FACTORS LIMITING EDAPHIC ALGAL BIOMASS AND PRODUCTIVITY IN A GEORGIA SALT MARSH1,2

A “planted core” system was developed to test the effect of short term (1–2 weeks) experimental manipulation of environmental parameters on edaphic microalgae under field conditions. A large number of small cores (surface area = 7 cm²) were collected, randomized and replanted in the marsh in fiddler crab exclosures with appropriate experimental treatments. Daily enrichment of the cores with NH₄⁺ resulted in significant increases in edaphic primary productivity and levels of chlorophyll a in both summer and winter seasons in the short-Spartina marsh. Enrichment with a complete nutrient solution caused no further increases. Nutrient enrichment of creekbank sediments was much less stimulatory to the resident algal assemblage. In both sites, but especially in the creekbank, the removal of fiddler crab grazers resulted in significant increases in chlorophyll a and productivity. Experimental manipulation of light intensity showed that the average light intensity reaching the sediment surface was saturating for chlorophyll production in the short-Spartina marsh. A reciprocal transplant experiment involving unfertilized cores from the short-Spartina marsh and creekbank marsh demonstrated that NH₄⁺ inputs occurring in the creekbank site rapidly alleviated nitrogen limitation of edaphic algae from short-Spartina marsh. Algae in creekbank cores incubated in the short-Spartina marsh were unable to sustain high productivity once the original standing stock of NH₄⁺ declined.     

Concentration and distribution of chlorophyll in fertilized plots in a Massachusetts salt marsh

The amount of algal chlorophyll on the mud surface of a salt marsh was not affected directly by additions of urea or sewage sludge fertilizer but was proportional to light reaching the surface. Fertilization affected the algae by increasing the standing crop of grass and theregy reducing the light underneath. Vertical distribution of total chlorophyll depended on the species and biomass of grass present. Algal chlorophyll was concentrated at the mud surface and horizontally was very patchy. Locally the quantity exceeded the amount found in other productive marine ecosystems.     

Effects of nitrogen,phosphorus and carbon enrichment on planktonic and periphytic algae in a softwater,oligotrophic lake in Florida,USA

The objective of this study was to investigate nutrient limitation of algal abundance in Anderson-Cue Lake, a softwater clear oligotrophic lake in north-central Florida. Nutrient diffusing clay pots and cylindrical enclosures were used in the field to test effects of different combinations of nitrogen, phosphorus, silica, and carbon on algal standing crop and composition of periphytic and planktonic algae, respectively. Effects of nutrient enrichment on periphytic algae were examined in two studies conducted 31 May – 8 July and 10 June – 15 July 1991. Nutrient effects on planktonic algae were examined in one study from 13 June – 1 July 1991. Planktonic and periphytic algal biovolume was significantly higher (p<0.05) when nitrogen and carbon were added in combination than with treatments without nitrogen, carbon, or nitrogen and carbon. Treatments with nitrogen and carbon combined resulted in lower algal diversity and dominance by coccoid green algae andScenedesmus. Results indicate that carbon and nitrogen can be limiting factors to algal growth in Anderson-Cue Lake and possibly other lakes of similar water quality.     

LONG-TERM EFFECTS OF MANIPULATING LIGHT INTENSITY AND NUTRIENT ENRICHMENT ON THE STRUCTURE OF A SALT MARSH DIATOM COMMUNITY1,2

The long-term effects of manipulating light intensity and nutrient enrichment on the structural characteristics of a diatom community inhabiting the sediments beneath a pure stand of dwarf Spartina alterniflora Loisel. were investigated over a yearly cycle. Clipping or shading the cord grass cover, or phosphorus enrichment caused significant decreases in both species diversity (H') and the number of diatom species, whereas nitrogen enrichment only significantly decreased the latter parameter. Of the 105 diatom taxa identified, only 10 were restricted to certain of the 12 study areas; and of these, 8 occurred exclusively in the clipped habitats. An analysis of variance (light × nutrient × collection date) involving 19 of the most abundant taxa revealed that certain experimental treatments had significant effects on the relative abundances of each and every taxon. However, attempts to group taxa with similar response patterns proved unsuccessful because of the frequent significance of the 3-way interaction term. Synthesis of these results with earlier work by the author showed that differences in structure of diatom communities inhabiting the sediments beneath the 3 dominant marsh grasses were not primarily caused by differences in reduction of light intensity by their grass canopies, and that clipping of the cord grass produced a shift in community structure towards that characteristic of a salt panne algal mat.     

Seasonal succession of planktonic and epiphytic algae in a canal in Southern England

The standing crop of phytoplankton in a canal in southern England remained low during 1973 and 1974, seldom exceeding 5 × 10⁴ cells/1. Since phosphorus, nitrogen and silicon occurred abundantly in the water, competition with higher plants for some other substance must have limited development. Although the standing crop of epiphytic algae associated withNasturtium officinale andGroenlandia densa seemed to be limited by the number of attachment sites, this factor was of little importance in the case of algae attached toCladophora glomerata. Achnanthes minutissima v.cryptocephala was always predominant in the epiphytic assemblages, representing 50–8o% by numbers of the flora. The limited pool of predominant epiphytic taxa may have restricted the communities' ability to adapt to fluctuations in environmental conditions. The grazing of isopods, amphipods and molluscs probably never limited algal densities.     

PRIMARY PRODUCTION OF EDAPHIC ALGAL COMMUNITIES IN A MISSISSIPPI SALT MARSH1

Primary production rates of edaphic algae associated with the sediments beneath four monospecific canopies of vascular plants were determined over an annual cycle in a Mississippi salt marsh. The edaphic algal flora was dominated by small, motile pennate diatoms. Algal production (as measured by ¹⁴C uptake) was generally highest in spring-early summer and lowest in fall. Hourly rates ranged from a low of 1.4 mg C/m² in Juncus roemerianus Scheele to a high of 163 mg C/m² beneath the Scirpus olneyi gray canopy. Stepwise multiple regressions identified a soil moisture index and chlorophyll a as the best environmental predictors of hourly production; light energy reaching the marsh surface and sediment and air temperature proved of little value. Adding the relative abundances of 33 diatom taxa to the set of independent variables only slightly increased R²; however, virtually all variables selected were diatom taxa. R² was only 0.38 for the Spartina alterniflora Loisel. habitat but ranged from 0.70 to 0.87 for the remaining three vascular plant zones. Annual rates of algal production (g C/m²) were estimated as follows: Juncus (28), Spartina (57), Distichlis spicata (L.) Greene (88), and Scirpus (151). The ratio of annual edaphic algal production to vascular plant net aerial production (EAP / VPP) was 10–12% for the first three habitats and 61% for Scirpus. Chlorophyll a concentrations, annual algal production rates, and EAP / VPP values were comparable to those determined in Texas, Delaware, and Massachusetts salt marshes but lower than those reported for Georgia and particularly California marshes.     

ECOLOGY OF THE EPIPELIC ALGAL COMMUNITIES IN MARION LAKE,BRITISH COLUMBIA1,2

Studies of the seasonal and spatial distribution of the epipelic algal standing crop and primary productivity were conducted in Marion Lake, British Columbia. Possible biological, chemical, and physical factors controlling the epipelic algal community dynamics were also investigated. The epipelic algal flora of the lake was very diverse, however, it can be generally considered as acidophilic associations of algae. The vertical distribution of the epipelic algae is partially influenced by the amount of light reaching the sediment, but also strongly influenced by the grazing of animals and erosion by wave action. Temperature, light, and grazing by animals all appear to influence the seasonal fluctuations in the algal standing crop. Concentrations of nutrients immediately above the sediment surface appear to be less important as controlling factors. The most important variables influenceing the primary productivity of the epipelic community are temperature, total algal standing crop, and light. Nutrients, again, appear to be less important as controlling factors. The study supports the idea that epipelic algal growth is high in shallow, low nutrient lakes and that the epipelic algal productivity is extremely important to the total energy budget of the lake.     

Predaceous insects may limit algal grazers in high-nutrient environments: implications for wastewater remediation in open bioreactors

Algal biofuel has potential as a source of renewable fuel and a tool for wastewater remediation. Open algal bioreactors fertilized with wastewater can have net energy gain but are vulnerable to colonization by algal grazers. However, colonizing predaceous insects may limit grazer impacts on algae. Here, we investigate the effects of grazers, predators, and invading algae species on algal production and community structure in high-nutrient environments. First, we grew diverse algal assemblages in treated municipal wastewater in a greenhouse with Daphnia grazers and different insect predators that were added experimentally. When Daphnia were present without predators, they eliminated suspended algae. But, dragonfly larvae [Odonata: Libellulidae] and backswimmers [Hemiptera: Notonectidae], but not larval diving beetles [Coloeoptera: Dytiscidae], suppressed Daphnia allowing suspended algae to persist. Second, we grew Chlorella algae in field tanks that were open or protected from natural invertebrate colonization and half the tanks received wild-collected plankton in a factorial design. Mosquito larvae [Culex sp.] readily colonized open tanks and reduced algal mass and dissolved phosphorus concentrations. Colonist addition to open tanks shifted algal functional and taxonomic composition but did not impact suspended algal production. Our study indicates that large numbers of grazer individuals can rapidly colonize open bioreactors. Experimentally added and naturally colonizing grazers altered algal community structure and reduced algal standing crops but may also aid in nutrient removal from wastewater-fed bioreactors. Effective operation of open algal bioreactors must consider cultivated algae species’ vulnerability to competition and local grazers as well as the ability of potential predators to both naturally disperse into bioreactors and to control grazers.     

DISTRIBUTION OF BLUEGREEN ALGAE IN A MISSISSIPPI GULF COAST SALT MARSH1

Edaphic bluegreen algal communities were sampled from five menotypic angiosperm zones in Grvelline Bay Marsh near Ocean Spring. Mississippi. Samples and environmental data were taken on a quarterly basis from October 1976 to June 1977 beneath the following marsh angiosperms: Distichlis spicata (L.) Green, Scirpus olneyi gray, SPartina patens (Aiton) Muhl., Sp. alterniflora Loisel., and Juncus roemerianus Scheele. Communities in all five zones were dominated by Schizothrix calcicola (Ag.) Gom. (sensu Drauet) throughout the study, while the subdominant bluegreens varied according to season. The number of individuals in all zones was greatest in the summer and lowest in the winter. An examination of the structure of the five edaphic communities indicated a single, nearly homogeneous community exists over the entire surface of the marsh shaded by a angiosperm canopy. Light intensity appears to be the major factor affecting the distribution of bluegreen algae in this salt marsh.     

The Availability of Condensed Phosphates for the Algal Strain Scenedesmus obtusiusculus Chodat,with Special Regard to Conditions Characteristic in Lake Balaton

The limiting factor in the water of Lake Balaton was calculated by means of Verduin's equation. If only the orthophosphate-phosphorus concentration is inserted into the equation, phosphorus is the primary limiting factor. If, however, total phosphorus is considered, the factor intensity of nitrogen will be the least, i. e. nitrogen will be the primary limiting-factor of plant growth. According to measurements during 1976, average values for the total dissolved phosphorus and orthophosphate-phosphorus content of Balaton Lake water were 15,66 mg/m³ and 7.66 mg/m³, respectively. Experiments with the algal strain Scenedesmus obtusiusculus Chod. (Chlorophyceae, Chlorococcales). were designed to test the availability of the condensed phosphorus form for the algae. The experiments were performed partly with synthetic polyphosphates, partly, with polyphosphates isolated from Balaton Lake water. The results showed, contrary to our expectations, that phosphates present in the condensed form (irrespective of their structure and degree of condensation). were not utilized by the algae under sterile conditions, i. e. in the absence of bacterial activity. In the light of the above it is recommended to consider only orthophosphate-phosphorus when calculating the limiting factor.     

A comparison of algal,macroinvertebrate, and fish assemblage indices for assessing low-level nutrient enrichment in wadeable Ozark streams

Biotic indices for algae, macroinvertebrates, and fish assemblages can be effective for monitoring stream enrichment, but little is known regarding the value of the three assemblages for detecting perturbance as a consequence of low-level nutrient enrichment. In the summer of 2006, we collected nutrient and biotic samples from 30 wadeable Ozark streams that spanned a nutrient-concentration gradient from reference to moderately enriched conditions. Seventy-three algal metrics, 62 macroinvertebrate metrics, and 60 fish metrics were evaluated for each of the three biotic indices. After a group of candidate metrics had been identified with multivariate analysis, correlation procedures and scatter plots were used to identify the four metrics having strongest relations to a nutrient index calculated from log transformed and normalized total nitrogen and total phosphorus concentrations. The four metrics selected for each of the three biotic indices were: algae—the relative abundance of most tolerant diatoms, the combined relative abundance of three species of Cymbella, mesosaprobic algae percent taxa richness, and the relative abundance of diatoms that are obligate nitrogen heterotrophs; macroinvertebrate—the relative abundance of intolerant organisms, Baetidae relative abundance, moderately tolerant taxa richness, and insect biomass; fish—herbivore and detritivore taxa richness, pool species relative abundance, fish catch per unit effort, and black bass (Micropterus spp.) relative abundance.All three biotic indices were negatively correlated to nutrient concentrations but the algal index had a higher correlation (rho = ?0.89) than did the macroinvertebrate and fish indices (rho = ?0.63 and ?0.58, respectively). Biotic index scores were lowest and nutrient concentrations were highest for streams with basins having the highest poultry and cattle production. Because of the availability of litter for fertilizer and associated increases in grass and hay production, cattle feeding capacity increases with poultry production. Studies are needed that address the synergistic effect of poultry and cattle production on Ozark streams in high production areas before ecological risks can be adequately addressed.     

The role of light availability and herbivory on algal responses to nutrient enrichment in a riparian wetland,Alaska

We investigated how the relative availability of solar radiation in the presence or absence of grazing alters the ability of benthic algae to respond to nutrient enrichment in an Alaskan marsh. We used a factorial mesocosm experiment that included nutrient enrichment (enriched or control), grazing (grazed or ungrazed), and light (unshaded or shaded) to simulate shading by macrophytes early and late in the growing season, respectively. We found stronger effects of grazers and nutrients compared to light on benthic algal biomass and taxonomic composition. Algal biomass increased in nutrient‐enriched treatments and was reduced by grazing. Shading did not have an effect on algal biomass or taxonomic composition, but the concentration of chl a per algal biovolume increased with shading, demonstrating the ability of algae to compensate for changes in light availability. Algal taxonomic composition was more affected by grazer presence than nutrients or light. Grazer‐resistant taxa (basal filaments of Stigeoclonium) were replaced by diatoms (Nitzschia) and filamentous green algae (Ulothrix) when herbivores were removed. The interacting and opposing influences of nutrients and grazing indicate that the algal community is under dual control from the bottom‐up (nutrient limitation) and from the top‐down (consumption by herbivores), although grazers had a stronger influence on algal biomass and taxonomic composition than nutrient enrichment. Our results suggest that low light availability will not inhibit the algal response to elevated nutrient concentrations expected with ongoing climate change, but grazers rapidly consume algae following enrichment, masking the effects of elevated nutrients on algal production.     

On the nitrogen and phosphorus removal in algal photobioreactors

This study shows results of nitrogen and phosphorus removal by microalgae (tertiary treatment) in a prototype of tubular photobioreactor tested under controlled and uncontrolled conditions. The wastewater was the supernatant coming from a secondary settler of a municipal wastewater activated sludge treatment plant without nitrification and denitrification units. The algal biomass was directly selected from the supernatant and it was principally composed of genus Scenedesmus (autochthonous algae). All the experiments evaluated both nitrogen and phosphorus removal and biomass and lipid production. A satisfactory nutrients removal - about 99.9% for the nitrogen and phosphorus - and a specific biomass productivity of 0.25 g/l d have been obtained in the indoor photobioreactor; less satisfactory results have been reached in the outdoor photobioreactor because of ambient condition instability and limiting nutrients concentration.     

Bioassays with Stigeoclonium Kütz. (Chlorophyceae) to identify nitrogen and phosphorus limitations

Eleven strains of the filamentous algae genus Stigeoclonium Kütz. (Chlorophyceae) were tested as bioassay organisms to identify nitrogen and phosphorus limitations to growth in a variety of waters. The assay results were related to the nutrient status of the waters during the year. Optimal growth was obtained at an inorganic phosphorus concentration of 0.65 mg P 1^?1 and an inorganic nitrogen concentration of 3.75 mg N 1^?1. The algal growth potentials were generally higher in November and February than in August. In summer, nitrogen became important as a limiting nutrient. When ratios of inorganic nitrogen to inorganic phosphorus (N/P) were above 6.2, phosphorus was primarily limiting and at lower N/P ratios nitrogen became the important limiting factor. Stigeoclonium proved to be a suitable bioassay organism to identify nutrient limitation in freshwaters.     

Filamentous algal communities related to pH in streams in The Trossachs, Scotland

• 1 A survey of acidified streams in the Loch Ard area of The Trossachs, Scotland, was carried out between 1986 and 1988, to determine the influence of physical and chemical factors on the distribution of benthic algae. Samples were taken on thirty-five occasions from fifteen sites on ten streams. Forty-nine operational taxa of filamentous algae were distinguished. Relative abundance of taxa in samples was scored on a seven-point scale. Estimates of standing crop were made by pigment extraction and ash-free dry weight (AFDW) determination from biomass developed on artificial substrata.
• 2 No evidence was found for seasonality in standing crop, nor for an increase in standing crop with a decrease in pH. Seasonality in relative abundance was evident for few taxa. Species richness and diversity were highly correlated with pH and correlated chemical variables.
• 3 Canonical correspondence analysis (CCA) carried out using the CANOCO program ordinates species and sites and relates them directly to environmental variables. The most important variable was found to be pH (or an acidification-related variable that is highly correlated with pH), with percentage forest cover and total oxidized nitrogen being of secondary importance. The effect of subdividing the data set, using data based on presence-absence as well as estimated relative abundance, was tested. A comparison of the effects on the distribution of site and species values on the pH vector showed that the results are robust, confirming the value of this semi-quantitative sampling method for use with a difficult algal group. It was possible to infer pH from algal community structure using CANOCO.
• 4 The pH ranking of species derived from CCA revealed that larger cell diameter taxa do not predominate at low pH, implying that reduction of invertebrate grazing was not an important determinant of algal community composition.

     

Nutrient concentrations in a Littorella uniflora community at higher CO2 concentrations and reduced light intensities

1. Oligotrophic softwater lakes represent a special type of aquatic ecosystem with unique plant communities where generalisations from other aquatic plant communities to rising CO₂ in the water column may not apply. 2. In the present study, we set up large in situ mesocosms and supporting laboratory experiments with isoetid vegetation (Littorella uniflora) where water column CO₂ and light could be manipulated in order to test whether (i) light and CO₂ availability affect nutrient concentrations in isoetid vegetation, and (ii) if changes in light and CO₂ climate affect fluxes of inorganic nitrogen (N) and phosphorus (P) from sediment to water column, which potentially could result in increased growth of epiphytic algae. 3. The results showed that the standing stocks of phosphorus and nitrogen in the L. uniflora vegetation were significantly influenced by CO₂ concentration and light intensity. Both standing stocks of P and N were significantly higher in the mesocosm treatments with high CO₂ concentration than in those at low CO₂ concentration. Similarly, standing stocks of P and N enhanced with increasing light intensity. 4. Measurements of nutrient fluxes both in the field and the laboratory did not show any significant release of nutrients to the water column from plants or sediments at any of the light or CO₂ treatments. However, mats of epiphytic algae developed from the beginning of June to late September and caused a light reduction for the isoetid vegetation. 5. Increasing CO₂ concentrations in the water column may over time potentially result in a change in soft water plant communities.     

An Analysis of the Nutrient Contribution by Phytoplankton Primary Production to the Food Web of a Small Lake

Phytoplankton primary production was measured for 148 days, from May 31 to October 25, 1971, in Sunfish Lake, Ontario, using the ¹⁴C light and dark bottle technique. The purpose of the experiment was not only to measure the rate of production but also to give some indication of the fates of the material produced by the algae. The following data were calculated for the sampling season: total gross production 57,770.6 mg C m ⁻²; algal respiration 31.74% of gross production; total net production 39,895.4 mg C m ⁻²; extracellular production 9.2% of total net production. The efficiencies of both gross and net total production were inversely correlated with light intensity. The rate of extracellular production was directly correlated with the rate of total production. The rate of respiration was not significantly correlated with either the rate of gross production or with the algal biomass as calculated from differential cell counts. The weekly rate of phytoplankton turnover was calculated by combining measurements of algal biomass with net particulate production measurements. The turnover rates for the phototrophic zone ranged from plus 6,788.5 mg C m ⁻² to minus 1,104.4 mg C m ⁻² The negative figure indicates an increase in algal biomass which can not be accounted for by primary production. The rate of phytoplankton turnover was compared with fluctuations in the standing crop of zooplankton but the correlation was not significant. Gross and net rates of extracellular production were estimated and compared. There was no significant difference between the two measurements and the implications of this are discussed with respect to other work in the field.     

CARBON,NITROGEN, AND PHOSPHORUS AND THE EUTROPHICATION OF FRESHWATER LAKES1

The question of nutrients responsible for eutrophication of freshwater lakes is reviewed, and recent additions to the literature on nutrient limitation are discussed. The paper by Lange is criticized on several grounds, including the facts that utilization of HCO₃^? by phytoplankton and the invasion of lake waters by atmospheric CO₂ are ignored as sources of photosynthetic carbon. The phosphorus and nitrogen concentrations used in Lange's experiments are far higher than values published by others for Lakes Erie and Ontario. Preliminary results of fertilizing a small oligotrophic lake with nitrogen and phosphorus are described. The standing crop of phytoplankton increased by 30–50 ×, while the P:N:C ratio in seston did not change from ratios found in unfertilized lakes. Other experiments done in water columns isolated with polyethylene film showed that addition of carbon did not increase the phytoplankton standing crop. Since the fertilized lake was initially lower in total CO₂ than any other recorded in the literature, it is concluded that carbon is unlikely to limit the standing crop of phytoplankton in almost any situation. Measurements of invasion of atmospheric gases to the fertilized lake by the Rn²²² technique were compared with phytoplankton production measurements, revealing that atmospheric invasion of CO₂ is sufficient to support the high phytoplankton standing crop in the epilimnion of the lake. Possible errors in interpretation of culture and bottle-bioassay experiments with respect to eutrophication are discussed.     

Grazing by Talorchestia longicornis on an algal mat in a new England salt marsh

Grazing experiments using ¹⁴C and an analysis of fecal pellets and gut contents established that the gammaridean amphipod, Talorchestia longicornis Say, ingests blue-green algae on algal mats in a Massachusetts salt marsh. This grazing had a measurable effect on the lower algal mat, where the density of T. longicornis was high. Exclusion of amphipods resulted in increases in chlorophyll a content, carbon incorporation, and nitrogen fixation. This effect was not seen on the upper mat where T. longicornis was less abundant. The assimilation efficiency of T. longicornis feeding on a diet consisting mainly of blue-green algae was surprisingly high (67 %) considering that blue-green algae are usually considered as a poor quality food for herbivores. The population of T. longicornis seems to be annual, with growth of the overwintered juveniles in spring and early summer.     

Standing crops and primary productivity of the epipelon of two small ponds in North Somerset,U. K.

Summary The primary productivity and the standing crop of the epipelic algal communities in two small ponds of contrasting nutrient status were measured over a 2 year period. Standing crops reflected the richer nutrient status of the eutrophic Abbot's Pond compared with the oligotrophic Priddy Pool as did the productivity results when the same depth category in both ponds was examined. Production resulted from an interrelationship among light intensity, standing crop size and other variable environmental factors in both pools, however light in Abbot's Pond appeared to be the most important factor in limiting standing crop size and production in the deeper depth categories.