PHOTOSYNTHESIS AND PHOTOSYNTHESIS-COUPLED RESPIRATION IN NATURAL BIOFILMS QUANTIFIED WITH OXYGEN MICROSENSORS1

Photosynthesis and respiration were analyzed in natural biofilms by use of O₂ microsensors. Depth profiles of gross photosynthesis were obtained from the rate of decrease in O₂ concentration during the first few seconds following extinction of light, and net photosynthesis of the photic zone was calculated from O₂ concentration gradients measured at steady state. Respiration within the photic zone was calculated as the difference between gross and net photosynthesis. Two types of biofilms were investigated: one dominated by diatoms, and one dominated by cyanobacteria. High O₂/CO₂ ratios caused increased respiration especially within the diatom biofilm, which could indicate that photorespiration was a dominant O₂-consuming process. The rate of respiration was constant within both biofilms during the first 4.6 s following extinction of light, even when respiration was stimulated by high O₂/CO₂ ratio. The assumption of a constant rate of respiration during the dark period is an essential one for the determination of gross photosynthetic activity by use of O₂ microsensors. We here present the first evidence to substantiate this assumption. The results strongly suggest that gross photosynthesis as measured by use of O₂ microsensors may include carbon equivalents that are subsequently lost through photorespiration. Computer modeling of photosynthesis profiles measured after 1.1, 1.6, and 2.6 s of dark incubation illustrated how the actual photosynthesis profile could have appeared if it had been possible to do the determination at time 0. Diffusion of O₂ during the up to 4.6-s long dark incubations did not affect gross photosynthetic rate when integrated over all depths, but the apparent vertical distribution of the photosynthetic activity was strongly affected.     

Temperature- and Light-Dependent Performance of the Cyanobacterium <Emphasis Type="Italic">Leptolyngbya Foveolarum</Emphasis> and the Diatom <Emphasis Type="Italic">Nitzschia Perminuta</Emphasis> in Mixed Biofilms

This paper investigates the role of species interactions as a mechanism determining the changing seasonal abundance of microphytobenthic species. Different kinds of interactions can occur between microphytobenthic species, e.g., interference competition (a species directly hindering the growth of another) or resource competition. If such interactions are strong, the capacity of species to exploit parts of the seasonal spectrum of temperature and light conditions could be greatly affected. A model system of two freshwater benthic phototrophs, the cyanobacterium Leptolyngbya foveolarum (Rabenhorst ex Gomont) Anagnostidis et Komárek and the diatom Nitzschia perminuta (Grunow) M. Pergallo, was used to study the capacity of each species to grow in ranges of temperature (7, 15 and 25 °C) and light (5, 40 and 200 μmol m⁻² s⁻¹) conditions in single-species and two-species cultures. Growth was followed for 14–17 days by measuring chlorophyll a and maximum photosynthetic capacity using spectrophotometry and pulse amplitude modulated (PAM) fluorimetry. A PHYTO-PAM fluorimeter facilitated simultaneous observations in mixed cultures on the two species. In mixed cultures, the diatom appeared to be a ‘cool season species’ (low temperature and low light intensity) and the cyanobacterium a ‘summer or autumn species’ (higher temperature and light intensities). This is different than predicted by monoculture experiments, where a wide range of optimal growth conditions was found. Two-species biofilm tests indicated inhibitory effects of the cyanobacterium on the diatom species, especially under conditions favorable to the cyanobacterium. High or low light intensities, increase of local pH caused by depletion of inorganic carbon, and limitation of other inorganic nutrients (resource competition) were examined as factors contributing to diatom inhibition, but none provided an acceptable explanation for observed growth patterns. Our results pointed towards interference competition.     

Particulate fatty acids in two small Siberian reservoirs dominated by different groups of phytoplankton

SUMMARY 1. We studied the composition of fatty acids (FAs) in the seston from two small freshwater reservoirs (Bugach and Lesnoi) with distinct periodicity of domination by cyanobacteria and eukaryotic algae during the growth season.
2. The diatoms in the both reservoirs were characterised by a high content of 14:0 and C16 unsaturated acids, whereas that of the essential FA 20:5ω3 [eicosapentanoic acid (EPA)] was low. The correlation between this polyunsaturated FA (PUFA) and diatom biomass was not significant in either reservoir. The percentage of 20:5ω3 in seston significantly correlated with the biomass of euglenophyta in Bugach and dinophyta in Lesnoi. Hence the diatoms, usually referred as a valuable food for zooplankton, were not an important source of the essential PUFA in these systems.
3. The dominant cyanobacteria in Bugach, and the green algae in Lesnoi, both contained the same marker acids: 18:3ω3 and 18:2ω6. Hence, a discrimination between these two phytoplanktonic groups on the basis of FA biomarkers may be difficult in some cases.
4. We found no significant correlation between the content of 20:5ω3 in seston and the biomass of the dominant daphniids in either reservoir. This is contrary to expectations, based on the literature, that EPA is generally important. Rather, the biomass of the two dominant Daphnia species in Bugach correlated strongly with the content of 18:3ω3 in the seston. The cyanobacteria were a probable source of this ω3 FA for Daphnia . We conclude that EPA is not always important for Daphnia populations although, in such cases, some other PUFA (e.g. 18:3ω3) might be related to their growth.     

Diel activity of Gammarus pulex (Crustacea) in a South Swedish stream: comparison of drift catches vs baited traps

The benthos of a perennially ice-covered Antarctic lake, Lake Hoare, contained three distinct signatures of lipophilic pigments. Cyanobacterial mats found in the moat at the periphery of the lake were dominated by the carotenoid myxoxanthophyll; carotenoids: chlorophyll a ratios in this high light environment ranged from 3 to 6.8. Chlorophyll c and fucoxanthin, pigments typical of golden-brown algae, were found at 10 to 20 m depths where the benthos is aerobic. Anaerobic benthic sediments at 20 to 30 m depths were characterized by a third pigment signature dominated by a carotenoid, tentatively identified as alloxanthin from planktonic cryptomonads, and by phaeophytin b from senescent green algae. Pigments were not found associated with alternating organic and sediment layers. As microzooplankton grazers are absent from this closed system and transformation rates are reduced at low temperatures, the benthos beneath the lake ice appears to contain a record of past phytoplankton blooms undergoing decay.     

A Novel Ca2+ Signaling Pathway Coordinates Environmental Phosphorus Sensing and Nitrogen Metabolism in Marine Diatoms

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Short-term response of monospecific and natural algal biofilms to copper exposure

The effect of copper additions (Cu ranging from 0 to 30?µM) on the photosynthesis of three different microalgal biofilms was studied to identify the factors that cause sensitivity differences between benthic and pelagic algae. The response of biofilms which colonized artificial substrata in the River Meuse was compared with those of two laboratory-grown monospecific biofilms, one consisting of the diatom Synedra ulna, and the other composed of a filament-forming cyanobacterium, Oscillatoria sp. The photosynthetic yield Φ_II (quantum efficiency of photosystem II) was studied with PAM (Pulse Amplitude Modulated) fluorimetry. S. ulna biofilms appeared to be the most sensitive to Cu, followed by the cyanobacteria, while natural biofilms, dominated by supposedly very sensitive diatom species such as Melosira varians and Diatoma vulgare, were the most resistant to Cu. In the highly productive biofilms, pH is suggested to play a role in lowering toxicity by helping the precipitation of cupric ions. Cu accumulation by the biofilms during the exposure period followed a linear relationship with Cu concentration, saturation not being observed; natural biofilms had an accumulation factor of 1–2.5?×?10³ relative to the concentrations in the water, while the diatoms growing unattached to the substratum had a higher concentration factor, up to 4.9?×?10³. It was concluded that the physical structure of the biofilm (package of cells and thickness), and not the species composition, was the main factor regulating the sensitivity of the biofilm to Cu toxicity during short-term exposures.     

Resource competition among saline-lake diatoms under varying N/P ratio, salinity and anion composition

1. A series of resource competition experiments were performed with four diatom taxa isolated from saline lakes in the Northern Great Plains of North America. Three variables were tested in a factorial design: N/P ratio (6 : 1, 12 : 1, 24 : 1, 48 : 1), salinity (5, 8, 11‰), and anion composition (sulphate versus bicarbonate-dominated medium).
2. Coexistence of Chaetoceros elmorei , Cyclotella quillensis , and Cymbella pusilla occurred at the two lowest N/P ratios. These three taxa have similar N requirements and thus competitive exclusion by the taxon with the lowest N requirement may have required more time.
3. Anomoeoneis costata , which had the highest nitrogen requirements and slowest growth rate of these four taxa, declined in all cases. Cyclotella biovolume concentration consistently declined at the two highest N/P ratios, indicating a higher P requirement, which is consistent with its distribution pattern.
4. In general, we found opposite trends with respect to all variables for Chaetoceros , a taxon that mainly inhabits sulphate systems, versus Cymbella , a bicarbonate-associated taxon. Our results suggest that Cymbella may be found mainly in bicarbonate systems because of a higher C requirement, whereas Chaetoceros may be less abundant in these systems, in part, because of competitive interactions with bicarbonate-associated taxa.     

Priority effects in a planktonic bloom-forming marine diatom

Priority effects occur when a species or genotype with earlier arrival has an advantage such that its relative abundance in the community or population is increased compared with later-arriving species. Few studies have dealt with this concept in the context of within-species competition. Skeletonema marinoi is a marine diatom that shows a high degree of genetic differentiation between populations over small geographical distances. To test whether historical events such as priority effects may have been important in inducing these patterns of population differentiation, we performed microcosm experiments with successive inoculation of different S. marinoi strains. Our results show that even in the absence of a numerical advantage, significant priority effects were evident. We propose that priority effects may be an important mechanism in initiating population genetic differentiation.     

Combining a regional climate model with a phytoplankton community model to predict future changes in phytoplankton in lakes

1. Linking a regional climate model (RCM) configured for contemporary atmospheric greenhouse gas concentrations, with a phytoplankton community model (PROTECH) produced realistic simulations of 20 years of recent phytoplankton data from Bassenthwaite Lake, in the North‐West of England. 2. Meteorological drivers were derived from the RCM to represent a future climate scenario involving a 1% per annum compound increase in atmospheric CO₂ concentrations until 2100. Using these drivers, PROTECH was run for another 20 year period representing the last two decades of the 21st century. 3. Comparison of these present and future simulations revealed likely impacts on the current seasonal phytoplankton development. Under future climate conditions, the simulated spring bloom showed an increase in cyanobacteria dominance caused by greater success of Planktothrix. Also, the summer cyanobacteria bloom declined earlier because of nutrient limitation caused by the increased spring growth. Overall productivity in the lake did not change. 4. Analysis showed that these predicted changes were driven by changes in water temperature, which were in turn triggered by the higher air temperatures predicted by the RCM.     

Response of grazing impacts of an algivorous fish (Pseudogastromyzon myersi: Balitoridae) to seasonal disturbance in Hong Kong streams

1. Manipulative experiments were carried out in four Hong Kong streams (two shaded, two unshaded) to investigate the impact of grazing by an algivorous fish, Pseudogastromyzon myersi, on benthic algal biomass and assemblage composition. Experiments were conducted and repeated during both the dry and wet seasons to determine whether spate‐induced disturbance modified any grazing effect. Treatments comprised fish exclusion and inclusion via closed and open cages, with a no‐cage treatment used as a control for the cage effect. Treatments were maintained for 4 weeks in each experimental run. 2. Grazing by P. myersi reduced benthic algal biomass and the organic matter content of periphyton in open cages and the no‐cage treatment relative to closed cages. The similarity between open‐cage and no‐cage treatments was evidence that the overall difference among treatments was caused by limiting fish access to closed cages and not merely an artifact of caging. Grazing effects were broadly similar in all streams, but there was a significant statistical interaction between treatments and seasons. 3. Analysis of dry‐season data matched the overall trend in inter‐treatment differences, confirming the effects of grazing by P. myersi on algal biomass and periphyton organic matter. Significant differences in algal assemblage composition between closed‐cage and no‐cage treatments during the dry season reflected reductions in the abundance of erect, stalked diatoms (Gomphonema) and filamentous cyanobacteria (Homeothrix). Removal of these vulnerable overstorey algae by P. myersi resulted in greater abundance of understorey diatoms (Achnanthes and Cocconeis) in the no‐cage treatment in all streams during the dry season. The composition of algal assemblages in open cages was intermediate between the other two treatments. 4. Although fish densities were greater in all streams during the wet season, spate‐induced disturbance obscured grazing effects and there were no significant differences among treatments attributable to fish grazing. Seasonal variation in impacts of P. myersi grazing provides support for the harsh‐benign hypothesis, and confirms that biotic factors are less important controls of stream algal biomass and assemblage structure during periods (i.e. the wet season in Hong Kong) when abiotic disturbances are frequent or intense.     

Artificial cell-immobilization: a model simulating immobilization in natural environments?

A study of the marine diatom Haslea ostrearia was performed in controlled culture conditions. Biofilm formation by cells grown in liquid medium was studied through macroscopic events and extracellular polysaccharides (EPS) production. Alcian blue staining and image analysis methods were used to quantify EPS production in regard to the usual pattern (lag, exponential and stationary phases) of microalgal growth. Simultaneously, an artificial immobilization of H. ostrearia cells was carried out using an agar layer. Samples from liquid medium and artificially immobilized cultures were studied in scanning electron microscopy. They revealed the structure of the entrapping material and progressive changes of the natural biofilm along culture ageing. The use of artificially immobilized cultures for solute diffusivity investigations is discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Effects of cadmium stress on periphytic diatom communities in indoor artificial streams

SUMMARY 1. Two experiments were performed with periphytic diatoms originating from the River Lot (France) and allowed to grow on clean substrata within indoor artificial streams. Three cadmium (Cd) levels (control, low Cd=10 μg L⁻¹ and high Cd=100 μg L⁻¹) were used to test the effects of Cd on (i) the settlement and development of diatom communities (Experiment 1, 4 weeks Cd exposure) and (ii) predeveloped communities grown over a 2-week period without contamination (Experiment 2, 2 weeks Cd exposure).
2. Experiment 1 revealed that growth and taxonomic composition of diatom communities clearly differed with Cd exposure. Biofilms were more adpressed to substrata under low Cd concentration, and were thinner and patchy under high Cd concentration.
3. Exposure of developed diatom communities (Experiment 2) to Cd revealed only minor variations in taxonomic composition, possibly linked to the protective role of the developed organic matrix against metal stress.
4. These results support the validity of periphytic diatom communities as indicators of metal pollution, although significant structural changes would take longer within developed communities.     

Benthic Algae in High Altitude Streams of the Alps – a Neglected Component of the Aquatic Biota

This is a review on benthic algae from streams situated above the tree line in the Alps. It integrates published and unpublished data from alpine streams in Austria and in the Trento Province (Northern Italy). The main focus is on the structural and taxonomic composition of benthic algae including macro- and micro-algae and their contribution to the epilithic biofilm and the stream food-web. The environmental conditions relevant to algae within the two major stream types, the glacial (glacier-fed) and non-glacial krenal (spring-fed) stream are discussed. The paper considers both the maximum possible structural complexity of transverse algae zonation in cascading alpine/subalpine stream segments, and the effects of glacial water on species richness in the Central Alps in Austria. Autecological data are given for 46 common diatoms from 42 sites in the Central Eastern and Southern Central Alps and for 30 algae in addition to diatoms for 22 streams in the Central Eastern Alps. Since data on responses of benthic algae to the harsh conditions in high altitude Alpine streams are very scarce, results from literature and the author’s experiences from these and other mountain stream types are synthesized to formulate major objectives for future research in benthic high altitude Alpine stream ecology.     

The radiation of spring snails of the genus Belgrandiella in Austria (Mollusca: Caenogastropoda: Hydrobiidae)

Grain density microautoradiography (MAR) was used to study uptake of dissolved amino acids (DFAA) in microalgal communities from a sandy bay on the west coast of Sweden. A mixture of fifteen ³H-labeled amino acids (final concentration 20–80 nmol l^–1) was added to sediment samples collected from two depths (0.5 and 4 m), on five occasions representing different seasons. On all sampling occasions, the microflora was dominated by diatoms (> 85% of the total biomass). Cyanobacteria occurred in the summer and autumn, but never dominated the biomass. Between 5 and 48% of all counted algal cells showed uptake (1–44% of algal biomass). Uptake was recorded for all majors microalgal groups (diatoms, cyanobacteria and autotrophic flagellates). Uptake was more frequent on the shallow site and the highest proportion of cells showing uptake occurred in May. Although uptake was common among both motile and attached growth forms, on several occasions uptake frequencies were higher for attached cells. As the attached fraction is frequently burried out of the photic zone, and not able to rapidly migrate towards light, this implies that heterotrophic capacity should be valuable. Also at species level, some taxa showed higher uptake frequencies than others, for example populations of motile diatom species that displayed a seasonal behaviour (blooming species), such as Nitzschia cf. dissipata and Cylindrotheca closterium. Other species showing frequent uptake were the coccoid colony-forming cyanobacterium Microcrocis sp., and the filamentous cyanobacterium Phormidium sp.. The results suggest that there is a high potential for DFAA uptake in the microphytobenthic community and that for some growth forms and species this could imply a competitive advantage.     

Associating particulate essential fatty acids of the ω3 family with phytoplankton species composition in a Siberian reservoir

1. We studied variation in the composition of fatty acids in the seston of a small freshwater reservoir with changes in phytoplankton composition during four growth seasons. We focused on the dynamics of the ω3 fatty acids because of their potential importance for zooplankton nutrition. 2. Total diatoms were related to the 20:5ω3 fatty acid (eicosapentaenoic, EPA) content in seston. Among two dominant diatom genera, Cyclotella was not associated with EPA content. In contrast, there was a significant correlation between Stephanodiscus and the percentage contribution and content of EPA throughout the study. Hence, freshwater diatoms can differ strongly in content of the essential EPA. 3. We considered abundant cyanobacteria as a potential source of 18:3ω3 fatty acid (linolenic, ALA) to aquatic food webs. Among four dominant cyanobacteria species, two (Anabaena flos‐aquae and Planktothrix agardhii) showed significant correlation with the ALA content of the seston, while the other two (Aphanizomenon flos‐aquae and Microcystis aeruginosa) did not. 4. Dinophyta had a relatively high level of 22:6ω3 (docosahexaenoic, DHA) for freshwater species and can be also a source of EPA to aquatic food webs. 5. Our results show that various species of diatoms as well as cyanobacteria can be of contrasting nutritional value for zooplankton because of their different content of the essential PUFAs. Diatoms, which are low in EPA, could not be considered as a valuable food, while some field populations of cyanobacteria might be valuable sources of essential ALA.