Crassulacean acid metabolism
Crassulacean acid metabolism, also known as CAM photosynthesis, is a
Historical background
Observations relating to CAM were first made by
Overview: a two-part cycle
CAM is an adaptation for increased efficiency in the use of water, and so is typically found in plants growing in arid conditions.
During the night
During the night, a plant employing CAM has its stomata open, allowing CO2 to enter and be fixed as organic acids by a
During the day
During the day, the stomata close to conserve water, and the CO2-storing organic acids are released from the vacuoles of the mesophyll cells. An enzyme in the stroma of chloroplasts releases the CO2, which enters into the Calvin cycle so that photosynthesis may take place.[citation needed]
Benefits
The most important benefit of CAM to the plant is the ability to leave most leaf stomata closed during the day.[8] Plants employing CAM are most common in arid environments, where water is scarce. Being able to keep stomata closed during the hottest and driest part of the day reduces the loss of water through evapotranspiration, allowing such plants to grow in environments that would otherwise be far too dry. Plants using only C3 carbon fixation, for example, lose 97% of the water they take up through the roots to transpiration - a high cost avoided by plants able to employ CAM.[9][What percentage is lost in CAM plants?]
Comparison with C4 metabolism
The
There are some C4/CAM intermediate species, such as Peperomia camptotricha, Portulaca oleracea, and Portulaca grandiflora. It was previously thought that the two pathways of photosynthesis in such plants could occur in the same leaves but not in the same cells, and that the two pathways could not couple but only occur side by side.[10] It is now known, however, that in at least some species such as Portulaca oleracea, C4 and CAM photosynthesis are fully integrated within the same cells, and that CAM-generated metabolites are incorporated directly into the C4 cycle.[11]
Biochemistry
Plants with CAM must control storage of CO2 and its reduction to branched carbohydrates in space and time.
At low temperatures (frequently at night), plants using CAM open their
In daylight, plants using CAM close their guard cells and discharge malate that is subsequently transported into chloroplasts. There, depending on plant species, it is cleaved into
. During the following cool night, PEP is finally exported into the cytoplasm, where it is involved in fixing carbon dioxide via malate.Use by plants
Plants use CAM to different degrees. Some are "obligate CAM plants", i.e. they use only CAM in photosynthesis, although they vary in the amount of CO2 they are able to store as organic acids; they are sometimes divided into "strong CAM" and "weak CAM" plants on this basis. Other plants show "inducible CAM", in which they are able to switch between using either the C3 or C4 mechanism and CAM depending on environmental conditions. Another group of plants employ "CAM-cycling", in which their stomata do not open at night; the plants instead recycle CO2 produced by respiration as well as storing some CO2 during the day.[5]
Plants showing inducible CAM and CAM-cycling are typically found in conditions where periods of water shortage alternate with periods when water is freely available. Periodic drought – a feature of semi-arid regions – is one cause of water shortage. Plants which grow on trees or rocks (as epiphytes or lithophytes) also experience variations in water availability. Salinity, high light levels and nutrient availability are other factors which have been shown to induce CAM.[5]
Since CAM is an adaptation to arid conditions, plants using CAM often display other
Aquatic CAM
CAM photosynthesis is also found in aquatic species in at least 4 genera, including: Isoetes, Crassula, Littorella, Sagittaria, and possibly Vallisneria,[14] being found in a variety of species e.g. Isoetes howellii, Crassula aquatica.
These plants follow the same nocturnal acid accumulation and daytime deacidification as terrestrial CAM species.[15] However, the reason for CAM in aquatic plants is not due to a lack of available water, but a limited supply of CO2.[14] CO2 is limited due to slow diffusion in water, 10000x slower than in air. The problem is especially acute under acid pH, where the only inorganic carbon species present is CO2, with no available bicarbonate or carbonate supply.
Aquatic CAM plants capture carbon at night when it is abundant due to a lack of competition from other photosynthetic organisms.[15] This also results in lowered photorespiration due to less photosynthetically generated oxygen.
Aquatic CAM is most marked in the summer months when there is increased competition for CO2, compared to the winter months. However, in the winter months CAM still has a significant role.[16]
Ecological and taxonomic distribution of CAM-using plants
The majority of plants possessing CAM are either
The only trees that can do CAM are in the genus Clusia;[18] species of which are found across Central America, South America and the Caribbean. In Clusia, CAM is found in species that inhabit hotter, drier ecological niches, whereas species living in cooler montane forests tend to be C3.[19] In addition, some species of Clusia can temporarily switch their photosynthetic physiology from C3 to CAM, a process known as facultative CAM. This allows these trees to benefit from the elevated growth rates of C3 photosynthesis, when water is plentiful, and the drought tolerant nature of CAM, when the dry season occurs.
Plants which are able to switch between different methods of carbon fixation include Portulacaria afra, better known as Dwarf Jade Plant, which normally uses C3 fixation but can use CAM if it is drought-stressed,[20] and Portulaca oleracea, better known as Purslane, which normally uses C4 fixation but is also able to switch to CAM when drought-stressed.[21]
CAM has
The following list summarizes the taxonomic distribution of plants with CAM:
Division | Class/Angiosperm group | Order | Family | Plant Type | Clade involved |
---|---|---|---|---|---|
Lycopodiophyta
|
Isoetopsida
|
Isoetales | Isoetaceae | hydrophyte | Isoetes[25] (the sole genus of class Isoetopsida) - I. howellii (seasonally submerged), I. macrospora, I. bolanderi, I. engelmannii, I. lacustris, I. sinensis, I. storkii, I. kirkii, I. taiwanensis. |
Pteridophyta
|
Polypodiopsida
|
Polypodiales | Polypodiaceae | epiphyte, lithophyte | CAM is recorded from Microsorum, Platycerium and Polypodium,[26] Pyrrosia and Drymoglossum[27] and Microgramma |
Pteridopsida
|
Polypodiales | Pteridaceae[28] | epiphyte | Vittaria[29] | |
Cycadophyta
|
Cycadopsida
|
Cycadales
|
Zamiaceae | Dioon edule[31] | |
Gnetophyta | Gnetopsida
|
Welwitschiales
|
Welwitschiaceae | xerophyte | Welwitschiales )
|
Magnoliophyta
|
magnoliids | Magnoliales | Piperaceae | epiphyte | Peperomia camptotricha[33] |
eudicots
|
Caryophyllales | Aizoaceae | xerophyte | widespread in the family; Mesembryanthemum crystallinum is a rare instance of an halophyte that displays CAM[34] | |
Cactaceae
|
xerophyte | Almost all cacti have obligate Crassulacean Acid Metabolism in their stems; the few cacti with leaves may have C3 Metabolism in those leaves;[35] seedlings have C3 Metabolism.[36] | |||
Portulacaceae | xerophyte | recorded in approximately half of the genera (note: Portulacaceae is paraphyletic with respect to Cactaceae and Didiereaceae)[37] | |||
Didiereaceae | xerophyte | ||||
Saxifragales | Crassulaceae | hydrophyte, xerophyte, lithophyte | Crassulacean acid metabolism is widespread among the (eponymous) Crassulaceae. | ||
eudicots (rosids) | Vitales
|
Vitaceae[38] | Cissus,[39] Cyphostemma | ||
Malpighiales | Clusiaceae | hemiepiphyte | Clusia[39][40] | ||
Euphorbiaceae[38] | CAM is found is some species of Synadenium . C4 photosynthesis is also found in Euphorbia (subgenus Chamaesyce).
| ||||
Passifloraceae[28] | xerophyte | Adenia[42] | |||
Geraniales | Geraniaceae | CAM is found in some succulent species of Pelargonium,[43] and is also reported from Geranium pratense[44] | |||
Cucurbitales | Cucurbitaceae | Xerosicyos danguyi,[45] Dendrosicyos socotrana,[46] Momordica[47] | |||
Celastrales | Celastraceae[48] | ||||
Oxalidales | Oxalidaceae[49] | Oxalis carnosa var. hirta[49] | |||
Brassicales | Moringaceae
|
Moringa[50] | |||
Salvadoraceae[49] | CAM is found in Salvadora persica.[49] Salvadoraceae were previously placed in order Celastrales, but are now placed in Brassicales. | ||||
Sapindales | Sapindaceae | Dodonaea viscosa | |||
Fabales | Fabaceae[49] | CAM is found in Prosopis juliflora (listed under the family Salvadoraceae in Sayed's (2001) table,[49]) but is currently in the family Fabaceae (Leguminosae) according to The Plant List[51]). | |||
Zygophyllaceae | Zygophyllum[50] | ||||
eudicots (asterids) | Ericales | Ebenaceae | |||
Solanales | Convolvulaceae | Ipomoea[citation needed] (Some species of Ipomoea are C3[39][52] - a citation is needed here.) | |||
Gentianales | Rubiaceae | epiphyte | Hydnophytum and Myrmecodia | ||
Apocynaceae | CAM is found in subfamily Asclepidioideae ( | ||||
Lamiales | Gesneriaceae | epiphyte | CAM was found Codonanthe crassifolia, but not in 3 other genera[56] | ||
Lamiaceae | Plectranthus marrubioides, Coleus[citation needed] | ||||
Plantaginaceae | hydrophyte | Littorella uniflora[25] | |||
Apiales | Apiaceae | hydrophyte | Lilaeopsis lacustris | ||
Asterales | Asteraceae[38] | some species of Senecio[57] | |||
monocots | Alismatales | Hydrocharitaceae | hydrophyte | Hydrilla,[38] Vallisneria | |
Alismataceae | hydrophyte | Sagittaria | |||
Araceae | Zamioculcas zamiifolia is the only CAM plant in Araceae, and the only non-aquatic CAM plant in Alismatales[58] | ||||
Poales | Bromeliaceae | epiphyte | |||
Cyperaceae | hydrophyte | Scirpus,[38] Eleocharis | |||
Asparagales | Orchidaceae
|
epiphyte | Orchidaceae has more CAM species than any other family (CAM Orchids) | ||
Agavaceae[40]
|
xerophyte | Agave,[39] Hesperaloe, Yucca and Polianthes[42] | |||
Asphodelaceae[38] | xerophyte | Aloe,[39] Gasteria,[39] and Haworthia | |||
Ruscaceae[38]
|
Sansevieria[39][49] (This genus is listed under the family Dracaenaceae in Sayed's (2001) table, but currently in the family Asparagaceae according to The Plant List), Dracaena[60] | ||||
Commelinales | Commelinaceae | Callisia,[39] Tradescantia, Tripogandra |
See also
- C2 photosynthesis
- C3 carbon fixation
- C4 carbon fixation
- RuBisCO
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