ACCUMULATION OF FERREDOXIN AND FLAVODOXIN IN A MARINE DIATOM IN RESPONSE TO FE

Despite recognition that Fe availability is significant in regulating oceanic production in some regions, the biogeochemistry of this trace element is poorly understood. To complement contemporary methods of analytical chemistry, we have used an immunological approach to monitor the Fe nutrition of marine phytoplankton. In prokaryotes and numerous microalgae, the redox catalyst ferredoxin is functionally replaced by flavodoxin during periods of Fe deficiency. In this study, antibodies were raised against ferredoxin purified from a marine diatom, and their utility as a diagnostic indicator was assessed. A species survey demonstrated broad reactivity with both pennate and centric diatoms and additionally with several nondiatom taxa. In batch cultures of the diatom Phaeodactylum tricornutum Bohlin, in which Fe levels were varied, accumulation of ferredoxin varied with the physiological state of the culture; in unimpaired cells (F_v/F_m≥ 0.65), ferredoxin levels were high, whereas levels dropped markedly in cells experiencing even slight photochemical impairment. Accumulation of flavodoxin varied inversely with that of ferredoxin. An experiment was performed to demonstrate the temporal pattern of accumulation of ferredoxin upon recovery from Fe limitation. Prior to Fe amendment, cells were physiologically impaired (chlorotic, F_v/F_m < 0.3) and contained flavodoxin but no detectable ferredoxin. Following addition of Fe, constraints on photochemistry were relaxed within hours. Coinciding with this, levels of flavodoxin declined, whereas ferredoxin was accumulated to high levels within 8 h.     

On the algal ecology of the lowland Iraqi marshes

An account is given of the algal vegetation in the near boat channels in the central area of the lowland Iraqi marshes. Sampling was carried out in February and September. The water here may be characterized as relatively turbid, slightly saline, highly calcareous and probably typically mesotrophic. Diatoms dominated the phytoplankton, many of the species being ones which are at least as characteristic of benthic communities. Periphyton was more conspicuous in February than September, with filamentous green algae especially common in February. Blue-green algae were more common in September; they frequently formed a felt with several different species and irregular deposition of calcite. Heterocystous blue-green algae were frequent in the periphyton but apparently not so in the plankton. A well-developed laminated community of blue-green algae and purple phototrophic bacteria occurred on the wooden hull of a boat.     

PHOTOSYNTHETIC PIGMENT ORGANIZATION IN DIATOMS (BACILLARIOPHYCEAE)1

The isolation and characterization of six pigment-protein complexes from five diatom species (Phaeodactylum tricornutum Bohlin, Chaetoceros gracilis Schutt, Nitzschia sp. Mono Lake, Nitzschia laevis Hust. and Thalassiosira pseudonana (Hust.) Hasle and Heimdal) was accomplished by membrane dissociation with digitonin followed by gel electrophoresis. Six analogous complexes obtained from all species were correlated in spectral characteristics and relative mass with complexes from higher plants obtained by the same procedure. The largest of these complexes, comprising about 15% of the total Chl a, contained reaction centers of Photosystem I (PSI) and antenna pigments (LHC₁).Some PSI complexes also separated from LHC₁ in the gel. For the first time in diatoms, a Photosystem II complex was isolated and identified from its position in the gels, absorption and fluorescence spectra, lack of P700, and enrichment carotene. Three antenna pigment-protein complexes in addition to LHC₁ occurred in varying proportion under different experimental conditions but in sum, they accounted for 70% of the total Chl a. All three bands were highly enriched in Chl c and fucoxanthin, although the ratio of Chl c/ xanthophyll decreased between the slowest migrating LHC₂ and fastest moving LHC₄ LHC₃ contained the highest proportio of pigment-protein and was composed primarily of polypeptides of about 18,000 D. Essentially all α- and β-carotene was bound to the reaction center complexes. The Nitzschia from Mono Lake differed from the other species in that PSI complexes could not be readily dissociated from its membrane by digitonin treatment, a characteristic which may reflect a different chloroplast membrane structure in this alga.     

Influence of the mixing regime on algal photosynthetic performance in laboratory scale enclosures

The photosynthesis of plankton sampled from the eutrophic Lake Loosdrecht was studied in Laboratory Scale Enclosures (LSEs) with regard to the rate of mixing. First, two LSEs were operated at different mixing rates. No significant differences in photosynthetic performance were found, with the exception of a depressed photosynthesis in the afternoon in the LSE which had a low mixing rate. Secondly, when mixing was stopped, the phytoplankton which stayed in the dark due to the steep light gradient in the LSE responded by changing its maximal photosynthetic capacity. The results show that the filamentous cyanobacteria in the lake can respond rapidly to changes in the depth of the mixed layer by altering their photosynthetic performance.This is publication 567 of the Centre for Estuarine and Coastal Ecology.     

A fluorometric method for the differentiation of algal populations in vivo and in situ

Fingerprints of excitation spectra of chlorophyll (Chl) fluorescence can be used to differentiate `spectral groups' of microalgae in vivo and in situ in, for example, vertical profiles within a few seconds. The investigated spectral groups of algae (green group, Chlorophyta; blue, Cyanobacteria; brown, Heterokontophyta, Haptophyta, Dinophyta; mixed, Cryptophyta) are each characterised by a specific composition of photosynthetic antenna pigments and, consequently, by a specific excitation spectrum of the Chl fluorescence. Particularly relevant are Chl a, Chl c, phycocyanobilin, phycoerythrobilin, fucoxanthin and peridinin. A laboratory-based instrument and a submersible instrument were constructed containing light-emitting diodes to excite Chl fluorescence in five distinct wavelength ranges. Norm spectra were determined for the four spectral algal groups (several species per group). Using these norm spectra and the actual five-point excitation spectrum of a water sample, a separate estimate of the respective Chl concentration is rapidly obtained for each algal group. The results of dilution experiments are presented. In vivo and in situ measurements are compared with results obtained by HPLC analysis. Depth profiles of the distribution of spectral algal groups taken over a time period of few seconds are shown. The method for algae differentiation described here opens up new research areas, monitoring and supervision tasks related to photosynthetic primary production in aquatic environments. This revised version was published online in June 2006 with corrections to the Cover Date.     

Aspects of Scotia Sea zooplankton

Information on small scale distributions of three species of Antarctic zooplankton is reviewed. Aggregations of the euphausiid Euphausia superba , the tunicate Salpa thompsoni , and the amphipod Parathemisto gaudichaudii are compared, and the manner in which such aggregations mav arise is discussed. A possible relationship between swarming and feeding activity in E. superba is suggested in which krill are thought to be dispersed whilst feeding but that on repletion they swarm. It is thought that this may account for this species' irregular spatial distribution as recorded bv previous expeditions. A further consequence of this theory is that during the Winter swarming will be minimal.     

THE RELATIVE PHOTOSYNTHETIC EFFICIENCY OF ASTERIONELLA FORMOSA HASS. (BACILLARIOPHYTA) IN NATURAL PLANKTON ASSEMBLAGES1

The relative photosynthetic efficiencies of net vs. nanoplankton and diatom vs. non-diatom plankton were estimated weekly in Woodcock Creek Reservoir, Pennsylvania, from April to November 1979, using carbon-14 in conjunction with size fractionation and germanium as a diatom inhibitor. The nanoplankton contributed a proportionally greater share of the community carbon assimilation than their share of the community biomass would indicate. Diatoms, specifically Asterionella formosa Hass. were photosynthetically inefficient and contributed significantly less than expected. The mechanism of periodic diatom dominance in lakes remains unexplained.     

Group-specific phytoplankton biomass/dissolved carbohydrate relationships in the Gulf of Trieste (Northern Adriatic)

Distribution of carbohydrates (CHO) and photosynthetic pigments werestudied in the Gulf of Trieste, northern Adriatic Sea, during the period ofsummer stratification with a special emphasis on determining the impact ofthe taxonomic composition and concentration of phytoplankton biomass on thecarbohydrate levels in the water column. Dissolved total carbohydrates(DTCHO), dissolved monosaccharides (DMCHO) as well as particulatecarbohydrates (PTCHO) were determined using the colorimetric MBTH-method,while pigment biomarkers of the phytoplankton biomass were determined byreversed-phase HPLC. Concentrations of the total CHO (dissolved+particulate) varied in a wide range from 173 μg Cl^™1 to 1552 μg C l^™1. The percentage ofPTCHO in the total CHO concentration was relatively low(4–25%), indicating that the main pool of CHO was in thedissolved phase. The contribution of DTCHO to the total dissolved organiccarbon (DOC) in late summeontribution r was highly variable(10–65%) with an average value of 20 ± 14%, whilein early summer this percentage was somewhat lower and less variable (range11–23%; average 17± 3%). Analyses of biomarkerpigments revealed a very high diversity and a rather heterogenous verticaland spatial distribution of the phytoplankton biomass during the period ofsummer stratification. In September 1994, the predominant taxonomic groupsof phytoplankton were prymnesiophytes, diatoms, silicoflagellates,cyanobacteria and, especially in the bottom layer, dinoflagellates. Arelatively good correlation (r² =0.51) found betweenDTCHO and chl a suggested that DTCHO were mainly of phytoplankton origin.Furthermore, a concomitant increase of DTCHO with peridinin and fucoxanthinindicated that dinoflagellates and diatoms had a decisive impact on CHOlevels in the water column. By contrast, early summer phytoplankton (June),which was dominated by prymnesiophytes, exhibited a comparatively lowerimpact on the CHO distribution. This revised version was published online in July 2006 with corrections to the Cover Date.     

AUXOSPORE FORMATION BY THE SILICA‐SINKING,OCEANIC DIATOM FRAGILARIOPSIS KERGUELENSIS (BACILLARIOPHYCEAE)1

Size restoration by the auxospore that develops from the zygote is a crucial stage in diatom life cycles. However, information on sexual events in pelagic diatom species is very limited. We report for the first time auxospore formation by the pennate diatom Fragilariopsis kerguelensis (O'Hara) Hustedt during an iron‐induced bloom in the Southern Ocean (EIFEX, European Iron Fertilization EXperiment). Auxospores of F. kerguelensis resembled those described for Pseudo‐nitzschia species. The auxospore was characterized by an outer coating, the perizonium; two caps, one at each distal end; and four chloroplasts, one at each end and two in the central part. Different stages of auxospore elongation were recorded, with a length of 24–91 μm, but only the largest auxospores contained the initial cell, whose apical axis ranged between 76 and 90 μm. Gametangial cell walls were often attached to the auxospores and ranged from 10 to 31 μm in length. Auxospore abundances were consistently higher in the fertilized patch, where an increase in the F. kerguelensis population was observed, as compared with surrounding waters.     

Responses of estuarine intertidal microphytobenthic algal assemblages to enhanced ultraviolet B radiation

As a result of ozone depletion, ground doses of ultraviolet B (UVB) radiation in the mid latitudes of the Northern Hemisphere have increased since the 1980s, and current predictions indicate no possible alleviation until at least post 2020. Mudflats and sandflats are important coastal-zone habitats, and support extensive biofilms of benthic microalgae (microphytobenthos). In intertidal situations, these assemblages are exposed to high levels of UVB radiation during periods of tidal exposure. Exposure of intertidal biofilms dominated by epipelic (mud-inhabiting) diatoms to 0, 0.18 or 0.35 W m⁻² UVB radiation for between 4 and 10 days resulted in no significant decreases in the maximum PSII quantum efficiency (F_v/F_m) throughout diel exposure periods. Although the quantum efficiency of electron transport (F_q′/F_m′) showed significant reductions early in some experiments, the major response was an increase in F_q′/F_m′ in UVB exposed biofilms. This increase in F_q′/F_m′ was suggestive of a protective vertical migration down into the sediment. Single-cell and whole biofilm fluorescence imaging demonstrated, for the first time, that motile diatoms are able to detect UVB radiation independently of UVA or photosynthetically active radiation (PAR) and migrate rapidly down (within 15 min) into the sediments to avoid it. This behavioural acclimation mechanism appears to prevent significant accumulation of UVB induced damage to the algae. UVB exposure had no significant effect of biofilm photosynthesis (measured by ¹⁴C carbon fixation), but did alter organic carbon allocation patterns, with significantly less new carbon allocated to intracellular storage (glucan) and extracellular colloidal carbohydrate fractions. Significant reductions in the sediment standing stocks of chlorophyll a (Chl a), colloidal carbohydrates, extracellular polymeric substances (EPS) were seen after 7 days of UVB exposure. This study showed that marine intertidal benthic diatoms use a behavioural strategy to avoid exposure to UVB and that this response is effective as a short-term protection mechanism against UVB damage. However, altered carbon allocation patterns feed forward over time into changes in biofilm biomass and sediment carbohydrate dynamics. This suggests that continual long-term exposure to UVB may impact on sediment carbon cycling and trophic interactions and on the stabilization of sediments by microalgal biofilms through their production of extracellular carbohydrates.     

Why is the South Orkney Island shelf (the world's first high seas marine protected area) a carbon immobilization hotspot?

The Southern Ocean archipelago, the South Orkney Islands (SOI), became the world's first entirely high seas marine protected area (MPA) in 2010. The SOI continental shelf (~44 000 km²), was less than half covered by grounded ice sheet during glaciations, is biologically rich and a key area of both sea surface warming and sea‐ice losses. Little was known of the carbon cycle there, but recent work showed it was a very important site of carbon immobilization (net annual carbon accumulation) by benthos, one of the few demonstrable negative feedbacks to climate change. Carbon immobilization by SOI bryozoans was higher, per species, unit area and ice‐free day, than anywhere‐else polar. Here, we investigate why carbon immobilization has been so high at SOI, and whether this is due to high density, longevity or high annual production in six study species of bryozoans (benthic suspension feeders). We compared benthic carbon immobilization across major regions around West Antarctica with sea‐ice and primary production, from remotely sensed and directly sampled sources. Lowest carbon immobilization was at the northernmost study regions (South Georgia) and southernmost Amundsen Sea. However, data standardized for age and density showed that only SOI was anomalous (high). High immobilization at SOI was due to very high annual production of bryozoans (rather than high densities or longevity), which were 2x, 3x and 5x higher than on the Bellingshausen, South Georgia and Amundsen shelves, respectively. We found that carbon immobilization correlated to the duration (but not peak or integrated biomass) of phytoplankton blooms, both in directly sampled, local scale data and across regions using remote‐sensed data. The long bloom at SOI seems to drive considerable carbon immobilization, but sea‐ice losses across West Antarctica mean that significant carbon sinks and negative feedbacks to climate change could also develop in the Bellingshausen and Amundsen seas.     

Exploring ocean biogeochemistry by single-cell microprobe analysis of protist elemental composition

The biogeochemical cycles of many elements in the ocean are linked by their simultaneous incorporation into protists. In order to understand these elemental interactions and their implications for global biogeochemical cycles, accurate measures of cellular element stoichiometries are needed. Bulk analysis of size-fractionated particulate material obscures the unique biogeochemical roles of different functional groups such as diatoms, calcifying protists, and diazotrophs. Elemental analysis of individual protist cells can be performed using electron, proton, and synchrotron X-ray microprobes. Here we review the capabilities and limitations of each approach and the application of these advanced techniques to cells collected from natural communities. Particular attention is paid to recent studies of plankton biogeochemistry in low-iron waters of the Southern Ocean. Single-cell analyses have revealed significant inter-taxa differences in phosphorus, iron, and nickel quotas. Differences in the response of autotrophs and heterotrophs to iron fertilization were also observed. Two-dimensional sub-cellular mapping indicates the importance of iron to photosynthetic machinery and of zinc to nuclear organelles. Observed changes in diatom silicification and cytoplasm content following iron fertilization modify our understanding of the relationship between iron availability and silicification. These examples demonstrate the advantages of studying ocean biogeochemistry at the level of individual cells.     

Seasonal changes in sedimenting material collected by high aspect ratio sediment traps operated in a holomictic eutrophic lake

Cylindrical sediment traps with an aspect ratio (height (60 cm): diameter (5.1 cm)) of 11.8 were located 1 m above the surface sediment by a rigid metal framework support. Traps were exposed in Lough Neagh for one year, from May 1978. Each trap collected between 11 and 12 cm of faintly laminated sediment. One 12 cm sediment column was examined using conventional palaeolimnological techniques of core extrusion and analysis. The algal record in trapped sediment is shown to correspond with successional changes in phytoplankton abundance in the lake during the year of study. The sediment accumulation rate measured by the traps is an order of magnitude greater than that measured in dated sediment cores and redeposited and inwashed sediment formed the bulk of trapped material. However, the value of these high aspect ratio traps is that they provide a continuous but qualitative account of compositional changes in sedimenting material through time. Their potential as long term biological monitoring devices is emphasized.     

Physiological ecology of Mesozoic polar forests in a high CO2 environment

Fossils show that coniferous forests extended into polar regions during the Mesozoic, a time when models and independent paleo-CO2 indicators suggest that the atmospheric CO2 concentration was at least double that of the present day. Consequently, such polar forests would have experienced high CO2 interacting with an extreme variation in light. Here we describe an experiment investigating this plant-environment interaction for extant tree species that were important components of polar forests, and give results from the first year of treatment. Specifically, we tested the hypotheses that growth in elevated CO2 (1) stimulates photosynthesis; (2) reduces photoinhibition during the polar summer; and (3) reduces respiration of above- and below-ground plant organs. Our results indicate that CO2 fertilization generally does not affect photosynthesis under continuous daylight characteristic of the polar summer but does increase it when the period of illumination is shorter. Growth in elevated CO2 did not alter the potential for photoinhibition. CO2 enrichment significantly reduced leaf and root respiration rates by 50 and 25 %, respectively, in a range of evergreen taxa. Incorporating these observed CO2 effects into numerical simulations using a process-based model of coniferous forest growth indicates that a high paleo-CO2 concentration would have increased the productivity of Cretaceous conifer forests in northern Alaska. This results from decreased respiratory costs that more than compensate for the absence of high CO2-high temperature interactions during the polar summer. The longer-term effects of CO2 enrichment on seasonal changes in the above- and below-ground carbon balance of trees are discussed.     

Phytoplankton of four rivers,the Tyne,Wear, Tees and Swale

An account is given of the phytoplankton in four relatively fast-flowing rivers in north-east England associated with a water transfer scheme. The total number of live cells (or units) ranged from 2 × 10² to 2 × 10⁴ cells ml^–1. Input of plankton from a reservoir did not appear to influence densities at a site 25 km downstream. Diatoms formed the majority of cells in most samples, with green algae second and cryptomonads third in importance. Among the diatoms, centrics were relatively more abundant at downstream sites. The ratios of live versus dead diatoms showed marked differences with season and between centric and pennate forms.