Cell Respiration - 1

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1 Cell Respiration - 1 All cells must do work to stay alive and maintain an ordered cellular environment. Movement of substances through membranes, manufacture of the organic molecules needed for structure and metabolism and cellular movement are some of the activities that require energy in each cell. Cell growth, development and reproduction all require energy. Cells obtain the energy to do work by oxidizing organic molecules, a process called cellular respiration. Although many organic molecules can be oxidized, glucose, the main product of photosynthesis, is the primary fuel molecule for the cells of living organisms. Cell respiration pathways are catabolic -- the end products have less energy than the reactants. Some of the energy released during cell respiration is heat energy; the rest is use to make molecules of ATP. Both autotrophs and heterotrophs must do cell respiration. In fact, the metabolic pathways used in the process of cellular respiration are the same in virtually all eukaryotic organisms. Recall that organisms that do photosynthesis (or properly, manufacture their own fuel molecules) are called autotrophs. Heterotrophs obtain their fuel molecules "pre-formed" by other organisms. Animals, fungi and many protists are heterotrophs, as are many bacteria. Plants and some protists are autotrophs, as are some bacteria. The metabolic pathways of cell respiration are variable, depending on the type of organism, the enzymes the organism has, and what the final product molecule in the cell respiration process is. We will focus on the metabolism of glucose in cell respiration, but we shall also discuss how alternative fuel molecules fit into the cell respiration. Most eukaryotic organisms are aerobic (oxygen requiring). In aerobic cellular respiration, which is the complete metabolism of glucose, electrons removed from glucose move down an electron transport system to a final electron acceptor, oxygen, hence, the emphasis on oxygen in cell respiration. Most organisms are obligate aerobes. They cannot survive without the oxygen needed for aerobic cell respiration. For complete aerobic respiration, glucose is broken down into water and carbon dioxide. This process requires oxygen. C 6 H 12 O 6 + 6O 2 6H 2 O + 6CO *kcal (ATP + Heat) {*720kcal within cells ( G= -720kcal)}

2 Cell Respiration - 2 Not all cell respiration is aerobic. Organisms that do cell respiration without oxygen are said to be anaerobic. Fuel molecules can be oxidized without oxygen to yield smaller amounts of ATP. The fermentations involve the partial breakdown of glucose without using oxygen. Many prokaryotes have a variety of fermentation pathways, using a number of different fuel molecules. The final electron acceptor for the fermentations is an organic molecule. In addition, if the final electron acceptor is an inorganic molecule other than oxygen, the process is called anaerobic respiration. All organisms do some type of anaerobic respiration or fermentation during times of oxygen deficit, although it may not be sufficient to sustain the organism's ATP needs. Some organisms are obligate anaerobes. They can not survive in the presence of oxygen. Other anaerobes are metabolic anaerobes; they lack the enzymes needed to do aerobic cell respiration. Some organisms will survive nicely in the absence of oxygen but will do aerobic respiration when oxygen is available.

3 Cell Respiration - 3 Aerobic Cell Respiration - An Overview As with many metabolic processes, cell respiration has a number of stages (three or four depending on who is describing the process) and can be used to obtain energy from a number of fuel molecules. By convention, and because it is the primary fuel molecule for most cellular respiration, we use glucose as our fuel to illustrate the respiration pathway. Glycolysis The initial stage of glucose metabolism, or cell respiration, is a process called glycolysis, which splits a glucose molecule into two molecules of pyruvate, a 3- carbon compound. Glycolysis occurs in the cytosol of the cell. What follows glycolysis depends on the presence or absence of oxygen and/or the enzymes needed. If oxygen is not available, or if the organism lacks enzymes needed for aerobic respiration, the pyruvate molecules will proceed with fermentations, or for some prokaryotes, anaerobic respiration, which we will discuss later. If oxygen is available and the organism has the enzymes to do aerobic respiration, the pyruvate molecules will be oxidized in the next stages of aerobic respiration. During the second (and third) stages of aerobic respiration: Pyruvate molecules are oxidized and lose a CO 2. The two-carbon molecules then enter the Krebs cycle, where more oxidations occur, releasing two more CO 2. The Krebs cycle occurs in the mitochondrial matrix. The final stage of aerobic respiration is the electron transport chain and the chemiosmotic synthesis of ATP. Since the energy to synthesize ATP is from the oxidation-reduction reactions, such synthesis is called oxidative phosphorylation. Oxygen is the final electron acceptor for the oxidation-reductions that start with NADH in the electron transport system. The electron transport system takes place in the inner membrane of the mitochondria. When oxygen is available, as much as 38 ATP can be generated from one glucose molecule

4 Cell Respiration - 4 Cellular Respiration - The Pathways Glycolysis Glucose is activated by two ATP-consuming reactions. The glucose molecule is phosphorylated in these reactions. The phosphorylated bonding sites are sufficiently unstable to start what is, from that point, a series of exergonic oxidations. Glucose is then broken into two molecules of the 3-carbon compound, Pyruvate. In addition: Two molecules of NADH are produced A net of two molecules of ATP are produced (Four molecules of ATP are made during Glycolysis, but 2 molecules are consumed in activating the glucose) Glycolysis always occurs in the cytosol of the cell. Incidentally, Glycolysis is the most widespread metabolic pathway in living organisms, today and evolutionarily. The earliest prokaryotes probably had the glycolysis pathway.

5 Cell Respiration - 5

6 Cell Respiration - 6 Summary of Glycolysis Glucose + 2ATP + 2NAD + + 2ADP + 2P i --> 2 Pyruvate + 2NADH + 4ATP* * Net gain of 2ATP Inputs Glucose 2 ATP* (And 2 NA D ADP + 2P i ) Outputs 2 Pyruvate 2 NADH 4 ATP* * Therefore the net energy yield is 2 ATP Note 1 The ATP generated is by substrate-level phosphorylation Note 2: All steps are catalyzed by enzymes Glycolysis occurs in the cytosol of the cell Glycolysis is the initial cell respiratory pathway for all eukaryotic organisms.

7 Cell Respiration - 7 After Glycolysis Aerobic Pathway Aerobic Cellular Respiration is comprised of two or three stages following Glycolysis (Some references consider the oxidation of pyruvate to be a part of the Krebs cycle.) Oxidation of Pyruvate to Acetyl-CoA The Krebs Cycle Electron Transport Chain and Oxidative phosphorylation These reactions occur within the mitochondria of the cell. Oxidation of Pyruvate to Form Acetyl-CoA The two Pyruvate molecules are transported into the inner matrix of the mitochondria via facilitated diffusion Each Pyruvate is oxidized releasing H + to reduce NAD + to NADH CO 2 is removed producing Acetyl (A 2-carbon compound) Acetyl combines with Co-enzyme A (formed from the B vitamin, pantothenic acid) to form Acetyl-CoA, which can enter the Krebs cycle. For one glucose molecule (two pyruvate molecules), we will obtain: 2 CO 2 2 NADH 2 Acetyl C0-A Note: When the level of ATP is high in a cell, the cell can convert acetyl-coa into lipid molecules that can be stored for later energy use. This is one way that excess calories, no matter the nutrient source, are converted to fat.

8 Cell Respiration - 8 The Krebs Cycle (Citric Acid or TCA Cycle) The Krebs cycle is a means to remove energy rich H + (with its electrons) (originally part of the glucose molecule) that can subsequently be used to generate ATP in electron transport via chemiosmosis. Essentially, the acids of the Krebs cycle are molecules, which in the right conditions (i.e., The Krebs Cycle), can be oxidized (That is donate H + with its electrons). Once the hydrogen is removed, carbon can also be removed as the waste product, CO 2. For each glucose molecule, two ATP are produced in the Krebs cycle by substrate-level phosphorylation, one for each acetyl Co-A molecule that enters the Krebs cycle. (Recall that the glucose molecule has already gone through glycolysis and been converted to two molecules of Pyruvate in the cytoplasm prior to starting the Krebs cycle.) A closer Look at the Krebs (Citric acid) Cycle Any cycle requires a substance to start the cycle (which will also be the end of the cycle). For the Krebs cycle the starter is Oxaloacetic acid (A 4-carbon acid), which is regenerated at the end of the cycle. (Note that the acids in this process are ionized, and the naming convention is to use the suffix -ate. For example, oxaloacetic acid is called oxaloacetate.) The enzymes needed to do the Krebs cycle are located in the mitochondrial matrix. Acetyl-CoA combines with Oxaloacetic acid to begin the cycle. For each turn of the Krebs cycle we will get 2 CO 2 given off plus the one in the preparation step when pyruvate is oxidized to acetyl = 4 CO 2 1 ATP produced (by substrate phosphorylation) 1 FADH 2 3 NADH plus the one in the preparation step when pyruvate is oxidized to acetyl = 4 NADH

9 Cell Respiration - 9 The Krebs Cycle Specifics The Krebs cycle will turn two times for each glucose molecule doing aerobic cellular respiration, since glycolysis produces two Pyruvate. Therefore, for each glucose molecule that we start with, the Krebs cycle in its two turns (including the preparation step of pyruvate acetyl will produce 6 CO2 2 ATP 2 FADH2 8 NADH

10 Cell Respiration - 10 Special Note on Vitamins and the Krebs Cycle Several vitamins function as precursors to coenzymes and energy transfer molecules involved in the Krebs cycle (as well as in nutrient interconversion so that fuel molecules other than glucose can be used in cell respiration). Here are a few: Coenzyme A is made from pantothenic acid NAD is made from niacin FAD is made from riboflavin Cobalamin (B 12 ) is needed for amino acid interconversion Biotin is used for conversion of fats for fuel molecules Pyroxidine (B 6 ) is used for amino acid interconversion and converting glycogen to glucose Thiamin is a coenzyme in removing CO 2 molecules Revisiting the Mitochondrion Prior to discussing the final stage of aerobic respiration, Electron Transport, let's review the structure of the mitochondrion. Recall that the mitochondrion has an outer smooth membrane and an inner deeply folded membrane. The folds are called cristae. The internal space of the mitochondrion is called the inner matrix. The space between the outer membrane and the inner membrane is the intermembrane space. The enzymes needed to do the Krebs cycle are located in the mitochondrial matrix. The enzymes and electron carrier complex for electron transport are located in the inner membrane.

11 Cell Respiration - 11 Electron Transport Chain and ATP Synthesis by Chemiosmosis Electrons can travel down an electron transport chain, releasing their energy in controlled bits. This energy can be used for the synthesis of ATP. The molecules of the electron transport chain and a protein complex, ATP Synthase, are found in the inner membrane of the mitochondria. ATP is produced by chemiosmosis using H + concentration and charge gradients to run the ATP Synthase pumps in the membrane during electron transport. The redox reactions of the electron transport chain are used to move, by active transport, Hydrogen ions (H + ) from the mitochondrial matrix through the inner membrane into the intermembrane space. Some of the carriers pick up both electrons and H + and release the H + on the opposite side of the membrane. The Electron carriers, FADH 2 and NADH, produced in the Krebs cycle (and in glycolysis), provide the electrons and Hydrogen ions needed to do the ATP synthesis. This concentration of H + in the intermembrane space establishes both a concentration and an electrical gradient that has an inherent (potential) energy value. (This is not insignificant. There can be as much as a 1000 X difference H + concentration on the different sides of the mitochondrion inner membrane). The accumulated H + ions, known as the proton-motive force, then diffuse through the channels of the ATP synthase protein complex. The protein complex of ATP synthase uses the exergonic flow of H + ions (both a concentration and charge gradient exist) to phosphorylate ADP, forming ATP in the mitochondrial matrix. ATP is synthesized in the thylakoid membranes of the chloroplast by a similar mechanism. Oxygen is required as the final electron (and Hydrogen) acceptor, producing water as the end product of aerobic cellular respiration as the H + and e- passed off the carriers combine with oxygen. (Recall that CO 2 is also a product of aerobic cellular respiration.) As a side note, certain poisons work by blocking electron transport. Rotenone blocks NADH. Cyanide and carbon monoxide block cytochrome c from reducing oxygen. Oligomycin blocks the flow of H + through the ATP synthase pump.

12 Cell Respiration - 12 How much ATP do we get from oxidizing glucose in aerobic cellular respiration? The electrons and H + from each NADH produced in the Krebs cycle and the oxidation of pyruvate to acetyl Co-A provides sufficient energy to produce a maximum of 3 ATP by chemiosmosis. The electrons and H + from each FADH 2 produced in the Krebs cycle provides sufficient energy to produce a maximum of 2 ATP by chemiosmosis (FAD is a lower energy electron transfer molecule and enters the transport chain in mid-chain, rather than at the start) The electrons and hydrogen from each NADH from Glycolysis provides sufficient energy to produce 2 ATP (The hydrogens and electrons have to be transferred from NADH in the cytoplasm to the mitochondria) 6 NADH from Krebs X 3 ATP each = 18 ATP 2 NADH from Pyruvate to Acetyl X 3 ATP each = 6 ATP 2 NADH from Glycolysis X 2 ATP each = 4 ATP 2 FADH 2 from Krebs X 2 ATP each = 4 ATP From Direct ATP synthesis (Substrate phosphorylation) 2 ATP directly from Krebs 2 ATP 2 ATP directly from glycolysis 2 ATP Maximum Total ATP from 1 glucose = 36 ATP Unfortunately, the maximum ATP is seldom realized, since the inner mitochondrial membrane is leaky to protons and some energy is used to move pyruvate from the cytoplasm into the mitochondrial matrix. Cells get about.5 ATP less per reduced carrier that enters electron transport than the maximum.

13 Cell Respiration - 13 Aerobic (Cellular) Respiration Summary The complete aerobic respiration of glucose requires three to four stages: Glycolysis Pyruvate Oxidation The Krebs cycle Electron transport phosphorylation Oxygen is the final electron acceptor in the electron transport system which combines with Hydrogen to form water Carbon Dioxide (CO 2 ) is released during aerobic respiration As much as 36 ATP can be produced for each glucose molecule Oxidation of pyruvate, the Krebs cycle and Electron transport occur in the mitochondria; Glycolysis occurs in the cytosol. All steps are catalyzed by enzymes The overwhelming majority of living organisms must do aerobic cellular respiration to stay alive. Other pathways (which we will discuss now) provide insufficient ATP to sustain life for most organisms.

14 Cell Respiration - 14 The Fermentations: Fate of Pyruvate in the Absence of Oxygen When no oxygen is available for aerobic cell respiration, eukaryotic organisms, and some prokaryotes, will complete glucose metabolism with the fermentation reactions, which are essentially an extension of glycolysis. Other prokaryotes may do anaerobic respiration. Fermentations For some microorganisms, prokaryotes and eukaryotes, fermentation is a way of life. Some lack the enzymes to do the Krebs cycle; for others, oxygen is toxic. These are the strict (or obligate) anaerobes. Others, such as yeasts and E. coli are facultative organisms. When oxygen is available, they do aerobic respiration. When oxygen is not, they perform a fermentation. NADH must be recycled constantly in cells. Like ATP, it cannot be stockpiled. NADH must use its electrons to reduce something and recover NAD + for more glycolysis. NADH's very high energy electrons can be used to make ATP only in the presence of oxygen. In the fermentations the NADH electrons produced in glycolysis are used to reduce Pyruvate to some other organic molecule, which becomes the final electron acceptor. No more ATP energy is obtained in the fermentation processes beyond the two ATP produced during glycolysis. In the Fermentations: Organic molecules serve as the electron acceptors for NADH. Among the Prokaryotes there are several different fermentation pathways. However only two pathways are found in Eukaryotic organisms: Alcoholic Fermentation Lactic Acid Fermentation Details of the Fermentations The Pyruvate from glycolysis functions as the electron acceptor for the NADH produced in glycolysis. NADH is used to reduce Pyruvate to some stable organic molecule, freeing the NAD + (or regenerating NAD + ). No additional ATP is produced. Two fermentation pathways are common in eukaryotes. The fermentation pathways are genetically determined. Humans, for example, do a lactic acid fermentation; yeasts do alcohol fermentation.

15 Cell Respiration - 15 Alcoholic Fermentation Lactic Acid Fermentation Anaerobic Electron Transport in Prokaryotes (Anaerobic Respiration) Some bacteria also have an electron transport system. In those bacterial, some inorganic substance, such as Sulfur or Nitrogen molecules, becomes the final electron acceptor, rather than oxygen. Of note are the methanogens, a significant source of methane production on earth. They reduce CO 2 to CH 4 using hydrogens from a number of organic molecules, including acids. The sulfur bacteria can reduce sulfates to hydrogen sulfide, and the first photosynthesis on earth oxidized H 2 S for its source of hydrogen rather than water. Some bacteria today still use this method of photosynthesis. Nitrogen and iron molecules also provide reducing power for anaerobic respiration. These processes are studied in microbiology.

16 Cell Respiration - 16 Versatility of Metabolic Pathways Those Other Fuel Molecules Other carbohydrates -----> Glucose -----> Glycolysis Proteins -----> Amino Acids > Pyruvate > Krebs Cycle or Proteins -----> Amino Acids > Krebs Cycle or Proteins -----> Amino Acids** -----> Glucose > Glycolysis ** Amino acids in this group are converted to pyruvate and metabolized "back" to glucose to provide glucose to brain and nervous system cells and developing red blood cells. Note: All amino acids must be deaminated prior to being used for fuel. Lipids > Glycerol > Glycolysis (Glyceraldehyde 3 Phosphate) Lipids > Fatty Acids -----> Acetyl -----> Krebs Cycle Alcohol -----> Acetyl -----> Krebs Cycle Some Notes Some of the steps in nutrient inter-conversion can work in both directions. Acids from the Krebs cycle can be used to synthesize some amino acids, and acetyl can be used to synthesize fatty acids. (About half the amino acids are non-essential in this sense; they can be made from other amino acids or from other acids in the cells.)

17 Cell Respiration - 17 The conversion of fatty acids to 2-carbon fragments that form acetyl is called β oxidation, and occurs in the mitochondrial matrix. ATP is needed for the conversion and one FADH 2 and NADH are produced along with each acetyl Co-A. (A 16-carbon fatty acid can yield 8 acetyl Co-A molecules.) Fats are more energy rich than carbohydrates. A gram of fat potentially can produce two times as much ATP as a gram of carbohydrate. Most moderate muscle activity, such as breathing and heart beat, routinely use a mixture of fats and carbohydrates. Unfortunately, use of fatty acids for fuel is a strictly aerobic process. All anaerobic respiration must have glucose. Also, fatty acid fragments can not normally cross the brain membrane barriers so that the brain does not use fats for fuel. During starvation or fasting, or when there is insufficient carbohydrate for energy needs, the body uses its protein from body tissues to supply fuel molecules to the brain and red blood cells. When fat reserves are mobilized in response to insufficient calories or insufficient carbohydrate in the diet, some of the fatty acid fragments combine to form ketone bodies rather than acetyl. These ketone bodies enter into circulation. Muscle and some other tissues can use ketone bodies for fuel, and ketone bodies can provide energy to some brain cells. However, some ketone bodies contain carboxyl groups forming keto acids that can cause ketosis, a condition that lowers the ph of the blood and impairs health. Regulating Cell Respiration Cells regulate cell respiration just as they regulate other metabolic activities. Cells that are metabolically more active will do more cell respiration (and generally have more mitochondria) to provide the ATP needed. When activity drops, the rate of ATP formation likewise diminishes. Fuel molecules used in cell respiration are also regulated. In general: When levels of carbohydrate are high, glucose is metabolized more than fats. As levels of glucose fall, stored glycogen (in animals) will be converted to glucose. When glucose supplies diminish, more fat is mobilized to supplement metabolic needs. As stated earlier, protein will be removed from body tissues when carbohydrate is unavailable to provide glucose for brain and nerve cells. When specific nutrients are high, biosynthesis pathways related to nutrient inter-conversion that would produce those nutrients are stopped.

18 Cell Respiration - 18 By the way, all excess calories are converted to fat. Fat to fat conversion is most efficient, and any fat consumed not needed for structural or fuel purposes is readily converted to adipose for storage. Excess carbohydrate beyond the maximum glycogen stores is converted to fat in an endergonic process. Some of the caloric value of the carbohydrate is lost in the conversion. Excess amino acids will be converted either to glucose, if carbohydrate reserves are low, or to adipose. Much of this conversion occurs in the respiratory pathway. Just as we convert fatty acids to acetyl to "feed" the Krebs cycle, acetyl not needed for the Krebs cycle is readily converted to fat. The mechanisms for most of these regulations involve feedback inhibition/activation. The relative amounts of ATP/ADP, NADH, some intermediates in the Krebs cycle regulate aerobic respiration rate by feedback. High ADP stimulates the enzyme, phosphofructokinase, that converts fructose 6-phosphate to fructose 1,6-bisphosphate, stopping glycolysis. High ATP levels inhibit the first step of the Krebs cycle as well as inhibiting phosphofructokinase. Low citrate levels in the Krebs cycle also stimulate phosphofructokinase. High NADH inhibits the enzyme, pyruvate decarboxylase, that oxidizes pyruvate to acetyl, thereby stopping the Krebs cycle.

19 Cell Respiration - 19 Generating Heat From Cell Respiration There are times when generating heat, rather than ATP is the needed. Organisms, such as bats, that experience torpor (a reduced metabolic state that results in lowered body temperature) need to increase body temperature rapidly when they wake from torpor. Mitochondria in special fat containing cells have a H+ transport protein the moves protons through the membrane by-passing the ATP synthase pump. Energy released by the oxidations generates heat instead. Arum lilies attract carrion beetles and flies for pollination by exuding odors that smell like carrion. The voodoo lily increases the temperature of the flower so that the odors volatilize and disperse better. The voodoo lily respiratory pathway bypasses the electron transport chain using a pathway that generates heat but does not pump protons across the mitochondrial membrane.

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