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Bioenergetics
Sengupta 1
Bioenergetics
1.Compare the difference between potential and kinetic energy.
2.Define the two laws of thermodynamics and explain how they relate to biological systems.
3.Explain how ATP functions as an energy shuttle. 4.Describe how enzymes speed up chemical reactions. 5.Describe how cellular respiration produces energy that can be stored in ATP.
6.Describe the three main stages of cellular respiration. 7.Discuss the importance of oxidative phosphorylation in producing ATP.
8.Compare ATP generation from substrate level phosphorylation and oxidative phosphorylation.
9.Compare cellular respiration and fermentation.
Learning Objectives
Energy Transformation in cells
• Cells are small units, a chemical factory, housing thousands of chemical reactions.
• Cells use these chemical reactions for – cell maintenance, – manufacture of cellular parts, and – cell replication.
Energy
Energy is the capacity to cause change or to perform work. There are two kinds of energy:
• Kinetic energy is the energy of motion. • Potential energy is energy that matter
possesses as a result of its location or structure.
Fuel Energy conversion Waste products
Gasoline
Oxygen
Oxygen
Glucose
+
+
+
+
Heat energy
Combustion Kinetic energy of movement
Energy conversion in a car
Energy conversion in a cell
Energy for cellular work
Cellular respiration
ATP ATP
Heat energy
Carbon dioxide
Carbon dioxide
Water
Water
• Heat or thermal energy, is a type of kinetic energy associated with the random movement of atoms or molecules.
• Light is kinetic energy, and can be harnessed to power photosynthesis.
• Electrical energy used in the nervous system. • Chemical energy is responsible for all the
biochemical reactions in the cell like ATP
Types of Energy used by Cells

Bioenergetics
Sengupta 2
Thermodynamics
• “Thermodynamics, science of the relationship between heat, work, temperature, and energy. In broad terms, thermodynamics deals with the transfer of energy from one place to another and from one form to another. The key concept is that heat is a form of energy corresponding to a definite amount of mechanical work”.
Encyclopedia Britannica
• Scientists use the word – system for the matter under study and – surroundings for the rest of the universe.
9
Biological Thermodynamics
• Quantitative study of the energy transductions that occur in and between living organisms, structures, and cells and of the nature and function of the chemical processes underlying these transductions.
• Biological thermodynamics may address the
question of whether the benefit associated with any particular phenotypic trait is worth the energy investment it requires.
Energy Transformations
Two laws govern energy transformations in living organisms.
– The first law of thermodynamics, energy in the universe is constant, (statement of the conservation of energy)and
– The second law of thermodynamics, energy conversions increase the disorder of the universe.
• Entropy is the measure of disorder, or randomness.
Energy Transforma.on Energy Transformations in Cells
• Cells use oxygen in reactions that release energy from fuel molecules.
• In cellular respiration, the chemical energy stored in organic molecules is converted to a form that the cell can use to perform work.

Bioenergetics
Sengupta 3
Energy and Chemical reactions
• Chemical reactions either – release energy (exergonic reactions)
or – require an input of energy and store
energy (endergonic reactions).
Exergonic reactions
Exergonic reactions release energy. – These reactions release the energy in covalent
bonds of the reactants – Burning wood releases the energy in glucose as
heat and light – Cellular respiration
• involves many steps, • releases energy slowly, and • uses some of the released energy to produce ATP.
Exergonic Reaction
Reactants
Energy Products
Amount of energy
released
Pot
entia
l ene
rgy
of m
olec
ules
Endergonic Reactions
Endergonic reaction – requires an input of energy and – yields products rich in potential energy.
Endergonic reactions – begin with reactant molecules that contain
relatively little potential energy but – end with products that contain more chemical
energy.
Endergonic Reaction
Reactants
Energy
Products
Amount of energy
required
Pot
entia
l ene
rgy
of m
olec
ules
Photosynthesis : endergonic reaction
Photosynthesis is a type of endergonic process.
– Energy-poor reactants, carbon dioxide, and water are used.
– Energy is absorbed from sunlight. – Energy-rich sugar molecules are produced.

Bioenergetics
Sengupta 4
Metabolism
• A living organism carries out thousands of endergonic and exergonic chemical reactions.
• Metabolism :The total of an organism’s chemical reactions.
• Metabolism = Anabolism + Catabolism • A metabolic pathway is a series of chemical
reactions that either – builds a complex molecule or – breaks down a complex molecule into simpler
compounds.
The Energy Cycle
Metabolic pathways
Breakdown Proteins to Amino Acids, Starch to Glucose
Synthesis Amino Acids to Proteins, Glucose to Starch
Energy Coupling
• Energy coupling uses the – energy released from exergonic reactions
to drive – essential endergonic reactions, – usually using the energy stored in ATP
molecules.
• ATP, adenosine triphosphate, powers
nearly all forms of cellular work.
• ATP consists of
– the nitrogenous base adenine,
– the five-carbon sugar ribose, and
– three phosphate groups.
The Energy Currency of the Cell Renewable cellular energy: ATP
• ATP is a renewable source of energy for the cell
• In the ATP cycle, energy released in an exergonic reaction, such as the breakdown of glucose,is used in an endergonic reaction to generate ATP

Bioenergetics
Sengupta 5
ATP drives cellular work
• Hydrolysis of ATP releases energy by transferring its third phosphate from ATP to some other molecule in a process called phosphorylation.
• Most cellular work depends on ATP energizing molecules by phosphorylating them.
Figure 5.12A_s1
Adenine P P P
Phosphate group
ATP: Adenosine Triphosphate
Ribose
ADP: Adenosine Diphosphate
P P P Energy
H2O Hydrolysis
Ribose
Adenine P P P
Phosphate group
ATP: Adenosine Triphosphate ATP drives cellular work
• There are three main types of cellular work: 1. chemical, 2. mechanical, and 3. transport
• ATP drives all three of these types of work
ATP ATP ATP
ADP ADP ADP P P P
P
P
P
P P P
Chemical work Mechanical work Transport work
Reactants
Motor protein
Solute
Membrane protein
Product
Molecule formed Protein filament moved Solute transported
HOW ENZYMES FUNCTION

Bioenergetics
Sengupta 6
Biological Catalysts
• Although biological molecules possess much potential energy, it is not released spontaneously. – An energy barrier must be overcome
before a chemical reaction can begin. – This energy is called the activation
energy (EA).
The Energy of Activation
• We can think of EA
– as the amount of energy needed for a reactant molecule to move “uphill” to a higher energy but an unstable state
– so that the “downhill” part of the reaction can begin.
• One way to speed up a reaction is to add heat, – which agitates atoms so that bonds break
more easily and reactions can proceed but – but could also kill a cell.
Enzymes lower energy of activation
Enzymes • are usually proteins, although some RNA
molecules can function as enzymes. • function as biological catalysts by lowering the
EA needed for a reaction to begin, • increase the speed of a reaction without being
consumed by the reaction
Activation energy barrier
Reactant
Products
Reaction without enzyme
Ene
rgy
Activation energy barrier reduced by enzyme
Reactant
Products
Reaction with enzyme
Ene
rgy
Enzyme
Metabolic reaction in a cell with and without enzyme
Reactants
Products
Ene
rgy
Progress of the reaction
a
b
c

Bioenergetics
Sengupta 7
Enzymes have a specific shape
• An enzyme – is very selective in the reaction it
catalyzes and – has a shape that determines the
enzyme’s specificity. • The specific reactant that an enzyme acts
on is called the enzyme’s substrate. • A substrate fits into a region of the enzyme
called the active site. • Enzymes are specific because their active
site fits only specific substrate molecules.
Figure 5.14_s2
2
1
Enzyme (sucrase)
Active site
Enzyme available with empty active site
Substrate (sucrose)
Substrate binds to enzyme with induced fit
Figure 5.14_s3
3
2
1
Enzyme (sucrase)
Active site
Enzyme available with empty active site
Substrate (sucrose)
Substrate binds to enzyme with induced fit
Substrate is converted to products
H2O
4
3
2
1
Products are released
Fructose
Glucose Enzyme (sucrase)
Active site
Enzyme available with empty active site
Substrate (sucrose)
Substrate binds to enzyme with induced fit
Substrate is converted to products
H2O
A �my�l�a�s �e
catalase lactase
Some common enzymes
http://mm.rcsb.org
Conditions for optimum enzyme activity
Every enzyme has optimal conditions under which it is most effective.
• Temperature affects molecular motion. – An enzyme’s optimal temperature produces the
highest rate of contact between the reactants and the enzyme’s active site.
– Most human enzymes work best at 35–40ºC. – But high temperature disrupts the structure
• The optimal pH for most enzymes is near neutrality.

Bioenergetics
Sengupta 8
Enzyme activity
Four Variables
Temperature
pH
Enzyme Concentration
Substrate Concentration
Effect of heat on enzyme activity
Denaturing protein
IF ACTIVE SITE CHANGES SHAPE: SUBSTRATE NO LONGER FITS
Even if temperature lowered – enzyme cannot regain its correct shape
Rat
e of
Rea
ctio
n
Temperature
0 20 30 50 10 40 60
40oC -denatures 5- 35oC Increase in Activity
<5oC - inactive
Environmental Conditions can Cause Changes in Shape (Denaturing)
Rat
e of
Rea
ctio
n
pH
1 3 4 2 5 6 7 8 9
Narrow pH optima
Disrupt Ionic bonds - Structure
Effect charged residues at active site
Cofactors and coenzymes
• Many enzymes require nonprotein helpers called cofactors, which – bind to the active site and – function in catalysis.
• Some cofactors are inorganic, such as zinc, iron, or copper.
• If a cofactor is an organic molecule, such as most vitamins, it is called a coenzyme.

Bioenergetics
Sengupta 9
Enzyme inhibitors
• A chemical that interferes with an
enzyme’s activity is called an inhibitor.
• Competitive inhibitors
– block substrates from entering the active
site and
– reduce an enzyme’s productivity.
Enzyme inhibitors
• Noncompetitive inhibitors
– bind to the enzyme somewhere other than the active site,
– change the shape of the active site, and
– prevent the substrate from binding.
Substrate
Enzyme
Allosteric site
Active site
Normal binding of substrate
Competitive inhibitor
Noncompetitive inhibitor
Enzyme inhibi.on Enzyme inhibitors regulate cell metabolism
• Enzyme inhibitors are important in regulating cell metabolism.
• In some reactions, the product may act as an inhibitor of one of the enzymes in the pathway that produced it. This is called feedback inhibition.
Feedback Inhibition
Feedback inhibition
Starting molecule
Product
Enzyme 1 Enzyme 2 Enzyme 3
Reaction 1 Reaction 2 Reaction 3 A B C D
Some drugs, pes.cides and poisons are enzyme inhibitors
• Many beneficial drugs act as enzyme inhibitors, including – Ibuprofen, inhibiting the production of
prostaglandins, – some blood pressure medicines, – some antidepressants, – many antibiotics, and – protease inhibitors used to fight HIV.
• Enzyme inhibitors have also been developed as pesticides and deadly poisons for chemical warfare.

Bioenergetics
Sengupta 10
The Marathoner and the Sprinter
The Marathoners and the Sprinters
Cellular respiration is the process by which cells produce energy aerobically
The different types of muscle fibers use different processes for making ATP
• Slow-twitch fibers undergo aerobic respiration (in the presence of O2) respiration are reddish in color as it contains
myoglobin
• Fast-twitch fibers undergo anaerobic respiration (in the absence of O2) respiration
Breathing and Cellular Respiration
Breathing
Lungs
Muscle cells carrying out Cellular Respiration
Bloodstream
Glucose + O2
CO2 + H2O + ATP
O2 CO2
CO2 O2
Connecting breathing and cellular respiration
Breathing and cellular respiration are closely related • Breathing brings O2 into the body from the
environment • O2 is distributed to cells in the bloodstream • In cellular respiration, mitochondria use O2 to
harvest energy and generate ATP • Breathing disposes of the CO2 produced as a
waste product of cellular respiration.
The human energy requirement
The human body uses energy from ATP for all its activities
– The body needs a continual supply of energy to maintain basic functioning: to breathe, to keep heart pumping and maintain body temperature.
– The brain requires 120 gm of glucose/day! – ATP supplies energy (kilocalories) for voluntary
activities
– An average adult human needs about 2,000 kcal of energy each day.
Cellular respiration and ATP molecules
• Cellular respiration is an exergonic process that transfers energy from the bonds in glucose to form ATP. ▪ Cellular respiration
– Can produce up to 32-34 ATP molecules from each glucose molecule and
– captures only about 34% of the energy originally stored in glucose.
▪ Other foods (organic molecules) can also be used as a source of energy.

Bioenergetics
Sengupta 11
Breaking down Glucose
Glucose Oxygen Water Carbon dioxide
C6H12O6 O2 H2O ATP 6 6 6
+ Heat
CO2
What is kcal?
• A kilocalorie (kcal) is – the quantity of heat required to raise the
temperature of 1 kilogram (kg) of water by 1oC,
– the same as a food Calorie, and – used to measure the nutritional values
indicated on food labels.
Activity kcal consumed per hour by a 67.5-kg (150-lb) person*
Running (8–9 mph)
Dancing (fast)
Bicycling (10 mph)
Swimming (2 mph)
Walking (4 mph)
Walking (3 mph)
Dancing (slow)
Driving a car
Sitting (writing)
*Not including kcal needed for body maintenance
979
510
490
408
341
245
204
61
28
How can energy be released from glucose?
• Energy can be released from glucose by simply burning it.
• The energy is dissipated as heat and light and is not available to living organisms.
How do cells tap energy from glucose?
• On the other hand, cellular respiration is the controlled breakdown of organic molecules.
• Energy is – gradually released in small amounts, – captured by a biological system, and – stored in ATP.
The redox reaction in cells
• Movement of electrons from one molecule to another is an oxidation-reduction reaction, or redox reaction. In a redox reaction, – the loss of electrons from one substance is
called oxidation, – the addition of electrons to another substance
is called reduction, – a molecule is oxidized when it loses one or
more electrons, and – reduced when it gains one or more electrons.

Bioenergetics
Sengupta 12
The Process
• A cellular respiration equation is helpful to show the changes in hydrogen atom distribution.
• Glucose – loses its hydrogen atoms and – becomes oxidized to CO2.
• Oxygen – gains hydrogen atoms and – becomes reduced to H2O.
The Redox Reaction in Cells
Glucose + Heat
C6H12O6 O2 CO2 H2O ATP 6 6 6
Loss of hydrogen atoms (becomes oxidized)
Gain of hydrogen atoms (becomes reduced)
Mitochondria in Action
Occurs in Cytoplasm
Occurs in Matrix
Occurs across Cristae
Where does Cellular Respiration Occur?
What are the three phases of Cellular Respiration ?
• Glycolysis • Citric Acid Cycle • Oxidative Phosphorylation
C6H12O6 CO2 O2 H2O
Glucose Oxygen Carbon dioxide
Water
+ 6 + 6 6
Reduction
Oxidation
Oxygen gains electrons (and hydrogens)
Glucose loses electrons (and hydrogens)

Bioenergetics
Sengupta 13
The role of Oxygen in Cellular Respiration
• Cellular respiration can produce up to 38 ATP molecules for each glucose molecule consumed.
• During cellular respiration, hydrogen and its bonding electrons change partners.
– Hydrogen and its electrons go from glucose to oxygen, forming water.
– This hydrogen transfer is why oxygen is so vital to cellular respiration.
Release of heat energy
1 2 H2 O2
H2O
Glucose and Oxygen
• Glucose
– loses its hydrogen atoms and – becomes oxidized to CO2.
• Oxygen – gains hydrogen atoms and – becomes reduced to H2O.
Controlled breaking of the glucose molecule to release energy
Glucose + Heat
C6H12O6 O2 CO2 H2O ATP 6 6 6
Loss of hydrogen atoms (becomes oxidized)
Gain of hydrogen atoms (becomes reduced)
The controlled breakdown of glucose
• There are other electron “carrier” molecules that function like NAD+. – They form an energy ‘staircase’ where the
electrons pass from one to the next down the next level.
– These electron carriers collectively are called the electron transport chain.
– As electrons are transported down the chain, ATP is generated.
The molecules in the process
I. Enzymes are necessary to oxidize glucose and other foods. II.NAD+
– is an important enzyme in oxidizing glucose,
– accepts electrons, and – becomes reduced to NADH.

Bioenergetics
Sengupta 14
• NAD+ picks up one hydrogen atom and another single electron from carbohydrate, becoming NADH.
• It will retain this form until it drops off its energetic electrons (and a proton) in a later stage of the energy-harvesting process.
NAD+ ATP and oxidative phosphorylation
NADH passes electrons to the electron transport chain.
– As electrons “move” from carrier to carrier and finally to O2, energy is released in small quantities.
– The energy released is used by the cell to make ATP.
The Electron Carrier NAD+
empty empty loaded
proton (oxidized)
used in later stage of respiration
used in later stage of respiration
goes to pick up more electrons
(reduced)
1. 2. NAD+ within a cell, along with two hydrogen atoms that are part of the food that is supplying energy for the body.
NAD+ is reduced to NAD by accepting an electron from a hydrogen atom. It also picks up another hydrogen atom to become NADH.
NADH carries the electrons to a later stage of respiration then drops them off, becoming oxidized to its original form, NAD+.
The energy gradient NADH
NAD+
Electron transport
chain
2e− Controlled release of energy for synthesis
of ATP
ATP
H+
2 H+ 2e−
H2O
O2 1/2
• Uses two ATP molecules per glucose to split the six-carbon glucose.
• Makes four additional ATP directly when enzymes transfer phosphate groups from fuel molecules to ADP. Glycolysis produces a net of two
molecules of ATP per glucose molecule.
Glycolysis: substrate level phosphorylation
Figure 6.7A
Glucose
2 Pyruvate
2 ADP
2 P 2 NAD+
2 NADH
2 H+ ATP 2

Bioenergetics
Sengupta 15
Glycolysis harvests chemical energy by oxidizing glucose to pyruvate
• ATP is formed in glycolysis by • substrate-level phosphorylation • during which
– an enzyme transfers a phosphate group from a substrate molecule to ADP and
– ATP is formed. • two NADH molecules are produced for each initial glucose
molecule and • ATP molecules are generated.
• The compounds that form between the initial reactant, glucose, and the final product, pyruvate, are called intermediates.
Substrate Level Phosphorylation
Enzyme Enzyme
P
P
ADP
P
ATP
Substrate Product
Pyruvate is chemically processed to enter the citric acid cycle
– A large, multienzyme complex catalyzes three reactions in the mitochondrial matrix
• A carbon atom is removed from pyruvate and released in CO2
• The remaining two-carbon compound is oxidized,
• molecule of NAD+ is reduced to NADH • Coenzyme A joins with the 2-carbon group to
produce acetyl CoA
From Glycolysis to Citric Acid Cycle Making Acetyl CoA
Pyruvate
Coenzyme A
Acetyl coenzyme A
NAD+ NADH H+
CoA
CO2 3
2
1
The citric acid cycle
• The citric acid cycle – is also called the Krebs cycle (after the German-
British researcher Hans Krebs, who worked out much of this pathway in the 1930s),
• Citric acid cycle processes two molecules of acetyl CoA for each initial glucose.
• Thus, after two turns of the citric acid cycle, the overall yield per glucose molecule is – 2 ATP, – 6 NADH and – 2 FADH2.
Acetyl CoA
Citric Acid Cycle
CoA CoA
CO2 2
3
3
NAD+
3 H+
NADH
ADP ATP P
FAD
FADH2

Bioenergetics
Sengupta 16
Figure 6.9B_s1 CoA
CoA
1
Acetyl CoA
Oxaloacetate
Citric Acid Cycle
2 carbons enter cycle
Step Acetyl CoA stokes the furnace.
1
Figure 6.9B_s2
NADH
NADH
NAD+
NAD+
H+
H+
CO2
CO2
ATP ADP P
CoA CoA
3 2 1
3
1
2
Acetyl CoA
Oxaloacetate
Citric Acid Cycle
2 carbons enter cycle
Citrate
leaves cycle
Alpha-ketoglutarate
leaves cycle
Step Acetyl CoA stokes the furnace.
Steps – NADH, ATP, and CO2 are generated during redox reactions.
Figure 6.9B_s3
NADH
NADH
NAD+
NAD+ NAD+ NADH
H+
H+
H+
CO2
CO2
ATP ADP P
FAD
FADH2
CoA CoA
3 2 1 4 5
3 4
5
1
2
Acetyl CoA
Oxaloacetate
Citric Acid Cycle
2 carbons enter cycle
Citrate
leaves cycle
Alpha-ketoglutarate
leaves cycle
Succinate
Malate
Step Acetyl CoA stokes the furnace.
Steps – NADH, ATP, and CO2 are generated during redox reactions.
Steps – Further redox reactions generate FADH2 and more NADH.
3 NAD+
ADP + P
3 NADH
FADH2 FAD
Acetic acid
Citric acid
Acceptor molecule
Citric Acid Cycle
ATP
2 CO2
INPUT OUTPUT
Phase 3: Oxidative phosphorylation
• Occurs in the cristae of mitochondria
• The molecules of the electron transport chain are built into the inner membranes of mitochondria.
• Uses the energy released by electrons “falling” down the electron transport chain to pump H+ across a membrane
• Harnesses the energy of the H+ gradient through chemiosmosis, producing ATP
– These ions store potential energy.
– requires an adequate supply of oxygen.
Making ATP in Bulk
• Electrons from NADH and FADH2 travel down the electron transport chain to O2.
• Oxygen picks up H+ to form water.
• Energy released by these redox reactions is used to pump H+ from the mitochondrial matrix into the intermembrane space.
• In chemiosmosis, the H+ diffuses back across the inner membrane through ATP synthase complexes, driving the synthesis of ATP.

Bioenergetics
Sengupta 17
The Electron Transport Chain
Oxidative Phosphorylation
Electron Transport Chain Chemiosmosis
Mito- chondrial matrix
Inner mito- chondrial membrane
Intermem- brane space
Electron flow
Protein complex of electron carriers
Mobile electron carriers ATP
synthase
NADH NAD+ 2 H+ FADH2 FAD
O2 H2O
ADP P ATP
1 2
H+
H+
H+
H+
H+ H+
H+
H+
H+ H+
H+
I
II
III IV
Oxidative Phosphorylation
H+
H+
H+ H+
H+
H+
H+ H+
H+
H+
H+
Oxidative Phosphorylation
Electron Transport Chain Chemiosmosis
I
II
IIIIV
NADH NAD+ 2 H+ FADH2 FAD
O2 H2O
ADP P ATP
2 1
Electron flow
Protein complex of electron carriers
Mobile electron carriers ATP
synthase
ATP synthase
http://www.mrc-mbu.cam.ac.uk/research/atp-synthase
• ATP synthase is a huge molecular complex (>500,000 daltons) embedded in the inner membrane of mitochondria. • Its function is to convert the energy of protons (H+) moving down their concentration gradient into the synthesis of ATP. • 3 to 4 protons moving through this machine is enough to convert a molecule of ADP and Pi (inorganic phosphate) into a molecule of ATP. • One ATP synthase complex can generate >100 molecules of ATP each second. http://mm.rcsb.org
Interrupting cellular respiration can have both harmful and beneficial effects
Three categories of cellular poisons obstruct the process of oxidative phosphorylation. These poisons
1. block the electron transport chain (for example, rotenone, cyanide, and carbon monoxide),
2. inhibit ATP synthase (for example, the antibiotic oligomycin), or
3. make the membrane leaky to hydrogen ions (called uncouplers, examples include dinitrophenol).
Poisons of the Cellular Respiration
ATP synthase
NAD+ NADH
FADH2 FAD
H+
H+ H+
H+
H+ H+ H+ H+
2 H+
H2O ADP ATP P
O2 2 1
Rotenone Cyanide, carbon monoxide
Oligomycin
DNP
Review: Each molecule of glucose yields many molecules of ATP
• The total yield is about 32 ATP molecules per glucose molecule.
• This is about 34% of the potential energy of a glucose molecule.
• In addition, water and CO2 are produced.

Bioenergetics
Sengupta 18
The overview of cellular respiration
NADH High-energy electrons
carried by NADH
GLYCOLYSIS Glucose Pyruvate
Cytoplasm
ATP Substrate-level phosphorylation
Substrate-level phosphorylation
CITRIC ACID CYCLE
CO2 CO2
ATP
NADH FADH2 and
ATP
Mitochondrion
Oxidative phosphorylation
OXIDATIVE PHOSPHORYLATION
(Electron Transport and Chemiosmosis)
Energy Harvest
2 ATP
2 ATP
34 ATP
glycolysis
38 ATP maximum per glucose molecule
Krebs cycle
electron transport
chain
glucose
(b) In schematic terms
(a) In metaphorical terms
mitochondrion H2O
O2
2 FADH2
2 NADH
2 NADH
6 NADH CO2
CO2 glucose derivatives
cytosol
glycolysis
Krebs cycle
electron transport
chain
insert 1 glucose
2 energy tokens
2 energy tokens
32 energy tokens
reactants products
The Energy Harvest
Each molecule of glucose yields many molecules of ATP
– Glycolysis and the citric acid cycle together yield four ATP per glucose molecule
– Oxidative phosphorylation, using electron transport and chemiosmosis, yields 34 ATP per glucose
– These numbers are maximum – Some cells may lose a few ATP to NAD+ or FAD
shuttles
FERMENTATION: ANAEROBIC HARVESTING OF ENERGY
Fermentation enables cells to produce ATP without oxygen
• Fermentation is a way of harvesting chemical energy that does not require oxygen. Fermentation – takes advantage of glycolysis, – produces two ATP molecules per glucose, and – reduces NAD+ to NADH.
• The trick of fermentation is to provide an anaerobic path for recycling NADH back to NAD+.
Fermentation enables cells to produce ATP without oxygen
• Your muscle cells and certain bacteria can oxidize NADH through lactic acid fermentation, in which – NADH is oxidized to NAD+ and – pyruvate is reduced to lactate.

Bioenergetics
Sengupta 19
Fermentation enables cells to make ATP without oxygen
• Lactate is carried by the blood to the liver, where it is converted back to pyruvate and oxidized in the mitochondria of liver cells.
• The dairy industry uses lactic acid fermentation by bacteria to make cheese and yogurt.
• Other types of microbial fermentation turn – soybeans into soy sauce and – cabbage into sauerkraut.
Lactic acid Fermentation
2 NAD+
2 NADH
2 NAD+
2 NADH
2 Lactate
2 Pyruvate
Glucose
2 ADP
2 ATP
2 P
Gly
coly
sis
Alcohol Fermentation
• The baking and winemaking industries have used alcohol fermentation for thousands of years.
• In this process yeasts (single-celled fungi) – oxidize NADH back to NAD+ and – convert pyruvate to CO2 and ethanol.
Alcohol Fermentation
2 NAD+
2 NADH
2 NAD+
2 NADH
2 Ethanol
2 Pyruvate
Glucose
2 ADP
2 ATP
2 P
Gly
coly
sis
2 CO2
Food
Polysaccharides Fats Proteins
Sugars Glycerol Fatty acids Amino acids
Glycolysis Acetyl CoA
Citric Acid Cycle
Electron Transport
ATP

Figure 6.UN02
cellular work
chemiosmosis
H+ gradient
glucose and organic fuels
Cellular respiration
generates has three stages oxidizes uses
produce some
produces many
to pull electrons down
H+ diffuse through
ATP synthase
pumps H+ to create uses
uses by a process called
energy for
(a)
(b)
(c)
(d)
(e)
(f) (g)
to