http://wc.pima.edu/~bfiero/tucsonecology/plants/plants_photosynthesis.

htm

TYPES OF PHOTOSYNTHESIS
This page introduces you to photosynthesis and explains the three types of
photosynthesis and their relevance for desert adaptation.

Concepts:

• Photosynthesis: the bonding together of CO2 (carbon dioxide) with H2O

(water) to make CH2O (sugar) and O2 (oxygen), using the sun's energy.
The sugar contains the stored energy and serves as the raw material from
which other compounds are made. Respiration is the opposite of
photosynthesis -- the stored energy in the sugar is released in the
presence of oxygen, and this reaction releases the CO2 and H2O
originally jammed together by the sun's energy.
• Stomata: the "pores" in leaves (and stems) through which CO2 is taken in
and O2 is released during photosynthesis. Plants control when stomata
are open or closed and the width of the opening (formed by two guard
cells that expand and contract to open and close the space between
them).
• Transpiration: the water that evaporates out of stomata when they are
open. This pulls more water and nutrients up to the top of the plant, but
causes the plant to lose water and potentially dehydrate.
• Water Use Efficiency (WUE): How good the plant is at bringing in carbon
dioxide for photosynthesis without losing much water out of its stomata.
More specifically, it is the ratio of carbon dioxide intake to water lost
through transpiration.
• Photorespiration: Under high light and high heat, the enzyme
(RUBISCO) that grabs carbon dioxide for photosynthesis may grab
oxygen instead, causing repiration to occur instead of photosynthesis,
thus causing a slowing of the production of sugars from photosynthesis.

The three types of photosynthesis are C3, C4, and CAM. C3 photosynthesis is
the typical photosynthesis that most plants use and that everyone learns about in
school (it was all we knew about until a few decades ago). C4 and CAM
photosynthesis are both adaptations to arid conditions because they result in
better water use efficiency. In addition, CAM plants can "idle," saving precious
energy and water during harsh times, and C4 plants can photosynthesize faster
under the desert's high heat and light conditions than C3 plants because they
use an extra biochemical pathway and special anatomy to reduce
photorespiration. Below are the details.

C3 Photosynthesis : C3 plants.
 • Called C3 because the CO2 is first incorporated into a 3-carbon
compound.
• Stomata are open during the day.
• RUBISCO, the enzyme involved in photosynthesis, is also the enzyme
involved in the uptake of CO2.
• Photosynthesis takes place throughout the leaf.
• Adaptive Value: more efficient than C4 and CAM plants under cool and
moist conditions and under normal light because requires less machinery
(fewer enzymes and no specialized anatomy)..
• Most plants are C3.

C4 Photosynthesis : C4 plants.

• Called C4 because the CO2 is first incorporated into a 4-carbon

compound.
• Stomata are open during the day.
• Uses PEP Carboxylase for the enzyme involved in the uptake of CO2.
This enzyme allows CO2 to be taken into the plant very quickly, and then it
"delivers" the CO2 directly to RUBISCO for photsynthesis.
• Photosynthesis takes place in inner cells (requires special anatomy called
Kranz Anatomy)
• Adaptive Value:
o Photosynthesizes faster than C3 plants under high light intensity
and high temperatures because the CO2 is delivered directly to
RUBISCO, not allowing it to grab oxygen and undergo
photorespiration.
o Has better Water Use Efficiency because PEP Carboxylase brings
in CO2 faster and so does not need to keep stomata open as much
(less water lost by transpiration) for the same amount of CO2 gain
for photosynthesis.
• C4 plants include several thousand species in at least 19 plant families.
Example: fourwing saltbush pictured here, corn, and many of our summer
annual plants.

CAM Photosynthesis : CAM plants. CAM stands for Crassulacean Acid

Metabolism

• Called CAM after the plant family in which it was first found
(Crassulaceae) and because the CO2 is stored in the form of an acid
before use in photosynthesis.
• Stomata open at night (when evaporation rates are usually lower) and are
usually closed during the day. The CO2 is converted to an acid and stored
during the night. During the day, the acid is broken down and the CO 2 is
released to RUBISCO for photosynthesis
• Adaptive Value:
 o Better Water Use Efficiency than C3 plants under arid conditions
due to opening stomata at night when transpiration rates are lower
(no sunlight, lower temperatures, lower wind speeds, etc.).
o May CAM-idle. When conditions are extremely arid, CAM plants
can just leave their stomata closed night and day. Oxygen given off
in photosynthesis is used for respiration and CO2 given off in
respiration is used for photosynthesis. This is a little like a perpetual
energy machine, but there are costs associated with running the
machinery for respiration and photosynthesis so the plant cannot
CAM-idle forever. But CAM-idling does allow the plant to survive
dry spells, and it allows the plant to recover very quickly when
water is available again (unlike plants that drop their leaves and
twigs and go dormant during dry spells).
• CAM plants include many succulents such as cactuses and agaves and
also some orchids and bromeliads

http://faculty.weber.edu/sharley/2104/photosynthesis.html

2. The Calvin Cycle

There are three phases to the Calvin cycle: carboxylation, reduction, and
regeneration (Figure 2)
A. Carboxylation
Carbon dioxide and RuBP (a C5 sugar) are combined to give two molecules of
PGA (a C3 acid).
The enzyme that catalyzes this reaction is abbreviated rubisco.
B. Reduction
This phase uses the 2/3 of the ATP and all of the NADPH produced during the
light reactions.
Each PGA molecule is reduced to PGald (a C3 sugar).
C. Regeneration
This phase uses the last of the ATP to turn a bunch of PGald molecules into a
bunch of RuBP molecules to regenerate the cycle.
D. what the abbreviations stand for:
rubisco = RuBP carboxylase and oxygenase
RuBP = ribulose-1,5-bisphosphate
PGA = 3-phosphoglyceric acid (3-phosphoglycerate)
PGald = 3-phosphoglyceraldehyde (also known as glyceraldehyde-3-phosphate)
ATP = adenosine triphosphate
ADP = adenosine diphosphate
NADP+/NADPH = nicotinamide adenine dinucleotide phosphate
(oxidized/reduced)
Once you have lots of PGald, it can be used to make glucose, fructose, sucrose,
and starch.
(The reactions of respiration, besides providing a means to change the stored
calories of sugars into useable energy, let a cell start the process of converting
carbons from carbohydrates into a variety of molecules: amino acids,
nucleotides, pigments, hormones, etc.)

1950s: Melvin Calvin and associates: used 14C (available for labeling
experiments after WWII) and 2-D paper chromatography (developed as a
technique in the 1940s) to identify the carbon fixation steps of photosynthesis.
The first stable product to contain the label was PGA. At later stages, the label
appeared in triose-P and then the various sugars of the regeneration steps
(reductive pentose phosphate pathway). They used Chlorella, a unicellular green
alga, as their experimental organism. With Chlorella, they were able to take
small samples at various time intervals and rapidly kill all the cells simultaneously
to stop the reactions.

Photosynthesis as a Leaf Event

Besides looking at photosynthesis as a chloroplast event, you need to remember

that it is also a leaf event.
Inside of a leaf are three tissues (functional collections of cells):
1. epidermis
holes for gas exchange called stomata (guard cells open and close the holes)
covered by a wax layer called cuticle
2. vascular tissue
xylem + phloem together in a vascular bundle (vein)
3. mesophyll (ground tissue)
tightly packed layer of cells = palisade mesophyll
loose cell layer with lots of air spaces = spongy mesophyll
Most plants open their stomata during the day (light) so CO2 enters the leaf for
photosynthesis. Downside: water evaporates out of the stomata whenever they
are open. Evaporation is fastest when the temperatures are highest, which would
also be during the day. The stomata close at night when photosynthesis is not
going on (no need to let in CO2).
Some plants have a system that lets them open their stomata at night to collect
and store CO2. During the day, they can close their stomata to conserve water,
but still do photosynthesis. These plants are known as CAM plants. CAM ==
Crassulacean acid metabolism. CAM was first discovered in members of the
Crassulaceae family. CAM has since been found in many angiosperm families
(both monocots and dicots), a seedless vascular plant, and a gymnosperm.
CAM plants grow in arid habitats: deserts, alpine regions, as epiphytes. CAM
plants have at least some succulence (water storing). Two CAM plants are
important from the money end of things: pineapple and orchids.
PEP = phosphoenolpyruvate, a C3 acid.
CO2 can be attached to PEP by the enzyme PEP carboxylase.
At night, the stomata are open. Starch is broken down to produce PEP. PEP
combines with CO2 to form a C4 acid. This C4 acid is stored in the vacuole.
During the day, the stomata close. The C4 acid is broken down to release CO2
and a C3 acid. The C3 acid is converted back to starch. The CO2 enters the
Calvin cycle. (Figure 3)
CAM is estimated to occur in ~ 10% of plant species. C3 photosynthesis (where
the only carbon reactions are the Calvin cycle ones) occurs in ~ 89% of species.
(C3 plants include wheat, rice, daisies, petunias, roses, fruit trees, and conifers.)
The remaining ~1% do C4 photosynthesis. Although C4 accounts for only a
fraction of the photosynthesis it attracts a lot of study because (1) it is a highly
efficient form of photosynthesis and (2) it accounts for the high productivity of
such major crops as corn, sugar cane, sorghum, and millet.
rubisco = RuBP carboxylase and oxygenase
O2 + RuBP ------> PGA + phosphoglycolyate (C2 acid )
In a series of reactions that occur in the chloroplast, peroxisome, and
mitochondria, 2 molecules of phosphoglycolate are converted to PGA + CO2.
(Figure 4) The CO2 that is released by this process because of the oxygenase
reaction if rubisco is photorespiration: extra CO2 released in the light compared
to plants under the same conditions in the dark. Photorespiration is highest
under conditions of high temperature, high light intensity, and low water. Under
these conditions, a C3 plant might lose 50% of its carbon via photorespiration.
How can you decrease photorespiration? Keep rubisco away from O2. Some
plants do this by engaging in C4 photosynthesis.
C4 plants have a distinctive leaf anatomy. There is a prominent ring of cells
around the vascular bundles = the bundle sheath. The mesophyll cells form a
ring that is tightly appressed to the bundle sheath cells. Kranz anatomy.
In a mesophyll cell, CO2 and PEP combine to form a C4 acid. The C4 acid is sent
to a bundle sheath cell. In the bundle sheath cell, the CO2 is released from the C4
acid and enters the Calvin cycle. The C3 acid that remains goes back to the
mesophyll cell, is made back into PEP, and is ready to carry more CO2. (Rubisco
is located only in the bundle sheath cells.) So, a CO2 shuttle system delivers CO2
to rubisco; the leaf anatomy keeps O2 away from the bundle sheaths. Result ==>
no photorespiration. (Figure 5)
Downside to C4: the CO2 shuttle is not a free ride. It adds 2 ATP to the standard
3 ATP (for the Calvin cycle) needed per CO2. So C4 is only cost effective for
plants in an environment where photorespiration could be so great as to be
detrimental.

1965: Kortschack, Hartt, and Burr repeated the experiments done by Calvin’s
group, using sugar cane as the experimental organism. The results indicated
that sugar cane made malate (C4 acid) as the first stable product, not PGA (C3
acid). Soon afterwards, Hatch and Slack confirmed these results and showed
that this form of photosynthesis was found in a number of plants, both monocots
and dicots.