This subproject is one of many research subprojects utilizing the resources provided by a Center grant funded by NIH/NCRR. The subproject and investigator (PI) may have received primary funding from another NIH source, and thus could be represented in other CRISP entries. The institution listed is for the Center, which is not necessarily the institution for the investigator. The initial source of Earth s organic carbon is atmospheric or dissolved CO2, which is fixed by enzymes found in green plants, algae and autotrophic bacteria. The carbon is then spread through the food chain to all other living organisms. The most important enzyme responsible for fixing carbon is ribulose bisphosphate carboxylase/oxygenase (RuBisCO), which catalyzes the first step in carbon fixation via the Calvin-Benson-Bassham cycle - the covalent attachment of CO2 to ribulose-1,5-bisphosphate and its subsequent cleavage into two molecules of 3-phosphoglycerate. Based on the immense biomass of photosynthetic and chemoautotrophic organisms on Earth, RuBisCO is estimated to be the most abundant protein known. All cyanobacteria and some chemoautotrophic bacteria enhance their CO2 fixation by sequestering RuBisCO into polyhedral bodies called carboxysomes. Halothiobacillus neapolitanus carboxysomes are delimited by a proteinaceous shell and are filled with RuBisCO. In previous electron microscopic studies, carboxysomes appeared hexagonal with a granular interior, a diameter of approximately 120 nm and a shell thickness of between 3 to 4 nm 8; 9. Although the enhancement of carbon dioxide fixation by the carboxysome has been firmly established, the exact mechanism has not yet been elucidated. Although much has been published on the occurrence, physiology, biochemistry and genetics of these microcompartments/organelles, only two major reports have analyzed the structure or symmetry of purified carboxysomes. Peters concluded that the carboxysomes of Nitrobacter agilis obey icosahedral symmetry, based on negative stain projection images and heavy-metal shadowing of critical-point-dried carboxysomes Holthuijzen and co-workers, on the other hand, reported the shape of carboxysomes of H. neapolitanus to be dodecahedral. We propose to use cryoEM to determine the molecular structure of ca. rboxysomes of H. neapolitanus
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