Fractionation of carbon isotopes in oxygenic photosynthesis be less fractionated in C (ie, δC is relatively less negative) compared to C3 plants with low water-use efficiency. C4 plants have developed the C4 carbon fixation pathway to conserve water loss, thus are more prevalent in hot, sunny, and dry climates. These plants differ from C3 plants because the Calvin Cycle in C4 plants produces a four-carbon molecule, malate (rather than 3-phosphoglycerate), after fixation of CO. Malate, the four-carbon compound is the namesake of "C4" photosynthesis. This pathway allows C4 photosynthesis to efficiently shuttle CO to the RuBisCO enzyme and maintain high concentrations of CO within bundle sheath cells.
Evolution of photosynthesis as an electron donor. Green nonsulfur bacteria used various amino and other organic acids. Purple nonsulfur bacteria used a variety of nonspecific organic molecules. Fossils of what are thought to be filamentous photosynthetic organisms have been dated at 3.4 billion years old. More recent studies, reported in March 2018, also suggest that photosynthesis may have begun about 3.4 billion years ago. The main source of oxygen in the atmosphere is oxygenic photosynthesis, and its first appearance is sometimes referred to as the oxygen catastrophe. Geological evidence suggests that oxygenic photosynthesis, such as that in cyanobacteria, became important during the Paleoproterozoic