Seasonal changes in the digestive enzyme levels of the amphipod Corophium volutator (Pallas) in relation to diet

The diet of the amphipod Corophium volutator (Pallas) living on extensive intertidal mud flats in the Bay of Fundy, eastern Canada, was investigated by direct and indirect methods. Microscopic observation of gut contents revealed macrophyte detritus but few diatom frustules. Laboratory cultures showed good survival on a diet of benthic diatoms, while on a diet of Spartina detritus survival was poor, although significantly better than the survival of starved animals. The ratios of gut enzyme activities, amylase to protease and amylase to laminarinase, changed markedly with the season. In summer the ratio was similar to that of animals cultured on a diet of benthic diatoms; in spring it resembled that of animals cultured on Spartina detritus, while in winter it was similar to that of starved animals. It is concluded that in summer, rapid growth is made possible by feeding on benthic diatoms, while Spartina detritus is an inferior food source which aids survival at other times of the year.     

Behavioural versus physiological photoprotection in epipelic and epipsammic benthic diatoms

Benthic diatoms are dominant primary producers in intertidal marine sediments, which are characterized by widely fluctuating and often extreme light conditions. To cope with sudden increases in light intensity, benthic diatoms display both behavioural and physiological photoprotection mechanisms. Behavioural photoprotection is restricted to raphid pennate diatoms, which possess a raphe system that enables motility and hence positioning in sediment light gradients (e.g. via vertical migration into the sediment). The main physiological photoprotection mechanism is to dissipate excess light energy as heat, measured as Non-Photochemical Quenching (NPQ) of chlorophyll fluorescence. A trade-off between vertical migration and physiological photoprotection (NPQ) in benthic diatoms has been hypothesized before, but this has never been formally tested. We exposed five epipelic diatom species (which move in between sediment particles) and four epipsammic diatom species (which live in close association with individual sand grains) to high light conditions, and characterized both NPQ and the relative magnitude of the migratory response to high light. Our results reveal the absence of a significant downward migratory response in an araphid diatom, but also in several raphid epipsammic diatoms, while all epipelic species showed a significant migratory response upon high light exposure. In all epipsammic species the upregulation of NPQ was rapid and pronounced; NPQ relaxation in low light conditions, however, occurred faster in the araphid diatom, compared with the raphid epipsammic species. In contrast, all epipelic species lacked a strong and flexible NPQ response and showed higher susceptibility to photodamage when not able to migrate. While overall our results support the vertical migration-NPQ trade-off, the lack of strong relationships between the capacity for vertical migration and NPQ within the epipsammic and epipelic groups suggests that other factors as well, such as cell size, substrate type and photoacclimation, may influence photoprotective strategies.     

Role of benthic diatoms in the implementation of the Urban Wastewater Treatment Directive in the River Wear, North-East England

The use of diatoms to monitor rivers in the UK was stimulated largely bythe requirements of the European Union's Urban Wastewater TreatmentDirective. The Trophic Diatom Index (TDI) was developed to assist theinterpretation of benthic diatom assemblages from UK rivers. This paperdescribes how data collected from one river, the River Wear in N-E England,sampled over a five-year period, was used to provide information about thepotential benefits of nutrient removal on major sewage treatment works.Althoughthe TDI indicated a clear increase in eutrophication with distance downstream,interpretation was confused by considerable within-site variability.Multivariate techniques such as TWINSPAN, when used alongside diatom indices,provide a more rigorous interpretation of longitudinal changes. The potentialfor using statistical power analysis to design sensitive monitoring programsusing benthic diatoms is also described.     

Frequency distribution studies of epipelic diatoms along an intertidal shore

Live and dead cell counts of epipelic diatoms were analysed at 16 sites along a transect crossing a saltmarsh, sandflat, and mudflat at Berrow Flats, Somerset, U.K. Replicate monthly samples were taken from January 1982 to June 1984. The ratio of dead to live cells varied according to position along the transect. Temporal changes of the ratios did not display a cyclic seasonal pattern. The clustered mosaic of live cells and empty frustules on surface sediments was quantitatively assessed. Graphing variance against seasonal mean cell counts showed that the variance was proportional to the square of the mean. The Neyman Type A equation is suggested as a possible model to explain this relationship.     

Short-term variations in particulate matter sedimentation off Kapp Norvegia,Weddell Sea,Antarctica: relation to water mass advection,ice cover,plankton biomass and feeding activity

Summary A multi-cup sediment trap was deployed at 250m in the shelf area off Kapp Norvegia, Weddell Sea (630 m water depth) to determine the relative importance of water mass advection, sea ice movement, phytoplankton biomass and plankton feeding. Short-term fluctuations in sedimentation were determined using a sampling frequency of 2.7 days over 54 days during January and February 1988. Three periods of enhanced sedimentation were associated with water mass exchange, settling of diatoms following break-up of ice cover and release of fecal matter by krill feeding on particulate matter derived from phytoplankton and ice algae. An initial sedimentation pulse (28 Jan) was mainly due to sinking pelagic diatoms and krill fecal strings containing algae released from sea ice passing over the trap position. The ¹³C-composition of the sedimented organic carbon was about-24. The isotope ratio decreased sharply by about 5.5 at the end of the first pulse indicating the source of sinking matter becoming pelagic diatoms of the retreating ice-edge. At this time the diatom Corethron criophilum contributed a very high proportion of the organic flux causing an increase of the opal/C_org ratios. The second pulse (6 Feb) was due to empty diatom frustules, minipellets and small planktonic aggregates. Much of the organic carbon was transported by round fecal pellets. During the third pulse (14 Feb), round fecal pellets transported even more; the percentage of C. criophilum to the diatom organic carbon flux was more than 80% (>2mg C m^–2 day^–1).Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation     

Effects of estuarine infauna on sediment stability and particle sedimentation

Secretions produced by the two estuarine benthic invertebrates Corophium volutator and Nereis diversicolor stabilise sediments by increasing their shear strength, and reduce sedimentation of previously resuspended particles. The secretions consist of 1 to 2 m threads which bind the particles together. They are used by both species in the production of complex burrow systems in the top 15 cm of sediment. The burrow systems have been demonstrated by a new resin impregnation technique. These results have widespread implications for the stability and erosion of estuarine sediments.     

Eukaryotic pathogens (Chytridiomycota and Oomycota) infecting marine microphytobenthic diatoms – a methodological comparison

Using sediment samples from the Solthörn tidal flat (southern North Sea, Germany), collected in bi‐weekly intervals from June to July 2012, a range of qualitative and quantitative screening methods for oomycete and chytrid pathogens infecting benthic diatoms were evaluated. Pre‐treatment of sediment samples using short ultrasound pulses and gradient centrifugation, in combination with CalcoFluor White, showed the best results in the visualization of both pathogen groups. The highest number of infected benthic diatoms was observed in mid July (5.8% of the total benthic diatom community). Most infections were caused by chytrids and, in a few cases, oomycetes (Lagenisma Drebes (host: Coscinodiscus radiatus Ehrenberg) and Ectrogella Zopf (hosts: Dimeregramma minor in Pritchard and Gyrosigma peisonis). Among the chytrids, sporangium morphology indicated the presence of five different morphotypes, infecting mainly epipelic taxa of the orders Naviculales (e.g., Navicula digitoradiata) and Achnanthales (e.g., Achnanthes brevipes Agardh). The presence of multiple pathogens in several epipelic diatom taxa suggests a significant role for fungal parasitism in affecting microphytobenthic diatom succession.     

Ultra‐sensitive immunoassay biosensors using hybrid plasmonic‐biosilica nanostructured materials

We experimentally demonstrate an ultra‐sensitive immunoassay biosensor using diatom biosilica with self‐assembled plasmonic nanoparticles. As the nature‐created photonic crystal structures, diatoms have been adopted to enhance surface plasmon resonances of metal nanoparticles on the surfaces of diatom frustules and to increase the sensitivity of surface‐enhanced Raman scattering (SERS). In this study, a sandwich SERS immunoassay is developed based on the hybrid plasmonic‐biosilica nanostructured materials that are functionalized with goat anti‐mouse IgG. Our experimental results show that diatom frustules improve the detection limit of mouse IgG to 10 pg/mL, which is ?100× better than conventional colloidal SERS sensors on flat glass.

Ultra‐sensitive immunoassay biosensor using diatom biosilica with self‐assembled plasmonic nanoparticles.     

Diatoms of the Peatbog Sediments from Shemya Island (Aleutian Islands,United States)

Results of a study of diatoms from Holocene peatbog sediments from Shemya Island (Aleutian Islands, United States) are presented. The column of peat sediments (385 cm depth) was investigated (formation of peat sediments began more than 9500 years ago). Sixty-seven taxa of 31 genera, 17 families, eight orders, and three classes were identified. According to results of taxonomical and eco-geographical analyses, most taxa belong to the order Naviculales and family Pinnulariaceae and are benthic organisms with cosmopolitan distribution. Forms with different quality of frustule preservation were revealed. Centric diatoms demonstrate the best preservation in sediment. Of special interest is the dissolution of frustules in a water body with presumably low pH value. Dynamics of the pattern of species relative abundance was studied. Several zones with characteristic diatom complexes are distinguished. Analysis of distribution of diatoms in the column showed that, apparently, this locality was a shallow oligotrophic reservoir with a relatively low pH in the past. Water level changed occasionally, but trophic level did not change over the period of the reservoir’s existence.     

Spatial patterns of benthic diatoms,carbohydrates and mud on a tidal flat in the Ems-Dollard estuary

The chlorophyll a content and two operational fractions of carbohydrate (water extractable and EDTA extractable) were measured every three months during one year along transects on a tidal flat in the Ems-Dollard estuary (The Netherlands). Chlorophyll a was used as an indicator of microphytobenthos biomass, which was composed predominantly of epipelic diatoms. Both carbohydrate fractions correlated significantly with chlorophyll a. EDTA extractable carbohydrates were more resistant towards degradation than the water extractable fraction. During most of the year, concentrations of chlorophyll a and carbohydrates were low, but in June, high concentrations of up to 90 g chlorophyll a/g sediment were found in a narrow zone running parallel to the channel. Maximum concentrations of water extractable carbohydrates and EDTA extractable carbohydrates ranged between 800–1200 and 600–800 g/g sediment, respectively. The mud content was high (± 90%) at the margin of the tidal flat. This was not limited to the growth season of the diatoms, but was observed throughout the year. This indicated that the high mud content at the mudflat margin was mainly caused by hydrodynamic factors, and not by biostabilization. In June, exceptionally high diatom densities were found in sediment with a high mud content. There was only minor evidence that biostabilization by epipelic diatoms lead to a further increase in the mud content of the sediment.     

Distribution of benthic diatoms in relation to environmental variables in lowland streams

The composition (% relative abundance) of diatom assemblages from soft bottom sediments was studied at 75 sites situated in 46 rivers, brooks, and ditches in the islands of Hiiumaa and Saaremaa and in the lowland of West Estonia. Although the total number of recorded taxa was 205, the studied diatom assemblages consisted of 54, 55, 48, and 50 constant species in the drainage basin of Moonsund, the Gulf of Riga, Hiiumaa, and Saaremaa, respectively. The habitats of the dominating taxa were heterogenous and the most widespread species were Achnanthidium minutissimum, Martyana martyi, Meridion circulare, Cocconeis placentula, Planothidium lanceolatum, and Amphora pediculus. The Shannon–Weaver diversity (H) index varied from 2.09 to 4.63. Multivariate analyses were used to identify the environmental variables governing the composition and structure of the benthic diatom assemblage. Principal Component Analysis (PCA) and Redundancy Analysis (RDA), based on 56 most abundant taxa, indicated differences in the composition and structure of diatom assemblages between different drainage basins as well as between the upper and lower stream courses. In the headwaters there prevailed small epipsammic diatoms (Martyana, Planothidium, Staurosira, Staurosirella). Different motile epipelic species from the genera Amphora, Navicula, Nitzschia, etc. were distributed abundantly in the lower courses of the streams. There was a positive correlation between order of the stream site and trophic level of water (R=0.35; p<0.05). Along a river system, the increasing order of the stream was accompanied by higher trophic level of water.     

Predicting Epipelic Diatom Exopolymer Concentrations in Intertidal Sediments from Sediment Chlorophyll a

Abstract In many intertidal cohesive—sediment habitats, epipelic diatoms are the dominant microphytobenthic organisms. In such sediments, concentrations of colloidal carbohydrate [including the exopolymeric substances (EPS) produced by diatoms during motility] are closely correlated with the biomass (chlorophyll a) of epipelic diatoms. A model describing this relationship (log (conc. coll. carbo. + 1) = 1.40 + 1.02(log (chl. a conc. + 1)) was derived from published data. It was validated against published and unpublished data from 6 different estuaries, and accounted for 64.6% of the variation in sediment colloidal carbohydrate concentrations. The model was valid for intertidal habitats with cohesive sediments where epipelic diatoms constituted >50% of the microphytobenthic assemblage. In sites with noncohesive sediments, or where the microphytobenthic assemblage was dominated by other algal groups, the model was not applicable. The mean percentage of EPS in colloidal carbohydrate extracts varied between 11 and 37% for axenic cultures of epipelic diatoms (with higher values obtained during stationary phase), and between 22.7% and 24.3% for natural sediments dominated by epipelic diatoms. Assuming an EPS percentage of 25% in colloidal extracts yielded an EPS chl. a ratio of 2.62:1. Maximum rates of EPS production in diatom cultures occurred at the beginning of stationary phase (1.6–5.09 μg EPS μg⁻¹ chl a d⁻¹), with Nitzschia sigma having a significantly (P < 0.05) higher rate of production than N. frustulum, Navicula perminuta and Surirella ovata. Similar rates of EPS production were measured in the field. The dynamics of EPS production and loss on mudflats is discussed, with reference to the model and these production rates. Received: 25 February 1997; Accepted: 23 May 1997     

Continuous culture with deep seawater of a benthic food diatom Nitzschia sp.

A continuous culture system for a benthic food diatom Nitzschia sp. wasestablished by using properties of high nutrient and clean of deep seawater(DSW). DSW collected from 320 m depth in Muroto City, Japan, was introducedinto a glass-pipe bioreactor (14 cm length, 3 cm diam.) containing glassbeads of 0.5 cm diam. as substrata for the alga, and it was incubated at18°C · 80Em^–2sec^–1 · L:D=14:10. The chlorophyll a yield of benthicdiatoms in a reactor as a unit of surface area of the substratum was only0.001–0.003 g cm^–2 when the flow rate of DSW was 0(batch culture conditions). However, when DSW was supplied continuously to areactor, the yield increased to 1.4 g-chl.a cm^–2 alongwith the increase in flow rate of DSW. Moreover, amounts of chl.a washed outof the system were negligible, 0.0014 to 0.0045%, even though theflow rate of DSW was as much as 25 times h^–1, suggesting thatsloughing of benthic diatoms from the substratum was minimized. Although theyield of diatoms fluctuated significantly at the time that the DSW wascollected, the variation could be minimized by increasing the flow rate ofDSW. These results indicate that the continuous culturing system with DSWsupports the stable and effective mass culture of benthic food diatom.     

Diatom elemental and morphological changes in response to iron limitation: a brief review with potential paleoceanographic applications

Diatoms are a major group of phytoplankton that account for approximately 40% of the ocean carbon fixation and the vast majority of biogenic silica production through the construction of their cell walls (termed frustules). These frustules accumulate and are partially preserved in the ocean sediments. Diatom growth and nutrient utilization in high‐nitrate, low‐chlorophyll regions of the world’s oceans are mostly regulated by iron availability. Diatoms acclimate to iron limitation by decreasing cell size. The associated increase in surface area‐to‐volume ratio and decrease in diffusive boundary layer thickness may improve nutrient uptake kinetics. In parallel, cellular silicon (Si) contents are elevated in iron‐limited diatoms relative to nitrogen (N) and carbon (C). Variations in degree of silicification and nutritional requirements of iron‐limited diatoms have been hypothesized to account for higher cellular Si and/or lower cellular N and C, respectively. However, in some diatoms, frustule silicification does not significantly change when cells are iron‐limited. Instead, changes in the Si‐containing valve surface area relative to volume within these diatoms is hypothesized to be responsible for the variations in the cellular Si : N and Si : C ratios. In particular, some examined iron‐limited pennate diatoms have reduced widths relative to their lengths (i.e. lower length‐normalized widths, LNW) compared to iron‐replete cells. In the pennate diatom Fragilariopsis kerguelensis, the mean LNWs of valves preserved in sediments throughout the Southern Ocean (a well‐characterized iron‐limited region) is positively correlated with satellite‐derived, climatological net primary productivity in the overlying waters. Because of the specific morphological changes in pennate diatom frustules in response to iron availability, the valve morphometerics (e.g. LNWs) can potentially be used as a diagnostic tool for iron‐limited diatom growth and relative changes in the Si : N (and Si : C) ratios in extant diatom assemblages as well as those preserved in the sediments.     

Phytoplankton distribution in relation to sea ice,hydrography and nutrients in the northwestern Weddell Sea in early spring 1988 during EPOS

Summary Phytoplankton biomass and distribution of major phytoplankton groups were investigated in relation to sea ice conditions, hydrography and nutrients along three north-south transects in the north western Weddell Sea in early spring 1988 during the EPOS Study (European Polarstern Study), Leg 1. Three different zones along the transects could be distinguished: 1) the Open Water Zone (OWZ) from 58° to 60°S with high chlorophyll a concentrations up to 3.5 g l^–1; 2) the Marginal Ice Zone (MIZ) from 60° to about 62.5° with chlorophyll a concentrations between 0.1 and 0.3 g l^–1, and 3) the closed pack-ice zone (CPI) from 62.5° to 63.2°S with chlorophyll a concentrations below 0.1 gl^–1. Nutrient concentrations increased towards the south showing winter values under the closed pack-ice. Centric diatoms such as Thalassiosira gravida and Chaetoceros neglectum forming large colonies dominated the phytoplankton assemblage in terms of biomass in open water together with large, long chain forming, pennate diatoms, whereas small pennate diatoms such as Nitzschia spp., and nanoflagellates prevailed in ice covered areas. Fairly low concentrations of phytoplankton cells were encountered at the southernmost stations and many empty diatom frustules were found in the samples. The enhanced phytoplankton biomass in the Weddell-Scotia-Confluence area is achieved through sea ice melting in the frontal zone of two different water masses, the Weddell and the Scotia Sea surface waters.Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation     

Influence of bacteria and salinity on diatom biogenic silica dissolution in estuarine systems

Dissolution of diatom biogenic silica (bSiO₂) in estuaries and its control by water salinity and bacteria were investigated using the river euryhaline species Cyclotella meneghiniana as a model. Laboratory-controlled bioassays conducted at different salinities with an estuarine bacteria inoculum showed a faster dissolution of diatom bSiO₂ at the lowest salinity where bacteria were the most abundant. However in another experiment, salinity increase clearly enhanced the dissolution of cleaned frustules (organic matter free). The presence of active bacteria might therefore predominate on the effect of salinity for freshly lysed diatoms whereas salinity might rather control dissolution of organic-matter-free frustule remains. Incubation of cultivated diatoms at different protease concentrations revealed that high proteolytic activities had little effect on bSiO₂ dissolution at a 1-month scale in spite of an efficient removal of organic matter from the frustules. Altogether it is hypothesized that bacterial colonization increases bSiO₂ dissolution by creating a microenvironment at the diatom surface with high ectoproteolytic activity but also via the release of metabolic byproducts since the presence of organic matter seems generally to facilitate diatom bSiO₂ dissolution.     

Auxosporulation in Paralia guyana MacGillivary (Bacillariophyta) and Possible New Insights into the Habit of the Earliest Diatoms

Background

Diatoms are one of the most ecologically important aquatic micro-eukaryotes. As a group unambiguously recognized as diatoms, they seem to have appeared relatively recently with a limited record of putative remains from oldest sediments. In contrast, molecular clock estimates for the earliest possible emergence of diatoms suggest a considerably older date. Depending on the analysis, Paralia and Leptocylindrus have been recovered within the basal molecular divergences of diatoms. Thus these genera may be in the position to inform on characters that the earliest diatoms possessed.

Findings

Here we present auxospore development and structure of initial and post-auxospore cells in a representative of the ancient non-polar centric genus Paralia. Their initial frustules showed unusual, but not unprecedented, spore-like morphology. Similarly, initial frustules of Leptocylindrus have been long considered resting spores and a unique peculiarity of this genus. However, even though spore-like in appearance, initial cells of Paralia readily resumed mitotic divisions. In addition, Paralia post-auxospore cells underwent several rounds of mitoses in a multi-step process of building a typical, “perfect” vegetative valve. This degree of heteromorphy immediately post-auxosporulation is thus far unknown among the diatoms.

Implications

A spore-related origin of diatoms has already been considered, most recently in the form of the “multiplate diploid cyst” hypothesis. Our discovery that the initial cells in some of the most ancient diatom lineages are structurally spore-like is consistent with that hypothesis because the earliest diatoms may be expected to look somewhat similar to their ancestors. We speculate that because the earliest diatoms may have appeared less diatom-like and more spore-like, they could have gone unrecognized as such in the Triassic/Jurassic sediments. If correct, diatoms may indeed be much older than the fossil record indicates, and possibly more in line with some molecular clock predictions.     

Seasonal changes in silicon content of diatoms estimated from the ratio of particulate silicon to diatom volume under silicon sufficiency in diatom-rich Lake Barato

To clarify the changes in Si content of diatoms, the particulate silicon (PSi) concentration and total diatom volume (TDV) were determined in Lake Barato, Japan, from April to July 1998–2000. The soluble reactive silicon (SRSi) concentration decreased markedly with the rapid increase in TDV in May and June in all three years, although the value did not fall below that at which diatom growth might be limited. The proliferation of small discoid diatoms contributed to the decrease in SRSi concentration each year. The Si content of diatoms may not be constant as indicated by the changes in PSi:TDV ratio. The low PSi:TDV ratio and the fact that PSi concentration was lower than diatom PSi concentration (calculated from the volume of diatom species) accompanying the decrease in TDV suggests the possibility of a disturbance in the silicification in May and June 1999. These parameter changes accompanying the increase in TDV suggest that the silicification did not catch up with the cell division in early April 1998, early May 1999, and mid-June 2000. In addition, the PSi:TDV ratio increased rapidly and showed large fluctuations in July 1998 and 1999. This may have been caused by a change in dominant species from small discoid diatoms to Aulacoseira granulata because of the differences in Si content per unit cell volume.     

Morphology and ultrastructure of teratological forms of the diatoms Stephanodiscus niagarae and S. parvus (Bacillariophyceae) from Hamilton Harbour (Lake Ontario,Canada)

Teratological forms of Stephanodiscus niagarae Ehrenb. and S. parvus Stoermer & Håkansson were observed during a study of diatoms preserved in a radiometrically dated core from Hamilton Harbour (Lake Ontario, Canada). Morphological features and ultrastructures of both species were studied under the light and scanning electron microscope. The valve structure of abnormal forms of S. niagarae appears to be weakly silicified, especially in the central area. The shape of satellite pores are very irregular in comparison with the round shape of the normal specimen. The central fultoportulae are characterized as small tubes extending out of the external valve. Two types of abnormal frustules are present in the population of S. parvus studied. In one type valves are underdeveloped and only the siliceous layer and ribs are present, and in the other type the valves are strongly silicified and the areolae are almost completely occluded.The teratological forms of both species appeared in the core sediments after 1911, and both became the dominant components of diatom assemblages after 1970. Their occurrence and increased abundance coincides with records of heavy metal pollution in the harbour.     

Gonadal neoplasms in wild carp-goldfish hybrids from the Welland River near Niagara Falls,Canada

An experimental approach was taken to examine the processes of detritus decomposition in river sediments.Addition of macrophyte detritus (Alternanthera philoxeroides (Mart.) Griseb) to river sediment resultedin an increase in carbon mineralization from a sbasal rate of 200 up to 500 mg C m^–2 d^–1.Carbon mineralization after addition of oak detritus was only slightly higher than mineralization in sediments thatreceived no addition ( 200 mg C m^–2 d^–1). Bacterial biomass and production in sediments \s+ alligator-weedwere higher than in sediments + oak or zero-addition. In these experiments the major fate of addedalligatorweed was mineralization. For alligatorweed detritus, microbial metabolism depletes organic carbonrather than leading to increases in food quality. Therefore, a pulse input of alligatorweed detritus would notbe available as a long-term source of organic carbon. Oak detritus was not rapidly decomposed, and sopersists in these sediments for a longer period.