oxidation" - and this is carried out through a series of metabolic pathways. Complex chemical transformations in the cell occur in a series of separate reactions to form each pathway, and each reaction is catalyzed by a specic enzyme. Interestingly, metabolic pathways are similar in all organisms, from bacteria to humans. In eukaryotes (plants and animals) many of the metabolic pathways are compartmentalized, with certain reactions occurring in specic organelles. Basically, cells trap free energy released from the breakdown (metabolism) of glucose. This energy gets trapped in the ATP as it converts from ADP to ATP by the addition of phosphate.! ! There are 3 main pathways for harvesting energy from glucose:! Glycolysis - begins glucose metabolism in all cells to produce 2 molecules of pyruvate. Occurs outside of mitochondria, usually in cytoplasm.! ! Cellular Respiration - uses oxygen from the environment and converts each pyruvate to three molecules of carbon dioxide while trapping the energy released in this process in ATP. There are 3 sub-pathways of

cellular respiration - pyruvate oxidation, the citric acid (Krebs or Tricarboxylic Acid) cycle and the electron transport chain. Occurs in di#erent sub-compartments of mitochondria.! ! Fermentation - converts pyruvate into lactic acid or ethanol; does not need oxygen. It is not as e"cient as cellular respiration; it occurs in the cytoplasm.! ! In terms of how much ATP is produced, you can look in your basic texts and assess how many ATPs are used versus how many are produced for each aspect of metabolism! ! Answer 2:! ATP is created through a complex enzyme-driven process. There are a couple of ways this works in cells:! ! -glycolysis, in which glucose is broken up into two subunits, called pyruvate, which creates two units of ATP per molecule of glucose. This happens in the cytoplasm, in both animal and plant cells.! ! -respiration, in which pyruvate is combined with oxygen to form carbon dioxide and water, which creates a lot of ATP per unit of pyruvate (16 I think). Obviously, glycolysis has to happen rst, in order to make the pyruvate. This happens in the mitochondrion, which likewise exists in both animals and plants.! ! -light-dependent photosynthesis, in which electrons are cycled around photosynthetic pigments after being jostled around by mid-high-energy photons (light particles), which lter through an electron pump that makes ATP. This happens in chloroplasts, and as such only occurs in plants.! ! Note: there is one more major (eukaryotic) ATP-generating reaction, called "fermentation", which takes pyruvate and turns it into carbon dioxide and ethyl alcohol. Neither animals nor plants can do this, but fungi can do it.! ! There is also light-independent photosynthesis, which USES ATP and creates glucose from carbon dioxide and water, producing oxygen as a waste product. This is really an energy storage mechanism, so that the

organism doing it can later burn the glucose through glycolysis and respiration. This also happens in chloroplasts, so plants can, and animals can't.! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! !

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Energy Production In A Cell

(Chapter 25 Metabolism)

Large food molecules contain a lot of potential energy in the form of chemical bonds but it requires a lot of work to liberate the energy. Cells need a quick easy way to get energy for anabolism: this is done with ATP. ATP is an unstable molecule, the bonds of which are easy to break making it a useful source of energy for cells. ATP ! ADP + P + free energy from food Food energy + ADP + P ! ATP Catabolic reactions generate energy to make ATP, and the ATP energy is used to drive anabolic reactions, such as metabolic turnover (replacement of cell parts), growth and cell division, and special functions (such as secretion, absorption, contraction, or signaling). Metabolism = the sum of all chemical reactions in the body; catabolism + anabolism All energy production begins in the cytosol of the cell. Large molecules are catabolized into smaller molecules, but very little energy is produced: Proteins ! amino acids Triglycerides ! fatty acids and glycerol Carbohydrates ! short carbon chains These smaller molecules are then absorbed and processed in reactions inside the mitochondria. 40% of the energy is captured to produce ATP from ADP and the remaining 60% escapes as heat (used to maintain body temperature). Oxidation-Reduction Reactions (Redox Rxns) Oxidation = the removal of electrons (or addition of oxygen) Reduction = the addition of electrons These reactions are always coupled: one molecule must be oxidized while another is reduced. A-e + B ! A + B-e (A is oxidized while B is reduced) The reduced molecule gains energy while the oxidized molecule loses energy. Cells more commonly perform dehydrogenation reactions where a hydrogen (1 proton + 1 electron) is exchanged instead of a free electron, but this is still a redox reaction. Catabolism of large molecules results in reduced carrier compounds, which are then oxidized to generate ATP.

Amy Warenda Czura, Ph.D.

SCCC BIO130 Chapter 25 Cellular Respiration Handout

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ATP Production Generation of ATP involves the addition of a phosphate to ADP and can be accomplished one of two ways: 1. Substrate Level Phosphorylation: a high-energy phosphate is transferred directly from a substrate to ADP thus forming ATP. 2. Oxidative Phosphorylation: electrons are transferred from an organic compound to a cofactor carrier molecule (e.g. NAD+). The electrons are passed through other carriers (the electron transport chain) to a final acceptor (oxygen) and the passing of the electrons releases energy that is harvested to add a phosphate to ADP in a process called chemiosmosis. Carbohydrate Catabolism Carbohydrates are the primary source of cellular energy for most organisms. Glucose is the most commonly used carbohydrate and will always be used first. Fatty acids will be used second when glucose is in short supply, and then more rarely amino acids can be utilized. Glucose can be catabolized for ATP production in two ways: 1. Cellular respiration: requires oxygen to serve as the final electron acceptor in a series of redox reactions that generate ATP by oxidative phosphorylation. This is the most efficient method of ATP production (1 glucose generates 36 ATP) and involves reaction performed inside the mitochondria. 2. Fermentation: requires an organic molecule to serve as the final electron acceptor and can be done in the absence of oxygen. ATP is synthesized using substrate level phosphorylation, which is less efficient (1 glucose generates 2 ATP). In humans this results in the production of lactic acid. Fermentation reactions are carried out in the cytoplasm. Aerobic Respiration Of Glucose C6H12O6 + 6 O2 ! 6 CO2 + 6 H2O (energy from 1 glucose ! 36 ATP) Three stages of aerobic respiration: 1. Glycolysis: oxidation of glucose to pyruvic acid with some ATP and NADH produced. 2. Citric Acid Cycle: oxidation of acetyl to carbon dioxide with some ATP, NADH and FADH2 produced. 3. Electron Transport Chain: NADH and FADH2 are oxidized providing electrons for redox reactions ultimately reduce oxygen to generate ATP. The majority of the ATP is produced at this step. *(NAD (nicotinamide adenine dinucleotide) and FAD (flavin adenine dinucleotide) are coenzymes that function to transport electrons in the form of hydrogen: NAD+ carries 2 electrons but only one proton, FAD carries 2 complete hydrogen atoms)

Amy Warenda Czura, Ph.D.

SCCC BIO130 Chapter 25 Cellular Respiration Handout

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Glycolysis a.k.a the Embden-Myerhof Pathway Glycolysis is an anaerobic process (does not involve oxygen) that occurs in the cytoplasm. It consists of a 10-step metabolic pathway that catabolizes and oxidizes 1 glucose molecule into 2 pyruvic acid molecules and generates 2 molecules of ATP by substrate level phosphorylation. Many cells can survive on glycolysis alone (called fermentation) but it is not very efficient, and fermentation generates lactic acid as a waste product, which will need to be removed and processed to prevent a drastic alteration in pH and loss of homeostasis. Two stages: 1. Preparatory Stage: two ATP molecules are used to phosphorylate one 6-carbon glucose and catabolize it into two 3-carbon molecules. 2. Energy Conservation Stage: The two 3-carbon molecules are oxidized to generate two 3carbon pyruvic acid molecules. At the same time two NAD+ molecules are reduced to two NADH molecules and four ATP molecules are produced by substrate level phosphorylation. Summary of glycolysis: 1 Glucose + 2 NAD+ + 2 ADP + 2 P ! 2 Pyruvic acid + 2 NADH + 2 H+ + 2 ATP Decarboxylation (Preparation for the Citric Acid Cycle) This is the first step in the aerobic process of glucose metabolism. This and all subsequent steps (Citric Acid Cycle and Electron Transport) will occur only when oxygen is available and will take place inside the mitochondria. The 3-carbon pyruvic acid is decarboxylated into carbon dioxide and a 2-carbon acetyl. The acetyl is attached to Coenzyme A (which serves only as a carrier) and one NAD+ is reduced to NADH (this will happen twice: 1 glucose generates 2 pyruvic acid molecules). Decarboxylation reactions occur in the matrix of the mitochondria. Summary of decarboxylation: 2 Pyruvic acid + 2 NAD+ + 2 CoA ! 2 Acetyl CoA + 2 CO2 + 2 NADH Citric Acid Cycle a.k.a. The Krebs Cycle or The Tricarboxylic Acid Cycle This part of the aerobic metabolism of glucose involves 8 enzymatic reactions occurring in the mitochondrial matrix that function to reduce the coenzymes NAD+ and FAD. The 2-carbon acetyl is attached to a 4-carbon oxaloacetic acid creating a 6-carbon citric acid. Oxidation and decarboxylation reactions occur which catabolize the 6-carbon citric acid back into a 4-carbon oxaloacetic acid and two carbon dioxide molecules. At the same time three NAD+ and one FAD are reduced into three NADH and one FADH2 respectively, and one ATP is produced by substrate level phosphorylation. (Remember: 1 glucose ! 2 pyruvic acid ! 2 acetyl so this cycle will run twice.) Summary of the Citric Acid Cycle: 2 Acetyl Co A + 6 NAD+ + 2 FAD + 2 ADP + 2 P + 4 H2O ! 2 CoA + 4 CO2 + 6 NADH + 4 H+ + 2 FADH2 + 2 ATP

Amy Warenda Czura, Ph.D.

SCCC BIO130 Chapter 25 Cellular Respiration Handout

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Electron Transport Chain This is the truly aerobic part of the aerobic metabolism of glucose as this is where the oxygen is utilized. Oxidative phosphorylation occurs on a membrane, the mitochondrial cristae, to generate most of the ATP produced from glucose. Coenzymes from the previous reactions pass electrons to a series of electron carrier molecules, which carry out redox reactions resulting in the chemiosmotic generation of ATP. There are three classes of carrier molecules: 1. FMN (flavin mononucleotide): protein + flavin coenzyme 2. CoEnzyme Q : nonprotein 3. Cytochromes: protein + an iron group (most common) Events of the electron transport chain: 1. NAD+ and FAD collected energy in the form of hydrogens (electrons) from organic molecules during Glycolysis, Decarboxylation, and the Citric Acid Cycle becoming the reduced forms NADH and FADH2 2. NADH and FADH2 are oxidized and pass hydrogens (electrons and protons) to the electron transport chain consisting of flavoproteins, cytochromes, and coenzyme Q. As electrons are passed along the chain, protons are pushed out through the membrane. This sets up a concentration gradient with protons (positive charge) on the outside and electrons (negative charge) on the inside. 3. At the end of the chain the electrons are accepted by oxygen creating an anion (O-) inside, which has a strong affinity for the cations (H+) outside. 4. Chemiosmosis generates ATP: H+ from the outside moves toward O- on the inside through special membrane channels that are coupled to ATP synthase, and the high-energy diffusion of H+ drives the reaction ADP + P ! ATP. The energy from 1 NADH from glycolysis can generate 2 ATP, the energy from 1 NADH from decarboxylation and the Citric Acid Cycle can generate 3 ATP, and that from 1 FADH2 can generate 2 ATP for a total of 32 ATP. 5. H+ combines with O- inside the mitochondria creating water (H2O). Summary of Electron Transport: 2 NADH from Glycolysis + 2 NADH from Decarboxylation + 6 NADH from Citric Acid Cycle + 2 FADH2 from Citric Acid Cycle + 6 O2 + 32 ADP + 32 P ! 12 H2O + 32 ATP + 10 NAD+ + 2 FAD Final Summary For Aerobic Respiration: C6H12O6 + 6 O2 + 36 ADP + 36 P ! 6 CO2 + 6 H2O + 36 ATP (In most Eukaryotic cells, 36 ATP are produced from 1 glucose molecule: 2 from Glycolysis and 2 from the Citric Acid Cycle by substrate level phosphorylation, and 32 from Electron Transport by oxidative phosphorylation. Glycolysis occurs in the cytoplasm, the Citric Acid Cycle occurs in the matrix of the mitochondria, and Electron Transport occurs on the cristae of the mitochondria. Some energy is lost as electrons are carried from the cytoplasm across the mitochondrial membrane and thus the NADH from Glycolysis can generate only four total ATP instead of the 3 each observed for NADH created inside the mitochondria.)

Amy Warenda Czura, Ph.D.

SCCC BIO130 Chapter 25 Cellular Respiration Handout

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Cellular Respiration Review: 1. Glycolysis: - anaerobic, occurs in cytoplasm - 1 glucose oxidized into 2 pyruvic acids - 2 ATP produced by substrate level phosphorylation - 2 NADH produced by reduction of NAD via oxidation of glucose - If no O2 available, pyruvic acid reduced to lactic acid (fermentation) - Erythrocytes (RBCs) ! glycolysis only (no mitochondria!) - Skeletal muscle ! fermentation when no O2 - Neurons and cardiac muscle cannot ferment, need O2 2. Decarboxylation + Krebs / Citric Acid Cycle: -occur in matrix of mitochondria -Decarboxylation: - 2 pyruvic acid decarboxylated and oxidized into 2 acetyl Co A + 2 CO2 with 2 NADH -Citric Acid Cycle: - 2 acetyl combined with 2 oxaloacetic acids creating 2 citric acids - citric acid decarboxylated and oxidized ! 4 CO2, 6 NADH, 2 FADH2 - 2 ATP generated by substrate level phosphorylation 3. Electron Transport: - aerobic, occurs on cristae of mitochondria - NADH and FADH2 reduced during glycolysis and citric acid cycle are oxidized - electrons are passed to cytochromes, finally accepted by oxygen - 32 ATP generated by chemiosmosis / oxidative phosphorylation - 12 H2O produced as waste from oxidation of oxygen

* With oxygen, 1 glucose will produce 36 ATP in most human tissue cells. * Without oxygen, 1 glucose will produce 2 ATP in human tissue cells that are capable of fermentation (not neurons or cardiac muscle).

Amy Warenda Czura, Ph.D.

SCCC BIO130 Chapter 25 Cellular Respiration Handout

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Lipolysis (Lipid Catabolism) The process of lipid catabolism hydrolyzes triglycerides, the storage form of fat, into glycerol and three fatty acids. The glycerol is converted into pyruvic acid in the cytoplasm and catabolized through the Citric Acid Cycle in the mitochondria. The fatty acids are catabolized by Beta-oxidation in the mitochondria to be entered into the Citric Acid Cycle as two-carbon fragments. For each two-carbon fragment of fatty acid produced by Beta-oxidation, the cell can generate 17 molecules of ATP. This is 1.5 times the energy production (when compared carbon to carbon) as with glucose. Although lipolysis generates more energy, it requires more oxygen and occurs much more slowly than equal carbohydrate metabolism. Protein and Amino Acid Catabolism For proteins to be used for energy production, they must first be broken down into individual amino acids. Then for the amino acids to be used in the Citric Acid Cycle they must first have the amino group (NH2) removed in a process called deamination, which requires vitamin B6. The amino group is removed in conjunction with a hydrogen creating ammonia (NH3) which is very toxic. The liver must then convert the ammonia to urea, which is relatively harmless, for excretion by the kidney. The remaining amino acid carbon chains are then used at various stages in the Citric Acid Cycle to generate ATP. The amount of ATP produced varies with the type of amino acid, depending on which step into the Citric Acid Cycle it was entered. Catabolism of amino acids is not a practical source of quick energy and is typically only used in starvation situations. Proteins are harder to break apart than carbohydrates or lipids, their catabolism generates toxic waste products, and they are the structural and functional parts of every cell, and thus tend to only be used when no other energy source is available. Instead, amino acids are usually simply recycled by hydrolysis of peptide bonds in one protein, to be reassembled by dehydration synthesis into the next. Nucleic Acid Catabolism DNA is never catabolized for energy. RNA can be broken down into simple sugars and nitrogenous bases. The sugars are metabolized in glycolysis but only the pyrimidine bases, uracil and cytosine, can be processed into the Citric Acid Cycle. The purines, adenine and guanine, are deaminated and excreted as uric acid making RNA metabolism very inefficient. Typically nucleotides are simply recycled into new nucleic acid molecules and are not used for energy production.

Amy Warenda Czura, Ph.D.

SCCC BIO130 Chapter 25 Cellular Respiration Handout

! ! Home! Cellular Respiration! ! Introduction! ! What is Cellular Respiration?! ! Cellular respiration allows organisms to use (release) energy stored in the chemical bonds of glucose (C6H12O6). The energy in glucose is used to produce ATP. Cells use ATP to supply their energy needs. Cellular respiration is therefore a process in which the energy in glucose is transferred to ATP.! ! ! ! In respiration, glucose is oxidized and thus releases energy. Oxygen is reduced to form water.! ! The carbon atoms of the sugar molecule are released as carbon dioxide (CO2).! ! The complete breakdown of glucose to carbon dioxide and water requires two major steps: 1) glycolysis and 2) aerobic respiration. Glycolysis produces two ATP. Thirty-four more ATP are produced by aerobic pathways if oxygen is present.! ! In the absence of oxygen, fermentation reactions produce alcohol or lactic acid but no additional ATP.! ! ! ! Review of Electron Carriers! ! NAD+ + 2H --> NADH + H+! ! FAD + 2H --> FADH2!

! Glycolysis! ! During glycolysis, glucose (C6) is broken down to two molecules of pyruvate (C3). (Note that compounds that end in "___ate" can be called "___ic acid". For example, lactate is lactic acid and malate is malic acid.)! ! Glycolysis occurs in the cytoplasm (cytosol) and does not require oxygen.! ! There are ten steps in glycolysis and each one is catalyzed by a specic enzyme. A brief summary of these reactions is presented here.! ! 2 ATP molecules are used to phosphorylate and activate compounds that will eventually become converted to pyruvate (or pyruvic acid) (see diagram below).! ! Two hydrogen atoms are removed by NAD+ forming 2 NADH (see diagram).! ! Additional phosphorylation results in intermediate 3-carbon molecules with 2 phosphate groups.! ! Four ATP are produced by substrate-level phosphorylation. Recall that substrate-level phosphorylation is the production of ATP using energy from other high-energy compounds but without the use of the electron transport system in the mitochondria.! ! ! ! The net yield of ATP in glycolysis is 2 for each glucose molecule (2 are used but 4 are produced).! ! ! ! Some bacteria have alternative energy-producing reactions. Two of these are the pentose phosphate pathway and the Entner-Doudoro# pathway.! ! Formation of Acetyl CoA!

! Pyruvate produced by glycolysis (see above) enters the mitochondrion by active transport and is converted to acetyl CoA as shown below. The remainder of the reactions of cellular respiration occur in the mitochondrion.! ! pyruvate (C3) --> acetyl CoA (C2) + CO2! ! A carbon atom is removed from each of the pyruvate molecules forming a two-carbon compound and CO2.! ! Each of the two-carbon compounds are oxidized forming NADH from NAD+.! ! Coenzyme A is attached to each of the two-carbon compounds producing two acetyl CoA molecules.! ! ! ! Citric Acid Cycle! ! The citric acid cycle can be summarized by either of the diagrams below. The diagram below occurs twice, once for each acetyl CoA.! ! ! ! Coenzyme A is removed when the two-carbon compound is attached to a four-carbon compound producing a six-carbon compound (citrate).! ! Each citrate molecule undergoes a series of reactions that removes 2 carbon atoms which are released as CO2. In addition, 3 NADH, 1 ATP, and 1 FADH2 are produced. In addition, the four-carbon compound that began the cycle is regenerated.! ! The diagram below also summarizes the citric acid cycle.! ! ! !

Electron Transport and Oxidative Phosphorylation! ! Mitochondrion structure! ! The inner membrane forms folds called cristae. These folds contain the carriers of the electron transport system.! ! Acetyl CoA formation and the citric acid cycle occur in the inner space called the matrix.! ! The space outside the inner membrane is the intermembrane space. The electron transport system pumps hydrogen ions (H+) into this space for oxidative phosphorylation.! ! Oxidative Phosphorylation! ! The electron transport system is found in the mitochondrion and chloroplast of eucaryotes and in the plasma membrane of procaryotes. It consists of a series of carrier molecules which pass electrons from a highenergy compound to a nal low-energy electron acceptor. Energy is released during these oxidation-reduction reactions to produce ATP.! ! The discussion below applies to the mitochondria of eucaryotes.! ! NADH or FADH2 bring electrons to the electron transport system in the mitochondria.! ! The system contains membrane-bound electron carriers that pass electrons from one to another. When a carrier reduces another, some of the energy that is released as a result of that reduction is used to pump hydrogen ions across the membrane into the intermembrane space. The remaining energy is used to reduce the next carrier.! ! As a result of the electron transport system, hydrogen ions become concentrated in the intermembrane space. These concentrated ions contain energy much like a dam. The enzyme ATP synthase is able to use the energy of this osmotic gradient to produce ATP as the hydrogen ions move under osmotic pressure through the enzyme back into the matrix of

the mitochondrion.! ! Oxygen is the nal electron acceptor. The low-energy electrons that emerge from the electron transport system are taken up by O2. The negatively charged oxygen molecules take up protons from the medium and form water (2H+ + 2e- + 1/2 O2 --> H2O).! ! ! ! Summary of Glycolysis and Cellular Respiration! ! ! ! Glycolysis! ! During glycolysis, glucose (C6) is converted to two pyruvates (C3).! ! C-C-C-C-C-C --> C-C-C + C-C-C! ! Formation of Acetyl CoA! ! One acetyl CoA is formed for each pyruvate produced by glycolysis (see the step above).! ! C-C-C --> C-C + CO2! ! pyruvate --> acetyl CoA + CO2! ! Citric Acid Cycle! ! C-C --> 2 CO2! ! The citric acid cycle produces NADH, FADH2, and ATP.! ! NADH and FADH2 carry electrons to the electron transport system.! ! Electron Transport System! !

In the electron transport system, NADH and FADH2 are oxidized and the energy is used to produce ATP.! ! Total ATP yield per glucose! ! Conversions! ! NADH produced in the cytoplasm produces two to three ATP by the electron transport system.! ! NADH produced in the mitochondria produces approximately three ATP.! ! FADH2 adds its electrons to the electron transport system at a lower level than NADH, so it produces approximately two ATP.! ! Glycolysis! ! 2 ATP! ! 2 NADH (= 4 ATP; these are converted to ATP in the mitochondria during cellular respiration)! ! Formation of Acetyl CoA! ! 2 NADH (= 6ATP)! ! Citric Acid Cycle! ! 6 NADH (= 18 ATP)! ! 2 FADH2 (= 4 ATP)! ! 2 ATP! ! Total Yield! ! Glycolysis produces 2 ATP; aerobic respiration produces 34 more ATP! !

Pathway$ Substrate-Level! Phosphorylation$ Oxidative! Phosphorylation$ Total! ATP! Glycolysis$ 2 ATP$ 2 NADH = 4 - 6 ATP$6 - 8! CoA$ $ 2 NADH = 6 ATP$6! Citric Acid Cycle$2 ATP$ 6 NADH = 18 ATP! 2 FADH2 = 4 ATP$ 24! TOTAL$ ! 4 ATP! ! 32 ATP! ! 36 - 38! Fermentation! ! Without oxygen, cellular respiration could not occur because oxygen serves as the nal electron acceptor in the electron transport system. The electron transport system would therefore not be available.! ! Glycolysis can occur without oxygen. Although glycolysis does not require oxygen, it does require NAD+. Cells without oxygen available need to regenerate NAD+ from NADH so that in the absence of oxygen, at least some ATP can be made by glycolysis.! ! To regenerate NAD+ from NADH, the electrons from NADH are added to pyruvate to produce alcohol (plants, yeast) or lactate (animals, bacteria).! ! The total ATP yield of fermentation comes from glycolysis; 2 ATP molecules are produced per glucose.! ! ! ! Usefulness of Fermentation! ! Anaerobic exercise! ! During vigorous exercise, oxygen is consumed faster than it is needed.

Additional ATP energy is provided to the muscles by glycolysis and the result is a buildup of lactate in the muscles.! ! When lactate builds up, the blood pH drops and the muscles fatigue.! ! At rest, lactate is converted back to pyruvate (the oxygen debt is repaid). This is why you continue to breathe hard after you have nished running or rapid stair climbing.! ! Yeast! ! Yeast produce alcohol which accumulates in their environment. As the concentration of alcohol in their environment increases, it becomes more and more toxic to them. Beer and wine have a maximum alcohol concentration because a higher concentration will kill the yeast cells.! ! Evolution of Cellular Respiration! ! Early cells probably fermented organic molecules in the oceans.! ! Today, nearly all organisms show some form of fermentation which indicates that it evolved early in evolutionary history.! ! Evolution typically operates by building upon or adding to what is already there. Aerobic respiration appears to have been added to fermentation.! ! Summary! ! Glycolysis! ! Two ATP molecules are used to phosphorylate and activate glucose.! ! Two hydrogen atoms are removed by NAD+ forming 2 NADH.! ! Four ATP molecules are produced by substrate-level phosphorylation.! ! The net yield of ATP is two; two are used and four are produced.! !

Fermentation! ! Fermentation is needed to regenerate NAD+ from NADH so that at least some ATP can be made in glycolysis.! ! Electrons from NADH are added to pyruvate (reduction) to produce alcohol (plants, yeast) or lactate (animals, bacteria)! ! Aerobic Respiration! ! Aerobic respiration occurs when oxygen is available.! ! pyruvate --> CO2 + H2O! ! It occurs in the mitochondrion.! ! NAD+ and FAD carry electrons to the electron transport system.! ! In the electron transport system, NADH and FADH2 are used to produce ATP as electrons are passed from one carrier to another. Eventually the electrons combine with hydrogen ions and oxygen (reduction) to form water.! ! ! ! Click the icon (left) to see an enlarged, color version of the diagram above.! ! Review Exercise! ! Draw each of the following. Draw only the carbon skeletons. Do not name the molecules. Draw carbon dioxide as CO2.! ! glycolysis! formation of acetyl CoA! citric acid cycle! Add the following names:! !

glucose! pyruvate! acetyl CoA! Add the following:! ! ADP, ATP! NAD+, NADH! FAD, FADH2! Fermentation! ! Draw the carbon skeletons of the molecules involved in fermentation in animals.! ! Name the compounds involved.! ! Add ATP and NADH where appropriate.! !