Algae have adopted two primary strategies to maximize the performance of Rubisco in photosynthetic CO2 fixation. This has included either the development of a CO2-concentrating mechanism (CCM), based at the level of the chloroplast, or the evolution of the kinetic properties of Rubisco. This review examines the potential diversity of both Rubisco and chloroplast-based CCMs across algal divisions, including both green and nongreen algae, and seeks to highlight recent advances in our understanding of the area and future areas for research. Overall, the available data show that Rubisco enzymes from algae have evolved a higher affinity for CO2 when the algae have adopted a strategy for CO2 fixation that does not utilise a CCM. This appears to be true of both Green and Red Form I Rubisco enzymes found in green and nongreen algae, respectively. However, the Red Form I Rubisco enzymes present in nongreen algae appear to have reduced oxygenase potential at air level of O2. This has resulted in a photosynthetic physiology with a reduced potential to be inhibited by O2 and a reduced need to deal with photorespiration. In the limited number of microalgae that have been examined, there is a strong correlation between the existence of a high-affinity CCM physiology and the presence of pyrenoids in all algae, highlighting the potential importance of these chloroplast Rubisco-containing bodies. However, in macroalgae, there is greater diversity in the apparent relationships between pyrenoids and chloroplast features and the CCM physiology that the species shows. There are many examples of microalgae and macroalgae with variations in the presence and absence of pyrenoids as well as single and multiple chloroplasts per cell. This occurs in both green and nongreen algae and should provide ample material for extending studies in this area. Future research into the function of the pyrenoid and other chloroplast features, such as thylakoids, in the operation of a chloroplast-based CCM needs to be addressed in a diverse range of algal species. This should be approached together with assessment of the coevolution of Rubisco, particularly the evolution of Red Form I Rubisco enzymes, which appear to achieve superior kinetic characteristics when compared with the Rubisco of C3 higher plants, which are derived from green algal ancestors.Key words: Rubisco, CO2-concentrating mechanism, carbonic anhydrase, aquatic photosynthesis, algae, pyrenoids, inorganic carbon.

Cyanobacteria have evolved a significant environmental adaptation, known as a CO(2)-concentrating-mechanism (CCM), that vastly improves photosynthetic performance and survival under limiting CO(2) concentrations. The CCM functions to transport and accumulate inorganic carbon actively (Ci; HCO(3)(-), and CO(2)) within the cell where the Ci pool is utilized to provide elevated CO(2) concentrations around the primary CO(2)-fixing enzyme, ribulose bisphosphate carboxylase-oxygenase (Rubisco). In cyanobacteria, Rubisco is encapsulated in unique micro-compartments known as carboxysomes. Cyanobacteria can possess up to five distinct transport systems for Ci uptake. Through database analysis of some 33 complete genomic DNA sequences for cyanobacteria it is evident that considerable diversity exists in the composition of transporters employed, although in many species this diversity is yet to be confirmed by comparative phenomics. In addition, two types of carboxysomes are known within the cyanobacteria that have apparently arisen by parallel evolution, and considerable progress has been made towards understanding the proteins responsible for carboxysome assembly and function. Progress has also been made towards identifying the primary signal for the induction of the subset of CCM genes known as CO(2)-responsive genes, and transcriptional regulators CcmR and CmpR have been shown to regulate these genes. Finally, some prospects for introducing cyanobacterial CCM components into higher plants are considered, with the objective of engineering plants that make more efficient use of water and nitrogen.

SUMMARY Cyanobacteria are the globally dominant photoautotrophic lineage. Their success is dependent on a set of adaptations collectively termed the CO 2 -concentrating mechanism (CCM). The purpose of the CCM is to support effective CO 2 fixation by enhancing the chemical conditions in the vicinity of the primary CO 2 -fixing enzyme, d -ribulose 1,5-bisphosphate carboxylase/oxygenase (RubisCO), to promote the carboxylase reaction and suppress the oxygenase reaction. In cyanobacteria and some proteobacteria, this is achieved by encapsulation of RubisCO within carboxysomes, which are examples of a group of proteinaceous bodies called bacterial microcompartments. Carboxysomes encapsulate the CO 2 -fixing enzyme within the selectively permeable protein shell and simultaneously encapsulate a carbonic anhydrase enzyme for CO 2 supply from a cytoplasmic bicarbonate pool. These bodies appear to have arisen twice and undergone a process of convergent evolution. While the gross structures of all known carboxysomes are ostensibly very similar, with shared gross features such as a selectively permeable shell layer, each type of carboxysome encapsulates a phyletically distinct form of RubisCO enzyme. Furthermore, the specific proteins forming structures such as the protein shell or the inner RubisCO matrix are not identical between carboxysome types. Each type has evolutionarily distinct forms of the same proteins, as well as proteins that are entirely unrelated to one another. In light of recent developments in the study of carboxysome structure and function, we present this review to summarize the knowledge of the structure and function of both types of carboxysome. We also endeavor to cast light on differing evolutionary trajectories which may have led to the differences observed in extant carboxysomes.

Past increases in yield potential of wheat have largely resulted from improvements in harvest index rather than increased biomass. Further large increases in harvest index are unlikely, but an opportunity exists for increasing productive biomass and harvestable grain. Photosynthetic capacity and efficiency are bottlenecks to raising productivity and there is strong evidence that increasing photosynthesis will increase crop yields provided that other constraints do not become limiting. Even small increases in the rate of net photosynthesis can translate into large increases in biomass and hence yield, since carbon assimilation is integrated over the entire growing season and crop canopy. This review discusses the strategies to increase photosynthesis that are being proposed by the wheat yield consortium in order to increase wheat yields. These include: selection for photosynthetic capacity and efficiency, increasing ear photosynthesis, optimizing canopy photosynthesis, introducing chloroplast CO(2) pumps, increasing RuBP regeneration, improving the thermal stability of Rubisco activase, and replacing wheat Rubisco with that from other species with different kinetic properties.

No abstract