Photosynthesis: The Role of Light

The heart of photosynthesis as it occurs in most autotrophs consists of two key processes:

• the removal of hydrogen (H) atoms from water molecules (photolysis) : light
reaction
• the reduction of carbon dioxide (CO2) by these hydrogen atoms to form
organic molecules. (carbon fixation/Calvin cycle) : dark reaction

Light Reaction
The electrons (e−) and protons (H+) that make up hydrogen atoms are stripped away
separately from water molecules.

2H2O - → 4e− + 4H+ + O2

The electrons serve two functions:

• They reduce NADP+ to NADPH for use in the Calvin Cycle.

• They set up an electrochemical charge that provides the energy for pumping
protons from the stroma of the chloroplast into the interior of the thylakoid.

The protons also serve two functions:

• They participate in the reduction of NADP+ to NADPH.

• As they flow back out from the interior of the thylakoid (by facilitated diffusion),
passing down their concentration gradient), the energy they give up is harnessed
to the conversion of ADP to ATP.
• Because it is drive by light, this process is called photophosphorylation.

ADP + Pi - → ATP

The ATP provides the second essential ingredient for running the Calvin Cycle.

The removal of electrons from water molecules and their transfer to NADP+ requires
energy. The electrons are moving from a redox potential of about +0.82 volt in water to
−0.32 volt in NADPH. Thus enough energy must be available to move them against a
total potential of 1.14 volts. Where does the needed energy come from? Light.

The Chloroplast

Chloroplasts contain a system of thylakoid membranes surrounded by a fluid stroma.

Chloroplasts

A typical plant cell (e.g., in the palisade layer of a leaf) might contain as many as 50
chloroplasts.

The chloroplast is made up of 3 types of membrane:

1. A smooth outer membrane which is freely permeable to molecules.

2. A smooth inner membrane which contains many transporters: integral
membrane proteins that regulate the passage in an out of the chloroplast of
o small molecules like sugars
o proteins synthesized in the cytoplasm of the cell but used within the
chloroplast
3. A system of thylakoid membranes

Thylakoids

• The thylakoid membranes enclose a lumen: a system of vesicles (that may all be
interconnected).
• At various places within the chloroplast these are stacked in arrays called grana
(resembling a stack of coins).
• Four types of protein assemblies are embedded in the thylakoid membranes:
1. Photosystem I which includes chlorophyll and carotenoid molecules
2. Photosystem II which also contains chlorophyll and carotenoid molecules
3. Cytochromes b and f
4. ATP synthase

These carry out the so-called light reactions of photosynthesis.

• The thylakoid membranes are surrounded by a fluid stroma.

 The stroma contains:

o all the enzymes, e.g., RUBISCO, needed to carry out the "dark" reactions
of photosynthesis; that is, the conversion of CO2 into organic molecules like
glucose.
o A number of identical molecules of DNA, each of which carries the
complete chloroplast genome. The genes encode some - but not all - of the
molecules needed for chloroplast function. The others are
 transcribed from genes in the nucleus of the cell
 translated in the cytoplasm and
 transported into the chloroplast.

The electron micrograph on the left

(courtesy of Kenneth R. Miller) shows
the inner surface of a thylakoid
membrane. Each particle may represent
one photosystem II complex. In the
functioning chloroplast, these particles
may not be as highly ordered as seen
here.

Six different complexes of integral membrane proteins are embedded in the thylakoid
membrane. The exact structure of these complexes differs from group to group (e.g.,
plant vs. alga) and even within a group (e.g., illuminated in air or underwater). But, in
general, one finds:

1. Photosystem I

The structure of photosystem I in a cyanobacterium ("blue-green alga") has been

completely worked out. It probably closely resembles that of plants as well.

It is a homotrimer with each subunit in the trimer containing:

• 12 different protein molecules bound to

• 96 molecules of chlorophyll a
o 2 molecules of the reaction center chlorophyll P700
o 4 accessory molecules closely associated with them
o 90 molecules that serve as antenna pigments
• 22 carotenoid molecules
• 4 lipid molecules
• 3 clusters of Fe4S4 + 2 phylloquinones
 •

Chlorophylls and Carotenoids

Chlorophylls

Two types of chlorophyll are found in plants and the green algae.

• chlorophyll a and
• chlorophyll b

In the chloroplast, both types are associated with integral membrane proteins in the
thylakoid membrane.

Note the system of alternating single and double bonds (white bars) that run around the
porphyrin ring. Although I am forced to draw the single and double bonds in fixed
positions, actually the "extra" electrons responsible for the double bonds are not fixed
between any particular pair of carbon atoms but instead are free to migrate around the ring.
This property enables these molecules to absorb light.

Both chlorophylls absorb light most strongly in the red and violet parts of the spectrum.
Green light is absorbed poorly. Thus when white light shines on chlorophyll-containing
structures like leaves, green light is transmitted and reflected and the structures appear
green.

Carotenoids

Chloroplasts also contain carotenoids. These are also pigments with colors ranging from
red to yellow.

Carotenoids absorb light most strongly in the blue portion of the spectrum. They thus
enable the chloroplast to trap are larger fraction of the radiant energy falling on it.

Carotenoids are often the major pigments in flowers and fruits. The red of a ripe tomato
and the orange of a carrot are produced by their carotenoids.

In leaves, the carotenoids are usually masked by the chlorophylls. In the autumn, as the
quantity of chlorophyll in the leaf carotenoid
declines, the carotenoids become visible and produce the
yellows and reds of autumn foliage.

beta-carotene, one of the most abundant carotenoids. Note again the system of alternating
single and double bonds that in this molecule runs along the hydrocarbon chain that
connects the two benzene rings. As in chlorophyll, the electrons of the double bonds
actually migrate though the chain and also make this molecule an efficient absorber of
light.

2. Photosystem II

Photosystem II is also a complex of

• as many as 20 different protein molecules bound to

• 50 or more chlorophyll a molecules
o 2 molecules of the reaction center chlorophyll P680
o 2 accessory molecules close to them
o 2 molecules of pheophytin (chlorophyll without the Mg++)
o the remaining molecules of chlorophyll a serve as antenna pigments.
• some half dozen carotenoid molecules. These also serve as antenna pigments.
• 2 molecules of plastoquinone

3. Light-Harvesting Complexes (LHC)

• LHC-I associated with photosystem I

• LHC-II associated with photosystem II

These contain several protein molecules associated with scores of chlorophylls — both
chlorophyll a and chlorophyll b. These LHCs also act as antenna pigments harvesting
light and passing its energy on to their respective photosystems.

4.Cytochromes b6 and f

5.ATP synthase

How the System Works

• Light is absorbed by the antenna pigments of Photosystems II and I

• The absorbed energy is transferred to the reaction center pigment, P680 in
Photosystem II, P700 in Photosystem I
• Activation of P680 removes an electron from it.
• With its resulting positive charge, P680 is sufficiently electronegative that it can
remove electrons from water.
• These electrons are transferred (by way of plastoquinone — PQ in the figure) to
the cytochrome b6/f complex where they provide the energy for
chemiosmosis(ATP).
 • Activation of P700 enables it to pick up electrons from the cytochrome b6/f
complex (by way of plastocyanin — PC in the figure) and raise them to a
sufficiently high redox potential that, after passing through ferredoxin (Fd in the
figure),
• they can reduce NADP+ to NADPH.

The sawtooth shifts in redox potential as electrons pass from P680 to NADP+ have caused
this system to be called the Z-Scheme (although as I have drawn the diagram, it looks
more like an "N"). It is also called noncyclic photophosphorylation because it produces
ATP in a one-way process (unlike cyclic photophosphorylation and pseudocyclic
photophosphorylation described below).

Chemiosmosis in Chloroplasts

The energy released as electrons pass down the gradient between Photosystem II and
Photosystem I is harnessed by the cytochrome b6/f complex to pump protons (H+)
against their concentration gradient from the stroma of the chloroplast into the interior of
the thylakoid (an example of active transport). As their concentration increases inside
(which is the same as saying that the pH of the interior decreases), a strong diffusion
gradient is set up. The only exit for these protons is through the ATP synthase complex.
As in mitochondria, the energy released as these protons flow down their gradient is
harnessed to the synthesis of ATP. The process is called chemiosmosis and is an
example of facilitated diffusion
 chemiosmosis

Each CO2 taken up by the Calvin cycle) requires:

o 2 NADPH molecules and

o 3 ATP molecules
• Each molecule of oxygen released by the light reactions supplies the 4 electrons
needed to make 2 NADPH molecules.
 • The chemiosmosis driven by these 4 electrons as they pass through the PQ +
cytochrome b6/f complex liberates only enough energy to pump 12 protons into
the interior of the thylakoid.
• But in order to make 3 molecules of ATP, the ATPase in chloroplasts appears to
have 14 protons (H+) pass through it.
• So there appears to be a deficit of 2 protons.
• How is this deficit to be made up?
• One likely answer: cyclic photophosphorylation.

In cyclic photophosphorylation,

• the electrons expelled by the energy of light absorbed by photosystem I pass, as

normal, to ferredoxin,
• but instead of then going on to reduce NADP+ to NADPH, they pass to
plastoquinone and on back into the cytochrome b6/f complex.
• Here the energy each electron liberates pumps 2 protons (H+) into the interior of
the thylakoid — enough to make up the deficit left by noncyclic
photophosphorylation.

This process is truly cyclic because no outside source of electrons is required. Like the
photocell in a light meter, photosystem I is simply using light to create a flow of current.
The only difference is that instead of using the current to move the needle on a light
meter, the chloroplast uses the current to help synthesize ATP.

Antenna Pigments
Chlorophylls a and b differ slightly in the wavelengths of light that they absorb best
(although both absorb red and blue much better than yellow and green). Carotenoids help
fill in the gap by strongly absorbing green light. The entire complex ensures that most of
the energy of light will be trapped and passed on to the reaction center chlorophylls.

Photosynthesis: Pathway of Carbon Fixation

Photosynthesis is the synthesis of organic molecules using the energy of light. For the
sugar glucose (one of the most abundant products of photosynthesis) the equation is:

6CO2 + 12H2O → C6H12O6 + 6H2O + 6O2

Light provides the energy to:

• transfer electrons from water to nicotinamide adenine dinucleotide phosphate

(NADP+) forming NADPH;
• generate ATP.
ATP and NADPH provide the energy and electrons to reduce carbon dioxide (CO2) to
organic molecules.

The Steps
• CO2 combines with the phosphorylated 5-carbon sugar ribulose bisphosphate.
• This reaction is catalyzed by the enzyme ribulose bisphosphate carboxylase
oxygenase (RUBISCO)(an enzyme which can fairly claim to be the most
abundant protein on earth).
• The resulting 6-carbon compound breaks down into two molecules of 3-
phosphoglyceric acid (PGA).
• The PGA molecules are further phosphorylated (by ATP) and are reduced (by
NADPH) to form phosphoglyceraldehyde (PGAL).
• Phosphoglyceraldehyde serves as the starting material for the synthesis of glucose
and fructose.
• Glucose and fructose make the disaccharide sucrose, which travels in solution to
other parts of the plant (e.g., fruit, roots).
• Glucose is also the monomer used in the synthesis of the polysaccharides starch
and cellulose.
 The first stable compound
are 3 PG, So they named
C3 cycle or Calvin cycle

The graphic shows the steps in the fixation of carbon dioxide during photosynthesis.
These steps were worked out by Melvin Calvin and his colleagues at the University of
California and, for this reason, are named the Calvin cycle.

All the reactions of carbon fixation occur in the stroma of the chloroplast.

Photorespiration and C4 Plants •

All plants carry on photosynthesis by

• adding carbon dioxide (CO2) to a phosphorylated 5-carbon sugar called ribulose

bisphosphate.
• This reaction is catalyzed by the enzyme ribulose bisphosphate carboxylase
oxygenase (RUBISCO).
• The resulting 6-carbon compound breaks down into two molecules of 3-
phosphoglyceric acid (PGA).
• These 3-carbon molecules serve as the starting material for the synthesis of
glucose and other food molecules.
• The process is called the Calvin cycle and the pathway is called the C3 pathway.
• When temperature rise, photosynthesis increase while the stomata close, which
then lead to a decrease of CO2 and increase O2, as a result of photosynthesis.
High O2 concentration lead to photorespiration.

Photorespiration

As its name suggests, RUBISCO catalyzes two different reactions:

• adding CO2 to ribulose bisphosphate — the carboxylase activity

• adding O2 to ribulose bisphosphate — the oxygenase activity.

Which one predominates depends on the relative concentrations of O2 and CO2 with

• (1)high CO2, low O2 favoring the carboxylase action,

• (2)high O2, low CO2 favoring the oxygenase action.

The light reactions of photosynthesis liberate oxygen and more oxygen dissolves in the
cytosol of the cell at higher temperatures. Therefore,

• high light intensities and

• high temperatures (above ~ 30°C)

favor the second (2) reaction, which is photorespiration

The details of photorespiration

• The uptake of O2 by RUBISCO forms:

o the 3-carbon molecule 3-phosphoglyceric acid — just as in the Calvin
cycle
o the 2-carbon molecule glycolate.
• The glycolate enters peroxisomes where it uses O2 to form intermediates that
• enter mitochondria where they are broken down to CO2.
So this process uses O2 and liberates CO2 as cellular respiration does which is why it is
called photorespiration.

2 glycolate ------- CO2 + P-glycerate (Sugar)

It undoes the good anabolic work of photosynthesis, reducing the net productivity of the
plant for 25 %

For this reason, much effort — so far largely unsuccessful — has gone into attempts to
alter crop plants so that they carry on less photorespiration.

The problem may solve itself. If atmospheric CO2 concentrations continue to rise, due to
increase of organic combustion, perhaps this will enhance the net productivity of the
world's crops by reducing losses to photorespiration.

C4 Plants
Over 8000 species of angiosperms, scattered among 18 different families, have developed
adaptations which minimize the losses to photorespiration.

They all use a supplementary method of CO2 uptake which forms a 4-carbon molecule
instead of the two 3-carbon molecules of the Calvin cycle. Hence these plants are called
C4 plants. (Plants that have only the Calvin cycle are thus C3 plants.)

• Some C4 plants — called CAM plants — separate their C3 and C4 cycles by time.
CAM plants are discussed below.
• Other C4 plants have structural changes in their leaf anatomy so that
o their C4 and C3 pathways are separated in different parts of the leaf with
o RUBISCO sequestered where the CO2 level is high; the O2 level low.
The details of the C4 cycle

• After entering through stomata, CO2 diffuses into a mesophyll cell.

o Being close to the leaf surface, these cells are exposed to high levels of O2,
but
o have no RUBISCO so cannot start photorespiration (nor the dark reactions
of the Calvin cycle).
• Instead the CO2 is inserted into a 3-carbon compound (C3) called
phosphoenolpyruvic acid (PEP) forming
• the 4-carbon compound oxaloacetic acid (C4).
• Oxaloacetic acid is converted into malic acid or aspartic acid (both have 4
carbons), which is
• transported (by plasmodesmata) into a bundle sheath cell. Bundle sheath cells
o are deep in the leaf so atmospheric oxygen cannot diffuse easily to them;
o often have thylakoids with reduced photosystem II complexes (the one
that produces O2).
o Both of these features keep oxygen levels
low.
• Here the 4-carbon compound is broken down into
o carbon dioxide, which enters the Calvin
cycle to form sugars and starch.
o pyruvic acid (C3), which is transported back
to a mesophyll cell where it is converted
back into PEP.

These C4 plants are well adapted to (and likely to be found

in) habitats with high daytime temperatures and intense
sunlight.
Some examples:

• crabgrass
• corn (maize)
• sugarcane
• sorghum
C4 cells in C3 plants

The ability to use the C4 pathway has evolved repeatedly in different families of
angiosperms. Perhaps the potential is in them all.

A report in the 24 January 2002 issue of Nature (by Julian M. Hibbard and W. Paul
Quick) describes the discovery that tobacco, a C3 plant, has cells capable of fixing carbon
dioxide by the C4 path. These cells are clustered around the veins (containing xylem and
phloem) of the stems and also in the petioles of the leaves. In this location, they are far
removed from the stomata that could provide atmospheric CO2. Instead, they get their
CO2 and/or the 4-carbon malic acid in the sap that has been brought up in the xylem from
the roots.

If this turns out to be true of many C3 plants, it would explain why it has been so easy for
C4 plants to evolve from C3 ancestors

CAM Plants

These are also C4 plants but instead of segregating the C4 and C3 pathways in different
parts of the leaf, they separate them in time instead. (CAM stands for crassulacean acid
metabolism because it was first studied in members of the plant family Crassulaceae.)

At night,

• CAM plants take in CO2 through their open stomata (they tend to have reduced
numbers of them).
• The CO2 joins with PEP to form the 4-carbon oxaloacetic acid.
• This is converted to 4-carbon malic acid that accumulates during the night in the
central vacuole of the cells.

In the morning,

• the stomata close (thus conserving moisture as well as reducing the inward
diffusion of oxygen).
• The accumulated malic acid leaves the vacuole and is broken down to release
CO2.
• The CO2 is taken up into the Calvin (C3) cycle.

These adaptations also enable their owners to thrive in conditions of

• high daytime temperatures

• intense sunlight
 • low soil moisture.

Some examples of CAM plants:

• cacti
• Bryophyllum
• the pineapple and all epiphytic bromeliads
• sedums
• the "ice plant" that grows in sandy parts of the scrub forest biome

C4 Diatoms
On 26 October 2000, Nature reported the discovery of both the C3 and C4 pathways in a
marine diatom. In this unicellular organism, the two paths are kept separate by having
the C4 path in the cytosol, and the C3 path confined to the chloroplast. The presence of a
C4 pathway probably reflects the frequent low concentrations of CO2 in ocean waters.

WHY STUDY PHOTOSYNTHESIS?

By Devens Gust, Ph.D.

Professor of Chemistry and Biochemistry

Center for the Study of Early Events in Photosynthesis

What is photosynthesis?

Photosynthesis is arguably the most important biological process on earth. By liberating

oxygen and consuming carbon dioxide, it has transformed the world into the hospitable
environment we know today. Directly or indirectly, photosynthesis fills all of our food
requirements and many of our needs for fiber and building materials. The energy stored
in petroleum, natural gas and coal all came from the sun via photosynthesis, as does the
energy in firewood, which is a major fuel in many parts of the world. This being the case,
scientific research into photosynthesis is vitally important. If we can understand and
control the intricacies of the photosynthetic process, we can learn how to increase crop
yields of food, fiber, wood, and fuel, and how to better use our lands. The energy-
harvesting secrets of plants can be adapted to man-made systems which provide new,
efficient ways to collect and use solar energy. These same natural "technologies" can help
point the way to the design of new, faster, and more compact computers, and even to new
medical breakthroughs. Because photosynthesis helps control the makeup of our
atmosphere, understanding photosynthesis is crucial to understanding how carbon dioxide
and other "greenhouse gases" affect the global climate. In this document, we will briefly
explore each of the areas mentioned above, and illustrate how photosynthesis research is
critical to maintaining and improving our quality of life.

Photosynthesis and food. All of our biological energy needs are met by the plant
kingdom, either directly or through herbivorous animals. Plants in turn obtain the energy
to synthesize foodstuffs via photosynthesis. Although plants draw necessary materials
from the soil and water and carbon dioxide from the air, the energy needs of the plant are
filled by sunlight. Sunlight is pure energy. However, sunlight itself is not a very useful
form of energy; it cannot be eaten, it cannot turn dynamos, and it cannot be stored. To be
beneficial, the energy in sunlight must be converted to other forms. This is what
photosynthesis is all about. It is the process by which plants change the energy in sunlight
to kinds of energy that can be stored for later use. Plants carry out this process in
photosynthetic reaction centers. These tiny units are found in leaves, and convert light
energy to chemical energy, which is the form used by all living organisms. One of the
major energy-harvesting processes in plants involves using the energy of sunlight to
convert carbon dioxide from the air into sugars, starches, and other high-energy
carbohydrates. Oxygen is released in the process. Later, when the plant needs food, it
draws upon the energy stored in these carbohydrates. We do the same. When we eat a
plate of spaghetti, our bodies oxidize or "burn" the starch by allowing it to combine with
oxygen from the air. This produces carbon dioxide, which we exhale, and the energy we
need to survive. Thus, if there is no photosynthesis, there is no food. Indeed, one widely
accepted theory explaining the extinction of the dinosaurs suggests that a comet, meteor,
or volcano ejected so much material into the atmosphere that the amount of sunlight
reaching the earth was severely reduced. This in turn caused the death of many plants and
the creatures that depended upon them for energy.

Photosynthesis and energy. One of the carbohydrates resulting from photosynthesis is

cellulose, which makes up the bulk of dry wood and other plant material. When we burn
wood, we convert the cellulose back to carbon dioxide and release the stored energy as
heat. Burning fuel is basically the same oxidation process that occurs in our bodies; it
liberates the energy of "stored sunlight" in a useful form, and returns carbon dioxide to
the atmosphere. Energy from burning "biomass" is important in many parts of the world.
In developing countries, firewood continues to be critical to survival. Ethanol (grain
alcohol) produced from sugars and starches by fermentation is a major automobile fuel in
Brazil, and is added to gasoline in some parts of the United States to help reduce
emissions of harmful pollutants. Ethanol is also readily converted to ethylene, which
serves as a feedstock to a large part of the petrochemical industry. It is possible to convert
cellulose to sugar, and then into ethanol; various microorganisms carry out this process. It
could be commercially important one day.

Our major sources of energy, of course, are coal, oil and natural gas. These materials are
all derived from ancient plants and animals, and the energy stored within them is
chemical energy that originally came from sunlight through photosynthesis. Thus, most
of the energy we use today was originally solar energy!

Photosynthesis, fiber, and materials. Wood, of course, is not only burned, but is an
important material for building and many other purposes. Paper, for example, is nearly
pure photosynthetically produced cellulose, as is cotton and many other natural fibers.
Even wool production depends on photosynthetically-derived energy. In fact, all plant
and animal products including many medicines and drugs require energy to produce, and
that energy comes ultimately from sunlight via photosynthesis. Many of our other
materials needs are filled by plastics and synthetic fibers which are produced from
petroleum, and are thus also photosynthetic in origin. Even much of our metal refining
depends ultimately on coal or other photosynthetic products. Indeed, it is difficult to
name an economically important material or substance whose existence and usefulness is
not in some way tied to photosynthesis.

Photosynthesis and the environment. Currently, there is a lot of discussion concerning

the possible effects of carbon dioxide and other "greenhouse gases" on the environment.
As mentioned above, photosynthesis converts carbon dioxide from the air to
carbohydrates and other kinds of "fixed" carbon and releases oxygen to the atmosphere.
When we burn firewood, ethanol, or coal, oil and other fossil fuels, oxygen is consumed,
and carbon dioxide is released back to the atmosphere. Thus, carbon dioxide which was
removed from the atmosphere over millions of years is being replaced very quickly
through our consumption of these fuels. The increase in carbon dioxide and related gases
is bound to affect our atmosphere. Will this change be large or small, and will it be
harmful or beneficial? These questions are being actively studied by many scientists
today. The answers will depend strongly on the effect of photosynthesis carried out by
land and sea organisms. As photosynthesis consumes carbon dioxide and releases
oxygen, it helps counteract the effect of combustion of fossil fuels. The burning of fossil
fuels releases not only carbon dioxide, but also hydrocarbons, nitrogen oxides, and other
trace materials that pollute the atmosphere and contribute to long-term health and
environmental problems. These problems are a consequence of the fact that nature has
chosen to implement photosynthesis through conversion of carbon dioxide to energy-rich
materials such as carbohydrates. Can the principles of photosynthetic solar energy
harvesting be used in some way to produce non-polluting fuels or energy sources? The
answer, as we shall see, is yes.

Why study photosynthesis?

Because our quality of life, and indeed our very existence, depends on photosynthesis, it
is essential that we understand it. Through understanding, we can avoid adversely
affecting the process and precipitating environmental or ecological disasters. Through
understanding, we can also learn to control photosynthesis, and thus enhance production
of food, fiber and energy. Understanding the natural process, which has been developed
by plants over several billion years, will also allow us to use the basic chemistry and
physics of photosynthesis for other purposes, such as solar energy conversion, the design
of electronic circuits, and the development of medicines and drugs. Some examples
follow.

Photosynthesis and agriculture. Although photosynthesis has interested mankind for

eons, rapid progress in understanding the process has come in the last few years. One of
the things we have learned is that overall, photosynthesis is relatively inefficient. For
example, based on the amount of carbon fixed by a field of corn during a typical growing
season, only about 1 - 2% of the solar energy falling on the field is recovered as new
photosynthetic products. The efficiency of uncultivated plant life is only about 0.2%. In
sugar cane, which is one of the most efficient plants, about 8% of the light absorbed by
the plant is preserved as chemical energy. Many plants, especially those that originate in
the temperate zones such as most of the United States, undergo a process called
photorespiration. This is a kind of "short circuit" of photosynthesis that wastes much of
the plants' photosynthetic energy. The phenomenon of photorespiration including its
function, if any, is only one of many riddles facing the photosynthesis researcher.

If we can fully understand processes like photorespiration, we will have the ability to
alter them. Thus, more efficient plants can be designed. Although new varieties of plants
have been developed for centuries through selective breeding, the techniques of modern
molecular biology have speeded up the process tremendously. Photosynthesis research
can show us how to produce new crop strains that will make much better use of the
sunlight they absorb. Research along these lines is critical, as recent studies show that
agricultural production is leveling off at a time when demand for food and other
agricultural products is increasing rapidly.

Because plants depend upon photosynthesis for their survival, interfering with
photosynthesis can kill the plant. This is the basis of several important herbicides, which
act by preventing certain important steps of photosynthesis. Understanding the details of
photosynthesis can lead to the design of new, extremely selective herbicides and plant
growth regulators that have the potential of being environmentally safe (especially to
animal life, which does not carry out photosynthesis). Indeed, it is possible to develop
new crop plants that are immune to specific herbicides, and to thus achieve weed control
specific to one crop species.

Photosynthesis and energy production. As described above, most of our current energy
needs are met by photosynthesis, ancient or modern. Increasing the efficiency of natural
photosynthesis can also increase production of ethanol and other fuels derived from
agriculture. However, knowledge gained from photosynthesis research can also be used
to enhance energy production in a much more direct way. Although the overall
photosynthesis process is relatively wasteful, the early steps in the conversion of sunlight
to chemical energy are quite efficient. Why not learn to understand the basic chemistry
and physics of photosynthesis, and use these same principles to build man-made solar
energy harvesting devices? This has been a dream of chemists for years, but is now close
to becoming a reality. In the laboratory, scientists can now synthesize artificial
photosynthetic reaction centers which rival the natural ones in terms of the amount of
sunlight stored as chemical or electrical energy. More research will lead to the
development of new, efficient solar energy harvesting technologies based on the natural
process.

The role of photosynthesis in control of the environment. How does photosynthesis in

temperate and tropical forests and in the sea affect the quantity of greenhouse gases in the
atmosphere? This is an important and controversial issue today. As mentioned above,
photosynthesis by plants removes carbon dioxide from the atmosphere and replaces it
with oxygen. Thus, it would tend to ameliorate the effects of carbon dioxide released by
the burning of fossil fuels. However, the question is complicated by the fact that plants
themselves react to the amount of carbon dioxide in the atmosphere. Some plants, appear
to grow more rapidly in an atmosphere rich in carbon dioxide, but this may not be true of
all species. Understanding the effect of greenhouse gases requires a much better
knowledge of the interaction of the plant kingdom with carbon dioxide than we have
today. Burning plants and plant products such as petroleum releases carbon dioxide and
other byproducts such as hydrocarbons and nitrogen oxides. However, the pollution
caused by such materials is not a necessary product of solar energy utilization. The
artificial photosynthetic reaction centers discussed above produce energy without
releasing any byproducts other than heat. They hold the promise of producing clean
energy in the form of electricity or hydrogen fuel without pollution. Implementation of
such solar energy harvesting devices would prevent pollution at the source, which is
certainly the most efficient approach to control.

Photosynthesis and electronics. At first glance, photosynthesis would seem to have no

association with the design of computers and other electronic devices. However, there is
potentially a very strong connection. A goal of modern electronics research is to make
transistors and other circuit components as small as possible. Small devices and short
connections between them make computers faster and more compact. The smallest
possible unit of a material is a molecule (made up of atoms of various types). Thus, the
smallest conceivable transistor is a single molecule (or atom). Many researchers today are
investigating the intriguing possibility of making electronic components from single
molecules or small groups of molecules. Another very active area of research is
computers that use light, rather than electrons, as the medium for carrying information. In
principle, light-based computers have several advantages over traditional designs, and
indeed many of our telephone transmission and switching networks already operate
through fiber optics. What does this have to do with photosynthesis? It turns out that
photosynthetic reaction centers are natural photochemical switches of molecular
dimensions. Learning how plants absorb light, control the movement of the resulting
energy to reaction centers, and convert the light energy to electrical, and finally chemical
energy can help us understand how to make molecular-scale computers. In fact, several
molecular electronic logic elements based on artificial photosynthetic reaction centers
have already been reported in the scientific literature.

Photosynthesis and medicine. Light has a very high energy content, and when it is
absorbed by a substance this energy is converted to other forms. When the energy ends
up in the wrong place, it can cause serious damage to living organisms. Aging of the skin
and skin cancer are only two of many deleterious effects of light on humans and animals.
Because plants and other photosynthetic species have been dealing with light for eons,
they have had to develop photoprotective mechanisms to limit light damage. Learning
about the causes of light- induced tissue damage and the details of the natural
photoprotective mechanisms can help us can find ways to adapt these processes for the
benefit of humanity in areas far removed from photosynthesis itself. For example, the
mechanism by which sunlight absorbed by photosynthetic chlorophyll causes tissue
damage in plants has been harnessed for medical purposes. Substances related to
chlorophyll localize naturally in cancerous tumor tissue. Illumination of the tumors with
light then leads to photochemical damage which can kill the tumor while leaving
surrounding tissue unharmed. Another medical application involves using similar
chlorophyll relatives to localize in tumor tissue, and thus act as dyes which clearly
delineate the boundary between cancerous and healthy tissue. This diagnostic aid does
not cause photochemical damage to normal tissue because the principles of
photosynthesis have been used to endow it with protective agents that harmlessly convert
the absorbed light to heat.

Conclusions
The above examples illustrate the importance of photosynthesis as a natural process and
the impact that it has on all of our lives. Research into the nature of photosynthesis is
crucial because only by understanding photosynthesis can we control it, and harness its
principles for the betterment of mankind. Science has only recently developed the basic
tools and techniques needed to investigate the intricate details of photosynthesis. It is now
time to apply these tools and techniques to the problem, and to begin to reap the benefits
of this research.