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Notes: Abstract Photosynthesis was studied in four species of red marine macroalgae: Palmaria palmata, Laurencia pinnatifida, Lomentaria articulata and Delesseria sanguinea. The rate of O2 evolution for submersed photosynthesis was measured as a function of incident photon flux density at normal pH and inorganic carbon concentration (pH 8.0, 2 mol m−3), and as a function of inorganic carbon concentration at pH 8.0 at saturating and at limiting photon flux density. The rate of CO2 uptake was measured for emersed photosynthesis as a function of CO2 partial pressure at saturating photon flux density. Previous pH-drift results suggest that Palmaria and Laurencia are able to use HCO inf3 sup− as well as CO2 whereas Lomentaria and Delesseria are restricted to CO2. None of the algae are saturated by 2 mol m−3 inorganic carbon at high light (400 μmol m−2 s−1) but are saturated at low light (35 μmol m−2 s−1). The inorganic C concentration at which half the light-saturated rate of O2 evolution is achieved is higher for Palmaria and Laurencia (1.51 and 1.85 mol m−3) than for Lomentaria and Delesseria (0.772 and 0.841 mol m−3). The lower values for the latter two species could reflect their putative restriction to CO2. If expressed in terms of CO2, the half-saturation values yield 7.2 and 7.8 mmol m−3 respectively, which are very similar to values obtained previously during pH-drift experiments but at lower concentrations of HCO inf3 sup− , consistent with restriction to CO2. The photosynthetic conductance (m s−1), calculated from the initial slope for photosynthesis at low concentrations of inorganic carbon, correlates with the suggested ability to extract inorganic carbon based on pH-drift results. Calculations made assuming that CO2 is the only species diffusing across the boundary layer are consistent with boundary layer thicknesses of 20 and 19 μm for Lomentaria and Delesseria respectively, which is feasible given the rapid water movement in the experiments. For Laurencia however, an unreasonably small boundary layer thickness of 6 μm is necessary to explain the flux, which indicates co-diffusion by HCO inf3 sup− . In the apparent absence of external carbonic anhydrase, direct uptake of HCO inf3 sup− , rather than external conversion to CO2 is indicated in this species. In air, the CO2 concentration at which photosynthesis is half-maximal increases in the same order as the ability to raise pH in drift experiments. At 21 kPa the CO2 compensation partial pressures for Palmaria and Laurencia at 0.56 and 1.3 Pa are low enough to suggest a carbon-concentrating mechanism is operating, while those of Lomentaria at 1.8 Pa and particularly that of Delesseria at 4.5 Pa could be explained without a carbon-concentrating mechanism. The algae tested (all except Delesseria) showed more O2 evolution than could be accounted for with a photosynthetic quotient of 1.0 and uncatalysed conversion of HCO inf3 sup− to CO2 outside the cell in high light at pH 8.0 when high algal fresh weight per unit medium was used. These results are concordant with other data suggesting use of HCO inf3 sup− by Palmaria and Laurencia, but discordant with the rest of the available information in indicating use of HCO inf3 sup− by Lomentaria. The reason for this is unclear. The lightsaturated rate of O2 evolution on an algal area basis and the photon flux density needed to saturate photosynthesis were related partly to the habitat from which the seaweeds were collected, but more strongly to the ability to use HCO inf3 sup− . Values for the two users of HCO inf3 sup− , Palmaria (population used was intertidal; also occurs subtidally) and Laurencia (intertidal/shaded intertidal), were greater than for Lomentaria (shaded intertidal), which was greater than Delesseria (subtidal), both of which are believed to be restricted to CO2. In accordance with earlier δ13C data and, for Delesseria, estimates of the achieved growth rates in situ, carbon is likely to be saturating and use of HCO inf3 sup− is unlikely to occur in the normal low-light habitats of Lomentaria and Delesseria. Analysis of N-use efficiencies show that they are closer to the low-CO2-affinity Laminariales than the high-CO2-affinity Fucaceae.

Notes: Summary The natural abundance13C/12C ratios (as δ13C) of organic matter of marine macroalgae from Fife and Angus (East Scotland) were measured for comparison with the species' ability to use CO2 and HCO 3 - for photosynthesis, as deduced from previously published pH-drift measurements. There was a clear difference in δ13C values for species able or unable to use HCO 3 - . Six species of Chlorophyta, 12 species of Phaeophyta and 8 species of Rhodophyta that the pH-drift data suggested could use HCO 3 - had δ13C values in the range -8.81‰ to -22.55‰. A further 6 species of Rhodophyta which the pH-drift data suggested could only use CO2 had δ13C values in the range -29.90‰ to-34.51‰. One of these six species (Lomentaria articulata) is intertidal; the other five are subtidal and so have no access to atmospheric CO2 to complicate the analysis. For these species, calculations based on the measured δ13C of the algae, the δ13C of CO2 in seawater, and the known13C/12C discrimination of CO2 diffusion and RUBISCO carboxylation suggest that only 15–21% of the limitation to photosynthesisin situ results from CO2 diffusion from the bulk medium to the plastids; the remaining 79–85% is associated with carboxylation reactions (and, via feedback effects, down-stream processes). This analysis has been extended for one of these five species,Delesseria sanguinea, by incorporating data onin situ specific growth rates, respiratory rates measured in the laboratory, and applying Fick's law of diffusion to calculate a boundary layer thickness of 17–24 μm. This value is reasonable for aDelesseria sanguinea frondin situ. For HCO 3 - -using marine macroalgae the range of δ13C values measured can be accommodated by a CO2 efflux from algal cells which range from 0.306 of the gross HCO 3 - influx forEnteromorpha intestinalis (δ13C=-8.81‰) in a rockpool to 0.787 forChondrus crispus (δ13C=-22.55‰). The relatively high computed CO2 efflux for those HCO 3 - -users with the more negative δ13C values implies a relatively high photon cost of C assimilation; the observed photon costs can be accommodated by assuming coupled, energy-independent inorganic carbon influx and efflux. The observed δ13C values are also interpreted in terms of water movement regimes and obtaining CO2 from the atmosphere. Published δ13C values for freshwater macrophytes were compared with the ability of the species to use CO2 and HCO 3 - and again there was an apparent separation in δ13C values for these two groups. δ13C values obtained for marine macroalgae for which no pH-drift data are available permit predictions, as yet untested, as to whether they use predominantly CO2 or HCO 3 -