The many types of carbonic anhydrases in photosynthetic organisms

Carbonic anhydrases (CAs) are enzymes that catalyze the interconversion of CO₂ and HCO₃^-. In nature, there are multiple families of CA, designated with the Greek letters α through θ. CAs are ubiquitous in plants, algae and photosynthetic bacteria, often playing essential roles in the CO₂ concentrating mechanisms (CCMs) which enhance the delivery of CO₂ to Rubisco. As algal CCMs become better characterized, it is clear that different types of CAs are playing the same role in different algae. For example, an α-CA catalyzes the conversion of accumulated HCO₃^- to CO₂ in the green alga Chlamydomonas reinhardtii, while a θ-CA performs the same function in the diatom Phaeodactylum tricornutum. In this review we argue that, in addition to its role of delivering CO₂ for photosynthesis, other metabolic roles of CA have likely changed as the Earth's atmospheric CO₂ level decreased. Since the algal and plant lineages diverged well before the decrease in atmospheric CO₂, it is likely that plant, algae and photosynthetic bacteria all adapted independently to the drop in atmospheric CO₂. In light of this, we will discuss how the roles of CAs may have changed over time, focusing on the role of CA in pH regulation, how CAs affect CO₂ supply for photosynthesis and how CAs may help in the delivery of HCO₃^- for other metabolic reactions.