AP Bio Gussler Unit 3 Enzymes, Photosynthesis and Respiration Energy Conversion

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Cellular Respiration Introduction Comparison with photosynthesis PHOTOSYNTHESIS In cholorophyll-bearing cells In the presence of light Carbon dioxide and water Reduced carbon compounds, oxygen, and water Light Energy stored RESPIRATION In all cells All the time Reduced (high energy) carbon compounds and oxygen Carbon dioxide and water Chemical bonds Energy released Oxidation of carbon compounds NAD and FAD

Where? When? Input Output

Energy sources Energy result Chemical Reduction of carbon compounds reaction Energy carrier(s) NADP

II. Some main points on respiration -respiration releases energy for use by the organism - respiration involves four biochemical systems Anaerobic 1. glycolysis 2. fermentation Aerobic cellular respiration 1. Krebs cycle including conversion reactions (pyruvate to acetyl CoA) 2. Electron transport chain -respiration pathways are different for prokaryotes

AP Bio Gussler Unit 3 Enzymes, Photosynthesis and Respiration Page 2 -Respiration is about 40% efficient (Comparable to man-made machines 25%) III. Summary of ATP Yield Source Glycolvsis Krebs Cycle Electron Transport System (28 -34 approx) Total Energy Yield 2 ATP 2 ATP 32 ATP 36 ATP

IV. Energy transformations detail - Terminology: 1. Electron acceptor molecules: -Background: electrons not easily removed from covalent bonds -Since most biological molecules are covalent bonds it is hard for redox reactions to occur, so usually in a living system: 2. Oxidation = removal of hydrogen atom and release of energy Reduction = gain of hydrogen atom (therefore energy is stored in chemical bond) 3. The enzyme dehydrogenase strips hydrogen atoms from organic molecules it removes two protons and two electrons(two hydrogens). The high energy electrons are picked up by electron acceptor molecules. NAD+ (nicotinamide adenine dinucloetide) is common acceptor and coenzyme derived from vitamin B-3 Niacin is a water-soluble vitamin, which is also
known as nicotinic acid or vitamin B3. Nicotinamide is the derivative of niacin and used by the body to form the coenzymes nicotinamide adenine dinucleotide (NAD) and nicotinamide adenine dinucleotide phosphate (NADP).

-When NAD+ is in oxidized state it is positively charged, so when itaccepts two electrons, it also accepts one proton to form hydrogen -Oxidized state = NAD+ (lower energy molecule) Reduced state = NADH (higher energy molecule)

-the extra H+ ion goes to the solution in the cell. -NADH can then deliver high energy electron where they are needed like to the electron transport chain 4. aerobic with oxygen anaerobic without oxygen 5. Chemical Reactions: Redox (coupled reaction-one must accept,other must give) - Oxidation - atom or molecule loses an electron and energy released - Reduction - atom or molecule gains an electron and energy stored 6. Electron transport chain: (membranes of mitochondria- cristae)

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Unit 3 Enzymes, Photosynthesis and Respiration Page 3 Produces the most ATP (32 or 34) Cytochromes are hydrogen acceptors (protein + iron) which have a heme group and are similar to hemoglobin Cytochromes and enzymes make up ETS As electrons pass from one cytochrome to the next energy is given off - some is lost as heat, some change shape allowing H+ ions to pass thru creating gradient (proton channel opens) Products - ATP + water

a. High energy electrons are provided from the foods we eat. Food molecules are oxidized and electron carrier molecules are reduced. b. Electron carrier molecules pass the energized electrons along an electron transport chain where the acquired energy from glucose (or other foods) is harvested and is used to phosphorylate ADP to ATP c. Protons transported into outer membrane region of mitochondrion to create concentration gradient difference. pH and concentration gradient that is established across the membrane creates potential energy to produce ATP d. At each step in the electron transport chain, electrons start at a molecule where the electrons have more energy and move to one that has less, releasing energy slowly -NADH transfers its electrons to FMN ( a flavoprotein) near start of ETC -FADH2 transfers its electrons to ubiquinone (carrier Q) at a lower

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Unit 3 Enzymes, Photosynthesis and Respiration Page 4 energy level ( another coenzyme from vitamin B2 riboflavin) e. This slow release of energy, from one electron acceptor to the next helps living organisms to control energy release. If release happens to quickly too much energy is released as heat and light and is lost to the organism f. Oxygen is the final electron acceptor. It is very electronegative, meaning it steals electrons easily from the final molecule in the ETC. Each oxygen molecule combines with two hydrogen ions (from solution) and two electron to make water. 2e- + 2H+ + O2 H2O g. poisons can irreversibly inhibit the action of the molecules in the ETC. Cyanide
ions bind to the iron atom of the enzyme cytochrome c oxidase (also known as aa3) in the fourth complex in the mitochondrial membrane in the mitochondria of cells. This denatures the enzyme, and the final transport of electrons from cytochrome c oxidase to oxygen cannot be completed. As a result, the electron transport chain is disrupted, meaning that the cell can no longer aerobically produce ATP for energy.

8. Chemiosmosis: (process of creating ATP) High energy electrons are used to create proton concentration gradient. Hydrogen ions would diffuse back into the matrix of the mitochondrion but can not cross the membrane except at points bridged by enzyme that produces ATP (ATP synthase). As protons move through enzyme, energy is released creating ATP (Oxidative phosphorylation) 4. Substrate level phosphorylation an enzyme transfers a phosphate from an organic molecule to ADP much simpler! Occurs in cytoplasm in glycolysis Also how ATP is produced in Krebs cycle

V. Cellular Respiration - Oxidation reaction; chemical energy in food is released(Exergonic) CH2O + Oxygen C6 H 12O 6 + 6 O2

Carbon Dioxide + Water + energy 6CO2 + 6H2O + energy (ATP and heat)

- Process in cells by which energy is released from food in the form of ATP - Glycolysis occurs first a. Glucose to pyruvate and 2NADH b. Net production of 2ATP c. No CO2 produced, no oxygen required - Pyruvate can be converted to several different products depending on conditions in the cell a. Aerobic respiration - most efficient, CO2 and H2O waste products

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Unit 3 Enzymes, Photosynthesis and Respiration Page 5 b. Alcohol fermentation - occurs in yeast cells without oxygen c. Lactic acid fermentation - occurs in organisms when oxygen present in insufficient or small quantities - Feedback inhibition controls the rate of respiration a. phosphofructokinase catazlyzes step 3 in glycolysis is the pacemaker of respiration because allosterically inhibited by ATP and citrate b. phosphofructokinase stimulated by AMP

A. Glycolysis (lysis or breakdown of glucose) First stage in all respiration pathways Takes place in cytoplasm Input= glucose (six carbon molecule) + 2 ATP Products= 2 pyruvic acids + 4 ATP's + 2 NADH2 No oxygen required (anaerobic)

ATPs used to begin process; 4 ATPs produced as energy is transferred from broken chemical bonds- NET PRODUCTION = 2 ATP B. Exercise: 1. More efficient metabolic pathway to produce energy - Aerobic respiration *term aerobic exercise comes from this 2. Increased exercise increases bodys demand for oxygen which is carried by blood and pumped by the heart. a. Increased blood volume from demands of exercise b. Increased efficiency of heart muscle c. Increased efficiency of lungs to obtain oxygen C. Aerobic Respiration: 1. Efficient metabolic pathway for breakdown of carbohydrates, proteins and lipids producing between 25 and 38 ATP molecules per molecule of substance broken down 2. Aerobic - in presence of oxygen

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Unit 3 Enzymes, Photosynthesis and Respiration

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3. Preparatory conversion: Pyruvate is converted in mitochondrion to acetyl CoA which enters the Krebs cycle. Acetyl CoA is the molecule that links glycolysis and the Krebs cycle. Acetyl CoA is a two carbon molecule. Products for each pyruvic acid: 1 carbon dioxide, 1 NADH, 1 Acetyl CoA 4. Krebs cycle (matrix of mitochondrion) a. Hydrogens and electrons are removed from carbon compounds and used in the electron transport chain to make ATP by chemiosmotic phosphorylation b. For each acetyl CoA that enter the Krebs cycle, two CO2 are released c. Second electron carrier - flavin adenine dinucleotide FAD also used d. ATPs produced in Krebs cycle by substrate level phosphorylation e. 28 - 32 are produced by chemiosmotic phosphorylation - inner membrane of mitochondrion

f. Other molecules (fats, proteins or other carbohydrates) can enter Glycolysis or the Krebs cycle at different points and be used for energy production -proteins digested to amino acids, then amino groups removed (in liver ammonia turned into ures), then can be converted to pyruvate or acetyl CoA; fats release more energy than carbs or proteins (remember dehydrogenase removes Hydrogens!)

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Unit 3 Enzymes, Photosynthesis and Respiration

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g. Intermediate products in the Krebs cycle can be used as building blocks to make other organic molecules that are needed by the organism **if the Krebs cycle does not occur due to a lack of oxygen, then pyruvate will build up. The build up of pyruvate signals muscle cells to undergo lactic acid fermentation to get rid of excess pyruvate and to provide NAD+ so glycolysis can occur. Products: As citric acid is broken down in the Krebs cycle the following products are released (for each molecule of Acetyl CoA that enters): 2 carbon dioxides 3 NADH 1 ATP 1 FADH2 (1 molecule of oxaloacetic acid remains for recycling) ETC takes electrons from NADH and FADH2 to produce about 32 to 34 ATP for each glucose!

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Unit 3 Enzymes, Photosynthesis and Respiration

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D. Anaerobic respiration 1. Occurs in environment with little or no oxygen 2. Purpose to recycle NAD+ to be reused in glycolysis so more ATP can be produced 3. Only ATPs produced in this metabolic pathway are 2 ATPs produced in glycolysis - barely enough to sustain life - inefficient! 4. Two forms: a. Alcohol fermentation - pyruvate broken down to produce ethyl alcohol (ETOH) and CO2 and NAD+ to recycle in glycolysis * Used to make bread rise * Used to make alcoholic beverages * occurs in yeast * Products for 2 pyruvate: 2NAD+, 2 ethyl alcohol, 2CO2 b. Lactic acid fermentation - pyruvate broken down to produce lactate or lactic acid and NAD+ to recycle in glycolysis (NO CO2) *occurs in skeletal muscles of vertebrates and in bacteria *responsible for sour milk *occurs during strenuous exercise when in oxygen debt

AP Bio Gussler Unit 3 Enzymes, Photosynthesis and Respiration Photosynthesis I. Photosynthesis in Nature

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Chloroplasts are the sites of photosynthesis in plants Plants and other autotrophs are the producers of the biosphere (heterotrophs must get their energy from autotrophs) Plants are photosynthetic autotrophs (other organisms are chemosynthetic autotrophs or chemoautotrops) 6CO2 + 12H2O C6H12O6 + 6H2O + 6O2 Leaf structure: epidermis top waxy layer and bottom layer with stoma (not stroma) or stomata (plural); mesophyll palisade mesophyll (for capturing sunlight on top) and spongy mesophyll (for gas exchange); chloroplast stroma (fluid) and thylakoids stacked into grana

I.

The Pathways of Photosynthesis Evidence that chloroplasts split water molecules enabled researchers to track atoms through photosynthesis The light reactions and the Calvin cycle (the two parts of photosynthesis) cooperate in converting light energy to the chemical energy of food

III. The light reactions transform solar energy to chemical energy of ATP and NADPH

The Nature of Sunlight

-Visible portion of spectrum is used in photosynthesis -The amount of energy contained in any kind of light is inversely proportional to the wavelength and directly proportional to the frequency. *violet - shorter wavelength, more energy

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Unit 3 Enzymes, Photosynthesis and Respiration *red - longer wavelength, less energy Photosynthetic pigments

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-Chlorophyll due to its molecular structure absorbs most wavelengths except green and yellow which are reflected. chlorophyll a blue/green chlorophyll b yellow/green -carotenoids (red, orange and yellow) are pigments are present in smaller number. In fall when leaves cease to produce chlorophyll you can see other colors which had been masked by chlorophyll. More pigments absorb more of light spectrum trapping energy. Can also offer photoprotective service Photosystems: Light-harvesting complexes of the thylakoid Non-cyclic electron flow (produces all products necessary) Cyclic Electron flow (produced only ATP) Water is the source of hydrogen needed to reduce carbon dioxide to form sugars. H+ and electrons from chlorophyll are used to produce ATP and NADPH.

Light-Dependent Reactions: (membranes of thylakoid) 1. Wavelengths of light are absorbed by chlorophyll arranged in photosystems. Photosystems are numerous pigment(chlorophyll) and protein molecules And electron acceptor molecules together. 2. Light strikes a photosystem and two electrons are energized and as electrons return to their normal level, the energy can be converted to useful chemical energy for photosynthesis.(*an electron acceptor molecule NADPH nicotinamide adenine dinucleotide phosphate) 3. Electron transport chain: (membranes of thylakoid) a. Photosystem collects enough radiant energy to energize electrons in chlorophyll molecule to higher level. Electron acceptor takes electron (hydrogen becomes electron and protons) b. Electron carrier molecules pass the energized electrons along an electron transport chain where the acquired energy from light is harvested and is used to phosphorylate ADP to ATP c. Protons transported into interior region of thylakoid to create concentration gradient difference. pH gradient that is established across thylakoid membrane creates potential energy to produce ATP. 4. Terminology: a. Photolysis -water splitting which occurs when light energy trapped in electrons H2O 2H + + 2 e- + O2 b. Electron acceptor molecules in photosynthesis: -When hydrogen and its energy removed from chlorophyll it is picked up by acceptor molecule (NADP+) -When NADP+ is in oxidized state it is positively charged, so when it accepts electrons, they unite with the protons to form hydrogen Oxidized state = NADP+ Reduced state = NADPH c. Chlorophyll a and chlorophyll b differ slightly in their chemical structure and absorb different wavelengths of light. Chlorophyll a absorbs colors other than green and chlorophyll b absorbs blue and orange light. Why are plants green? Other pigment such as caroteniods may absorb excess light that would harm the chlorophyll

AP Bio Gussler Unit 3 Enzymes, Photosynthesis and Respiration Page 11 d. Chemiosmosis: Light energy used to create proton concentration gradient. Hydrogen ions would diffuse out but can not cross the membrane except at points bridged by enzyme that produces ATP (ATP synthase). As protons move through enzyme, energy is released. e. Photosystems: like antennae for light waves Photosystem I - 200 to 300 pigment molecules absorbing light in 700 nanometer range Photosystem II - 200 to 300 pigment molecules absorbing 680 nm range 5. Non cyclic Photophosphorylation a. Involves both photosystems b. One way flow of electrons from water to NADP+ c. Reduction of NADP+ requires 2 electrons. Photosystem I can give only one electron at a time and becomes positively charged after doing so. An electron must be restored to it and is donated by Photosystem II

d. Photosystem II: -donates 2 electrons to photosystem I -produces ATP from the proton gradient that is established -gets its missing electrons from the water that is split because P680 absorbs light energy, becomes positively charges and exerts a strong pull on electrons in water. Water electrons donated to P680, protons transferred to NADP+ and oxygen is released e. Photosystem I -donates energy and electrons to reduce NADP+ to NADPH f. Products: NADPH,ATP(from chemiosmosis at CF1 complex), 2H +, 2 e-, O2

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Unit 3 Enzymes, Photosynthesis and Respiration

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6. Cyclic photophosphorylation -when NADP+ or water is low -uses only Photosystem I -produces ONLY ATP (MUST have NADPH for Calvin to be completed, so NO carbs produced) IV. The Calvin-Benson cycle uses ATP and NADPH to convert CO2 to sugar Sugar is high energy glyceraldehyde-3-phosphate, which is a 3 carbon precursor to glucose and other sugars, fats or amino acids (Calvin-Benson is a C-3 pathway of sugar production) Alternative mechanisms of carbon fixation have evolved in hot, arid climates Photorespiration C4 plants CAM Photosynthesis is the biospheres metabolic foundation

Calvin Cycle: 1. Fixing of atmospheric carbon to form organic molecules a. ATP, NADPH and carbon dioxide used to produce high energy compounds b. Calvin-Benson Cycle: C3 cycle -Two turns of cycle required to make one glucose -CARBON FIXATION:RuBP (ribulose biphosphate-5 carbon sugar) combines with CO2 rubisco (RuBP carboylase) is the enzyme that catalizes this reaction -REDUCTION: G-3-P (glyceraldehyde 3 phosphate) can become *glucose, other sugar or organic molecule or *REGENERATION: it can become RuBP to keep cycle going -ADP and NADP+ can be used again in light reactions -For release of one G-3-P for each cycle 9 ATP consumed 6 NADPH consumed

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Unit 3 Enzymes, Photosynthesis and Respiration

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2. C4 cycle (alternative method) a. Can take up carbon dioxide more efficiently than other plants b. Fixing of carbon dioxide occurs not with RuBP, but oxaloacetate a four carbon organic molecule (i.e. C4) c. The structural difference in these plants allow them to enhance concentrations of carbon dioxide d. High temperature environments and higher light optimum (Can be more successful in hot environments because they can keep stomata closed and still undergo photosynthesis.) e. CO2 is fixed on oxaloacetate (4 carbon) and is converted through a series of reactions to pyruvate(3 carbon) and carbon dioxide. This CO2 can be concentrated in areas that the plant needs it to undergo the Calvin Cycle. PEP + CO2 oxaloacetate ( in presence of PEP carboxylase)

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Unit 3 Enzymes, Photosynthesis and Respiration Page 14 3. CAM plants (Crassulacean acid metabolism) (alternative method) a. cacti, pineapple etc (succulents) are adapted to very dry environments b. open stoma only during the night c. fix CO2 onto variety of organic acidslater released for Calvin cycle d. photosynthesis can still occur during the day even though stoma are closed because of the carbon dioxide stored in the plant

climate Examples Enzyme CO2 fixation Stoma Isolation

C3 Normal Most Rubisco PGA Open when enough water None

C4 High temp, high light Corn, sugar cane, crabgrass PEP carboxylase PEP + CO2 oxaloacetate malate (C4) Enough water but shorter time Spatial CO2 absorbed in mesophll; bundle sheath cells undergo conversion to pyruvate and C3 Little O2 in bundle sheat cells so little photoresp. : Increased rate photo allows reduced open time for stomata Conversion Mitochondria Pyruvate Acetyl CoA, CO2 and NADH Krebs Matrix

CAM Very cry Succulents, cactus, pineapple PEP carboxylase PEP + CO2 oxaloacetate Malic acid Only open at night Temporal fixation at night, malic acid stored in vacuoles in mesophyll converted to CO2 when needed

Where Reactants Products

Glycolysis Cytoplasm Glucose ATP, NADH, pyruvate

Actyl CoA and OAA NADH, FADH, ATP, CO2, OAA

ETC Cristae membrane NADH, FADH ATP and O2 and water