chapter 6 how cells harvest chemical energy...
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© 2012 Pearson Education, Inc. Lecture by Edward J. Zalisko
PowerPoint Lectures for Campbell Biology: Concepts & Connections, Seventh Edition Reece, Taylor, Simon, and Dickey
Chapter 6 How Cells Harvest Chemical Energy Introduction
In eukaryotes, cellular respiration
– harvests energy from food,
– yields large amounts of ATP, and
– Uses ATP to drive cellular work.
A similar process takes place in many prokaryotic organisms.
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Figure 6.0_1 Chapter 6: Big Ideas
Cellular Respiration: Aerobic Harvesting
of Energy
Stages of Cellular Respiration
Fermentation: Anaerobic Harvesting of Energy
Connections Between Metabolic Pathways
CELLULAR RESPIRATION: AEROBIC HARVESTING
OF ENERGY
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6.1 Photosynthesis and cellular respiration provide energy for life
Life requires energy.
In almost all ecosystems, energy ultimately comes from the sun.
In photosynthesis,
– some of the energy in sunlight is captured by chloroplasts,
– atoms of carbon dioxide and water are rearranged, and
– glucose and oxygen are produced.
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6.1 Photosynthesis and cellular respiration provide energy for life
In cellular respiration
– glucose is broken down to carbon dioxide and water and
– the cell captures some of the released energy to make ATP.
Cellular respiration takes place in the mitochondria of eukaryotic cells.
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Figure 6.1
Sunlight energy
ECOSYSTEM
Photosynthesis in chloroplasts
Cellular respiration in mitochondria
(for cellular work)
Heat energy
Glucose CO2
H2O O2
ATP
Figure 6.1_1
Sunlight energy
ECOSYSTEM
Photosynthesis in chloroplasts
Cellular respiration in mitochondria
(for cellular work)
Heat energy
Glucose CO2
H2O O2
ATP
6.2 Breathing supplies O2 for use in cellular respiration and removes CO2
Respiration, as it relates to breathing, and cellular respiration are not the same.
– Respiration, in the breathing sense, refers to an exchange of gases. Usually an organism brings in oxygen from the environment and releases waste CO2.
– Cellular respiration is the aerobic (oxygen requiring) harvesting of energy from food molecules by cells.
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Figure 6.2
Breathing
Lungs
Bloodstream CO2 O2
O2 CO2
Muscle cells carrying out Cellular Respiration
Glucose + O2 CO2 + H2O + ATP
6.3 Cellular respiration banks energy in ATP molecules
Cellular respiration is an exergonic process that transfers energy from the bonds in glucose to form ATP.
Cellular respiration
– produces up to 32 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.
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Figure 6.3
Glucose Oxygen Water Carbon dioxide
C6H12O6 O2 H2O ATP 6 6 6
+ Heat
CO2
3
6.4 CONNECTION: The human body uses energy from ATP for all its activities
The average adult human needs about 2,200 kcal of energy per day.
– About 75% of these calories are used to maintain a healthy body.
– The remaining 25% is used to power physical activities.
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6.4 CONNECTION: The human body uses energy from ATP for all its activities
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.
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Figure 6.4
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
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6.5 Cells tap energy from electrons “falling” from organic fuels to oxygen
The energy necessary for life is contained in the arrangement of electrons in chemical bonds in organic molecules.
An important question is how do cells extract this energy?
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6.5 Cells tap energy from electrons “falling” from organic fuels to oxygen
When the carbon-hydrogen bonds of glucose are broken, electrons are transferred to oxygen.
– Oxygen has a strong tendency to attract electrons.
– An electron loses potential energy when it “falls” to oxygen.
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6.5 Cells tap energy from electrons “falling” from organic fuels to oxygen
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.
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6.5 Cells tap energy from electrons “falling” from organic fuels to oxygen
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.
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6.5 Cells tap energy from electrons “falling” from organic fuels to oxygen
The 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.
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6.5 Cells tap energy from electrons “falling” from organic fuels to oxygen
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.
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Figure 6.5A
Glucose + Heat
C6H12O6 O2 CO2 H2O ATP 6 6 6
Loss of hydrogen atoms (becomes oxidized)
Gain of hydrogen atoms (becomes reduced)
6.5 Cells tap energy from electrons “falling” from organic fuels to oxygen
Enzymes are necessary to oxidize glucose and other foods.
NAD+
– is an important enzyme in oxidizing glucose,
– accepts electrons, and
– becomes reduced to NADH.
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Figure 6.5B
Becomes oxidized
Becomes reduced
2H
2H NAD+ NADH
H+
H+
2 2 (carries
2 electrons)
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6.5 Cells tap energy from electrons “falling” from organic fuels to oxygen
There are other electron “carrier” molecules that function like NAD+.
– They form a staircase where the electrons pass from one to the next down the staircase.
– These electron carriers collectively are called the electron transport chain.
– As electrons are transported down the chain, ATP is generated.
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Figure 6.5C
Controlled release of energy for synthesis of ATP
NADH
NAD+
H+
H+ O2
H2O
2
2
2
ATP
Electron transport chain
2 1
STAGES OF CELLULAR RESPIRATION
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6.6 Overview: Cellular respiration occurs in three main stages
Cellular respiration consists of a sequence of steps that can be divided into three stages.
– Stage 1 – Glycolysis
– Stage 2 – Pyruvate oxidation and citric acid cycle
– Stage 3 – Oxidative phosphorylation
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6.6 Overview: Cellular respiration occurs in three main stages
Stage 1: Glycolysis
– occurs in the cytoplasm,
– begins cellular respiration, and
– breaks down glucose into two molecules of a three-carbon compound called pyruvate.
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6.6 Overview: Cellular respiration occurs in three main stages
Stage 2: The citric acid cycle
– takes place in mitochondria,
– oxidizes pyruvate to a two-carbon compound, and
– supplies the third stage with electrons.
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6.6 Overview: Cellular respiration occurs in three main stages
Stage 3: Oxidative phosphorylation
– involves electrons carried by NADH and FADH2,
– shuttles these electrons to the electron transport chain embedded in the inner mitochondrial membrane,
– involves chemiosmosis, and
– generates ATP through oxidative phosphorylation associated with chemiosmosis.
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Figure 6.6
NADH
NADH FADH2
ATP ATP ATP
CYTOPLASM
Glycolysis
Electrons carried by NADH
Glucose Pyruvate Pyruvate Oxidation
Citric Acid Cycle
Oxidative Phosphorylation
(electron transport and chemiosmosis)
Mitochondrion
Substrate-level phosphorylation
Substrate-level phosphorylation
Oxidative phosphorylation
Figure 6.6_1
NADH
NADH FADH2
ATP ATP ATP
CYTOPLASM
Glycolysis
Electrons carried by NADH
Glucose Pyruvate Pyruvate Oxidation
Citric Acid Cycle
Oxidative Phosphorylation
(electron transport and chemiosmosis)
Mitochondrion
Substrate-level phosphorylation
Substrate-level phosphorylation
Oxidative phosphorylation
6.7 Glycolysis harvests chemical energy by oxidizing glucose to pyruvate
In glycolysis,
– a single molecule of glucose is enzymatically cut in half through a series of steps,
– two molecules of pyruvate are produced,
– two molecules of NAD+ are reduced to two molecules of NADH, and
– a net of two molecules of ATP is produced.
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Figure 6.7A
Glucose
2 Pyruvate
2 ADP
2 P 2 NAD+
2 NADH
2 H+ ATP 2
6.7 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.
The compounds that form between the initial reactant, glucose, and the final product, pyruvate, are called intermediates.
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Figure 6.7B
Enzyme Enzyme
P
P
ADP
P
ATP
Substrate Product
6.7 Glycolysis harvests chemical energy by oxidizing glucose to pyruvate
The steps of glycolysis can be grouped into two main phases.
– In steps 1–4, the energy investment phase,
– energy is consumed as two ATP molecules are used to energize a glucose molecule,
– which is then split into two small sugars that are now primed to release energy.
– In steps 5–9, the energy payoff,
– two NADH molecules are produced for each initial glucose molecule and
– ATP molecules are generated.
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Figure 6.7Ca_s1 Glucose
Glucose 6-phosphate
Fructose 6-phosphate
Fructose 1,6-bisphosphate
ENERGY INVESTMENT
PHASE
P P
P
P
ADP
ADP
ATP
ATP
Step Steps – A fuel molecule is energized, using ATP.
3
3
2
1
1
Figure 6.7Ca_s2 Glucose
Glucose 6-phosphate
Fructose 6-phosphate
Fructose 1,6-bisphosphate
Glyceraldehyde 3-phosphate (G3P)
ENERGY INVESTMENT
PHASE
P P
P P
P
P
ADP
ADP
ATP
ATP
Step Steps – A fuel molecule is energized, using ATP.
Step A six-carbon intermediate splits into two three-carbon intermediates.
4
4
3
3
2
1
1
Figure 6.7Cb_s1
5 5 5 Step A redox reaction generates NADH.
ENERGY PAYOFF PHASE
1,3-Bisphospho- glycerate
NADH NADH
NAD+ NAD+
H+ H+
P P
P
P P
P
P P
Figure 6.7Cb_s2
6 6 6
5 5 5
9
9 9
8 8
7 7
Step A redox reaction generates NADH.
Steps – ATP and pyruvate are produced.
ENERGY PAYOFF PHASE
1,3-Bisphospho- glycerate
3-Phospho- glycerate
2-Phospho- glycerate
Phosphoenol- pyruvate (PEP)
Pyruvate
NADH NADH
NAD+ NAD+
H+ H+
ADP ADP
ADP ADP
ATP ATP
ATP ATP
H2O H2O
P P
P
P P
P
P
P
P
P P
P
P
P
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6.8 Pyruvate is oxidized prior to the citric acid cycle
The pyruvate formed in glycolysis is transported from the cytoplasm into a mitochondrion where
– the citric acid cycle and
– oxidative phosphorylation will occur.
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6.8 Pyruvate is oxidized prior to the citric acid cycle
Two molecules of pyruvate are produced for each molecule of glucose that enters glycolysis.
Pyruvate does not enter the citric acid cycle, but undergoes some chemical grooming in which
– a carboxyl group is removed and given off as CO2,
– the two-carbon compound remaining is oxidized while a molecule of NAD+ is reduced to NADH,
– coenzyme A joins with the two-carbon group to form acetyl coenzyme A, abbreviated as acetyl CoA, and
– acetyl CoA enters the citric acid cycle. © 2012 Pearson Education, Inc.
Figure 6.8
Pyruvate
Coenzyme A
Acetyl coenzyme A
NAD+ NADH H+
CoA
CO2 3
2
1
6.9 The citric acid cycle completes the oxidation of organic molecules, generating many NADH and FADH2 molecules
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),
– completes the oxidation of organic molecules, and
– generates many NADH and FADH2 molecules.
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Figure 6.9A Acetyl CoA
Citric Acid Cycle
CoA CoA
CO2 2
3
3
NAD+
3 H+
NADH
ADP ATP P
FAD
FADH2
6.9 The citric acid cycle completes the oxidation of organic molecules, generating many NADH and FADH2 molecules
During the citric acid cycle
– the two-carbon group of acetyl CoA is added to a four-carbon compound, forming citrate,
– citrate is degraded back to the four-carbon compound,
– two CO2 are released, and
– 1 ATP, 3 NADH, and 1 FADH2 are produced.
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6.9 The citric acid cycle completes the oxidation of organic molecules, generating many NADH and FADH2 molecules
Remember that the 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.
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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.
6.10 Most ATP production occurs by oxidative phosphorylation
Oxidative phosphorylation
– involves electron transport and chemiosmosis and
– requires an adequate supply of oxygen.
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6.10 Most ATP production occurs by oxidative phosphorylation
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.
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Figure 6.10
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
Figure 6.10_1
H+
H+
H+
H+ H+
H+
H+ H+
H+
H+
H+
Oxidative Phosphorylation
Electron Transport Chain Chemiosmosis
I
II
III IV
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
6.11 CONNECTION: 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).
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Figure 6.11
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
6.11 CONNECTION: Interrupting cellular respiration can have both harmful and beneficial effects
Brown fat is – a special type of tissue associated with the generation
of heat and
– more abundant in hibernating mammals and newborn infants.
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6.11 CONNECTION: Interrupting cellular respiration can have both harmful and beneficial effects
In brown fat, – the cells are packed full of mitochondria,
– the inner mitochondrial membrane contains an uncoupling protein, which allows H+ to flow back down its concentration gradient without generating ATP, and
– ongoing oxidation of stored fats generates additional heat.
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6.12 Review: Each molecule of glucose yields many molecules of ATP
Recall that the energy payoff of cellular respiration involves
1. glycolysis,
2. alteration of pyruvate,
3. the citric acid cycle, and
4. oxidative phosphorylation.
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6.12 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.
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Figure 6.12
NADH
FADH2
NADH FADH2 NADH
or NADH
Mitochondrion CYTOPLASM
Electron shuttles across membrane
Glycolysis Glucose
2 Pyruvate
Pyruvate Oxidation 2 Acetyl
CoA
Citric Acid Cycle
Oxidative Phosphorylation
(electron transport and chemiosmosis)
Maximum per glucose:
by substrate-level phosphorylation
by substrate-level phosphorylation
by oxidative phosphorylation
2 2
2
2
6 2
ATP + 2 about +
28 ATP About ATP 32
ATP + 2
FERMENTATION: ANAEROBIC HARVESTING OF ENERGY
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6.13 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+.
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6.13 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.
© 2012 Pearson Education, Inc. Animation: Fermentation Overview
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6.13 Fermentation enables cells to produce 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.
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Figure 6.13A
2 NAD+
2 NADH
2 NAD+
2 NADH
2 Lactate
2 Pyruvate
Glucose
2 ADP
2 ATP
2 P
Gly
coly
sis
6.13 Fermentation enables cells to produce ATP without oxygen
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.
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Figure 6.13B
2 NAD+
2 NADH
2 NAD+
2 NADH
2 Ethanol
2 Pyruvate
Glucose
2 ADP
2 ATP
2 P
Gly
coly
sis
2 CO2
6.13 Fermentation enables cells to produce ATP without oxygen
Obligate anaerobes
– are poisoned by oxygen, requiring anaerobic conditions, and
– live in stagnant ponds and deep soils.
Facultative anaerobes
– include yeasts and many bacteria, and
– can make ATP by fermentation or oxidative phosphorylation.
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6.14 EVOLUTION CONNECTION: Glycolysis evolved early in the history of life on Earth
Glycolysis is the universal energy-harvesting process of life.
The role of glycolysis in fermentation and respiration dates back to
– life long before oxygen was present,
– when only prokaryotes inhabited the Earth,
– about 3.5 billion years ago.
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6.14 EVOLUTION CONNECTION: Glycolysis evolved early in the history of life on Earth
The ancient history of glycolysis is supported by its
– occurrence in all the domains of life and
– location within the cell, using pathways that do not involve any membrane-bounded organelles.
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CONNECTIONS BETWEEN METABOLIC PATHWAYS
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6.15 Cells use many kinds of organic molecules as fuel for cellular respiration
Although glucose is considered to be the primary source of sugar for respiration and fermentation, ATP is generated using
– carbohydrates,
– fats, and
– proteins.
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6.15 Cells use many kinds of organic molecules as fuel for cellular respiration
Fats make excellent cellular fuel because they
– contain many hydrogen atoms and thus many energy-rich electrons and
– yield more than twice as much ATP per gram than a gram of carbohydrate or protein.
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Figure 6.15
Food, such as peanuts
Sugars Glycerol Fatty acids Amino acids
Amino groups
Oxidative Phosphorylation
Citric Acid
Cycle Pyruvate Oxidation
Acetyl CoA
ATP
Glucose G3P Pyruvate Glycolysis
Carbohydrates Fats Proteins
6.16 Food molecules provide raw materials for biosynthesis
Cells use intermediates from cellular respiration for the biosynthesis of other organic molecules.
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Figure 6.16
Carbohydrates Fats Proteins
Cells, tissues, organisms
Amino acids Fatty acids Glycerol Sugars
Amino groups
Citric Acid
Cycle
Pyruvate Oxidation
Acetyl CoA
ATP needed to drive biosynthesis
ATP
Glucose Synthesis Pyruvate G3P Glucose
6.16 Food molecules provide raw materials for biosynthesis
Metabolic pathways are often regulated by feedback inhibition in which an accumulation of product suppresses the process that produces the product.
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