Photorespiration

Photorespiration (also known as the oxidative photosynthetic carbon cycle or C2

cycle) refers to a process in plant metabolism where the enzyme Rubisco
oxygenates RuBP, wasting some of the energy produced by photosynthesis.
Photorespiration is a wasteful pathway that competes with the Calvin cycle. It
begins when rubisco acts on oxygen instead of carbon dioxide. 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).
Photorespiration is also a. biochemical process in plants in which, especially under
conditions of water stress, oxygen inhibits the Calvin cycle, the carbon fixation
portion of photosynthesis.

Photorespiration begins in chloroplast when rubisco attaches o2 to RuBP in its

oxygenase reaction. Two molecules are produced: a three-carbon compound, 3-
PGA and a two-carbon compound, phosphoglycolate.

RuBP+02 GP+PG+C02

Why Plant Face Photorespiration

Photorespiration results in the light-dependent uptake of oxygen and release of

carbon dioxide and is associated with the synthesis and metabolism of a small
molecule called glycolate. Photorespiration takes place in green plants at the
same time that photosynthesis does. Because in photosynthesis carbon dioxide is
taken in, and in photorespiration carbon dioxide is given off, these two processes
work against each other.

The end result is that photorespiration decreases the net amount of carbon
dioxide which is converted into sugars by a photosynthesizing plant. By interfering
with photosynthesis in this way, photorespiration may significantly limit the
growth rate of some plants.

Photorespiration reduces the efficiency of photosynthesis because:

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-ATP is used up.

-RuBP is regenerated

-phosphoglycolate is removed.

RuBP oxygenase-carboxylase (Ribulose-1,5-bisphosphate carboxylase-oxygenase),

a key enzyme in photosynthesis. In the process of carbon fixation, rubisco
incorporates carbon dioxide into an organic molecule during the first stage of the
Calvin cycle. Rubisco is so important to plants that it makes up to 30% more of the
soluble protein in a typical plant leaf. But rubisco also has always using CO2 as a
substrate, it sometimes picks up O2 instead.
This side reaction initiates a pathway called photorespiration, which, rather than
fixing carbon, actually leads to the loss of already-fixed carbon as CO2.
Photorespiration wastes energy and decreases sugar synthesis, so when rubisco
initiates this pathway, it's committing a serious molecular mess.
So, In conclusion Photorespiration is the process of light-dependent uptake of
molecular oxygen (O2) concomitant with release of carbon dioxide (CO 2) from
organic compounds. And also the process of light-dependent uptake of molecular
oxygen (O2) concomitant with release of carbon dioxide (CO2) from organic
compounds.

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Difference Between Photorespiration & Respiration
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Photorespiration is a type of respiration that occurs in plant in the presence of
light and at higher concentrations of oxygen. The photorespiration is also called
c2 cycle or glycolate metabolism.
Respiration (also called normal or dark respiration) is a metabolic pathway which
releases energy rich molecules by the breakdown of sugar molecules such as
glucose.
Even though the photorespiration and respiration occur in the presence of
oxygen, the two pathways are independent and show considerable differences.

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 Photorespiration vs Respiration

Photorespiration Respiration
1 Also called c2 cycle or glycolate cycle. Also called normal or dark cycle.

2 Occurs only in green plants. Occurs in all aerobic and anaerobic

organisms.
3 It takes place only in the presence of light. It occurs both in light and dark.
4 Commonly occurs in c3 plants. Occurs in all plants.
5 It consumes energy. It liberates energy.
6 Does not involve glycolysis, Krebs cycle Aerobic respiration is completed in
and ETS. glycolysis, Krebs cycle and ETS.
7 Is completed in three organelles- Is completed in cytoplasm and
chloroplast, peroxisomes and mitochondria.
mitochondria.
8 End products are CO2 and PGA. End products are CO2 and water
9 No phosphorylation occurs Substrate level and oxidative
phosphorylation occurs
10 Release co2 at the end of Co2 is released at several places.
photorespiration.
11 One molecule of ammonia is produced. No ammonia is produced during
respiration
12 It produces H2O2 H2O2 is not formed
13 It is an energy wastage process. It is an energy producing process.
14 It decreases the net productivity of green It increases the net productivity of
plants green plants.
Advantage & Disadvantage of photorespiration

Disadvantages of Photorespiration:

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Photorespiration reduces the efficiency of photosynthesis as in this process o2 is
used to oxidize RUBP in formation of 3PGA and 2 carbon compound
phosphoglycolic acid.
Biochemical studies indicate that photorespiration consumes ATP and NADPH, the
high-energy molecules made by the light reactions of photosynthesis.
It is also considered as a wasteful process as extra energy is consumed for o2
fixation in the form of ATP but on other hand the pathway reuses ¾ of the carbon
Phosphoglycerate by regenerating 3GPA.
Advantages of photorespiration:
Photorespiration removes toxic metabolic intermediates.
Photorespiration is a major source of h2o2 in plants.
Photorespiration connects metabolic compartments of the cell and facilitate
transport among organelles as peroxisome, mitochondria and chloroplast.

Biological Adaptation to Minimize photorespiration

The C4 and CAM pathways are two adaptations—beneficial features arising by

natural selection—that allow certain species to minimize photorespiration. These
pathways work by ensuring that Rubisco always encounters high concentrations
of CO2 making it unlikely to bind to O2.
C4 and CAM photosynthesis both use the enzyme Phosphoenolpyruvate
carboxylase (PEPC) to add CO2 to a 4-Carbon sugar. PEPC is faster than RUBISCO,
and more selective for CO2.

C4 plants capture carbon dioxide in their mesophyll cells (using an enzyme

called phosphoenolpyruvate carboxylase which catalyzes the combination of
carbon dioxide with a compound called phosphoenolpyruvate (PEP)), forming
oxaloacetate. This oxaloacetate is then converted to malate and is transported
into the bundle sheath cells (site of carbon dioxide fixation by RUBISCO)
where oxygen concentration is low to avoid photorespiration.

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Maize uses the C4 pathway, minimizing photorespiration

This ability to avoid photorespiration makes these plants more hardy than other
plants in dry and hot environments.
CAM plants, such as cacti and succulent plants, also use the enzyme PEP
carboxylase to capture carbon dioxide, but only at night. Crassulacean acid
metabolism allows plants to conduct most of their gas exchange in the cooler
night-time air, sequestering carbon in 4-carbon sugars which can be released to
the photosynthesizing cells during the day.

Types Of Plants That Undergo Photorespiration

Photorespiration occurs in c3 plants.

WHY DOES PHOTORESPIRATION NOT OCCUR IN C4 PLANTS?
In c4 plants photorespiration does not occur because they have a mechanism that
increases the concentration of co2 at the enzyme site. This takes place when the
c4 acid from the mesophyll is broken down in the bundle sheath cells to release
co2. This results in increasing the intercellular concentration of co2.

Photorespiration is absent only in C4 plants (e.g. - sugarcane, maize,

sorghum, Atriplex, Amaranthus) .  They achieve this by

1. Carrying out photolysis and activity of RUBISCO enzyme at different

sites.
2. Effective pumping of CO2.

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 3. Thick and impermeable wall of bundle sheath cells for gaseous
exchange.

Photorespiration occurs in c3 plants.

Peanuts, coconuts, sugar beets, tobacco, spinach, soybeans, Wheat, tomato, oats,
rice, barley, mango, neem flower, bell pepper and most trees are c3 plants. most
lawn grasses such as rye and fescue are also c3 plants that undergo the process
photorespiration.

WHEN DOES PHOTORESPIRATION OCCUR?

Photorespiration usually occurs when there is high concentration of oxygen.

Under these circumstances, RUBISCO, the enzyme that catalyzes the
carboxylation of RuBP during the first step of Calvin cycle, functions such as an
oxygenase.
Or in other word photorespiration occurs when the co2 levels inside a leaf
become low.it occurs when rubisco( RuBP carboxylase) joins oxygen to RuBP in
the first step of carbon cycle rather that carbon dioxide.
Photorespiration generally occurs on hot, dry sunny days. When its too hot or
dry, the leaves run out of water and cannot continue the light reaction. Then the
plants close their stomata, inhibiting co2 uptake and increasing the o2
concentration in leaves.
Plants do not perform photorespiration at night. But they can if they are only
provided with artificial light of the corresponding wavelengths.

The C3 Pathway In plants

Plants which use only the Calvin cycle for fixing the carbon dioxide from the air
are known as C3 plants.C3 pathway is also known as Calvin cycle.
Calvin cycle occurs in three steps:
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 -carboxylation
-reduction
-regeneration

In the first step, the two molecules of 3-phosphoglyceric acid (PGA) are produced
with the help of the enzyme called RuBP carboxylase. Later in the second and
third steps, the ATP and NADPH phosphorylate the 3-PGA and ultimately
produces glucoses. Then the cycle restarts again by regeneration of RuBP.
About 85% of plant species are c3 plants. They include the cereal grains: wheat,
rice, barley, oats, peanuts, cotton etc...
The entire process, from light energy to sugar production occurs within the
chloroplast.

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C3 plants have the disadvantage that in hot dry conditions their photosynthetic
efficiency suffers because of photorespiration. When the c02 concentration in
chloroplasts drops below about 50 ppm, the catalyst rubisco that helps to fix
carbon begins to fix oxygen instead. This is highly wasteful .

The C4 Pathway in plants

Every photosynthetic plant follows Calvin cycle, but in some plants, there is a
primary stage to the Calvin Cycle known as C4 pathway. It is also known as hatch
and slack pathway. C4 carbon fixation or the Hatch–Slack pathway is one of
three known photosynthetic processes of carbon fixation in plants. It owes the
names to the 1960's discovery by Marshall Davidson Hatch and Charles Roger
Slack that some plants, when supplied with 14CO2, incorporate the 14C label
into four-carbon molecules first.
Plants in tropical desert regions commonly follow the C4 pathway. Here, a 4-
carbon compound called oxaloacetic acid (OAA) is the first product by carbon
fixation.
In c4 plants, the light-dependent reactions and the Calvin cycle are physically
separated, with the light-dependent reactions occurring in the mesophyll cells
(spongy tissue in the middle of the leaf) and the Calvin cycle occurring in special
cells around the leaf veins. These cells are called bundle-sheath cells.
C4 plants are highly efficient at:
-low co2 concentration
-high light intensity
-high temperature
Co2+PEP Oxaloacetate malate

Pyruvate

sugar

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C4 plants use a two stage process were co2 is fixed in thin walled mesophyll cells
to form a 4-carbon intermediate, typically malate (malic acid ). The reaction
involves phosphonyl pyruvate (PEP) which fixes co2 in a reaction catalyzed by PEP
carboxylate. It forms oxaloacetic acid (OAA) which is quickly converted to malic
acid. The 4-carbon acid is actively pumped across the cell membrane into a thick
walled sheath cell where it is split to co2 and a 3-carbon compound.
So the 4 phases in c4 are:
1. CO2 is fixed to produce a four-carbon molecule (malate or aspartate).
2. The molecule exits the cell and enters the bundle sheath cells.
3. It is then broken down into CO2 and pyruvate. CO2 enters the Calvin cycle
to produce carbohydrates.
4. Pyruvate reenters the mesophyll cell, where it is reused to produce
malate or aspartate.
C4 fixation is an addition to the ancestral and more common C3 carbon fixation.
The main carboxylating enzyme in C3 photosynthesis is called Rubisco, and
catalyzes two distinct reactions, with CO2 (carboxylation), and with oxygen
(oxygenation), which gives rise to the wasteful process of photorespiration. C4
photosynthesis reduces photorespiration by concentrating CO2 around Rubisco.
To ensure that Rubisco works in an environment where there is a lot of carbon
dioxide and very little oxygen, C4 leaves generally differentiate two partially
isolated compartments called mesophyll cells and bundle-sheath cells.
Corn, sugarcane and some shrubs are examples of plants that follow the C4
pathway. Calvin pathway is a common pathway in both C3 plants and C4 plants,
but it takes place only in the mesophyll cells of the C3 Plants but not in the C4
Plants.

The CAM Pathway in Plants

Crassulacean acid metabolism, also known as CAM is a carbon fixation pathway

that evolved in some plants as an adaption to arid conditions (drought).
The stomata in the leaves closed during the day to reduce evapotranspiration and
produce sugar, but open at night to collect carbon dioxide.

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The co2 is stored as the four-carbon acid malate, and then react with Rubisco
produce sugar through photosynthesis at day.
CAM plants fix carbon dioxide during the night, going to a similar process as c4
pathway until creates malate.
Malate is stored in a vacuole during day time.

Some plants that are adapted to dry environments, such as cacti and pineapples,
use the crassulacean acid metabolism (CAM) pathway to minimize
photorespiration. This name comes from the family of plants, the Crassulaceae, in
which scientists first discovered the pathway.

Instead of separating the light-dependent reactions and the use of co2 in the
Calvin cycle in space, CAM plants separate these processes in time. At night, CAM
plants open their stomata, allowing co2 to diffuse into the leaves. This co2 is fixed

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into oxaloacetate by PEP carboxylase, then converted to malate or another type
of organic acid.

The organic acid is stored inside vacuoles until the next day. In the daylight, the
CAM plants do not open their stomata, but they can still photosynthesize. That's
because the organic acids are transported out of the vacuole and broken down to
release co2, which enters the Calvin cycle. His controlled release maintains a high
concentration of co2 around rubisco.

plant species that use CAM photosynthesis not only avoid photorespiration, but
are also very water-efficient. Their stomata only open at night, when humidity
tends to be higher and temperatures are cooler, both factors that reduce water
loss from leaves. CAM plants are typically dominant in very hot, dry areas, like
deserts.

Some examples of CAM plants include pineapples and cactus.

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 Difference between C3, C4 & CAM pathways

C3 pathway C4 pathway CAM pathway

1. Produces, a three- Produces an four-carbon Gathers sunlight during
carbon compound via compound which split into the day and fix co2 at
the Calvin cycle. a three-carbon compound night.
for the Calvin cycle.
2. Occurs in all plants. Occurs in around 3% of Occurs in plants
vascular plants adapted to dry
environments
3 Occurs in mesophyll Occurs in mesophyll cells Occurs in mesophylls.
cells. and bundle sheath cells.
4 First stable product: 3- First stable product: First stable product: 3-
PGA oxaloacetate/malate PGA at daytime oaa at
night/malate
5. Dark reaction requires: Dark reaction requires: 12 Dark reaction requires:
12 NADPH and 18 ATP. NADPH and 30 ATP. 12 NADPH and 39 atps.
6 Optimal temperature: Optimal temperature:30-40 Optimal
15-25 temperature:>40
7. Stomata: open Stomata: open Stomata: close
8. Leaf anatomy: typical Leaf anatomy: kranz Leaf anatomy:
xeromorphic
9. Does not have special Minimize photorespiration Minimizes
features to combat by performing carbon photorespiration by
photorespiration. dioxide fixation and Calvin performing co2 fixation
cycle in separate cells. and Calvin cycle in
separate times.
10. Carboxylation enzyme: Carboxylation enzyme: PEP Carboxylation enzyme:
RuBP carboxylase carboxylase in mesophyll PEP carboxylase in the
and RuBP carboxylase in dark and RuBP
bundle sheath cells carboxylase in the day
time
11. Initial co2 acceptor: Initial co2 acceptor: PEP Initial co2 acceptor:PEP
RuBP
12. 89% of plant species <1% plant species 10% species of plants
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 worldwide worldwide worldwide
13. Only c3 type of cycle Both c3 and c4 type of cycle C3 during day
C4 during night
14. C02 compensation point 0-100 ppm 0-5 ppm
25-100 ppm
15. Co2: ATP: NADPH 1:3:2 1:5:2 1:6.5:2
16. Cool, wet environments Hot, sunny environments Very hot, dry
environments

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