Stephen C. Maberly

Fluxes of methane and carbon dioxide from a small productive lake to the atmosphere

The fluxes of CH4 and CO2 to the atmosphere, and the relative contributions of ebullition and molecular diffusion, were determined for a small hypertrophic freshwater lake (Priest Pot, UK) over the period May to October 1997. The average total flux of CH4 and CO2 (estimated from 7 sites on the lake) was approximately 52 mmol m−2 d−1 and was apportioned 12 and 40 mmol m−2 d−1 toCH4 and CO2 respectively. Diffusion across the air-water interface accounted for the loss of 0.4and 40 mmol m−2 d−1 of CH4 and CO2 respectively whilst the corresponding figures for ebullition losses were 12.0 (CH4) and 0.23 (CO2) mmol m−2 d−1. Most CH4 (96%) was lost by ebullition, and most CO2 (99%) by diffusive processes. The ebullition of gas, measured at weekly intervals along a transect of the lake, showed high spatial and temporal variation. The CH4 content of the trapped gas varied between 44 and 88% (by volume) and was highest at the deepest points. Pulses of gas ebullition were detected during periods of rapidly falling barometric pressure. Therelevance of the measurements to global estimates ofcarbon emission from freshwaters are discussed.

Algal evolution in relation to atmospheric CO2: carboxylases, carbon-concentrating mechanisms and carbon oxidation cycles

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 CO2 assimilation. The high CO2 and (initially) O2-free conditions permitted the use of a Rubisco with a high maximum specific reaction rate. As CO2 decreased and O2 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 CO2 affinity and CO2/O2 selectivity correlated with decreased CO2-saturated catalytic capacity and/or for CO2-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-CO2 episode followed by one or more lengthy high-CO2 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-CO2 ocean. More investigations, including studies of genetic adaptation, are needed.

Algal and aquatic plant carbon concentrating mechanisms in relation to environmental change.

Carbon dioxide concentrating mechanisms (also known as inorganic carbon concentrating mechanisms; both abbreviated as CCMs) presumably evolved under conditions of low CO2 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 CO2 (e.g. the Palaeocene–Eocene Thermal Maximum) to indicate how organisms with CCMs responded. Present and predicted environmental change in terms of increased CO2 and temperature are leading to increased CO2 and HCO3− and decreased CO32− 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 CO2 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 CO2 affinity, decreased iron availability increases CO2 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.

Hypothesis: The Rate and Scale of Dispersal of Freshwater Diatom Species is a Function of their Global Abundance

We have analysed the geographical records of a representative selection of extant diatom species from a freshwater pond. The more often a species is recorded in the ecological literature, the greater is its apparent global distribution. One explanation is that the frequently recorded species are globally abundant, whereas species that are infrequently recorded are globally rare. We suggest a model in which random dispersal is the dominant force driving large-scale distribution of species, with the rate and scale of dispersal largely determined by global population size. Thus species that are locally rare or abundant are likewise rare or abundant worldwide. It is predicted that many of the rarer diatom species will, with additional sampling effort, be shown to have wide geographical distribution, but this requires intensive studies focused on revealing species that are normally cryptic. The argument in favour of endemic diatom species is untenable, because it is not possible to disprove their existence elsewhere in the biosphere.

The British river of the future: How climate change and human activity might affect two contrasting river ecosystems in England

The possible effects of changing climate on a southern and a north-eastern English river (the Thames and the Yorkshire Ouse, respectively) were examined in relation to water and ecological quality throughout the food web. The CLASSIC hydrological model, driven by output from the Hadley Centre climate model (HadCM3), based on IPCC low and high CO(2) emission scenarios for 2080 were used as the basis for the analysis. Compared to current conditions, the CLASSIC model predicted lower flows for both rivers, in all seasons except winter. Such an outcome would lead to longer residence times (by up to a month in the Thames), with nutrient, organic and biological contaminant concentrations elevated by 70-100% pro-rata, assuming sewage treatment effectiveness remains unchanged. Greater opportunities for phytoplankton growth will arise, and this may be significant in the Thames. Warmer winters and milder springs will favour riverine birds and increase the recruitment of many coarse fish species. However, warm, slow-flowing, shallower water would increase the incidence of fish diseases. These changing conditions would make southern UK rivers in general a less favourable habitat for some species of fish, such as the Atlantic salmon (Salmo salar). Accidental or deliberate, introductions of alien macrophytes and fish may change the range of species in the rivers. In some areas, it is possible that a concurrence of different pressures may give rise to the temporary loss of ecosystem services, such as providing acceptable quality water for humans and industry. An increasing demand for water in southern England due to an expanding population, a possibly reduced flow due to climate change, together with the Water Framework Directive obligation to maintain water quality, will put extreme pressure on river ecosystems, such as the Thames.

Phagotrophy in the origins of photosynthesis in eukaryotes and as a complementary mode of nutrition in phototrophs: relation to Darwin's insectivorous plants

Darwin performed innovative observational and experimental work on the apparently paradoxical occurrence of carnivory in photosynthetic flowering plants. The nutritional use of particulate organic material which also supplies other elements is now known to be widespread in free-living algae as well as in organisms with endosymbiotic algae and with kleptoplastids. In addition to this direct nutritional role, phagotrophy, in the broad sense of internalization of photosynthetic organisms by a eukaryote, is essential for the occurrence of present-day endosymbiotic algae and kleptoplastid-containing protists, and was essential for the origin of plastids themselves. The endosymbiotic phenomena involving photosynthetic organisms clearly played a major role in combining genomes providing different metabolic functions, but, in our opinion, this does not demand a re-appraisal of evolution by natural selection. That the balance of physiological optimization for competition for resources and minimization of losses (e.g. through predation) is a fine one, and thus subject to a complex selective process, is illustrated by the diversity of mixotrophic strategies in extant phytoplankton.

INORGANIC CARBON ACQUISITION BY CHRYSOPHYTES

Twelve species, representing 12 families of the chrysophytes sensu lato, were tested for their ability to take up inorganic carbon. Using the pH‐drift technique, CO2 compensation points generally varied between 1 and 20 μmol · L−1 with a mean concentration of 5 μmol · L−1. Neither pH nor alkalinity affected the CO2 compensation point. The concentration of oxygen had a relatively minor effect on CO2‐uptake kinetics, and the mean CO2 compensation point calculated from the kinetic curves was 3.6 μmol · L−1 at 10–15 kPa starting oxygen partial pressure and 3.8 μmol · L−1 at atmospheric starting oxygen partial pressure (21 kPa). Similarly, uptake kinetics were not affected by alkalinity, and hence concentration of bicarbonate. Membrane inlet mass spectrometry (MIMS) in the presence and absence of acetazolamide suggested that external carbonic anhydrase in Dinobryon sertularia Ehrenb. and Synura petersenii Korschikov was either very low or absent. Rates of net HCO3− uptake were very low (∼5% of oxygen evolution) using MIMS and decreased rather than increased with increasing HCO3− concentration, suggesting that it was not a real uptake. The CO2 compensation points determined by MIMS for CO2 uptake and oxygen evolution were similar to those determined in pH‐drift and were >1 μmol · L−1. Overall, the results suggest that chrysophytes as a group lack a carbon‐concentrating mechanism (CCM), or an ability to make use of bicarbonate as an alternative source of inorganic carbon. The possible evolutionary and ecological consequences of this are briefly discussed.

Linking nutrient limitation and water chemistry in upland lakes to catchment characteristics

The relationship between catchment characteristics and lake water chemistry, phytoplankton and periphyton biomass, and phytoplankton and periphyton nutrient limitation was investigated for 30 upland lakes in the U.K. These catchment characteristics included the proportion of different land cover categories in the catchment and some hydrological information. Multiple regression models could predict alkalinity, pH, total dissolved phosphorus, dissolved inorganic nitrogen, dissolved organic nitrogen, dissolved organic carbon and phytoplankton chlorophyll a from the proportional contribution of between two and six land cover categories within the catchment and explain between 42 and 73% of the variance. Phosphorus limitation was positively linked to the proportion of shrub-heath and bracken in the catchment, and negatively linked to the proportion of pasture. Nitrogen limitation was positively linked to the proportion of marsh and rough grass, deciduous and mixed woodland, and negatively linked to the proportion of rough pasture, shrub heath and bare ground in the catchment. Nitrogen limitation decreased and phosphorus limitation increased with catchment slope, although the correlation between land cover classes and slope was not significant. The results suggest that map-based data can be used to predict water chemistry and nutrient limitation in upland lakes.

Geoengineering in lakes: welcome attraction or fatal distraction?

Abstract The use of geoengineering techniques for phosphorus management offers the promise of greater and quicker chemical and ecological recovery. It can be attractive when used with other restoration measures but should not be considered a panacea. The range of materials being proposed for use as well as the in-lake processes targeted for manipulation continues to grow. With increasing political imperatives to meet regulatory goals for water quality, we recommend a coordinated approach to the scientific understanding, costs, and integration of geoengineering with other approaches to lake management.

Comparative sequence analysis of CP12, a small protein involved in the formation of a Calvin cycle complex in photosynthetic organisms

CP12, a small intrinsically unstructured protein, plays an important role in the regulation of the Calvin cycle by forming a complex with phosphoribulokinase (PRK) and glyceraldehyde-3-phosphate dehydrogenase (GAPDH). An extensive search in databases revealed 129 protein sequences from, higher plants, mosses and liverworts, different groups of eukaryotic algae and cyanobacteria. CP12 was identified throughout the Plantae, apart from in the Prasinophyceae. Within the Chromalveolata, two putative CP12 proteins have been found in the genomes of the diatom Thalassiosira pseudonana and the haptophyte Emiliania huxleyi, but specific searches in further chromalveolate genomes or EST datasets did not reveal any CP12 sequences in other Prymnesiophyceae, Dinophyceae or Pelagophyceae. A species from the Euglenophyceae within the Excavata also appeared to lack CP12. Phylogenetic analysis showed a clear separation into a number of higher taxonomic clades and among different forms of CP12 in higher plants. Cyanobacteria, Chlorophyceae, Rhodophyta and Glaucophyceae, Bryophyta, and the CP12-3 forms in higher plants all form separate clades. The degree of disorder of CP12 was higher in higher plants than in the eukaryotic algae and cyanobacteria apart from the green algal class Mesostigmatophyceae, which is ancestral to the streptophytes. This suggests that CP12 has evolved to become more flexible and possibly take on more general roles. Different features of the CP12 sequences in the different taxonomic groups and their potential functions and interactions in the Calvin cycle are discussed.