Interactions between carbon dioxide and oxygen in the photosynthesis of three species of marine red macroalgae

Summary

Red algae have the highest known selectivity factor (S_rel) for CO₂ over O₂ of ribulose bisphosphate carboxylase-oxygenase (RUBISCO). This allows the prediction that a red alga relying on diffusive supply of CO₂ to RUBISCO from air-equilibrated solution should have less O₂ inhibition of photosynthesis than would an otherwise similar non-red alga with a lower S_rel of RUBISCO. Furthermore, RUBISCO shows an increased S_rel values at low temperatures. The prediction that O ₂inhibition of photosynthesis should be small for marine red algae relying on diffusive CO₂ entry growing in the North Sea with an annual temperature range of 4–16°C was tested in O₂ electrode experiments at 12°C. Phycodrys rubens and Plocamium cartilagineum, which rely on diffusive CO₂ entry showed, as predicted, only a small inhibition at lower inorganic C concentrations. Palmaria palmata, which has a CO₂ concentrating mechanism, had the expected negligible O ₂ inhibition of photosynthesis at any inorganic C concentration except (non-significantly) for saturating inorganic C.     

Present and potential uses of the natural abundance of stable isotopes in plant science, with illustrations from the marine environment

The use of stable isotope natural abundance measurements in plant ecophysiological research is discussed in the context of studies of ¹³C/¹²C ratios in marine plants, with emphasis on the uniqueness of the information given by natural abundance measurements and of the importance of complementary data obtained by other techniques in making full use of the natural abundance data. (1) Inorganic C acquisition and assimilation in marine plants can involve diffusive entry of CO₂, or the occurrence of a CO₂-concentrating mechanism frequently involving active HCO₃^? influx. For diffusive CO₂ entry, the δ¹³C measurements can give unique information on the fractional limitation of photosynthesis by CO₂ transport which, with photosynthetic rate measurements, can be used to compute transport conductances. For active HCO₃^?, influx, the δ¹³C values uniquely permit computation of the ratio of the bidirection fluxes (influx/efflux) which, with photon yield data, can be used to given information on the mechanism of the efflux. The analyses are absolutely dependent on external (non-δ¹³C) data distinguishing between diffusive CO₂ entry and the occurrence of a CO₂ concentrating mechanism. (2) δ¹³C measurements on marine photolithotrophs and on members of other trophic levels collected from the sea can give unique data on food webs, with measurements of δ values for other isotopes and compositional data adding precision to the interpretations. (3) Measurements of in situδ¹³C values for extant marine photolithotrophs, compared with δ¹³C values for ancient atmospheric CO₂, can give unique information on the mechanism of atmospheric CO₂ draw-down at the start of glacials; other information permits more concrete conclusions to be drawn.     

Seasonal CO2 exchange patterns of developing peach (Prunus persica) fruits in response to temperature, light and CO2 concentration

CO₂ exchange rates per unit dry weight, measured in the field on attached fruits of the late-maturing Cal Red peach cultivar, at 1200 μmol photons m^?2S^?1 and in dark, and photosynthetic rates, calculated by the difference between the rates of CO₂ evolution in light and dark, declined over the growing season. Calculated photosynthetic rates per fruit increased over the season with increasing fruit dry matter, but declined in maturing fruits apparently coinciding with the loss of chlorophyll. Slight net fruit photosynthetic rates ranging from 0. 087 ± 0. 06 to 0. 003 ± 0. 05 nmol CO₂ (g dry weight)^?1 S^?1 were measured in midseason under optimal temperature (15 and 20°C) and light (1200 μmol photons m^?2 S^?1) conditions. Calculated fruit photosynthetic rates per unit dry weight increased with increasing temperatures and photon flux densities during fruit development. Dark respiration rates per unit dry weight doubled within a temperature interval of 10°C; the mean seasonal O₁₀ value was 2. 03 between 20 and 30°C. The highest photosynthetic rates were measured at 35°C throughout the growing season. Since dark respiration rates increased at high temperatures to a greater extent than CO₂ exchange rates in light, fruit photosynthesis was apparently stimulated by high internal CO₂ concentrations via CO₂ refixation. At 15°C, fruit photosynthetic rates tended to be saturated at about 600 μmol photons m^?2 S^?1. Young peach fruits responded to increasing ambient CO₂ concentrations with decreasing net CO₂ exchange rates in light, but more mature fruits did not respond to increases in ambient CO₂. Fruit CO₂ exchange rates in the dark remained fairly constant, apparently uninfluenced by ambient CO₂ concentrations during the entire growing season. Calculated fruit photosynthetic rates clearly revealed the difference in CO₂ response of young and mature peach fruits. Photosynthetic rates of younger peach fruits apparently approached saturation at 370 μl CO₂1^?2. In CO₂ free air, fruit photosynthesis was dependent on CO₂ refixation since CO₂ uptake by the fruits from the external atmosphere was not possible. The difference in photosynthetic rates between fruits in CO₂-free air and 370 μl CO₂ 1^?1 indicated that young peach fruits were apparently able to take up CO₂ from the external atmosphere. CO₂ uptake by peach fruits contributed between 28 and 16% to the fruit photosynthetic rate early in the season, whereas photosynthesis in maturing fruits was supplied entirely by CO₂ refixation.     

Membrane-Associated Polypeptides Induced in Chlamydomonas by Limiting CO(2) Concentrations

Chlamydomonas reinhardtii and other unicellular green algae have a high apparent affinity for CO₂, little O₂ inhibition of photosynthesis, and reduced photorespiration. These characteristics result from operation of a CO₂-concentrating system. The CO₂-concentrating system involves active inorganic carbon transport and is under environmental control. Cells grown at limiting CO₂ concentrations have inorganic carbon transport activity, but cells grown at 5% CO₂ do not. Four membrane-associated polypeptides (M_r 19, 21, 35, and 36 kilodaltons) have been identified which either appear or increase in abundance during adaptation to limiting CO₂ concentrations. The appearance of two of the polypeptides occurs over roughly the same time course as the appearance of the CO₂-concentrating system activity in response to CO₂ limitation.     

TRANSPORT OF INORGANIC CARBON AND THE ‘CO2 CONCENTRATING MECHANISM’ IN CHLORELLA EMERSONII (CHLOROPHYCEAE)1

Chlorella emersonii Shihira et Krauss var. emersonii exhibits ‘C₄-like’ gas exchange characteristics when grown at air levels of CO₂, but is ‘C₃-like’ when grown with extra CO₂. The total inorganic carbon concentration, and the free CO₂ concentration, averaged over the cell interior are higher in air-adapted cells than can be accounted for by passive CO₂ equilibration from the medium and the mean intracellular pH value. The ‘extra’ inorganic C in the air-grown cells probably cannot all be accounted for in terms of binding to proteins and requires an active transport process to account for it. The electrical potential of the cell interior becomes more negative when the ‘CO₂ concentrating mechanism’ is operative; this is most readily explained if the active step in inorganic C accumulation is primary active uniport of HCO₃^?. Since the ‘CO₂ concentrating mechanism’ can operate when CO₂ is the species crossing the outer permeation barrier, it is suggested that the site of active HCO₃^? transport in Chlorella (and other eukaryotes) is the chloroplast envelope, and the plasmalemma in cyanobacteria. This scheme explains the obligatory role of the de-repressed carbonic anhydrase in C₄-like photosynthesis in algae, but some other data support an explanation of C₄-like photosynthesis in terms of special properties of carbonic anhydrase as a carbon donor to RuBP carboxylase-oxygenase.     

The induction of the CO2-concentrating mechanism is correlated with the formation of the starch sheath around the pyrenoid of Chlamydomonas reinhardtii

The pyrenoid is a prominent proteinaceous structure found in the stroma of the chloroplast in unicellular eukaryotic algae, most multicellular algae, and some hornworts. The pyrenoid contains the enzyme ribulose-1,5-bisphosphate carboxylase/oxygenase and is sometimes surrounded by a carbohydrate sheath. We have observed in the unicellular green alga Chlamydomonas reinhardtii Dangeard that the pyrenoid starch sheath is formed rapidly in response to a decrease in the CO₂ concentration in the environment. This formation of the starch sheath occurs coincidentally with the induction of the CO₂-concentrating mechanism. Pyrenoid starch-sheath formation is partly inhibited by the presence of acetate in the growth medium under light and low-CO₂ conditions. These growth conditions also partly inhibit the induction of the CO₂-concentrating mechanism. When cells are grown with acetate in the dark, the CO₂-concentrating mechanism is not induced and the pyrenoid starch sheath is not formed even though there is a large accumulation of starch in the chloroplast stroma. These observations indicate that pyrenoid starch-sheath formation correlates with induction of the CO₂-concentrating mechanism under low-CO₂ conditions. We suggest that this ultrastructural reorganization under lowCO₂ conditions plays a role in the CO₂-concentrating mechanism C. reinhardtii as well as in other eukaryotic algae.     

A pyrenoid-based carbon-concentrating mechanism is present in terrestrial bryophytes of the class Anthocerotae

It has been widely accepted that carbon assimilation in bryophytes is exclusively based on the conventional C₃ photosynthetic pathway. The occurrence of biochemical CO₂-concentrating mechanisms (C₄ or Crassulacean acid metabolism), which have developed in plants in the last 20–100 million years, has been discounted for bryophytes from studies of the carbon isotope composition (¹³C) of organic material. In contrast cyanobacteria and many algae show active accumulation of dissolved inorganic carbon via biophysical CO₂-concentrating mechanisms which are also found in the photobiont partners in certain lichens. The presence of a pyrenoid, a granular particle within the chloroplast, has been linked with CO₂-concentrating mechanism activity in green algae and lichens and we now show that such a mechanism is categorically associated with the occurrence of a pyrenoid in bryophytes belonging to the class of Anthocerotae. These observations have significant evolutionary implications for the development of terrestrial photosynthesis during the colonisation of the land, raising the intriguing question of why the pyrenoid-based CO₂-concentrating mechanism did not persist in the terrestrial environment.Abbreviations and Symbols CCM carbon-concentrating mechanism - DIG dissolved inorganic carbon (CO₂+HCO ₃ ^- +CO ₂ ^- ) - DW dry weight - K_0.5 external concentration of CO₂ at which half-maximal rates of CO₂ assimilation are reached - Rubisco ribulose-l,5-bisphosphate carboxylase-oxygenase - carbon isotope discrimination (%) - ¹³C carbon isotope ratio (%) This work was supported by the Natural Environment Research Council (GR3/8813) and the Leverhulme Trust. We thank Prof. A. Roy Perry (National Museum of Wales, Cardiff), Dr. B. Coppins and Mr. D. Long (Royal Botanic Garden Edinburgh) for access to herbarium specimens and Mr. M. Fletcher for providing living bryophytes.     

Carbonic anhydrase activity and inorganic carbon fluxes in low- and high-C1 cells of Chlamydomonas reinhardtü and Scenedesmus obliquus

Carbonic anhydrase (CA) activity associated with high- and low-dissolved inorganic carbon (C₁) grown cells was examined in whole cells by measuring ¹⁸O exchange from doubly labeled CO₂ (¹³C¹⁸O¹⁸O). Both algal species showed the presence of extracellular (periplasmic) as well as intracellular CA activity, which were both greatly increased in low-C₁ cells. The periplasmic CA activity was at least 40-fold higher in lowcompared to high-C₁ cells in both C. reinhardtii and S. obliquus. while low-C₁ cells of S. obliquus showed the highest activity of internal CA. The CA inhibitor ethoxyzolamide showed a strong inhibition of the C₁ uptake process in both C. reinhardtii and S. obliquus as in cyanobacteria. which may indicate that the nature of the primary uptake process is similar in both green algae and cyanobacteria. By using a mass spectrometnc disequilibrium technique it was possible to separate the C₁ fluxes of net HCO^?₃-uptake and net CO₂-uptake during steady-state photosynthesis in high- and Sow-C₁ grown cells of Chlamydomonas reinhardtii (WT. 2137+) and Scenedesmus obliquus (WT. D3). It was found that both high- and low-C₁ cells of the two algae can utilize both CO₂ and HCO^?₃ for photosynthesis, although low-C₁ cells have a higher affinity for the uptake of both C₁ species. Induction at low-C₁ causes an increase in the affinity of both species for HCO^?₃ and CO₂; changes in net CO₂-uptake were, however, significantly greater.     

Carbon fixation and carbon availability in marine phytoplankton

It is widely believed that inorganic C does not limit the rate of short-term photosynthesis, the net productivity, or the maximum biomass, of marine phytoplankton. This lack of inorganic C restriction is less widely believed to hold for phytoplankton in many low alkalinity freshwaters or for seaweed in nutrient-enriched rock pools. These views are examined in the context of the physical chemistry of the inorganic C system in natural waters and of the ways in which various taxa of phytoplankton deal with inorganic C and discriminate between ¹²C and ¹³C. Using this information to interpret data obtained in the ocean or in freshwater suggests that short-term photosynthesis, production rate, and achieved biomass, of phytoplankton are rarely limited by inorganic C supply but, rather, that the widely suggested factors of limited light, nitrogen or phosphorus supply are the resource inputs which restrict productivity. Global change, by increasing atmospheric CO₂ partial pressure and global mean temperatures, is likely to increase the mean CO₂ concentration in the atmosphere, but the corresponding change in the oceans will be much less. There are, however, genotypic differences in the handling of inorganic C among the diversity of marine phytoplankton, and in impact on use of limiting nutrients, so increases in the mean CO₂ and HCO₃ ^- concentrations in surface ocean waters could cause changes in species composition. However, the rarity of inorganic C limitation of marine phytoplankton short-term photosynthesis, net productivity, or the maximum biomass, in today's ocean means that global change is unlikely to increase these three values in the ocean.     

Growth, photosynthetic properties and Rubisco activities and amounts of marine macroalgae grown under current and elevated seawater CO2 concentrations

Growth rates, photosynthetic responses and the activity, amount and CO₂ affinity of ribulose‐1,5‐bisphosphate carboxylase/oxygenase (Rubisco) were determined for common marine macroalgae grown in seawater (containing 14.5 ± 2.1 µM CO₂) or CO₂‐enriched seawater (averaging 52.8 ± 19.2 µM CO₂). The algae were grown in 40 L fiberglass tanks (outdoor) for 4–15 weeks and in a field experimental setup for 5 days. Growth rates of the species studied (representing the three major divisions, i.e. Chlorophyta, Rhodophyta and Phaeophyta) were generally not significantly affected by the increased CO₂ concentrations in the seawater medium. Rubisco characteristics of algae cultivated in CO₂‐enriched seawater were similar to those of algae grown in nonenriched seawater. The lack of response of photosynthetic traits in these aquatic plants is likely to be because of the presence of CO₂ concentrating mechanisms (CCMs) which rely on HCO₃^– utilization, the inorganic carbon (Ci) form that dominates the total Ci pool available in seawater. Significant changes on the productivity of these particular marine algae species would not be anticipated when facing future increasing atmospheric CO₂ levels.     

CO<Subscript>2</Subscript> sequestration in plants: lesson from divergent strategies

Most organisms inhabiting earth feed directly or indirectly on the products synthesized by the reaction of photosynthesis, which at the current atmospheric CO₂ levels operates only at two thirds of its peak efficiency. Restricting the photorespiratory loss of carbon and thereby improving the efficiency of photosynthesis is seen by many as a good option to enhance productivity of food crops. Research during last half a century has shown that several plant species developed CO₂-concentrating mechanism (CCM) to restrict photorespiration under lower concentration of available CO₂. CCMs are now known to be operative in several terrestrial and aquatic plants, ranging from most advanced higher plants to algae, cyanobacteria and diatoms. Plants with C₄ pathway of photosynthesis (where four-carbon compound is the first product of photosynthesis) or crassulacean acid metabolism (CAM) may consistently operate CCM. Some plants however can undergo a shift in photosynthetic metabolism only with change in environmental variables. More recently, a shift in plant photosynthetic metabolism is reported at high altitude where improved efficiency of CO₂ uptake is related to the recapture of photorespiratory loss of carbon. Of the divergent CO₂ assimilation strategies operative in different oraganisms, the capacity to recapture photorespiratory CO₂ could be an important approach to develop plants with efficient photosynthetic capacity.     

Exogenous inorganic carbon sources for photosynthesis in seawater by members of the Fucales and the Laminariales (Phaeophyta): ecological and taxonomic implications

Summary Characteristics of inorganic carbon assimilation by photosynthesis in seawater were investigated in six species of the Fucales (five Fucaceae, one Cystoseiraceae) and four species of the Laminariales (three Laminariaceae, one Alariaceae) from Arbroath, Scotland. All of the algae tested could photosynthesise faster at high external pH values than the uncatalysed conversion of HCO ₃ ^- to CO₂ can occur, i.e. can use external HCO ₃ ^- . They all had detectable extracellular carbonic anhydrase activity, suggesting that HCO ₃ ^- use could involve catalysis of external CO₂ production, a view supported to some extent by experiments with an inhibitor of carbonic anhydrase. All of the algae tested had CO₂ compensation concentrations at pH 8 which were lower than would be expected from diffusive entry of CO₂ supplying RUBISCO as the initial carboxylase, consistent with the operation of energized entry of HCO ₃ ^- and / or CO₂ acting as a CO₂ concentrating mechanism. Quantitative differences among the algae examined were noted with respect to characteristics of inorganic C assimilation. The most obvious distinction was between the eulittoral Fucaceae, which are emersed for part of, or most of, the tidal cycle, and the other three families (Cystoseiraceae, Laminariaceae, Alariaceae) whose representatives are essentially continually submersed. The Fucaceae examined are able to photosynthesise at high pH values, and have lower CO₂ compensation concentrations, and lower K_1/2 values for inorganic C use in photosynthesis, at pH 8, than the other algae tested. Furthermore, the Fucaceae are essentially saturated with inorganic C for photosynthesis at the normal seawater concentration at pH 8 and 10°C. These characteristics are consistent with the dominant role of a CO₂ concentrating mechanism in CO₂ acquisition by these plants. Other species tested have characteristcs which suggest a less effective HCO ₃ ^- use and CO₂ concentrating mechanism, with the Laminariaceae being the least effective; unlike the Fucaceae, photosynthesis by these algae is not saturated with inorganic C in normal seawater. Taxonomic and ecological implications of these results are considered in relation to related data in the literature.     

CO2 uptake and transport in leaf mesophyll cells

Abstract The acquisition of inorganic carbon for photosynthetic assimilation by leaf mesophyll cells and chloroplasts is discussed with particular reference to membrane permeation of CO₂ and HCO^?₃. Experimental evidence indicates that at the apoplast pH normally experienced by leaf mesophyll cells (pH 6–7) CO₂ is the principal species of inorganic carbon taken up. Uptake of HCO^?₃ may also occur under certain circumstances (i.e. pH 8.5), but its contribution to the net flux of inorganic carbon is small and HCO^?₃ uptake does not function as a CO₂-concentrating mechanism. Similarly, CO₂ rather than HCO^?₃ appears to be the species of inorganic carbon which permeates the chloroplast envelope. In contrast to many C₃ aquatic plants and C₄ plants, C₃ terrestrial plants lack specialized mechanisms for the acquisition and transport of inorganic carbon from the intercellular environment to the site of photosynthetic carboxylation, but rely upon the diffusive uptake of CO₂.     

Quantification of the rate of CO2 formation in the periplasmic space of microalgae during photosynthesis. A comparison of whole-cell rate constants for CO2 and HCO3– uptake among three species of the green alga Chlorella

As previously described, the absolute rate of photosynthesis due to a limited concentration of dissolved inorganic carbon at alkaline pH, where the rate of CO₂ formation is strictly limited, plotted as a function of chlorophyll (Chl) concentration, will take the form of a rectangular hyperbola combined with a linear rate directly proportional to [Chl], which are, respectively, due to the contribution of CO₂ and HCO₃^– to photosynthesis. This model represents that the mathematical asymptote of absolute rate of photosynthesis versus cell density is described by the whole-cell rate constant for HCO₃^– uptake and the maximum rate of CO₂ formation in the extracellular space. This means that any trace modification of the CO₂ formation rate outside the cell will alter the photosynthetic rate and should be detectable experimentally. In air-grown Chlorella ellipsoidea and C. kessleri and in high CO₂-grown C. saccharophila, the graph of the absolute rate of photosynthesis against [Chl] clearly followed the mathematical model described above and the actual CO₂ formation rates outside the cells were not significantly different from the calculated rates. It also indicated that the whole-cell rate constants for CO₂ and HCO₃^– uptake in air-grown C. ellipsoidea and C. saccharophila were similar at ≈ 300 and 2·0 mm³μg^–1 Chl min^–1, respectively, whereas those in air-grown C. kessleri were ≈ 550 and 15 mm³μg^–1 Chl min^–1. These results indicate that no acidification of the periplasmic space occurs, and there is no trace activity of external carbonic anhydrase in these microalgae.     

The sensitivity of C3 photosynthesis to increasing CO2 concentration: a theoretical analysis of its dependence on temperature and background CO2 concentration

The atmospheric CO₂ concentration has increased from the pre-industrial concentration of about 280 μmol mol⁻¹ to its present concentration of over 350 μmol mol⁻¹, and continues to increase. As the rate of photosynthesis in C₃ plants is strongly dependent on CO₂ concentration, this should have a marked effect on photosynthesis, and hence on plant growth and productivity. The magnitude of photo-synthetic responses can be calculated based on the well-developed theory of photosynthetic response to intercellular CO₂ concentration. A simple biochemically based model of photosynthesis was coupled to a model of stomatal conductance to calculate photosynthetic responses to ambient CO₂ concentration. In the combined model, photosynthesis was much more responsive to CO₂ at high than at low temperatures. At 350 μmol mol⁻¹, photosynthesis at 35°C reached 51% of the rate that would have been possible with non-limiting CO₂, whereas at 5°C, 77% of the CO₂ non-limited rate was attained. Relative CO₂ sensitivity also became smaller at elevated CO₂, as CO₂ concentration increased towards saturation. As photosynthesis was far from being saturated at the current ambient CO₂ concentration, considerable further gains in photosynthesis were predicted through continuing increases in CO₂ concentration. The strong interaction with temperature also leads to photosynthesis in different global regions experiencing very different sensitivities to increasing CO₂ concentrations.     

Inorganic carbon acquisition by eukaryotic algae: four current questions

The phylogenetically and morphologically diverse eukaryotic algae are typically oxygenic photolithotrophs. They have a diversity of incompletely understood mechanisms of inorganic carbon acquisition: this article reviews four areas where investigations continue. The first topic is diffusive CO₂ entry. Most eukaryotic algae, like all cyanobacteria, have inorganic carbon concentrating mechanisms (CCMs). The ancestral condition was presumably the absence of a CCM, i.e. diffusive CO₂ entry, as found in a small minority of eukaryotic algae today; however, it is likely that, as is found in several cases, this condition is due to a loss of a CCM. There are a number of algae which are in various respects intermediate between diffusive CO₂ entry and occurrence of a CCM: further study is needed on this aspect. A second topic is the nature of cyanelles and their role in inorganic carbon assimilation. The cyanelles (plastids) of the euglyphid amoeba Paulinella have been acquired relatively recently by endosymbiosis with genetic integration of an α-cyanobacterium with a Form 1A Rubisco. The α-carboxysomes in the cyanelles are presumably involved in a CCM, but further investigation is needed.Also called cyanelles are the plastids of glaucocystophycean algae, but is it now clear that these were derived from the β-cyanobacterial ancestor of all plastids other than that of Paulinella. The resemblances of the central body of the cyanelles of glaucocystophycean algae to carboxysomes may not reflect derivation from cyanobacterial β-carboxysomes; although it is clear that these algae have CCMs but these are now well characterized. The other two topics concern CCMs in other eukaryotic algae; these CCMs arose polyphyletically and independently of the cyanobacterial CCMs. It is generally believed that eukaryotic algal, like cyanobacterial, CCMs are based on active transport of an inorganic carbon species and/or protons, and they have C₃ biochemistry. This is the case for the organism considered as the third topic, i.e. Chlamydomonas reinhardtii, the eukaryotic alga with the best understood CCM. This CCM involves HCO₃ ^? conversion to CO₂ in the thylakoid lumen so the external inorganic carbon must cross four membranes in series with a final CO₂ effux from the thylakoid. More remains to be investigated about this CCM. The final topic is that of the occurrence of C₄-like metabolism in the CCMs of marine diatoms. Different conclusions have been reached depending on the organism investigated and the techniques used, and several aspects require further study.     

Adaptation of Chlamydomonas reinhardtii High-CO(2)-Requiring Mutants to Limiting CO(2)

Photosynthetic characteristics of four high-CO₂-requiring mutants of Chlamydomonas reinhardtii were compared to those of wild type before and after a 24-hour exposure to limiting CO₂ concentrations. The four mutants represent two loci involved in the CO₂-concentrating system of this unicellular alga. All mutants had a lower photosynthetic affinity for inorganic carbon than did the wild type when grown at an elevated CO₂ concentration, indicating that the genetic lesion in each is expressed even at elevated CO₂ concentrations. Wild type and all four mutants exhibited adaptive responses to limiting CO₂ characteristic of the induction of the CO₂-concentrating system, resulting in an increased affinity for inorganic carbon only in wild type. Although other components of the CO₂-concentrating system were induced in these mutants, the defective component in each was sufficient to prevent any increase in the affinity for inorganic carbon. It was concluded that the genes corresponding to the ca-1 and pmp-1 loci exhibit at least partially constitutive expression and that all components of the CO₂-concentrating system may be required to significantly affect the photosynthetic affinity for inorganic carbon.     

Carbon isotope ratios of photolithotrophs from allt meall nan damh,a burn at ardeonaig,Perthshire, and their ecophysiological significance

Summary

δ¹³C measurements were made of dissolved inorganic C, and of submerged benthic cyanobacteria, algae and bryophytes, from Allt Meall nan Damh, a burn at Ardeonaig, Perthshire. The δ¹³C of the CO₂, HCO3/- and CO3/2- components of the inorganic C were computed, and the Δ values of the organic C in the photolithotrophs were then calculated relative to dissolved CO₂. The decreasing order of A values in the Ardeonaig Burn is Lemanea and bryophytes ≥ green macroalgae and Audouinella > diatom mats, which is the same as in the Dighty Burn. However, the Δ values of Lemanea and the bryophytes, which depend on diffusive CO₂ entry, are lower at Ardeonaig than in the Dighty Burn, suggesting greater diffusive limitation to photosynthesis in the Ardeonaig Burn. It is not easy to relate this difference in Δ values in Lemanea to the higher C:N atomic ratio in the Ardeonaig Burn (21.2 ± 0.64) than in the Dighty Burn (9.5–11.0). The Δ values relative to HCO3/- for the HCO3/--using diatom mat in the Ardeonaig Burn is also lower than that in the Dighty Burn; this is consistent with a greater diffusion limitation of photosynthesis in the thicker mats in the Ardeonaig Burn. The δ¹³C of a Lyngbya mat overlying a Lemanea population stranded by low summer water levels indicates that some of the C fixed by the HCO3/--using Lyngbya comes from respiration of low-δ¹³C inorganic C by the Lemanea which is shaded by the Lyngbya. The δ¹³C values of Mesotaenium in its mucilage sheath on a thinly vegetated bank is suggestive of predominant use of the higher CO₂ concentrations with lower δ¹³C from groundwater rather than of atmospheric CO₂ yielding lower dissolved CO₂ concentrations with a higher δ¹³C value.     

Photosynthetic response to globally increasing CO2 of co‐occurring temperate seagrass species

Photosynthesis of most seagrass species seems to be limited by present concentrations of dissolved inorganic carbon (DIC). Therefore, the ongoing increase in atmospheric CO₂ could enhance seagrass photosynthesis and internal O₂ supply, and potentially change species competition through differential responses to increasing CO₂ availability among species. We used short‐term photosynthetic responses of nine seagrass species from the south‐west of Australia to test species‐specific responses to enhanced CO₂ and changes in HCO₃^?. Net photosynthesis of all species except Zostera polychlamys were limited at pre‐industrial compared to saturating CO₂ levels at light saturation, suggesting that enhanced CO₂ availability will enhance seagrass performance. Seven out of the nine species were efficient HCO₃^? users through acidification of diffusive boundary layers, production of extracellular carbonic anhydrase, or uptake and internal conversion of HCO₃^?. Species responded differently to near saturating CO₂ implying that increasing atmospheric CO₂ may change competition among seagrass species if co‐occurring in mixed beds. Increasing CO₂ availability also enhanced internal aeration in the one species assessed. We expect that future increases in atmospheric CO₂ will have the strongest impact on seagrass recruits and sparsely vegetated beds, because densely vegetated seagrass beds are most often limited by light and not by inorganic carbon.     

Terrestrial higher-plant response to increasing atmospheric [CO2] in relation to the global carbon cycle

Terrestrial higher plants exchange large amounts of CO₂ with the atmosphere each year; c. 15% of the atmospheric pool of C is assimilated in terrestrial-plant photosynthesis each year, with an about equal amount returned to the atmosphere as CO₂ in plant respiration and the decomposition of soil organic matter and plant litter. Any global change in plant C metabolism can potentially affect atmospheric CO₂ content during the course of years to decades. In particular, plant responses to the presently increasing atmospheric CO₂ concentration might influence the rate of atmospheric CO₂ increase through various biotic feedbacks. Climatic changes caused by increasing atmospheric CO₂ concentration may modulate plant and ecosystem responses to CO₂ concentration. Climatic changes and increases in pollution associated with increasing atmospheric CO₂ concentration may be as significant to plant and ecosystem C balance as CO₂ concentration itself. Moreover, human activities such as deforestation and livestock grazing can have impacts on the C balance and structure of individual terrestrial ecosystems that far outweigh effects of increasing CO₂ concentration and climatic change. In short-term experiments, which in this case means on the order of 10 years or less, elevated atmospheric CO₂ concentration affects terrestrial higher plants in several ways. Elevated CO₂ can stimulate photosynthesis, but plants may acclimate and (or) adapt to a change in atmospheric CO₂ concentration. Acclimation and adaptation of photosynthesis to increasing CO₂ concentration is unlikely to be complete, however. Plant water use efficiency is positively related to CO₂ concentration, implying the potential for more plant growth per unit of precipitation or soil moisture with increasing atmospheric CO₂ concentration. Plant respiration may be inhibited by elevated CO₂ concentration, and although a naive C balance perspective would count this as a benefit to a plant, because respiration is essential for plant growth and health, an inhibition of respiration can be detrimental. The net effect on terrestrial plants of elevated atmospheric CO₂ concentration is generally an increase in growth and C accumulation in phytomass. Published estimations, and speculations about, the magnitude of global terrestrial-plant growth responses to increasing atmospheric CO₂ concentration range from negligible to fantastic. Well-reasoned analyses point to moderate global plant responses to CO₂ concentration. Transfer of C from plants to soils is likely to increase with elevated CO₂ concentrations because of greater plant growth, but quantitative effects of those increased inputs to soils on soil C pool sizes are unknown. Whether increases in leaf-level photosynthesis and short-term plant growth stimulations caused by elevated atmospheric CO₂ concentration will have, by themselves, significant long-term (tens to hundreds of years) effects on ecosystem C storage and atmospheric CO₂ concentration is a matter for speculation, not firm conclusion. Long-term field studies of plant responses to elevated atmospheric CO₂ are needed. These will be expensive, difficult, and by definition, results will not be forthcoming for at least decades. Analyses of plants and ecosystems surrounding natural geological CO₂ degassing vents may provide the best surrogates for long-term controlled experiments, and therefore the most relevant information pertaining to long-term terrestrial-plant responses to elevated CO₂ concentration, but pollutants associated with the vents are a concern in some cases, and quantitative knowledge of the history of atmospheric CO₂ concentrations near vents is limited. On the whole, terrestrial higher-plant responses to increasing atmospheric CO₂ concentration probably act as negative feedbacks on atmospheric CO₂ concentration increases, but they cannot by themselves stop the fossil-fuel-oxidation-driven increase in atmospheric CO₂ concentration. And, in the very long-term, atmospheric CO₂ concentration is controlled by atmosphere-ocean C equilibrium rather than by terrestrial plant and ecosystem responses to atmospheric CO₂ concentration.