Photorespiration

Photorespiration
• Photorespiration (also known as the oxidative photosynthetic carbon cycle,
or C2 photosynthesis) refers to a process in plant metabolism where the
enzyme RuBisCO oxygenates RuBP, wasting some of the energy produced
by photosynthesis.The desired reaction is the addition of carbon dioxide to
RuBP (carboxylation), a key step in the Calvin–Benson cycle, but
approximately 25% of reactions by RuBisCO instead add oxygen to RuBP
(oxygenation), creating a product that cannot be used within the Calvin–
Benson cycle.
• This process reduces the efficiency of photosynthesis, potentially reducing
photosynthetic output by 25% in C3 plants. Photorespiration involves a
complex network of enzyme reactions that exchange metabolites between
chloroplasts, leaf peroxisomes and mitochondria.
• The oxygenation reaction of RuBisCO is a wasteful
process because 3-phosphoglycerate (G3P) is created at
a reduced rate and higher metabolic cost compared with
RuBP carboxylase activity. While photorespiratory
carbon cycling results in the formation of G3P
eventually, around 25% of carbon fixed by
photorespiration is re-released as CO2 and nitrogen, as
ammonia. Ammonia must then be detoxified at a
substantial cost to the cell. Photorespiration also incurs
a direct cost of one ATP and one NAD(P)H.
• While it is common to refer to the entire process as
photorespiration, technically the term refers only to the
metabolic network which acts to rescue the products of
the oxygenation reaction (phosphoglycolate).
 Oxygenation
• The first step of photorespiration is oxygenation. It
is catalyzed by RUBISCO and converts RUBP to
one molecule of 3PGA and one molecule of 2PG,
which is a 2 carbon compound with one phosphate
group.
• The oxygenation reaction is influenced by
environmental factors. In normal air at 25°C, a well
watered plant fixes oxygen once for every 3 carbon
dioxide fixations. If the temperature increases or the
plant is short of water, however, the occurrence of
oxygenation increases.
1- Temperature
• Increasing temperature increases oxygenation in two ways:
• The first of these is that RUBISCO becomes less specific for CO 2 as the
temperature rises. Apparently increasing temperature reduces the
specificity of the active site of the enzyme such that it performs
oxygenation more frequently.
•  
The second way in which temperature affects oxygenation is by its
effect on the relative solubilities of O2 and CO2. Both O2 and CO2 must
dissolve in the water of a leaf cell before they are available to react with
RUBISCO. The solubility of any gas in water decreases as temperature
increases so that there is less O2 and CO2 dissolved in a cells when they
are at higher temperature. The solubility of O 2 decreases with
temperature less than the solubility of CO 2, however. This means that as
temperature rises, the ratio of CO2 to O2dissolved in the cell solution
decreases. This decreasing ratio favors oxygenation
2- Lack of water
• As a plant becomes short of water, it closes its
stomata. Inside a leaf with closed stomata, O 2 is
produced by Photosystem II and builds up. CO 2 is
consumed by carboxylation and drops to low
levels. Thus, the ratio of CO2 to O2 inside a leaf
becomes very low when stomata are closed,
favoring oxygenation.
3. High CO2 abolishes oxygenation
• Increasing CO2 concentration to 1% abolishes oxygenation.
At this concentration, CO2 outcompetes O2 for the active
site of RUBISCO. This effect is significant in two ways:
•
First, the concentration of CO2 in the Earth's atmosphere is
rising because of deforestation and the burning of fossil
fuels. It has risen by 30% since 1850. As CO2 rises,
photorespiration and its metabolic costs to plants
decrease.
• Second, some plants and algae have evolved ways to avoid
photorespiration by increasing the concentration of CO2 in
the vicinity of RUBISCO
 Processing of 2 phosphoglycolate
• 3 PGA is converted back to RUBP by the Calvin
cycle during its normal operation. 2 PG cannot be
converted to RUBP by the Calvin cycle and is
instead processed to 3 PGA by an elaborate series
of steps that involve 3 different organelles: the
chloroplast, the mitochondrion, and the
peroxisome
The important steps of processing that occur in each
organelle are:
i. Chloroplast steps
The original oxygenation reaction by RUBISCO
occurs in the chloroplast. This is where O2 is
consumed by photorespiration. The 2PG that is
formed is dephosphorylated and exported as glycolate
to the peroxisome.
The final steps of processing also occur in the
chloroplast. Glycerate is converted to 3PGA at the
expense of one ATP. This ATP is part of the energetic
cost of photorespiration.
Peroxisome steps
• In the peroxisome, glycolate from the chloroplast is converted to glyoxylate. This
reaction consumes O2 and generates hydrogen peroxide. The O2 is released again,
however, as the hydrogen peroxide is converted to water by the enzyme catalase.
• Glyoxylate is converted to the amino acid glycine with donation of an amino group that
ultimately comes from the amino acid serine. These are the "trans amination" reactions
of photorespiration. They shift amino groups from one molecule to another.
 iii. Mitochondrion steps
• The mitochondrion is where 2 carbon amino acids are converted into 3 carbon amino
acids. 2 glycine molecules, 2 C each, are combined to make one 3 carbon serine. In the
process, 1 NH3 is released and so is 1 CO2. This is the CO2 produced by
photorespiration. The enzyme that catalyzes this reaction is glycine decarboxylase.
Serine is exported to the peroxisome where its amino group is removed, converting it to
glycerate. The glycerate is then exported to the chloroplast where it is converted to
3PGA at the expense of one ATP. It should be noted that transport of photorespiration
intermediates between organelles requires a special membrane transporter for every
different molecule and organelle.
• Overall, 1 3PGA is recovered / 2PG processed, thus 3/4 of the carbon lost by
oxygenation is recovered. The NH3 is released and so is 1 CO2 released by
photorespiration must be reassimilated, with some expense of ATP and reducing power.
Also, the phosphorylation of glycerate back to 3 PGA costs one ATP per reaction.
The costs of photorespiration
• In normal air at 25°C, photorespiration
decreases the efficiency of CO2 assimilation by
40%. This decrease in efficiency results from
the use of ATP to phosphorylate glycerate and
other costs associated with glycolate
processing
The benefits of photorespiration
• Photorespiration is either a necessary evil of plant
metabolism or it may have some adaptive function
that is not apparent. Some have proposed that
photorespiration allows plant leaves to use up
excess light energy and reduce photooxidative
damage when the plant is water-stressed and the
stomata are closed. There are many plants,
however, that do no photorespiration and are not
especially sensitive to drought. C4 plants are an
example of these.