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C
3
, C
4
and CAM. Regulation of The Activity of
Photosynthesis

C
4

A number of plants display an increased and more efficient net photosynthesis
during strong light intensities. A prime example are the Gramineae of warmer
regions like maize or sugar-cane.
At the beginning of the sixties observed H. KORTSCHAK (Hawaiian Sugar
Planter's Association) that the first product of photosynthesis in sugar-cane is
not the C
3
unit 3-phosphoglycerate but a unit with four C-atoms. The
Australian plant physiologist M. D. HATCH and his English colleague C. R.
SLACK confirmed this result and identified the compound as oxaloacetate
(OAA). It is produced by the addition of one molecule of carbon dioxide
to phosphoenolpyruvate (PEP). The cycle is also known as the HATCH-
SLACK-cycle or the C
4
cycle. Plants with this cycle are called C
4
-plants
(and CAM plants, respectively) in contrast to C
3
plants where the carbon
dioxide is directly fed into the CALVIN cycle. The oxaloacetate is usually
converted into malate of which the carbon dioxide is split off again with the
help of an enzyme.


This carbon dioxide is now bound by ribulose-1,5-diphosphate and
assimilated viathe CALVIN cycle. :
Some species use malate instead of aspartate
oxaloacetate + L-glutamate > aspartate + alpha-ketoglutarate.
The reversible binding of carbon dioxide has the function to accumulate and
store CO
2
. The process consumes energy, so that it could also be spoken of a
carbon dioxide pump. It should be mentioned that the HATCH-SLACK cycle
requires two molecules of ATP are per fixed carbon dioxide.






Photosynthesis of C
4
plants. CO
2
is bound to phosphoenolpyruvate (PEP)
in mesophyll cells. The product is oxaloacetate. The next step generates malate.
In the cells of the vascular bundle sheath, the 'Kranz' cells, is carbon dioxide
split off the malate and fed into the CALVIN cycle. The pyruvate is transported
back into the mesophyll cells (active transport) and is with the help of
additional ATP phosphorylated to PEP.




The anatomy of C
4
leaves with so-called 'Kranz' cells differs fundamentally
from that of C
3
plants. The chloroplasts of C
3
plants are of homogeneous
structure, while two types of chloroplasts occur in C
4
plants. The mesophyll
cells contain normal chloroplasts, that of the vascular bundle sheath
have chloroplasts without grana , i.e. they are partially impaired in function.
This peculiarity does not affect the CALVIN cycle, it concerns only the light
reactions of photosynthesis. The first binding of carbon dioxide (the HATCH-
SLACK reaction) occurs in the mesophyll cells, the incorporation into
carbohydrates (the CALVIN cycle) in the cells of the vascular bundle sheath.
Both processes of photosynthesis are spatially separated.

The Crassulacean Acid Metabolism (CAM)
CAM is the abbreviation of Crassulacean acid metabolism. The name points at
the fact that this pathway occurs mainly in Crassulacean species (and other
succulent plants). The chemical reaction of the carbon dioxide accumulation is
similar to that of C
4
plants but here are carbon dioxide fixation and its
assimilation not separated spatially but in time. CAM plants occur mainly in
arid regions. The opening of the stomata to take up carbon dioxide is always
connected with large losses of water. To inhibit this loss during intense sun (the
transpiration via the cuticle remains intact) has a mechanism developed that
allows the uptake of carbon dioxide during the night. The prefixed carbon
dioxide is stored in the vacuoles as malate (and isocitrate) and is used during
the daytime for photosynthesis.

Which Metabolism Goes With Which Conditions?
The enzyme that catalyzes the primary carbon dioxide fixation of C
4
and CAM
plants is phosphoenolpyruvate carboxylase (PEPC). Its affinity for carbon
dioxide is by far higher than that of Rubisco, the first enzyme of the CALVIN
cycle. As a consequence are C
4
plants able to use even trace amounts of carbon
dioxide. PEPC occurs in small amounts (roughly 2 - 3 %) also in C
3
plants,
where it, too, has a key position in the metabolic regulation.
Carbon dioxide yield of C
4
and
C
3
plants of open grasslands in
different parts of the world. In
temperate regions is the rather low
light intensity decisive for the
disadvantage of C
4
plants. C
3
plants
have an advantage due to their low
rate of photorespiration and because
they need no energy for the previous
fixation of CO
2
. (J. R. EHRLICHER,
1978).



In growing roots (of maize seedlings, for example) does PEPC help to supply
the lipid synthesis with NADH + H
+
. The following reactions take place:
1. phosphoenolpyruvate + HCO
3
- > oxaloacetate + P
i

2. oxaloacetate > malate. - During this step is NADH + H
+
oxidized
to NAD
+
.
3. malate > pyruvate + CO
2
.
During the last reaction is NAD reduced to NADH + H
+
.
In the root nodules of leguminosae is nitrogen fixed. Enough carbon bodies
have to be supplied in order to incorporate the ammonia produced by the
bacteria. the CO
2
-binding via PEPC-reaction thus is an important supplement
Furthermore produces the PEPC intermediates of the citric acid
cycle(oxaloacetate and / or malate), a back-up in case of shortages. The activity
of PEPC is controlled by extern factors, the day length is decisive. In some
cases have different isoenzymes be found in different tissues. Their production
is controlled by different triggers.
C
3
plants can loose up to 20 percent of the carbon fixed in the CALVIN
cycle at intense radiation. Under strong light is the photorespiration 1.5 - 3.5
times as high as the usual respiration in darkness. In C
4
plants becomes the
photorespiration drastically reduced, it may even not be detectable any more. In
other words:
The net rate of photosynthesis (and consequently also the net production of
biomass) of C
4
plants is far larger at high light intensities than that of C
3
plants.
The optimal temperature of photosynthesis is below that of the respiration in
darkness. As a consequence are losses caused by respiration larger at high than
at low temperatures. Where light is a limiting factor and temperatures are low
(i.e. in temperate climatic zones) have C
3
plants the advantage, C
4
plant do
hardly occur (one of the exceptions is Spartina townsendii). C
4
plants, nearly
always herbs or shrubs, are more successful in the open country of warmer
zones.
It has to be mentioned that two molecules of ATP are consumed in the
HATCH-SLACK cycle
C
4
plants belong to numerous, phylogenetically not related monocotyledonous
and dicotyledonous families. Moreover have C
4
activities also been detected in
the blue-green alga Anacystis nidulans as well as in some dinoflagellates.
Since the alternative C
3
or C
4
is accompanied by considerable changes of the
leaf anatomy has it to be assumed that the genetic potential for both pathways is
quite common in the plant kingdom and that, depending on the ecological
needs, one way is chosen by a species while a related species may choose the
other one.
A well-studied example is the genus Atriplex, where both ways are realized.
The C
3
plants belong to one phylogenetic group, the C
4
plants to another. In
some cases can hybrids of C
3
and C
4
species be generated.




Influence of different parameters on the efficiency of the carbon dioxide uptake
(ordinate) of a C
3
plant (Atriplex patula, yellow line) and a C
4
plant (Atriplex
rosea, green line). Measured parameters (from left to right): light intensity, leaf
temperature and concentration of carbon dioxide within the intercellular space
(according to O. BJRKMAN and J. BERRY, 1973).

In several plant species of the genera Zea, Mollugo, Moricandia, Flaveria, etc.
occur both types of CO
2
fixation within one plant. In younger plants is usually
the C
3
-, in older ones the C
4
pathway taken. The amount of C
4
is controlled by
environmental factors.
CAM: Advantages and Disadvantages. CAM has been detected in more than
1000 angiosperms of 17 different families. It is usually accompanied by
succulence, though not all Crassulaceae, for example, display CAM and
succulence is no precondition of CAM. Tillandsia usneoides of the bromelia
family is not succulent, but uses CAM. Mesemryanthemum crystallinum (a
plant with succulent leaves) can use the C
3
pathway but switches to CAM when
growing in saline soils. Under experimental conditions can the shift be
achieved by increasing the NaCl concentration of the nutrient medium (K.
WINTER and D. J. von WILLERT, 1972). While the advantage of C
4
plants
comes in useful under high light intensities, is the degree of the CAM influence
in CAM plants regulated mainly by temperature, atmospheric humidity and
salinity. Both strong and weak CAM plants are known. In weak CAM plants
becomes CAM only apparent at certain differences between day and night
temperature. CAM plants that store a lot of malate and due to the thus high
osmotic value also a lot of water, are usually less frost resistant than C
3
plants.
Because of the high concentration of acid are they less heat resistant, too.
Species of arid regions are therefore forced to break their pool of malate down
during the daytime (R. LSCH and H. KAPPEN, Universitt Kiel, 1985).
Usually do the C
4
pathway and CAM exclude each other. An exception is the
succulent C
4
dicotyledon Portulaca oleracea that is able to choose the optimal
pathway. under natural conditions


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