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Chapter 9Chapter 9
Cellular Respiration
Fig. 9-2
Lightenergy
ECOSYSTEM
Photosynthesis in chloroplasts
CO2 + H2O
Cellular respirationin mitochondria
Organicmolecules+ O2
ATP powers most cellular work
Heatenergy
ATP
I. Catabolic Pathways and Production of ATP
• Fermentation = partial degradation of sugars that occurs without O2
• Aerobic respiration uses organic molecules and O2 and yields ATP
• Anaerobic respiration = similar to aerobic but consumes compounds other than O2
• Cellular respiration includes aerobic and anaerobic
• C6H12O6 + 6 O2 6 CO2 + 6 H2O + Energy (ATP + heat)
II. Redox Reactions: Oxidation and Reduction
• Transfer of electrons, releases energy stored in organic molecules, used to synthesize ATP
A. Principle of Redox
• Chemical reactions that transfer electrons between reactants are called oxidation-reduction reactions, or redox reactions
• Oxidation = substance loses electrons, oxidized
• Reduction = substance gains electrons, reduced (the amount of + charge is reduced)
• Electron donor = reducing agent
• Electron receptor = oxidizing agent
Fig. 9-UN1
becomes oxidized(loses electron)
becomes reduced(gains electron)
Fig. 9-UN2
becomes oxidized
becomes reduced
Fig. 9-3
Reactants
becomes oxidized
becomes reduced
Products
Methane(reducing
agent)
Oxygen(oxidizing
agent)
Carbon dioxide Water
Fig. 9-UN3
becomes oxidized
becomes reduced
B. NAD+ and the Electron Transport Chain
• Glucose is broken down in a series of steps
• Electrons are first transferred to NAD+, a coenzyme
• As an electron acceptor, NAD+ functions as an oxidizing agent
• Each NADH (the reduced form of NAD+) represents stored energy that is tapped to make ATP
• How NAD works
Fig. 9-4
Dehydrogenase
Reduction of NAD+
Oxidation of NADH
2 e– + 2 H+
2 e– + H+
NAD+ + 2[H]
NADH
+
H+
H+
Nicotinamide(oxidized form)
Nicotinamide(reduced form)
• NADH passes the electrons to the electron transport chain
• ETC 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 regenerates ATP
Fig. 9-5
Fre
e en
erg
y, G
Fre
e en
erg
y, G
(a) Uncontrolled reaction
H2O
H2 + 1/2 O2
Explosiverelease of
heat and lightenergy
(b) Cellular respiration
Controlledrelease ofenergy for
synthesis ofATP
2 H+ + 2 e–
2 H + 1/2 O2
(from food via NADH)
ATP
ATP
ATP
1/2 O22 H+
2 e–E
lectron
transp
ort
chain
H2O
III. Stages of Cellular Respiration: A Preview
• Cellular respiration has three stages:
– Glycolysis (breaks down glucose into two molecules of pyruvate)
– Citric acid cycle (completes the breakdown of glucose)
– Oxidative phosphorylation (accounts for most of the ATP synthesis)
Fig. 9-6-1
Substrate-levelphosphorylation
ATP
Cytosol
Glucose Pyruvate
Glycolysis
Electronscarried
via NADH
Fig. 9-6-2
Mitochondrion
Substrate-levelphosphorylation
ATP
Cytosol
Glucose Pyruvate
Glycolysis
Electronscarried
via NADH
Substrate-levelphosphorylation
ATP
Electrons carriedvia NADH and
FADH2
Citricacidcycle
Fig. 9-6-3
Mitochondrion
Substrate-levelphosphorylation
ATP
Cytosol
Glucose Pyruvate
Glycolysis
Electronscarried
via NADH
Substrate-levelphosphorylation
ATP
Electrons carriedvia NADH and
FADH2
Oxidativephosphorylation
ATP
Citricacidcycle
Oxidativephosphorylation:electron transport
andchemiosmosis
• Oxidative phosphorylation (powered by redox reactions) generates most of the ATP (about 90% of total from cellular resp)
• Other 10% of ATP is formed in glycolysis and the citric acid cycle by substrate-level phosphorylation
Fig. 9-7
Enzyme
ADP
PSubstrate
Enzyme
ATP+
Product
Concept 9.2: Glycolysis harvests chemical energy by oxidizing glucose to pyruvate
• Glycolysis (“splitting of sugar”) breaks down glucose into two molecules of pyruvate, 2 ATP and 2 NADH (high energy e- carriers)
• Glycolysis occurs in the cytoplasm and has two major phases:
– Energy investment phase
– Energy payoff phase
• How glycolysis works
Fig. 9-8
Energy investment phase
Glucose
2 ADP + 2 P 2 ATP used
formed4 ATP
Energy payoff phase
4 ADP + 4 P
2 NAD+ + 4 e– + 4 H+ 2 NADH + 2 H+
2 Pyruvate + 2 H2O
2 Pyruvate + 2 H2OGlucoseNet
4 ATP formed – 2 ATP used 2 ATP
2 NAD+ + 4 e– + 4 H+ 2 NADH + 2 H+
Fig. 9-9-1
ATP
ADP
Hexokinase1
ATP
ADP
Hexokinase1
Glucose
Glucose-6-phosphate
Glucose
Glucose-6-phosphate
Fig. 9-9-2
Hexokinase
ATP
ADP
1
Phosphoglucoisomerase2
Phosphogluco-isomerase
2
Glucose
Glucose-6-phosphate
Fructose-6-phosphate
Glucose-6-phosphate
Fructose-6-phosphate
1
Fig. 9-9-3
Hexokinase
ATP
ADP
Phosphoglucoisomerase
Phosphofructokinase
ATP
ADP
2
3
ATP
ADP
Phosphofructo-kinase
Fructose-1, 6-bisphosphate
Glucose
Glucose-6-phosphate
Fructose-6-phosphate
Fructose-1, 6-bisphosphate
1
2
3
Fructose-6-phosphate
3
Fig. 9-9-4
Glucose
ATP
ADP
Hexokinase
Glucose-6-phosphate
Phosphoglucoisomerase
Fructose-6-phosphate
ATP
ADP
Phosphofructokinase
Fructose-1, 6-bisphosphate
Aldolase
Isomerase
Dihydroxyacetonephosphate
Glyceraldehyde-3-phosphate
1
2
3
4
5
Aldolase
Isomerase
Fructose-1, 6-bisphosphate
Dihydroxyacetonephosphate
Glyceraldehyde-3-phosphate
4
5
Fig. 9-9-52 NAD+
NADH2
+ 2 H+
2
2 P i
Triose phosphatedehydrogenase
1, 3-Bisphosphoglycerate
6
2 NAD+
Glyceraldehyde-3-phosphate
Triose phosphatedehydrogenase
NADH2
+ 2 H+
2 P i
1, 3-Bisphosphoglycerate
6
2
2
Fig. 9-9-62 NAD+
NADH2
Triose phosphatedehydrogenase
+ 2 H+
2 P i
2
2 ADP
1, 3-Bisphosphoglycerate
Phosphoglycerokinase2 ATP
2 3-Phosphoglycerate
6
7
2
2 ADP
2 ATP
1, 3-Bisphosphoglycerate
3-Phosphoglycerate
Phosphoglycero-kinase
2
7
Fig. 9-9-7
3-Phosphoglycerate
Triose phosphatedehydrogenase
2 NAD+
2 NADH+ 2 H+
2 P i
2
2 ADP
Phosphoglycerokinase
1, 3-Bisphosphoglycerate
2 ATP
3-Phosphoglycerate2
Phosphoglyceromutase
2-Phosphoglycerate2
2-Phosphoglycerate2
2
Phosphoglycero-mutase
6
7
8
8
Fig. 9-9-82 NAD+
NADH2
2
2
2
2
+ 2 H+
Triose phosphatedehydrogenase2 P i
1, 3-Bisphosphoglycerate
Phosphoglycerokinase
2 ADP
2 ATP
3-Phosphoglycerate
Phosphoglyceromutase
Enolase
2-Phosphoglycerate
2 H2O
Phosphoenolpyruvate
9
8
7
6
2 2-Phosphoglycerate
Enolase
2
2 H2O
Phosphoenolpyruvate
9
Fig. 9-9-9
Triose phosphatedehydrogenase
2 NAD+
NADH2
2
2
2
2
2
2 ADP
2 ATP
Pyruvate
Pyruvate kinase
Phosphoenolpyruvate
Enolase2 H2O
2-Phosphoglycerate
Phosphoglyceromutase
3-Phosphoglycerate
Phosphoglycerokinase
2 ATP
2 ADP
1, 3-Bisphosphoglycerate
+ 2 H+
6
7
8
9
10
2
2 ADP
2 ATP
Phosphoenolpyruvate
Pyruvate kinase
2 Pyruvate
10
2 P i
IV. Citric acid cycle completes oxidation
• With O2, pyruvate enters mitochondrion
• Pyruvate must be converted to acetyl CoA, which links the cycle to glycolysis
Fig. 9-10
CYTOSOL MITOCHONDRION
NAD+ NADH + H+
2
1 3
Pyruvate
Transport protein
CO2Coenzyme A
Acetyl CoA
• Citric acid cycle (Krebs cycle) takes place within mito matrix
• Cycle oxidizes organic fuel derived from pyruvate, generating 1 ATP, 3 NADH, and 1 FADH2 per turn (and 2 CO2, cutting up “sugar” even more)
Fig. 9-11
Pyruvate
NAD+
NADH
+ H+Acetyl CoA
CO2
CoA
CoA
CoA
Citricacidcycle
FADH2
FAD
CO22
3
3 NAD+
+ 3 H+
ADP + P i
ATP
NADH
Fig. 9-12-8
Acetyl CoA
CoA—SH
Citrate
H2O
IsocitrateNAD+
NADH
+ H+
CO2
-Keto-glutarate
CoA—SH
CO2NAD+
NADH
+ H+SuccinylCoA
CoA—SH
P i
GTP GDP
ADP
ATP
Succinate
FAD
FADH2
Fumarate
CitricacidcycleH2O
Malate
Oxaloacetate
NADH
+H+
NAD+
1
2
3
4
5
6
7
8
• Overview Glycolysis - Citric Acid Cycle
V. Electron Transport Chain
• NADH and FADH2 donate e- s to the electron transport chain, which powers ATP synthesis via oxidative phosphorylation
• In the cristae of mitochondrion
• Carriers alternate reduced and oxidized states as they accept and donate e-
• e- s drop in free energy as they go down the chain and are passed to O2, forming H2O
Fig. 9-13
NADH
NAD+2FADH2
2 FADMultiproteincomplexesFAD
Fe•S
FMN
Fe•S
Q
Fe•S
Cyt b
Cyt c1
Cyt c
Cyt a
Cyt a3
IV
Fre
e en
erg
y (G
) r e
lat i
ve t
o O
2 (
kcal
/mo
l)
50
40
30
20
10 2
(from NADHor FADH2)
0 2 H+ + 1/2 O2
H2O
e–
e–
e–
• e- s are transferred from NADH or FADH2 to the electron transport chain
• Electron transport chain generates no ATP
• Function is to break the large free-energy drop from food to O2 into smaller steps that release energy in manageable amounts
• Electron Transport Chain and Oxi Phos (3min)
VI. Chemiosmosis: Energy-Coupling
• e- transfer in ETC causes proteins to pump H+ from mito matrix to intermembrane space
• H+ then moves back across membrane, passing through channels in ATP synthase
• ATP synthase uses the exergonic (energy releasing) flow of H+ to drive phosphorylation of ATP (ADP to ATP)
• Chemiosmosis = use of energy in a H+ gradient to drive cellular work
• H+ gradient is a proton-motive force, emphasizing its capacity to do work
Fig. 9-14
INTERMEMBRANE SPACE
Rotor
H+
Stator
Internalrod
Cata-lyticknob
ADP+P ATP
i
MITOCHONDRIAL MATRIX
Fig. 9-16
Protein complexof electroncarriers
H+
H+H+
Cyt c
Q
V
FADH2 FAD
NAD+NADH
(carrying electronsfrom food)
Electron transport chain
2 H+ + 1/2O2H2O
ADP + P i
Chemiosmosis
Oxidative phosphorylation
H+
H+
ATP synthase
ATP
21
• ETC / Oxi Phos Overview
VII. ATP Production by Cellular Respiration
• During cellular respiration, most energy flows in this sequence:
glucose NADH electron transport chain proton-motive force ATP
• About 40% of energy in a glucose molecule is transferred to ATP during cellular respiration (about 36 / 38 ATP)
Fig. 9-17
Maximum per glucose: About36 or 38 ATP
+ 2 ATP+ 2 ATP + about 32 or 34 ATP
Oxidativephosphorylation:electron transport
andchemiosmosis
Citricacidcycle
2AcetylCoA
Glycolysis
Glucose2
Pyruvate
2 NADH 2 NADH 6 NADH 2 FADH2
2 FADH2
2 NADHCYTOSOL Electron shuttles
span membrane
or
MITOCHONDRION
• Cell Respiration Part 2 (9min)
• Cellular Respiration Overview - Bozeman (14min)
VIII. Fermentation and anaerobic respiration
• W/o 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, ex. sulfate
• Fermentation uses phosphorylation instead of an electron transport chain to generate ATP
IX. Types of Fermentation
• Fermentation = glycolysis plus reactions that regenerate NAD+ (can be reused by glycolysis)
• Alcohol fermentation = pyruvate is converted to ethanol in two steps, w/ the first releasing CO2
• Yeast = used in brewing, winemaking, and baking
Fig. 9-18a
2 ADP + 2 P i 2 ATP
Glucose Glycolysis
2 Pyruvate
2 NADH2 NAD+
+ 2 H+CO2
2 Acetaldehyde2 Ethanol
(a) Alcohol fermentation
2
• Lactic acid fermentation = pyruvate is reduced to NADH, forming lactate, w/ no release of CO2
• Some fungi and bacteria = cheese and yogurt
• Human muscle cells = generate ATP when O2 is scarce
Fig. 9-18b
Glucose
2 ADP + 2 P i 2 ATP
Glycolysis
2 NAD+ 2 NADH+ 2 H+
2 Pyruvate
2 Lactate
(b) Lactic acid fermentation
X. Fermentation and Aerobic Respiration Compared
• Both use glycolysis to oxidize glucose to pyruvate
• 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
• Obligate anaerobes = carry out fermentation or anaerobic respiration and cannot survive in the presence of O2
• Facultative anaerobes (Yeast and many bacteria) = can survive using either fermentation or cellular respiration
– Pyruvate is a fork in the metabolic road that leads to two alternative catabolic routes
Fig. 9-19Glucose
Glycolysis
Pyruvate
CYTOSOL
No O2 present:Fermentation
O2 present:
Aerobic cellular respiration
MITOCHONDRION
Acetyl CoAEthanolor
lactateCitricacidcycle
XI. Evolutionary Significance of Glycolysis
• Glycolysis occurs in nearly all organisms
• Glycolysis probably evolved in ancient prokaryotes before there was O2 in atmosphere
XII. Versatility of Catabolism
• Catabolic pathways funnel electrons from many kinds of organic molecules into cellular respiration
• Glycolysis accepts a wide range of carbohydrates
• Proteins = digested to amino acids; amino groups can feed glycolysis / citric acid cycle
• Fats = digested to glycerol (used in glycolysis) and fatty acids (used in generating acetyl CoA, by beta oxidation)
• An oxidized gram of fat produces more than 2xs as much ATP as an oxidized gram of carbohydrate
Fig. 9-20Proteins
Carbohydrates
Aminoacids
Sugars
Fats
Glycerol Fattyacids
Glycolysis
Glucose
Glyceraldehyde-3-
Pyruvate
P
NH3
Acetyl CoA
Citricacidcycle
Oxidativephosphorylation
XIII. Regulation by Feedback Mechanisms
• Feedback inhibition is most common mechanism for control
• [ATP] begins to drop, resp speeds up; when ATP increases, resp slows down
• Control of catabolism is based on regulating the activity of enzymes at points in pathway
Fig. 9-21Glucose
GlycolysisFructose-6-phosphate
Phosphofructokinase
Fructose-1,6-bisphosphateInhibits
AMP
Stimulates
Inhibits
Pyruvate
CitrateAcetyl CoA
Citricacidcycle
Oxidativephosphorylation
ATP
+
––