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Palaeolimnological records provide valuable information about how phytoplankton respond to long-term drivers of environmental change. Traditional palaeolimnological tools such as microfossils and pigments are restricted to taxa that leave sub-fossil remains, and a method that can be applied to the wider community is required. Sedimentary DNA (sedDNA), extracted from lake sediment cores, shows promise in palaeolimnology, but validation against data from long-term monitoring of lake water is necessary to enable its development as a reliable record of past phytoplankton communities. To address this need, 18S rRNA gene amplicon sequencing was carried out on lake sediments from a core collected from Esthwaite Water (English Lake District) spanning ~105 years. This sedDNA record was compared with concurrent long-term microscopy-based monitoring of phytoplankton in the surface water. Broadly comparable trends were observed between the datasets, with respect to the diversity and relative abundance and occurrence of chlorophytes, dinoflagellates, ochrophytes and bacillariophytes. Up to 20% of genera were successfully captured using both methods, and sedDNA revealed a previously undetected community of phytoplankton. These results suggest that sedDNA can be used as an effective record of past phytoplankton communities, at least over timescales of <100 years. However, a substantial proportion of genera identified by microscopy were not detected using sedDNA, highlighting the current limitations of the technique that require further development such as reference database coverage. The taphonomic processes which may affect its reliability, such as the extent and rate of deposition and DNA degradation, also require further research., (© 2023 The Authors. Molecular Ecology Resources published by John Wiley & Sons Ltd.)

Species richness is a key ecological characteristic that influences numerous ecosystem functions. Here we analyse the patterns and possible causes of phytoplankton taxon richness in seasonal datasets from twenty contrasting lakes in the English Lake District over six years and near-weekly datasets over 33 years from Windermere. Taxon richness was lowest in winter and highest in summer or autumn in all of the lakes. Observed richness was very similar to richness estimated from coverage and sampling effort, implying that it closely reflected true seasonal patterns. Summer populations were dominated by Chlorophyta and functional groups X1, F, N and P (sensu Reynolds). In Windermere, weekly taxon richness was strongly positively correlated with surface water temperature, as was the number of functional groups and the number of taxa per functional group. Turnover in richness of taxa and functional groups were positively correlated and both were related to surface temperature. This suggests that high taxon richness in summer is linked to higher water temperature, promoting a turnover in richness of taxa and functional groups in these lakes. However, since the number of taxa per unit concentration of chlorophyll a decreased with increasing concentration of chlorophyll a, competition might occur when abundance is high., (Copyright © 2022 The Authors. Published by Elsevier GmbH.. All rights reserved.)

Seagrass meadows are one of the most productive ecosystems on the planet, but their photosynthesis rate may be limited by carbon dioxide but mitigated by exploiting the high concentration of bicarbonate in the ocean using different active processes. Seagrasses are declining worldwide at an accelerating rate because of numerous anthropogenic pressures. However, rising ocean concentrations of dissolved inorganic carbon, caused by increases in atmospheric carbon dioxide, may benefit seagrass photosynthesis. Here we compare the ability of two seagrass from the Mediterranean Sea, Posidonia oceanica (L.) Delile and Zostera marina L., to use carbon dioxide and bicarbonate at light saturation, and model how increasing concentrations of inorganic carbon affect their photosynthesis rate. pH-drift measurements confirmed that both species were able to use bicarbonate in addition to carbon dioxide, but that Z. marina was more effective than P. oceanica . Kinetic experiments showed that, compared to Z. marina , P. oceanica had a seven-fold higher affinity for carbon dioxide and a 1.6-fold higher affinity for bicarbonate. However, the maximal rate of bicarbonate uptake in Z. marina was 2.1-fold higher than in P. oceanica . In equilibrium with 410 ppm carbon dioxide in the atmosphere, the modelled rates of photosynthesis by Z. marina were slightly higher than P. oceanica , less carbon limited and depended on bicarbonate to a greater extent. This greater reliance by Z. marina is consistent with its less depleted 13 C content compared to P. oceanica . Modelled photosynthesis suggests that both species would depend on bicarbonate alone at an atmospheric carbon dioxide partial pressure of 280 ppm. P. oceanica was projected to benefit more than Z. marina with increasing atmospheric carbon dioxide partial pressures, and at the highest carbon dioxide scenario of 1135 ppm, would have higher rates of photosynthesis and be more saturated by inorganic carbon than Z. marina . In both species, the proportional reliance on bicarbonate declined markedly as carbon dioxide concentrations increased and in P. oceanica carbon dioxide would become the major source of inorganic carbon., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 Maberly, Stott and Gontero.)

Climate change and other anthropogenic stressors have led to long-term changes in the thermal structure, including surface temperatures, deepwater temperatures, and vertical thermal gradients, in many lakes around the world. Though many studies highlight warming of surface water temperatures in lakes worldwide, less is known about long-term trends in full vertical thermal structure and deepwater temperatures, which have been changing less consistently in both direction and magnitude. Here, we present a globally-expansive data set of summertime in-situ vertical temperature profiles from 153 lakes, with one time series beginning as early as 1894. We also compiled lake geographic, morphometric, and water quality variables that can influence vertical thermal structure through a variety of potential mechanisms in these lakes. These long-term time series of vertical temperature profiles and corresponding lake characteristics serve as valuable data to help understand changes and drivers of lake thermal structure in a time of rapid global and ecological change., (© 2021. The Author(s).)

Lake ecosystems, and the organisms that live within them, are vulnerable to temperature change 1-5 , including the increased occurrence of thermal extremes 6 . However, very little is known about lake heatwaves-periods of extreme warm lake surface water temperature-and how they may change under global warming. Here we use satellite observations and a numerical model to investigate changes in lake heatwaves for hundreds of lakes worldwide from 1901 to 2099. We show that lake heatwaves will become hotter and longer by the end of the twenty-first century. For the high-greenhouse-gas-emission scenario (Representative Concentration Pathway (RCP) 8.5), the average intensity of lake heatwaves, defined relative to the historical period (1970 to 1999), will increase from 3.7 ± 0.1 to 5.4 ± 0.8 degrees Celsius and their average duration will increase dramatically from 7.7 ± 0.4 to 95.5 ± 35.3 days. In the low-greenhouse-gas-emission RCP 2.6 scenario, heatwave intensity and duration will increase to 4.0 ± 0.2 degrees Celsius and 27.0 ± 7.6 days, respectively. Surface heatwaves are longer-lasting but less intense in deeper lakes (up to 60 metres deep) than in shallower lakes during both historic and future periods. As lakes warm during the twenty-first century 7,8 , their heatwaves will begin to extend across multiple seasons, with some lakes reaching a permanent heatwave state. Lake heatwaves are likely to exacerbate the adverse effects of long-term warming in lakes and exert widespread influence on their physical structure and chemical properties. Lake heatwaves could alter species composition by pushing aquatic species and ecosystems to the limits of their resilience. This in turn could threaten lake biodiversity 9 and the key ecological and economic benefits that lakes provide to society.

Globally, lake surface water temperatures have warmed rapidly relative to air temperatures, but changes in deepwater temperatures and vertical thermal structure are still largely unknown. We have compiled the most comprehensive data set to date of long-term (1970-2009) summertime vertical temperature profiles in lakes across the world to examine trends and drivers of whole-lake vertical thermal structure. We found significant increases in surface water temperatures across lakes at an average rate of + 0.37 °C decade -1 , comparable to changes reported previously for other lakes, and similarly consistent trends of increasing water column stability (+ 0.08 kg m -3 decade -1 ). In contrast, however, deepwater temperature trends showed little change on average (+ 0.06 °C decade -1 ), but had high variability across lakes, with trends in individual lakes ranging from - 0.68 °C decade -1 to + 0.65 °C decade -1 . The variability in deepwater temperature trends was not explained by trends in either surface water temperatures or thermal stability within lakes, and only 8.4% was explained by lake thermal region or local lake characteristics in a random forest analysis. These findings suggest that external drivers beyond our tested lake characteristics are important in explaining long-term trends in thermal structure, such as local to regional climate patterns or additional external anthropogenic influences.

The freshwater monocot Ottelia alismoides is the only known species to operate three CO2-concentrating mechanisms (CCMs): constitutive bicarbonate (HCO3-) use, C4 photosynthesis, and facultative Crassulacean acid metabolism, but the mechanism of HCO3- use is unknown. We found that the inhibitor of an anion exchange protein, 4,4'-diisothio-cyanatostilbene-2,2'-disulfonate (DIDS), prevented HCO3- use but also had a small effect on CO2 uptake. An inhibitor of external carbonic anhydrase (CA), acetazolamide (AZ), reduced the affinity for CO2 uptake but also prevented HCO3- use via an effect on the anion exchange protein. Analysis of mRNA transcripts identified a homologue of solute carrier 4 (SLC4) responsible for HCO3- transport, likely to be the target of DIDS, and a periplasmic α-carbonic anhydrase 1 (α-CA1). A model to quantify the contribution of the three different pathways involved in inorganic carbon uptake showed that passive CO2 diffusion dominates inorganic carbon uptake at high CO2 concentrations. However, as CO2 concentrations fall, two other pathways become predominant: conversion of HCO3- to CO2 at the plasmalemma by α-CA1 and transport of HCO3- across the plasmalemma by SLC4. These mechanisms allow access to a much larger proportion of the inorganic carbon pool and continued photosynthesis during periods of strong carbon depletion in productive ecosystems., (© The Author(s) 2020. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Diatoms belong to a major, diverse and species-rich eukaryotic clade, the Heterokonta, within the polyphyletic chromalveolates. They evolved as a result of secondary endosymbiosis with one or more Plantae ancestors, but their precise evolutionary history is enigmatic. Nevertheless, this has conferred them with unique structural and biochemical properties that have allowed them to flourish in a wide range of different environments and cope with highly variable conditions. We review the effect of pH, light and dark, and CO 2 concentration on the regulation of carbon uptake and assimilation. We discuss the regulation of the Calvin-Benson-Bassham cycle, glycolysis, lipid synthesis, and carbohydrate synthesis at the level of gene transcripts (transcriptomics), proteins (proteomics) and enzyme activity. In contrast to Viridiplantae where redox regulation of metabolic enzymes is important, it appears to be less common in diatoms, based on the current evidence, but regulation at the transcriptional level seems to be widespread. The role of post-translational modifications such as phosphorylation, glutathionylation, etc., and of protein-protein interactions, has been overlooked and should be investigated further. Diatoms and other chromalveolates are understudied compared to the Viridiplantae, especially given their ecological importance, but we believe that the ever-growing number of sequenced genomes combined with proteomics, metabolomics, enzyme measurements, and the application of novel techniques will provide a better understanding of how this important group of algae maintain their productivity under changing conditions., (Copyright © 2020 Launay, Huang, Maberly and Gontero.)

The concentration of dissolved organic matter (DOM) in freshwaters is increasing in large areas of the world. In addition to carbon, DOM contains nitrogen and phosphorus and there is growing concern that these organic nutrients may be bioavailable and contribute to eutrophication. However, relatively few studies have assessed the potential for dissolved organic nitrogen (DON) or dissolved organic phosphorus (DOP) compounds to be bioavailable to natural river phytoplankton communities at different locations or times. Temporal and spatial variations in uptake, relative to environmental characteristics were examined at six riverine sites in two contrasting catchments in the UK. This study also examined how the uptake by riverine phytoplankton of four DON and four DOP compounds commonly found in rivers, varied with concentration. Total nitrogen (TN) and phosphorus (TP) concentrations, the proportion of inorganic nutrient species, and nutrient limitation varied temporally and spatially, as did the potential for DON and DOP uptake. All eight of the DOM compounds tested were bioavailable, but to different extents. Organic nutrient use depended on the concentration of the organic compound supplied, with simple compounds (urea and glucose-6-phosphate) supporting algal growth even at very low concentrations. DON use was negatively correlated with the TN and ammonia concentration and DOP use was negatively correlated with soluble reactive phosphorus (SRP) and dissolved organic carbon (DOC) concentration. The evidence indicates that DOM in rivers has been overlooked as a potential source of nutrients to phytoplankton and therefore as an agent of eutrophication., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2020 The Authors. Published by Elsevier B.V. All rights reserved.)

Background and Aims: Ottelia alismoides (Hydrocharitaceae) is a freshwater macrophyte that, unusually, possesses three different CO2-concentrating mechanisms. Here we describe its leaf anatomy and chloroplast ultrastructure, how these are altered by CO2 concentration and how they may underlie C4 photosynthesis., Methods: Light and transmission electron microscopy were used to study the anatomy of mature leaves of O. alismoides grown at high and low CO2 concentrations. Diel acid change and the activity of phosphoenolpyruvate carboxylase were measured to confirm that CAM activity and C4 photosynthesis were present., Key Results: When O. alismoides was grown at low CO2, the leaves performed both C4 and CAM photosynthesis whereas at high CO2 leaves used C4 photosynthesis. The leaf comprised an upper and lower layer of epidermal cells separated by a large air space occupying about 22 % of the leaf transverse-section area, and by mesophyll cells connecting the two epidermal layers. Kranz anatomy was absent. At low CO2, chloroplasts in the mesophyll cells were filled with starch even at the start of the photoperiod, while epidermal chloroplasts contained small starch grains. The number of chloroplasts in the epidermis was greater than in the mesophyll cells. At high CO2, the structure was unchanged but the thicknesses of the two epidermal layers, the air space, mesophyll and the transverse-section area of cells and air space were greater., Conclusions: Leaves of O. alismoides have epidermal and mesophyll cells that contain chloroplasts and large air spaces but lack Kranz anatomy. The high starch content of mesophyll cells suggests they may benefit from an internal source of CO2, for example via C4 metabolism, and are also sites of starch storage. The air spaces may help in the recycling of decarboxylated or respired CO2. The structural similarity of leaves at low and high CO2 is consistent with the constitutive nature of bicarbonate and C4 photosynthesis. There is sufficient structural diversity within the leaf of O. alismoides to support dual-cell C4 photosynthesis even though Kranz anatomy is absent., (© The Author(s) 2020. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Carbonic anhydrases (CAs) exist in all kingdoms of life. They are metalloenzymes, often containing zinc, that catalyze the interconversion of bicarbonate and carbon dioxide-a ubiquitous reaction involved in a variety of cellular processes. So far, eight classes of apparently evolutionary unrelated CAs that are present in a large diversity of living organisms have been described. In this review, we focus on the diversity of CAs and their roles in photosynthetic microalgae. We describe their essential role in carbon dioxide-concentrating mechanisms and photosynthesis, their regulation, as well as their less studied roles in non-photosynthetic processes. We also discuss the presence in some microalgae, especially diatoms, of cambialistic CAs (i.e., CAs that can replace Zn by Co, Cd, or Fe) and, more recently, a CA that uses Mn as a metal cofactor, with potential ecological relevance in aquatic environments where trace metal concentrations are low. There has been a recent explosion of knowledge about this well-known enzyme with exciting future opportunities to answer outstanding questions using a range of different approaches.

The effect of dams on dissolved inorganic carbon (DIC) transport and riverine ecosystems is unclear in karst cascade reservoirs. Here, we analyzed water samples from a karst river system with seven cascade reservoirs along the Wujiang River, southwestern China, during one hydrological year. From upstream to downstream, the average concentration of DIC increased from 2.2 to 2.6 mmol/L and its carbon isotope composition (δ 13 C DIC ) decreased from -8.0 to -10‰. Meanwhile, the air temperature (Ta) increased from 20.3 °C to 26.7 °C and 10 °C to 13.7 °C in the warm and cold seasons, respectively. The results suggest that a cascade of dams has a stronger effect on DIC dynamics and retention than a single dam. The good correlation between Ta/HRT (hydraulic retention time) and Δ[DIC] as well as Δ[δ 13 C DIC ] mean that Ta and HRT affected the magnitude of the damming effect by altering changes in concentration of DIC and δ 13 C DIC in the reservoir compared to the inflowing water. In particular, daily regulated reservoirs with short retention times acted more like river corridors and had a smaller effect on carbon dynamics, so modulating retention time might be used reduce the effect of dams on the riverine ecosystem., (Copyright © 2019 Elsevier B.V. All rights reserved.)

Water temperature is critical for the ecology of lakes. However, the ability to predict its spatial and seasonal variation is constrained by the lack of a thermal classification system. Here we define lake thermal regions using objective analysis of seasonal surface temperature dynamics from satellite observations. Nine lake thermal regions are identified that mapped robustly and largely contiguously globally, even for small lakes. The regions differed from other global patterns, and so provide unique information. Using a lake model forced by 21 st century climate projections, we found that 12%, 27% and 66% of lakes will change to a lower latitude thermal region by 2080-2099 for low, medium and high greenhouse gas concentration trajectories (Representative Concentration Pathways 2.6, 6.0 and 8.5) respectively. Under the worst-case scenario, a 79% reduction in the number of lakes in the northernmost thermal region is projected. This thermal region framework can facilitate the global scaling of lake-research.

Hydropower development is the key strategy in many developing countries for energy supply, climate-change mitigation and economic development. However, it is commonly assumed that river dams retain nutrients and therefore reduce downstream primary productivity and fishery catches, compromising food security and causing trans-boundary disputes. Contrary to expectation, here we found that a cascade of reservoirs along the upper Mekong River increased downstream bioavailability of nitrogen and phosphorus. The dams caused phytoplankton density to increase with hydraulic residence time and stratification of the stagnant reservoirs caused hypoxia at depth. This allowed the release of bioavailable phosphorus from the sediment and an increase in dissolved inorganic nitrogen as well as a shift in nitrogen species from nitrate to ammonium, which were transported downstream by the discharge of water from the base of the dam. Our findings provide a new perspective on the environmental impacts of river dams on nutrient cycling and ecosystem functioning, with potential implications for sustainable development of hydropower worldwide., (© The Author(s) 2020. Published by Oxford University Press on behalf of China Science Publishing & Media Ltd.)

The role of sediment-bound organic phosphorus (P o ) as an additional nutrient source is a component of internal P budgets in lake system that is usually neglected. Here we examined the relative importance of sediment P o to internal P load and the role of bioavailable P o in algal growth in Lake Erhai, China. Lake Erhai sediment extractable P o accounted for 11-43% (27% average) of extractable total P, and bioavailable P o accounted for 21-66% (40%) of P o . The massive storage of bioavailable P o represents an important form of available P, essential to internal loads. The bioavailable P o includes mainnly labile monoester P and diester P was identified in the sequential extractions by H 2 O, NaHCO 3 , NaOH, and HCl. 40% of H 2 O-P o , 39% of NaHCO 3 -P o , 43% of NaOH-P o , and 56% of HCl-P o can be hydrolyzed to labile monoester and diester P, suggesting that the bioavailability of P o fractions was in decreasing order as follows: HCl-P o  > NaOH-P o  > H 2 O-P o  > NaHCO 3 -P o . It is implied that traditional sequential fractionation of P o might overestimate the availability of labile P o in sediments. Furthermore, analysis of the environmental processes of bioavailable P o showed that the stabler structure of dissloved organic matter (DOM) alleviated the degradation and release of diester P, abundant alkaline phosphatase due to higher algal biomass promoted the degradation of diester P. The stability of DOM structure and the degradation of diester P might responsible for the spatial differences of labile monoester P. The biogeochemical cycle of bioavailable P o replenishs available P pools in overlying water and further facilitate algal growth during the algal blooms. Therefore, to control the algal blooms in Lake Erhai, an effective action is urgently required to reduce the accumulation of P o in sediments and interrupt the supply cycle of bioavailable P o to algal growth., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

Unlike in land plants, photosynthesis in many aquatic plants relies on bicarbonate in addition to carbon dioxide (CO 2 ) to compensate for the low diffusivity and potential depletion of CO 2 in water. Concentrations of bicarbonate and CO 2 vary greatly with catchment geology. In this study, we investigate whether there is a link between these concentrations and the frequency of freshwater plants possessing the bicarbonate use trait. We show, globally, that the frequency of plant species with this trait increases with bicarbonate concentration. Regionally, however, the frequency of bicarbonate use is reduced at sites where the CO 2 concentration is substantially above the air equilibrium, consistent with this trait being an adaptation to carbon limitation. Future anthropogenic changes of bicarbonate and CO 2 concentrations may alter the species compositions of freshwater plant communities., (Copyright © 2019 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works.)

Cyanobacterial blooms are an increasing threat to water quality and global water security caused by the nutrient enrichment of freshwaters. There is also a broad consensus that blooms are increasing with global warming, but the impacts of other concomitant environmental changes, such as an increase in extreme rainfall events, may affect this response. One of the potential effects of high rainfall events on phytoplankton communities is greater loss of biomass through hydraulic flushing. Here we used a shallow lake mesocosm experiment to test the combined effects of: warming (ambient vs. +4°C increase), high rainfall (flushing) events (no events vs. seasonal events) and nutrient loading (eutrophic vs. hypertrophic) on total phytoplankton chlorophyll-a and cyanobacterial abundance and composition. Our hypotheses were that: (a) total phytoplankton and cyanobacterial abundance would be higher in heated mesocosms; (b) the stimulatory effects of warming on cyanobacterial abundance would be enhanced in higher nutrient mesocosms, resulting in a synergistic interaction; (c) the recovery of biomass from flushing induced losses would be quicker in heated and nutrient-enriched treatments, and during the growing season. The results supported the first and, in part, the third hypotheses: total phytoplankton and cyanobacterial abundance increased in heated mesocosms with an increase in common bloom-forming taxa-Microcystis spp. and Dolichospermum spp. Recovery from flushing was slowest in the winter, but unaffected by warming or higher nutrient loading. Contrary to the second hypothesis, an antagonistic interaction between warming and nutrient enrichment was detected for both cyanobacteria and chlorophyll-a demonstrating that ecological surprises can occur, dependent on the environmental context. While this study highlights the clear need to mitigate against global warming, oversimplification of global change effects on cyanobacteria should be avoided; stressor gradients and seasonal effects should be considered as important factors shaping the response., (© 2019 The Authors. Global Change Biology Published by John Wiley & Sons Ltd.)

An increase of dissolved organic carbon (DOC) in inland waters has been reported across the northern temperate region but the effects of this on whole lake ecosystems, often combined with other anthropogenic stressors like nutrient inputs and warming, are poorly known. The effects of these changes on different component of the ecosystem were assessed in an experiment using twenty-four large (3000L) outdoor mesocosms simulating shallow lakes. Two different temperature regimes (ambient and ambient +4 °C) combined with three levels of organic matter (OM, added as filtered peaty water), simulating the DOC increase that is predicted to take place over the next 4 to 21 years were used. Neither temperature nor OM had significant effects on net ecosystem production, respiration or gross primary production. Phytoplankton chlorophyll a concentration was not significantly affected by warming, however in summer, autumn and winter it was significantly higher in mesocosms receiving intermediate OM levels (July-Feb DOC concentrations 2-6 mg L -1 ). Summer cyanobacterial blooms were highest in intermediate, and lowest in the highest OM treatments. OM concentration also influenced total macroinvertebrate abundance which was greater in spring and summer in mesocosms with intermediate and high OM. Fish abundance was not significantly affected by OM concentration, but abundance was greater in ambient (55 fish subsample -1 ) compared to heated mesocosms (17 fish subsample -1 ) and maximum abundance occurred two weeks later compared to heated mesocosms. The results suggest that changes in OM may have a greater effect on shallow lakes than temperature and that phytoplankton, especially cyanobacteria, benefit from intermediate OM concentrations, therefore, nuisance algal blooms might increase in relatively clear shallow eutrophic lakes where DOC concentrations increase., (Copyright © 2019 Elsevier B.V. All rights reserved.)

Most aquatic photoautotrophs depend on CO 2 -concentrating mechanisms (CCMs) to maintain productivity at ambient concentrations of CO 2 , and carbonic anhydrase (CA) plays a key role in these processes. Here we present different lines of evidence showing that the protein LCIP63, identified in the marine diatom Thalassiosira pseudonana, is a CA. However, sequence analysis showed that it has a low identity with any known CA and therefore belongs to a new subclass that we designate as iota-CA. Moreover, LCIP63 unusually prefers Mn 2+ to Zn 2+ as a cofactor, which is potentially of ecological relevance since Mn 2+ is more abundant than Zn 2+ in the ocean. LCIP63 is located in the chloroplast and only expressed at low concentrations of CO 2 . When overexpressed using biolistic transformation, the rate of photosynthesis at limiting concentrations of dissolved inorganic carbon increased, confirming its role in the CCM. LCIP63 homologs are present in the five other sequenced diatoms and in other algae, bacteria, and archaea. Thus LCIP63 is phylogenetically widespread but overlooked. Analysis of the Tara Oceans database confirmed this and showed that LCIP63 is widely distributed in marine environments and is therefore likely to play an important role in global biogeochemical carbon cycling.

Blooms of cyanobacteria are a current threat to global water security that is expected to increase in the future because of increasing nutrient enrichment, increasing temperature and extreme precipitation in combination with prolonged drought. However, the responses to multiple stressors, such as those above, are often complex and there is contradictory evidence as to how they may interact. Here we used broad scale data from 494 lakes in central and northern Europe, to assess how cyanobacteria respond to nutrients (phosphorus), temperature and water retention time in different types of lakes. Eight lake types were examined based on factorial combinations of major factors that determine phytoplankton composition and sensitivity to nutrients: alkalinity (low and medium-high), colour (clear and humic) and mixing intensity (polymictic and stratified). In line with expectations, cyanobacteria increased with temperature and retention time in five of the eight lake types. Temperature effects were greatest in lake types situated at higher latitudes, suggesting that lakes currently not at risk could be affected by warming in the future. However, the sensitivity of cyanobacteria to temperature, retention time and phosphorus varied among lake types highlighting the complex responses of lakes to multiple stressors. For example, in polymictic, medium-high alkalinity, humic lakes cyanobacteria biovolume was positively explained by retention time and a synergy between TP and temperature, while in polymictic, medium-high alkalinity, clear lakes only retention time was identified as an explanatory variable. These results show that, although climate change will need to be accounted for when managing the risk of cyanobacteria in lakes, a "one-size fits-all" approach is not appropriate. When forecasting the response of cyanobacteria to future environmental change, including changes caused by climate and local management, it will be important to take this differential sensitivity of lakes into account., (© 2018 John Wiley & Sons Ltd.)

The freshwater macrophyte, Ottelia alismoides, is a bicarbonate user performing C4 photosynthesis in the light, and crassulacean acid metabolism (CAM) when acclimated to low CO 2 . The regulation of the three mechanisms by CO 2 concentration was studied in juvenile and mature leaves. For mature leaves, the ratios of phosphoenolpyruvate carboxylase (PEPC) to ribulose-bisphosphate carboxylase/oxygenase (Rubisco) are in the range of that of C4 plants regardless of CO 2 concentration (1.5-2.5 at low CO 2 , 1.8-3.4 at high CO 2 ). In contrast, results for juvenile leaves suggest that C4 is facultative and only present under low CO 2 . pH-drift experiments showed that both juvenile and mature leaves can use bicarbonate irrespective of CO 2 concentration, but mature leaves have a significantly greater carbon-extracting ability than juvenile leaves at low CO 2 . At high CO 2 , neither juvenile nor mature leaves perform CAM as indicated by lack of diurnal acid fluctuation. However, CAM was present at low CO 2 , though the fluctuation of titratable acidity in juvenile leaves (15-17 µequiv g -1 FW) was slightly but significantly lower than in mature leaves (19-25 µequiv g -1 FW), implying that the capacity to perform CAM increases as leaves mature. The increased CAM activity is associated with elevated PEPC activity and large diel changes in starch content. These results show that in O. alismoides, carbon-dioxide concentrating mechanisms are more effective in mature compared to juvenile leaves, and C4 is facultative in juvenile leaves but constitutive in mature leaves.

The global proliferation of harmful algal blooms poses an increasing threat to water resources, recreation and ecosystems. Predicting the occurrence of these blooms is therefore needed to assist water managers in making management decisions to mitigate their impact. Evaluation of the potential for forecasting of algal blooms using the phytoplankton community model PROTECH was undertaken in pseudo-real-time. This was achieved within a data assimilation scheme using the Ensemble Kalman Filter to allow uncertainties and model nonlinearities to be propagated to forecast outputs. Tests were made on two mesotrophic lakes in the English Lake District, which differ in depth and nutrient regime. Some forecasting success was shown for chlorophyll a, but not all forecasts were able to perform better than a persistence forecast. There was a general reduction in forecast skill with increasing forecasting period but forecasts for up to four or five days showed noticeably greater promise than those for longer periods. Associated forecasts of phytoplankton community structure were broadly consistent with observations but their translation to cyanobacteria forecasts was challenging owing to the interchangeability of simulated functional species., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Extreme weather can have a substantial influence on lakes and is expected to become more frequent with climate change. We explored the influence of one particular extreme event, Storm Ophelia, on the physical and chemical environment of England's largest lake, Windermere. We found that the substantial influence of Ophelia on meteorological conditions at Windermere, in particular wind speed, resulted in a 25-fold increase (relative to the study-period average) in the wind energy flux at the lake-air interface. Following Ophelia, there was a short-lived mixing event in which the Schmidt stability decreased by over 100 Jm -2 and the thermocline deepened by over 10 m during a 12-h period. As a result of changes to the strength of stratification, Ophelia also changed the internal seiche regime of Windermere with the dominant seiche period increasing from ~ 17 h pre-storm to ~ 21 h post-storm. Following Ophelia, there was an upwelling of cold and low-oxygenated waters at the southern-end of the lake. This had a substantial influence on the main outflow of Windermere, the River Leven, where dissolved oxygen concentrations decreased by ~ 48%, from 9.3 to 4.8 mg L -1 , while at the mid-lake monitoring station in Windermere, it decreased by only ~ 3%. This study illustrates that the response of a lake to extreme weather can cause important effects downstream, the influence of which may not be evident at the lake surface. To understand the impact of future extreme events fully, the whole lake and downstream-river system need to be studied together.

In Plantae, the Calvin-Benson-Bassham (CBB) cycle is highly regulated and most of its enzymes have been thoroughly studied. Since diatoms arose as a result of secondary endosymbiosis with one or more Plantae ancestors, their precise evolutionary history is enigmatic and complex resulting in biochemical variations on the original CBB cycle theme. The Rubisco Michaelis constant for CO 2 is higher in diatoms than land plants and the nuclear-encoded Rubisco activase in Plantae is replaced by an analogous chloroplast-encoded CbbX (Calvin-Benson-Bassham protein X) in diatoms. In the CBB cycle reduction phase, phosphoglycerate kinase in diatoms is redox-regulated and similar to that in red algae; however, glyceraldehyde phosphate dehydrogenase (GAPDH) is not redox-regulated, unlike in Plantae. The phosphoribulokinase (PRK)-GAPDH-CP12 complex found in many photosynthetic organisms has not yet been found in diatoms, but a ferredoxin-NADP reductase (FNR)-GAPDH-CP12 complex has been found in one species. In the CBB cycle regeneration phase, sedoheptulose 1,7-bisphosphatase and PRK are not redox-regulated in diatoms, unlike in Plantae. Regulation at the transcriptional level seems to be important in diatoms. CBB cycle enzyme properties appear to be variable among diatoms, but this view relies on results from a few model species: a greater range of diatoms need to be studied to test this.This article is part of the themed issue 'The peculiar carbon metabolism in diatoms'., (© 2017 The Author(s).)

The presence of CO2-concentrating mechanisms (CCMs) is believed to be one of the characteristics that allows diatoms to thrive in many environments and to be major contributors to global productivity. Here, the type of CCM and the responses to variable CO2 concentration were studied in marine and freshwater diatoms. At 400 ppm, there was a large diversity in physiological and biochemical mechanisms among the species. While Phaeodactylum tricornutum mainly used HCO3-, Thalassiosira pseudonana mainly used CO2. Carbonic anhydrase was an important component of the CCM in all species and C4 metabolism was absent, even with T. weissflogii. For all species, at 20 000 ppm, the affinity for dissolved inorganic carbon was lower than at 400 ppm CO2 and the reliance on CO2 was higher. Despite the difference in availability of inorganic carbon in marine and fresh waters, there were only small differences in CCMs between species from the two environments, and Navicula pelliculosa behaved similarly when grown in the two environments. The results suggest that species-specific differences are great, and more important than environmental differences in determining the nature and effectiveness of the CCM in diatoms., (© The Author 2017. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Ottelia alismoides is a constitutive C4 plant and bicarbonate user, and has facultative crassulacean acid metabolism (CAM) at low CO2. Acclimation to a factorial combination of light and CO2 showed that the ratio of phosphoenolpyruvate carboxylase (PEPC) to ribulose-bisphosphate carboxylase/oxygenase (Rubisco) (>5) is in the range of that of C4 plants. This and short-term response experiments showed that the activity of PEPC and pyruvate phosphate dikinase (PPDK) was high even at the end of the night, consistent with night-time acid accumulation and daytime carbon fixation. The diel acidity change was maximal at high light and low CO2 at 17-25 µequiv g-1 FW. Decarboxylation proceeded at ~2-3 µequiv g-1 FW h-1, starting at the beginning of the photoperiod, but did not occur at high CO2; the rate was greater at high, compared with low light. There was an inverse relationship between starch formation and acidity loss. Acidity changes account for up to 21% of starch production and stimulate early morning photosynthesis, but night-time accumulation of acid traps <6% of respiratory carbon release. Ottelia alismoides is the only known species to operate CAM and C4 in the same tissue, and one of only two known aquatic species to operate CAM and bicarbonate use., (© The Author 2017. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

In aquatic environments, the concentration of inorganic carbon is spatially and temporally variable and CO2 can be substantially oversaturated or depleted. Depletion of CO2 plus low rates of diffusion cause inorganic carbon to be more limiting in aquatic than terrestrial environments, and the frequency of species with a CO2-concentrating mechanism (CCM), and their contribution to productivity, is correspondingly greater. Aquatic photoautotrophs may have biochemical or biophysical CCMs and exploit CO2 from the sediment or the atmosphere. Though partly constrained by phylogeny, CCM activity is related to environmental conditions. CCMs are absent or down-regulated when their increased energy costs, lower CO2 affinity, or altered mineral requirements outweigh their benefits. Aquatic CCMs are most widespread in environments with low CO2, high HCO3-, high pH, and high light. Freshwater species are generally less effective at inorganic carbon removal than marine species, but have a greater range of ability to remove carbon, matching the environmental variability in carbon availability. The diversity of CCMs in seagrasses and marine phytoplankton, and detailed mechanistic studies on larger aquatic photoautotrophs are understudied. Strengthening the links between ecology and CCMs will increase our understanding of the mechanisms underlying ecological success and will place mechanistic studies in a clearer ecological context., (© The Author 2017. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

The concentration of CO 2 in many aquatic systems is variable, often lower than the K M of the primary carboxylating enzyme Rubisco, and in order to photosynthesize efficiently, many algae operate a facultative CO 2 concentrating mechanism (CCM). Here we measured the responses of a marine diatom, Thalassiosira pseudonana, to high and low concentrations of CO 2 at the level of transcripts, proteins and enzyme activity. Low CO 2 caused many metabolic pathways to be remodeled. Carbon acquisition enzymes, primarily carbonic anhydrase, stress, degradation and signaling proteins were more abundant while proteins associated with nitrogen metabolism, energy production and chaperones were less abundant. A protein with similarities to the Ca 2+ / calmodulin dependent protein kinase II&#95;association domain, having a chloroplast targeting sequence, was only present at low CO 2 . This protein might be a specific response to CO 2 limitation since a previous study showed that other stresses caused its reduction. The protein sequence was found in other marine diatoms and may play an important role in their response to low CO 2 concentration., Competing Interests: The authors declare no competing financial interests.

The effect of CO 2 supply is likely to play an important role in algal ecology. Since inorganic carbon (C i ) acquisition strategies are very diverse among microalgae and C i availability varies greatly within and among habitats, we hypothesized that C i acquisition depends on the pH of their preferred natural environment (adaptation) and that the efficiency of C i uptake is affected by CO 2 availability (acclimation). To test this, four species of green algae originating from different habitats were studied. The pH-drift and C i uptake kinetic experiments were used to characterize C i acquisition strategies and their ability to acclimate to high and low CO 2 conditions and high and low pH was evaluated. Results from pH drift experiments revealed that the acidophile and acidotolerant Chlamydomonas species were mainly restricted to CO 2 , whereas the two neutrophiles were efficient bicarbonate users. CO 2 compensation points in low CO 2 -acclimated cultures ranged between 0.6 and 1.4 μM CO 2 and acclimation to different culture pH and CO 2 conditions suggested that CO 2 concentrating mechanisms were present in most species. High CO 2 acclimated cultures adapted rapidly to low CO 2 condition during pH-drifts. C i uptake kinetics at different pH values showed that the affinity for C i was largely influenced by external pH, being highest under conditions where CO 2 dominated the C i pool. In conclusion, C i acquisition was highly variable among four species of green algae and linked to growth pH preference, suggesting that there is a connection between C i acquisition and ecological distribution., (© 2016 Phycological Society of America.)

Carbon (C) and silicon (Si) biogeochemical cycles are important factors in the regulation of atmospheric CO 2 concentrations and hence climate change. Theoretically, these elements are linked by chemical weathering and organism stoichiometry, but this coupling has not been investigated in freshwaters. Here we compiled data from global rivers and lakes in the United States of America and the United Kingdom, in order to characterize the stoichiometry between the biogeochemical cycles of C and Si. In rivers this coupling is confirmed by a significant relationship between HCO 3 - /Na + and DSi/Na + , and DSi:HCO 3 - ratio can reflect the mineral source of chemical weathering. In lakes, however, these characteristic ratios of chemical weathering are altered by algal activity. The lacustrine Si:C atomic ratio is negative feedback regulation by phytoplankton, which may result in this ratio in algal assemblages similar to that in water column. And this regulation suggests lacustrine photosynthetic C fixation in this equilibrium state is quantitative and depends on the DSi concentration. These findings provide new insights into the role of freshwaters in global C and Si biogeochemical cycles.

The release of phosphorus (P) from bed sediments to the overlying water can delay the recovery of lakes for decades following reductions in catchment contributions, preventing water quality targets being met within timeframes set out by environmental legislation (e.g. EU Water Framework Directive: WFD). Therefore supplementary solutions for restoring lakes have been explored, including the capping of sediment P sources using a lanthanum (La)-modified bentonite clay to reduce internal P loading and enhance the recovery process. Here we present results from Loch Flemington where the first long-term field trial documenting responses of phytoplankton community structure and abundance, and the UK WFD phytoplankton metric to a La-bentonite application was performed. A Before-After-Control-Impact (BACI) analysis was used to distinguish natural variability from treatment effect and confirmed significant reductions in the magnitude of summer cyanobacterial blooms in Loch Flemington, relative to the control site, following La-bentonite application. However this initial cyanobacterial response was not sustained beyond two years after application, which implied that the reduction in internal P loading was short-lived; several possible explanations for this are discussed. One reason is that this ecological quality indicator is sensitive to inter-annual variability in weather patterns, particularly summer rainfall and water temperature. Over the monitoring period, the phytoplankton community structure of Loch Flemington became less dominated by cyanobacteria and more functionally diverse. This resulted in continual improvements in the phytoplankton compositional and abundance metrics, which were not observed at the control site, and may suggest an ecological response to the sustained reduction in filterable reactive phosphorus (FRP) concentration following La-bentonite application. Overall, phytoplankton classification indicated that the lake moved from poor to moderate ecological status but did not reach the proxy water quality target (i.e. WFD Good Ecological Status) within four years of the application. As for many other shallow lakes, the effective control of internal P loading in Loch Flemington will require further implementation of both in-lake and catchment-based measures. Our work emphasizes the need for appropriate experimental design and long-term monitoring programmes, to ascertain the efficacy of intervention measures in delivering environmental improvements at the field scale., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

Ecological and biogeochemical processes in lakes are strongly dependent upon water temperature. Long-term surface warming of many lakes is unequivocal, but little is known about the comparative magnitude of temperature variation at diel timescales, due to a lack of appropriately resolved data. Here we quantify the pattern and magnitude of diel temperature variability of surface waters using high-frequency data from 100 lakes. We show that the near-surface diel temperature range can be substantial in summer relative to long-term change and, for lakes smaller than 3 km2, increases sharply and predictably with decreasing lake area. Most small lakes included in this study experience average summer diel ranges in their near-surface temperatures of between 4 and 7°C. Large diel temperature fluctuations in the majority of lakes undoubtedly influence their structure, function and role in biogeochemical cycles, but the full implications remain largely unexplored.

Diatoms are widespread in aquatic ecosystems where they may be limited by the supply of inorganic carbon. Their carbon dioxide-concentrating mechanisms (CCMs) involving transporters and carbonic anhydrases (CAs) are well known, but the contribution of a biochemical CCM involving C4 metabolism is contentious. The CCM(s) present in the marine-centric diatom, Thalassiosira pseudonana, were studied in cells exposed to high or low concentrations of CO2 , using a range of approaches. At low CO2 , cells possessed a CCM based on active uptake of CO2 (70% contribution) and bicarbonate, while at high CO2 , cells were restricted to CO2 . CA was highly and rapidly activated on transfer to low CO2 and played a key role because inhibition of external CA produced uptake kinetics similar to cells grown at high CO2 . The activities of phosphoenolpyruvate (PEP) carboxylase (PEPC) and the PEP-regenerating enzyme, pyruvate phosphate dikinase (PPDK), were lower in cells grown at low than at high CO2 . The ratios of PEPC and PPDK to ribulose bisphosphate carboxylase were substantially lower than 1, even at low CO2 . Our data suggest that the kinetic properties of this species results from a biophysical CCM and not from C4 type metabolism., (© 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.)

Conventional modes of environmental governance, which typically exclude those stakeholders that are most directly linked to the specific place, frequently fail to have the desired impact. Using the example of lake water management in Loweswater, a small hamlet within the English Lake District, we consider the ways in which new "collectives" for local, bottom-up governance of water bodies can reframe problems in ways which both bind lay and professional people to place, and also recast the meaning of "solutions" in thought-provoking ways.

The productivity and ecological distribution of freshwater plants can be controlled by the availability of inorganic carbon in water despite the existence of different mechanisms to ameliorate this, such as the ability to use bicarbonate. Here we took advantage of a short, natural gradient of CO2 concentration, against a background of very high and relatively constant concentration of bicarbonate, in a spring-fed river, to study the effect of variable concentration of CO2 on the ability of freshwater plants to use bicarbonate. Plants close to the source, where the concentration of CO2 was up to 24 times air equilibrium, were dominated by Berula erecta. pH-drift results and discrimination against (13)C were consistent with this and the other species being restricted to CO2 and unable to use the high concentration of bicarbonate. There was some indication from stable (13)C data that B. erecta may have had access to atmospheric CO2 at low water levels. In contrast, species downstream, where concentrations of CO2 were only about 5 times air-equilibrium were almost exclusively able to use bicarbonate, based on pH-drift results. Discrimination against (13)C was also consistent with bicarbonate being the main source of inorganic carbon for photosynthesis in these species. There was, therefore, a transect downstream from the source of increasing ability to use bicarbonate that closely matched the decreasing concentration of CO2. This was produced largely by altered species composition, but partly by phenotypic changes in individual species., (Crown Copyright © 2014. Published by Elsevier GmbH. All rights reserved.)

Climate-induced shifts in the timing of life-history events are a worldwide phenomenon, and these shifts can de-synchronize species interactions such as predator-prey relationships. In order to understand the ecological implications of altered seasonality, we need to consider how shifts in phenology interact with other agents of environmental change such as exploitation and disease spread, which commonly act to erode the demographic structure of wild populations. Using long-term observational data on the phenology and dynamics of a model predator-prey system (fish and zooplankton in Windermere, UK), we show that age-size truncation of the predator population alters the consequences of phenological mismatch for offspring survival and population abundance. Specifically, age-size truncation reduces intraspecific density regulation due to competition and cannibalism, and thereby amplifies the population sensitivity to climate-induced predator-prey asynchrony, which increases variability in predator abundance. High population variability poses major ecological and economic challenges as it can diminish sustainable harvest rates and increase the risk of population collapse. Our results stress the importance of maintaining within-population age-size diversity in order to buffer populations against phenological asynchrony, and highlight the need to consider interactive effects of environmental impacts if we are to understand and project complex ecological outcomes.

Two freshwater macrophytes, Ottelia alismoides and O. acuminata, were grown at low (mean 5 μmol L(-1)) and high (mean 400 μmol L(-1)) CO2 concentrations under natural conditions. The ratio of PEPC to RuBisCO activity was 1.8 in O. acuminata in both treatments. In O. alismoides, this ratio was 2.8 and 5.9 when grown at high and low CO2, respectively, as a result of a twofold increase in PEPC activity. The activity of PPDK was similar to, and changed with, PEPC (1.9-fold change). The activity of the decarboxylating NADP-malic enzyme (ME) was very low in both species, while NAD-ME activity was high and increased with PEPC activity in O. alismoides. These results suggest that O. alismoides might perform a type of C4 metabolism with NAD-ME decarboxylation, despite lacking Kranz anatomy. The C4-activity was still present at high CO2 suggesting that it could be constitutive. O. alismoides at low CO2 showed diel acidity variation of up to 34 μequiv g(-1) FW indicating that it may also operate a form of crassulacean acid metabolism (CAM). pH-drift experiments showed that both species were able to use bicarbonate. In O. acuminata, the kinetics of carbon uptake were altered by CO2 growth conditions, unlike in O. alismoides. Thus, the two species appear to regulate their carbon concentrating mechanisms differently in response to changing CO2. O. alismoides is potentially using three different concentrating mechanisms. The Hydrocharitaceae have many species with evidence for C4, CAM or some other metabolism involving organic acids, and are worthy of further study.

Diatoms are a widespread and ecologically important group of heterokont algae that contribute c. 20% to global productivity. Previous work has shown that regulation of their key Calvin cycle enzymes differs from that of the Plantae, and that in crude extracts, glyceraldehyde-3-phosphate dehydrogenase (GAPDH) can be inhibited by nicotinamide adenine dinucleotide phosphate reduced (NADPH) under oxidizing conditions. The freshwater diatom, Asterionella formosa, was studied using enzyme kinetics, chromatography, surface plasmon resonance, mass spectrometry and sequence analysis to determine the mechanism behind this GAPDH inhibition. GAPDH interacted with ferredoxin-nicotinamide adenine dinucleotide phosphate (NADP) reductase (FNR) from the primary phase of photosynthesis, and the small chloroplast protein, CP12. Sequences of copurified GAPDH and FNR were highly homologous with published sequences. However, the widespread ternary complex among GAPDH, phosphoribulokinase and CP12 was absent. Activity measurements under oxidizing conditions showed that NADPH can inhibit GAPDH-CP12 in the presence of FNR, explaining the earlier observed inhibition within crude extracts. Diatom plastids have a distinctive metabolism, including the lack of the oxidative pentose phosphate pathway, and so cannot produce NADPH in the dark. The observed down-regulation of GAPDH in the dark may allow NADPH to be rerouted towards other reductive processes contributing to their ecological success., (© 2014 The Authors. New Phytologist © 2014 New Phytologist Trust.)

The WHAM-FTOX model quantifies the combined toxic effects of protons and metal cations towards aquatic organisms through the toxicity function (FTOX), a linear combination of the products of organism-bound cation and a toxic potency coefficient for each cation. We describe the application of the model to predict an observable ecological field variable, species richness of pelagic lake crustacean zooplankton, studied with respect to either acidification or the impacts of metals from smelters. The fitted results give toxic potencies increasing in the order H(+) < Al < Cu < Zn < Ni. In general, observed species richness is lower than predicted, but in some instances agreement is close, and is rarely higher than predictions. The model predicts recovery in agreement with observations for three regions, namely Sudbury (Canada), Bohemian Forest (Czech Republic) and a subset of lakes across Norway, but fails to predict observed recovery from acidification in Adirondack lakes (USA)., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

Phenological changes have been observed globally for marine, freshwater and terrestrial species, and are an important element of the global biological 'fingerprint' of climate change. Differences in rates of change could desynchronize seasonal species interactions within a food web, threatening ecosystem functioning. Quantification of this risk is hampered by the rarity of long-term data for multiple interacting species from the same ecosystem and by the diversity of possible phenological metrics, which vary in their ecological relevance to food web interactions. We compare phenological change for phytoplankton (chlorophyll a), zooplankton (Daphnia) and fish (perch, Perca fluviatilis) in two basins of Windermere over 40 years and determine whether change has differed among trophic levels, while explicitly accounting for among-metric differences in rates of change. Though rates of change differed markedly among the nine metrics used, seasonal events shifted earlier for all metrics and trophic levels: zooplankton advanced most, and fish least, rapidly. Evidence of altered synchrony was found in both lake basins, when combining information from all phenological metrics. However, comparisons based on single metrics did not consistently detect this signal. A multimetric approach showed that across trophic levels, earlier phenological events have been associated with increasing water temperature. However, for phytoplankton and zooplankton, phenological change was also associated with changes in resource availability. Lower silicate, and higher phosphorus, concentrations were associated with earlier phytoplankton growth, and earlier phytoplankton growth was associated with earlier zooplankton growth. The developing trophic mismatch detected between the dominant fish species in Windermere and important zooplankton food resources may ultimately affect fish survival and portend significant impacts upon ecosystem functioning. We advocate that future studies on phenological synchrony combine data from multiple phenological metrics, to increase confidence in assessments of change and likely ecological consequences., (© 2013 John Wiley & Sons Ltd.)

Phoslock(®) is increasingly used worldwide to control sediment phosphorus (P) release and cyanobacterial blooms despite the fact that little is known about its mode of action in lake bed sediments. This study quantified the effects of Phoslock(®) on sediment elemental composition and P fractionation (one year pre- and post-application of 170 g Phoslock(®) m(-2)) in an attempt to address these knowledge gaps. Post-application, sediment La content was significantly higher in the top 10 cm of the sediment compared to pre-application conditions. Mass balance calculations indicated that the applied mass of La had the potential to bind 25% of potentially release-sensitive P (Pmobile; sum 'labile P', 'reductant-soluble P' and 'organic P' fraction) present in the top 4 cm or 10% of Pmobile present in the top 10 cm of the sediment. Assessing variation in sediment P partitioning indicated that the application caused a significant increase in the mass of P present in the more refractory 'apatite bound P' fraction between post-application month 4 and 7 compared to Pmobile. This suggests that Phoslock(®) controls sediment P release by increasing the mass of P permanently bound in the sediment. To address uncertainty in estimating product dose required to control sediment P release we conducted laboratory assays using intact sediment cores to which we added serial additions of Phoslock(®) under either aerobic or anaerobic conditions. The laboratory experiment indicated that the original dose to Loch Flemington was sufficient to control sediment P release under aerobic conditions but that significant P release will occur should prolonged anaerobic conditions persist. However, Phoslock(®) may be a viable option to control sediment P-release under anaerobic conditions which would require an estimated additional application of up to 510 g Phoslock(®) m(-2). A conceptual model is proposed for the use of P-capping agents in lake remediation projects which is likely to increase cost-effectiveness and reduce non-target effects by applying multiple smaller doses compared to a single high dose., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

Many proteins contain disordered regions under physiological conditions and lack specific three-dimensional structure. These are referred to as IDPs (intrinsically disordered proteins). CP12 is a chloroplast protein of approximately 80 amino acids and has a molecular mass of approximately 8.2-8.5 kDa. It is enriched in charged amino acids and has a small number of hydrophobic residues. It has a high proportion of disorder-promoting residues, but has at least two (often four) cysteine residues forming one (or two) disulfide bridge(s) under oxidizing conditions that confers some order. However, CP12 behaves like an IDP. It appears to be universally distributed in oxygenic photosynthetic organisms and has recently been detected in a cyanophage. The best studied role of CP12 is its regulation of the Calvin cycle responsible for CO2 assimilation. Oxidized CP12 forms a supramolecular complex with two key Calvin cycle enzymes, GAPDH (glyceraldehyde-3-phosphate dehydrogenase) and PRK (phosphoribulokinase), down-regulating their activity. Association-dissociation of this complex, induced by the redox state of CP12, allows the Calvin cycle to be inactive in the dark and active in the light. CP12 is promiscuous and interacts with other enzymes such as aldolase and malate dehydrogenase. It also plays other roles in plant metabolism such as protecting GAPDH from inactivation and scavenging metal ions such as copper and nickel, and it is also linked to stress responses. Thus CP12 seems to be involved in many functions in photosynthetic cells and behaves like a jack of all trades as well as being a master of the Calvin cycle.

Oxygenic photosynthesis evolved at least 2.4 Ga; all oxygenic organisms use the ribulose bisphosphate carboxylase-oxygenase (Rubisco)-photosynthetic carbon reduction cycle (PCRC) rather than one of the five other known pathways of autotrophic CO(2) assimilation. The high CO(2) and (initially) O(2)-free conditions permitted the use of a Rubisco with a high maximum specific reaction rate. As CO(2) decreased and O(2) increased, Rubisco oxygenase activity increased and 2-phosphoglycolate was produced, with the evolution of pathways recycling this inhibitory product to sugar phosphates. Changed atmospheric composition also selected for Rubiscos with higher CO(2) affinity and CO(2)/O(2) selectivity correlated with decreased CO(2)-saturated catalytic capacity and/or for CO(2)-concentrating mechanisms (CCMs). These changes increase the energy, nitrogen, phosphorus, iron, zinc and manganese cost of producing and operating Rubisco-PCRC, while biosphere oxygenation decreased the availability of nitrogen, phosphorus and iron. The majority of algae today have CCMs; the timing of their origins is unclear. If CCMs evolved in a low-CO(2) episode followed by one or more lengthy high-CO(2) episodes, CCM retention could involve a combination of environmental factors known to favour CCM retention in extant organisms that also occur in a warmer high-CO(2) ocean. More investigations, including studies of genetic adaptation, are needed.

Lanthanum-modified bentonite clay (Phoslock(®) is a lake remediation tool designed to strip dissolved phosphorus (P) from the water column and increase the sediment P-sorption capacity. This study investigated short term alterations in sediment elemental composition and sediment P-fractions based on sediment cores taken 2 days before and 28 days following the application of 24 t of Phoslock® to a 9 ha, man-made reservoir. Following the application, sediment lanthanum (La) content increased significantly (p < 0.05; n = 4) in the top 8 cm of the sediment, thereby theoretically increasing sediment P-binding capacity on the whole reservoir scale by 250 kg. Mass balance calculations were used to estimate the theoretical binding of release-sensitive P (P(mobile); sum of 'labile P', 'reductant-soluble P' and 'organic P' fraction) by La across the top 4 cm and 10 cm depth of sediment. The amended mass of La in the sediment had the potential to bind 42% of P(mobile) present in the top 4 cm or 17% of P(mobile) present in the top 10 cm. However, with the exception of a significant increase (p<0.05; n=4) in the 'residual P' fraction in the top 2 cm, sediment P-fractions, including P(mobile,) did not differ significantly following the Phoslock® application. Experimental P-adsorption studies indicated P-saturation values for Phoslock® of 21,670 mg P kg⁻¹ Phoslock®. Sequential extraction of P from saturated Phoslock® under laboratory conditions indicated that around 21% of P bound by Phoslock® was release-sensitive, while around 79% of bound P was unlikely to be released under reducing or common pH (5-9) conditions in shallow lakes. Applying Phoslock® is, therefore, likely to increase the P-sorption capacity of sediments under reducing conditions., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

Carbon dioxide concentrating mechanisms (also known as inorganic carbon concentrating mechanisms; both abbreviated as CCMs) presumably evolved under conditions of low CO(2) availability. However, the timing of their origin is unclear since there are no sound estimates from molecular clocks, and even if there were, there are no proxies for the functioning of CCMs. Accordingly, we cannot use previous episodes of high CO(2) (e.g. the Palaeocene-Eocene Thermal Maximum) to indicate how organisms with CCMs responded. Present and predicted environmental change in terms of increased CO(2) and temperature are leading to increased CO(2) and HCO(3)(-) and decreased CO(3)(2-) and pH in surface seawater, as well as decreasing the depth of the upper mixed layer and increasing the degree of isolation of this layer with respect to nutrient flux from deeper waters. The outcome of these forcing factors is to increase the availability of inorganic carbon, photosynthetic active radiation (PAR) and ultraviolet B radiation (UVB) to aquatic photolithotrophs and to decrease the supply of the nutrients (combined) nitrogen and phosphorus and of any non-aeolian iron. The influence of these variations on CCM expression has been examined to varying degrees as acclimation by extant organisms. Increased PAR increases CCM expression in terms of CO(2) affinity, whilst increased UVB has a range of effects in the organisms examined; little relevant information is available on increased temperature. Decreased combined nitrogen supply generally increases CO(2) affinity, decreased iron availability increases CO(2) affinity, and decreased phosphorus supply has varying effects on the organisms examined. There are few data sets showing interactions amongst the observed changes, and even less information on genetic (adaptation) changes in response to the forcing factors. In freshwaters, changes in phytoplankton species composition may alter with environmental change with consequences for frequency of species with or without CCMs. The information available permits less predictive power as to the effect of the forcing factors on CCM expression than for their overall effects on growth. CCMs are currently not part of models as to how global environmental change has altered, and is likely to further alter, algal and aquatic plant primary productivity.