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Derived copy of Bis2A 07.5 The
Calvin Cycle or Using Light Energy
∗
to Make Organic Molecules
Erin Easlon
Based on Bis2A 07.5 The Calvin Cycle or Using Light Energy to Make Organic Molecules† by
OpenStax College
Mitch Singer
This work is produced by OpenStax-CNX and licensed under the
Creative Commons Attribution License 4.0‡
Abstract
By the end of this section, you will be able to:
• Describe the Calvin cycle
• Dene carbon xation
• Explain how photosynthesis works in the energy cycle of all living organisms
1 Intro into Carbon Fixation
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 convert light energy to chemical energy in the form
of a high energy electron that then enters the electron transport chain to produce ATP and NADPH, see
module as described in Module 6.3.
In photosynthesis, the plant cells use the ATP and NADPH formed
2
during photophosphorylation, the light reactions, to reduce CO
to sugar, (as we will see, specically G3P)
in what is called the dark reactions. While we all familiar with this process in green plants, I want to point
out that this process had its origins in the bacterial world. ATP and NADPH can be made during anoxygenic
2
photophosphorylation and CO
into reduced sugars for the cell. In this module we will go over the general
reactions of the Calvin Cycle, a reductive pathway that incorporates CO
2
into cellular material. In many
ways it is the reverse of the oxidative pentose phosphate pathway. One exercise to keep in mind is to look
for the similarities between these two pathways.
∗ Version
1.1: Jul 18, 2015 9:59 am -0500
† http://cnx.org/content/m56025/1.2/
‡ http://creativecommons.org/licenses/by/4.0/
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A Note from the instructor
As with the modules on glycolysis, the TCA cycle, and the Pentose Phosphate Pathway, there is a lot of
material in this module. AS with these other modules, I do not expect you to memorize specic names of
compounds or enzymes. However, as in those other modules I will give include the names for completeness.
For exams I will always provide you with the pathways we discuss in class and in the BioStax Biology text
modules. What you need to be able to do is understand the under lying principles for each reaction. We will
go over in lecture problems that will be similar to those I will ask of you on exams. Do not be overwhelmed
with specic enzyme names and specic structures. What you should know are the general types of enzymes
used and the types of structures found. If you have any questions please ask.
2 FIXING CARBON INTO BIOLOGICAL MOLECULES
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 photosynthetic bacteria, such as Cyanobacteria, and purple non-sulfur bacteria, as well as and plants
Carbon
Fixation", the incorporation of inorganic carbon (CO2 ) into organic molecules, glyceraldehyde-3-phosphate
the energy (ATP) and reducing power (NADPH) obtained from photophosphorylation is coupled to "
(G3P) and eventually into glucose. Organisms that can obtain all of their required carbon from an inorganic
2
source (CO )are refereed to as
autotrophs,
while those organisms that require organic forms of carbon,
such as glucose or protein, are refereed to as
carbon xation is called the
2
the reduction of CO
Calvin Cycle
heterotrophs.
The the biological pathway that leads to
and is a reductive pathway (consumes energy) which leads to
to G3P.
3 The Calvin Cycle
2
In plants, carbon dioxide (CO ) enters the leaves through stomata, where it diuses over short distances
2
through intercellular spaces until it reaches the mesophyll cells. Once in the mesophyll cells, CO
diuses
into the stroma of the chloroplastthe 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 (Figure 1). 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.
While the process is similar in bacteria, there are no specic organelles that house the Calvin Cycle genes
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. In fact,
there is strong evidence that supports the hypothesis that the origin of
between cyanobacteria and early plant cells.
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chloroplasts comes from a symbiosis
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Figure 1: 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.
The light-independent reactions of the Calvin cycle can be organized into three basic stages: xation,
reduction, and regeneration.
3.1 Stage 1: 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 2. RuBP has ve atoms of carbon,
anked by two phosphates.
:
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Figure 2: 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.
Which of the following statements is true?
a.In photosynthesis, oxygen, carbon dioxide, ATP, and NADPH are reactants. GA3P and water
are products.
b.In photosynthesis, chlorophyll, water, and carbon dioxide are reactants. GA3P and oxygen
are products.
c.In photosynthesis, water, carbon dioxide, ATP, and NADPH are reactants. RuBP and oxygen
are products.
d.In photosynthesis, water and carbon dioxide are reactants. GA3P and oxygen are products.
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
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3CO
5
+ 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 organic molecules.
3.2 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). That is a reduction reaction because it involves the gain of electrons by
3-PGA. Recall that a
reduction is the gain of an electron by an atom or molecule.
ATP and NADPH are used.
Six molecules of both
For ATP, energy is released with the loss of the terminal phosphate atom,
+
converting it into ADP; for NADPH, both energy and a hydrogen atom are lost, converting it into NADP .
Both of these molecules return to the nearby light-dependent reactions to be reused and reenergized.
3.3 Stage 3: Regeneration
Interestingly, at this point, only one of the G3P molecules leaves the Calvin cycle and is sent to the cytoplasm
to contribute to the formation of other compounds needed by the plant. Because the G3P exported from
the chloroplast 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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This link
1 leads to an animation of the Calvin cycle.
6
Click stage 1, stage 2, and then stage 3 to see
G3P and ATP regenerate to form RuBP.
:
Photosynthesis
During the evolution of photosynthesis, a major shift occurred from the bacterial type of photosynthesis that involves only one photosystem and is typically anoxygenic (does not generate oxygen)
into modern oxygenic (does generate oxygen) photosynthesis, employing two photosystems. This
modern oxygenic photosynthesis is used by many organismsfrom giant tropical leaves in the rainforest to tiny cyanobacterial cellsand the process and components of this photosynthesis remain
largely the same. Photosystems absorb light and use electron transport chains to convert energy
into the chemical energy of ATP and NADH. The subsequent light-independent reactions then
assemble carbohydrate molecules with this energy.
1 http://openstaxcollege.org/l/calvin_cycle
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Photosynthesis in desert plants has evolved adaptations that conserve water. In the harsh dry heat,
every drop of water must be used to survive. Because stomata must open to allow for the uptake of
2
CO , water escapes from the leaf during active photosynthesis. Desert plants have evolved processes
to conserve water and deal with harsh conditions.
adapt to living with less water.
A more ecient use of CO
2
allows plants to
Some plants such as cacti (Figure 3) can prepare materials for
photosynthesis during the night by a temporary carbon xation/storage process, because opening
the stomata at this time conserves water due to cooler temperatures. In addition, cacti have evolved
the ability to carry out low levels of photosynthesis without opening stomata at all, an extreme
mechanism to face extremely dry periods.
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Figure 3: 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)
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4 The Energy Cycle
Whether the organism is a bacterium, plant, or animal, all living things access energy by breaking down
carbohydrate molecules. But if plants make carbohydrate molecules, why would they need to break them
2
down, especially when it has been shown that the gas organisms release as a waste product (CO ) acts
as a substrate for the formation of more food in photosynthesis? Remember, living things need energy to
perform life functions. In addition, an organism can either make its own food or eat another organism
either way, the food still needs to be broken down. Finally, in the process of breaking down food, called
2
cellular respiration, heterotrophs release needed energy and produce waste in the form of CO
gas.
In nature, there is no such thing as waste. Every single atom of matter and energy is conserved, recycling
over and over innitely. Substances change form or move from one type of molecule to another, but their
constituent atoms never disappear (Figure 4).
CO
2
is no more a form of waste than oxygen is wasteful to photosynthesis.
Both are byproducts of
reactions that move on to other reactions. Photosynthesis absorbs light energy to build carbohydrates in
chloroplasts, and aerobic cellular respiration releases energy by using oxygen to metabolize carbohydrates
in the cytoplasm and mitochondria.
Both processes use electron transport chains to capture the energy
necessary to drive other reactions. These two powerhouse processes, photosynthesis and cellular respiration,
function in biological, cyclical harmony to allow organisms to access life-sustaining energy that originates
millions of miles away in a burning star humans call the sun.
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Figure 4: Photosynthesis consumes carbon dioxide and produces oxygen. Aerobic respiration consumes
oxygen and produces carbon dioxide. These two processes play an important role in the carbon cycle.
(credit: modication of work by Stuart Bassil)
5 Section Summary
Using the energy carriers formed in the rst steps of photosynthesis, the light-independent reactions, or the
Calvin cycle, take in CO
2
2
from the environment. An enzyme, RuBisCO, catalyzes a reaction with CO
and
another molecule, RuBP. After three cycles, a three-carbon molecule of G3P leaves the cycle to become part
of a carbohydrate molecule. The remaining G3P molecules stay in the cycle to be regenerated into RuBP,
2
which is then ready to react with more CO .
cellular respiration.
Photosynthesis forms an energy cycle with the process of
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.
chloroplasts and mitochondria.
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Therefore, plants contain both
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6 Additional Links
Khan Academy Links
• Calvin Cycle2
Chemwiki links
•
Calvin Cycle
3
YouTube Videos
•
•
4
3D animation of photosynthesis in plants
5
Calvin Cycle
7 Art Connections
Exercise 1
(Solution on p. 13.)
Figure 2 Which of the following statements is true?
a. In photosynthesis, oxygen, carbon dioxide, ATP, and NADPH are reactants. G3P and water
are products.
b. In photosynthesis, chlorophyll, water, and carbon dioxide are reactants. G3P and oxygen are
products.
c. In photosynthesis, water, carbon dioxide, ATP, and NADPH are reactants. RuBP and oxygen
are products.
d. In photosynthesis, water and carbon dioxide are reactants. G3P and oxygen are products.
8 Review Questions
Exercise 2
(Solution on p. 13.)
Which molecule must enter the Calvin cycle continually for the light-independent reactions to take
place?
a. RuBisCO
b. RuBP
c. 3-PGA
d. CO
2
Exercise 3
(Solution on p. 13.)
Which order of molecular conversions is correct for the Calvin cycle?
a. RuBP + G3P
→
→
3-PGA
CO2
c. RuBP + CO2
d. CO2
→
→
sugar
→ G3P
→ [RuBisCO] 3-PGA →
→ 3-PGA → RuBP → G3P
b. RuBisCO
RuBP
G3P
2 https://www.khanacademy.org/science/biology/cellular-molecular-biology/photosynthesis/v/photosynthesis-calvin-cycle
3 http://chemwiki.ucdavis.edu/Biological_Chemistry/Metabolism/Catabolism
4 https://www.youtube.com/watch?v=YeD9idmcX0w
5 https://www.youtube.com/watch?v=E_XQR800AgM
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Exercise 4
(Solution on p. 13.)
Where in eukaryotic cells does the Calvin cycle take place?
a. thylakoid membrane
b. thylakoid lumen
c. chloroplast stroma
d. granum
Exercise 5
(Solution on p. 13.)
Which statement correctly describes carbon xation?
a. the conversion of CO
2
into an organic compound
b. the use of RuBisCO to form 3-PGA
c. the production of carbohydrate molecules from G3P
d. the formation of RuBP from G3P molecules
e. the use of ATP and NADPH to reduce CO
2
9 Free Response
Exercise 6
(Solution on p. 13.)
Why is the third stage of the Calvin cycle called the regeneration stage?
Exercise 7
(Solution on p. 13.)
Which part of the light-independent reactions would be aected if a cell could not produce the
enzyme RuBisCO?
Exercise 8
(Solution on p. 13.)
Why does it take three turns of the Calvin cycle to produce G3P, the initial product of photosynthesis?
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Solutions to Exercises in this Module
to Exercise (p. 11)
Figure 2 D
to Exercise (p. 11)
D
to Exercise (p. 11)
C
to Exercise (p. 12)
C
to Exercise (p. 12)
A
to Exercise (p. 12)
Because RuBP, the molecule needed at the start of the cycle, is regenerated from G3P.
to Exercise (p. 12)
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. 12)
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: 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 2: carbon xation
process of converting inorganic CO
Denition 3: reduction
2
gas into organic compounds
gain of electron(s) by an atom or molecule
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