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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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Photosynthesis
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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Photosynthesis
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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Photosynthesis
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
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Photosynthesis
Photosynthesis
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Photosynthesis
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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+
Photosynthesis
Photosynthesis
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Photosynthesis
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Chemiosmotic Theory
Chemiosmotic theory describes the mechanism by which ADP is phosphorylated to
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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Photosynthesis
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.
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Photosynthesis
In plants, ferredoxin is a protein accepts electrons from photosystem I and carries them to
ferredoxin-NADP+ reductase.
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+
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Photosynthesis
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.
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Photosynthesis
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.
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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Photosynthesis
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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Photosynthesis
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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Photosynthesis
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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Photosynthesis
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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Photosynthesis
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.
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Photosynthesis
Enzyme
Triose phosphate Isomerase
TI
EP
Epimerase
AL
Aldolase
IS
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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Photosynthesis
Review
Photosynthesis is the process by which water and light energy are used to fix
inorganic carbon dioxide into glucose. The complete equation is
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Photosynthesis
Photosynthesis
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