Carboxysomes are bacterial organelles consisting of polyhedral protein shells filled with the enzyme
Ribulose-1,5-biphosphate carboxylase/oxygenase (
RuBisCO) -the predominant enzyme in
carbon fixation and the
rate limiting enzyme in the
Calvin Cycle-and a
carbonic anhydrase. Carboxysomes are thought to have evolved as a consequence of the increase in oxygen concentration in the ancient atmosphere; this is because oxygen is a competing substrate to carbon dioxide in the RuBisCO reaction. To overcome the inefficiency of RuBisCO, carboxysomes concentrate carbon dioxide inside the shell by means of co-localized carbonic anhydrase activity, which produces carbon dioxide from the bicarbonate that diffuses into the carboxysome. The resulting production of carbon dioxide near RuBisCO decreases the proportion of ribulose-1,5-bisphosphate oxygenation and thereby avoids costly
photorespiratory reactions. The surrounding shell provides a barrier to carbon dioxide loss, helping to increase its concentration around RuBisCO. The carboxysome is an essential part of the
carbon-concentrating mechanism (CCM).