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Electron shuttles                                                MITOCHONDRION
           span membrane               2 NADH
                                          or
                                       2 FADH2

      2 NADH                               2 NADH                  6 NADH   2 FADH2



     Glycolysis                Pyruvate oxidation                                     Oxidative
                                                        Citric                    phosphorylation:
Glucose      2 Pyruvate          2 Acetyl CoA           acid                      electron transport
                                                        cycle                            and
                                                                                    chemiosmosis




          + 2 ATP                                       + 2 ATP                   + about 26 or 28 ATP


                                                       About
                           Maximum per glucose:     30 or 32 ATP

CYTOSOL




                      Cellular Respiration

                                     Dr. Kristen Walker
The Structure and Hydrolysis of ATP
 • ATP (adenosine triphosphate) is cell’s energy
   shuttle
 • ATP is composed of ribose (a sugar), adenine
   (a nitrogenous base), and three phosphate
   groups
Figure 8.8a




                             Adenine




          Phosphate groups
                                 Ribose


 (a) The structure of ATP
• The bonds between the phosphate groups
  of ATP’s tail can be broken by hydrolysis
• Energy is released from ATP when the
  terminal phosphate bond is broken
• This release of energy comes from the
  chemical change to a state of lower free
  energy, not from the phosphate bonds
  themselves
Figure 8.8b




               Adenosine triphosphate (ATP)




                                               Energy


   Inorganic     Adenosine diphosphate (ADP)
   phosphate

 (b) The hydrolysis of ATP
How the Hydrolysis of ATP Performs Work
 • The three types of cellular work (mechanical,
   transport, and chemical) are powered by the
   hydrolysis of ATP
 • In the cell, the energy from the exergonic reaction
   (release of free energy) of ATP hydrolysis can be
   used to drive an endergonic reaction (absorbs free
   energy)
 • Overall, the coupled reactions are exergonic
• ATP drives endergonic reactions by phosphorylation,
  transferring a phosphate group to some other
  molecule, such as a reactant
• The recipient molecule is now called a
  phosphorylated intermediate
Figure 8.10
                      Transport protein             Solute



                ATP                                               ADP   Pi

                                    P                    Pi

                                            Solute transported
              (a) Transport work: ATP phosphorylates transport proteins.

                      Vesicle                Cytoskeletal track




                ATP                                               ADP   Pi
                                  ATP




                            Motor protein      Protein and
                                              vesicle moved
              (b) Mechanical work: ATP binds noncovalently to motor
                  proteins and then is hydrolyzed.
The Regeneration of ATP
 • ATP is a renewable resource that is regenerated by
   addition of a phosphate group to adenosine
   diphosphate (ADP)
 • Energy to phosphorylate ADP comes from catabolic
   reactions in the cell
    • Catabolic - release energy
    • Anabolic - require energy
 • ATP cycle is a revolving door through which energy
   passes during its transfer from catabolic to anabolic
   pathways
Figure 8.11




                          ATP    H2O




 Energy from                           Energy for cellular
 catabolism (exergonic,                work (endergonic,
 energy-releasing         ADP   Pi     energy-consuming
 processes)                            processes)
Life Is Work
 • Living cells require energy from outside
   sources
 • Some animals, such as the chimpanzee, obtain
   energy by eating plants, and some animals
   feed on other organisms that eat plants
• Energy flows into an ecosystem as sunlight
  and leaves as heat
• Photosynthesis generates O2 and organic
  molecules, which are used in cellular
  respiration
• Cells use chemical energy stored in organic
  molecules to regenerate ATP, which powers
  work
Figure 9.2

                      Light
                      energy

             ECOSYSTEM



                             Photosynthesis
                             in chloroplasts
                CO2 + H2O                           Organic
                                                             + O2
                                                   molecules
                            Cellular respiration
                             in mitochondria




                                          ATP powers
                                   ATP
                                          most cellular work

                         Heat
                         energy
Catabolic pathways yield energy by
oxidizing organic fuels
 • Several processes are central to cellular
   respiration and related pathways
Catabolic Pathways and Production of ATP

 • 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
• 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)
Redox Reactions: Oxidation and Reduction
 • The transfer of electrons during chemical
   reactions releases energy stored in organic
   molecules
 • This released energy is ultimately used to
   synthesize ATP
The Principle of Redox
 • Chemical reactions that transfer electrons
   between reactants are called oxidation-reduction
   reactions, or redox reactions
 • In oxidation, a substance loses electrons, or is
   oxidized
 • In reduction, a substance gains electrons, or is
   reduced (the amount of positive charge is
   reduced)
Figure 9.UN01




                becomes oxidized
                 (loses electron)

                          becomes reduced
                           (gains electron)
Oxidation of Organic Fuel Molecules During
Cellular Respiration
 • During cellular respiration, the fuel (such as
   glucose) is oxidized, and O2 is reduced
Figure 9.UN03




                becomes oxidized


                             becomes reduced
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
 • 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
 • Each NADH (the reduced form of NAD+)
   represents stored energy that is tapped to
   synthesize ATP
Figure 9.4




             NAD+                                                        NADH
                                                     Dehydrogenase
                                                    Reduction of NAD+

                                      (from food)   Oxidation of NADH
                    Nicotinamide                                        Nicotinamide
                    (oxidized form)                                     (reduced form)
• NADH passes the electrons to the electron
  transport chain
• Unlike an uncontrolled reaction, the electron
  transport chain passes electrons in a series of
  steps instead of one explosive reaction
• O2 pulls electrons down the chain in an energy-
  yielding tumble
• The energy yielded is used to regenerate ATP
The Stages of Cellular Respiration:
A Preview
 • Harvesting of energy from glucose has three
   stages
     – Glycolysis (breaks down glucose into two
       molecules of pyruvate)
     – The citric acid cycle (completes the
       breakdown of glucose) also called Krebs
       cycle
     – Oxidative phosphorylation (accounts for
       most of the ATP synthesis)
Figure 9.UN05




 1. Glycolysis (color-coded teal throughout the chapter)
 2. Pyruvate oxidation and the citric acid cycle
    (color-coded salmon)
 3. Oxidative phosphorylation: electron transport and
    chemiosmosis (color-coded violet)
Figure 9.6-1




               Electrons
                carried
               via NADH




               Glycolysis

    Glucose            Pyruvate




   CYTOSOL                        MITOCHONDRION




                 ATP

         Substrate-level
        phosphorylation
Figure 9.6-2




               Electrons                                Electrons carried
                carried                                  via NADH and
               via NADH                                      FADH2



                                   Pyruvate
               Glycolysis                           Citric
                                   oxidation
                                                    acid
    Glucose            Pyruvate   Acetyl CoA        cycle




   CYTOSOL                            MITOCHONDRION




                 ATP                                ATP

         Substrate-level                        Substrate-level
        phosphorylation                        phosphorylation
Figure 9.6-3




               Electrons                                Electrons carried
                carried                                  via NADH and
               via NADH                                      FADH2



                                                                         Oxidative
                                   Pyruvate
               Glycolysis                           Citric           phosphorylation:
                                   oxidation
                                                    acid             electron transport
    Glucose            Pyruvate   Acetyl CoA        cycle                   and
                                                                       chemiosmosis



   CYTOSOL                            MITOCHONDRION




                 ATP                                ATP                          ATP

         Substrate-level                        Substrate-level               Oxidative
        phosphorylation                        phosphorylation              phosphorylation
• The process that generates most of the ATP is
  called oxidative phosphorylation because it is
  powered by redox reactions




                 BioFlix: Cellular Respiration
• Oxidative phosphorylation accounts for almost
  90% of the ATP generated by cellular
  respiration
• A smaller amount of ATP is formed in glycolysis
  and the citric acid cycle
• For each molecule of glucose degraded to CO2
  and water by respiration, the cell makes up to
  32 molecules of ATP
Glycolysis harvests chemical energy by
oxidizing glucose to pyruvate
 • 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
 • Glycolysis occurs whether or not O2 is present
Figure 9.8

             Energy Investment Phase
                                 Glucose



                           2 ADP + 2 P          2 ATP used



             Energy Payoff Phase

                           4 ADP + 4 P          4 ATP formed


                   2 NAD+ + 4 e− + 4 H+         2 NADH + 2 H+


                                               2 Pyruvate + 2 H2O

             Net
                                 Glucose    2 Pyruvate + 2 H2O
             4 ATP formed − 2 ATP used      2 ATP
                     2 NAD+ + 4 e− + 4 H+   2 NADH + 2 H+
After pyruvate is oxidized, the citric acid
cycle completes the energy-yielding oxidation
of organic molecules
 • In the presence of O2, pyruvate enters the
   mitochondrion (in eukaryotic cells) where the
   oxidation of glucose is completed
Oxidation of Pyruvate to Acetyl CoA
 • Before the citric acid cycle can begin, pyruvate
   must be converted to acetyl Coenzyme A
   (acetyl CoA), which links glycolysis to the citric
   acid cycle
 • This step is carried out by a multi-enzyme
   complex that catalyses three reactions
Figure 9.10




                                        MITOCHONDRION
              CYTOSOL       CO2    Coenzyme A

                        1                   3



                                   2

                            NAD+       NADH + H+   Acetyl CoA
   Pyruvate

   Transport protein
The Citric Acid Cycle
 • The citric acid cycle, also called the Krebs
   cycle, completes the break down of pyruvate
   to CO2
 • The cycle oxidizes organic fuel derived from
   pyruvate, generating 1 ATP, 3 NADH, and 1
   FADH2 per turn
 • The citric acid cycle has eight steps, each
   catalyzed by a specific enzyme
Figure 9.11
              Pyruvate

                                                 CO2
                          NAD   +

                                                 CoA
                         NADH
                         + H+       Acetyl CoA
                                           CoA


                                                  CoA




                                         Citric
                                         acid
                                         cycle            2 CO2

                 FADH2                                    3 NAD+

                    FAD                                  3 NADH
                                                          + 3 H+
                                             ADP + P i

                                      ATP
During oxidative phosphorylation,
chemiosmosis couples electron transport to
ATP synthesis
  • Following glycolysis and the citric acid cycle,
    NADH and FADH2 account for most of the
    energy extracted from food
  • These two electron carriers donate electrons to
    the electron transport chain, which powers ATP
    synthesis via oxidative phosphorylation
The Pathway of Electron Transport
 • The electron transport chain is in the inner
   membrane (cristae) of the mitochondrion
 • Most of the chain’s components are proteins,
   which exist in multiprotein complexes
 • The carriers alternate reduced and oxidized
   states as they accept and donate electrons
 • Electrons drop in free energy as they go down
   the chain and are finally passed to O2, forming
   H2O
Figure 9.13
                                                                 NADH
                                                          50
                                                               2 e−
                                                                      NAD+
                                                                                     FADH2

                                                                                 2 e−    FAD                   Multiprotein




              Free energy (G) relative to O2 (kcal/mol)
                                                          40   FMN
                                                                             I                                 complexes
                                                                                 Fe• S   II
                                                                       Fe• S
                                                                                 Q
                                                                                                     III
                                                                                     Cyt b
                                                                                             Fe• S
                                                          30
                                                                                                     Cyt c1
                                                                                                                                 IV
                                                                                                              Cyt c
                                                                                                                      Cyt a
                                                                                                                              Cyt a3
                                                          20




                                                                                                                              2 e−
                                                          10
                                                                                                       (originally from
                                                                                                       NADH or FADH2)


                                                           0                                                     2 H+ + 1/2 O2




                                                                                                                                 H2O
• Electrons are transferred from NADH or FADH2
  to the electron transport chain
• Electrons are passed through a number of
  proteins including cytochromes (each with an
  iron atom) to O2
• The electron transport chain generates no ATP
  directly
• It breaks the large free-energy drop from food to
  O2 into smaller steps that release energy in
  manageable amounts
Chemiosmosis: The Energy-Coupling
Mechanism
 • Electron transfer in the electron transport chain
   causes proteins to pump H+ from the
   mitochondrial matrix to the intermembrane space
 • H+ then moves back across the membrane,
   passing through the proton, ATP synthase
 • ATP synthase uses the exergonic flow of H+ to
   drive phosphorylation of ATP
 • This is an example of chemiosmosis, the use of
   energy in a H+ gradient to drive cellular work
Figure 9.14
               INTERMEMBRANE SPACE


                              H+
                                      Stator
                     Rotor




               Internal
               rod

              Catalytic
              knob



                 ADP
                  +
                  Pi               ATP

               MITOCHONDRIAL MATRIX
Figure 9.15




                                                                        H+
                                         H   +




  Protein     H   +


  complex                                                                                      H+
                                                     Cyt c
  of electron
  carriers
                                                                   IV
                                  Q
                                      III
              I
                                                                                                    ATP
                             II                                                                     synth-
                                                      2 H + + 1 /2 O2        H2O                    ase
                             FADH2 FAD

    NADH              NAD+
                                                                                   ADP + P i        ATP
  (carrying electrons
  from food)                                                                                   H+

                             1 Electron transport chain                                2 Chemiosmosis
                                                 Oxidative phosphorylation
• The energy stored in a H+ gradient across a
  membrane couples the redox reactions of the
  electron transport chain to ATP synthesis
• The H+ gradient is referred to as a proton-
  motive force, emphasizing its capacity to do
  work
An Accounting of ATP Production by
Cellular Respiration
 • During cellular respiration, most energy
   flows in this sequence:
   glucose → NADH → electron transport
   chain → proton-motive force → ATP
 • About 34% of the energy in a glucose
   molecule is transferred to ATP during
   cellular respiration, making about 32 ATP
Figure 9.16




               Electron shuttles                                                  MITOCHONDRION
               span membrane                2 NADH
                                               or
                                            2 FADH2

              2 NADH                             2 NADH                  6 NADH   2 FADH2



          Glycolysis                Pyruvate oxidation                                      Oxidative
                                                              Citric                    phosphorylation:
   Glucose       2 Pyruvate            2 Acetyl CoA           acid                      electron transport
                                                              cycle                            and
                                                                                          chemiosmosis




              + 2 ATP                                         + 2 ATP                   + about 26 or 28 ATP


                                                             About
                                   Maximum per glucose:   30 or 32 ATP

  CYTOSOL
Fermentation and anaerobic
respiration enable cells to produce ATP
without the use of oxygen
 • Most cellular respiration requires O2 to produce
   ATP
 • Without O2, the electron transport chain will
   cease to operate
 • In that case, glycolysis couples with
   fermentation or anaerobic respiration to
   produce ATP
• Anaerobic respiration uses an electron
  transport chain with a final electron acceptor
  other than O2, for example sulfate
• Fermentation uses substrate-level
  phosphorylation instead of an electron
  transport chain to generate ATP
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
• In alcohol fermentation, pyruvate is converted
  to ethanol in two steps, with the first releasing
  CO2
• Alcohol fermentation by yeast is used in
  brewing, winemaking, and baking
Figure 9.17a



                           2 ADP + 2 P i         2 ATP



                Glucose             Glycolysis


                                                             2 Pyruvate

                             2 NAD +           2 NADH               2 CO2
                                             + 2 H+




               2 Ethanol                                 2 Acetaldehyde

       (a) Alcohol fermentation
• In lactic acid fermentation, pyruvate is reduced
  to NADH, forming lactate as an end product,
  with no release of CO2
• Lactic acid fermentation by some fungi and
  bacteria is used to make cheese and yogurt
• Human muscle cells use lactic acid
  fermentation to generate ATP when O2 is
  scarce
Figure 9.17b



                           2 ADP + 2 P i         2 ATP



                Glucose             Glycolysis




                             2 NAD +          2 NADH
                                            + 2 H+
                                                         2 Pyruvate




               2 Lactate

          (b) Lactic acid fermentation
Comparing Fermentation with Anaerobic
and Aerobic Respiration
 • All use glycolysis (net ATP = 2) to oxidize glucose
   and harvest chemical energy of food
 • In all three, NAD+ is the oxidizing agent that
   accepts electrons during glycolysis
 • 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 32 ATP per glucose
   molecule; fermentation produces 2 ATP per
   glucose molecule
• 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
Figure 9.18
                                 Glucose


                                      Glycolysis
               CYTOSOL

                                 Pyruvate
                No O2 present:          O2 present:
                Fermentation             Aerobic cellular
                                           respiration



                                                         MITOCHONDRION
                 Ethanol,                   Acetyl CoA
                lactate, or
              other products
                                                         Citric
                                                         acid
                                                         cycle
The Evolutionary Significance of Glycolysis
 • Ancient prokaryotes are thought to have used
   glycolysis long before there was oxygen in the
   atmosphere
 • Very little O2 was available in the atmosphere
   until about 2.7 billion years ago, so early
   prokaryotes likely used only glycolysis to
   generate ATP
 • Glycolysis is a very ancient process
Glycolysis and the citric acid cycle connect
to many other metabolic pathways

  • Gycolysis and the citric acid cycle are major
    intersections to various catabolic and anabolic
    pathways
The Versatility of Catabolism
 • Catabolic pathways funnel electrons from many
   kinds of organic molecules into cellular
   respiration
 • Glycolysis accepts a wide range of
   carbohydrates
 • Proteins must be digested to amino acids;
   amino groups can feed glycolysis or the citric
   acid cycle
• Fats are digested to glycerol (used in
  glycolysis) and fatty acids (used in generating
  acetyl CoA)
• Fatty acids are broken down by beta oxidation
  and yield acetyl CoA
• An oxidized gram of fat produces more than
  twice as much ATP as an oxidized gram of
  carbohydrate
Figure 9.19
                    Proteins     Carbohydrates            Fats


                    Amino           Sugars           Glycerol Fatty
                    acids                                     acids


                                   Glycolysis
                                    Glucose


                               Glyceraldehyde 3- P

              NH3                   Pyruvate


                                  Acetyl CoA




                                      Citric
                                      acid
                                      cycle




                                   Oxidative
                                phosphorylation
Biosynthesis (Anabolic Pathways)
 • The body uses small molecules to build other
   substances
 • These small molecules may come directly
   from food, from glycolysis, or from the citric
   acid cycle
Regulation of Cellular Respiration via
Feedback Mechanisms
 • Feedback inhibition is the most common
   mechanism for control
 • If ATP concentration begins to drop,
   respiration speeds up; when there is plenty
   of ATP, respiration slows down
 • Control of catabolism is based mainly on
   regulating the activity of enzymes at
   strategic points in the catabolic pathway

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Week 9

  • 1. Electron shuttles MITOCHONDRION span membrane 2 NADH or 2 FADH2 2 NADH 2 NADH 6 NADH 2 FADH2 Glycolysis Pyruvate oxidation Oxidative Citric phosphorylation: Glucose 2 Pyruvate 2 Acetyl CoA acid electron transport cycle and chemiosmosis + 2 ATP + 2 ATP + about 26 or 28 ATP About Maximum per glucose: 30 or 32 ATP CYTOSOL Cellular Respiration Dr. Kristen Walker
  • 2. The Structure and Hydrolysis of ATP • ATP (adenosine triphosphate) is cell’s energy shuttle • ATP is composed of ribose (a sugar), adenine (a nitrogenous base), and three phosphate groups
  • 3. Figure 8.8a Adenine Phosphate groups Ribose (a) The structure of ATP
  • 4. • The bonds between the phosphate groups of ATP’s tail can be broken by hydrolysis • Energy is released from ATP when the terminal phosphate bond is broken • This release of energy comes from the chemical change to a state of lower free energy, not from the phosphate bonds themselves
  • 5. Figure 8.8b Adenosine triphosphate (ATP) Energy Inorganic Adenosine diphosphate (ADP) phosphate (b) The hydrolysis of ATP
  • 6. How the Hydrolysis of ATP Performs Work • The three types of cellular work (mechanical, transport, and chemical) are powered by the hydrolysis of ATP • In the cell, the energy from the exergonic reaction (release of free energy) of ATP hydrolysis can be used to drive an endergonic reaction (absorbs free energy) • Overall, the coupled reactions are exergonic
  • 7. • ATP drives endergonic reactions by phosphorylation, transferring a phosphate group to some other molecule, such as a reactant • The recipient molecule is now called a phosphorylated intermediate
  • 8. Figure 8.10 Transport protein Solute ATP ADP Pi P Pi Solute transported (a) Transport work: ATP phosphorylates transport proteins. Vesicle Cytoskeletal track ATP ADP Pi ATP Motor protein Protein and vesicle moved (b) Mechanical work: ATP binds noncovalently to motor proteins and then is hydrolyzed.
  • 9. The Regeneration of ATP • ATP is a renewable resource that is regenerated by addition of a phosphate group to adenosine diphosphate (ADP) • Energy to phosphorylate ADP comes from catabolic reactions in the cell • Catabolic - release energy • Anabolic - require energy • ATP cycle is a revolving door through which energy passes during its transfer from catabolic to anabolic pathways
  • 10. Figure 8.11 ATP H2O Energy from Energy for cellular catabolism (exergonic, work (endergonic, energy-releasing ADP Pi energy-consuming processes) processes)
  • 11. Life Is Work • Living cells require energy from outside sources • Some animals, such as the chimpanzee, obtain energy by eating plants, and some animals feed on other organisms that eat plants
  • 12. • Energy flows into an ecosystem as sunlight and leaves as heat • Photosynthesis generates O2 and organic molecules, which are used in cellular respiration • Cells use chemical energy stored in organic molecules to regenerate ATP, which powers work
  • 13. Figure 9.2 Light energy ECOSYSTEM Photosynthesis in chloroplasts CO2 + H2O Organic + O2 molecules Cellular respiration in mitochondria ATP powers ATP most cellular work Heat energy
  • 14. Catabolic pathways yield energy by oxidizing organic fuels • Several processes are central to cellular respiration and related pathways
  • 15. Catabolic Pathways and Production of ATP • 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
  • 16. • 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)
  • 17. Redox Reactions: Oxidation and Reduction • The transfer of electrons during chemical reactions releases energy stored in organic molecules • This released energy is ultimately used to synthesize ATP
  • 18. The Principle of Redox • Chemical reactions that transfer electrons between reactants are called oxidation-reduction reactions, or redox reactions • In oxidation, a substance loses electrons, or is oxidized • In reduction, a substance gains electrons, or is reduced (the amount of positive charge is reduced)
  • 19. Figure 9.UN01 becomes oxidized (loses electron) becomes reduced (gains electron)
  • 20. Oxidation of Organic Fuel Molecules During Cellular Respiration • During cellular respiration, the fuel (such as glucose) is oxidized, and O2 is reduced
  • 21. Figure 9.UN03 becomes oxidized becomes reduced
  • 22. 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 • 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 • Each NADH (the reduced form of NAD+) represents stored energy that is tapped to synthesize ATP
  • 23. Figure 9.4 NAD+ NADH Dehydrogenase Reduction of NAD+ (from food) Oxidation of NADH Nicotinamide Nicotinamide (oxidized form) (reduced form)
  • 24. • NADH passes the electrons to the electron transport chain • Unlike an uncontrolled reaction, the electron transport chain passes electrons in a series of steps instead of one explosive reaction • O2 pulls electrons down the chain in an energy- yielding tumble • The energy yielded is used to regenerate ATP
  • 25. The Stages of Cellular Respiration: A Preview • Harvesting of energy from glucose has three stages – Glycolysis (breaks down glucose into two molecules of pyruvate) – The citric acid cycle (completes the breakdown of glucose) also called Krebs cycle – Oxidative phosphorylation (accounts for most of the ATP synthesis)
  • 26. Figure 9.UN05 1. Glycolysis (color-coded teal throughout the chapter) 2. Pyruvate oxidation and the citric acid cycle (color-coded salmon) 3. Oxidative phosphorylation: electron transport and chemiosmosis (color-coded violet)
  • 27. Figure 9.6-1 Electrons carried via NADH Glycolysis Glucose Pyruvate CYTOSOL MITOCHONDRION ATP Substrate-level phosphorylation
  • 28. Figure 9.6-2 Electrons Electrons carried carried via NADH and via NADH FADH2 Pyruvate Glycolysis Citric oxidation acid Glucose Pyruvate Acetyl CoA cycle CYTOSOL MITOCHONDRION ATP ATP Substrate-level Substrate-level phosphorylation phosphorylation
  • 29. Figure 9.6-3 Electrons Electrons carried carried via NADH and via NADH FADH2 Oxidative Pyruvate Glycolysis Citric phosphorylation: oxidation acid electron transport Glucose Pyruvate Acetyl CoA cycle and chemiosmosis CYTOSOL MITOCHONDRION ATP ATP ATP Substrate-level Substrate-level Oxidative phosphorylation phosphorylation phosphorylation
  • 30. • The process that generates most of the ATP is called oxidative phosphorylation because it is powered by redox reactions BioFlix: Cellular Respiration
  • 31. • Oxidative phosphorylation accounts for almost 90% of the ATP generated by cellular respiration • A smaller amount of ATP is formed in glycolysis and the citric acid cycle • For each molecule of glucose degraded to CO2 and water by respiration, the cell makes up to 32 molecules of ATP
  • 32. Glycolysis harvests chemical energy by oxidizing glucose to pyruvate • 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 • Glycolysis occurs whether or not O2 is present
  • 33. Figure 9.8 Energy Investment Phase Glucose 2 ADP + 2 P 2 ATP used Energy Payoff Phase 4 ADP + 4 P 4 ATP formed 2 NAD+ + 4 e− + 4 H+ 2 NADH + 2 H+ 2 Pyruvate + 2 H2O Net Glucose 2 Pyruvate + 2 H2O 4 ATP formed − 2 ATP used 2 ATP 2 NAD+ + 4 e− + 4 H+ 2 NADH + 2 H+
  • 34. After pyruvate is oxidized, the citric acid cycle completes the energy-yielding oxidation of organic molecules • In the presence of O2, pyruvate enters the mitochondrion (in eukaryotic cells) where the oxidation of glucose is completed
  • 35. Oxidation of Pyruvate to Acetyl CoA • Before the citric acid cycle can begin, pyruvate must be converted to acetyl Coenzyme A (acetyl CoA), which links glycolysis to the citric acid cycle • This step is carried out by a multi-enzyme complex that catalyses three reactions
  • 36. Figure 9.10 MITOCHONDRION CYTOSOL CO2 Coenzyme A 1 3 2 NAD+ NADH + H+ Acetyl CoA Pyruvate Transport protein
  • 37. The Citric Acid Cycle • The citric acid cycle, also called the Krebs cycle, completes the break down of pyruvate to CO2 • The cycle oxidizes organic fuel derived from pyruvate, generating 1 ATP, 3 NADH, and 1 FADH2 per turn • The citric acid cycle has eight steps, each catalyzed by a specific enzyme
  • 38. Figure 9.11 Pyruvate CO2 NAD + CoA NADH + H+ Acetyl CoA CoA CoA Citric acid cycle 2 CO2 FADH2 3 NAD+ FAD 3 NADH + 3 H+ ADP + P i ATP
  • 39. During oxidative phosphorylation, chemiosmosis couples electron transport to ATP synthesis • Following glycolysis and the citric acid cycle, NADH and FADH2 account for most of the energy extracted from food • These two electron carriers donate electrons to the electron transport chain, which powers ATP synthesis via oxidative phosphorylation
  • 40. The Pathway of Electron Transport • The electron transport chain is in the inner membrane (cristae) of the mitochondrion • Most of the chain’s components are proteins, which exist in multiprotein complexes • The carriers alternate reduced and oxidized states as they accept and donate electrons • Electrons drop in free energy as they go down the chain and are finally passed to O2, forming H2O
  • 41. Figure 9.13 NADH 50 2 e− NAD+ FADH2 2 e− FAD Multiprotein Free energy (G) relative to O2 (kcal/mol) 40 FMN I complexes Fe• S II Fe• S Q III Cyt b Fe• S 30 Cyt c1 IV Cyt c Cyt a Cyt a3 20 2 e− 10 (originally from NADH or FADH2) 0 2 H+ + 1/2 O2 H2O
  • 42. • Electrons are transferred from NADH or FADH2 to the electron transport chain • Electrons are passed through a number of proteins including cytochromes (each with an iron atom) to O2 • The electron transport chain generates no ATP directly • It breaks the large free-energy drop from food to O2 into smaller steps that release energy in manageable amounts
  • 43. Chemiosmosis: The Energy-Coupling Mechanism • Electron transfer in the electron transport chain causes proteins to pump H+ from the mitochondrial matrix to the intermembrane space • H+ then moves back across the membrane, passing through the proton, ATP synthase • ATP synthase uses the exergonic flow of H+ to drive phosphorylation of ATP • This is an example of chemiosmosis, the use of energy in a H+ gradient to drive cellular work
  • 44. Figure 9.14 INTERMEMBRANE SPACE H+ Stator Rotor Internal rod Catalytic knob ADP + Pi ATP MITOCHONDRIAL MATRIX
  • 45. Figure 9.15 H+ H + Protein H + complex H+ Cyt c of electron carriers IV Q III I ATP II synth- 2 H + + 1 /2 O2 H2O ase FADH2 FAD NADH NAD+ ADP + P i ATP (carrying electrons from food) H+ 1 Electron transport chain 2 Chemiosmosis Oxidative phosphorylation
  • 46. • The energy stored in a H+ gradient across a membrane couples the redox reactions of the electron transport chain to ATP synthesis • The H+ gradient is referred to as a proton- motive force, emphasizing its capacity to do work
  • 47. An Accounting of ATP Production by Cellular Respiration • During cellular respiration, most energy flows in this sequence: glucose → NADH → electron transport chain → proton-motive force → ATP • About 34% of the energy in a glucose molecule is transferred to ATP during cellular respiration, making about 32 ATP
  • 48. Figure 9.16 Electron shuttles MITOCHONDRION span membrane 2 NADH or 2 FADH2 2 NADH 2 NADH 6 NADH 2 FADH2 Glycolysis Pyruvate oxidation Oxidative Citric phosphorylation: Glucose 2 Pyruvate 2 Acetyl CoA acid electron transport cycle and chemiosmosis + 2 ATP + 2 ATP + about 26 or 28 ATP About Maximum per glucose: 30 or 32 ATP CYTOSOL
  • 49. Fermentation and anaerobic respiration enable cells to produce ATP without the use of oxygen • Most cellular respiration requires O2 to produce ATP • Without O2, the electron transport chain will cease to operate • In that case, glycolysis couples with fermentation or anaerobic respiration to produce ATP
  • 50. • Anaerobic respiration uses an electron transport chain with a final electron acceptor other than O2, for example sulfate • Fermentation uses substrate-level phosphorylation instead of an electron transport chain to generate ATP
  • 51. 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
  • 52. • In alcohol fermentation, pyruvate is converted to ethanol in two steps, with the first releasing CO2 • Alcohol fermentation by yeast is used in brewing, winemaking, and baking
  • 53. Figure 9.17a 2 ADP + 2 P i 2 ATP Glucose Glycolysis 2 Pyruvate 2 NAD + 2 NADH 2 CO2 + 2 H+ 2 Ethanol 2 Acetaldehyde (a) Alcohol fermentation
  • 54. • In lactic acid fermentation, pyruvate is reduced to NADH, forming lactate as an end product, with no release of CO2 • Lactic acid fermentation by some fungi and bacteria is used to make cheese and yogurt • Human muscle cells use lactic acid fermentation to generate ATP when O2 is scarce
  • 55. Figure 9.17b 2 ADP + 2 P i 2 ATP Glucose Glycolysis 2 NAD + 2 NADH + 2 H+ 2 Pyruvate 2 Lactate (b) Lactic acid fermentation
  • 56. Comparing Fermentation with Anaerobic and Aerobic Respiration • All use glycolysis (net ATP = 2) to oxidize glucose and harvest chemical energy of food • In all three, NAD+ is the oxidizing agent that accepts electrons during glycolysis • 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 32 ATP per glucose molecule; fermentation produces 2 ATP per glucose molecule
  • 57. • 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
  • 58. Figure 9.18 Glucose Glycolysis CYTOSOL Pyruvate No O2 present: O2 present: Fermentation Aerobic cellular respiration MITOCHONDRION Ethanol, Acetyl CoA lactate, or other products Citric acid cycle
  • 59. The Evolutionary Significance of Glycolysis • Ancient prokaryotes are thought to have used glycolysis long before there was oxygen in the atmosphere • Very little O2 was available in the atmosphere until about 2.7 billion years ago, so early prokaryotes likely used only glycolysis to generate ATP • Glycolysis is a very ancient process
  • 60. Glycolysis and the citric acid cycle connect to many other metabolic pathways • Gycolysis and the citric acid cycle are major intersections to various catabolic and anabolic pathways
  • 61. The Versatility of Catabolism • Catabolic pathways funnel electrons from many kinds of organic molecules into cellular respiration • Glycolysis accepts a wide range of carbohydrates • Proteins must be digested to amino acids; amino groups can feed glycolysis or the citric acid cycle
  • 62. • Fats are digested to glycerol (used in glycolysis) and fatty acids (used in generating acetyl CoA) • Fatty acids are broken down by beta oxidation and yield acetyl CoA • An oxidized gram of fat produces more than twice as much ATP as an oxidized gram of carbohydrate
  • 63. Figure 9.19 Proteins Carbohydrates Fats Amino Sugars Glycerol Fatty acids acids Glycolysis Glucose Glyceraldehyde 3- P NH3 Pyruvate Acetyl CoA Citric acid cycle Oxidative phosphorylation
  • 64. Biosynthesis (Anabolic Pathways) • The body uses small molecules to build other substances • These small molecules may come directly from food, from glycolysis, or from the citric acid cycle
  • 65. Regulation of Cellular Respiration via Feedback Mechanisms • Feedback inhibition is the most common mechanism for control • If ATP concentration begins to drop, respiration speeds up; when there is plenty of ATP, respiration slows down • Control of catabolism is based mainly on regulating the activity of enzymes at strategic points in the catabolic pathway

Notes de l'éditeur

  1. Figure 8.8 The structure and hydrolysis of adenosine triphosphate (ATP).
  2. Figure 8.8 The structure and hydrolysis of adenosine triphosphate (ATP).
  3. Figure 8.10 How ATP drives transport and mechanical work.
  4. Figure 8.11 The ATP cycle.
  5. For the Discovery Video Space Plants, go to Animation and Video Files.
  6. Figure 9.2 Energy flow and chemical recycling in ecosystems.
  7. Figure 9.UN01 In-text figure, p. 164
  8. Figure 9.UN03 In-text figure, p. 165
  9. Figure 9.4 NAD  as an electron shuttle.
  10. Figure 9.UN05 In-text figure, p. 167
  11. Figure 9.6 An overview of cellular respiration.
  12. Figure 9.6 An overview of cellular respiration.
  13. Figure 9.6 An overview of cellular respiration.
  14. Figure 9.8 The energy input and output of glycolysis.
  15. Figure 9.10 Oxidation of pyruvate to acetyl CoA, the step before the citric acid cycle.
  16. Figure 9.11 An overview of pyruvate oxidation and the citric acid cycle.
  17. Figure 9.13 Free-energy change during electron transport.
  18. Figure 9.14 ATP synthase, a molecular mill.
  19. Figure 9.15 Chemiosmosis couples the electron transport chain to ATP synthesis.
  20. Figure 9.16 ATP yield per molecule of glucose at each stage of cellular respiration.
  21. Figure 9.17 Fermentation.
  22. Figure 9.17 Fermentation.
  23. Figure 9.18 Pyruvate as a key juncture in catabolism.
  24. Figure 9.19 The catabolism of various molecules from food.