Chapter 12 | Photosynthesis 217

12 | PHOTOSYNTHESIS
12.1 | Overview of Photosynthesis

The sun not only provides the energy for the plants themselves, but for all other creatures on the planet.
Indeed, ancient photosynthesis provided the fossil fuel energy that we use to generate electricity or power our
cars. Each cell in every organism runs on the chemical energy found mainly in carbohydrate molecules (food),
and the majority of these molecules are produced by one process: photosynthesis. Through photosynthesis,
certain organisms convert solar energy (sunlight) into chemical energy, which is then used to build carbohydrate
molecules. The energy used to hold these molecules together is released when an organism breaks down food
during cellular respiration. Cells then use this energy to perform work, such as movement.
The energy that is harnessed from photosynthesis enters the ecosystems of our planet continuously and is
transferred from one organism to another until almost all of the harvested energy is transferred and released as
heat energy. Therefore, directly or indirectly, the process of photosynthesis provides most of the energy required
by living things on earth (Figure 12.1).

Figure 12.1 This image shows the role of photosynthesis is the flow of energy through and ecosystem. Light energy
enters the system through photosynthesis and leaves primarily as heat energy once the energy is used by organisms.
(image by Eva Horne and Robert Bear)

Photosynthesis also results in the release of oxygen into the atmosphere. In short, to eat and breathe, humans
depend almost entirely on the organisms that carry out photosynthesis.

Solar Dependence and Food Production

Some organisms can carry out photosynthesis, whereas others cannot. An autotroph is an organism that
can produce its own food. The Greek roots of the word autotroph mean “self” (auto) “feeder” (troph). Plants
are the best-known autotrophs, but others exist, including certain types of bacteria and algae (Figure 12.2).
Oceanic algae contribute enormous quantities of food and oxygen to global food chains. Plants are also
photoautotrophs, a type of autotroph that uses sunlight and carbon from carbon dioxide to synthesize chemical
energy in the form of carbohydrates. All organisms carrying out photosynthesis require sunlight.
218 Chapter 12 | Photosynthesis

Figure 12.2 (a) Plants, (b) algae, and (c) certain bacteria, called cyanobacteria, are photoautotrophs that can carry
out photosynthesis. Algae can grow over enormous areas in water, at times completely covering the surface. (credit
a: Steve Hillebrand, U.S. Fish and Wildlife Service; credit b: "eutrophication&hypoxia"/Flickr; credit c: NASA; scale-bar
data from Matt Russell)

Heterotrophs are organisms incapable of photosynthesis that must therefore obtain energy and carbon from
food by consuming other organisms. The Greek roots of the word heterotroph mean “other” (hetero) “feeder”
(troph), meaning that their food comes from other organisms. Even if the food organism is another animal, this
food traces its origins back to autotrophs and the process of photosynthesis. Humans are heterotrophs, as are
all animals. Heterotrophs depend on autotrophs, either directly or indirectly. Deer and wolves are heterotrophs.
A deer obtains energy by eating plants. A wolf eating a deer obtains energy that originally came from the plants
eaten by that deer. The energy in the plant came from photosynthesis, and therefore it is the only autotroph
in this example. Using this reasoning, all food eaten by humans also links back to autotrophs that carry out
photosynthesis.

The Flow of Energy

Whether the organism is a bacterium, plant, or animal, all living things access energy by breaking down
carbohydrate molecules. But if plants make carbohydrate molecules, why would they need to break them down,
especially when it has been shown that the gas organisms release as a “waste product” (CO2) acts as a
substrate for the formation of more food in photosynthesis? Remember, living things need energy to perform
life functions. In addition, an organism can either make its own food or eat another organism—either way, the
food still needs to be broken down. Finally, in the process of breaking down food, called cellular respiration,
heterotrophs release needed energy and produce “waste” in the form of CO2 gas.
In nature, there is no such thing as waste. Every single atom of matter and energy is conserved, recycling over
and over infinitely. Substances change form or move from one type of molecule to another, but their constituent
atoms never disappear.
CO2 is no more a form of waste than oxygen is form of waste from photosynthesis. Both are byproducts
of reactions and move on to other reactions. Photosynthesis absorbs light energy to build carbohydrates in
chloroplasts, and aerobic cellular respiration releases energy by using oxygen to take metabolize carbohydrates
in the cytoplasm and mitochondria(Figure 12.3). Both processes use electron transport chains to capture the
energy necessary to drive the reactions, because breaking down a substance requires energy. These two
powerhouse processes, photosynthesis and cellular respiration, function in biological, cyclical harmony to allow
organisms to access life-sustaining energy that originates millions of miles away in a burning star humans call
the sun.

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Chapter 12 | Photosynthesis 219

Figure 12.3 Photosynthesis which occurs in the chloroplast consumes carbon dioxide and water while producing
carbohydrates (glucose) and oxygen while Aerobic Cellular respiration which occurs in the mitochondria consumes
glucose and oxygen while producing carbohydrates. (Image by Eva Horne and Robert Bear)

Main Structures and Summary of Photosynthesis

Photosynthesis requires sunlight, carbon dioxide, and water as starting reactants (Figure 12.4). After the
process is complete, photosynthesis releases oxygen and produces carbohydrate molecules, most commonly
glucose. These sugar molecules contain the energy that living things need to survive.
220 Chapter 12 | Photosynthesis

Figure 12.4 Photosynthesis uses solar energy, carbon dioxide, and water to release oxygen and to produce energy-
storing sugar molecules.

The complex reactions of photosynthesis can be summarized by the chemical equation shown in Figure 12.5.

Figure 12.5 The process of photosynthesis can be represented by an equation, wherein carbon dioxide and water
produce sugar and oxygen using energy from sunlight. Since the products have a higher energy level than the
reactants, photosynthesis is an endergonic reaction. (Image by Robert Bear)

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Chapter 12 | Photosynthesis 221

Although the equation looks simple, the many steps that take place during photosynthesis are actually quite
complex, as in the way that the reaction summarizing cellular respiration represented many individual reactions.
Before learning the details of how photoautotrophs turn sunlight into food, it is important to become familiar with
the physical structures involved.
In plants, photosynthesis takes place primarily in leaves, which consist of many layers of cells and have
differentiated top and bottom sides. The process of photosynthesis occurs not on the surface layers of the leaf,
but rather in a middle layer called the mesophyll (Figure 12.6). The gas exchange of carbon dioxide and oxygen
occurs through small, regulated openings called stomata.
In all autotrophic eukaryotes, photosynthesis takes place inside an organelle called a chloroplast. In plants,
chloroplast-containing cells exist in the mesophyll. Chloroplasts have a double (inner and outer) membrane.
Within the chloroplast is a third membrane that forms stacked, disc-shaped structures called thylakoids.
Embedded in the thylakoid membrane are molecules of chlorophyll, a pigment (a molecule that absorbs light)
through which the entire process of photosynthesis begins. Chlorophyll is responsible for the green color of
plants. The thylakoid membrane encloses an internal space called the thylakoid space. Other types of pigments
exist that can carry out photosynthesis, but chlorophyll is by far the most common. As shown in Figure 12.6, a
stack of thylakoids is called granum, and the space surrounding the granum is called stroma (not to be confused
with stomata, the openings on the leaves).

Figure 12.6 Not all cells of a leaf carry out photosynthesis. Cells within the middle layer of a leaf have chloroplasts,
which contain the photosynthetic apparatus. (credit "leaf": modification of work by Cory Zanker)

The Two Parts of Photosynthesis

Photosynthesis takes place in two stages: the light-dependent reactions and the Calvin cycle (Figure 12.7). In
the light-dependent reactions, which take place at the thylakoid membrane, chlorophyll absorbs energy from
sunlight and then converts it into chemical energy with the use of water. The light-dependent reactions release
oxygen from the hydrolysis of water as a byproduct. In the Calvin cycle, which takes place in the stroma,
the chemical energy derived from the light-dependent reactions drives both the capture of carbon in carbon
222 Chapter 12 | Photosynthesis

dioxide molecules and the subsequent assembly of sugar molecules. The two reactions use carrier molecules
to transport the energy from one to the other. The carriers that move energy from the light-dependent reactions
to the Calvin cycle reactions can be thought of as “full” because they bring energy. After the energy is released,
the “empty” energy carriers return to the light-dependent reactions to obtain more energy.

Figure 12.7 The process of photosynthesis is divided to two stages that are linked by the energy and electron carriers
ATP and NADPH. (Image by Eva Horne and Robert Bear)

12.2 | The Light-Dependent Reactions

The light from the sun powers every cell on the planet, allowing plants to make food that the rest of us can also
partake of. How can light be used to make food? When a person turns on a lamp, electrical energy becomes
light energy. Like all other forms of kinetic energy, light can travel, change form, and be harnessed to do work. In
the case of photosynthesis, light energy is converted into chemical energy, which photoautotrophs use to build
carbohydrate molecules (Figure 12.8). However, autotrophs only use a few specific components of sunlight.

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Chapter 12 | Photosynthesis 223

Figure 12.8 Photoautotrophs can capture light energy from the sun, converting it into the chemical energy used to
build food molecules. (credit: Gerry Atwell)

What Is Light Energy?

The sun emits an enormous amount of electromagnetic radiation (solar energy). Humans can see only a fraction
of this energy, which portion is therefore referred to as “visible light.” The manner in which solar energy travels is
described as waves. Scientists can determine the amount of energy of a wave by measuring its wavelength, the
distance between consecutive points of a wave. A single wave is measured from two consecutive points, such
as from crest to crest or from trough to trough (Figure 12.9).

Figure 12.9 The wavelength of a single wave is the distance between two consecutive points of similar position (two
crests or two troughs) along the wave.

Visible light constitutes only one of many types of electromagnetic radiation emitted from the sun and other stars.
Scientists differentiate the various types of radiant energy from the sun within the electromagnetic spectrum. The
electromagnetic spectrum is the range of all possible frequencies of radiation (Figure 12.10). The difference
between wavelengths relates to the amount of energy carried by them.
224 Chapter 12 | Photosynthesis

Figure 12.10 The sun emits energy in the form of electromagnetic radiation. This radiation exists at different
wavelengths, each of which has its own characteristic energy. All electromagnetic radiation, including visible light, is
characterized by its wavelength.

Absorption of Light
Light energy initiates the process of photosynthesis when pigments absorb the light. Organic pigments, whether
in the human retina or the chloroplast thylakoid, have a narrow range of energy levels that they can absorb.
Energy levels lower than those represented by red light are insufficient to raise an orbital electron to a
populatable, excited (quantum) state. Energy levels higher than those in blue light will physically tear the
molecules apart, called bleaching. So retinal pigments can only “see” (absorb) 700 nm to 400 nm light, which is
therefore called visible light. For the same reasons, plants pigment molecules absorb only light in the wavelength
range of 700 nm to 400 nm; plant physiologists refer to this range for plants as photosynthetically active radiation.
The visible light seen by humans as white light actually exists in a rainbow of colors. Certain objects, such as
a prism or a drop of water, disperse white light to reveal the colors to the human eye. The visible light portion
of the electromagnetic spectrum shows the rainbow of colors, with violet and blue having shorter wavelengths,
and therefore higher energy. At the other end of the spectrum toward red, the wavelengths are longer and have
lower energy (Figure 12.11).

Figure 12.11 The colors of visible light do not carry the same amount of energy. Violet has the shortest wavelength
and therefore carries the most energy, whereas red has the longest wavelength and carries the least amount of energy.
(credit: modification of work by NASA)

Understanding Pigments
Different kinds of pigments exist, and each has evolved to absorb only certain wavelengths (colors) of visible
light Figure 12.12. Pigments reflect or transmit the wavelengths they cannot absorb, making them appear in the

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Chapter 12 | Photosynthesis 225

corresponding color.
Chlorophylls and carotenoids are the two major classes of photosynthetic pigments found in plants and algae;
each class has multiple types of pigment molecules. There are five major chlorophylls: a, b, c and d and a
related molecule found in prokaryotes called bacteriochlorophyll. Chlorophyll a and chlorophyll b are found
in higher plant chloroplasts and will be the focus of the following discussion. With dozens of different forms,
carotenoids are a much larger group of pigments. The carotenoids found in fruit—such as the red of tomato
(lycopene), the yellow of corn seeds (zeaxanthin), or the orange of an orange peel (β-carotene)—are used as
advertisements to attract seed dispersers. In photosynthesis, carotenoids function as photosynthetic pigments
that are very efficient molecules for the disposal of excess energy. Many carotenoids reside in the thylakoid
membrane, absorb excess energy, and safely dissipate that energy as heat.

Figure 12.12 (a) Chlorophyll a, (b) chlorophyll b, and (c) β-carotene are hydrophobic organic pigments found in
the thylakoid membrane. Chlorophyll a and b, which are identical except for the part indicated in the red box, are
responsible for the green color of leaves. β-carotene is responsible for the orange color in carrots. Each pigment has
(d) a unique absorbance spectrum.

Many photosynthetic organisms have a mixture of pigments; using them, the organism can absorb energy from
a wider range of wavelengths. Not all photosynthetic organisms have full access to sunlight. Some organisms
grow underwater where light intensity and quality decrease and change with depth. Other organisms grow in
competition for light. Plants on the rainforest floor must be able to absorb any bit of light that comes through,
because the taller trees absorb most of the sunlight and scatter the remaining solar radiation.

How Light-Dependent Reactions Work

The overall function of light-dependent reactions is to convert solar energy into chemical energy in the form of
NADPH and ATP. This chemical energy supports the Calvin cycle and fuels the assembly of sugar molecules.
The light-dependent reactions are depicted in Figure 12.14. Protein complexes and pigment molecules work
together to produce NADPH and ATP.
226 Chapter 12 | Photosynthesis

Figure 12.13 A photosystem consists of a light-harvesting complex and a reaction center. Pigments in the light-
harvesting complex pass light energy to two special chlorophyll a molecules in the reaction center. The light excites
an electron from the chlorophyll a pair, which passes to the primary electron acceptor. The excited electron must then
be replaced. In (a) photosystem II, the electron comes from the splitting of water, which releases oxygen as a waste
product. In (b) photosystem I, the electron comes from the chloroplast electron transport chain discussed below.

The actual step that converts light energy into chemical energy takes place in a multiprotein complex called a
photosystem Figure 12.13, two types of which are found embedded in the thylakoid membrane, photosystem
II (PSII) and photosystem I (PSI) (Figure 12.14). The two complexes differ on the basis of what they oxidize
(that is, the source of the low-energy electron supply) and what they reduce (the place to which they deliver their
energized electrons).
Both photosystems have the same basic structure; a number of antenna proteins to which the chlorophyll
molecules are bound surround the reaction center where the photochemistry takes place. Each photosystem
is serviced by the light-harvesting complex, which passes energy from sunlight to the reaction center; it consists
of multiple antenna proteins that contain a mixture of 300–400 chlorophyll a and b molecules as well as other
pigments like carotenoids. The absorption of a single photon or distinct quantity or “packet” of light by any of
the chlorophylls pushes that molecule into an excited state. In short, the light energy has now been captured
by biological molecules but is not stored in any useful form yet. The energy is transferred from chlorophyll to
chlorophyll until eventually (after about a millionth of a second), it is delivered to the reaction center. Up to this
point, only energy has been transferred between molecules, not electrons.

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Chapter 12 | Photosynthesis 227

Figure 12.14 In the photosystem II (PSII) reaction center, energy from sunlight is used to extract electrons from water.
The electrons travel through the chloroplast electron transport chain to photosystem I (PSI), which reduces NADP+ to
NADPH. The electron transport chain moves protons across the thylakoid membrane into the lumen. At the same time,
splitting of water adds protons to the lumen, and reduction of NADPH removes protons from the stroma. The net result
is a low pH in the thylakoid lumen, and a high pH in the stroma. ATP synthase uses this electrochemical gradient to
make ATP.

The reaction center contains a pair of chlorophyll a molecules with a special property. Those two chlorophylls
can undergo oxidation upon excitation; they can actually give up an electron in a process called a photoact. It
is at this step in the reaction center, this step in photosynthesis, that light energy is converted into an excited
electron. All of the subsequent steps involve getting that electron onto the electron carrier NADP+ for delivery to
the Calvin cycle where the electron is deposited onto carbon for long-term storage in the form of a carbohydrate.
PSII and PSI are two major components of the photosynthetic electron transport chain, which also includes
the cytochrome complex, a group of reversibly oxidizable and reducible proteins that forms part of the electron
transport chain between PSII and PSI.
The reaction center of PSII (called P680) delivers its high-energy electrons, one at a time, to a series of proteins
and electron carriers (or primary electron acceptors, which are pigments or other organic molecules in the
reaction center that accept energized electrons from the reaction center) that sits between it and PSI. P680’s
missing electron is replaced by extracting a low-energy electron from water; thus, water is split and PSII is re-
reduced after every photoact. Splitting one H2O molecule releases two electrons, two hydrogen atoms, and one
atom of oxygen. Splitting two molecules is required to form one molecule of diatomic O2 gas. About 10 percent
of the oxygen is used by mitochondria in the leaf to support oxidative phosphorylation. The remainder escapes
to the atmosphere where it is used by aerobic organisms to support respiration.
As electrons move through the proteins that reside between PSII and PSI, they lose energy. That energy is used
to move hydrogen atoms from the stromal side of the membrane to the thylakoid lumen. Those hydrogen atoms,
plus the ones produced by splitting water, accumulate in the thylakoid lumen and will be used synthesize ATP
in a later step. Because the electrons have lost energy prior to their arrival at PSI, they must be re-energized by
PSI, hence, another photon is absorbed by the PSI antenna. That energy is relayed to the PSI reaction center
(called P700). P700 is oxidized and sends a high-energy electron to NADP+ to form NADPH. Thus, PSII captures
the energy to make ATP, and PSI captures the energy to reduce NADP+ into NADPH. The two photosystems
work in concert, in part, to guarantee that the production of NADPH will roughly equal the production of ATP.
Other mechanisms exist to fine tune that ratio to exactly match the chloroplast’s constantly changing energy
needs.
228 Chapter 12 | Photosynthesis

Overview of Light-Dependent Reaction

Light energy is harvested and transformed into short term chemical energy during the light-dependent reactions.
The pigments in the reaction center of photosytem II absorb light energy and excite electrons into the electron
transport chain. To replace the electrons excited in photosystem II, water is spilt releasing electrons, H+ ions
and oxygen gas. As the electrons move through the electron transport chain, energy, release by the movement
of electrons, is used to produce a H+ ion gradient inside the thylakoid. The process of chemiosmosis uses
the H+ concentration to produce ATP. After flowing through the electron transport chain, the electrons enter
photosystem I. The reaction center of photosystem I absorbs more light energy and excites electrons. These
energized electrons reduce NADP+, to NADPH. See (Figure 12.15)below for a visual representation of the light-
dependent reactions.

Figure 12.15 This is an image of the Light-Dependent Reactions of photosynthesis. (Image by Eva Horne and Robert
Bear)

12.3 | Calvin Cycle

After the energy from the sun is converted into chemical energy and temporarily stored in ATP and NADPH
molecules, the cell has the fuel needed to build carbohydrate molecules for long-term energy storage. The
products of the light-dependent reactions, ATP and NADPH, have lifespans in the range of millionths of seconds,
whereas the products of the light-independent reactions (carbohydrates and other forms of reduced carbon)
can survive for hundreds of millions of years. Carbohydrates, like the cellulose that makes up the bulk of most
plants, obviously contain carbon. Where does the carbon come from? It comes from the air, in the form of carbon
dioxide, the gas that is a waste product of respiration in microbes, fungi, plants, and animals.

The Calvin Cycle

In plants, carbon dioxide (CO2) enters the leaves through stomata, where it diffuses over short distances through
intercellular spaces until it reaches the mesophyll cells. Once in the mesophyll cells, CO2 diffuses into the stroma
of the chloroplast—the site of light-independent reactions of photosynthesis. These reactions actually have
several names associated with them. Another term, the Calvin cycle, is named for the man who discovered
it, and because these reactions function as a cycle. Others call it the Calvin-Benson cycle to include the name
of another scientist involved in its discovery. The most outdated name is dark reactions (the term originally

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Chapter 12 | Photosynthesis 229

used by Melvin Calvin, who got the Nobel Prize for elucidating these reactions), because light is not directly
required (Figure 12.16). However, the term dark reactions can be misleading because it implies incorrectly that
the reaction only occurs at night or is independent of light, which is why it has faded from everyday usage.

Figure 12.16 Light reactions harness energy from the sun to produce chemical bonds, ATP, and NADPH. These
energy-carrying molecules are made in the stroma where carbon fixation takes place. Work by Eva Horne and Robert
A. Bear

The light-independent reactions of the Calvin cycle can be organized into three basic stages: fixation, reduction,
and regeneration.
Stage 1: Fixation
In the stroma, in addition to CO2, two other components are present to initiate the light-independent reactions: an
enzyme called ribulose bisphosphate carboxylase (RuBisCO), and molecules of ribulose bisphosphate (RuBP),
as shown in Figure 12.17. RuBisCO catalyzes a reaction between CO2 and RuBP. For each CO2 molecule that
reacts with one RuBP, two molecules of phosphoglycerate (PGA) form.
Stage 2: Reduction
ATP and NADPH are used to convert the six molecules of PGA into six molecules of a chemical called
phosphoglyceraldehyde (PGAL). That is a reduction reaction because it involves the gain of electrons (from
NADPH) by PGA. Recall that a reduction is the gain of an electron by an atom or molecule. Six molecules of
both ATP and NADPH are used; making glucose is obviously an energy-intensive activity.
Stage 3: Regeneration
Interestingly, at this point, only one of the PGAL molecules leaves the Calvin cycle and is sent to the cytoplasm
to contribute to the formation of other compounds needed by the plant. Because the PGAL exported from
the chloroplast has three carbon atoms, it takes three “turns” of the Calvin cycle to fix enough net carbon to
export one PGAL. But each turn makes two PGAL, thus three turns make six PGAL. One is exported while the
remaining five PGAL molecules remain in the cycle and are used to regenerate RuBP, which enables the system
to prepare for more CO2 to be fixed. Three more molecules of ATP are used in these regeneration reactions.
Overview of Calvin Cycle
During the Calvin cycle, energy and electrons harvested in the light-dependent reactions are used to produce
carbohydrates i.e. glucose. There are three stages in the Calvin cycle. The first stage is carbon fixation, CO2
from the atmosphere is attached to an organic molecule RuBP during this stage. The second stage is the carbon
230 Chapter 12 | Photosynthesis

reduction, the energy and electrons in ATP and NADPH are used to produce carbohydrates (glucose). The last
stage is the regeneration stage, energy from ATP is used to regenerate the first substrate of the cycle (RUBP).
See (Figure 12.17) below for a review of the Calvin Cycle.

Figure 12.17 This image represents the Calvin Cycle. (Image by Eva Horne and Robert Bear)

Photosynthesis
During the evolution of photosynthesis, a major shift occurred from the bacterial type of photosynthesis
that involves only one photosystem and is typically anoxygenic (does not generate oxygen) into modern
oxygenic (does generate oxygen) photosynthesis, employing two photosystems. This modern oxygenic
photosynthesis is used by many organisms—from giant tropical leaves in the rainforest to tiny
cyanobacterial cells—and the process and components of this photosynthesis remain largely the same.
Photosystems absorb light and use electron transport chains to convert energy into the chemical energy of
ATP and NADH. The subsequent light-independent reactions then assemble carbohydrate molecules with
this energy.
Photosynthesis in desert plants has evolved adaptations that conserve water. In the harsh dry heat, every
drop of water must be used to survive. Because stomata must open to allow for the uptake of CO2, water
escapes from the leaf during active photosynthesis. Desert plants have evolved processes to conserve
water and deal with harsh conditions. A more efficient use of CO2 allows plants to adapt to living with
less water. Some plants such as cacti (Figure 12.18) can prepare materials for photosynthesis during the
night by a temporary carbon fixation/storage process, because opening the stomata at this time conserves
water due to cooler temperatures. In addition, cacti have evolved the ability to carry out low levels of
photosynthesis without opening stomata at all, an extreme mechanism to face extremely dry periods.

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Chapter 12 | Photosynthesis 231

Figure 12.18 The harsh conditions of the desert have led plants like these cacti to evolve variations of the light-
independent reactions of photosynthesis. These variations increase the efficiency of water usage, helping to conserve
water and energy. (credit: David A. Rintoul)

Overview of Photosynthesis
Photosynthesis converts light energy to chemical energy in two stages, the light-dependent reactions and the
Calvin cycle. By exploring these two sets of reactions, we learned how photons of light energy are turned
into food by photosynthesis. The light-dependent reactions harvest the light energy to make ATP and to
transfer electrons from H2O to NADP+ forming NADPH and Oxygen gas. The energy and electrons in ATP and
NADPH are used in the Calvin cycle to produce glucose from carbon dioxide. The sunlight energy entering the
chloroplasts becomes stored as the chemical bonds in the organic molecules. See (Figure 12.19) below for a
review of photosynthesis.
232 Chapter 12 | Photosynthesis

Figure 12.19 The process of photosynthesis is divided to two stages that are linked by the energy and electron carriers
ATP and NADPH. The light-dependent reactions split water and releases oxygen as a byproduct, and these reactions
convert light energy to chemical energy (ATP and NADPH). The Calvin cycle uses the energy in ATP and NADPH and
produces carbohydrates by fixing CO2 a byproduct of aerobic cellular respiration. (Image by Eva Horne and Robert
Bear)

What is the fate of the carbohydrates produced by photosynthesis? About 50% of the carbohydrates are used
by the plant for aerobic cellular respiration in their mitochondria. The other 50% of the carbohydrates are the
building blocks for the biological macromolecules the make up plant cells that you learned about in module 3. As
you may realize, these biological macromolecules are the food we eat, and you are what you eat. Taking this a
bit further, all the activities you do from reading this text to sleeping require energy and that energy comes from
the Sun.

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