Carbon fixation

Cyanobacteria such as these carry out photosynthesis. Their emergence foreshadowed the evolution of many photosynthetic plants, changing the earth's atmosphere. Carbon fixation is the conversion of inorganic carbon (carbon dioxide) to organic compounds by living organisms. The most prominent example is photosynthesis, although chemosynthesis is another form of carbon fixation that can take place in the absence of sunlight. Organisms that grow by fixing carbon are called autotrophs. Autotrophs include photoautotrophs, which synthesize organic compounds using the energy of sunlight, and lithoautotrophs, which synthesize organic compounds using the energy of inorganic oxidation. Heterotrophs are organisms that grow using the carbon fixed by autotrophs. The organic compounds are used by heterotrophs to produce energy and to build body structures. "Fixed carbon", "reduced carbon", and "organic carbon" are equivalent terms for various organic compounds.[1]

Net vs gross CO2 fixation

Graphic showing net annual amounts of CO2 fixation by land and sea-based organisms. It is estimated that approximately 258 billion tons of carbon dioxide are converted by photosynthesis annually. The majority of the fixation occurs in marine environments, especially areas of high nutrients. The gross amount of carbon dioxide fixed is much larger since approximately 40% is consumed by respiration in the evenings following each day of photosynthesis.[1] Given the scale of this process, it is understandable that RuBisCO is the most abundant protein on earth.

Overview of pathways
Six autotrophic carbon fixation pathways are known as of 2011. The Calvin cycle fixes carbon in the chloroplasts of plants and algae, and in the cyanobacteria. It also fixes carbon in the anoxygenic photosynthetic proteobacteria called purple bacteria, and in some non-phototrophic proteobacteria.[2]

Oxygenic photosynthesis
In photosynthesis, energy from sunlight drives the carbon fixation pathway. Oxygenic photosynthesis is used by the primary producersplants, algae, and cyanobacteria. They contain the pigment chlorophyll, and use the Calvin cycle to fix carbon autotrophically. The process works like this:

2H2O 4e- + 4H+ + O2 CO2 + 4e- + 4H+ CH2O + H2O In the first step, water is dissociated into electrons, protons, and free oxygen. This allows the use of water, one of the most abundant substances on Earth, as an electron donoras a source of reducing power. The release of free oxygen is a side-effect of enormous consequence. The first step uses the energy of sunlight to oxidize water to O2, and, ultimately, to produce ATP ADP + Pi ATP + H2O

and the reductant, NADPH NADP+ + 2e- + 2H+ NADPH + H+

The second step, called the Calvin cycle, the actual fixation of carbon dioxide is carried out. This process consumes ATP and NADPH. The Calvin cycle in plants accounts for the preponderance of carbon fixation on land. In algae and cyanobacteria, it accounts for the preponderance of carbon fixation in the oceans. The Calvin cycle converts carbon dioxide into sugar, as triose phosphate (TP), which is glyceraldehyde 3-phosphate (GAP) together with dihydroxyacetone phosphate (DHAP): 3 CO2 + 12 e- + 12 H+ + Pi TP + 4 H2O An alternative perspective accounts for NADPH (source of e-) and ATP: 3 CO2 + 6 NADPH + 6 H+ + 9 ATP + 5 H2O TP + 6 NADP+ + 9 ADP + 8 Pi The formula for inorganic phosphate (Pi) is HOPO32- + 2H+. Formulas for triose and TP are C2H3O2-CH2OH and C2H3O2-CH2OPO32- + 2H+

Evolutionary considerations
Somewhere between 3.5 and 2.3 billion years ago, cyanobacteria evolved oxygenic photosynthesis.[3][4][5]

Carbon concentrating mechanisms

Many photosynthetic organisms have acquired inorganic carbon concentrating mechanisms (CCM), which increase the concentration of carbon dioxide available to the initial carboxylase of the Calvin cycle, the enzyme RuBisCO. The benefits of CCM include increased tolerance to low external concentrations of inorganic carbon, and reduced loses to photorespiration. CCM can make plants more tolerant of heat and water stress. Carbon concentrating mechanisms use the enzyme carbonic anhydrase (CA), which catalyze both the dehydration of bicarbonate to carbon dioxide and the hydration of carbon dioxide to bicarbonate

HCO3- + H+

CO2 + H2O

Lipid membranes are much less permeable to bicarbonate than to carbon dioxide. To capture inorganic carbon more effectively, some plants have adapted the anaplerotic reaction HCO3- + H+ + PEP OAA + Pi catalyzed by PEP carboxylase (PEPC), to carboxylate phosphoenolpyruvate (PEP) to oxaloacetate (OAA) which is a C4 dicarboxylic acid. CAM plants CAM plants that use Crassulacean acid metabolism as an adaptation for arid conditions. CO2 enters through the stomata during the night and is converted into the 4-carbon compound, malic acid, which releases CO2 for use in the Calvin cycle during the day, when the stomata are closed. The jade plant (Crassula ovata) and cacti are typical of CAM plants. Sixteen thousand species of plants use CAM.[6] These plants have a carbon isotope signature of -20 to -10 .[7] C4 plants C4 plants preface the Calvin cycle with reactions that incorporate CO2 into one of the 4-carbon compounds, malic acid or aspartic acid. C4 plants have a distinctive internal leaf anatomy. Tropical grasses, such as sugar cane and maize are C4 plants, but there are many broadleaf plants that are C4. Overall, 7600 species of terrestrial plants use C4 carbon fixation, representing around 3% of all species.[8] These plants have a carbon isotope signature of -16 to -10 .[7] C3 plants The large majority of plants are C3 plants. They are so-called to distinguish them from the CAM and C4 plants, and because the carboxylation products of the Calvin cycle are 3-carbon compounds. They lack C4 dicarboxylic acid cycles, and therefore have higher carbon dioxide compensation points than CAM or C4 plants. C3 plants have a carbon isotope signature of -24 to 33.[