Operation Of CO2-concentrating Mechanism Research Articles

Diversity of CO2 Concentrating Mechanisms in Macroalgae Photosynthesis: A Case Study of Ulva sp.

Many algae respond to the CO2 limitation in seawater by inducing a CO2 concentrating mechanism (CCM) to obtain sufficient inorganic carbon to meet their photosynthetic needs, and Ulva sp. is a model population suitable for studying the ecological adaptability of macroalgae. As the dominant species of green tide disaster, Ulva sp. often faces strong inorganic carbon restriction due to its rapid growth and high population density and must have evolved a variety of carbon acquisition strategies, such as CCM, to overcome these limitations. This paper briefly summarizes the position and function of the important components of CCM (inorganic carbon transporters, carbonic anhydrase, Rubisco, and pyrenoid) and introduces several indexes suitable for evaluating the relative function of CCMs in macroalgae from the aspects of affinity between photosynthesis and Rubisco for CO2, and carbonic anhydrase inhibitor. The methods of judging the carbon sequestration pathway of Ulva sp., the CCM responses of diversity under different carbon sources, and the related genes that may be involved in the operation of CCMs were summarized. This work could provide a reference for revealing the CCMs of macroalgae and lay a foundation for further research on the inorganic carbon utilization strategy of the Ulva sp.

Pyrenoid Starch Sheath Is Required for LCIB Localization and the CO2-Concentrating Mechanism in Green Algae.

Aquatic photosynthetic organisms induce a CO2-concentrating mechanism (CCM) to overcome the difficulty of acquiring inorganic carbon under CO2-limiting conditions. As part of the CCM, the CO2-fixing enzyme Rubisco is enriched in the pyrenoid located in the chloroplast, and, in many green algae, several thick starch plates surround the pyrenoid to form a starch sheath. In Chlamydomonas reinhardtii, low-CO2-inducible protein B (LCIB), which is an essential factor for the CCM, displays altered cellular localization in response to a decrease in environmental CO2 concentration, moving from dispersed throughout the chloroplast stroma to around the pyrenoid. However, the mechanism behind LCIB migration remains poorly understood. Here, we report the characteristics of an Isoamylase1-less mutant (4-D1), which shows aberrant LCIB localization and starch sheath formation. Under very-low-CO2 conditions, 4-D1 showed retarded growth, lower photosynthetic affinities against inorganic carbon, and a decreased accumulation level of the HCO3 - transporter HLA3. The aberrant localization of LCIB was also observed in another starch-sheathless mutant sta11-1, but not in sta2-1, which possesses a thinned starch sheath. These results suggest that the starch sheath around the pyrenoid is required for the correct localization of LCIB and for the operation of CCM.

Adaptation of a Chlamydomonas mutant with reduced rate of photorespiration to different concentrations of CO2

It has been widely accepted that Chlamydomonas reinhardtii cells utilize inorganic carbon very efficiently for photosynthesis by operating a CO2-concentrating mechanism (CCM) under conditions of limited CO2. To help define the mechanism, 7FR2N, one of the suppressor double mutants of phosphoglycolate phosphatase-deficient (pgp1) mutants that have a reduced photorespiration rate (RPR) was crossed with wild-type strains to generate the strain N21 as a single RPR mutant. The comparison of photosynthetic characteristics with wild-type strains after the cells adapted to different concentrations of CO2 revealed that photosynthetic affinity for inorganic carbon was higher than that in wild-type strains after adaptation to concentrations between 50 µL·L–1 CO2 and 5% CO2. Chlorophyll fluorescence parameters were also compared, and the biggest difference between N21 and the wild-type strains was observed in the photochemical quenching and effective quantum yield of photosystem II (ΔF/Fm′) at the CO2 compensation point. These values in N21 increased in a similar manner to the photosynthetic affinity for CO2, and increased significantly when the cells adapted to low-CO2 levels, whereas the values in the wild-type strains were apparently lower without any significant changes, regardless of the CO2 concentrations to which they were adapted. Although it was not clear if a nonphotochemical quenching parameter (NPQ) in N21 was higher than that in wild-type strains, NPQ increased coincidentally with the increase in photosynthetic affinity for inorganic carbon when the CO2 concentrations to which the strains were adapted decreased, in both the mutant and wild-type strain, suggesting that this form of NPQ reflects the operation of CCM in certain conditions. Possible candidates for the RPR mutation and the relationship between CCM and photosynthetic electron flow are discussed.Key words: Chlamydomonas reinhardtii, chlorophyll fluorescence, CO2-concentrating mechanism, low-CO2 responsive gene, phosphoglycolate phosphatase, photorespiration.

The diversity and coevolution of Rubisco, plastids, pyrenoids, and chloroplast-based CO2-concentrating mechanisms in algae

Algae have adopted two primary strategies to maximize the performance of Rubisco in photosynthetic CO2fixation. 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 CO2when the algae have adopted a strategy for CO2fixation 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 O2and 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 C3higher plants, which are derived from green algal ancestors.Key words: Rubisco, CO2-concentrating mechanism, carbonic anhydrase, aquatic photosynthesis, algae, pyrenoids, inorganic carbon.

Evidence for the functioning of photosynthetic CO2-concentrating mechanisms in lichens containing green algal and cyanobacterial photobionts

The photosynthetic properties of a range of lichens containing both green algal (11 species) and cyanobacterial (6 species) photobionts were examined with the aim of determining if there was clear evidence for the operation of a CO2-concentrating mechanism (CCM) within the photobionts. Using a CO2-gas-exchange system, which allowed resolution of fast transients, evidence was obtained for the existence of an inorganic carbon pool which accumulated in the light and was released in the dark. The pool was large (500–1000 nmol · mg Chl) in cyanobacterial lichens and about tenfold smaller in green algal lichens. In Hypogymnia physodes (L.) Nyl., which contains the green alga Trebouxia jamesii, a small inorganic carbon pool was rapidly formed in the light. Carbon dioxide was released from this pool into the gas phase upon darkening within about 20 s when photosynthesis was inhibited by the carbon-reduction-cycle inhibitor glycolaldehyde. In the absence of this inhibitor, release appeared to be obscured by carboxylation of ribulose bisphosphate. The kinetics of CO2 uptake and release were monophasic. The operation of an active CCM could be distinguished from passive accumulation and release accompanying the reversible light-dependent alkalization of the stroma by the presence of saturation characteristics with respect to external CO2. In Peltigera canina (L.) Willd., which contains the cyanobacterium Nostoc sp., a larger CO2 pool was taken up over a longer period in the light and the release of this pool in the dark was slow, lasting 3–5 min. This pool also accumulated in the presence of glycolaldehyde, and under these conditions the CO2 release was biphasic. In both species, photosynthesis at low CO2 was inhibited by the carbonic-anhydrase inhibitor ethoxyzolamide (EZ). Inhibition could be reversed fully or to a considerable extent by high CO2. In Peltigera, EZ decreased both the accumulation of the CO2 pool by the CCM and the rate of photosynthesis. Free-living cultures of Nostoc sp. showed a similar effect of EZ on photosynthesis, although it was more dramatic than that seen with the lichen thalli. In contrast, in Hypogymnia, EZ actually increased the size of the CO2 pool, although it inhibited photosynthesis. This effect was also seen when glycolaldehyde was present together with EZ. Surprisingly, EZ did not alter the kinetics of either CO2 uptake or release. Taken together, the evidence indicates the operation in cyanobacterial lichens of a CCM which is capable of considerable elevation of internal CO2 and is similar to that reported for free-living cyanobacteria. The CCM of green algal lichens accumulates much less CO2 and is probably less effective than that which operates in cyanobacterial lichens.