chapter 10 cell respiration - ju medicine · cellular respiration during cellular respiration, the...
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Lecture Presentations by
Nicole Tunbridge and
Kathleen Fitzpatrick
Chapter 10
Cell Respiration
© 2018 Pearson Education Ltd.
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Life Is Work
▪ Living cells require energy from outside sources to
do work
▪ The work of the cell includes assembling polymers,
membrane transport, moving, and reproducing
▪ Animals can obtain energy to do this work by feeding
on other animals or photosynthetic organisms
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Figure 10.1
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Figure 10.1a
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▪ Energy flows into an ecosystem as sunlight and
leaves as heat
▪ The chemical elements essential to life are recycled
▪ Photosynthesis generates O2 and organic
molecules, which are used in cellular respiration
▪ Cells use chemical energy stored in organic
molecules to generate ATP, which powers work
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Figure 10.2
Lightenergy
ECOSYSTEM
Photosynthesisin chloroplasts
Cellular respirationin mitochondria
CO2 + H2O + O2Organic
molecules
ATP
Heatenergy
ATP powersmost cellular work
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BioFlix: The Carbon Cycle
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Concept 10.1: Catabolic pathways yield energy
by oxidizing organic fuels
▪ Catabolic pathways release stored energy by
breaking down complex molecules
▪ Electron transfer plays a major role in these
pathways
▪ These processes are central to cellular respiration
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Catabolic Pathways and Production of ATP
▪ 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
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▪ 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)
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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
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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)
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Figure 10.UN01
becomes oxidized(loses electron)
becomes reduced(gains electron)
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Figure 10.UN02
becomes oxidized
becomes reduced
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▪ The electron donor is called the reducing agent
▪ The electron receptor is called the oxidizing agent
▪ Some redox reactions do not transfer electrons but
change the electron sharing in covalent bonds
▪ An example is the reaction between methane and O2
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Figure 10.3
Reactants
becomes oxidized
Methane(reducing
agent)
Products
Energy
becomes reduced
Oxygen(oxidizing
agent)
Carbon dioxide Water
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Oxidation of Organic Fuel Molecules During
Cellular Respiration
▪ During cellular respiration, the fuel (such as glucose)
is oxidized, and O2 is reduced
▪ Organic molecules with an abundance of hydrogen
are excellent sources of high-energy electrons
▪ Energy is released as the electrons associated with
hydrogen ions are transferred to oxygen, a lower
energy state
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Figure 10.UN03
becomes oxidized
becomes reduced
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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
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Figure 10.4
NAD+
DehydrogenaseNADH
2[H](from food)
Reduction of NAD+
Oxidation of NADH
Nicotinamide(oxidized form)
Nicotinamide(reduced form)
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Figure 10.4a
NAD+
Nicotinamide(oxidized form)
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Figure 10.4b
NAD+
DehydrogenaseNADH
2[H](from food)
Reduction of NAD+
Oxidation of NADH
Nicotinamide(oxidized form)
Nicotinamide(reduced form)
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Figure 10.UN04
Dehydrogenase
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▪ 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
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Figure 10.5
H2 2 H + ½ O2
2 H+ + 2 e–
Controlledrelease of
energy
ATP
ATP
ATP
2 e–
2 H+½ O2
Fre
e e
nerg
y,
G
Fre
e e
nerg
y,
G
Explosiverelease of
energy
H2O H2O
(a) Uncontrolled reaction (b) Cellular respiration
½ O2+
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The Stages of Cellular Respiration: A Preview
▪ Harvesting of energy from glucose has three stages
1. Glycolysis (breaks down glucose into two
molecules of pyruvate)
2. The citric acid cycle (completes the breakdown of
glucose)
3. Oxidative phosphorylation (accounts for most of
the ATP synthesis)
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Figure 10.UN05
GLYCOLYSIS (color-coded blue throughout the chapter)
PYRUVATE OXIDATION and the CITRIC ACID CYCLE
(color-coded light orange and dark orange)
OXIDATIVE PHOSPHORYLATION: Electron transport and
chemiosmosis (color-coded purple)
1.
2.
3.
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Figure 10.6_1
Electronsvia NADH
GLYCOLYSIS
Glucose Pyruvate
CYTOSOL MITOCHONDRION
ATP
Substrate-level
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Figure 10.6_2
Electronsvia NADH
Electronsvia NADH and FADH2
GLYCOLYSIS
Glucose Pyruvate
PYRUVATEOXIDATION
Acetyl CoA
CITRICACID
CYCLE
CYTOSOL MITOCHONDRION
ATP
Substrate-level
ATP
Substrate-level
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Figure 10.6_3
Electronsvia NADH
Electronsvia NADH and FADH2
GLYCOLYSIS
Glucose Pyruvate
PYRUVATEOXIDATION
Acetyl CoA
CITRICACID
CYCLE
OXIDATIVEPHOSPHORYLATION
(Electron transportand chemiosmosis)
CYTOSOL MITOCHONDRION
ATP
Substrate-level
ATP
Substrate-level
ATP
Oxidative
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BioFlix: Cellular Respiration
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▪ The process that generates almost 90% of the ATP
is called oxidative phosphorylation because it is
powered by redox reactions
▪ A smaller amount of ATP is formed in glycolysis and
the citric acid cycle by substrate-level
phosphorylation
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Figure 10.7
Enzyme
ADP
P
Enzyme
ATP
Substrate
Product
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▪ For each molecule of glucose degraded to CO2 and
water by respiration, the cell makes up to 32
molecules of ATP
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▪ We can use money as an analogy for cellular
respiration:
▪ Glucose is like a larger-denomination bill—it is worth a
lot, but it is hard to spend
▪ ATP is like a number of smaller-denomination bills of
equivalent value—they can be spent more easily
▪ Cellular respiration cashes in a large denomination of
energy (glucose) for the small change of many
molecules of ATP
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Concept 10.2: Glycolysis harvests chemical
energy by oxidizing glucose to pyruvate
▪ Glycolysis (“sugar splitting”) 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
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Figure 10.UN06
GLYCOLYSISPYRUVATEOXIDATION
CITRICACID
CYCLE
OXIDATIVE PHOSPHORYL-
ATION
ATP
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Figure 10.8
Energy Investment Phase
Glucose
2 ADP + 22 ATP used
Energy Payoff Phase
4 ADP + 4
2 NAD+ + 4 e– + 4 H+
4 ATP formed
NADH
2 Pyruvate + 2 H2O
NetGlucose
4 ATP formed – 2 ATP used
2 NAD+ + 4 e– + 4 H+
2 Pyruvate + 2 H2O
2 ATP
2 NADH + 2 H+
P
P
2 + 2 H+
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Figure 10.9a
GLYCOLYSIS: Energy Investment Phase
Glyceraldehyde3-phosphate (G3P)
GlucoseATP Glucose
6-phosphateFructose
6-phosphateATP
Fructose1,6-bisphosphate
Aldolase
Isomerase
ADPADP
Phosphogluco-
isomerase
Hexokinase Phospho-
fructokinaseDihydroxyacetonephosphate (DHAP)
5
432
1
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Figure 10.9aa_1
GLYCOLYSIS: Energy Investment Phase
Glucose
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Figure 10.9aa_2
GLYCOLYSIS: Energy Investment Phase
ATPGlucose
Glucose
6-phosphate
ADP
Hexokinase
1
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Figure 10.9aa_3
GLYCOLYSIS: Energy Investment Phase
ATPGlucose
Glucose
6-phosphate
ADP
Fructose
6-phosphate
Hexokinase Phosphogluco-isomerase
12
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Figure 10.9ab_1
GLYCOLYSIS: Energy Investment Phase
Fructose6-phosphate
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Figure 10.9ab_2
GLYCOLYSIS: Energy Investment Phase
Fructose6-phosphate
ATP
ADP
Fructose1,6-bisphosphate
Phospho-fructokinase
3
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Figure 10.9ab_3
GLYCOLYSIS: Energy Investment Phase
Glyceraldehyde3-phosphate (G3P)
Fructose6-phosphate
ATP
ADP
Fructose1,6-bisphosphate
Phospho-fructokinase
Aldolase
Isomerase
Dihydroxyacetonephosphate (DHAP)
34
5
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Figure 10.9b
GLYCOLYSIS: Energy Payoff Phase
2 ATP
2 ADP
2
2 NAD+
2 NADH
+ 2 H+
2 H2O
2 2 2
ATP2 ADP
2
Glycer-
aldehyde
3-phosphate
(G3P)
Triosephosphate
dehydrogenase
Phospho-glycerokinase
Phospho-glyceromutase
Enolase
1,3-Bisphospho-
glycerate
3-Phospho-
glycerate
Pyruvatekinase
2-Phospho-
glycerate
Phosphoenol-
pyruvate (PEP)
Pyruvate
2
2 Pi
109
876
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Figure 10.9ba_1
GLYCOLYSIS: Energy Payoff Phase
Glyceraldehyde3-phosphate (G3P)
Isomerase
AldolaseDihydroxyacetonephosphate (DHAP)
4
5
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Figure 10.9ba_2
GLYCOLYSIS: Energy Payoff Phase
Glyceraldehyde3-phosphate (G3P)
2
2 NAD+ + 2 H+
Isomerase
Aldolase
Triosephosphate
dehydrogenaseP i
Dihydroxyacetonephosphate (DHAP)
1,3-Bisphospho-glycerate
2
4
5 6
2
NADH
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Figure 10.9ba_3
GLYCOLYSIS: Energy Payoff Phase
ATP
2 ADPGlyceraldehyde
3-phosphate (G3P) 2 NAD+ + 2 H+ 2
Isomerase
Aldolase
Triosephosphate
dehydrogenaseP i
Phospho-glycerokinase
Dihydroxyacetonephosphate (DHAP)
1,3-Bisphospho-glycerate
3-Phospho-glycerate
2
2
4
5 67
2
2 NADH
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Figure 10.9bb_1
GLYCOLYSIS: Energy Payoff Phase
2
3-Phospho-glycerate
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Figure 10.9bb_2
GLYCOLYSIS: Energy Payoff Phase
2 H2O
2
Phospho-glyceromutase
2
Enolase
2
3-Phospho-glycerate
2-Phospho-glycerate
Phosphoenol-pyruvate (PEP)
89
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Figure 10.9bb_3
GLYCOLYSIS: Energy Payoff Phase
2 H2O
2
Phospho-glyceromutase
2
2 ADP
ATP
22
Enolase
2
Pyruvatekinase
3-Phospho-glycerate
2-Phospho-glycerate
Phosphoenol-pyruvate (PEP)
Pyruvate8
910
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Concept 10.3: After pyruvate is oxidized, the
citric acid cycle completes the energy-yielding
oxidation of organic molecules
▪ In the presence of O2, pyruvate enters a
mitochondrion (in eukaryotic cells), where the
oxidation of glucose is completed
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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 multienzyme complex
that catalyzes three reactions
1. Oxidation of pyruvate and release of CO2
2. Reduction of NAD+ to NADH
3. Combination of the remaining two-carbon fragment
and coenzyme A to form acetyl CoA
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Figure 10.UN07
GLYCOLYSISPYRUVATEOXIDATION
CITRICACID
CYCLE
OXIDATIVE PHOSPHORYL-
ATION
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Figure 10.10
CYTOSOL
NAD+
CO2
MITOCHONDRION
Coenzyme A
NADH Acetyl CoAPyruvate
Transport protein
+ H+
1
2
3
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The Citric Acid Cycle
▪ The citric acid cycle, also called the Krebs cycle,
completes the breakdown of pyruvate to CO2
▪ The cycle oxidizes organic fuel derived from
pyruvate, generating 1 ATP, 3 NADH, and 1 FADH2
per turn
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▪ The citric acid cycle has eight steps, each catalyzed
by a specific enzyme
▪ The acetyl group of acetyl CoA joins the cycle by
combining with oxaloacetate, forming citrate
▪ The next seven steps decompose the citrate back to
oxaloacetate, making the process a cycle
▪ The NADH and FADH2 produced by the cycle relay
electrons extracted from food to the electron
transport chain
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Figure 10.UN08
ATP
GLYCOLYSISPYRUVATEOXIDATION
CITRICACID
CYCLE
OXIDATIVE PHOSPHORYL-
ATION
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Figure 10.11
PYRUVATE OXIDATION
Pyruvate(from glycolysis,2 molecules per glucose)
CO2
CoA
NADH
+ H+
NAD+
Acetyl CoA
CoA
NADH
NAD+
CoA
CITRICACID
CYCLE
FADH2
FAD
ADP + P i
ATP
2
2 NAD+
NADH
+ H+
CO2
2
+ 2 H+
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Figure 10.11a
PYRUVATE OXIDATION
Pyruvate(from glycolysis,2 molecules per glucose)
NAD+
NADH
+ H+Acetyl CoA
CoA
CO2
CoA
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Figure 10.11b
Acetyl CoA
CoA
NADH
+ H+
NAD+
CoA
CITRICACID
CYCLE
FADH2
FAD
ADP + P
ATP
2
2 NAD+
2
+ 2 H+
i
NADH
CO2
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Figure 10.12_1
Acetyl CoA
CoA-SH
Oxaloacetate
Citrate
1
CITRICACID
CYCLE
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Figure 10.12_2
Acetyl CoA
CoA-SH
H2O
Oxaloacetate
CitrateIsocitrate
1
2
CITRICACID
CYCLE
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Figure 10.12_3
Acetyl CoA
CoA-SH
H2O
Oxaloacetate
CitrateIsocitrate
NAD+
+ H+
α-Ketoglutarate
1
2
3CITRICACID
CYCLE
NADH
CO2
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Figure 10.12_4
Acetyl CoA
CoA-SH
H2O
Oxaloacetate
CitrateIsocitrate
CoA-SH
NAD+
+ H+
α-Ketoglutarate
NAD+
+ H+
SuccinylCoA
1
2
3
4
CITRICACID
CYCLE
NADH
CO2
NADH
CO2
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Figure 10.12_5
Acetyl CoA
CoA-SH
H2O
Oxaloacetate
CitrateIsocitrate
CITRICACID
CYCLECoA-SH
NAD+
+ H+
CoA-SH
α-Ketoglutarate
NAD+
+ H+Succinate
ADP
ATP
GTP SuccinylCoA
GDP
1
2
3
5
4
Pi
NADH
CO2
NADH
CO2
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Figure 10.12_6
Acetyl CoA
CoA-SH
H2O
Oxaloacetate
CitrateIsocitrate
CITRICACID
CYCLEFumarate CoA-SH
NAD+
+ H+
CoA-SH
α-Ketoglutarate
FADNAD+
+ H+Succinate
ADP
ATP
GTP SuccinylCoA
GDP
1
2
3
6
5
4
Pi
NADH
CO2
FADH2
NADH
CO2
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Figure 10.12_7
Acetyl CoA
CoA-SH
H2O
Oxaloacetate
Malate CitrateIsocitrate
H2O
CITRICACID
CYCLEFumarate CoA-SH
NAD+
+ H+
CoA-SH
α-Ketoglutarate
FADNAD+
+ H+Succinate
ADP
ATP
GTP SuccinylCoA
GDP
1
2
37
6
5
4
Pi
NADH
CO2
FADH2
NADH
CO2
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Figure 10.12_8
Acetyl CoA
CoA-SH
NADH
+ H+
NAD+
H2O
Oxaloacetate
Malate CitrateIsocitrate
H2O
CITRICACID
CYCLEFumarate CoA-SH
NAD+
NADH
+ H+
CO2
CoA-SH
α-Ketoglutarate
FADH2
FADNAD+
NADH
+ H+
CO2
Succinate
ADP
ATP
GTP SuccinylCoA
GDP
1
2
3
8
7
6
5
4
Pi
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Figure 10.12a
Acetyl CoA
CoA-SH
H2O
Oxaloacetate
Citrate
Isocitrate
2
1
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Figure 10.12b
Isocitrate
CoA-SH
NAD+
NADH
+ H+
CO2
α-Ketoglutarate
NAD+
NADH
+ H+
CO2
SuccinylCoA
3
4
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Figure 10.12c
Fumarate
CoA-SH
FADH2
FAD
Succinate
GTP
ADP
ATP
P i
SuccinylCoA
GDP
6
5
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Figure 10.12d
NADH
+ H+
NAD+
Malate
Oxaloacetate
H2O
Fumarate
8
7
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Concept 10.4: 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
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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
▪ Electrons drop in free energy as they go down the
chain and are finally passed to O2, forming H2O
▪ Electron carriers alternate between reduced and
oxidized states as they accept and donate electrons
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▪ 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
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Figure 10.UN09
ATP
GLYCOLYSISPYRUVATEOXIDATION
CITRICACID
CYCLE
OXIDATIVE PHOSPHORYL-
ATION
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Figure 10.13
NADH
50
NAD+
FADH2
Fre
e e
ne
rgy (
G)
rela
tive
to
O2
(kc
al/
mo
l)
40 FMN I
Fe•S
FAD
III
Fe•S
Cyt c1
Cyt c
Electron transport chain 20
Cyt a
Cyt a3
IV
Fe•S
Q
Cyt b
30
10 2
02 H+ + ½
(most electronegative)
H2O
O2
e–
2 e–
2 e–
(least electronegative)
II
Complexes I-IV
each consist of multiple proteins with electroncarriers.
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Figure 10.13a
NADH
NAD+
FADH2
FMNI
Fe•S
FAD
Complexes I-IVeach consist of multiple proteins with electroncarriers.
III
Fe•S
Cyt c1
Cyt c
Electron transport chain
Cyt a
Cyt a3
IV
Fe•S
Q
Cyt b
2 e–
2 e–
(least electronegative)
II
2 e–
20
30
10
Fre
e e
nerg
y (
G)
rela
tive t
o O
2(k
cal/
mo
l)
40
50
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Figure 10.13b
Cyt c1
Cyt c
20
Cyt a
Cyt a3
IV
30
10 2
02 H+ + ½
(most electronegative)
H2O
O2
e–
Fre
e e
ne
rgy (
G)
rela
tive
to
O2
(kc
al/m
ol)
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Chemiosmosis: The Energy-Coupling
Mechanism
▪ The energy released as electrons are passed down
the electron transport chain is used to pump H+ from
the mitochondrial matrix to the intermembrane space
▪ H+ then moves down its concentration gradient back
across the membrane, passing through the protein
complex ATP synthase
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▪ H+ moves into binding sites on the rotor of ATP
synthase, causing it to spin in a way that catalyzes
phosphorylation of ADP to ATP
▪ This is an example of chemiosmosis, the use of
energy in a H+ gradient to drive cellular work
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Figure 10.14
INTERMEMBRANE SPACE
Rotor
H+ Stator
Internal rod
Catalytic knob
MITOCHONDRIAL MATRIX
ADP
+
P i ATP
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Video: ATP Synthase 3-D Structure, Top View
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Video: ATP Synthase 3-D Structure, Side View
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▪ Certain electron carriers in the electron transport
chain accept and release H+ along with the electrons
▪ In this way, 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
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Figure 10.15
Proteincomplexof electroncarriers
H+
Cyt c
ATPsynthase
Q
I
II
FADH2
NADH NAD+
III
IV
2 H+ + ½ O2 H2O
ADP + P i
H+
ATP
(carrying electronsfrom food)
Electron transport chain Chemiosmosis
Oxidative phosphorylation
2
FAD
H+H+
H+
1
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Figure 10.15a
Proteincomplexof electroncarriers
H+
Cyt c
Q
I
II
FADH2
NADH NAD+
III
IV
2 H+ + ½ O2 H2O
(carrying electronsfrom food)
Electron transport chain
FAD
H+H+
1
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Figure 10.15b
ATPsynthase
ADP + P i
H+
ATP
Chemiosmosis2
H+
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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
▪ The rest of the energy is lost as heat
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Figure 10.16
Electron shuttlesspan membrane
CYTOSOL
2 NADH
2 NADH
or
2 FADH2
2 NADH 6 NADH
MITOCHONDRION
2 FADH2
GLYCOLYSIS
Glucose 2 Pyruvate
PYRUVATEOXIDATION
2 Acetyl CoA
CITRICACID
CYCLE
OXIDATIVEPHOSPHORYLATION
(Electron transportand chemiosmosis)
+ 2 ATP
Maximum per glucose:
+ 2 ATP
About30 or 32 ATP
+ about 26 or 28 ATP
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Figure 10.16a
Electron shuttlesspan membrane 2 NADH
or
2 FADH2
2 NADH
GLYCOLYSIS
Glucose 2Pyruvate
+ 2 ATP© 2018 Pearson Education Ltd.
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Figure 10.16b
2 NADH 6 NADH 2 FADH2
PYRUVATEOXIDATION
2 Acetyl CoA
CITRICACID
CYCLE
+ 2 ATP
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Figure 10.16c
2 NADH
or
2 FADH2
2 NADH 6 NADH 2 FADH2
OXIDATIVEPHOSPHORYLATION
(Electron transportand chemiosmosis)
+ about 26 or 28 ATP
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Figure 10.16d
Maximum per glucose:About
30 or 32 ATP
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▪ There are three reasons why the number of ATP is
not known exactly
1. Photophosphorylation and the redox reactions are
not directly coupled; the ratio of NADH to ATP
molecules is not a whole number
2. ATP yield varies depending on whether electrons
are passed to NAD+ or FAD in the mitochondrial
matrix
3. The proton-motive force is also used to drive other
kinds of work
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Concept 10.5: Fermentation and anaerobic
respiration enable cells to produce ATP without
the use of oxygen
▪ Most cellular respiration depends on electronegative
oxygen to pull electrons down the transport chain
▪ Without oxygen, the electron transport chain will
cease to operate
▪ In that case, glycolysis couples with anaerobic
respiration or fermentation to produce ATP
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▪ Anaerobic respiration uses an electron transport
chain with a final electron acceptor other than
oxygen, for example, sulfate
▪ Fermentation uses substrate-level phosphorylation
instead of an electron transport chain to generate
ATP
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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
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▪ In alcohol fermentation, pyruvate is converted to
ethanol in two steps
▪ The first step releases CO2 from pyruvate
▪ The second step produces NAD+ and ethanol
▪ Alcohol fermentation by yeast is used in brewing,
winemaking, and baking
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Figure 10.17
2 ADP + 2 P i 2 2 ADP + 2 P i 2
Glucose GLYCOLYSIS Glucose GLYCOLYSIS
2 Pyruvate
2 NAD+ 2 NADH
+ 2 H+
2 2 NAD+ 2 NADH
+ 2 H+
2 Pyruvate
NAD+ REGENERATIONNAD+ REGENERATION
2 Ethanol
(a) Alcohol fermentation
2 Acetaldehyde 2 Lactate
(b) Lactic acid fermentation
ATP ATP
CO2
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Figure 10.17a
Glucose GLYCOLYSIS
2 Pyruvate
2 NADH
+ 2 H+
2 Ethanol
(a) Alcohol fermentation
2 Acetaldehyde
NAD+ 2
NAD+ REGENERATION
2 ADP ATP+ 2 P i 2
CO22
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▪ In lactic acid fermentation, pyruvate is reduced by
NADH, forming NAD+ and lactate as end products,
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 during strenuous exercise when O2 is
scarce
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Figure 10.17b
2 ADP ATP
Glucose GLYCOLYSIS
NAD+ NADH
+ 2 H+
2 Pyruvate
NAD+
2 Lactate
(b) Lactic acid fermentation
+ 2 P i 2
REGENERATION
2 2
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Animation: Fermentation Overview
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Comparing Fermentation with Anaerobic and
Aerobic Respiration
▪ All use glycolysis (net ATP = 2) to oxidize glucose
and harvest the chemical energy of food
▪ In all three, NAD+ is the oxidizing agent that accepts
electrons during glycolysis
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▪ The processes have different mechanisms for
oxidizing NADH to NAD+:
▪ In fermentation, an organic molecule (such as
pyruvate or acetaldehyde) acts as a final electron
acceptor
▪ In cellular respiration, electrons are transferred to the
electron transport chain
▪ Cellular respiration produces 32 ATP per glucose
molecule; fermentation produces 2 ATP per glucose
molecule
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▪ 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
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Figure 10.18
Glucose
Pyruvate
Glycolysis
No O2 present:Fermentation
O2 present:Aerobic cellularrespiration
CYTOSOL
MITOCHONDRION
Ethanol,lactate, or
other products
Acetyl CoA
CITRICACID
CYCLE
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The Evolutionary Significance of Glycolysis
▪ Glycolysis is an ancient process
▪ Early prokaryotes likely used glycolysis to produce
ATP before O2 accumulated in the atmosphere
▪ Used in both cellular respiration and fermentation, it
is the most widespread metabolic pathway on Earth
▪ This pathway occurs in the cytosol so does not
require the membrane-bound organelles of
eukaryotic cells
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Concept 10.6: 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
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The Versatility of Catabolism
▪ Catabolic pathways funnel electrons from many
kinds of organic molecules into cellular respiration
▪ Glycolysis accepts a wide range of carbohydrates
including starch, glycogen, and several
disaccharides
▪ Proteins that are used for fuel must be digested to
amino acids and their amino groups must be
removed
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▪ Fats are digested to glycerol (used to produce
compounds needed for glycolysis) and fatty acids
▪ Fatty acids are broken down by beta oxidation and
yield acetyl CoA, NADH, and FADH2
▪ An oxidized gram of fat produces more than twice as
much ATP as an oxidized gram of carbohydrate
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Figure 10.19_1
Proteins
Aminoacids
Carbohydrates
Sugars
Fats
Glycerol Fattyacids
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Figure 10.19_2
Proteins
Aminoacids
Carbohydrates
Sugars
Fats
Glycerol Fattyacids
GLYCOLYSIS
Glucose
Glyceraldehyde 3-
NH3 Pyruvate
P
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Figure 10.19_3
Proteins
Aminoacids
Carbohydrates
Sugars
Fats
Glycerol Fattyacids
GLYCOLYSIS
Glucose
NH3 Pyruvate
Acetyl CoA
Glyceraldehyde 3- P
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Figure 10.19_4
Proteins
Aminoacids
Carbohydrates
Sugars
Fats
Glycerol Fattyacids
GLYCOLYSIS
Glucose
NH3 Pyruvate
Acetyl CoA
CITRICACID
CYCLE
Glyceraldehyde 3- P
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Figure 10.19_5
Proteins
Aminoacids
Carbohydrates
Sugars
Fats
Glycerol Fattyacids
GLYCOLYSIS
Glucose
NH3 Pyruvate
Acetyl CoA
CITRICACID
CYCLE
OXIDATIVEPHOSPHORYLATION
Glyceraldehyde 3- P
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Biosynthesis (Anabolic Pathways)
▪ The body uses small molecules from food to build
other their own molecules such as proteins
▪ 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
▪ Feedback inhibition is the most common mechanism
for metabolic 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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Figure 10.20Glucose
AMPGLYCOLYSIS
Fructose 6-phosphate
Inhibits
Phosphofructokinase–
Fructose 1,6-bisphosphateInhibits
Stimulates+
Pyruvate
ATPAcetyl CoA
Citrate
CITRICACID
CYCLE
Oxidativephosphorylation
–
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Figure 10.UN10a
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Figure 10.UN10b
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Figure 10.UN11
OutputsInputs
Glucose
GLYCOLYSIS
2 Pyruvate 2 2 NADHATP
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Figure 10.UN12
CO2 26 FADH2
82 ATP NADH
OutputsInputs
2 Pyruvate 2 Acetyl CoA
2 OxaloacetateCITRIC
ACIDCYCLE
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Figure 10.UN13
H+
H+H+INTERMEMBRANE
SPACE
MITOCHONDRIAL MATRIXNADH NAD+
(carrying electrons from food)
Proteincomplexof electroncarriers
FADH2FAD
I
II
III
QIV
Cyt c
2 H+ + ½ O2 H2O
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Figure 10.UN14
INTER-MEMBRANESPACE
MITO-
CHONDRIAL
MATRIXATPsynthase
ATPADP P H+i+
H+
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Figure 10.UN15
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Figure 10.UN16
Ph
osp
ho
fru
cto
kin
as
eacti
vit
y
Fructose 6-phosphateconcentration
High ATPconcentration
Low ATPconcentration
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Figure 10.UN17
Time
pH
dif
fere
nce
acro
ss m
em
bra
ne
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Figure 10.UN18
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