section_path: Light-independent reactions/Carbon concentrating mechanisms/On land
section_name: On land
paragraph_text: On land
  
  In hot and dry conditions, plants close their stomata to prevent water loss. Under these conditions, CO will decrease and oxygen gas, produced by the light reactions of photosynthesis, will increase, causing an increase of photorespiration by the oxygenase activity of ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBisCO) and decrease in carbon fixation. Some plants have evolved mechanisms to increase the CO concentration in the leaves under these conditions.
  
  Plants that use the C carbon fixation process chemically fix carbon dioxide in the cells of the mesophyll by adding it to the three-carbon molecule phosphoenolpyruvate (PEP), a reaction catalyzed by an enzyme called PEP carboxylase, creating the four-carbon organic acid oxaloacetic acid. Oxaloacetic acid or malate synthesized by this process is then translocated to specialized bundle sheath cells where the enzyme RuBisCO and other Calvin cycle enzymes are located, and where CO released by decarboxylation of the four-carbon acids is then fixed by RuBisCO activity to the three-carbon 3-phosphoglyceric acids. The physical separation of RuBisCO from the oxygen-generating light reactions reduces photorespiration and increases CO fixation and, thus, the photosynthetic capacity of the leaf. C plants can produce more sugar than C plants in conditions of high light and temperature. Many important crop plants are C plants, including maize, sorghum, sugarcane, and millet. Plants that do not use PEP-carboxylase in carbon fixation are called C plants because the primary carboxylation reaction, catalyzed by RuBisCO, produces the three-carbon 3-phosphoglyceric acids directly in the Calvin-Benson cycle. Over 90% of plants use C carbon fixation, compared to 3% that use C carbon fixation; however, the evolution of C in over sixty plant lineages makes it a striking example of convergent evolution. C photosynthesis, which involves carbon-concentration by selective breakdown of photorespiratory glycine, is both an evolutionary precursor to C and a useful carbon-concentrating mechanism in its own right.
  
  Xerophytes, such as cacti and most succulents, also use PEP carboxylase to capture carbon dioxide in a process called Crassulacean acid metabolism (CAM). In contrast to C metabolism, which spatially separates the CO fixation to PEP from the Calvin cycle, CAM temporally separates these two processes. CAM plants have a different leaf anatomy from C plants, and fix the CO at night, when their stomata are open. CAM plants store the CO mostly in the form of malic acid via carboxylation of phosphoenolpyruvate to oxaloacetate, which is then reduced to malate. Decarboxylation of malate during the day releases CO inside the leaves, thus allowing carbon fixation to 3-phosphoglycerate by RuBisCO. CAM is used by 16,000 species of plants.
  
  Calcium-oxalate -accumulating plants, such as Amaranthus hybridus and Colobanthus quitensis, show a variation of photosynthesis where calcium oxalate crystals function as dynamic carbon pools, supplying carbon dioxide (CO) to photosynthetic cells when stomata are partially or totally closed. This process was named alarm photosynthesis. Under stress conditions (e.g., water deficit), oxalate released from calcium oxalate crystals is converted to CO by an oxalate oxidase enzyme, and the produced CO can support the Calvin cycle reactions. Reactive hydrogen peroxide (H O), the byproduct of oxalate oxidase reaction, can be neutralized by catalase. Alarm photosynthesis represents a photosynthetic variant to be added to the well-known C4 and CAM pathways. However, alarm photosynthesis, in contrast to these pathways, operates as a biochemical pump that collects carbon from the organ interior (or from the soil) and not from the atmosphere.