C4 and Krantz anatomy

(a) describe the anatomical structure of C4 leaf (Krantz anatomy) in comparison to C3 leaf;

(b) explain carbon dioxide fixation in C4 plants and Crassulacean Acid Metabolism (CAM) plants;

differentiate the metabolism of C3, C4 and CAM plants.

When is photorespiration the highest?
• Not when photosynthesis is highest. Because if the stoma is open the
oxygen produced will diffuse out of the mesophyll.
• Photorespiration is the highest when light intensity is high AND plant
is experiencing water stress that ia sufficient to cause the plant to
close its stomata
• Under these two conditions oxygen produced by the light dependent
reaction accumulates in the mesophyll, AND the co2 in the mesophyll
air spaces are used up by the light independent reaction in a C3 plant.
• The concentration of oxygen in the mesophyll is relatively high
compared to co2 concentration. Relative concentration of co2 is low.
• The relative turnover number for oxygen and co2 at rubisco depends
What is a C3 and a C4 plant
the first product of carbon fixation in a 3 carbon compound, that is
glycerate 3 phosphate in a C3 plant.

the first product of carbon fixation in a 4 carbon compound,

oxaloacetate in a C4 plant
Krantz (crown) anatomy of leaf of C4
• Leaf have parallel veins, monocot/ monocotyledonous plant ()
• Some monocot plants are C4 plants
• Circle
• Concentric (share the same centre) cylinders in the leaf, but appear like concentric circles
in the leaf cross section
• Inner ( not exposed to /shield from atmospheric oxygen) cylinder is the bundle sheath
cells
• Outer (significance is the cell is exposed to carbon dioxide from the atmosphere) cylinder
is made up of the mesophyll cells. No air spaces between adjacent mesophyll cells. The
only air space is the substomatal air space
• A leaf will have parallel veins (vascular bundles), each vein is enclose in these two
concentric cylinders of cells (two layers of cells)
• Every mesophyll cell has a contact surface with a bundle sheath cell
• Bundle sheath cells and mesophyll cells are connected by plasmodesmata at the surface
of contact
Hatch -Slack pathway
The first step is carried out by an enzyme present only in
mesophyll cells called PEP carboxylase. This enzyme
adds CO2 to phosphoenolpyruvate (PEP), forming the
four-carbon product oxaloacetate.

Oxaloacetate is reduced to malate using a molecule of NADPH

PEP carboxylase has a much higher affinity for CO2 than does rubisco and no affinity for O2. Therefore, PEP carboxylase
can fix carbon efficiently when rubisco cannot—that is, when it is hot and dry and stomata are partially closed, causing CO 2
concentration in the leaf to be lower and O 2 concentration to be relatively higher.
2
After the CO2 is fixed in the mesophyll cells, the four carbon products (malate in the example shown in
Figure 10.20) are exported to bundle-sheath cells through plasmodesmata (see Figure 6.29).
3
Within the bundle-sheath cells, the four-carbon
compounds (malate) release CO ,(produce high
2

concentration) which is re-fixed into organic material

by rubisco and the Calvin cycle.
The same reaction regenerates pyruvate, which is
transported to mesophyll cells.
There, ATP is used to convert pyruvate to PEP, which
can accept addition of another CO , allowing the
2

reaction cycle to continue.

Why is photosynthesis in C4 plant faster than
in the C3 plant
• PEP carboxylase has a much higher affinity for CO2 than does rubisco.
• Smaller KM . That the enzyme reach V max at lower substrate (CO2 )
concentration
PEP carboxylase no affinity for oxygen (do not use chemical symbol in sentences)
No photorespiration will occur in the mesophyll cell.
Efficiency of PEP carboxylase is translated into high concentration of malate in the
bundle sheath cells, resulting in high concentration of carbon dioxide in bundle
sheath cell.
Photorespiration in the bundle sheath cell is prevented (competitively) or very
reduced due to the high concentration of carbon dioxide
This ATP can be thought of, in a sense, as the “price” of concentrating
CO in the bundle-sheath cells. To generate this extra ATP, bundle-sheath
2

cells carry out cyclic electron flow, the process described earlier in this
chapter (see Figure 10.16).
In fact, these cells (BUNDLE SHEATH) contain PS I but no PS II, so
cyclic electron flow is their only photosynthetic mode of generating ATP.
(without PS II bundle sheath cell do not carry out non-cyclic
photophosphorylation, therefore do not produce oxygen) In effect, the
mesophyll cells of a C plan
4

bundle sheath cells are separated from the substomatal air space by a
layer of mesophyll cells and thick cell wall
Advantages and disadvantages of C4 and C3 carbon
fixation
•C4 plants grow better than C3 plants under hot, dry conditions
when plants must close their stomata to conserve water – with
stomata closed, CO2 levels in the interior of the leaf fall, and O2
levels rise.
•C3 plants grow better than C4 plants under cool, moist
conditions when plants can open their stomata, because C3 plants
do not incur the additional cost of ATP of C4 carbon fixation.
 Analysing and reflecting on the Hatch-Slack pathway

In effect, the mesophyll cells of a C4 plant pump CO2 into the bundle-sheath cells, keeping the CO2
concentration in those cells high enough for rubisco to bind CO2 rather than O2.

The cyclic series of reactions involving PEP carboxylase and the regeneration of PEP can be
thought of as an ATP powered pump that concentrates CO2. In this way, C4 photosynthesis
spends ATP energy to minimize photorespiration and enhance sugar production. This adaptation
is especially advantageous in hot regions with intense sunlight, where stomata
partially close during the day, and it is in such environments that C4 plants evolved and thrive
today.
Why do c4 plants lose some of its advantage
in temperate climate
• Temperate climate occur at higher latitudes
• Light intensity at higher latitude is lower
• C3 plants can keep their stomata open in cooler climates
C4 photosynthesis is considered more efficient than C3
photosynthesis because it uses less water and resources. On our
planet today, the world population and demand for food are
rapidly increasing. At the same time, the amount of land suitable
for growing crops is decreasing due to the effects of global
climate change, which include an increase in sea level as well
as a hotter, drier climate in many regions. To address issues of
food supply, scientists in the Philippines have been working
on genetically modifying rice—an important food staple that
is a C3 crop—so that it can instead carry out C4 photosynthesis.
Results so far seem promising, and these researchers estimate that the
yield of C4 rice might be 30–50% higher than C3 rice
 CAM Plants
A second photosynthetic adaptation to arid conditions has
evolved in many succulent (water-storing) plants, numerous
cacti, pineapples, and representatives of several other
plant families.
These plants open their stomata during the night and close them during
the day,
just the reverse of how
other plants behave. Closing stomata during the day helps
desert plants conserve water, but it also prevents CO2 from entering the
leaves.
 What is the advantage of opening the stomata during the night?
Facts about atmospheric conditions of night compared to day
Higher humidity than day.
Lower temperature than during the day

Rate of transpiration is at night is lower.

Rate of water loss per hour is less than day but the gain of co2 is same as
day.
What is the challenge of fixing co2 at night
but not reducing the organic acid produced?
• Organic acid reduce the pH of the cytoplasm, stroma of chloroplast
• Can / may denature proteins
• The solution// adaptation
• Store the malate in the vacuole where there are no proteins
• Vacuole has a capacity limit. This limit the growth rate of CAM plants
• (pineapple is expensive, cacti have thorns to defend their biomass
from herbivores. Recovering from herbivory is slow)
 NIGHT
During the night, when their stomata are open, these plants take up CO2 and
incorporate it into a variety of organic acids. PEP is used to fix carbon dioxide .
Oxaloacetate formed. OAA is reduced to malate. Malate is stored in the
vacuole.
This mode of carbon fixation is called crassulacean acid metabolism, or CAM,
after the plant family Crassulaceae, the succulents in which the process was
first discovered.
The mesophyll cells of CAM plants store the organic acids they make during
the night in their vacuoles until morning, when the stomata close.
DAY
During the day, when the light dependent reactions can supply ATP and
NADPH for the Calvin cycle, CO2 is released from the organic acids made
the night before to become incorporated into sugar in the
 Similarity
Notice in Figure 10.21 that the CAM pathway is similar to
the C4 pathway in that CO2 is first incorporated into organic
intermediates before it enters the Calvin cycle.
The difference
is that in C4 plants, the initial steps of carbon fixation are
separated structurally from the Calvin cycle, whereas in CAM
plants, the two steps occur within the same cell but at separate
times. / temporal separation (Keep in mind that CAM, C4, and C3 plants
all eventually use the Calvin cycle to make sugar from carbon dioxide.)
m uses PEP carboxylase to initially make a 4-carbon compound, that then releases CO2 to rubisco in leaf cells that are exposed to little oxygen. While this mechanism reduces the oxyge

af fall, and O2 levels rise.

xation.