photorespiration.
These pathways work by ensuring that Rubisco always
C3, C4, and CAM plants encounters high concentrations of , making it unlikely to bind to .
Google Classroom Microsoft Teams In the rest of this article, we'll take a closer look at the and CAM pathways
and see how they reduce photorespiration.
How the C4 and CAM pathways help minimize photorespiration.
plants
Key points: A "normal" plant—one that doesn't have photosynthetic adaptations to reduce
photorespiration—is called a plant. The first step of the Calvin cycle is the
Photorespiration is a wasteful pathway that occurs when the Calvin
fixation of carbon dioxide by rubisco, and plants that use only this "standard"
cycle enzyme rubisco acts on oxygen rather than carbon dioxide.
mechanism of carbon fixation are called plants, for the three-carbon
compound (3-PGA) the reaction produces. About of the plant species on the
The majority of plants are plants, which have no special features to
planet are plants, including rice, wheat, soybeans and all trees.
combat photorespiration.
plants minimize photorespiration by separating initial fixation and the
Calvin cycle in space, performing these steps in different cell types.
Crassulacean acid metabolism (CAM) plants minimize photorespiration
and save water by separating these steps in time, between night and day.
Introduction
High crop yields are pretty important—for keeping people fed, and also for
keeping economies running. If you heard there was a single factor that
reduced the yield of wheat by and the yield of soybeans by in the United
States, for instance, you might be curious to know what it was.
plants
As it turns out, the factor behind those (real-life) numbers is photorespiration.
In plants, the light-dependent reactions and the Calvin cycle are physically
This wasteful metabolic pathway begins when rubisco, the carbon-fixing
separated, with the light-dependent reactions occurring in the mesophyll cells
enzyme of the Calvin cycle, grabs rather than . It uses up fixed carbon,
(spongy tissue in the middle of the leaf) and the Calvin cycle occurring in
wastes energy, and tends to happens when plants close their stomata (leaf
special cells around the leaf veins. These cells are called bundle-
pores) to reduce water loss. High temperatures make it even worse.
sheath cells.
Some plants, unlike wheat and soybean, can escape the worst effects of
To see how this division helps, let's look at an example of photosynthesis in
photorespiration. The and CAM pathways are two adaptations—beneficial
action. First, atmospheric is fixed in the mesophyll cells to form a simple, -
features arising by natural selection—that allow certain species to minimize
carbon organic acid (oxaloacetate). This step is carried out by a non-rubisco
enzyme, PEP carboxylase, that has no tendency to bind . Oxaloacetate is CAM plants
then converted to a similar molecule, malate, that can be transported in to the
Some plants that are adapted to dry environments, such as cacti and
bundle-sheath cells. Inside the bundle sheath, malate breaks down, releasing
pineapples, use the crassulacean acid metabolism (CAM) pathway to
a molecule of . The is then fixed by rubisco and made into sugars via the
minimize photorespiration. This name comes from the family of plants, the
Calvin cycle, exactly as in photosynthesis.
Crassulaceae, in which scientists first discovered the pathway.
Image credit: "Crassulaceae," by Guyon Morée (CC BY 2.0).
This process isn't without its energetic price: ATP must be expended to return
the three-carbon “ferry” molecule from the bundle sheath cell and get it ready Instead of separating the light-dependent reactions and the use of in the
to pick up another molecule of atmospheric . However, because the Calvin cycle in space, CAM plants separate these processes in time. At night,
mesophyll cells constantly pump into neighboring bundle-sheath cells in the CAM plants open their stomata, allowing to diffuse into the leaves. This is
form of malate, there’s always a high concentration of relative to right fixed into oxaloacetate by PEP carboxylase (the same step used by plants),
around rubisco. This strategy minimizes photorespiration. then converted to malate or another type of organic acid.
The pathway is used in about of all vascular plants; some examples are The organic acid is stored inside vacuoles until the next day. In the daylight,
crabgrass, sugarcane and corn. plants are common in habitats that are hot, the CAM plants do not open their stomata, but they can still photosynthesize.
but are less abundant in areas that are cooler. In hot conditions, the benefits That's because the organic acids are transported out of the vacuole and
of reduced photorespiration likely exceed the ATP cost of moving from the broken down to release , which enters the Calvin cycle. This controlled
mesophyll cell to the bundle-sheath cell. release maintains a high concentration of around rubisco.
Type Separation of initial fixation Stomata Best adapted to
and Calvin cycle open
No separation Day Cool, wet
environments
Between mesophyll and bundle- Day Hot, sunny
sheath cells (in space) environments
CAM Between night and day (in time) Night Very hot, dry
environments
The CAM pathway requires ATP at multiple steps (not shown above), so like
photosynthesis, it is not an energetic "freebie." However, 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.
Comparisons of , , and CAM plants
, and CAM plants all use the Calvin cycle to make sugars from . These
pathways for fixing have different advantages and disadvantages and make
plants suited for different habitats. The mechanism works well in cool
environments, while and CAM plants are adapted to hot, dry areas.
Both the and CAM pathways have evolved independently over two dozen
times, which suggests they may give plant species in hot climates a
significant evolutionary advantage.
