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Photosynthesis Process

The chemical equation for photosynthesis

in this chemical equation:The reactants are carbon dioxide and water


The products are glucose, oxygen, and water
Light energy is converted into chemical energy, which is stored in the glucose molecule.
For photosynthesis to take place must be present chlorophyll, enzymes and light.
1-Chlorophyll
Chlorophyll is a green pigment found inside the chloroplasts of cells. Chlorophyll traps light
energy that is used to split water molecules.

Light Absorption By Chlorophyll A

2-A coenzyme is a molecule that can transfer hydrogen and electrons from one reaction to
another. In photosynthesis, the coenzyme NADP transfers hydrogen and electrons from
light to dark reactions. In humans, many vitamins function as coenzymes.
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The formation of glucose during photosynthesis takes place in two major steps known as
the light and dark reactions. Each reaction requires enzymes. Many chemicals used to kill
plants work by blocking key enzymes that are needed in photosynthesis.

3-Light can be separated into several different wavelengths: red, orange, yellow, green, blue,
indigo, and violet (ROY G BIV). The red wavelength of light is the best for photosynthesis.
Green is the worst wavelength for photosynthesis because the leaf reflects this wavelength.
During the light reactions, light energy trapped by chlorophyll is used to split water molecules
(photolysis). When water is split, oxygen and hydrogen are produced. The oxygen molecules
exit the leaf through the stomates and enter the atmosphere. The hydrogen atoms combine
with molecules of the coenzyme NADP (nicotinamide adenine dinucleotide phosphate) to
form NADPH + H+, which is needed in the dark reactions. A coenzyme transfers hydrogen
and electrons from one reaction to another. (Reminder: The hydrogen atom has a proton and
an electron.) This movement of electrons is known as the electron transport system (ETS)

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Then, oxygen produced during photosynthesis comes from water molecules due to light
energy splits water molecules, producing oxygen and hydrogen. This process is called
photolysis.

Central role of adenosine 5_-triphosphate (ATP) in metabolism, the energy needed for the synthesis of ATP from
adenosine 5_-diphosphate (ADP) and inorganic phosphate (Pi)

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Chloroplasts are the sites where both the light-dependent and light-independent
reactions of photosynthesis occur. Chloroplasts consist of an outside phospholipid
bilayer membrane called the stroma, within the stroma are stacks of membrane
layers are thylakoids; an entire stack of thylakoids is a granum (plural, grana).
Within the thylakoids are the light-absorbing pigments and enzymes for the lightdependent reactions. In the surrounding stroma are the enzymes for the CalvinBenson cycle. Thus, the light reactions occur on the thylakoid membranes,
and the dark reactions occur in the stroma

Two photosynthetic reactions


Photosynthesis involves two chemical events, termed the light and dark reactions.
The process of photosynthesis is regulated to take place when an organism absorbs
visible light. The light reactions refer to the set of reactions in which the energy of
absorbed light is used to generate ATP and reducing power (NADPH). The dark
reactions use this reducing power and energy to fix carbon, that is, to convert
carbon dioxide to glucose. Biochemically, converting CO2 to glucose without light
is possible if a supply of reducing equivalents and ATP are available.

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Noncyclic Photophosphorylation
Photophosphorylation is the process of making ATP from ADP and Pi
(phosphorylation) using energy derived from light (photo). Noncyclic
photophosphorylation begins with PS II and follows the steps:
1. Photosystem II. Electrons trapped by P680 in photosystem II are energized by light.
two electrons are shown moving up, signifying an increase in their energy.
2. Primary electron acceptor. Two energized electrons are passed to a molecule
called the primary electron acceptor. This electron acceptor is called primary
because it is the first in a chain of electron acceptors.
3. Electron transport chain. Electrons pass through an electron transport chain. This
chain consists of proteins that pass electrons from one carrier protein to the next.
Some carrier proteins, like ferredoxin and cytochrome, include nonprotein parts
containing iron.
4. Phosphorylation. As the two electrons move down the electron transport chain,
they lose energy. The energy lost by the electrons as they pass along the electron
transport chain is used to phosphorylate, on average, about 1.5 ATP molecules.
5. Photosystem I. The electron transport chain terminates with PS I (with P700). Here
the electrons are again energized by sunlight and passed to a primary electron
acceptor (different from the one associated with PS II).
6. NADPH. The two electrons pass through a short electron transport chain. At the
end of the chain, the two electrons combine with NADP+ and H+ to form NADPH.
NADPH is a coenzyme. Since the electrons have a considerable amount of energy
left, NADPH is an energy-rich molecule.
7. Photolysis. The two electrons that originated in PS II are now incorporated into
NADPH. The loss of these two electrons from PS II is replaced when H2O is split into
two electrons, 2 H+ and 12 O2. The process is called photolysis and literally means
decomposition (lysis) by light (photo). A manganese-containing protein complex
catalyzes the reaction. The two electrons from H2O replace the lost electrons from PS
II. One of the H+ provides the H in NADPH. In summary, photophosphorylation takes
the energy in light and the electrons in H2O to make the energy-rich molecules ATP
and NADPH. Because the reactions require light, they are often called the lightdependent reactions or, simply, light reactions. The following equation informally
summarizes the process:
H2O + ADP + Pi + NADP+ + light

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ATP + NADPH + O2 + H+

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Chemiosmotic Theory
Chemiosmotic theory describes the mechanism by which ADP is phosphorylated to
ATP

1. H+ ions (protons) accumulate inside thylakoids. The light reactions of


photosynthesis occur in the membranes of the thylakoids. As photolysis occurs, H+ are
created and released to the inside of the thylakoids (the oxygen is released to the
outside). Also, H+ and electrons they pass along the electron transport chain
between PS II and PS I. These H+ come from the stroma (outside the thylakoids)
and are released to the inside of the thylakoids.
2. A pH and electrical gradient across the thylakoid membrane is created. As H+
accumulate inside the thylakoid, the pH decreases. Since some of these H+ come from
outside the thylakoids (from the stroma), the H+ concentration decreases in the
stroma and its pH increases. This creates a pH gradient consisting of differences in the
concentration of H+ across the thylakoid membrane from a stroma pH 8 to a
thylakoid pH 5. Since H+ are positively charged, their accumulation on the inside of
the thylakoid creates an electric gradient (or voltage) as well.
3. ATP synthases generate ATP. The pH and electrical gradient represent potential
energy and channel proteins, called ATP synthases, allow the H+ to flow through
the thylakoid membrane and out to the stroma. The energy generated by the
passage of the H+ (like the water through turbines in a dam) provides the energy for
the ATP synthases to phosphorylate ADP to ATP. The passage of about three H+ is
required to generate one ATP.

Light Reactions
The photosynthetic scheme begins when the pigments of the light absorb a photon of
visible light. The absorption of light excites the pigment to a higher energy state. The
pigments transfer their excitation energy to the reaction center chlorophyll molecule.
Two types of reaction centers exist, generally called Photosystem I (PSI) and
Photosystem II (PSII). Thus, two types of reactions make up photosynthesis, the
light and dark reactions while two more types of reactions compose the light reaction
those carried out by PSI and PSII. Absorption of a photon causes either type of
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chlorophyll to become more easily oxidized that is, to give up an electron. The
electron given up by either chlorophyll molecule flows through an electron transport
chain, just as the electron given up by a mitochondrial cytochrome flows through the
mitochondrial electron transport chain. Just as in mitochondrial electron transport, the
flow of electrons leads to a proton gradient, which is used to synthesize ATP. Unlike
mitochondrial electron transport, the terminal electron acceptor isnt oxygen but
rather NADP.

Photosystem II
The photosystem pigment of PSII is a form of chlorophyll termed P680, because it is
a pigment that absorbs light with a wavelength of 680 nanometers. Absorption of a
photon by P680 leads to the excited form of the pigment, called P680*. P680* but not
ground-state P680 gives up an electron to another molecule, plastiquinone.

The light-induced electron transfer in photosynthesis drives protons into the thylakoid lumen.
The excess protons flow out of the lumen through ATP synthase to generate ATP in the
stroma.

The Roles of PSI and PSII


Photosystem I (PSI) is defined as containing reaction center chlorophylls with maximal red
light absorption at 700 nm; PSI is not involved in O2 evolution.
Photosystem II (PSII) functions in O2 evolution, using reaction centers that exhibit maximal
red light absorption at 680 nm.
All photosynthetic cells contain some form of photosystem
Pathway of Electron Flow From H2O to NADP+ in Photosynthesis

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In plants, ferredoxin is a protein accepts electrons from photosystem I and carries them to
ferredoxin-NADP+ reductase.

The absorption of light by photosystem II (P680) and photosystem I (P700).


Abbreviations: Ph, pheophytin; QA and QB, plastoquinone-binding proteins; Pc,
plastocyanin; A0 and A1, acceptors of electrons from P700*; Fd, ferredoxin; Mn, manganese
Roles of the two photosystems, PSI and PSII.

From the quinones, the electron is transferred to plastocyanin and then to cytochrome
bf. The two H+ ions (protons) left behind remain in the thylakoid lumen. As the
electrons move down this electron transport chain, protons are pumped into the
thylakoid lumen. Eventually the transported electron is given up to the oxidized P700
chlorophyll of Photosystem I.
This movement of an electron to PSI from P680* leaves P680 in a non-excited,
oxidized state. Oxidized P680 must be reduced to give up another electron. Hydrogen
atoms derived from H2O reduce it according to the reaction:
2 H2O + [Manganese Center]oxidized O2 + [Manganese Center]reduced + 4 H+

This oxidation transfers four electrons to the Manganese Center, a complex


metalloprotein, which then donates the electrons through an intermediate to oxidized
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P680. The protons derived from water are transported into the thylakoid lumen. The
protons pumped into the thylakoid lumen by PSII are used to make ATP through the
action of coupling factor, in a mechanism similar to that of mitochondrial ATP
synthesis.

Photosystem I
Just as the absorption of a photon of light converts P680 to P680*, which is a better
reductant, so too does the absorption of light convert the chlorophyll of PSI to a
species that gives up an electron more easily. P700* donates an electron to a series of
mobile quinones and then to a ferredoxin protein. The ferredoxin reduces NADP to
NADPH. This provides reducing power for conversion of CO2 to carbohydrate.
2 Fdreduced + NADP+ + H+ 2 Fdoxidized + NADPH

After P700* donates the electron to the quinones, oxidized P700 accepts an electron
from the cytochrome bf, which is the acceptor of the electron from Photosystem II.
The reduced P700 is now poised to absorb a photons energy and the cycle can start
over.

The Dark Reactions


Light is needed in photosynthesis to split water. All further reactions take place in the absence
of light and are called the dark reactions. This series of cyclic reactions was discovered by
Melvin Calvin and is often called the Calvin cycle. The Calvin cycle begins when three
carbon dioxide molecules combine with three molecules of a 5-carbon compound called
ribulose diphosphate (RDP); the resulting three 6-carbon compound molecules are
unstable and break to form six 3-carbon compound molecules called phosphoglycerate
(PGA). Hydrogen from the light reactions combines with PGA to form six new 3-carbon
compounds called phosphoglyceraldehyde (PGAL). Three carbon dioxide molecules enter
at the start of the cycle to be followed by an additional three molecules of carbon dioxide.
Each turn around the cycle can spin off a PGAL to eventually form glucose. An easy way to
understand the Calvin cycle is to follow the number of carbon atoms around the cycle.

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PGAL is the true end product of photosynthesis. Molecules of PGAL can combine and
reorganize to form additional RDP molecules, or two PGAL molecules can combine to form
glucose. PGAL can also be used to produce proteins and lipids.

How can PGAL (phosphoglyceraldehyde) form RDP (ribulose diphosphate)?


The key to correctly answering this question is to keep your eye on the number of carbon
atoms. PGAL is a 3-carbon compound. Each turn around the Calvin cycle produces 6 PGAL
molecules for a total of 18 carbon atoms. Now subtract 3 carbon atoms for 1 molecule of
PGAL that spins off to eventually form glucose. This leaves us with 15 carbon atoms. These
atoms are rearranged to form 3 molecules of RDP, which is a 5-carbon compound. The total
number of carbon atoms remains the same.

Calvin-Benson Cycle
The Calvin-Benson cycle fixes CO2. The Calvin-Benson cycle must repeat six
times, and use 6 CO2 molecules
1. Carboxylation: 6 CO2 combine with 6 RuBP to produce 12 PGA. The enzyme
RuBP carboxylase, or rubisco, catalyzes the merging of CO2 and RuBP (ribulose
bisphosphate). The Calvin-Benson cycle is referred to as C3 photosynthesis because
the first product formed, PGA (phosphoglycerate), contains three carbon atoms. Other
names are the Calvin cycle and the carbon reduction cycle.
2. Reduction: 12 ATP and 12 NADPH are used to convert 12 PGA to 12 PGAL.
The energy in the ATP and NADPH molecules is incorporated into PGAL (glyceride
3-phosphate), thus making PGAL a very energy-rich molecule. ADP, Pi, and NADP+
are released and then re-energized in noncyclic photophosphorylation.
3. Regeneration: 6 ATP are used to convert 10 PGAL to 6 RuBP. Regenerating
the 6 RuBP originally used to combine with 6 CO2 allows the cycle to repeat.
4. Carbohydrate synthesis. Note that 12 PGAL were created in step 2, but only 10
were used in step 3. What happened to the remaining two? These two remaining
PGAL are used to build glucose, a common energy-storing molecule. Other
monosaccharides like fructose and maltose can also be formed. In addition, glucose
molecules can be combined to form disaccharides like sucrose and polysaccharides
like starch and cellulose
You should recognize that no light is directly used in the Calvin-Benson cycle. Thus,
these reactions are often called the light-independent reactions or even the dark
reactions. But be carefulthe process cannot occur in the dark. This is because it is
dependent upon the energy from ATP and NADPH, and these two energy-rich
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molecules can be created only during photophosphorylation, which can occur only in
light.
In summary, the Calvin-Benson cycle takes CO2 from the atmosphere and the energy
in ATP and NADPH to create a glucose molecule. Of course, the energy in ATP and
NADPH represents energy from the sun captured during photophosphorylation. The
Calvin-Benson cycle can be informally summarized as follows:
6CO2+18ATP+12NADPH + H 18ADP + 18Pi +12NADP + 1 glucose
Keep in mind that the reactions that occur during photosynthesis (and in any
biosynthetic pathway) do not occur spontaneously. Every product formed in every
reaction is catalyzed by an enzyme. In some reactions, coenzymes or metal-ion
cofactors may also be involved.

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Carbohydrate Synthesis
The dark reactions use ATP and NADPH to convert CO2 into carbohydrate. The first
step is fixing CO2 into organic carbon. The basicreaction is addition of CO2 to a
phosphorylated acceptor. This step requires no direct input of energy. Two types of
plants exist, which use different acceptor molecules. In so-called C-3 plants, the
acceptor is a 5-carbon, doubly phosphorylated acceptor, and two 3-carbon
phosphorylated compounds are formed. In C-4 plants, the acceptor is
phosphoenolpyruvate, and the carboxylation makes the 4-carbon acid oxaloacetatic
acid and releases inorganic phosphate. The ATP and NADPH from the light reactions
are used for making the acceptors and converting the first products into glucose.

C-3 Photosynthesis
This pathway is sometimes called the Calvin-Benson cycle, after the biochemists who elucidated
it. The 5-carbon, doubly phosphorylated carbohydrate, ribulose bisphosphate is the acceptor for
CO2; the enzyme is called ribulose-bisphosphate carboxylase/oxygenase (called Rubisco).

The initial product of the reaction is unstable and quickly falls apart to yield two molecules of
3-phosphoglycerate. The rest of the Calvin cycle is involved in inter conversion of carbohydrates
to make glucose (or starch) and the regeneration of the ribulose-bisphosphate acceptor. The first
step is the phosphorylation of 3-phosphoglycerate by the same reactions involved in
gluconeogenesis.

The glyceraldehyde-3-phosphate can be converted into the 6-carbon sugar phosphate called
fructose-6-phosphate by the reactions of triose phosphate isomerase, aldolase, and fructose
bisphosphase.

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Note that the last step is irreversible, just as it is in gluconeogenesis. In shorthand, the
reactions to this point are: 5 + 1 = 3 + 3 = 6
Regeneration of ribulose bisphosphate occurs by the same reactions that occur in the
hexose-monophosphate shunt. The hexose monophosphate shunt interconverts 3-, 4-,
5-, 6- and 7-carbon sugar phosphates. In the formation of one molecule of glucose
from CO2, carrying out the previous reactions six times is necessary. Making the six
molecules of ribulose-bisphosphate required for fixing six CO2 occurs by the
following sets of reactions:Reaction 1: Transketolase transfers a 2-carbon unit

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Reaction 2: Transaldolase transfers a 3-carbon unit

Reaction 3: Transketolase transfers a 2-carbon unit

Remembering all these reactions can be difficult. A useful mnemonic exists to help you keep track
of the number of carbons in each of the three reactions:

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The 5-carbon sugar phosphates are interconverted by the action of epimerase and
isomerase to yield ribulose-5-phosphate, which is phosphorylated by the enzyme
ribulose phosphate kinase to make RuBP, the acceptor of CO2. Ribulose phosphate
kinase is active only when a cystine disulfide on the enzyme is reduced to two
cysteines. An electron carrier, thioredoxin, carries out this reduction, and is then itself
reduced by electrons from NADPH. Because the action of Photosystems I and II
forms NADPH, this reduction ensures that ribulose bisphosphate is made only when
enough light exists to support Photosynthesis. In other words, the light and dark
reactions are coupled.

Summary of Calvin cycle

Glyceraldehyde-3-P may be converted to other CHO metabolites (e.g., fructose-6-P,


glucose-1-P), energy stores (e.g., sucrose, starch), or cell wall constituents (e.g.,
cellulose). Plant cells can use glyceraldehyde-3-P as Csource for synthesis of fatty
acids & amino acids.

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Enzyme
Triose phosphate Isomerase
TI

EP

Epimerase

AL

Aldolase

IS

Isomerase Ribose Phosphate

FB

Fructose-1,6-bisphosphatase

PK

Phospho-ribulokinase

SB

Sedoheptulose-Bisphosphatase

TK

Transketolase

The free energy of cleavage of ~P bonds of ATP, and reducing power of NADPH,
are used to fix and reduce CO2 to form carbohydrate. Enzymes and intermediates of
the Calvin Cycle are located in the chloroplast stroma.

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Review
Photosynthesis is the process by which water and light energy are used to fix
inorganic carbon dioxide into glucose. The complete equation is

Various pigments exist in the photosynthetic membranes of plants. These include


chlorophyll a and b and various carotenoids. The purpose of a variety of pigments is
to absorb light energy (photons) of different wavelengths. The absorption spectrums
of the pigments overlap so that they maximize the energy absorbed. When light
energy is absorbed, the pigment becomes energized.
The energy is then bounced around among pigments until it is absorbed by either of
two special kinds of chlorophyll a, P700 and P680.
P700 and P680 are organized into separate photosystems (pigment systems) I and II.
Cyclic photophosphorylation involves photosystem I. This process is a more primitive
form of photosynthesis. In cyclic photophosphorylation, the energy absorbed by the
pigment system is captured by the P700. As a result, two electrons are excited to a
higher energy level, where they are absorbed by a primary electron acceptor. From
here, the electrons are passed through an electron transport chain from electron
acceptor to electron acceptor (some of which are cytochromes).
During this transit, the energy loss of the electrons is used to bond a phosphate group
to ADP, making it ATP (adenosine triphosphate). The process is called
phosphorylation, and the result is that energy (originally from light) from the two
electrons is trapped in an ATP molecule. The cycle is completed when the two
electrons return to the photosystem pigments.
Noncyclic photophosphorylation is a more advanced system involving both
photosystems I and II. It begins when chlorophyll P680 in photosystem II traps photon
energy and energizes two electrons. These two electrons are passed to a primary
electron acceptor, then through an electron transport chain producing on average, 1.5
ATP, and are then finally returned to photosystem I. Here the electrons are energized
again (by light), and are received by another primary electron acceptor. These two
electrons then combine with 2 H+ to form NADPH.
Meanwhile, a water molecule is split (photolysis) producing 2 electrons, 2 H+ and
12 O2. The 2 electrons replace the electrons energized from photosystem II. The
oxygen is released. One of the H+ is used to make the NADPH + H+.
The dark reaction (Calvin-Benson cycle) combines CO2, NADPH, and ATP to form
PGAL and RuBP. It takes 6 CO2 to create 2 PGAL, so the cycle repeats 6 times to
produce 2 PGAL. The 2 PGAL form glucose. Glucose can then be used to make
various other carbohydrates such as sucrose and starch, or it can be broken down to
release its store of energy in the form of ATP to drive metabolic activities.

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