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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.

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.

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


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.

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.

6.4 CONNECTION: The human body uses energy from ATP for all its

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.

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.

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?

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.

Energy can be released from glucose by simply burning it.

The energy is dissipated as heat and light and is not available to living

On the other hand, cellular respiration is the controlled breakdown of organic


Energy is gradually released in small amounts, captured by a biological

system, and stored in ATP.

The movement of electrons from one molecule to another is an oxidationreduction 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.

A cellular respiration equation is helpful to show the changes in hydrogen

atom distribution.

Glucose loses its hydrogen atoms and

becomes oxidized to CO2.


gains hydrogen atoms and

becomes reduced to H2O.

Enzymes are necessary to oxidize glucose and other foods.


is an important enzyme in oxidizing glucose,

accepts electrons, and

becomes reduced to NADH.

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


As electrons are transported down the chain, ATP is generated.

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

Stage 1: Glycolysis

occurs in the cytoplasm,

begins cellular respiration, and

breaks down glucose into two molecules of a three-carbon compound

called pyruvate.

Stage 2: The citric acid cycle

takes place in mitochondria,

oxidizes pyruvate to a two-carbon compound, and

supplies the third stage with electrons.

Stage 3: Oxidative phosphorylation

involves electrons carried by NADH and FADH 2,

shuttles these electrons to the electron transport chain embedded in

the inner mitochondrial membrane,

involves chemiosmosis, and

generates ATP through oxidative phosphorylation associated with


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.

ATP is formed in glycolysis by substrate-level phosphorylation during


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.

The steps of glycolysis can be grouped into two main phases.

In steps 14, 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 59, the energy payoff,

two NADH molecules are produced for each initial glucose

molecule and

ATP molecules are generated.

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.

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 CO 2,

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.

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.

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.

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.

6.10 Most ATP production occurs by oxidative phosphorylation

Oxidative phosphorylation

involves electron transport and chemiosmosis and

requires an adequate supply of oxygen.

Electrons from NADH and FADH2 travel down the electron transport chain to

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.

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).

Brown fat is
1. a special type of tissue associated with the generation of heat and
2. more abundant in hibernating mammals and newborn infants.

In brown fat,
1. the cells are packed full of mitochondria,
2. the inner mitochondrial membrane contains an uncoupling protein,
which allows H+ to flow back down its concentration gradient without
generating ATP, and
3. ongoing oxidation of stored fats generates additional heat.

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.

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.

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+.

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.

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.

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.

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.

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.

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.

6.15 Cells use many kinds of organic molecules as fuel for cellular

Although glucose is considered to be the primary source of sugar for

respiration and fermentation, ATP is generated using


fats, and


Fats make excellent cellular fuel because they

contain many hydrogen atoms and thus many energy-rich electrons


yield more than twice as much ATP per gram than a gram of
carbohydrate or protein.

6.16 Food molecules provide raw materials for biosynthesis

Cells use intermediates from cellular respiration for the biosynthesis of other
organic molecules.

Metabolic pathways are often regulated by feedback inhibition in which an

accumulation of product suppresses the process that produces the product.

7.1 Autotrophs are the producers of the biosphere


make their own food through the process of photosynthesis,

sustain themselves, and

do not usually consume organic molecules derived from other


Photoautotrophs use the energy of light to produce organic molecules.

Chemoautotrophs are prokaryotes that use inorganic chemicals as their

energy source.

Heterotrophs are consumers that feed on

plants or

animals, or

decompose organic material.

Photosynthesis in plants

takes place in chloroplasts,

converts carbon dioxide and water into organic molecules, and

releases oxygen.

7.2 Photosynthesis occurs in chloroplasts in plant cells

Chloroplasts are the major sites of photosynthesis in green plants.


is an important light-absorbing pigment in chloroplasts,

is responsible for the green color of plants, and

plays a central role in converting solar energy to chemical energy.

Chloroplasts are concentrated in the cells of the mesophyll, the green tissue
in the interior of the leaf.

Stomata are tiny pores in the leaf that allow

carbon dioxide to enter and

oxygen to exit.

Veins in the leaf deliver water absorbed by roots.

Chloroplasts consist of an envelope of two membranes, which

enclose an inner compartment filled with a thick fluid called stroma


contain a system of interconnected membranous sacs called



are often concentrated in stacks called grana and

have an internal compartment called the thylakoid space, which has

functions analogous to the intermembrane space of a mitochondrion.

Thylakoid membranes also house much of the machinery that converts

light energy to chemical energy.

Chlorophyll molecules

are built into the thylakoid membrane and

capture light energy.

7.3 SCIENTIFIC DISCOVERY: Scientists traced the process of

photosynthesis using isotopes

Scientists have known since the 1800s that plants produce O 2. But does this
oxygen come from carbon dioxide or water?

For many years, it was assumed that oxygen was extracted from CO 2
taken into the plant.

However, later research using a heavy isotope of oxygen, 18O,

confirmed that oxygen produced by photosynthesis comes from H 2O.

Experiment 1: 6 CO2 12 H2O C6H12O6 6 H2O 6 O2

Experiment 2: 6 CO2 12 H2O C6H12O6 6 H2O 6 O2

7.4 Photosynthesis is a redox process, as is cellular respiration

Photosynthesis, like respiration, is a redox (oxidation-reduction) process.

CO2 becomes reduced to sugar as electrons along with hydrogen ions

from water are added to it.

Water molecules are oxidized when they lose electrons along with
hydrogen ions.

Cellular respiration uses redox reactions to harvest the chemical energy

stored in a glucose molecule.

This is accomplished by oxidizing the sugar and reducing O 2 to H2O.

The electrons lose potential as they travel down the electron transport
chain to O2.

In contrast, the food-producing redox reactions of photosynthesis

require energy.

In photosynthesis,

light energy is captured by chlorophyll molecules to boost the energy

of electrons,

light energy is converted to chemical energy, and

chemical energy is stored in the chemical bonds of sugars.

7.5 Overview: The two stages of photosynthesis are linked by ATP and

Photosynthesis occurs in two metabolic stages.

1. The light reactions occur in the thylakoid membranes. In these

water is split, providing a source of electrons and giving off

oxygen as a by-product,

ATP is generated from ADP and a phosphate group, and

light energy is absorbed by the chlorophyll molecules to drive

the transfer of electrons and H+ from water to the electron
acceptor NADP+ reducing it to NADPH.

NADPH produced by the light reactions provides the electrons for

reducing carbon in the Calvin cycle.

2. The second stage is the Calvin cycle, which occurs in the stroma of
the chloroplast.

The Calvin cycle is a cyclic series of reactions that assembles

sugar molecules using CO2 and the energy-rich products of the
light reactions.

During the Calvin cycle, CO2 is incorporated into organic

compounds in a process called carbon fixation.

After carbon fixation, enzymes of the cycle make sugars by

further reducing the carbon compounds.

The Calvin cycle is often called the dark reactions or lightindependent reactions, because none of the steps requires light

7.6 Visible radiation absorbed by pigments drives the light reactions

Sunlight contains energy called electromagnetic energy or electromagnetic


Visible light is only a small part of the electromagnetic spectrum,

the full range of electromagnetic wavelengths.

Electromagnetic energy travels in waves, and the wavelength is the

distance between the crests of two adjacent waves.

Light behaves as discrete packets of energy called photons.

A photon is a fixed quantity of light energy.

The shorter the wavelength, the greater the energy.


absorb light and

are built into the thylakoid membrane.

Plant pigments

absorb some wavelengths of light and

reflect or transmit other wavelengths.

We see the color of the wavelengths that are transmitted. For example,
chlorophyll transmits green wavelengths.

Chloroplasts contain several different pigments, which absorb light of

different wavelengths.

Chlorophyll a absorbs blue-violet and red light and reflects green.

Chlorophyll b absorbs blue and orange and reflects yellow-green.


broaden the spectrum of colors that can drive photosynthesis


provide photoprotection, absorbing and dissipating excessive

light energy that would otherwise damage chlorophyll or interact
with oxygen to form reactive oxidative molecules.

7.7 Photosystems capture solar energy

Pigments in chloroplasts absorb photons (capturing solar power), which

increases the potential energy of the pigments electrons and

sends the electrons into an unstable state.

These unstable electrons

drop back down to their ground state, and as they do,

release their excess energy as heat.

Within a thylakoid membrane, chlorophyll and other pigment molecules

absorb photons and

transfer the energy to other pigment molecules.

In the thylakoid membrane, chlorophyll molecules are organized along with

other pigments and proteins into photosystems.

A photosystem consists of a number of light-harvesting complexes

surrounding a reaction-center complex.

A light-harvesting complex contains various pigment molecules bound to


Collectively, the light-harvesting complexes function as a light-gathering


The light energy is passed from molecule to molecule within the


Finally it reaches the reaction center where a primary electron acceptor

accepts these electrons and consequently becomes reduced.

This solar-powered transfer of an electron from the reaction-center

pigment to the primary electron acceptor is the first step in the
transformation of light energy to chemical energy in the light reactions.

Two types of photosystems (photosystem I and photosystem II) cooperate in

the light reactions.

Each type of photosystem has a characteristic reaction center.

Photosystem II, which functions first, is called P680 because its

pigment absorbs light with a wavelength of 680 nm.

Photosystem I, which functions second, is called P700 because it

absorbs light with a wavelength of 700 nm.

7.8 Two photosystems connected by an electron transport chain generate


In the light reactions, light energy is transformed into the chemical energy of

To accomplish this, electrons are

removed from water,

passed from photosystem II to photosystem I, and

accepted by NADP+, reducing it to NADPH.

Between the two photosystems, the electrons

move down an electron transport chain and

provide energy for the synthesis of ATP.

The products of the light reactions are


ATP, and


7.9 Chemiosmosis powers ATP synthesis in the light reactions

Interestingly, chemiosmosis is the mechanism that

is involved in oxidative phosphorylation in mitochondria and

generates ATP in chloroplasts.

ATP is generated because the electron transport chain produces a

concentration gradient of hydrogen ions across a membrane.

In photophosphorylation, using the initial energy input from light,

the electron transport chain pumps H+ into the thylakoid space, and

the resulting concentration gradient drives H + back through ATP

synthase, producing ATP.

How does photophosphorylation compare with oxidative phosphorylation?

Mitochondria use oxidative phosphorylation to transfer chemical

energy from food into the chemical energy of ATP.

Chloroplasts use photophosphorylation to transfer light energy into the

chemical energy of ATP.

7.10 ATP and NADPH power sugar synthesis in the Calvin cycle

The Calvin cycle makes sugar within a chloroplast.

To produce sugar, the necessary ingredients are

atmospheric CO2 and

ATP and NADPH generated by the light reactions.

The Calvin cycle uses these three ingredients to produce an energy-rich,

three-carbon sugar called glyceraldehyde-3-phosphate (G3P).

A plant cell may then use G3P to make glucose and other organic molecules.

The steps of the Calvin cycle include

carbon fixation,


release of G3P, and

regeneration of the starting molecule ribulose bisphosphate (RuBP).

7.11 EVOLUTION CONNECTION: Other methods of carbon fixation have

evolved in hot, dry climates

Most plants use CO2 directly from the air, and carbon fixation occurs when the
enzyme rubisco adds CO2 to RuBP.

Such plants are called C3 plants because the first product of carbon fixation
is a three-carbon compound, 3-PGA.

In hot and dry weather, C3 plants

close their stomata to reduce water loss but

prevent CO2 from entering the leaf and O2 from leaving.

As O2 builds up in a leaf, rubisco adds O 2 instead of CO2 to RuBP, and a

two-carbon product of this reaction is then broken down in the cell.

This process is called photorespiration because it occurs in the light,

consumes O2, and releases CO2.

But unlike cellular respiration, it uses ATP instead of producing it.

C4 plants have evolved a means of

carbon fixation that saves water during photosynthesis while

optimizing the Calvin cycle.

C4 plants are so named because they first fix CO 2 into a four-carbon


When the weather is hot and dry, C4 plants keep their stomata mostly closed,
thus conserving water.

Another adaptation to hot and dry environments has evolved in the CAM
plants, such as pineapples and cacti.

CAM plants conserve water by opening their stomata and admitting CO 2

only at night.

CO2 is fixed into a four-carbon compound,

which banks CO2 at night and

releases it to the Calvin cycle during the day.

7.12 Review: Photosynthesis uses light energy, carbon dioxide, and water
to make organic molecules

Most of the living world depends on the food-making

machinery of photosynthesis.

The chloroplast

integrates the two stages of photosynthesis and

makes sugar from CO2.

About half of the carbohydrates made by photosynthesis are

consumed as fuel for cellular respiration in the mitochondria of plant

Sugars also serve as the starting material for making other organic
molecules, such as proteins, lipids, and cellulose.

Excess food made by plants is stockpiled as starch in roots, tubers,

seeds, and fruits.

7.13 CONNECTION: Photosynthesis may moderate global climate change

The greenhouse effect operates on a global scale.

Solar radiation includes visible light that penetrates the

Earths atmosphere and warms the planets surface.

Heat radiating from the warmed planet is absorbed by gases in

the atmosphere, which then reflects some of the heat back to

Without the warming of the greenhouse effect, the Earth would

be much colder and most life as we know it could not exist.

The gases in the atmosphere that absorb heat radiation are called
greenhouse gases. These include

water vapor,

carbon dioxide, and


Increasing concentrations of greenhouse gases have been linked to

global climate change (also called global warming), a slow but
steady rise in Earths surface temperature.

Since 1850, the atmospheric concentration of CO2 has increased by

about 40%, mostly due to the combustion of fossil fuels including


oil, and


The predicted consequences of continued warming include

melting of polar ice,

rising sea levels,

extreme weather patterns,


increased extinction rates, and

the spread of tropical diseases.

Widespread deforestation has aggravated the global warming

problem by reducing an effective CO2 sink.

Global warming caused by increasing CO2 levels may be reduced by

limiting deforestation,

reducing fossil fuel consumption, and

growing biofuel crops that remove CO2 from the atmosphere.

7.14 SCIENTIFIC DISCOVERY: Scientific study of Earths ozone layer has

global significance

Solar radiation converts O2 high in the atmosphere to ozone (O3),

which shields organisms from damaging UV radiation.

Industrial chemicals called CFCs have caused dangerous thinning of

the ozone layer, but international restrictions on CFC use are
allowing a slow recovery.