Photosynthesis

Photosynthesis is a process by which plants use solar energy to make food molecules
from CO2 and water. It is the most important chemical process on earth for it provides
food supply for virtually all organisms.
Plant protoplasts capture light energy and convert it to chemical energy that is
stored in glucose and other organic molecules.
Photosynthesis is given by the following equation, which is a simplified version of a
very complex process.

Photosynthesis consists of two reactions, each with multiple steps. These are the
light-dependent and dark-dependent reactions. Light-dependent reactions occur in
the presence of light and generate adenosine triphosphate (ATP) and nicotinamide
adenine dinucleotide phosphate (NADPH). Dark-dependent reactions do not require
light but use ATP and NADPH generated during the light reactions to fix carbon dioxide
and make carbohydrates.
Photosynthetic pigments
When light strikes matter, it can be absorbed, radiated or reflected. Substances that
absorb visible light are called pigments. Different pigments absorb light of different
wavelengths. If a pigment absorbs all the wavelengths of light, we see black.
However, if a pigment absorbs all the other colours and reflects yellow or green, we
see the colour that is reflected. For example, we see green when we look at a leaf
because chlorophyll absorbs red and blue light while reflecting green light.

Absorption spectra of chlorophyll a and b

The absorption spectrum of chlorophyll a (the form of chlorophyll most important in
photosynthesis) has two peaks, corresponding to wavelengths for blue and red light
(the colours chlorophyll a absorbs best). Chlorophyll a can participate directly in the
light reactions that convert solar energy to chemical energy. Other pigments can
absorb light and transmit energy to chlorophyll a. One of these is chlorophyll b, which
is almost identical to chlorophyll a, except for a small difference. Chlorophyll a is
grass-green, whereas chlorophyll b is yellow-green.
Other pigments
The chloroplast has other pigments. These are carotenoids, which are of various
shades of yellow and orange. Chlorophyll a, chlorophyll b and the carotenoids are
found in the thylakoid membrane. The location of the chlorophyll a in the pigment is
called the reaction centre. The other chlorophyll molecules and the molecules of
chlorophyll b and the carotenoids collectively function as light gathering antennae
that absorb photons (light energy) and pass them to the reaction centre. For example,
if a photon of light strikes a carotenoid or chlorophyll b, the energy is conveyed to
chlorophyll a.

Photosynthesis takes place entirely within chloroplasts. Like mitochondria,

chloroplasts have a double membrane: an inner membrane and an outer membrane.
The inner membrane, which is relatively smooth, encloses a fluid matrix called the
stroma. In addition, chloroplasts have a third membrane called the thylakoid
membrane. This is folded into thin vesicles (the thylakoids), enclosing small spaces
called the thylakoid lumen. The thylakoid vesicles are often layered in stacks called
grana (single, granum).

Structure of chloroplast
Photosystems
Photosystems are light harnessing units in the thylakoid membrane. A photosystem
consists of an antenna complex, a reaction centre and a primary acceptor of
energized electrons. Two types of photosystems are found in the thylakoid membrane:
photosystem I and photosystem II, in order of their discovery. At the reaction centre
of photosystem 1, is a specialized chlorophyll a molecule called P700, so called
because the light the pigment absorbs best has a wavelength of 700 nm, which is in
the far-red part of the spectrum. At the reaction centre of photosystem II is a

specialized chlorophyll a molecule designated P680, with a peak absorption spectrum

of 680 nm, also in the red part of the spectrum.

Photosystem I and Photosystem II

Light-dependent reactions
Light reactions take place in the light and in the thylakoid membrane of chloroplasts.
The key reactions of the light reactions are (1) the absorption of light energy (2) the
excitation of electrons and (3) the formation of ATP and NADPH. There are two
possible routes for light reactions: cyclic and noncyclic routes. Both of these reactions
are also known as photophosphorylation because they use light energy to
phosphorylate (add inorganic phosphate) to a molecule of ADP to obtain ATP.
Cyclic Photophosphorylation
In cyclic photophosphorylation, light energy is trapped by pigment molecules in the
P700 antenna system and the excited state eventually reaches the reaction centre
P700. Energized electrons from P700 move from one acceptor molecule to the next
releasing free energy at each step and eventually returning to the P700 molecule.
Some of the energy released as the electron is eased down the energy gradient is used
by the cell to transport H+ ions across the thylakoid membrane creating an H+ ion
gradient. The energy of this gradient is used to phosphorylate ADP into ATP. Because
the electrons are returned to the chlorophyll molecules from which they originated,
this process is termed cyclic photophosphorylation. ATP is the only product of this
process.
Noncyclic Photophosphorylation
Noncyclic photophosphorylation involves both photosystem I
and photosystem II. When a photon energy is received by P680, electrons become
highly excited so that a few actually leave the photosystem II molecule, leaving

holes in reaction centre P680. These empty spaces (holes) are then filled by
electrons coming from two molecules of water split in a process called photolysis.
Oxygen produced by photolysis diffuses out of the cell and escapes into the air
through the leaf stomata. This is the oxygen that sustains life on earth.
2H20 4H+ +4e- + O2
The electrons are received by P700, the reaction centre for photosystem I. However,
prior to their arrival, the pigment complex in reaction centre P700 has absorbed a
photon of light energizing its electrons some of which are captured by an acceptor
molecule. The resulting vacancy in P700 can now be filled by electrons from
photosystem II. The excited electron of photosystem I reduces nicotinamide adenine
dinucleotide phosphate (NADP+) to NADPH.
Therefore, light reactions are concerned with (a) the production of oxygen from
water and (2) formation of energy rich compounds NADPH and ATP needed during the
synthesis of carbohydrates from CO2 in the dark reactions.
The Dark Reactions
Dark reactions (also known as the Calvin cycle) are involved in the final stage of
photosynthesis, which is the synthesis of sugar, using CO 2, ATP, and hydrogen atoms
from NADPH. These reactions occur in the stroma of the chloroplast and do not
require light. Carbon dioxide from the atmosphere diffuses into the leaf through the
stomata and enters the stroma of the chloroplast. It then binds with a 5-carbon sugar
called ribulose bisphosphate (RuDP), present in the leaf. The reaction is catalyzed by
an enzyme called rubisco carboxylate. This enzyme is present in large quantities in
the stroma of chloroplasts.
The product is unstable 6-carbon compound, which immediately splits into two
molecules of a 3-carbon compound called phosphoglyceric acid (PGA. Each PGA is
phosphorylated by ATP and reduced by hydrogen from NADP. The resulting energy-rich
3-carbon compound is called phosphoglyceraldehyde (PGAL). This is a stable sugar
and,
in a way, forms the final product of photosynthesis.
PGAL is a 3-carbon sugar and this is the reason photosynthesis is sometimes referred
to as a C3 photosynthesis. Three carbon compounds like PGAL are the simplest
carbohydrates that combine to form larger, more complex monosaccharides such as
glucose and fructose. Glucose molecules in turn combine to form the storage
carbohydrate starch. To form the 6-carbon molecule of glucose, takes six
CO2 molecules and six turns of the Calvin cycle.
Leaves as Organs of Photosynthesis

Photosynthesis can occur in all green parts of a plant but in most plants the leaf is
the main photosynthetic part. The structure of a dicot leaf consists of a stalk or
petiole, which attaches the leaf to the plant and a flat, broad blade. The blade
contains a complex system of veins, which serve as transport channels for water and
nutrients. Because of the flatness of the blade, the leaf exposes a large surface area
to sunlight.
A transverse section of the leaf shows that the outer surface is formed of epidermal
layers, usually one but sometimes two or more layers thick. A waxy cuticle covers the
outer surfaces of both the upper and lower epidermal layers. The chief function of
the epidermis is the protection of the internal tissues of the leaf from excessive loss
of water, invasion by fungi and mechanical injury. Most epidermal cells do not contain
chloroplasts. The cuticle is transparent and permits light to pass through to the inner
cells of the leaf.

Internal structure of a leaf

The region between the upper and lower epidermis is called the mesophyll and is usually
divided into an upper palisade mesophyll consisting of cells arranged vertically, and a
spongy mesophyll made up of irregularly shaped, loosely packed cells with air spaces
between them. The air spaces communicate with the outside via tiny pores on the
leaf called stomata (singular, stoma). The palisade cells are filled with chloroplasts
and it in palisade cells where most of the photosynthesis occurs. The spongy layer
cells contain chloroplasts but it seems their major function is temporary storage of
food substances manufactured by the leaves.
A system of veins known as vascular bundles branch from a central vein to the rest of
the blade forming a structural framework for the leaf and serving as transport

channels. Each vein contains xylem and phloem vascular tissues, and each is usually
surrounded by a bundle sheath, made up of very tightly packed cells that leave very
little space between them.
Adaptation to Water Conservation
C3, C4 and CAM are three types of photosynthetic pathways that are adapted to water
conservation. C3 photosynthesis is the typical photosynthesis that most plants use and
that most of us learn about at school. C4 and CAM photosynthesis are adaptations to
arid conditions because they result in better water conservation by plants.
C3 plants
Most plants use the C3 pathway to incorporate CO2 into the Krebs cycle during the
making of sugar compounds. These plants incorporate CO2 from the air directly into
the Calvin cycle. They derive their name from the fact that the first organic
compound produced is the 3-carbon compound, PGA. They require moderate sunlight
intensity, moderate temperatures, high CO2 concentration, about 200 ppm or more
and plenty of ground water. C3 plants open their stomata during the day. However,
during a very dry, hot day, they close the stomata to prevent water loss and avoid
wilting. Another characteristic of these plants is the fact that photosynthesis takes
place throughout the leaf.
One of the problems of growing C3 plants is that dry weather can reduce the rate of
photosynthesis and decrease crop productivity. On a dry, hot day, a C 3 plant closes its
stomata. This adaptation at water conservation prevents CO2 from entering the leaf
and O2 from leaving the leaf. This can lead to very low levels of CO2 and high
concentrations of O2 in the leaf.
When this happens, the first enzyme of the Calvin cycle, called rubisco, incorporates
O2 instead of CO2 and the Calvin cycle produces a 2-C compound called
phosphoglycolate instead of its usual 3-C product. The plant cell then breaks the 2-C
compound down to CO2 and water. This process is called photorespiration. Unlike
photosynthesis, photorespiration is wastage of energy because it produces no sugar
molecules. Unlike cellular respiration, it does not produce ATP.
C3 plants include soyabeans, oats, wheat and rice.
C4 Plants
C4 plants represent about 5% of the worlds plant biomass. They are called C4 plants
because CO2 is first incorporated into a 4-carbon compound. C4 plants have developed
a mechanism that delivers CO2 to the rubisco enzyme efficiently. They use their
specific leaf anatomy called Kranz Anatomy, where chloroplasts exist not only in the
mesophyll cells in the outer part of their leaves but also in the bundle sheath cells as

well. Instead of the direct fixation in the Calvin cycle, CO2 is converted to a 4-carbon
organic acid that has the ability to regenerate CO2 in the chloroplasts of the bundle
sheath cells. Bundle sheath cells can then utilise this CO2 to generate carbohydrates
by the usual C3 pathway.
Examples of C4 plants include corn, sugarcane, sorghum and finger millet.
CAM Plants
CAM plants are called after the plant family in which they were first found
(Crassulaceae) and because CO2 is stored in form of acid before use in
photosynthesis. Characteristics of CAM plants include:

Stomata open at night when evaporation is low to prevent water loss and
remain closed during the day

CO2 is converted to an acid and stored during the night. During the day, the
acid is broken down and CO2 is released to rubisco for photosynthesis.
Significance of CAM metabolism is that the use of water is more efficient than in C 3
plants under hot dry conditions. This is because the stomata open only at night when
the transpiration is at its lowest and remain closed during the day when transpiration
is highest.
However, under very hot, dry conditions, CAM plants keep their stomata closed day
and night. Oxygen given off in photosynthesis is used for respiration and CO 2 produced
by respiration is used for photosynthesis. This CAM idling as it is called allows the
plant to survive harsh dry spells and recover quickly once water becomes available.
CAM plants often display xerophytic (water conservation) characteristics e.g. thick,
reduced leaves with a low surface area, thick cuticle and stomata sunken into pits.
Some shed leaves during the dry season; others (succulents) store water in vacuoles.
They also use N2 very efficiently. Because the stomata are closed during the day, they
are less efficient at absorbing CO2.
C4 and CAM plants have competitive advantage over C3 plants under conditions of
drought, high temperatures, and limited nitrogen or carbon dioxide. Both have
overcome the tendency of rubisco (the first enzyme in the Calvin cycle) to
photorespire or waste energy by using O2 to break down carbon compounds to CO2.
However, C4 fixation still uses more energy in form of ATP than C3 fixation.