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Plant physiologists set about comprehending the genesis of the C4 photosynthetic pathway after its discovery by Hatch and Slack. They discovered that a sophisticated combination of morphological and biochemical adaptations allowed the plant to concentrate CO2 around RuBisCO to achieve maximum efficiency. We categorize the evolutionary events leading to C4 photosynthesis, beginning with anoxygenic photosynthesis and the evolution of RuBisCO to the cooling of Earth by the Great Oxygenation Event that led to the oxygenic photosynthesis. The evolutionary descent of the C4 plants is a phenomenon that occurred around 30 million years ago. Due to industrialization and population growth, improved photosynthetic efficiency and carbon fixation of C4 plants could contest the current global scenario of rising CO2 concentration. C3 crops engineered with C4 traits, implemented on a large scale, could impact the climate globally. Here we discuss the various strategies used to introduce C4 traits in the C3 plants and the potential techniques to be considered for successful hybridization. [ABSTRACT FROM AUTHOR]

Hydrogen isotope ratios (δ²H) represent an important natural tracer of metabolic processes, but quantitative models of processes controlling H-fractionation in aquatic photosynthetic organisms are lacking. Here, we elucidate the underlying physiological controls of ²H/¹H fractionation in algal lipids by systematically manipulating temperature, light, and CO2(aq) in continuous cultures of the haptophyte Gephyrocapsa oceanica. We analyze the hydrogen isotope fractionation in alkenones (αalkenone), a class of acyl lipids specific to this species and other haptophyte algae. We find a strong decrease in the αalkenone with increasing CO2 (aq) and confirm αalkenone correlates with temperature and light. Based on the known biosynthesis pathways, we develop a cellular model of the δ²H of algal acyl lipids to evaluate processes contributing to these controls on fractionation. Simulations show that longer residence times of NADPH in the chloroplast favor a greater exchange of NADPH with ²H-richer intracellular water, increasing αalkenone. Higher chloroplast CO2(aq) and temperature shorten NADPH residence time by enhancing the carbon fixation and lipid synthesis rates. The inverse correlation of αalkenone to CO2(aq) in our cultures suggests that carbon concentrating mechanisms (CCM) do not achieve a constant saturation of CO2 at the Rubisco site, but rather that chloroplast CO2 varies with external CO2(aq). The pervasive inverse correlation of αalkenone with CO2(aq) in the modern and preindustrial ocean also suggests that natural populations may not attain a constant saturation of Rubisco with the CCM. Rather than reconstructing growth water, αalkenone may be a powerful tool to elucidate the carbon limitation of photosynthesis. [ABSTRACT FROM AUTHOR]

Aquatic photosynthetic microorganisms, cyanobacteria and microalgae, account for almost half of the world's photosynthesis. They absorb carbon di oxide (CO2) as the major substrate to support photosynthesis, the beginning of energy flow into living organisms and one of the primary processes comprising the global carbon cycle. Among all photosynthetic mechanisms, inorganic carbon transport into ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBisCO) is one of the major limiting steps in photosynthetic carbon fixation which involves active transport of HCO3−, CO2and/or H+, or an energized biochemical mechanism. In fact, a unique system “carbon concentrating mechanism (CCM)” manages the inorganic carbon assimilation, accumulation of CO2around RuBisCO, and utilization in algal cells. However, the information on mechanism of CO2uptake and fixation inside the algal cells is limited. In order to make strategies for enhancement of CO2fixation, understanding of CCM is crucial. Thus, this review provides an overview of advances in CCM research, the comparative state of the art and reports on the CO2uptake model in cyanobacteria and microalgae. The review also discusses the challenges and future perspectives associated with algal CCM research. [ABSTRACT FROM AUTHOR]

The gammaproteobacterium Thiomicrospira crunogena XCL-2 is an aerobic sulfur-oxidizing hydrothermal vent chemolithoautotroph that has a CO concentrating mechanism (CCM), which generates intracellular dissolved inorganic carbon (DIC) concentrations much higher than extracellular, thereby providing substrate for carbon fixation at sufficient rate. This CCM presumably requires at least one active DIC transporter to generate the elevated intracellular concentrations of DIC measured in this organism. In this study, the half-saturation constant ( K) for purified carboxysomal RubisCO was measured (276 ± 18 µM) which was much greater than the K of whole cells (1.03 µM), highlighting the degree to which the CCM facilitates CO fixation under low CO conditions. To clarify the bioenergetics powering active DIC uptake, cells were incubated in the presence of inhibitors targeting ATP synthesis (DCCD) or proton potential (CCCP). Incubations with each of these inhibitors resulted in diminished intracellular ATP, DIC, and fixed carbon, despite an absence of an inhibitory effect on proton potential in the DCCD-incubated cells. Electron transport complexes NADH dehydrogenase and the bc complex were found to be insensitive to DCCD, suggesting that ATP synthase was the primary target of DCCD. Given the correlation of DIC uptake to the intracellular ATP concentration, the ABC transporter genes were targeted by qRT-PCR, but were not upregulated under low-DIC conditions. As the T. crunogena genome does not include orthologs of any genes encoding known DIC uptake systems, these data suggest that a novel, yet to be identified, ATP- and proton potential-dependent DIC transporter is active in this bacterium. This transporter serves to facilitate growth by T. crunogena and other Thiomicrospiras in the many habitats where they are found. [ABSTRACT FROM AUTHOR]

As all plastids that have been investigated so far can be traced back to endosymbiotic uptake of cyanobacteria by heterotrophic host cells, they accordingly show a high similarity regarding photosynthesis, which includes both the photosystems and the biochemical reactions around the CO 2 fixation via the Calvin–Bassham cycle. Major differences between the different algal and plant groups may include the presence or absence of carbon concentrating mechanisms, pyrenoids, Rubisco activases, carbonic anhydrases as well as differences in the regulation of the Calvin–Bassham cycle. This review describes the diversity of primary carbon fixation steps in algae and plants and the respective regulatory mechanisms. [ABSTRACT FROM AUTHOR]

An acid pH in the lumen of chloroplast thylakoids is necessary in order to derive the required amount of CO2 to account for the observed rates of carbon fixation. We point out that the endosymbiotic derivation of the chloroplast from a cyanobacterium would have resulted in the lumen of the thylakoid having an acid pH. The thylakoids of cyanobacteria are continuous with the plasma membrane, resulting in the lumen of the thylakoid being open to the outside of the cell. Endosymbiosis resulted in the cyanobacterium being taken up into a food vacuole of a protozoan. The vacuole would have had an acid pH, probably around pH 5, so the endosymbiotic bacterium would have been surrounded by an environment with an acidic pH. The lumen of the thylakoids would have been at an acid pH since they were open to the exterior of the cell, and to the contents of the vacuole. [ABSTRACT FROM AUTHOR]

Many autotrophic microorganisms are likely to adapt to scarcity in dissolved inorganic carbon (DIC; CO2 + HCO3- + CO32-) with CO2 concentrating mechanisms (CCM) that actively transport DIC across the cell membrane to facilitate carbon fixation. Surprisingly, DIC transport has been well studied among cyanobacteria and microalgae only. The deep-sea vent gammaproteobacterial chemolithoautotroph Thiomicrospira crunogena has a low-DIC inducible CCM, though the mechanism for uptake is unclear, as homologs to cyanobacterial transporters are absent. To identify the components of this CCM, proteomes of T. crunogena cultivated under low- and high-DIC conditions were compared. Fourteen proteins, including those comprising carboxysomes, were at least 4-fold more abundant under low-DIC conditions. One of these proteins was encoded by Tcr_0854; strains carrying mutated copies of this gene, as well as the adjacent Tcr_0853, required elevated DIC for growth. Strains carrying mutated copies of Tcr_0853 and Tcr_0854 overexpressed carboxysomes and had diminished ability to accumulate intracellular DIC. Based on reverse transcription (RT)-PCR, Tcr_0853 and Tcr_0854 were cotranscribed and upregulated under low-DIC conditions. The Tcr_0853-encoded protein was predicted to have 13 transmembrane helices. Given the mutant phenotypes described above, Tcr_0853 and Tcr_0854 may encode a two-subunit DIC transporter that belongs to a previously undescribed transporter family, though it is widespread among autotrophs from multiple phyla. [ABSTRACT FROM AUTHOR]