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in concentrations far exceeding that of the surrounding medium. Badger and Price first suggested the function of the pyrenoid to be analogous to that of the carboxysome in cyanobacteria, in being associated with CCM activity. CCM activity in algal and cyanobacterial photobionts of lichen associations was also identified using gas exchange and carbon isotope discrimination and associated with the pyrenoid by Palmqvist and Badger et al. The Hornwort CCM was later characterized by Smith and Griffiths.

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