PHOTOSYNTHESIS

All living organisms, from simple bacteria to complex plants and animals, require free energy
to carry out cellular processes, such as growth and reproduction. Organisms use various
strategies to capture, store, transform, and transfer free energy, including photosynthesis.
Photosynthesis allows organisms to access enormous amounts of free energy from the sun and
transform it to the chemical energy of sugars.
Although all organisms carry out some form of cellular respiration, only certain organisms,
called photoautotrophs, can perform photosynthesis. Examples of photoautotrophs include
plants, algae, some unicellular eukaryotes, and cyanobacteria. They require the presence of
chlorophyll, a specialized pigment that absorbs certain wavelengths of the visible light
spectrum to harness free energy from the sun.
Photosynthesis is a process where components of water and carbon dioxide are used to
assemble carbohydrate molecules and where oxygen waste products are released into the
atmosphere. In eukaryotes, the reactions of photosynthesis occur in chloroplasts; in
prokaryotes, such as cyanobacteria, the reactions are less localized and occur within
membranes and in the cytoplasm.
Photosynthesis is essential to all life on earth; both plants and animals
depend on it. It is the only biological process that can capture energy that
originates in outer space (sunlight) and convert it into chemical
compounds (carbohydrates) that every organism uses to power its
metabolism. In brief, the energy of sunlight is captured and used to
energize electrons, whose energy is then stored in the covalent bonds of
sugar molecules. How long lasting and stable are those covalent bonds?
The energy extracted today by the burning of coal and petroleum
products represents sunlight energy captured and stored by
photosynthesis almost 200 million years ago.
Plants, algae, and a group of bacteria called cyanobacteria are the
only organisms capable of performing photosynthesis . Because they
use light to manufacture their own food, they are called
photoautotrophs (literally, “self-feeders using light”). Other
organisms, such as animals, fungi, and most other bacteria, are
termed heterotrophs (“other feeders”), because they must rely on
the sugars produced by photosynthetic organisms for their energy
needs. A third very interesting group of bacteria synthesize sugars,
not by using sunlight’s energy, but by extracting energy from
inorganic chemical compounds; hence, they are referred to as
chemoautotrophs.
Photoautotrophs including (a) plants, (b) algae, and (c) cyanobacteria synthesize their organic
compounds via photosynthesis using sunlight as an energy source. Cyanobacteria and planktonic
algae can grow over enormous areas in water, at times completely covering the surface. In a (d) deep
sea vent, chemoautotrophs, such as these (e) thermophilic bacteria, capture energy from inorganic
compounds to produce organic compounds. The ecosystem surrounding the vents has a diverse array
of animals, such as tubeworms, crustaceans, and octopi that derive energy from the bacteria. (credit
a: modification of work by Steve Hillebrand, U.S. Fish and Wildlife Service; credit b: modification of
work by "eutrophication&hypoxia"/Flickr; credit c: modification of work by NASA; credit d: University
of Washington, NOAA; credit e: modification of work by Mark Amend, West Coast and Polar Regions
Undersea Research Center, UAF, NOAA)
The importance of photosynthesis is not just that it can capture sunlight’s
energy. A lizard sunning itself on a cold day can use the sun’s energy to
warm up. Photosynthesis is vital because it evolved as a way to store the
energy in solar radiation (the “photo-” part) as energy in the carbon-carbon
bonds of carbohydrate molecules (the “-synthesis” part). Those
carbohydrates are the energy source that heterotrophs use to power the
synthesis of ATP via respiration. Therefore, photosynthesis powers 99
percent of Earth’s ecosystems. When a top predator, such as a wolf, preys on
a deer , the wolf is at the end of an energy path that went from nuclear
reactions on the surface of the sun, to light, to photosynthesis, to
vegetation, to deer, and finally to wolf.
Photosynthesis is a multi-step process that requires sunlight, carbon dioxide
(which is low in energy), and water as substrates . After the process is
complete, it releases oxygen and produces glyceraldehyde-3-phosphate (GA3P),
simple carbohydrate molecules (which are high in energy) that can
subsequently be converted into glucose, sucrose, or any of dozens of other
sugar molecules. These sugar molecules contain energy and the energized
carbon that all living things need to survive
Photosynthesis uses solar energy, carbon
dioxide, and water to produce energy-
storing carbohydrates. Oxygen is
generated as a waste product of
photosynthesis.
Although the equation looks simple, the many steps that take
place during photosynthesis are actually quite complex.
In plants, photosynthesis generally takes place in leaves, which consist of
several layers of cells. The process of photosynthesis occurs in a middle layer
called the mesophyll.
The gas exchange of carbon dioxide and oxygen occurs through small,
regulated openings called stomata (singular: stoma), which also play roles in
the regulation of gas exchange and water balance.
The stomata are typically located on the underside of the leaf, which helps to
minimize water loss. Each stoma is flanked by guard cells that regulate the
opening and closing of the stomata by swelling or shrinking in response to
osmotic changes
In all autotrophic eukaryotes, photosynthesis takes
place inside an organelle called a chloroplast. For
plants, chloroplast containing cells exist in the
mesophyll. Chloroplasts have a double membrane
envelope (composed of an outer membrane and an
inner membrane). Within the chloroplast are stacked,
disc-shaped structures called thylakoids. Embedded in
the thylakoid membrane is chlorophyll, a pigment
(molecule that absorbs light) responsible for the initial
interaction between light and plant material, and
numerous proteins that make up the electron
transport chain. The thylakoid membrane encloses an
internal space called the thylakoid lumen. As shown in
Figure , a stack of thylakoids is called a granum, and
the liquid-filled space surrounding the granum is
called stroma or “bed” (not to be confused with
stoma or “mouth,” an opening on the leaf epidermis)
Photosynthesis takes place in two sequential stages: the light-dependent
reactions and the light independent-reactions. In the light-dependent reactions,
energy from sunlight is absorbed by chlorophyll and that energy is converted
into stored chemical energy. In the light-independent reactions, the chemical
energy harvested during the light-dependent reactions drives the assembly of
sugar molecules from carbon dioxide. Therefore, although the light-independent
reactions do not use light as a reactant, they require the products of the light-
dependent reactions to function. In addition, several enzymes of the light-
independent reactions are activated by light. The light-dependent reactions
utilize certain molecules to temporarily store the energy: These are referred to
as energy carriers. The energy carriers that move energy from light-dependent
reactions to light-independent reactions can be thought of as “full” because they
are rich in energy.
After the energy is released, the “empty” energy carriers return to the light-
dependent reaction to obtain more energy. Figure illustrates the components inside
the chloroplast where the light-dependent and light-independent reactions take
place.
Photosynthesis consists of two stages: the light-dependent reactions and
the light-independent reactions or Calvin cycle. The light-dependent
reactions occur when light is available. The overall equation for
photosynthesis shows that is it a redox reaction; carbon dioxide is
reduced and water is oxidized to produce oxygen
The light-dependent reactions occur in the thylakoid membranes of
chloroplasts, whereas the Calvin cycle occurs in the stroma of chloroplasts.
Embedded in the thylakoid membranes are two photosystems (PS I and PS II),
which are complexes of pigments that capture solar energy.
Chlorophylls a and b absorb violet, blue, and red wavelengths from the visible
light spectrum and reflect green.
The carotenoid pigments absorb violet-blue-green light and reflect yellow-to-
orange light.
Environmental factors such as day length and temperature influence which
pigments predominant at certain times of the year.
Although the two photosystems run simultaneously, it is easier to explore
them separately. Let’s begin with photosystem II.
A photon of light strikes the antenna pigments of PS II to initiate photosynthesis. In
the noncyclic pathway, PS II captures photons at a slightly higher energy level than PS
I. (Remember that shorter wavelengths of light carry more energy.) The absorbed
energy travels to the reaction center of the antenna pigment that contains chlorophyll
a and boosts chlorophyll a electrons to a higher energy level. The electrons are
accepted by a primary electron acceptor protein and then pass to the electron
transport chain also embedded in the thylakoid membrane. The energy absorbed in
PS II is enough to oxidize (split) water, releasing oxygen into the atmosphere; the
electrons released from the oxidation of water replace the electrons that were
boosted from the reaction center chlorophyll.
As the electrons from the reaction center chlorophyll pass through the series of electron carrier
proteins, hydrogen ions (H+ ) are pumped across the membrane via chemiosmosis into the interior of
the thylakoid.
This action builds up a high concentration of H+ ions, and as they flow through ATP synthase,
molecules of ATP are formed. These molecules of ATP will be used to provide free energy for the
synthesis of carbohydrate in the Calvin cycle, the second stage of photosynthesis.
The electron transport chain connects PS II and PS I.
Similar to the events occurring in PS II, this second photosystem absorbs a second photon of light,
resulting in the formation of a molecule of NADPH from NADP+. The energy carried in NADPH also is
used to power the chemical reactions of the Calvin cycle.
The 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. The carbohydrate
molecules made will have a backbone of carbon
atoms. Where does the carbon come from? It
comes from carbon dioxide, the gas that is a
waste product of respiration in microbes, fungi,
plants, and animals
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, because light is not directly required .
However, the term dark reaction can be misleading because it implies
incorrectly that the reaction only occurs at night or is independent of light,
which is why most scientists and instructors no longer use it.