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BIOLOGYA Global Approach
Campbell • Reece • Urry • Cain • Wasserman • Minorsky • Jackson
© 2015 Pearson Education Ltd
TENTH EDITION
Global Edition
Lecture Presentation by Nicole Tunbridge andKathleen Fitzpatrick
10
Cell Respiration
© 2015 Pearson Education Ltd
Life Is Work
a) Living cells require energy from outside sources
b) Some animals, such as the giraffe, obtain energy by eating plants, and some animals feed on other organisms that eat plants
© 2015 Pearson Education Ltd
Figure 10.1
© 2015 Pearson Education Ltd
a) Energy flows into an ecosystem as sunlight and leaves as heat
b) Photosynthesis generates O2 and organic molecules, which are used in cellular respiration
c) Cells use chemical energy stored in organic molecules to generate ATP, which powers work
© 2015 Pearson Education Ltd
Figure 10.2
Lightenergy
Organicmolecules O2CO2 H2O+
Photosynthesisin chloroplasts
Cellular respirationin mitochondria
ECOSYSTEM
ATP powersmost cellular workATP
Heatenergy
+
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BioFlix: The Carbon Cycle
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Concept 10.1: Catabolic pathways yield energy by oxidizing organic fuels
a) Catabolic pathways release stored energy by breaking down complex molecules
b) Electron transfer plays a major role in these pathways
c) These processes are central to cellular respiration
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Catabolic Pathways and Production of ATP
a) The breakdown of organic molecules is exergonic
b)Fermentation is a partial degradation of sugars that occurs without O2
c) Aerobic respiration consumes organic molecules and O2 and yields ATP
d) Anaerobic respiration is similar to aerobic respiration but consumes compounds otherthan O2
© 2015 Pearson Education Ltd
a) Cellular respiration includes both aerobic and anaerobic respiration but is often used to refer to aerobic respiration
b) 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)
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Redox Reactions: Oxidation and Reduction
a) The transfer of electrons during chemical reactions releases energy stored in organic molecules
b) This released energy is ultimately used to synthesize ATP
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The Principle of Redox
a) Chemical reactions that transfer electrons between reactants are called oxidation-reduction reactions, or redox reactions
b) In oxidation, a substance loses electrons,or is oxidized
c) In reduction, a substance gains electrons, or is reduced (the amount of positive charge is reduced)
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Stepwise Energy Harvest via NAD+ and the Electron Transport Chain
a) In cellular respiration, glucose and other organic molecules are broken down in a series of steps
b) Electrons from organic compounds are usually first transferred to NAD+, a coenzyme
c) As an electron acceptor, NAD+ functions as an oxidizing agent during cellular respiration
d) Each NADH (the reduced form of NAD+) represents stored energy that is tapped to synthesize ATP
© 2015 Pearson Education Ltd
Figure 10.4
NAD+
2 e− + 2 H+
2[H](from food)
Nicotinamide(oxidized form)
Reduction of NAD+
2 e− + H+
NADH
Nicotinamide(reduced form)
Oxidation of NADHH+
H+
Dehydrogenase
© 2015 Pearson Education Ltd
Figure 10.UN04
Dehydrogenase
© 2015 Pearson Education Ltd
a) NADH passes the electrons to the electron transport chain
b) Unlike an uncontrolled reaction, the electron transport chain passes electrons in a series of steps instead of one explosive reaction
c) O2 pulls electrons down the chain in an energy-yielding tumble
d) The energy yielded is used to regenerate ATP
© 2015 Pearson Education Ltd
Figure 10.5
H2 +½ O2 2 H + ½ O2
2 H+
2 e−
2 e−2 H+ +
H2O
½ O2
Controlledrelease of
energy
Electron transport
chain
ATP
ATP
ATPExplosive
release
Cellular respirationUncontrolled reaction(a) (b)
Free
ene
rgy,
G
Free
ene
rgy,
G
H2O
© 2015 Pearson Education Ltd
The Stages of Cellular Respiration: A Preview
a) Harvesting of energy from glucose has three stages
a)Glycolysis (breaks down glucose into two molecules of pyruvate)
b)The citric acid cycle (completes the breakdown of glucose)
c)Oxidative phosphorylation (accounts for most of the ATP synthesis)
© 2015 Pearson Education Ltd
Figure 10.UN05
1.2.
3.
GLYCOLYSIS (color-coded blue throughout the chapter)PYRUVATE OXIDATION and the CITRIC ACID CYCLE(color-coded orange)
OXIDATIVE PHOSPHORYLATION: Electron transport andchemiosmosis (color-coded purple)
© 2015 Pearson Education Ltd
Figure 10.6-1
Electronsvia NADH
ATP
CYTOSOL MITOCHONDRION
Substrate-level
GLYCOLYSIS
Glucose Pyruvate
© 2015 Pearson Education Ltd
Figure 10.6-2
Electronsvia NADH
Electronsvia NADHand FADH2
ATP ATP
CYTOSOL MITOCHONDRION
Substrate-level Substrate-level
GLYCOLYSIS PYRUVATEOXIDATION CITRIC
ACIDCYCLEAcetyl CoAGlucose Pyruvate
© 2015 Pearson Education Ltd
Figure 10.6-3
Electronsvia NADH
Electronsvia NADHand FADH2
ATP ATP ATP
CYTOSOL MITOCHONDRION
Substrate-level Substrate-level Oxidative
GLYCOLYSIS PYRUVATEOXIDATION CITRIC
ACIDCYCLE
OXIDATIVEPHOSPHORYLATION
(Electron transportand chemiosmosis)
Acetyl CoAGlucose Pyruvate
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BioFlix: Cellular Respiration
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a) The process that generates most of the ATP is called oxidative phosphorylation because it is powered by redox reactions
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a) Oxidative phosphorylation accounts for almost 90% of the ATP generated by cellular respiration
b) A smaller amount of ATP is formed in glycolysis and the citric acid cycle by substrate-level phosphorylation
c) For each molecule of glucose degraded to CO2 and water by respiration, the cell makes up to 32 molecules of ATP
© 2015 Pearson Education Ltd
Figure 10.7
Enzyme Enzyme
Substrate
Product
ATP
ADP
P
© 2015 Pearson Education Ltd
Concept 10.2: Glycolysis harvests chemical energy by oxidizing glucose to pyruvate
a) Glycolysis (“sugar splitting”) breaks down glucose into two molecules of pyruvate
b) Glycolysis occurs in the cytoplasm and has two major phases
a)Energy investment phase
b)Energy payoff phase
c) Glycolysis occurs whether or not O2 is present
© 2015 Pearson Education Ltd
Figure 10.UN06
GLYCOLYSIS PYRUVATEOXIDATION
CITRICACID
CYCLE
OXIDATIVEPHOSPHORYL-
ATION
ATP
© 2015 Pearson Education Ltd
Figure 10.8
Energy Investment Phase
Glucose
Energy Payoff Phase
Net
2 ATP used 2 ADP + 2 P
4 ADP + 4 P 4 ATP formed
NAD+ 4 e−2 + + 4 H+ 2 H+2 NADH
Pyruvate2
2
2
2
2
2
2
2 H+H+
4
4
Glucose Pyruvate
ATPATP usedATP formedH2O
NADH
−
2 NAD+
+
+
+
++ +
2 H2O
4e−
© 2015 Pearson Education Ltd
Concept 10.3: After pyruvate is oxidized, the citric acid cycle completes the energy-yielding oxidation of organic molecules
a) In the presence of O2, pyruvate enters the mitochondrion (in eukaryotic cells) where the oxidation of glucose is completed
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Oxidation of Pyruvate to Acetyl CoA
a) 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
b) This step is carried out by a multienzyme complex that catalyses three reactions
© 2015 Pearson Education Ltd
Figure 10.UN07
GLYCOLYSIS PYRUVATEOXIDATION
CITRICACID
CYCLE
OXIDATIVEPHOSPHORYL-
ATION
© 2015 Pearson Education Ltd
Figure 10.10
CYTOSOL
Pyruvate
Transport protein
MITOCHONDRION
Acetyl CoANAD+ H+NADH +
CO2Coenzyme A
1
2
3
© 2015 Pearson Education Ltd
The Citric Acid Cycle
a) The citric acid cycle, also called the Krebs cycle, completes the break down of pyruvate to CO2
b) The cycle oxidizes organic fuel derived from pyruvate, generating 1 ATP, 3 NADH, and 1 FADH2 per turn
© 2015 Pearson Education Ltd
Figure 10.11
PYRUVATE OXIDATIONPyruvate (from glycolysis,2 molecules per glucose)
NADH
NAD+CO2
CoA
CoA+ H+
CoA
CoA
CO2
CITRICACID
CYCLE
FADH2
FAD
ATP
ADP + P i
NAD+
+ 3 H+
NADH3
3
2
Acetyl CoA
© 2015 Pearson Education Ltd
Figure 10.11a
PYRUVATE OXIDATIONPyruvate (from glycolysis,2 molecules per glucose)
NADH
NAD+CO2
CoA
CoA+ H+ Acetyl CoA
© 2015 Pearson Education Ltd
Figure 10.11b
CoA
CoA
CO2
CITRICACID
CYCLEFADH2
FAD
ATPADP + P i
NAD+
+ 3 H+
NADH3
3
2
Acetyl CoA
© 2015 Pearson Education Ltd
a) The citric acid cycle has eight steps, each catalyzed by a specific enzyme
b) The acetyl group of acetyl CoA joins the cycle by combining with oxaloacetate, forming citrate
c) The next seven steps decompose the citrate back to oxaloacetate, making the process a cycle
d) The NADH and FADH2 produced by the cycle relay electrons extracted from food to the electron transport chain
© 2015 Pearson Education Ltd
Figure 10.UN08
GLYCOLYSIS PYRUVATEOXIDATION
CITRICACID
CYCLE
OXIDATIVEPHOSPHORYL-
ATION
ATP
© 2015 Pearson Education Ltd
Figure 10.12-1
Acetyl CoA
Oxaloacetate
CoA-SH
Citrate
CITRICACID
CYCLE
1
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Figure 10.12-2
Acetyl CoA
H2O
Oxaloacetate
CoA-SH
Citrate
CITRICACID
CYCLE
Isocitrate
1
2
© 2015 Pearson Education Ltd
Figure 10.12-3
Acetyl CoA
H2O
+ H+
Oxaloacetate
CoA-SH
Citrate
-Ketoglutarate
NAD+
CO2
NADHCITRICACID
CYCLE
Isocitrate
1
2
3
© 2015 Pearson Education Ltd
Figure 10.12-4
Acetyl CoA
H2O
+ H+
Oxaloacetate
CoA-SH
Citrate
-KetoglutarateCoA-SH
NAD+
CO2
NADH
CO2NAD+
NADH+ H+
SuccinylCoA
CITRICACID
CYCLE
Isocitrate
1
2
4
3
© 2015 Pearson Education Ltd
Figure 10.12-5
Acetyl CoA
H2O
+ H+
Oxaloacetate
CoA-SH
Citrate
-KetoglutarateCoA-SH
NAD+
CO2
NADH
CO2NAD+
NADH+ H+
SuccinylCoA
ATP
ADP
GTP GDPSuccinate
CITRICACID
CYCLE
Pi
Isocitrate
1
2
5
4
3
CoA-SH
© 2015 Pearson Education Ltd
Figure 10.12-6
Acetyl CoA
H2O
+ H+
Oxaloacetate
CoA-SH
Citrate
-KetoglutarateCoA-SH
NAD+
CO2
NADH
CO2NAD+
NADH+ H+
SuccinylCoA
ATP
ADP
GTP GDPSuccinateFAD
FADH2
Fumarate
CITRICACID
CYCLE
Pi
Isocitrate
1
2
5
6 4
3
CoA-SH
© 2015 Pearson Education Ltd
Figure 10.12-7
Acetyl CoA
H2O
+ H+
Oxaloacetate
CoA-SH
Citrate
-KetoglutarateCoA-SH
NAD+
CO2
NADH
CO2NAD+
NADH+ H+
SuccinylCoA
ATP
ADP
GTP GDPSuccinateFAD
FADH2
Fumarate
H2O
Malate
CITRICACID
CYCLE
Pi
Isocitrate
1
2
5
6 4
37
CoA-SH
© 2015 Pearson Education Ltd
Figure 10.12-8
Acetyl CoA
H2O
+ H+
Oxaloacetate
CoA-SH
Citrate
-KetoglutarateCoA-SH
NAD+
CO2
NADH
CO2NAD+
NADH+ H+
SuccinylCoA
ATP
ADP
GTP GDPSuccinateFAD
FADH2
Fumarate
H2O
Malate
CITRICACID
CYCLE
+ H+
NAD+
Pi
NADH
Isocitrate
1
2
5
6 4
37
8
CoA-SH
© 2015 Pearson Education Ltd
Concept 10.4: During oxidative phosphorylation, chemiosmosis couples electron transport to ATP synthesis
a) Following glycolysis and the citric acid cycle, NADH and FADH2 account for most of the energy extracted from food
b) These two electron carriers donate electrons to the electron transport chain, which powers ATP synthesis via oxidative phosphorylation
© 2015 Pearson Education Ltd
The Pathway of Electron Transport
a) The electron transport chain is in the inner membrane (cristae) of the mitochondrion
b) Most of the chain’s components are proteins,which exist in multiprotein complexes
c) The carriers alternate reduced and oxidized states as they accept and donate electrons
d) Electrons drop in free energy as they go down the chain and are finally passed to O2, forming H2O
© 2015 Pearson Education Ltd
Figure 10.UN09
GLYCOLYSISCITRICACID
CYCLE
PYRUVATEOXIDATION
ATP
OXIDATIVEPHOSPHORYL-
ATION
© 2015 Pearson Education Ltd
Figure 10.13
NADH
e−2 NAD+
FADH2
FAD2 e−
FMNFe•S
Q
Fe•S
Cyt bFe•S
Cyt c1
Cyt cCyt a
Cyt a3
e−2(originally from
NADH or FADH2)
2 H+ + ½ O2
H2O
MultiproteincomplexesI
II
III
IV
50
40
30
20
10
0
Free
ene
rgy
(G) r
elat
ive
to O
2 (kc
al/m
ol)
© 2015 Pearson Education Ltd
Figure 10.13a
50
40
30
20
10
Free
ene
rgy
(G) r
elat
ive
to O
2 (kc
al/m
ol)
NADH
e−2 NAD+
FADH2
Fe•S
e−
FMNFe•S
QCyt b
FADIII
FAD Multiproteincomplexes
III
Fe•SCyt c1
Cyt cCyt a
Cyt a3
IV
2
e−2
© 2015 Pearson Education Ltd
Figure 10.13b
30
20
10
Free
ene
rgy
(G) r
elat
ive
to O
2 (kc
al/m
ol)
0
Cyt c1
Cyt cCyt a
Cyt a3
IV
e−2
2 H+ + ½ O2
H2O
(originally fromNADH or FADH2)
© 2015 Pearson Education Ltd
a) Electrons are transferred from NADH or FADH2 to the electron transport chain
b) Electrons are passed through a number of proteins including cytochromes (each with an iron atom) to O2
c) The electron transport chain generates no ATP directly
d) It breaks the large free-energy drop from food to O2 into smaller steps that release energy in manageable amounts
© 2015 Pearson Education Ltd
Chemiosmosis: The Energy-Coupling Mechanism
a) Electron transfer in the electron transport chain causes proteins to pump H+ from the mitochondrial matrix to the intermembrane space
b) H+ then moves back across the membrane, passing through the protein complex, ATP synthase
c) ATP synthase uses the exergonic flow of H+ to drive phosphorylation of ATP
d) This is an example of chemiosmosis, the use of energy in a H+ gradient to drive cellular work
© 2015 Pearson Education Ltd
Figure 10.14
INTERMEMBRANESPACE
RotorH+ Stator
Internal rod
Catalyticknob
ADP+P i
MITOCHONDRIALMATRIX
ATP
© 2015 Pearson Education Ltd
Video: ATP Synthase 3-D Structure, Top View
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Video: ATP Synthase 3-D Structure, Side View
© 2015 Pearson Education Ltd
Figure 10.15
Proteincomplexof electroncarriers
(carryingelectronsfromfood)
ATPsynthase
Electron transport chain Chemiosmosis
Oxidative phosphorylation
2 H+ + ½ O2 H2O
ADP + P i
H+
ATP
H+
H+
H+
H+
Cyt c
QI
II
III
IV
FADFADH2
NADH NAD+
1 2
© 2015 Pearson Education Ltd
Figure 10.15a
Proteincomplexof electroncarriers
2 H+ + ½ O2 H2O
H+
NAD+
1
H+
H+
NADH
FADH2
Cyt cCyt c
IV
III
II
I
FAD
(carryingelectronsfromfood) Electron transport chain
Q
© 2015 Pearson Education Ltd
Figure 10.15b
H+
2
H+
ADP + P i ATP
Chemiosmosis
ATPsynthase
© 2015 Pearson Education Ltd
a) The energy stored in a H+ gradient across a membrane couples the redox reactions of the electron transport chain to ATP synthesis
b) The H+ gradient is referred to as a proton-motive force, emphasizing its capacity to do work
© 2015 Pearson Education Ltd
An Accounting of ATP Production by Cellular Respiration
a) During cellular respiration, most energy flows in this sequence:
glucose → NADH → electron transport chain → proton-motive force → ATP
b) About 34% of the energy in a glucose molecule is transferred to ATP during cellular respiration, making about 32 ATP
c) There are several reasons why the number of ATP is not known exactly
© 2015 Pearson Education Ltd
Figure 10.16
Electron shuttlesspan membrane
CYTOSOL 2 NADHor
2 FADH2
2 NADH 2 FADH26 NADH
MITOCHONDRION
OXIDATIVEPHOSPHORYLATION(Electron transportand chemiosmosis)
CITRICACID
CYCLE
PYRUVATEOXIDATION
2 Acetyl CoA
GLYCOLYSIS
Glucose 2Pyruvate
+ 2 ATP + 2 ATP
Maximum per glucose: About30 or 32 ATP
+ about 26 or 28 ATP
2 NADH
© 2015 Pearson Education Ltd
Concept 10.5: Fermentation and anaerobic respiration enable cells to produce ATP without the use of oxygen
a) Most cellular respiration requires O2 to produce ATP
b) Without O2, the electron transport chain will cease to operate
c) In that case, glycolysis couples with anaerobic respiration or fermentation to produce ATP
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a) Anaerobic respiration uses an electron transport chain with a final electron acceptor other than O2, for example sulfate
b) Fermentation uses substrate-level phosphorylation instead of an electron transport chain to generate ATP
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Types of Fermentation
a) Fermentation consists of glycolysis plus reactions that regenerate NAD+, which can be reused by glycolysis
b) Two common types are alcohol fermentation and lactic acid fermentation
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a) In alcohol fermentation, pyruvate is converted to ethanol in two steps
a)The first step releases CO2
b)The second step produces ethanol
b) Alcohol fermentation by yeast is used in brewing, winemaking, and baking
© 2015 Pearson Education Ltd
Figure 10.17
2 ADP + 2 P i 2 ATP
Glucose
2 NAD+ NADH2+ 2 H+
2 Pyruvate
CO22
2 Ethanol
(a) Alcohol fermentation
2 Acetaldehyde 2 LactateLactic acid fermentation
2 ADP + 2 P i 2 ATP
GLYCOLYSIS
NAD+
+ 2 H+
NADH22
2 Pyruvate
(b)
GLYCOLYSIS Glucose
© 2015 Pearson Education Ltd
Figure 10.17a
2 ADP + 2 P i
NAD+
+ 2 H+
GLYCOLYSISGlucose
2 ATP
22
2 Ethanol 2 Acetaldehyde
2 Pyruvate
(a) Alcohol fermentation
2 CO2NADH
© 2015 Pearson Education Ltd
Figure 10.17b
2 ADP + 2 P i
NAD+
+ 2 H+
GLYCOLYSISGlucose
2 ATP
22
Lactate(b) Lactic acid fermentation
NADH
2 Pyruvate
2
© 2015 Pearson Education Ltd
Animation: Fermentation Overview
© 2015 Pearson Education Ltd
a) In lactic acid fermentation, pyruvate is reduced by NADH, forming lactate as an end product, with no release of CO2
b) Lactic acid fermentation by some fungi and bacteria is used to make cheese and yogurt
c) Human muscle cells use lactic acid fermentation to generate ATP when O2 is scarce
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Comparing Fermentation with Anaerobic and Aerobic Respiration
a) All use glycolysis (net ATP = 2) to oxidize glucose and harvest chemical energy of food
b) In all three, NAD+ is the oxidizing agent that accepts electrons during glycolysis
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a) The processes have different mechanisms for oxidizing NADH:
a)In fermentation, an organic molecule (such as pyruvate or acetaldehyde) acts as a final electron acceptor
b)In cellular respiration electrons are transferred to the electron transport chain
b) Cellular respiration produces 32 ATP per glucose molecule; fermentation produces 2 ATP per glucose molecule
© 2015 Pearson Education Ltd
Figure 10.18Glucose
GlycolysisCYTOSOL
Pyruvate
No O2 present:Fermentation
O2 present: Aerobic cellular respiration
Ethanol,lactate, or
other products
MITOCHONDRIONAcetyl CoA
CITRICACID
CYCLE
© 2015 Pearson Education Ltd
The Evolutionary Significance of Glycolysis
a) Ancient prokaryotes are thought to have used glycolysis long before there was oxygen in the atmosphere
b) 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
c) Glycolysis is a very ancient process
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Concept 10.6: Glycolysis and the citric acid cycle connect to many other metabolic pathways
a) Gycolysis and the citric acid cycle are major intersections to various catabolic and anabolic pathways
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The Versatility of Catabolism
a) Catabolic pathways funnel electrons from many kinds of organic molecules into cellular respiration
b) Glycolysis accepts a wide range of carbohydrates
c) Proteins must be digested to amino acids; amino groups can feed glycolysis or the citric acid cycle
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a) Fats are digested to glycerol (used in glycolysis) and fatty acids (used in generating acetyl CoA)
b) Fatty acids are broken down by beta oxidation and yield acetyl CoA
c) An oxidized gram of fat produces more than twice as much ATP as an oxidized gram of carbohydrate
© 2015 Pearson Education Ltd
Figure 10.19-1
Proteins Carbohydrates Fats
Aminoacids
Sugars Glycerol Fattyacids
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Figure 10.19-2
Proteins Carbohydrates Fats
Aminoacids
Sugars Glycerol Fattyacids
GLYCOLYSISGlucose
Glyceraldehyde 3- P
PyruvateNH3
© 2015 Pearson Education Ltd
Figure 10.19-3
Proteins Carbohydrates Fats
Aminoacids
Sugars Glycerol Fattyacids
GLYCOLYSISGlucose
Glyceraldehyde 3- P
Pyruvate
Acetyl CoA
NH3
© 2015 Pearson Education Ltd
Figure 10.19-4
Proteins Carbohydrates Fats
Aminoacids
Sugars Glycerol Fattyacids
GLYCOLYSISGlucose
Glyceraldehyde 3- P
Pyruvate
Acetyl CoA
CITRICACID
CYCLE
NH3
© 2015 Pearson Education Ltd
Figure 10.19-5
Proteins Carbohydrates Fats
Aminoacids
Sugars Glycerol Fattyacids
GLYCOLYSISGlucose
Glyceraldehyde 3- P
Pyruvate
Acetyl CoA
CITRICACID
CYCLE
OXIDATIVEPHOSPHORYLATION
NH3
© 2015 Pearson Education Ltd
Biosynthesis (Anabolic Pathways)
a) The body uses small molecules to build other substances
b) These small molecules may come directly from food, from glycolysis, or from the citric acid cycle
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Regulation of Cellular Respiration via Feedback Mechanisms
a) Feedback inhibition is the most common mechanism for metabolic control
b) If ATP concentration begins to drop, respiration speeds up; when there is plenty of ATP, respiration slows down
c) Control of catabolism is based mainly on regulating the activity of enzymes at strategic points in the catabolic pathway
© 2015 Pearson Education Ltd
Figure 10.UN17