All plants carry on photosynthesis by
As its name suggests, RUBISCO catalyzes two different reactions:
- adding carbon dioxide (CO2) to a phosphorylated 5-carbon sugar called ribulose bisphosphate.
- This reaction is catalyzed by the enzyme ribulose bisphosphate carboxylase oxygenase (RUBISCO).
- The resulting 6-carbon compound breaks down into two molecules of 3-phosphoglyceric acid (PGA).
- These 3-carbon molecules serve as the starting material for the synthesis of glucose and other food molecules.
- The process is called the Calvin cycle and the pathway is called the C3 pathway.
Which one predominates depends on the relative concentrations of O2 and CO2 with
- adding CO2 to ribulose bisphosphate — the carboxylase activity
- adding O2 to ribulose bisphosphate — the oxygenase activity.
The light reactions of photosynthesis liberate oxygen and more oxygen dissolves in the cytosol of the cell at higher temperatures. Therefore,
- high CO2, low O2 favoring the carboxylase action,
- high O2, low CO2 favoring the oxygenase action.
favor the second reaction.
- high light intensities and
- high temperatures (above ~ 30°C)
So this process uses O2 and liberates CO2 as cellular respiration does which is why it is called photorespiration.
- The uptake of O2 by RUBISCO forms:
- the 3-carbon molecule 3-phosphoglyceric acid — just as in the Calvin cycle
- the 2-carbon molecule glycolate.
- The glycolate enters peroxisomes where it uses O2 to form intermediates that
- enter mitochondria where they are broken down to CO2.
It undoes the good anabolic work of photosynthesis, reducing the net productivity of the plant.
For this reason, much effort — so far largely unsuccessful — has gone into attempts to alter crop plants so that they carry on less photorespiration.
The problem may solve itself. If atmospheric CO2 concentrations continue to rise, perhaps this will enhance the net productivity of the world's crops by reducing losses to photorespiration.
Over 8000 species of angiosperms have developed adaptations which minimize the losses to photorespiration.
They all use a supplementary method of CO2 uptake which forms a 4-carbon molecule instead of the two 3-carbon molecules of the Calvin cycle. Hence these plants are called C4 plants. (Plants that have only the Calvin cycle are thus C3 plants.)
- Some C4 plants — called CAM plants — separate their C3 and C4 cycles by time. CAM plants are discussed below.
- Other C4 plants have structural changes in their leaf anatomy so that
These adaptations are described now.
- their C4 and C3 pathways are separated in different parts of the leaf with
- RUBISCO sequestered where the CO2 level is high; the O2 level low.
These C4 plants are well adapted to (and likely to be found in) habitats with
- After entering through stomata, CO2 diffuses into a mesophyll cell.
- Being close to the leaf surface, these cells are exposed to high levels of O2, but
- have no RUBISCO so cannot start photorespiration (nor the dark reactions of the Calvin cycle).
- Instead the CO2 is inserted into a 3-carbon compound (C3) called phosphoenolpyruvic acid (PEP) forming
- the 4-carbon compound oxaloacetic acid (C4).
- Oxaloacetic acid is converted into malic acid or aspartic acid (both have 4 carbons), which is
- transported (by plasmodesmata) into a bundle sheath cell. Bundle sheath cells
- are deep in the leaf so atmospheric oxygen cannot diffuse easily to them;
- often have thylakoids with reduced photosystem II complexes (the one that produces O2).
- Both of these features keep oxygen levels low.
- Here the 4-carbon compound is broken down into
- carbon dioxide, which enters the Calvin cycle to form sugars and starch.
- pyruvic acid (C3), which is transported back to a mesophyll cell where it is converted back into PEP.
- high daytime temperatures
- intense sunlight.
- corn (maize)
Although only ~3% of the angiosperms, C4 plants are responsible for ~25% of all the photosynthesis on land.
C4 cells in C3 plants
The ability to use the C4 pathway has evolved repeatedly in different families of angiosperms — a remarkable example of convergent evolution. Perhaps the potential is in all angiosperms.
A report in the 24 January 2002 issue of Nature (by Julian M. Hibbard and W. Paul Quick) describes the discovery that tobacco, a C3 plant, has cells capable of fixing carbon dioxide by the C4 path. These cells are clustered around the veins (containing xylem and phloem) of the stems and also in the petioles of the leaves. In this location, they are far removed from the stomata that could provide atmospheric CO2. Instead, they get their CO2 and/or the 4-carbon malic acid in the sap that has been brought up in the xylem from the roots.
If this turns out to be true of many C3 plants, it would explain why it has been so easy for C4 plants to evolve from C3 ancestors.
These are also C4 plants but instead of segregating the C4 and C3 pathways in different parts of the leaf, they separate them in time instead.
(CAM stands for crassulacean acid metabolism because it was first studied in members of the plant family Crassulaceae.)
In the morning,
- CAM plants take in CO2 through their open stomata (they tend to have reduced numbers of them).
- The CO2 joins with PEP to form the 4-carbon oxaloacetic acid.
- This is converted to 4-carbon malic acid that accumulates during the night in the central vacuole of the cells.
These adaptations also enable their owners to thrive in conditions of
- the stomata close (thus conserving moisture as well as reducing the inward diffusion of oxygen).
- The accumulated malic acid leaves the vacuole and is broken down to release CO2.
- The CO2 is taken up into the Calvin (C3) cycle.
Some examples of CAM plants:
On 26 October 2000, Nature reported the discovery of both the C3 and C4 pathways in a marine diatom. In this unicellular organism, the two paths are kept separate by having the C4 path in the cytosol, and the C3 path confined to the chloroplast. The presence of a C4 pathway probably reflects the frequent low concentrations of CO2 in ocean waters.
- high daytime temperatures
- intense sunlight
- low soil moisture.
29 December 2010