7 Photosynthesis: Using Light

to Make Food
Why Photosynthesis Matters

If you want �

to reduce the
rate of global
climate change,
plant a tree.

� Nearly all life on

Earth—including
you—can trace its
source of energy
back to the sun.

Protecting yourself �

from short
wavelengths of light
can be lifesaving.

140
 CHAPTER CONTENTS CHAPTER THREAD
The Basics of Photosynthesis 142 Biofuels
The Light Reactions: Converting Solar Energy to Chemical BIOLOGY AND SOCIETY A Greasy Crime Wave 141
Energy 144
THE PROCESS OF SCIENCE What Colors of Light Drive
The Calvin Cycle: Making Sugar from Carbon Dioxide 149 Photosynthesis? 145
EVOLUTION CONNECTION Creating a Better Biofuel
Factory 149

Biofuels BIOLOGY AND SOCIETY

A Greasy Crime Wave

In September 2013, police in Ocala, Florida, arrested two men and charged them with organized fraud and
grand theft. Their crime? The men were caught red-handed with more than 700 gallons of stolen used cook-
ing oil pilfered from a variety of local eateries. Why would anyone steal that nasty stuff? The reason is sim-
ple: Remnants of restaurant deep fryers, sometimes called “liquid gold,” fetch about $2 per pound when sold
to recyclers. That makes the burglars’ haul worth more than $5,000. Why is grease so valuable?
As fossil fuel supplies dwindle and prices rise, the need for reliable, renewable sources
of energy increases. In response, scientists are researching better ways to harness biofuels,
energy obtained from living material. Some researchers focus on burning plant matter
directly (wood pellet boilers, for example), and others focus on using plant material to
produce biofuels that can be burned.
There are several types of biofuels. Bioethanol is a type of alcohol (the same kind found
in alcoholic drinks) that is made from wheat, corn, sugar beets, and other food crops. Starch
made naturally by plants is converted to glucose and then fermented to ethanol by micro-
organisms such as single-celled algae. Bioethanol can be used directly as a fuel source in
specially designed vehicles, but it is more commonly used as a gasoline additive that can
increase fuel efficiency while decreasing vehicle emissions. You may have noticed a sticker
on a gas pump that declares the percentage of ethanol in that gasoline; most cars today run
on a blend of 85% gasoline and 15% ethanol. Many car manufacturers are producing “flex-
ible-fuel” vehicles that can run on any combination of gasoline and bioethanol. Although
bioethanol does reduce carbon emissions and is a renewable resource, its production raises
the prices of food crops (which become more expensive as acreage is diverted to biofuel
production).
Cellulosic ethanol is a form of bioethanol made from cellulose found in nonedible plant
material such as wood, grass, or scraps from crops. Biodiesel, the most common biofuel in
Europe, is made from plant oils such as recycled frying oil. Like bioethanol, it can be used on
its own or as an emissions-reducing additive to standard diesel. In a strange twist, rising Using biofuels. Biofuels are
values for diesel have sparked a greasy crime wave as thieves tap into this new and largely added to most gasoline in the
unguarded source of raw material. Today, only about 2.7% of the world’s fuel used for driving United States.
is provided by biofuels, but the International Energy Agency has set a goal of 25% by 2050.
When we derive energy from biofuels, we are actually tapping into the energy of the sun, which drives
photosynthesis in plants. Photosynthesis is the process by which plants use light to make sugars from carbon
dioxide—sugars that are food for the plant and the starting point for most of our own food. In this chap-
ter, we’ll first examine some basic concepts of photosynthesis; then we’ll look at the specific mechanisms
involved in this process.

141
 The Basics of Photosynthesis
CHAPTER 7
PHOTOSYNTHESIS: USING
LIGHT TO MAKE FOOD

The process of photosynthesis is the ultimate source chloroplasts (about 500,000 per square millimeter of
of energy for nearly every ecosystem on Earth. leaf surface—that’s equivalent to about 300 million chlo-
Photosynthesis is a process whereby plants, algae roplasts in a leaf the size of a standard postage stamp).
(which are protists), and certain bacteria trans- Their green color is from chlorophyll, a pigment (light-
Nearly all life on Earth— form light energy into chemical energy, using absorbing molecule) in the chloroplasts that plays a cen-
including you—can trace its carbon dioxide and water as starting materials tral role in converting solar energy to chemical energy.
source of energy back and releasing oxygen gas as a by-product. The Chloroplasts are concentrated in the interior cells of
to the sun. chemical energy produced via photosynthe- leaves (Figure 7.2), with a typical cell containing
sis is stored in the bonds of sugar molecules. 30–40!chloroplasts. Carbon dioxide (CO2) enters a
Organisms that generate their own organic matter leaf, and oxygen (O2) exits, by way of tiny pores called
from inorganic ingredients are called autotrophs (see stomata (singular, stoma, meaning “mouth”). The carbon
Chapter!6). Plants and other organisms that do this by dioxide that enters the leaf is the source of carbon for
photosynthesis—photoautotrophs—are the producers much of the body of the plant, including the sugars and
for most ecosystems (Figure 7.1). Not only do photo- starches that we eat. So the bulk of the body of a plant
autotrophs feed us, they also clothe us (as the source of derives from the air, not the soil. As proof of this idea,
cotton fibers), house us (wood), and provide energy for consider hydroponics, a means of growing plants using
warmth, light, and transportation (biofuels). only air and water; no soil whatsoever is involved. In
addition to carbon dioxide, photosynthesis requires water,
which is absorbed by the plant’s roots and transported to
Chloroplasts: Sites the leaves, where veins carry it to the photosynthetic cells.
Membranes within the chloroplast form the frame-
of Photosynthesis work where many of the reactions of photosynthe-
Photosynthesis in plants and algae occurs within light- sis occur. Like a mitochondrion, a chloroplast has a
absorbing organelles called chloroplasts (see Chapter!4, double-membrane envelope. The chloroplast’s inner
especially Figure 4.17). All green parts of a plant have membrane encloses a compartment filled with stroma,
chloroplasts and thus can carry out photosynthesis. a thick fluid. (It’s easy to confuse two terms associated
#Figure 7.1 A diversity of In most plants, however, the leaves have the most with photosynthesis: stomata are pores through which
photoautotrophs.

PHOTOSYNTHETIC AUTOTROPHS

Plants Photosynthetic Protists Photosynthetic Bacteria

(mostly on land) (aquatic) (aquatic)

LM 375×

Forest plants Kelp, a large, multicellular alga Micrograph of cyanobacteria

142
 gases are exchanged, and stroma is the fluid THE BASICS
within the chloroplast.) Suspended in the stroma OF PHOTOSYNTHESIS
are interconnected membranous sacs called
thylakoids. The thylakoids are concentrated
in stacks called grana (singular, granum). Inner and outer
membranes
The chlorophyll molecules that capture light
energy are built into the thylakoid mem-
branes. The structure of a chloroplast—

LM 2,000×
with its stacks of disks—aids its function
by providing a large surface area
for the reactions of Photosynthetic cells Vein
photosynthesis. (transports water
and nutrients) Stroma Thylakoid
Granum
Thylakoid
Interior cell space
Chloroplast

CHECKPOINT
Photosynthesis takes place
� Figure 7.2 Journey
within organelles called
into a leaf. This series of

Colorized TEM 1,000×

Stomata _________ using gases that
blowups takes you into a O2
leaf’s interior, then into a CO2 are exchanged via pores
plant cell, and finally into called _________.
a chloroplast, the site of Leaf cross section
Answer: chloroplasts; stomata

photosynthesis.
Energy Transformations

Energy
Within biological systems, information
stored in DNA is transmitted and
All biological systems depend on
obtaining, converting, and releasing
energy and matter.
Transformations
All biological systems, from
molecules to ecosystems, depend
on interactions between
An Overview of Photosynthesis
The following chemical equation, simplified to highlight sugar. Hydrogen is moved along with the electrons being
the relationship between photosynthesis and cellular transferred from water to carbon dioxide. This transfer # Figure 7.3 A road map
for photosynthesis. We’ll
respiration, provides a summary of the reactants and of hydrogen requires the chloroplast to split water mol-
use a smaller version of this
products of photosynthesis: ecules into hydrogen and oxygen. The hydrogen is trans- road map for orientation as we
ferred along with electrons to carbon dioxide to form take a closer look at the light
Light energy sugar. The oxygen escapes through stomata in leaves into reactions and the Calvin cycle.
the atmosphere as O2, a waste prod-
6 CO2 + 6 H 2O C 6H12O 6 + 6 O2
Photo- uct of photosynthesis. H 2O CO2
Chloroplast
Carbon Water synthesis Glucose Oxygen gas The overall equation
dioxide Light
Notice that the reactants of photosynthesis—carbon for photosynthesis is a
dioxide (CO2) and water (H2O)—are the same as the simple summary of a com-
waste products of cellular respiration (see Figure 6.2). plex process. Like many NADP+
Also notice that photosynthesis produces what respira- energy-producing processes
ADP
tion uses—glucose (C6H12O6) and oxygen (O2). In other within cells, photosynthesis is + P
words, photosynthesis recycles the “exhaust” of cellular a multistep chemical pathway, Light Calvin
cycle
respiration and rearranges its atoms to produce food with each step in the path pro- reactions

and oxygen. Photosynthesis is a chemical transforma- ducing products that are used as
ATP
tion that requires a lot of energy, and sunlight absorbed reactants in the next step. This is
by chlorophyll provides that energy. a clear example of one of biology’s – –
NADPH
Recall that cellular respiration is a process of electron major themes: the use of metabolic
transfer (see Chapter 6). A “fall” of electrons from food pathways to obtain, process, and
molecules to oxygen to form water releases the energy store energy. To help get a better over-
that mitochondria can use to make ATP (see Figure!6.9). view, let’s take a look at the two stages of O2
Sugar
The opposite occurs in photosynthesis: Electrons are photosynthesis: the light reactions and
boosted “uphill” and added to carbon dioxide to produce the Calvin cycle (Figure 7.3).

143
CHAPTER 7 In the light reactions, chlorophyll in the thylakoid Calvin cycle indirectly depends on light to produce
PHOTOSYNTHESIS: USING membranes absorbs solar energy (the “photo” part of sugar because it requires the supply of ATP!and
LIGHT TO MAKE FOOD
photosynthesis), which is then converted to the chemi- NADPH produced by the light reactions.
cal energy of ATP (the!molecule that drives most cel- The initial incorporation of carbon from CO2
lular work) and NADPH (an electron carrier). During into!organic compounds is called carbon fixation.
the light reactions, water is split, providing a source of This!process has important implications for global
electrons and giving off O2 gas as a by-product. climate, because the removal of carbon from the
CHECKPOINT The Calvin cycle uses the products of the air and its incorporation into plant material
light reactions to power the production of If you want to can help reduce the concentration of carbon
1. What molecules are the
inputs of photosynthesis? sugar from carbon dioxide (the “synthesis” reduce the rate dioxide in the atmosphere. Deforestation,
What molecules are the part of photosynthesis). The enzymes that of global climate which removes a lot of photosynthetic
outputs? drive the Calvin cycle are dissolved in the change, plant plant life, thereby reduces the ability of the
2. Name the two stages of stroma. ATP generated by the light reactions a tree. biosphere to absorb carbon. Planting new
photosynthesis in their provides the energy for sugar synthesis. And forests can have the opposite effect of fixing
proper order.
and O2 2. light reactions, Calvin cycle the NADPH produced by the light reactions carbon from the atmosphere, potentially reduc-
Answers: 1. CO2 and H2O; glucose provides the high-energy electrons that drive the ing the effect of the gases that contribute to global
synthesis of glucose from carbon dioxide. Thus, the climate change.

The Light Reactions: Converting Solar

Energy to Chemical Energy
Chloroplasts are solar-powered CO2
Increasing wavelength
Light H2O
sugar factories. Let’s look at
10–5 nm 10–3 nm 1 nm 103 nm 106 nm 1m 103 m
how they convert sunlight to NADP+
ADP
chemical energy. Light
+ P Calvin Gamma X-rays UV Infrared Micro- Radio
reactions cycle rays waves waves
ATP
Highest Lowest
– – energy energy
The Nature of NADPH

Sunlight O2
Sugar
Visible light

Sunlight is a type of energy called

radiation or electromagnetic energy. Electromagnetic 380 400 500 600 700 750

energy travels through space as rhythmic waves, like Wavelength (nm)

the ripples made by a pebble dropped into a pond.

The distance between the crests of two adjacent waves
is called a wavelength. The full range of radiation,
from the very short wavelengths of gamma rays to the Wavelength =
very long wavelengths of radio signals, is called the 580
nm
electromagnetic spectrum (Figure 7.4). Visible light is
the fraction of the spectrum that our eyes see as differ- � Figure 7.4 The electromagnetic spectrum. The middle of
ent colors. the figure expands the thin slice of the spectrum that is visible
When sunlight shines on a pigmented material, to us as different colors of light, from about 380 nanometers
certain wavelengths (colors) of the visible light are ab- (nm) to about 750 nm in wavelength. The bottom of the figure
shows electromagnetic waves of one particular wavelength of
sorbed and disappear from the light that is reflected by visible light.
the material. For example, we see a pair of jeans as blue
because pigments in the fabric absorb the other colors, botanists (biologists who study plants) discovered that
leaving only light in the blue part of the spectrum to only certain wavelengths of light are used by plants, as
be reflected from the fabric to our eyes. In the 1800s, we’ll see next.

144
 THE LIGHT REACTIONS:
Biofuels THE PROCESS OF SCIENCE CONVERTING SOLAR
ENERGY TO CHEMICAL
ENERGY

What Colors of Light Drive Variations of this classic experiment are still per-
formed today. For example, biofuel researchers test dif-
Photosynthesis? ferent species of algae to determine which wavelengths of
light result in optimal fuel production. Biofuel facilities of
In 1883, German biologist Theodor Engelmann made
the future may use a variety of species that take advantage
the observation that certain bacteria living in water
of the full spectrum of light that shines down on them.
tend to cluster in areas with higher oxygen concentra-
tions. He already knew that light passed through Light
� Figure 7.5 Investigating how light

a prism would separate into the different wavelengths wavelength affects photosynthesis.
Prism
When algal cells are placed on a microscope
(colors). Engelmann soon began to question whether
slide, oxygen-seeking bacteria migrate toward
he could use this information to determine which wave- algae exposed to certain colors of light. These
lengths of light work best for photosynthesis. results suggest that blue-violet and orange-red
Microscope slide
Engelmann’s hypothesis was that oxygen-seeking bac- wavelengths best drive photosynthesis, while
green wavelengths do so only a little bit.
teria would congregate near regions of algae performing
the most photosynthesis (and hence producing the most
oxygen). Engelmann began his experiment by laying a
string of freshwater algal cells within a drop of water on a
microscope slide. He then added oxygen-sensitive bacte-
ria to the drop. Next, using a prism, he created a spectrum
Number of bacteria

of light and shined it on the slide. His results, summa-

Bacteria Bacteria
rized in Figure 7.5, showed that most bacteria congre-
gated around algae exposed to red-orange and blue-violet
light, with very few bacteria moving to the area of green
light. Other experiments have since verified that chlo- Algal cells

roplasts absorb light mainly in the blue-violet and red-

orange part of the spectrum and that those wavelengths of 400 500 600 700
light are the ones mainly responsible for photosynthesis. Wavelength of light (nm)

Chloroplast Pigments
Light
The selective absorption of light by leaves
explains why they appear green to us;
Reflected
light of that color is poorly light
absorbed by chloroplasts Chloroplast
and is thus reflected or trans-
mitted toward the observer
(Figure 7.6). Energy cannot
be destroyed, so the absorbed
energy must be converted to
other forms. Chloroplasts
contain several different
pigments that absorb light
of different wavelengths. Absorbed
light

� Figure 7.6 Why are leaves green?

Transmitted
Chlorophyll and other pigments in light (detected
chloroplasts reflect or transmit green light by your eye)
while absorbing other colors.

145
CHAPTER 7 Chlorophyll a, the pigment that participates directly in When a pigment molecule absorbs a photon, one of
PHOTOSYNTHESIS: USING the light reactions, absorbs mainly blue-violet and red the pigment’s electrons gains energy. This electron is
LIGHT TO MAKE FOOD
light. A very similar molecule, chlorophyll!b, absorbs now said to be “excited”; that is, the electron has been
mainly blue and orange light. Chlorophyll b does not raised from its starting state (called the ground state)
participate directly in the light reactions, but it conveys to an excited state. The excited state is highly unstable,
absorbed energy to chlo- so an excited electron usually loses its excess energy
rophyll a, which then puts and falls back to its ground state almost immediately
the energy to work in the (Figure 7.8a). Most pigments release heat energy as
light reactions. their light-excited electrons fall back to their ground
Chloroplasts also con- state. (That’s why a surface with a lot of pigment, such
tain a family of yellow- as a black driveway, gets so hot on a sunny day.) But
orange pigments called some pigments emit light as well as heat after absorbing
carotenoids, which absorb
mainly blue-green light.
Some carotenoids have a Excited state
protective function: They
The electron falls to its
dissipate excess light en- Absorption of a
e– ground state, releasing
photon excites
ergy that would otherwise an electron.
heat and light.
damage chlorophyll. Some
carotenoids are human
Heat
nutrients: beta-carotene
� Figure 7.7 Photosynthetic (a bright orange/red pigment found in pumpkins, sweet
pigments. Falling autumn Light
potatoes, and carrots) is converted to vitamin A in the
temperatures cause a
decrease in the levels of body, and lycopene (a bright red pigment found in to-
Light
green chlorophyll within the matoes, watermelon, and red peppers) is an antioxidant (fluorescence)
foliage of leaf-bearing trees. that is being studied for potential anti-cancer properties.
This decrease allows the colors Photon
Additionally, the spectacular colors of fall foliage in some Ground
of the carotenoids to be seen. Chlorophyll state
parts of the world are due partly to the yellow-orange
molecule
light reflected from carotenoids (Figure 7.7). The decreas-
ing temperatures in autumn cause a decrease in the levels
CHECKPOINT (a) Absorption of a photon
of chlorophyll, allowing the colors of the longer-lasting
What is the specific name
carotenoids to be seen in all their fall glory.
of the pigment that absorbs
energy during the light All of these chloroplast pigments are built into the
reactions? thylakoid membranes (see Figure 7.2). There the pig-
Answer: chlorophyll a ments are organized into light-harvesting complexes
called photosystems, our next topic.

How Photosystems Harvest

Light Energy
Thinking about light as waves explains most of light’s
properties. However, light also behaves as discrete pack-
ets of energy called photons. A photon is a fixed quan-
tity of light energy. The shorter the wavelength of light,
the greater the energy of a photon. A photon of
violet light, for example, packs nearly twice as
Protecting yourself from much energy as a photon of red light. This is
short wavelengths of light
why short-wavelength light—such as ultravio-
can be lifesaving. (b) Fluorescence of a glow stick. Breaking a vial within a glow
let light and X-rays—can be damaging; pho- stick starts a chemical reaction that excites electrons within a
tons at these wavelengths carry enough energy fluorescent dye. As the electrons fall from their excited state to the
ground state, the excess energy is emitted as light.
to damage proteins and DNA, potentially leading to
cancerous mutations. � Figure 7.8 Excited electrons in pigments.

146
 THE LIGHT REACTIONS:
Chloroplast CONVERTING SOLAR
Cluster of pigment ENERGY TO CHEMICAL
molecules associated Photon ENERGY
with membrane proteins

Primary
electron
acceptor
e– Electron
Reaction
transfer Reaction- center
center
chlorophyll a

Pigment
� Figure 7.9 A photosystem: molecules
light-gathering molecules
that focus light energy onto Thylakoid membrane
Transfer of energy
a reaction center. Photosystem

photons. The fluorescent light emitted by a glow stick molecules functions as a light-gathering antenna. When a
is caused by a chemical reaction that excites electrons photon strikes one of the pigment molecules, the energy
CHECKPOINT
of a fluorescent dye (Figure 7.8b). The excited electrons jumps from molecule to molecule until it arrives at the
quickly fall back down to their ground state, releasing reaction center of the photosystem. The reaction center What is the role of a
reaction center during
energy in the form of fluorescent light. consists of chlorophyll a molecules that sit next to another
photosynthesis?
In the thylakoid membrane, chlorophyll molecules molecule called a primary electron acceptor. This primary chemical reactions.
are organized with other molecules into photosystems. electron acceptor traps the light-excited electron ( e– ) use this trapped energy to drive

Each photosystem has a cluster of a few hundred pig- from the chlorophyll a in the reaction center. Another
molecules to molecules that can
a light-excited photon from pigment
ment molecules, including chlorophylls a and b and team of molecules built into the thylakoid membrane then Answer: A reaction center transfers

some carotenoids (Figure 7.9). This cluster of pigment uses that trapped energy to make ATP and NADPH.

How the Light Reactions Generate ATP and NADPH

Primary NADP+
Two photosystems cooperate in the light reactions electron 2e–
(Figure 7.10). Photons excite electrons in the chlo- acceptor
rophyll of the first photosystem. These photons are Energy
3
to make ATP 2e–
then trapped by the primary electron acceptor. Primary
electron 2
This photosystem then replaces the lost electrons acceptor
– –
NADPH
by extracting new ones from water. This is the
step that releases O2 during photosyn- 2e–
Light
thesis. Energized electrons from Ele
ctro
the first photosystem pass down an nt
ran
spo
electron transport chain to the sec- Light rt c
hai
ond photosystem. The chloroplast n

uses the energy released by this

electron “fall” to make ATP. The Reaction-
center
second photosystem transfers its chlorophyll
1
light-excited electrons to NADP+,
reducing it to NADPH. Reaction-
center Second photosystem
H 2O chlorophyll

� Figure 7.10 The light reactions

2e–
of photosynthesis. The orange
First photosystem
arrows trace a light-driven flow of
electrons from H2O to NADPH. These 2 H+ + 1
O2
2
electrons also produce ATP.

147
CHAPTER 7 Figure 7.11 shows the location of the light reactions # Figure 7.12 The light reactions illustrated using
PHOTOSYNTHESIS: USING in the thylakoid membrane. The two photosystems and a hard-hat analogy.
LIGHT TO MAKE FOOD
the electron transport chain that connects them transfer
electrons from H2O to NADP+, producing NADPH.
Notice that the mechanism of ATP production during
e–
the light reactions is very similar to the mechanism we
saw in cellular respiration (see Figure 6.10). In both ATP

cases, an electron transport chain pumps hydrogen ions

(H+) across a membrane—the inner mitochondrial e– e–
– –
CHECKPOINT membrane in the case of respiration and the thylakoid NADPH

1. Why is water required as a membrane in photosynthesis. And in both cases, ATP

e–
reactant in photosynthesis? synthases use the energy stored by the H+ gradient to
(Hint: Review Figures 7.10 e–
make ATP. The main difference is that food provides the e–
and 7.11.)
high-energy electrons in cellular respiration, whereas
2. In addition to conveying

Photon
electrons between light-excited electrons flow down the transport chain Electron
photosystems, the during photosynthesis. The traffic of electrons shown transport
chain
electron transport chains in Figures 7.10 and 7.11 is analogous to the cartoon in
of chloroplasts provide Figure 7.12.
the energy for the We have seen how the light reactions absorb solar e–
synthesis of _________.
energy and convert it to the chemical energy of ATP and

Photon
NADPH. Notice again, however, that the light reactions
NAD+ to NADPH. 2. ATP
provides electrons for converting
Answers: 1. The splitting of water did not produce any sugar. That’s the job of the Calvin
cycle, as we’ll see next. First photosystem Second photosystem

# Figure 7.11 How the thylakoid

membrane converts light energy to the
chemical energy of NADPH and ATP.
Calvin
cycle

Light Light
– –
H+

:
NADPH
ATP
ADP + P
NADP+
H+

Stroma

Photosystem Electron transport chain Photosystem

Thylakoid ATP
membrane synthase

Inside thylakoid Electron flow

2e–
H+ H+
H 2O H+
H+ H+
1 O2
2

Thylakoid
membrane

148
The Calvin Cycle: Making Sugar
THE CALVIN CYCLE:
MAKING SUGAR
FROM CARBON DIOXIDE

from Carbon Dioxide CO2 (from air)

If chloroplasts are CO2

Light H2O
solar-powered sugar 1 An enzyme adds each CO2 to a
five-carbon sugar called RuBP. The
factories, then the +
NADP resulting molecule breaks into two
Calvin cycle is ADP P three-carbon molecules.
+ P Calvin
Light
the actual sugar- reactions cycle
ATP 4 Using energy from ATP, RuBP sugar Three-carbon molecule
manufacturing – – enzymes rearrange the
machinery. This NADPH remaining G3P sugars to
regenerate RuBP. ATP
process is called a P

Ribulos
O2
Sugar
cycle because its start- P
ing material is regenerated. ADP + P
ADP + P Calvin

isphate
With each turn of the cycle, there are chemical cycle – –
inputs and outputs. The inputs are CO2 from NADPH
ATP
the air as well as ATP and NADPH produced by
the light reactions. Using carbon from CO2, en- NADP+
ergy from ATP, and high-energy electrons from G3P sugar G3P sugar
NADPH, the Calvin cycle constructs an energy- P P
rich sugar molecule called glyceraldehyde
3-phosphate (G3P). The plant cell can then use 3 For every three molecules of CO2 2 Using energy from ATP and NADPH
G3P as the raw material to make the glucose that enter the cycle, the net output is produced by the light reactions,
one G3P sugar. The other G3P sugars enzymes convert each three-carbon
and other organic compounds (such as cellulose continue in the cycle. molecule to the three-carbon sugar G3P.
and starch) that it needs. Figure 7.13 presents
the basics of the Calvin cycle, emphasizing in-
puts and outputs. Each symbol represents a
� Figure 7.13 The Calvin cycle. With
carbon atom, and each P symbol represents a the energy of ATP and the electrons
phosphate group. G3P sugar
from NADPH, the carbons from CO2
are used to make a three-carbon sugar Glucose (and
P other organic
called G3P.
compounds)

Biofuels EVOLUTION CONNECTION

Creating a Better Biofuel Biomechanical engineers are working to solve this

Factory dilemma by turning to an obvious example: evolution by

natural selection. In nature, organisms with genes that
Throughout this chapter, we’ve studied how plants make them better suited to their local environment will,
convert solar energy to chemical energy via photosyn- on average, more often survive and pass those genes on
thesis. Such transformations are vital to our welfare and to the next generation. Repeated over many generations,
to Earth’s ecosystems. As discussed in the Biology and genes that enhance survival within that environment CHECKPOINT
Society section, scientists are attempting to tap into the will become more common, and the species evolves. What is the function of
“green energy” of photosynthesis to produce biofuels. When trying to solve an engineering problem, sci- NADPH in the Calvin cycle?
But the production of biofuels is highly inefficient. In entists can impose their own desired outcomes using
form G3P (a sugar).
electrons that are added to CO2 to
fact, it is usually far more costly to produce biofuels than a process called directed evolution (see the Process Answer: It provides the high-energy

to extract the equivalent amount of fossil fuels. of Science section in Chapter 5 for another example).

149
CHAPTER 7 During this process, scientists in the laboratory # Figure 7.14 Microscopic biofuel factories. This researcher is
PHOTOSYNTHESIS: USING (instead of the natural environment) determine monitoring a reaction chamber in which microscopic algae are using
LIGHT TO MAKE FOOD light to produce biofuels.
which organisms are the fittest. Directed evolu-
tion of biofuel production often involves mi-
croscopic algae (Figure 7.14) rather than plants
because algae are easier to manipulate and main-
tain within the laboratory. Furthermore, some al-
gae produce nearly half their own body weight in
hydrocarbons that are only a few chemical steps
away from useful biofuels.
In a typical directed evolution experiment,
the researcher starts with a large collection of
individual alga—sometimes naturally occurring
species and sometimes transgenic algae that
have been engineered to carry useful genes, such
as fungal genes for enzymes that break down
cellulose. The algae are exposed to mutation-
promoting chemicals. This produces a highly
varied collection of algae that can be screened
for the desired outcome: the ability to produce
the most useful biofuel in the largest quantity.
The tiny fraction of total algae that can best per-
form this task is grown and subjected to another
round of mutation and selection. After many
repetitions, the algae may slowly improve their
ability to efficiently produce biofuels. Many re-
search laboratories—some within major petro-
leum companies—are using such methods and
may someday produce an alga that can provide
the ultimate source of green energy, an achieve-
ment that would highlight how lessons from
natural evolution can be applied to improve
our!lives.

Chapter Review
Energy Transformation: An Overview of Photosynthesis
SUMMARY OF KEY CONCEPTS
Light energy
The Basics of Photosynthesis
Photosynthesis is a process whereby light energy is transformed into
chemical energy stored as bonds in sugars made from carbon dioxide and
water. 6 CO2 + 6 H2O C6H12O6 + 6 O2
Photosynthesis
Carbon Water Glucose Oxygen gas
Chloroplasts: Sites of Photosynthesis dioxide
Chloroplasts contain a thick fluid called stroma surrounding a network of
membranes called thylakoids.

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 CHAPTER REVIEW

The overall process of photosynthesis can be divided into two stages The Calvin Cycle: Making Sugar
connected by energy- and electron-carrying molecules: from Carbon Dioxide
Chloroplast
Within the stroma (fluid) of the chloroplast, carbon dioxide from the air and
CO2 ATP and NADPH produced during the light reactions are used to produce
Light H2O
G3P, an energy-rich sugar molecule that can be used to make glucose and
Stack of other organic molecules.
NADP+ Stroma
thylakoids
CO2
ADP
+ P Calvin
Light
reactions cycle
ATP
– –
NADPH
Sugar used for ATP ADP + P
• cellular respiration Calvin
O2 • cellulose – –
Sugar cycle
• starch NADPH NADP+
• other organic compounds

The Light Reactions: Converting Solar Energy

to Chemical Energy
The Nature of Sunlight
Visible light is part of the spectrum of electromagnetic energy. It travels G3P
Glucose and
through space as waves. Different wavelengths of light appear as different P other compounds
(such as cellulose
colors; shorter wavelengths carry more energy. and starch)

Chloroplast Pigments
Pigment molecules absorb light energy of certain wavelengths and reflect
other wavelengths. We see the reflected wavelengths as the color of the pig-
ment. Several chloroplast pigments absorb light of various wavelengths and For practice quizzes, BioFlix animations, MP3 tutorials, video tutors, and
convey it to other pigments, but it is the green pigment chlorophyll a that more study tools designed for this textbook, go to MasteringBiology®
participates directly in the light reactions.

How Photosystems Harvest Light Energy and How the

Light Reactions Generate ATP and NADPH SELF-QUIZ
1. The light reactions take place in the structures of the chloroplast called
NADP+
e– 2e–
the __________, while the Calvin cycle takes place in the __________.
ADP acceptor
ATP 2. In terms of the spatial organization of photosynthesis within the chloro-
e– 2e– plast, what is the advantage of the light reactions producing NADPH and
acceptor
– – ATP on the stroma side of the thylakoid membrane?
2e– Ele NADPH 3. Which of the following equations best summarizes photosynthesis?
ctro
nt
Photon
ran
spo
a. 6 CO2 + 6 H2O + 6 O2 → C6H12O6
Photon
rt c b. 6 CO2 + 6 H2O → C6H12O6 + 6 O2
hai
n
Chlorophyll c. 6 CO2 + 6 O2 → C6H12O6 + 6 H2O
d. C6H12O6 + 6 O2 → 6 CO2 + 6 H2O
H2O Chlorophyll 4. Explain how the name “photosynthesis” describes what this process
Second photosystem
2e– accomplishes.
First photosystem 5. Chlorophyll b primarily absorbs blue light. What color has chlorophyll
2 H+ + 12 O2 b got?
yellow

range 151
CHAPTER 7
PHOTOSYNTHESIS: USING
LIGHT TO MAKE FOOD

6. The carbon atoms that enter the Calvin cycle as CO2 eventually

combonydrates
Chloro-
asp or
end up in __________.
.

Absorption of light by
chloroplast pigments
phyll a Chlorophyll b
7. Which of the following are produced by reactions that take place in the
thylakoids and are consumed by reactions in the stroma?
a. CO2 and H2O Carotenoids
b. NADP+ and ADP
-c. ATP and NADPH
d. glucose and O2
400 500 600 700
8. The reactions of the Calvin cycle are not directly dependent on light,
and yet they usually do not occur at night. Why? Wavelength of light (nm)

9. Of the following metabolic processes, which one is common to

photosynthesis and cellular respiration?
a. reactions that convert light energy to chemical energy
b. reactions that split H2O molecules and release O2
-c. reactions that store energy by pumping H+ across membranes
d. reactions that convert CO2 to sugar
BIOLOGY AND SOCIETY
13. There is strong evidence that Earth is getting warmer because of an
Answers to these questions can be found in Appendix: Self-Quiz Answers.
intensified greenhouse effect resulting from increased CO2 emissions
from industry, vehicles, and the burning of forests. Global climate
change could influence agriculture, melt polar ice, and flood coastal
THE PROCESS OF SCIENCE regions. In response to these threats, 192 parties have accepted the
10. Oxygen is a highly reactive element that easily combines with (oxidizes) Kyoto Protocol, which calls for mandatory reductions of greenhouse
other elements and compounds. For example, it forms five different gas emissions in 30 industrialized nations by 2012. The United States
oxides with iron, which is by mass the most abundant element on Earth. has signed but not ratified (put into effect) the agreement, instead
Given its highly reactive nature, how is oxygen maintained at a level of proposing a more modest set of voluntary goals allowing businesses
16% in the Earth’s atmosphere? Based on the information in this chap- to decide whether they wish to participate and providing tax incentives
ter, draw a cycle of reactions that keeps the concentration of oxygen to encourage them to do so. The reasons given for rejecting the agree-
steady and describes its flow between different interconnected systems ment are that it might hurt the American economy and that some less
of the biosphere. Was the atmospheric concentration of oxygen higher or industrialized countries (such as India) are exempted from it, even
lower before the emergence of life? though they produce a lot of pollution. Do you agree with this decision?
11. Suppose you wanted to discover whether the oxygen atoms in the glu- In what ways might efforts to reduce greenhouse gases hurt the
cose produced by photosynthesis come from H2O or CO2. Explain how economy? How can those costs be weighed against the costs of global
you could use a radioactive isotope to find out. climate change? Should poorer nations carry an equal burden to reduce
12. Interpreting Data The graph above right is called an absorption spec- their emissions?
trum. Each line on the graph is made by shining light of varying wave- 14. In this age of ecological awareness, generation of energy from sunlight
lengths through a sample. For each wavelength, the amount of that light is gaining popularity. Drawbacks of this technology are the use of toxic
absorbed by the sample is recorded. This graph combines three such heavy metals in some photovoltaic cells and its still relatively low cost
measurements, one each for the pigments chlorophyll a, chlorophyll b, efficiency. Given that plants are experts in harvesting sunlight, would it
and the carotenoids. Notice that the graphs for the chlorophyll pigments not make sense to use their photosystems to convert sunlight into
match the data presented in Figure 7.5. Imagine a plant that lacks chloro- energy? Can you think of a device that performs this feat?
phyll and relies only on carotenoids for photosynthesis. What colors of light
would work best for this plant? How would this plant appear to your eye?

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