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BIG IDEA II Biological systems utilize free energy and molecular building blocks to grow, to reproduce and to maintain dynamic homeostasis. Enduring Understanding 2.A Growth, reproduction and maintenance of the organization of living systems require free energy and matter. Essential Knowledge 2.A.2 Organisms capture and store free energy for use in biological processes. PowerPoint® Lecture Presentations for Biology Eighth Edition Neil Campbell and Jane Reece Lectures by Chris Romero, updated by Erin Barley with contributions from Joan Sharp Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Essential Knowledge 2.A.2: Organisms capture and store free energy for use in biological processes. • Learning Objectives: – (2.4) The student is able to use representations to pose scientific questions about what mechanisms and structural features allow organisms to capture, store and use free energy. – (2.5) The student is able to construct explanations of the mechanisms and structural features of cells that allow organisms to capture, store or use free energy. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Autotrophs capture free energy from physical sources in the environment. • Photosynthetic organisms capture free energy present in sunlight. – 6CO2 + 6 H2O + light energy C6H12O6 + 6 O2 + 6 H2O – carbon dioxide + water + light energy sugar + oxygen + water • Chemosynthetic organisms capture free energy from small inorganic molecules present in their environment, and this process can occur in the absence of oxygen. – 6H2S + 6 H2O + 6 CO2 + 6 O2 C6H12O6 + 6 H2SO4 – hydrogen sulfide + water + carbon dioxide + oxygen sugar + sulfuric acid Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Autotrophs capture free energy from physical sources in the environment. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Heterotrophs capture free energy present in carbon compounds produced by other organisms. • Heterotrophs may metabolize carbohydrates, lipids and proteins by hydrolysis as sources of free energy. – C6H12O6 + 6 O2 6CO2 + 6 H2O + energy (ATP + heat) – organic compounds + oxygen carbon dioxide + water + energy • Fermentation produces organic molecules, including alcohol and lactic acid, and it occurs in the absence of oxygen. – C6H12O6 yeast 2 CH3CH2OH + 2 CO2 + heat – sugar yeast ethanol + carbon dioxide + heat Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Heterotrophs capture free energy present in carbon compounds produced by other organisms. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings The Role of Electron Acceptors in Energy-Capturing Processes • Different energy-capturing processes use different types of electron acceptors: – NADP+ in photosynthesis – Oxygen in cellular respiration • Chemical reactions that transfer electrons between reactants are called oxidation-reduction reactions, or redox reactions. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Catabolic Pathways & ATP Production • Catabolic Pathways yield energy by oxidizing organic fuels. • Several processes are central to cellular respiration and related pathways. • The breakdown of organic molecules is exergonic: – Fermentation is a partial degradation of sugars that occurs without O2. – Aerobic respiration consumes organic molecules and O2 and yields ATP. – Anaerobic respiration is similar to aerobic respiration but consumes compounds other than O2. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Cellular Respiration • Cellular respiration includes both aerobic and anaerobic respiration but is often used to refer to aerobic respiration • Although carbohydrates, fats, and proteins are all consumed as fuel, it is helpful to trace cellular respiration with the sugar glucose: • C6H12O6 + 6 O2 6 CO2 + 6 H2O + Energy (ATP + heat) – The transfer of electrons during chemical reactions releases energy stored in organic molecules – This released energy is ultimately used to synthesize ATP Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Oxidation of Organic Fuel Molecules During Cellular Respiration • During cellular respiration, the fuel (such as glucose) is oxidized, and O2 is reduced: Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Stepwise Energy Harvest via NAD+ and the Electron Transport Chain • In cellular respiration, glucose and other organic molecules are broken down in a series of steps. • Cellular respiration in eukaryotes involves a series of coordinated enzyme-catalyzed reactions that harvest free energy from simple carbohydrates. • Electrons from organic compounds are usually first transferred to NAD+, a coenzyme. • As an electron acceptor, NAD+ functions as an oxidizing agent during cellular respiration. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings The Stages of Cellular Respiration: A Preview • Cellular respiration has three stages: – Glycolysis (breaks down glucose into two molecules of pyruvate) – occurs in cytosol – The citric acid cycle (completes the breakdown of glucose) – occurs in mitochondrial matrix – Electron Transport/Oxidative Phosphorylation (accounts for most of the ATP synthesis) – occurs across inner membrane of mitochondria http://www.sumanasinc.com/webcontent/animations/content/cellularrespiration.html Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Figure 9.16 Review: how each molecule of glucose yields many ATP molecules during cellular respiration: http://www.wadsworthmedia.com/biology/0495119814_starr/big_picture/ch07_bp.html Review Questions from yesterday’s learning: • How are the equations for photosynthesis and chemosynthesis different (details) • What type of reaction is it when organisms break down sugars? • What types of reactions transfer electrons? (Compare the reactions) • Name the three steps in cellular respiration. • What is the net ATP production? Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Mitochondria Structure & Function Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Visual Overview of Cellular Respiration Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Term Comprehension: Substrate-Level Phosphorylation • A smaller amount of ATP is formed in glycolysis and the citric acid cycle by substrate-level phosphorylation. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Glycolysis http://highered.mcgrawhill.com/sites/0072507470/student_view0/chapter25/animation__how_glycolysis_works.html • Glycolysis rearranges the bonds in glucose molecules, releasing free energy to form ATP from ADP and inorganic phosphate, and resulting in the production of pyruvate. • Glycolysis harvests chemical energy by oxidizing glucose to pyruvate – it is the first stage of cellular respiration. • Glycolysis (“splitting of sugar”) breaks down glucose into two molecules of pyruvate. • Glycolysis occurs in the cytoplasm and has two major phases: – Energy investment phase – Energy payoff phase Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Glycolysis • Glycolysis occurs WITH or WITHOUT oxygen. • The first step is the phosphorylation of glucose (glucose molecule gains 2 inorganic phosphates) – this ACTIVATES the glucose to split. • The second step is the splitting of glucose – breaking it down into (2) 3-carbon molecules called pyruvic acid. – This process is achieved by stripping electrons and hydrogens from the unstable 3-C molecules (as well as the borrowed phosphates). • 2 ATPs are needed to produce 4 ATPs (energy investment and energy payoff phases). • A second product in glycolysis is 2 NADH, which results from the transfer of e- and H+ to the coenzyme NAD+. – Occurs in the cytoplasm – Net of 2 ATPs produced – 2 pyruvic acids formed – 2 NADH produced Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 10 Steps of Glycolysis Fig. 9-8 Energy investment phase Glucose 2 ADP + 2 P 2 ATP used 4 ATP formed Energy payoff phase 4 ADP + 4 P 2 NAD+ + 4 e– + 4 H+ 2 NADH + 2 H+ 2 Pyruvate + 2 H2O Net Glucose 4 ATP formed – 2 ATP used 2 NAD+ + 4 e– + 4 H+ 2 Pyruvate + 2 H2O 2 ATP 2 NADH + 2 H+ The “Intermediate” Step • The pyruvate produced during glycolysis is transported from the cytoplasm to the mitochondrion, where further oxidation occurs. • The conversion of pyruvate to acetyl CoA is the junction between glycolysis (step 1) and the Krebs cycle (step 2). • If oxygen is present, Pyruvate (3 C each) from glycolysis enters the mitochondrion. • Using Coenzyme A, each pyruvate is converted into a molecule of Acetyl CoA (2 C each). Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 9-10 CYTOSOL MITOCHONDRION NAD+ NADH + H+ 2 1 Pyruvate Transport protein 3 CO2 Coenzyme A Acetyl CoA The Citric Acid Cycle http://highered.mcgrawhill.com/sites/0072507470/student_view0/chapter25/animation__how_the_krebs_cycle_works__quiz_1_.html • In the Krebs cycle, carbon dioxide is released from organic intermediations ATP is synthesized from ADP and inorganic phosphate via substrate level phosphorylation and electrons are captured by coenzymes. • The citric acid (Krebs) cycle completes the energyyielding oxidation of organic molecules – and its events take place within the mitochondrial matrix. • The cycle oxidizes organic fuel derived from pyruvate, generating 1 ATP, 3 NADH, and 1 FADH2 per turn. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 9-11 Pyruvate CO2 NAD+ CoA NADH + H+ Acetyl CoA CoA CoA Citric acid cycle FADH2 2 CO2 3 NAD+ 3 NADH FAD + 3 H+ ADP + P i ATP Term Comprehension: Oxidative Phosphorylation • The process that generates most of the ATP during cellular respiration is called oxidative phosphorylation because it is powered by redox reactions of an electron transport chain. • Oxidative phosphorylation accounts for almost 90% of the ATP generated by cellular respiration. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings • What is the literal translation of glycolysis and why is it an appropriate name? • How many ATP are used and created from the process of glycolysis? • What is the intermediate product made in the citric acid cycle? • How many cycles are involved in the completion of the citric acid cycle? • What are the electron carriers which are produced in the first two portions of cellular respiration? Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Chemiosmosis & Electron Transport http://highered.mcgrawhill.com/sites/0072507470/student_view0/chapter25/animation__electron_transport_system_and_atp_synthesis__quiz_1_.html • Following the Krebs cycle, the electrons captured by NADH and FADH2 are passed to the electron transport chain: – The electron transport chain uses the high-energy electrons from the Krebs cycle to convert ADP to ATP. – Every time 2 high energy electrons transport down the ETC, their energy is used to transport H+ across the inner membrane of the mitochondria…this creates a + charge on the inside of the membrane and a – charge in the matrix of the mitochondria. – As a result of this charge difference, H+ ions escape through channel proteins called ATP synthase causing it to rotate. – Each time it rotates, the enzyme ATP synthase grabs a low energy ADP and attaches a phosphate, forming highenergy ATP. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings NADH 50 2 e– NAD+ FADH2 2 e– 40 FMN FAD Multiprotein complexes FAD Fe•S Fe•S Q Cyt b 30 Fe•S Cyt c1 I V Cyt c Cyt a Cyt a3 20 10 2 e– (from NADH or FADH2) 0 2 H+ + 1/2 O2 H2O • The electron transport chain captures free energy from electrons in a series of coupled reactions that establish an electrochemical gradient across membranes. • Electrons delivered by NADH and FADH2 are passed to a series of electron acceptors as they move toward the terminal electron acceptor, oxygen. • The passage of electrons is accompanied by the formation of a proton gradient across the inner mitochondrial membrane, with the membrane separating a region of high proton concentration from a region of low proton concentration. • The flow of protons back through membrane-bound ATP synthase by chemiosmosis generates ATP from ADP and inorganic phosphate (Pi). Fig. 9-14 INTERMEMBRANE SPACE H+ Stator Rotor Internal rod Catalytic knob ADP + P i ATP MITOCHONDRIAL MATRIX Fig. 9-16 H+ H+ H+ H+ Protein complex of electron carriers Cyt c V Q ATP synthase FADH2 NADH 2 H+ + 1/2O2 H2O FAD NAD+ ADP + P i (carrying electrons from food) ATP H+ 1 Electron transport chain Oxidative phosphorylation 2 Chemiosmosis Fig. 9-17 Electron shuttles span membrane CYTOSOL 2 NADH Glycolysis Glucose 2 Pyruvate MITOCHONDRION 2 NADH or 2 FADH2 6 NADH 2 NADH 2 Acetyl CoA + 2 ATP Citric acid cycle + 2 ATP Maximum per glucose: About 36 or 38 ATP 2 FADH2 Oxidative phosphorylation: electron transport and chemiosmosis + about 32 or 34 ATP Fermentation/Anaerobic Respiration • Fermentation and anaerobic respiration enable cells to produce ATP without the use of oxygen • Most cellular respiration requires O2 to produce ATP • Glycolysis can produce ATP with or without O2 (in aerobic or anaerobic conditions) • In the absence of O2, glycolysis couples with fermentation or anaerobic respiration to produce ATP – Anaerobic respiration uses an electron transport chain with an electron acceptor other than O2, for example sulfate – Fermentation uses phosphorylation instead of an electron transport chain to generate ATP Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Types of Fermentation • Fermentation consists of glycolysis plus reactions that regenerate NAD+, which can be reused by glycolysis – Two common types are alcohol fermentation and lactic acid fermentation Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 9-18 2 ADP + 2 Pi Glucose 2 ATP Glycolysis 2 Pyruvate 2 NAD+ 2 NADH + 2 H+ 2 CO2 2 Acetaldehyde 2 Ethanol (a) Alcohol fermentation 2 ADP + 2 Pi Glucose 2 ATP Glycolysis 2 NAD+ 2 NADH + 2 H+ 2 Pyruvate 2 Lactate (b) Lactic acid fermentation Fermentation and Aerobic Respiration Compared • Both processes use glycolysis to oxidize glucose and other organic fuels to pyruvate • The processes have different final electron acceptors: an organic molecule (such as pyruvate or acetaldehyde) in fermentation and O2 in cellular respiration • Cellular respiration produces 38 ATP per glucose molecule; fermentation produces 2 ATP per glucose molecule Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings The Anaerobes • Obligate anaerobes carry out fermentation or anaerobic respiration and cannot survive in the presence of O2 • Yeast and many bacteria are facultative anaerobes, meaning that they can survive using either fermentation or cellular respiration – In a facultative anaerobe, pyruvate is a fork in the metabolic road that leads to two alternative catabolic routes Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings The Versatility of Catabolism • Glycolysis and the citric acid cycle are major intersections to various catabolic and anabolic pathways. • Catabolic pathways funnel electrons from many kinds of organic molecules into cellular respiration. • Glycolysis accepts a wide range of carbohydrates. • In addition to carbohydrates, heterotrophs may metabolize lipids and proteins by hydrolysis as sources of free energy. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 9-20 Proteins Amino acids Carbohydrates Sugars Glycolysis Glucose Glyceraldehyde-3- P NH3 Pyruvate Acetyl CoA Citric acid cycle Oxidative phosphorylation Fats Glycerol Fatty acids Fig. 9-21 Glucose AMP Glycolysis Fructose-6-phosphate – Stimulates + Phosphofructokinase – Fructose-1,6-bisphosphate Inhibits Inhibits Pyruvate ATP Citrate Acetyl CoA Citric acid cycle Oxidative phosphorylation What do you know…. • Glycolysis • Kreb Cycle • ETC • Fermentation Pathways Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Overview of Cellular Respiration A. Glycolysis – *takes place in cytosol *breaks glucose into 2 molecules of pyruvate *produces a net of 2 ATP’s B. Krebs cycle - *takes place in mitochondrial matrix *makes a derivative of pyruvate into carbon dioxide *produces a net of 2 ATP’s C. ETC and Oxidative Phosphorylation – *takes place in inner membrane of mitochondrion *accepts e-’s from A and B via NADH *at end, e-’s are combined with H+ and oxygen to form water *forms a net of 34 ATP’s D. Fermentation Pathways *takes place in cytosol *2 types: Alcoholic and Lactic Acid Fermentation *stores most of the energy in chemical bonds *no production of ATP Energy Coupling H2O Following cellular respiration or fermentation, free energy becomes available for metabolism by the conversion of ATPADP, which is coupled to many steps in metabolic pathways. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings