The Calvin cycle proceeds in three stages: carboxylation, reduction, and regeneration
Carboxylation of the CO2 acceptor,
ribulose-1, 5-biphosphate, forming
two molecules of 3-phosphoglcerate.
Rubisco – the enzyme ribulose
biphosphate carboxylase/oxygenase
Reduction of 3-phosphoglycerate to
form glyceraldehyde-3-phosphate
which can be used in formation of
carbon compounds that are
translocated.
Regeneration of the CO2 acceptor
ribulose-1, 5-biphosphate from
glyceraldehyde-3-phosphate
RuBP
The affinity of Rubisco for CO2 is sufficiently high to
ensure rapid carboxylation at the low concentration
of CO2 found in photosynthesizing cells
Rubisco will also take O2 rather than CO2 and
oxygenate RuBP – called photorespiration.
The rate of operation of the Calvin Cycle can be
enhanced by increases in the concentration of its
intermediates. That is the cycle is autocatalytic.
Also, if there are insufficient intermediates
available, for example when a plant is transferred
from dark to light, then there is a lag, or induction
period, before photosynthesis reaches the level
that the light can sustain. (There can also be
enzyme induction.)
Increasing CO2
concentration in the
Basics of foliage photosynthesis atmosphere can
Dark Reaction increase the maximum
Light Reaction rate of photosynthesis in
Limiting Limiting
the short term
Saturation level.
sometimes called
photosynthetic
capacity.
Photosynthetic efficiency:
Increase in photosynthesis
per increase in irradiance
0
0
Compensation point
The irradiance at which CO uptake is zero
2
Measured light response curve of Abies amabilis first year foliage.
Shade foliage with low maximum value and low compensation point.
Observed assimilation rates
(µmolCO2/m2s) of Tsuga
heterophylla and Abies amabilis in
response to periods of 10 minutes
high light (1500µmol/m2s PPFD),
with 5 minutes intervals of
darkness (shaded parts in the
diagram) in between. Values
measured using 200 mol/s flow
rate.
Species differences in leaf photosynthesis
A has the highest
photosynthetic rate at
light saturation
B has the highest
photosynthetic efficiency
and the lowest
compensation point.
Units: μmol/m2/s
micro mols of CO2 per square
meter foliage per second
Another important measure is called Water Use Efficiency:
the ratio of photosynthesis achieved per unit of water lost.
Units: mmol/mol,
milli mols of CO milli [m] 0.001 (a thousandth)
2 micro [µ] 0.000 001 (a millionth)
per mol of water transpired
Old-growth species:
Tsuga
heterophylla Notice the
difference in
Pseudotsuga branch structure
menziesii
between the
species
Thuja plicata
Abies grandis
Upper Canopy Lower Canopy
Phot. Cap. Water Use Eff. Phot. Cap. Water Use Eff.
Douglas-fir
2
Pseudotsuga 13.1 μmol/m /s 6.2 mmol/mol 8.8 3.5
Western
hemlock 9.0 4.9 3.2 4.8
Tsuga
The problem of photorespiration
and the evolution of photosynthesis
When the enzyme Rubisco uses oxygen to breakdown
carbohydrate to CO2 rather than using CO2 to synthesize
carbohydrate
How some grasses have evolved a C4 metabolic
process and some desert plants have evolved
Crassulacean Acid Metabolism
Although Rubisco acts like a carboxylase in photosynthesis,
it can also act as an oxygenase when O2 is available.
O2 and CO2 compete for the same active site!
O2
P P P P
C-C-C-C-C C-C-C + C-C
RuBisCO
3-phospho 2-phospho
Ribulose 1, 5-biphosphate
glycerate glycerate
Enzyme
CO2
This becomes a problem when photosynthesis rates are
high, i.e. photosystem II produces lots of O2 .
This is called Photorespiration
In the presence of higher O2 levels,
photosynthesis rates are lower.
275 ppm CO2
The inhibition of
photosynthesis by O2 was first
73 ppm CO2
noticed by the German plant
physiologist, Otto Warburg, in
1920, and called the "Warburg
effect".
It is believed that photorespiration in plants has increased over
geologic time due to increasing atmospheric O2 concentration -the
product of photosynthetic organisms themselves.
C4 Photosynthesis
The first product of CO2 fixation is
malate (C4) in mesophyll cells, not
PGA as it is in C3 plants. This is
transported to bundle sheath cells
CO2 is released from malate in bundle
sheath cells, where it is fixed again by
Rubisco and the Calvin cycle proceeds.
PEP is recycled back to mesophyll
cells.
Decarboxylation of malate (CO2 release)
creates a higher concentration of CO2 in
bundle sheath cells than found in
photosynthetic cells of C3 plants.
This enables C4 plants to sustain higher
rates of photosynthesis. And, because
the concentration of CO2 relative to O2 in
bundle sheath cells is higher,
photorespiration rates are lower.
Anatomical separation of the Parenchyma filled Bundle sheath
with chloroplasts cells filled with
C4 photosynthesis
component processes chloroplasts.
CALVIN
REACTION SITE
Xylem
Phloem
Carbon
skeleton
compounds
return to
parenchyma
C4 acids synthesized in the
parenchyma move to the bundle
sheath
Crassulacean Acid Metabolism (CAM)
First discovered in succulents of the
Crassulacea: e.g.,sedums
Uses C4 pathways, but segregates CO2
assimilation and Calvin cycle between
day and night
CAM plants open their stomates at
night. This conserves H2O. CO2 is
assimilated into malic acid and stored
in high concentrations in cell vacuoles
During the day, stomates close, and
the stored malic acid is gradually
recycled to release CO2 to the Calvin
cycle
C3, majority of C4, e.g., sugar CAM,
species cane, corn
e.g., cacti
Leaf structure Bundle sheath Bundle sheath Mesophyll cells
cells lack cells have have large
chloroplasts chloroplasts vacuoles
Efficiency in Can be sun Ineffective CO2 capture
light or shade plants in shade at night
Typical habitat Requires Arid or Arid
characteristics relatively tropical environments
moist habitats regions
Productivity Moderate High Low
Optimum 30-40oC
15-25oC 35oC
Temperature
