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Notes: SUMMARY 1. Thirty small upland lakes in Cumbria, Wales, Scotland and Northern Ireland were visited three times between April and August 2000. On each occasion water chemistry was measured and phytoplankton bioassays were performed in the laboratory to assess growth-rate and yield limitation by phosphorus and nitrogen. In addition, yield limitation of periphyton growth was investigated twice, in situ, using nutrient-diffusing substrata.2. Over the whole season the percentage frequency of P, N and co-limitation was 24, 13 and 63%, respectively, for phytoplankton rate limitation and 20, 22 and 58%, respectively, for phytoplankton yield limitation.3. A clear response of periphyton yield to nutrient additions was found in 75% of all cases and of these, co-limitation was most common (54%). Average percentage frequency for P and N limitation was 26 and 20%, respectively.4. Phytoplankton and periphyton showed seasonal changes in nutrient limitation within sites. In particular, co-limitation became progressively more common as the season progressed.5. The response of phytoplankton growth rate to ammonium and nitrate addition was identical, but ammonium was a slightly better source of nitrogen than nitrate for phytoplankton yield on 7% and for periphyton yield on 10% of the occasions. However, the magnitude of the effect was small.6. The concentration of dissolved inorganic nitrogen (DIN) and the molar ratio of DIN to total dissolved phosphorus (TDP), appeared to be the main environmental factors controlling the extent of nitrogen or phosphorus limitation at a given site. Nitrogen limitation was more likely than phosphorus limitation where the DIN was 〈6.5 mmol m−3 and the ratio of DIN : TDP was 〈53. Co-limitation was the most likely outcome at a DIN concentration 〈13 mmol m−3 and at a DIN : TDP molar ratio 〈250. Above these values phosphorus limitation was most likely.7. The relatively high frequency of nitrogen limitation and co-limitation at higher N : P ratios than previously reported, may result from the inability of nitrogen-fixing cyanobacteria to thrive in these upland lakes where pH and the concentration of phosphorus tended to be low and where flushing rates tended to be high.

Notes: 1. Thirty-six years of winter meteorological and limnological measurements from four lakes in the English Lake District are analysed and related to variations in the North Atlantic Oscillation (NAO). Winter weather conditions were strongly influenced by the NAO with mild, wet winters being associated with strongly positive values of the NAO index (NAOI).2. Lake surface and bottom temperatures were strongly positively correlated with the NAOI, with the highest correlations being recorded in the shallower lakes.3. Variations in the NAOI also had a significant effect on the winter concentration of nitrate. In all the lakes, there was a significant negative correlation between the NAOI and the detrended winter concentration of nitrate. The key driving variable was the local air temperature, which appeared to limit the quantity of nitrate reaching the lake by increasing the amount assimilated in the surrounding catchment in mild winters.4. Dissolved reactive phosphorus (DRP) concentrations were not significantly correlated with the NAOI in the two larger basins but significant positive correlations were recorded in the two smaller lakes. The key driving variable was the local rainfall with higher DRP concentrations being recorded after heavy rain in the lakes with a short retention time.5. The NAOI-related changes in rainfall also influenced the phytoplankton. In wet winters the concentration of chlorophyll in the two smaller lakes with the shortest retention time was lower and the spring growth of Asterionella formosa was delayed in the smallest lake.6. These differential responses demonstrate how the large-scale effects associated with the NAO can be ‘filtered’ by the physical characteristics of a particular site.

Notes: 1. Thirty small upland lakes in Cumbria, Wales, Scotland and Northern Ireland were each visited once during June and July 2000. From each lake, samples of surface sediment epilithon, macrophytes and total dissolved nitrogen (TDN) were collected for nitrogen stable isotope analysis. As part of a wider programme, samples were also collected for chemical analysis and bioassays.2. Considerable variation was found in δ15N values in all measured nitrogen compartments. Some regional variation was evident but was generally weak. Sediment and epilithon δ15N were positively correlated with δ15N of TDN, suggesting that baseline nitrogen isotope ratios influence those in some organic matter compartments in the lakes.3. Sediment δ15N was higher when inorganic nitrogen concentration in the water was low, possibly reflecting reduced isotope fractionation under these conditions. However, this was not the case for epilithon or macrophytes. Sediment δ15N values were also negatively related to annual nitrogen deposition.4. Sediment, epilithon and macrophyte δ15N values all showed significant relations to nutrient limitation in the lakes as determined by algal bioassays. We suggest that sediment δ15N might be developed as a simple integrating measure of the degree of nitrogen limitation in lakes.

Notes: 1. A method for calculating and comparing the stoichiometric yields of phytoplankton standing crop (as chlorophyll a) is presented.2. Some worked examples show how the model distinguishes lakes in which the supportive capacity is constrained mainly by the available phosphorus or by the nitrogen supply or by the light availability. Sites are identified where control moves among the constraining factors.3.  An instance of lake management is given where the diagnosis, remedy and the remediation have conformed closely to the model predictions.

Notes: Rates of photosynthesis by the marine macroalga Ulva lactuca were measured in a factorial experiment at five concentrations of HCO3− and CO32- between 0·20 and 1·26 mol m−3, but very low concentrations of CO2. The results demonstrated that HCO3− was available for use, but an analysis of variance showed that CO32- had neither an inhibiting nor a stimulating effect on rates of photosynthesis over this concentration range. Over the experiment, pH varied from 8·46 to 10·06 and this also had no significant effect on rates of photosynthesis. The lack of a stimulatory effect of high concentrations of CO32- on the rate of photosynthesis at low concentrations of HCO3− was taken as circumstantial evidence for direct uptake of HCO3− rather than proton extrusion and external production of CO2. In the rockpools in which U. lactuca grows, pH values up to 10·35 have been recorded, and for much of the time, CO32- was the major form of inorganic carbon available. The apparent lack of an ability to use CO32- under these conditions suggests that direct use of CO32- as a source of inorganic carbon for photosynthesis is unlikely to be widespread.

Notes: Unlike behavioural responses to physical gradients, active responses to chemical gradients, and their physiological and ecological implications, have rarely been studied in freshwater phytoplanktonic flagellates. This study used microscale preference chambers to investigate the population and individual cell responses of five species exposed to four chemical gradients which commonly develop with depth in lakes: phosphate, oxygen, carbon dioxide and pH. Upon exposure of nutrient-replete and -depleted cells to a phosphate gradient, only nutrient-depleted cells of the autotrophic Chlamydomonas moewusii responded, accumulating at high concentrations. In contrast, all species responded to an oxygen gradient with Ceratium furcoides, Chlamydomonas moewusii, Dinobryon sertularia and Plagioselmis nannoplanctica preferring high concentrations whereas Euglena gracilis preferred low concentrations. In addition, all species displayed a strong affinity for carbon dioxide which was not mediated by detecting pH. Analysis of the swimming trajectories of individual cells showed that directed chemotaxes, rather than speed-dependent chemokineses, were responsible for the observed preferences. These complex and diverse species-dependent chemosensory responses may optimize photosynthesis, facilitate nutrient retrieval during migration, increase growth rate and may influence spatial and temporal distribution, contributing to the delineation of niche separation in phytoplanktonic flagellates.

Notes: The date of fry emergence over 30 years in a sea trout nursery stream, predicted by an individual-based model, correlated significantly (r=0·660, P〈0·001) with an index of the North Atlantic Oscillation. Water temperature is the main driving variable in the model and stream temperature also correlated significantly (r=0·662, P〈0·001) with the index, providing a probable causal link. Therefore, the inter-annual variations in emergence may not be unique to this one stream, but may be typical of other trout streams with similar climatic conditions.

Notes: Summary Thirty-four species of marine macroalgae from around St. Andrews, Scotland, have been assayed for their external activity and thirty-three species for their total activity of carbonic anhydrase. Activity was detected in all the Rhodophyta tested apart from Chondrus crispus, but was absent in Codium fragile, Enteromorpha sp. and Monostroma fuscum (Chlorophyta), and Alaria esculenta, Laminaria digitata, L. saccharina and L. hyperborea (Phaeophyta). Total activity of carbonic anhydrase per unit fresh weight tended to be higher in the Rhodophyta than in the Chlorophyta or Phaeophyta. External activity was present in two of the six Chlorophyta, four of the twelve Phaeophyta and four of the sixteen Rhodophyta tested. On average, when present, external carbonic anhydrase activity represented 2.7% of the total activity. A relationship was found between total carbonic anhydrase activity and habitat. Species from the high intertidal and the low-light subtidal habitats had significantly higher activity than species from the mid and low intertidal, rockpools, or high-light region of the subtidal. External carbonic anhydrase activity did not vary significantly with habitat. There appeared to be no strong relationship between carbonic anhydrase activity and the ability of a species to use HCO - 3 in photosynthesis under water.

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 -