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Bis2A 5.7: Photosynthesis and the
∗
Calvin Cycle
The BIS2A Team
This work is produced by OpenStax-CNX and licensed under the
Creative Commons Attribution License 4.0†
Abstract
This module will discuss how light energy is transformed into chemical energy in photosynthetic
organisms. The chemical energy, in the form of ATP and high energy electrons in NADH, is then used
to reduce carbon dioxide in a series of chemical reactions in the Calvin Cycle to produce sugars.
1 Section Summary
The light dependent reactions of photosynthesis couple the transfer of energy in light into chemical compounds through a series of redox reactions in an electron transport chain (review module 5.6). In the light
dependent reactions, both ATP and NADPH are generated. Using the energy carriers formed in the rst
steps of photosynthesis, the light-independent reactions, or the Calvin cycle, take in CO
2
from the environ-
2 and
ment and incorporate it into larger biomolecules. An enzyme, RuBisCO, catalyzes a reaction with CO
another molecule, RuBP to produce two three carbon sugars. This process is called carbon xation. After
three cycles, a three-carbon molecule of glyceraldehyde-3-phosphate (G3P), the same one we saw earlier in
glycolysis, leaves the cycle to become part of a carbohydrate molecule. The remaining G3P molecules stay in
2
the cycle to be regenerated into RuBP, which is then ready to react with more CO . Photosynthesis forms
an energy cycle with the process of cellular respiration.
Plants need both photosynthesis and respiration
for their ability to function in both the light and dark, and to be able to interconvert essential metabolites.
Therefore, plants contain both chloroplasts and mitochondria - more on these organelles soon.
2 Light Energy
The sun emits an enormous amount of electromagnetic radiation (solar energy). Scientists dierentiate the
various types of radiant energy from the sun within the electromagnetic spectrum. The
spectrum
electromagnetic
is the range of all possible frequencies of radiation. The human eye can only perceive a small
fraction of this energy and this portion of the electromagnetic spectrum is therefore referred to as visible
light.
Visible light constitutes only one of the many types of electromagnetic radiation emitted from the
sun and other stars.
∗ Version
1.1: Jan 27, 2016 10:52 pm -0600
† http://creativecommons.org/licenses/by/4.0/
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Figure 1: The sun emits energy in the form of electromagnetic radiation. This radiation exists at
dierent wavelengths, each of which has its own characteristic energy. All electromagnetic radiation,
including visible light, is characterized by its wavelength. Each type of electromagnetic radiation travels
at a particular wavelength. The longer the wavelength (or the more stretched out it appears in the
diagram), the less energy it carries. Short, tight waves carry the most energy. This may seem illogical,
but think of it in terms of a piece of moving a heavy rope. It takes little eort by a person to move
a rope in long, wide waves. To make a rope move in short, tight waves, a person would need to apply
signicantly more energy.
2.1 Light Absorption
Light energy initiates many light dependent biological process when pigments absorb photons of light. Organic pigments, whether in the human retina, chloroplast thylakoid, or microbial membrane often have
specic ranges of energy levels or wavelengths that they can absorb that are dependent on their chemical
makeup and structure. A pigment like the retinal in our eyes, when coupled with an opsin sensor protein,
sees (absorbs) light predominantly with wavelengths between 700 nm and 400 nm.
Because this range
denes the physical limits for what light we can see, we refer to it, as noted above, as the "visible range".
For similar reasons, plants pigment molecules absorb only light in the wavelength range of 700 nm to 400
nm; plant physiologists refer to this range for plants as photosynthetically active radiation.
The visible light seen by humans as white light is composed of a rainbow of colors, each with a characteristic wavelength. Certain objects, such as a prism or a drop of water, disperse white light to reveal the colors
to the human eye. In the visible spectrum, violet and blue light have shorter (higher energy) wavelengths
while the orange and red light have longer (lower energy) wavelengths.(Figure 2).
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Figure 2: The colors of visible light do not carry the same amount of energy. Violet has the shortest
wavelength and therefore carries the most energy, whereas red has the longest wavelength and carries
the least amount of energy. (credit: modication of work by NASA)
2.1.1 Understanding Pigments
Chlorophylls (including bacteriochlorophylls) and carotenoids are the two major classes of photosynthetic
lipid derived pigments found in bacteria, plants and algae; each class has multiple types of pigment molecules.
There are ve major chlorophylls:
a, b, c, d,
and
f.
Chlorophyll
a
is related to a class of more ancient
molecules found in bacteria called
bacteriochlorophylls.
found in bacteria and eukaryotes.
They are the red/orange/yellow pigments found in nature.
Carotenoids are also very ancient molecules,
They are
found in fruitsuch as the red of tomato (lycopene), the yellow of corn seeds (zeaxanthin), or the orange of
an orange peel (β -carotene)which are used as advertisements to attract seed dispersers (animals or insects
that may carry seeds elsewhere). In photosynthesis,
carotenoids function as photosynthetic pigments.
In
addition, when a leaf is exposed to full sun, that surface is required to process an enormous amount of
energy; if that energy is not handled properly, it can do signicant damage. Therefore, many carotenoids
help absorb excess energy in light and safely dissipate that energy as heat.
Each type of pigment can be identied by the specic pattern of wavelengths it absorbs from visible light,
which is the
a,
absorption spectrum.
chlorophyll
b,
The graph in Figure 3 shows the absorption spectra for chlorophyll
and a type of carotenoid pigment called
β -carotene
(which absorbs blue and green light).
Notice how each pigment has a distinct set of peaks and troughs, revealing a highly specic pattern of
absorption. Chlorophyll
a
absorbs wavelengths from either end of the visible spectrum (blue and red), but
not green. Because green is reected or transmitted, chlorophyll appears green. Carotenoids absorb in the
short-wavelength blue region, and reect the longer yellow, red, and orange wavelengths.
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Figure 3: (a) Chlorophyll a, (b) chlorophyll b, and (c) β -carotene are hydrophobic organic pigments
found in the thylakoid membrane. Chlorophyll a and b, which are identical except for the part indicated
in the red box, are responsible for the green color of leaves. β -carotene is responsible for the orange color
in carrots. Each pigment has (d) a unique absorbance spectrum.
Many photosynthetic organisms have a mixture of pigments which optimizes the organism's ability to
absorb energy from a wider range of wavelengths.
sunlight.
Not all photosynthetic organisms have full access to
Some organisms grow underwater where light intensity and available wavelengths decrease and
change, respectively, with depth. Other organisms grow in competition for light. Plants on the rainforest
oor must be able to absorb any bit of light that comes through, because the taller trees absorb most of the
sunlight and scatter the remaining solar radiation (Figure 4).
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Figure 4: Plants that commonly grow in the shade have adapted to low levels of light by changing the
relative concentrations of their chlorophyll pigments. (credit: Jason Hollinger)
2.1.2 Photophosphorylation an Overview:
Photophosphorylation is the process of transferring
the energy from light into chemicals, in particular
ATP and NADPH. The evolutionary roots of photophosphorylation are likely in the anaerobic world, between
3 billion and 1.5 billion years ago, when life was abundant in the absence of molecular oxygen. Photophosphorylation probably evolved relatively shortly after electron transport chains and
anaerobic respiration
began to provide metabolic diversity. The rst step of the process involves the absorption of a photon by a
pigment molecule. Light energy is transferred to the pigment and promotes electrons into a higher potential
energy state - termed an "excited state". The electrons are colloquially said to be "energized". In the excited
state, the pigment now has a very low reduction potential and can donate the "excited" electrons to other
carriers with greater reduction potentials.
These electron acceptors may in turn become donors to other
molecules with greater reduction potentials and in so doing form an electron transport chain. As electrons
pass from one electron carrier to another via red/ox reactions, these exergonic transfers can be coupled to
the endergonic transport (or pumping) of protons across a membrane to create an electrochemical gradient.
This electrochemical gradient generates a proton motive force whose exergonic drive to reach equilibrium
can be coupled to the endergonic production of ATP, via ATP synthase. As we will seen in more detail,
the electrons involved in this electron transport chain can have one of two fates: (1) they may be returned
to their initial source in a process called cyclic photophosphorylation or (2) they can be deposited onto a
close relative of NAD
+
+
called NADP .
If the electrons are deposited back on the original pigment in a
cyclic process, the whole process can start over. If, however, the electron is deposited onto NADP
+
to form
NADPH (**shortcut note - we didn't explicitly mention any protons but assume it is understood that they
are also involved**) the original pigment must regain an electron from somewhere else. This electron must
come from a source with a smaller reduction potential than the oxidized pigment and depending on the
2
2
system there are dierent possible sources, including H O, reduced sulfur compounds such as SH
0
elemental S .
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2.2 What happens when a compound absorbs a photon of light?
When a compound absorbs a photon of light, the compound is said to leave its ground state and become
"excited", in the sense that it has this extra energy. This is illustrated in gure 5 schematically.
Figure 5: A diagram of what happens to a molecule that absorbs a photon of light.
What are the fates of the "excited" electron? There are four possible outcomes, which are schematically
diagrammed in Figure 6 below. These options are:
1. The electron can relax to a lower orbital, transferring energy as heat.
2. The electron can relax to a lower orbital and transfer energy into a photon of light - a process known
as uorescence .
3. The energy can be transferred by resonance to a neighboring molecule as the e
-
returns to a lower
orbital.
this excited e donor to a proper e acceptor can lead to an exergonic electron transfer. In other words,
4. The energy can change the reduction potential such that the molecule can become an e donor. Linking
the excited state can be involved in Red/Ox reactions.
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Figure 6: What can happen to the energy absorbed by a molecule
As the excited electron decays back to its original orbit, the energy can be transferred in a variety of ways.
While many so called antenna or auxiliary pigments absorb light energy and transfer it to something known
as a reaction center (by mechanisms depicted in option III in gure 6) it is what happens at the reaction
center that we are most concerned with (option IV in gure 6). Here a chlorophyll or bacteriochlorophyll
molecule absorbs a photon's energy and an electron is excited.
This energy transfer is sucient to allow
the reaction center to donate the electron in a red/ox reaction to a second molecule.
photophosphorylation electron transport reactions.
This initiates the
The result is an oxidized reaction center that must
now be reduced in order to start the process again.
How this happens is the basis of electron ow in
photophosphorylation and will be described in detail below.
2.3 Simple Photophosphorylation Systems: Anoxygenic photophosphorylation
Introduction
Early in the evolution of photophosphorylation, these reactions evolved in anaerobic environments where
there was very little molecular oxygen available. Two sets of reactions evolved under these conditions, both
directly from anaerobic respiratory chains as described above.
These are known as the
light reactions
because they require the activation of an electron (an excited electron) from the absorption of a photon of
light by a reaction center pigment, such as bacteriochlorophyll. The light reactions are categorized either
as
cyclic or as noncyclic photophosphorylation, depending upon the nal state of the electron(s) removed
from the reaction center pigments. If the electron(s) return to the original pigment reaction center, such
as bacteriochlorophyll, this is cyclic photophosphorylation; the electrons make a complete circuit and is
+ to NADPH, the electron(s) are removed
diagramed in gure 8. If the electron(s) are used to reduce NADP
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from the pathway and end up on NADPH, this process is referred to as noncyclic since the electrons are no
longer part of the circuit. In this case the reaction center must be re-reduced before the process can happen
again. Therefore, an external electron source is required for noncylic photophosphorylation. In these systems
2
reduced forms of Sulfur, such as H S, which can be used as an electron donor and is diagrammed in gure 9.
To help you better understand the similarities of photophosphorylation to respiration, a redox tower (gure
7) has been provided that contains many commonly used compounds involved with photosphosphorylation.
Figure 7: Electron tower that has a variety of common photophosphorylation components. PSI and
PSII refer to Photosystems I and II of the oxygenic photophosphorylation pathways. For the examples
in Figure 8 and Figure 9 P840 is similar in reduction potential as is PSI.
Cyclic Photophosphorylation
In cyclic photophosphorylation the bacteriochlorophyll
ox .
and eject an electron to form bacteriochlorophyll
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red
molecule absorbs enough light energy to energize
The electron reduces a carrier molecule in the reac-
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tion center which in turn reduces a series of carriers via red/ox reactions.
carriers found in respiration.
+
suciently large, protons, H
ox
duce bacteriochlorophyll
These carriers are the same
If the change in reduction potential from the various red/ox reactions are
are translocated across the membrane. Eventually the electron is used to re-
and the whole process can start again. This is called cyclic photophosphorylation
because the electrons make a complete circuit: bacteriochlorophyll is the source of electrons and is the nal
electron acceptor. ATP is produced via the
F1 F0 ATPase.
The schematic in gure 8 below demonstrates
how cyclic photophosphorylation works.
Figure 8: Cyclic Photophosphorylation. The reaction center P840 absorbs light energy and becomes
excited, denoted with an *. The excited electron is ejected and used to reduce an FeS protein leaving an
oxidized reaction center. The electron its transferred to a quinone, then to a series of cytochromes which
in term is reduces the P840 reaction center. The process is cyclical. Note the gray array coming from the
FeS protein going to a ferridoxin (Fd), also in gray. This represents an alternative pathway the electron
can take and will be discussed below in non-cyclic photophosphorylation. NOTE the same electron that
leaves the P480 reaction center is not necessarily the same electron that eventually nds its way back to
reduce the oxidized P840.
note: The gure of cyclic photophosphorylation above depicts the ow of electrons in a respiratory
chain. How does this process help generate ATP?
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Non-cyclic photophosphorylation
In cyclic photophosphorylation electrons cycle from bacteriochlorophyll (or chlorophyll) to a series of electron
carriers and eventually back to bacteriochlorophyll (or chlorophyll): there is no loss of electrons, they stay
in the system.
In non-cyclic photophosphorylation the electrons are removed from the photosystem and
redox chain and they eventually end up on NADPH. That means there needs to be a source of electrons,
a source that has a smaller reduction potential than bacteriochlorophyll (or chlorophyll) that can donate
electrons to bacteriochlorophyll
ox
to reduce it.
An electron tower is proved above so you can see what
compounds can be used to reduce the oxidized form of bacteriochlorophyll. The second requirement, is that
when bacteriochlorophyll becomes oxidized and the electron is ejected it must reduce a carrier that has a
greater reduction potential than NADP/NADPH (see the electron tower). In this case, electrons can ow
from energized bacteriochlorophyll to NADP forming NADPH and oxidized bacteriochlorophyll. Electrons
are lost from the system and end up on NADPH, to complete the circuit bacteriochlorophyll
2
0
an external electron donor, such as H S or elemental S .
ox is reduced by
Figure 9: Non-cyclic photophosphorylation. In this example, the P840 reaction center absorbs light
energy and becomes energized, the emitted electron reduced a FeS protein and in turn reduces ferridoxin.
Reduced ferridoxin (Fdred ) can now reduce NADP to form NADPH. The electrons are now removed from
the system, nding their way to NADPH. The electrons need to be replaced on P840, which requires an
external electron donor. In this case, H2 S serves as the electron donor.
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note: It should be noted that per electron donated to the system, either NADPH or ATP is made,
but not both. This puts a limit on the versatility of the system. But what would happen if both
systems coexisted simultaneously? That is, if both ATP and NADPH could be formed from one
electron? Additionally, these systems require compounds such as reduced sulfur to act as electron
ox
donors, not necessarily widely found compounds. What would happen if a chlorophyll
2
2
had a reduction potential that was less than that of molecular O /H O?
molecule
3 Oxygenic Photophosphorylation
Generation of NADPH and ATP
The overall function of light-dependent reactions is to transfer solar energy into chemical compounds, largely
the the molecules NADPH and ATP. This energy supports the light-independent reactions and fuels the
assembly of sugar molecules. The light-dependent reactions are depicted in Figure 10. Protein complexes
and pigment molecules work together to produce NADPH and ATP.
note: Step back a little. Why is it a reasonable goal to want to make NADPH and ATP? In the
discussion of glycolysis and the TCA cycle the goal was to make ATP and NADH. What is the key
dierence? Perhaps how those molecules will be used? Something else?
Figure 10: A photosystem consists of a light-harvesting complex and a reaction center. Pigments in the
light-harvesting complex pass light energy to two special chlorophyll a molecules in the reaction center.
The light excites an electron from the chlorophyll a pair, which passes to the primary electron acceptor.
The excited electron must then be replaced. In (a) photosystem II, the electron comes from the splitting
of water, which releases oxygen as a waste product. In (b) photosystem I, the electron comes from the
chloroplast electron transport chain discussed below.
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The actual step that transfers light energy into the biomolecule takes place in a multiprotein complex called
a
photosystem, two types of which are found embedded in the thylakoid membrane, photosystem II
photosystem I (PSI) (Figure 11). The two complexes dier on the basis of what they oxidize
(PSII) and
(that is, the source of the low-energy electron supply) and what they reduce (the place to which they deliver
their energized electrons).
antenna proteins to which the chlororeaction center where the photochemistry takes place. Each
photosystem is serviced by the light-harvesting complex, which passes energy from sunlight to the reacBoth photosystems have the same basic structure; a number of
phyll molecules are bound surround the
tion center; it consists of multiple antenna proteins that contain a mixture of 300400 chlorophyll
molecules as well as other pigments like carotenoids. The absorption of a single
a
and
b
photon or distinct quantity
or packet of light by any of the chlorophylls pushes that molecule into an excited state. In short, the light
energy has now been captured by biological molecules but is not stored in any useful form yet. The energy
is transferred from chlorophyll to chlorophyll until eventually (after about a millionth of a second), it is
delivered to the reaction center. Up to this point, only energy has been transferred between molecules, not
electrons.
:
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Figure 11: In the photosystem II (PSII) reaction center, energy from sunlight is used to extract electrons
from water. The electrons travel through the chloroplast electron transport chain to photosystem I (PSI),
which reduces NADP+ to NADPH. The electron transport chain moves protons across the thylakoid
membrane into the lumen. At the same time, splitting of water adds protons to the lumen, and reduction
of NADPH removes protons from the stroma. The net result is a low pH in the thylakoid lumen, and a
high pH in the stroma. ATP synthase uses this electrochemical gradient to make ATP.
The reaction center contains a pair of chlorophyll
a molecules with a special property.
Those two chlorophylls
can undergo oxidation upon excitation; they can actually give up an electron in a process called a
activation.
photo
It is at this step in the reaction center, this step in photosynthesis, that light energy is transferred
into an excited electron. All of the subsequent steps involve getting that electron onto the energy carrier
NADPH for delivery to the Calvin cycle where the electron is deposited onto carbon for long-term storage
in the form of a carbohydrate.
transport chain,
PSII and PSI are two major components of the photosynthetic
which also includes the
cytochrome complex.
electron
The cytochrome complex, an enzyme
composed of two protein complexes, transfers the electrons from the carrier molecule plastoquinone (Pq) to
the protein plastocyanin (Pc), thus enabling both the transfer of protons across the thylakoid membrane and
the transfer of electrons from PSII to PSI.
The reaction center of PSII (called P680) delivers its high-energy electrons, one at a time, to the primary
electron acceptor, and through the electron transport chain (Pq to cytochrome complex to plastocyanine)
to PSI. P680's missing electron is replaced by extracting an electron from water; thus, water is split and PSII
2
is re-reduced after every photoact. Splitting one H O molecule releases two electrons, two hydrogen atoms,
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2 gas.
and one atom of oxygen. Splitting two molecules of water is required to form one molecule of diatomic O
About 10 percent of the oxygen is used by mitochondria in the leaf to support oxidative phosphorylation.
The remainder escapes to the atmosphere where it is used by aerobic organisms to support respiration.
As electrons move through the proteins that reside between PSII and PSI, they take part in exergonic
redox transfers. The free energy associated with the exergonic redox reaction is coupled to the endergonic
transport of hydrogen atoms from the stromal side of the membrane to the thylakoid lumen. Those hydrogen
atoms, plus the ones produced by splitting water, accumulate in the thylakoid lumen and will be used to
synthesize ATP in a later step.
Since the electrons on PSI now have a greater reduction potential than
when they started their trek (it is important to note that PSI unexcited sits lower on the redox tower than
NADP+/NADPH), they must be re-energized in PSI. Therefore, another photon is absorbed by the PSI
antenna. That energy is transferred to the PSI reaction center (called
an electron through several intermediate redox steps to NADP
+
P700).
P700 is oxidized and sends
to form NADPH. Thus, PSII captures
the energy in light and couples its transfer via redox reactions to the creation of a proton gradient. The
exergonic and controlled relaxation of this gradient can be coupled to the synthesis of ATP. PSI captures
+
energy in light and couples that, through a series of redox reactions, to reduce NADP
into NADPH. The
two photosystems work in concert, in part, to guarantee that the production of NADPH will be in the right
proportion to the production of ATP. Other mechanisms exist to ne tune that ratio to exactly match the
chloroplast's constantly changing energy needs.
3.1 Additional links
Some Additional Links on Photophosphorylation
• wiley presents photosynthesis1
• YouTube photosynthesis2
•
YouTube photosynthesis and electron tower usage by Dr. Easlon
3
4 Light Independent Reactions or Carbon Fixation
A short introduction
The general principle of carbon xation is that some cells under certain conditions can take inorganic carbon,
CO
2
(also referred to as mineralized carbon) and reduce it to a usable cellular form. Most of us are aware
2
that green plants can take up CO
2
and produce O
in a process known as photosynthesis. We have already
discussed photophosphorylation, the ability of a cell to transfer light energy onto chemicals and ultimately to
produce the energy carriers ATP and NADPH in a process known as the light reactions. In photosynthesis,
the plant cells use the ATP and NADPH formed during photophosphorylation to reduce CO
2
to sugar, (as
we will see, specically G3P) in what are called the dark reactions. While we appreciate that this process
happens in green plants, photosynthesis had its evolutionary origins in the bacterial world. In this module
we will go over the general reactions of the Calvin Cycle, a reductive pathway that incorporates CO
cellular material.
2
into
In photosynthetic bacteria, such as Cyanobacteria and purple non-sulfur bacteria, as well plants, the
energy (ATP) and
reducing power (NADPH) - a term used to describe electron carriers in their reduced
Carbon Fixation", the incorporation of inorganic
state - obtained from photophosphorylation is coupled to "
2
carbon (CO ) into organic molecules; initially as glyceraldehyde-3-phosphate (G3P) and eventually into
2
glucose. Organisms that can obtain all of their required carbon from an inorganic source (CO )are refereed
to as
autotrophs,
while those organisms that require organic forms of carbon, such as glucose or amino
acids, are refereed to as
heterotrophs.
The biological pathway that leads to carbon xation is called the
1 http://www.wiley.com/legacy/college/boyer/0470003790/animations/photosynthesis/photosynthesis.htm
2 https://www.youtube.com/watch?v=joZ1EsA5_NY
3 https://www.youtube.com/watch?v=joZ1EsA5_NY
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Calvin Cycle and is a reductive pathway (consumes energy/uses electrons) which leads to the reduction of
CO2 to G3P.
4.1 The Calvin Cycle: the reduction of CO2 to Glyceraldehyde 3-Phosphate
Figure 12: Light reactions harness energy from the sun to produce chemical bonds, ATP, and NADPH.
These energy-carrying molecules are made in the stroma where carbon xation takes place.
In plant cells, the Calvin cycle is located in the chloroplasts. While the process is similar in bacteria, there
are no specic organelles that house the Calvin Cycle and the reactions occur in the cytoplasm around a
complex membrane system derived from the plasma membrane. This
intracellular membrane system
can be quite complex and highly regulated. There is strong evidence that supports the hypothesis that the
origin of
chloroplasts from a symbiosis between cyanobacteria and early plant cells.
4.1.1 Stage 1: Carbon Fixation
2
In the stroma of plant chloroplasts, in addition to CO ,two other components are present to initiate the
light-independent reactions: an enzyme called ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBisCO),
and three molecules of ribulose bisphosphate (RuBP), as shown in Figure 13.
(RuBP) is composed of ve carbon atoms and includes two phosphates.
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Ribulose-1,5-bisphosphate
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Figure 13: The Calvin cycle has three stages. In stage 1, the enzyme RuBisCO incorporates carbon
dioxide into an organic molecule, 3-PGA. In stage 2, the organic molecule is reduced using electrons
supplied by NADPH. In stage 3, RuBP, the molecule that starts the cycle, is regenerated so that the
cycle can continue. Only one carbon dioxide molecule is incorporated at a time, so the cycle must be
completed three times to produce a single three-carbon GA3P molecule, and six times to produce a
six-carbon glucose molecule.
RuBisCO catalyzes a reaction between CO
2
2
and RuBP. For each CO
molecule that reacts with one RuBP,
two molecules of another compound (3-PGA) form. PGA has three carbons and one phosphate. Each turn
of the cycle involves only one RuBP and one carbon dioxide and forms two molecules of 3-PGA. The number
of carbon atoms remains the same, as the atoms move to form new bonds during the reactions (3 atoms from
2
3CO
+ 15 atoms from 3RuBP = 18 atoms in 3 atoms of 3-PGA). This process is called
2
because CO
carbonxation,
is xed from an inorganic form into an organic molecule.
4.1.2 Stage 2: Reduction
ATP and NADPH are used to convert the six molecules of 3-PGA into six molecules of a chemical called
glyceraldehyde 3-phosphate (G3P) - a carbon compound that also is produced in glycolysis. Six molecules of
both ATP and NADPH are used in the process. The exergonic process of ATP hydrolysis is in eect driving
+
the endergonic redox reactions, creating ADP and NADP . Both of these molecules return to the nearby
light-dependent reactions to be recycled back into ATP and NADPH.
4.1.3 Stage 3: Regeneration
Interestingly, at this point, only one of the G3P molecules leaves the Calvin cycle to contribute to the
formation of other compounds needed by the organism.
In plants, because the G3P exported from the
Calvin cycle has three carbon atoms, it takes three turns of the Calvin cycle to x enough net carbon to
export one G3P. But each turn makes two G3Ps, thus three turns make six G3Ps. One is exported while
the remaining ve G3P molecules remain in the cycle and are used to regenerate RuBP, which enables the
2
system to prepare for more CO
reactions.
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to be xed. Three more molecules of ATP are used in these regeneration
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Figure 14: The harsh conditions of the desert have led plants like these cacti to evolve variations of the
light-independent reactions of photosynthesis. These variations increase the eciency of water usage,
helping to conserve water and energy. (credit: Piotr Wojtkowski)
4.1.4 Free Response Questions
Exercise 1
(Solution on p. 19.)
Why is the third stage of the Calvin cycle called the regeneration stage?
Exercise 2
(Solution on p. 19.)
Which part of the light-independent reactions would be aected if a cell could not produce the
enzyme RuBisCO?
Exercise 3
(Solution on p. 19.)
Why does it take three turns of the Calvin cycle to produce G3P, the initial product of photosynthesis?
Prepare for the Test:
Create an energy story for each phase of the Calvin cycle.
http://cnx.org/content/m60007/1.1/
Classify the reactants and products and pay
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attention to where the energy is at the beginning of the reaction and the end of the reaction. At this point
you should be able to tell if a reaction is a REDOX reaction (does it have NADPH as a reactant or product?)
or if the reaction is endergonic or exergonic (is ATP created or used in the reaction?).
5 Additional Links of Interest
Khan Academy Links
• Calvin Cycle4
Chemwiki links
•
Calvin Cycle
5
YouTube Videos
•
•
6
3D animation of photosynthesis in plants
7
Calvin Cycle
4 https://www.khanacademy.org/science/biology/cellular-molecular-biology/photosynthesis/v/photosynthesis-calvin-cycle
5 http://chemwiki.ucdavis.edu/Biological_Chemistry/Metabolism/Catabolism
6 https://www.youtube.com/watch?v=YeD9idmcX0w
7 https://www.youtube.com/watch?v=E_XQR800AgM
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Solutions to Exercises in this Module
to Exercise (p. 17)
Because RuBP, the molecule needed at the start of the cycle, is regenerated from G3P.
to Exercise (p. 17)
None of the cycle could take place, because RuBisCO is essential in xing carbon dioxide.
Specically,
RuBisCO catalyzes the reaction between carbon dioxide and RuBP at the start of the cycle.
to Exercise (p. 17)
Because G3P has three carbon atoms, and each turn of the cycle takes in one carbon atom in the form of
carbon dioxide.
Glossary
Denition 1: absorption spectrum
range of wavelengths of electromagnetic radiation absorbed by a given substance
Denition 2: antenna protein
pigment molecule that directly absorbs light and transfers the energy absorbed to other pigment
molecules
Denition 3: carotenoid
photosynthetic pigment that functions to dispose of excess energy
Denition 4: chlorophyll a
form of chlorophyll that absorbs violet-blue and red light and consequently has a bluish-green
color; the only pigment molecule that performs the photochemistry by getting excited and losing
an electron to the electron transport chain
Denition 5: chlorophyll b
accessory pigment that absorbs blue and red-orange light and consequently has a yellowish-green
tint
Denition 6: cytochrome complex
group of reversibly oxidizable and reducible proteins that forms part of the electron transport chain
between photosystem II and photosystem I
Denition 7: electromagnetic spectrum
range of all possible frequencies of radiation
Denition 8: electron transport chain
group of proteins between PSII and PSI that pass energized electrons and use the energy released by
the electrons to move hydrogen ions against their concentration gradient into the thylakoid lumen
Denition 9: light harvesting complex
complex that passes energy from sunlight to the reaction center in each photosystem; it consists of
multiple antenna proteins that contain a mixture of 300400 chlorophyll
a
and
b
molecules as well
as other pigments like carotenoids
Denition 10: P680
reaction center of photosystem II
Denition 11: P700
reaction center of photosystem I
Denition 12: photoact
ejection of an electron from a reaction center using the energy of an absorbed photon
Denition 13: photon
distinct quantity or packet of light energy
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Denition 14: photosystem
group of proteins, chlorophyll, and other pigments that are used in the light-dependent reactions
of photosynthesis to absorb light energy and convert it into chemical energy
Denition 15: photosystem I
integral pigment and protein complex in thylakoid membranes that uses light energy to transport
+
electrons from plastocyanin to NADP
(which becomes reduced to NADPH in the process)
Denition 16: photosystem II
integral protein and pigment complex in thylakoid membranes that transports electrons from water
to the electron transport chain; oxygen is a product of PSII
Denition 17: primary electron acceptor
pigment or other organic molecule in the reaction center that accepts an energized electron from
the reaction center
Denition 18: reaction center
complex of chlorophyll molecules and other organic molecules that is assembled around a special
pair of chlorophyll molecules and a primary electron acceptor; capable of undergoing oxidation and
reduction
Denition 19: spectrophotometer
instrument that can measure transmitted light and compute the absorption
Denition 20: wavelength
distance between consecutive points of equal position (two crests or two troughs) of a wave in a
graphic representation; inversely proportional to the energy of the radiation
Denition 21: Calvin cycle
light-independent reactions of photosynthesis that convert carbon dioxide from the atmosphere into
carbohydrates using the energy and reducing power of ATP and NADPH
Denition 22: carbon xation
process of converting inorganic CO
Denition 23: reduction
2
gas into organic compounds
gain of electron(s) by an atom or molecule
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