 The Calvin Cycle: An Anabolic Cycle

During the light-independent reactions of photosynthesis, a.k.a., The Calvin

Cycle, the energy stored in ATP and NADP-H during the light-dependent
reactions is briefly released and then repackaged into the covalent bonds of
sugar.

 The sugar first produced by the Calvin Cycle is not 6-carbon glucose (this is
constructed later), but a 3-carbon sugar known as glyceraldehyde-3-
phosphate or G3P.

 The Calvin Cycle "spins" three times to make this 3-carbon sugar from three
molecules of CO2. The conversion of inorganic carbon from CO2 into the
carbons of an organic molecule, G3P, is known as carbon fixation.

 Remember that there are many Calvin Cycles going on in the stroma at any
given time, so many atoms of carbon are constantly being fixed quickly into
sugar.

 The Calvin Cycle can be reduce to three phases:

 I. Carbon Fixation

One CO2 molecule is attached to a 5-carbon sugar named ribulose

biphosphate (RuBP) by an enzyme named rubisco. (Rubisco may
be the most abundant protein on earth! It is certainly the most
abundant protein in the chloroplasts.)

The resulting 6-carbon sugar is extremely unstable, and it

immediately splits in half, forming two molecules of 3-carbon 3-
phospho glycerate.

 II. Reduction
o A phosphate group (from ATP) is attached to each 3-phospho
glycerate, forming 1,3-phosphoglycerate (guess where the
phosphates are attached!)
o NADP-H swoops in and reduces 1,3-phosphoglycerate, which loses
its phosphate group (in addition to gaining electrons) to become
our old pal, G3P.
 o For every six G3Ps made by the Calvin Cycle, five are recycled back
to regenerate new molecules of RuBp. Only ONE leaves the cycle
to be packaged for use by the plant.
o It takes two molecules of G3P (3 carbons) to make one molecule
of glucose (6 carbons).
o ADP and NADP+ formed during the Calvin Cycle are shuttled back
to the photosystems to be "recharged" with energy, and
converted to ATP and NADP-H, respectively.
 III. Regeneration of RuBP

Five molecules of G3P from the Calvin Cycle's Reduction phase

pass through a complex series of enzymatic reactions to yield
three molecules of RuBP. This costs the cell 3 more molecules of
ATP, but provides new "machinery" for the Calvin Cycle to
continue spinning

Carbon dioxide for photosynthesis enters via the stomates. But during the
hottest part of the day, stomates often close to prevent water loss. Even with
stomates partially open, CO2 levels drop rapidly. And this triggers a process
known as photorespiration. If the plant continues to attempt to fix CO2 when
its stomates are closed, the CO2 will become depleted, causing the cellular [O2]
to increase relative to cellular [CO2].

Photorespiration:
 As CO2 levels drop, the Calvin Cycle becomes "starved" for raw material.

 Rubisco has some affinity for O2 (albeit lower than for CO2).

 When CO2 is scarce, rubisco will bind to O2 to its ribulose-biphosphate

substrate, instead of the normal CO2.

 The resulting "deformed" product of this metabolic cycle, like the normal
one, is unstable. But instead of splitting into two molecules of 3-
phosphoglycerate, (the normal products), it splits into one molecule of 3-
phosphoglycerate and one molecule of a 2-carbon compound, 2-
phosphoglycolate.
 The 2-phosphoglycolate diffuses out of the chloroplast.

 When 2-phosphoglycerate enters the peroxisomes or mitochondria, it is

further split to release CO2.

 This process is called photorespiration because it occurs in the

light and generates CO2. But unlike cellular respiration, it
generates no ATP. (In fact, it consumes ATP.)

C3 plantsPlants that produce 3-carbon G3P via the Calvin Cycle are called C3 plants.
all of carbon fixation and photosynthesis happens in mesophyll cells just on the
surface of the leaf. C3 plants include most temperate plants (except many grasses)
—more than 95% of all earth’s plants.
 

 A unique leaf anatomy is associated with the C4 pathway: C4 plants have

two types of photosynthetic cells, bundle-sheath cells and mesophyll cells,
arranged in the leaf as shown here:

 The Calvin Cycle occurs only in the bundle-sheath cells.

 The mesophyll cells act as a "carbon dioxide pump," concentrating
carbon dioxide by transporting CO2 cleverly bound to other compounds.

But how?
 In the mesophyll cells, an enzyme known as PEP carboxylase binds free CO2
to phosphoenolpyruvate (PEP) to form 4-carbon oxaloacetate and other 4-
carbon compounds (e.g., malate).

 PEP carboxylase has a very high affinity for CO2, but NONE for oxygen gas. So
when rubisco is bamboozled by stomatal closure and oxygen, PEP carboxylase
can still sequester carbon dioxide.
 Via plasmodesmata, mesophyll cells transport their 4-carbon, CO2-carrying
products to the bundle-sheath cells.

 The 4-carbon compounds release their precious CO2 passengers, which then
enter the Calvin Cycle and can be incorporated into G3P and other products in
a normal fashion.

 This CO2 release generates pyruvate, which travels back to the mesophyll
cells and is converted to PEP (with energy expended).

 So this process is not without cost, but it keeps the Calvin Cycle going even
when CO2 concentration in the cell is low, while minimizing photorespiration.

But where do we get that extra ATP?

 Bundle sheath cells have only Photosystem I, which generates ATP, but no
NADP-H. Only cyclic electron flow is used to generate ATP in these cells.

In essence, C4 plants pump enough extra CO2 into the bundle sheath cells to
keep rubico loaded with CO2, at a cost of ATP.
This mechanism would be particularly advantageous in a hot, arid climate,
where stomates must close during the day to prevent desiccation, and it is in
these areas that C4 plants first evolved and still thrive.

The equation for the Calvin Cycle:

CO2 (Carbon dioxide in from stomata) + RuBP (Ribulose bisphosphate already in
plant) + the enzyme RUBISCO (Ribulose bisphosphate carboxylase) “fixes” carbon
from the atmosphere à 2PGA (phospholygerate)
 
PGA enters Calvin cycle in Mesophyll cells à more RuBP (to fix more CO2) + sugar
(CH2O)
 
 C3 are inefficient at CO2 fixation because RUBISCO has a greater affinity for
oxygen than CO2
 Mesophyll cells are packed with RUBISCO
 Stomata open during day (CO2, oxygen, and water can all flow out)
 
Photorespiration undoes CO2 assimilation
2PGA à CO2 + RuBP
increases when there is lots of O2, low levels of CO2, and increased
temperature
 
C4 plantsSome plants, across about 19 different families, have a clever trick that allows photosynthesis
while stomates are closed while minimizing photorespiration. These plants, known as C4 plants manufacture a
4-carbon compound that serves as a "shuttle" for CO2 to the Calvin Cycle's initial carbon fixation phase when
plants can't access atmospheric carbon dioxide.

carbon fixation and photosynthesis split between the mesophyll cells and bundle
sheath cells.
 
The equation:
In mesophyll (carbon fixation):
CO2 + PEP (phosphoenol pyruvate) + PEP carboxylase fixes carbon à OAA
(oxaloacetic acid)
 
OAA diffuses to bundle sheath cells
 
In bundle sheath (Calvin Cycle):
OAA à malic acid and aspartic acid is decarboxylated à CO2 + pyruvate
 
Then the Calvin Cycle CO2 + RuBP + the enzyme RUBISCO à 2PGA à RuBP +
CH2O
 
Pyruvate with ATP is moved back to mesophyll and turned into PEP (to fix more
CO2)
 
 Feature of many grasses (i.e. big blue stem back campus), corn, and many
arid/semi arid shrubs
  Calvin cycle in bundle sheath cells where there is no oxygen to be bound by
RUBISCO
 Very high concentration of CO2 in bundle sheath cells
 PEP carboxylase has a high affinity for CO2 so plants must open their stomata
less to get CO2 and hence lose less water (especially important in arid
regions)
 Low levels of photorespiration and higher net photosynthesis than C 3 because
of low photorespiration
 Costly adaptation because it requires lots of ATP (energy)—however, benefits
outweigh energy costs.
 Stomata are open during the day
 Fixation and the calvin cyle are physically separate
 
CAM Crassulacean acid metabolism Many succulent plants have evolved a different way to
allow normal photosynthesis even when stomates are closed. The plants in which this pathway was first
discovered are members of Family Crassulaceae, and the pathway, named for them is known as crassulacean
acid metabolism, or CAM.

During the day, CAM plants' stomates are closed. But at night they open, taking
up CO2 and incorporating it into a variety of organic acids for storage. These
acids are stored in the vacuoles of leaf mesophyll cells until daybreak.

As the light reactions start up in response to light, CO2 is released from the
organic acids. The energy from ATP and NADP-H from the light reactions can
now be used to fix that carbon, even though the stomates are closed.

photosynthesis takes place in the mesophyll cells, but carbon fixation (and opening
of stomata) takes place at night and the Calvin cycle happens during the day.

 C4 and CAM are similar in that they use organic intermediates to store CO2
for later release, concentrating it when the plant isn't open to the atmosphere.
 C4 and CAM are different in that C4 plants have two types of cells, and they
separate carbon dioxide storage cells (mesophyll) from Calvin Cycle-
performing cells (bundle-sheath). CAM plants perform both functions in the
same (mesophyll) cells).
 
The equations are the same as for C4 plants. Substitute “night” for mesophyll and
“day” for bundle sheath.
 
 Because the stomata are closed during the day, able to capture large levels of
CO2
 Adaptation found in succulents, cacti, and euphorbs for example that are
found in desert (very arid) environments.
 Very high water use efficiencies because stomata don’t open during the
hottest times of day when transpiration is the greatest.
 Also energy expensive