Interactions among nutrients, algae and invertebrates in a New Zealand mountain stream

SUMMARY. 1. Colonization of nutrient-diffusing substrata by periphyton and invertebrates was investigated at forested and open sites in a small, mountain stream and a spring in the South Island of New Zealand.
2. Substrata had colonization surfaces made from 100μm mesh plankton netting that enabled algal assemblages to be removed intact for scanning electron microscopy. They also allowed small volumes of solvent to be used for the extraction of photosynthetic pigments.
3. At all sites, periphyton assemblages were dominated by species of Achnanthes, Cocconeis and Gomphonema , and except in the forest in winter, periphyton biomass was always greater on enriched (N + P added) than control substrata.
4. Invertebrates colonizing diffusion substrata were principally larval Chironomidae (Orthocladiinac). No larvae were present in winter, but in three spring and summer trials mean larval densities were higher on nutrient-enriched than control substrata at all sites.
5. The inclusion of an insecticide Malathion in diffusion substrata, reduced insect colonization at open and forested sites. After 28 days, no concurrent increases in algal pigment concentration were observed on nutrienl-enriched or control substrata at the forested site. However, pigment concentrations were higher on substrata incorporating Malathion at the open site suggesting that algal standing crop was depressed by the activities of grazers.     

SPECTRAL FINGERPRINTING OF ALGAL COMMUNITIES: A NOVEL APPROACH TO BIOFILM ANALYSIS AND BIOMONITORING1

A new technique for spectral fingerprinting of major algal groups in the freshwater periphyton (i.e. cyanobacteria, green algae, and diatoms) was developed using confocal laser scanning microscopy. This technique used the differential spectral emission signatures of photosynthetic algae and allowed their spatially explicit quantification and community three‐dimensional reconstruction. Algal biovolume measurements, carried out with this technique, are superior to existing protocols involving chl and ash‐free dry mass assessments because they are nondestructive, localized, and specific at a group level. This technique can be used to generate depth profiles of the periphytic mat with various applications in aquatic ecology and biofilm analysis.     

Relationship of macrophyte-mediated changes in the water column to periphyton composition and abundance

SUMMARY. 1. Dense growths of Myriophyllum heterophyllum influenced temperature, dissolved oxygen, pH and light levels in the littoral waters of Lake Winnipesaukee, a soft-water. New England lake. Periphyton species composition and abundance (algal units cm' stem) were related to the macrophyte-mediated changes in the physicochemical environment.
2. Duringearlysummer, M. heterophyllum occupied onlythe lower part of the water column. Limnetic and littoral waters exchanged readily and were chemically similar. Early summer periphyton species composition was dominated by diatoms from the phytoplankton, entangled in the finely dissected leaves of M. heterophyllum.
3. By mid-summer, M. heterophyllum occupied the entire watercolumn of the littoral zone. The metabolic and photosynthetic activity and dense foliage of M. heterophyllum created marked vertical gradients in physicochemical conditions. Mid-day temperature, dissolved oxygen, pH and light levels were maximal in the surface waters of the M. heterophyllum mat during the summer. Concurrently, periphyton species composition shifted to blue-green and then to a filamentous green alga on the apex and mid-stem. On the lower stem, diatoms consistently dominated the periphyton.
4. Periphyton abundance on the apex was inversely related to apical elongation. Temporal fluctuations of periphyton abundance on the lower and mid-stem were small throughout the study. Periphyton abundance was lowest on the lower stem, where the deteriorating leaves provided less surface area for colonization.     

Effects of desiccation duration on the community structure and nutrient retention of short and long-hydroperiod Everglades periphyton mats

Responses of periphyton communities to different relevant durations of dry down were assessed. Long-hydroperiod sites within Everglades National Park remain wet for greater than 8 months of the year while short-hydroperiod mats are wet for fewer than 4 months of the year. Dry down duration of long and short-hydroperiod Everglades periphyton was manipulated from 0 to 1, 3, or 8 months after which periphyton was rewetted 1 month and examined for algal species composition. The effects of desiccation and rewetting on periphyton nutrient retention were also assessed. Relative abundance of diatoms declined from an average of 47% in the long-hydroperiod community at the start of the experiment to 24% after 1 month of desiccation and only 12% after 8 months of desiccation. Short-hydroperiod periphyton contained a lower proportion of diatoms at the outset (3%), which declined to less than 1% after the 8-month desiccation treatment. A significant increase in the filamentous cyanobacteria Schizothrix calcicola occurred in long-hydroperiod periphyton mats during this same period, but not in short-hydroperiod mats. Long-hydroperiod periphyton communities had a greater response to desiccation overall, but short-hydroperiod community structure responded to desiccation more rapidly. Because short-hydroperiod communities dry frequently, they appear to cope better to desiccating conditions than long-hydroperiod periphyton communities. This is indicated by the dominance of desiccation resistant algal taxa such as the cyanobacterial filaments S. calcicola and Scytonema hofmanni. Long-hydroperiod periphyton mat communities converge compositionally to short-hydroperiod periphyton communities after prolonged desiccation. Desiccation and rewetting caused long-hydroperiod periphyton to flux greater concentrations of nutrients than short-hydroperiod periphyton. Significant increases in efflux occurred from 1 to 8 months for total phosphorus (TP) and from 1 to 3 and 8 months for total nitrogen (TN) and total organic carbon (TOC). Thus, changes in periphyton mat community structure and function with altered hydroperiod may have long-term ecosystem effects.     

The relative importance of shading and nutrients on algal production in subtropical streams

1. We used artificial substrata in forested and open streams in South-East Queensland, Australia, to determine the relative importance of shading from riparian vegetation and of nutrients on periphyton growth, and whether nitrogen and/or phosphorus limited algal productivity.
2. Nutrient-diffusing substrata consisting of agar enriched with N, P and N + P, and controls without nutrients, were deployed in duplicate at 15 sites in headwater streams with riparian canopy cover ranging from 0 to 88%.
3. Shading was the over-riding factor controlling periphyton biomass accrual on the artificial substrata, with nutrients playing a relatively minor role. Microscopic examination of periphyton scrapings taken after 7 weeks revealed that diatoms dominated on the artificial substrata in shaded streams, whereas filamentous green algae dominated the algal assemblage in the more open canopy streams.
4. Whilst nutrients had little effect on the accrual of algal biomass compared with riparian shading, there was evidence that nitrogen, and not phosphorus, stimulated periphyton production in streams with sufficient light.     

Alterations in periphyton characteristics due to grazing in a Cascade foothill stream

SUMMARY 1. In situ experiments were conducted in a Washington stream to quantify the effects of grazing by a caddisfly larva, Dicosmoecus gilvipes (Trichoptera: Limnephilidae), and a mayfly nymph, Nixe rosea (Ephemeroptera: Heptageniidae) on periphyton biomass, structure, and function.
2. Dicosmoecus gilvipes reduced periphyton biomass from 92 mg m⁻²(as mean chlorophyll a ) to 33 mg m⁻². The grazed assemblage was less diverse and composed of smaller, closely attached diatoms, whereas there was a higher proportion of overstorey and filamenttius algae in the diverse, ungrazed periphyton.
3. By maintaining the periphyton community as a thin layer of diatoms, grazing by D. gilvipes appeared to promote a healthier, more vigorous community relative to the ungrazed mat, which became senescent in the latter part of the experiment.
4. Nixe rosea had little measurable effect on any characteristics of the periphyton measured. These nymphs apparently preferred small diatoms, which resulted in only micro-scale alterations in periphyton characteristics that were difficult to detect.
5. Biomass accrual of ungrazed and grazed periphyton was described by the logistic growth equation. Loss of biomass due to grazing by D. gilvipes or to senescence and sloughing were incorporated in the model to account for changes in grazed and ungrazed periphyton. respectively. Proposed mechanisms which described biomass accumulation were largely sup ported by model predictions.     

Influence of Algal Community Structure on Denitrification Rates in Periphyton Cultivated on Artificial Substrata

We conducted a field survey of periphyton cultivated on benthic mesh installations in freshwater aquatic systems, including two constructed wetlands and a pond, and also studied periphyton grown on a benthic mesh in laboratory mesocosms. The objectives of this study were to (1) determine if periphyton cultivated on benthic mesh denitrifies at higher rates than the underlying sediments and (2) determine if denitrification rates within periphyton vary with characteristics such as algal and bacterial community structure and biomass. We measured denitrification potential rates of field and laboratory periphyton by the acetylene inhibition method. We characterized algal community composition by algal identification and bacterial community composition by terminal restriction fragment length polymorphisms. Periphyton collected on benthic mesh from our field sites denitrified at significantly higher rates than the underlying sediments, regardless of sampling site or season. Results from both our field survey and laboratory studies indicated a significant, positive correlation between diatom presence and denitrification rate. In our laboratory studies, we found that periphyton with the highest diatom abundance showed the highest denitrification rates as well as a distinct bacterial community composition. These results suggest a synergistic relationship between diatoms and denitrifying bacteria that warrants further study.     

Perceptual scales of spatial heterogeneity of periphyton for freshwater snails

The hypothesis that individuals recognize their environment as homogeneous when the scales of spatial heterogeneity of resources are smaller than a certain scale, but can distinguish them as patches when they are larger than that scale was tested using freshwater snails ( Physa acuta ) in various distributions in periphyton environments. In a pattern of periphyton distribution in which the size of algal cells was 47 mm, individuals moved significantly more slowly on algal cells than on nonalgal cells. However, in other patterns in which the sizes of algal cells were 23.5 mm and 15.7 mm, the speeds of individuals on algal and nonalgal cells were not significantly different. These results support the hypothesis that individuals use the environment homogeneously when the scales of spatial heterogeneity of resources are smaller than a certain scale, but they can distinguish between patches when the scales are larger.     

The role of periphyton abundance and quality in the microdistribution of a stream grazer, Helicopsyche borealis (Trichoptera: Helicopsychidae)

SUMMARY. 1. A series of laboratory and field experiments was performed to determine if the preference of larvae of Helicopsyche borealis (Trichoptera: Helicopsychidae) for exposed rock surfaces in streams was related to patchiness of periphyton food.
2. At low current velocities in the laboratory larvae preferred food-saturated as opposed to food-depleted portions of substrata and drifted off the substratum in response to low food levels.
3. When offered a choice of three potential foods—a diatom, a green filamentous or a blue-green filamentous alga—larvae showed no preference.
4. Larval densities on artificial subtrata in field experiments were highest in areas of high algal chlorophyll a .
5. In laboratory experiments more adults emerged from cultures containing green algae or diatoms than cultures with blue-green algae or bacteria alone.
6. Periphyton quality was not as important as abundance in controlling H. borealis microdistribution. By selecting exposed surfaces of rocks, H. borealis larvae place themselves in the microhabitat most likely to contain both abundant and high quality food patches.     

Occurrence of phosphorus and its potential remohilization in the littoral sediments of a productive English lake

SUMMARY. 1. Vertical profiles of pH were measured at nine shallow water (<5m) locations in Esthwaite Water. These indicate strong gradients of pH near the sediment water interface suggesting a marked buffering capacity of the sediments.
2. Thirteen littoral sediment cores were horizontally sectioned and sequentially extracted (0.5 M NaHCO₃, 0.1 M NaOH, 1 M HCI) and analysed for soluble reactive phosphorus. The core sections were also analysed for total phosphorus and per cent organic content to determine the vertical and areal variability of phosphorus within the littoral sediments of Esthwaite Water.
3. The rate of release of phosphorus from intact sediment cores was measured in the laboratory as a function of the pH of overlying water, yielding the relationship log K=0.54 pH−3.94, K=mg Pm⁻²day⁻¹. The maximum release rate measured was 75 mg P m⁻² day⁻² at pH = 10.5.
4. Experiments on sediment slurries indicate that the release of phosphorus at pH 10 is rapid with approximately 50% of the total NaHCO₃+ NaOH extractable phosphorus being released within 3 h.
5. Phosphorus release from the littoral sediments may equal or exceed external sources plus hypolimnetic inputs during periods of high pH associated with times of maximum algal biomass.     

Low algal carbon content and its effect on the C : P stoichiometry of periphyton

1. We examined the contribution of algal cells to periphytic organic carbon and assessed the effects of variable biomass composition on the carbon : phosphorus (C : P) ratio of periphyton. We compiled more than 5000 published and unpublished observations of periphytic carbon : chlorophyll a (C : Chl) ratios, an index of algal prevalence, from a variety of substrata collected from lake and low‐salinity coastal habitats. In addition, we converted estimates of algal biovolume into algal C to obtain an independent measure of cellular algal carbon in periphyton. This information was used in a model relating periphyton C : P ratio to algal cellular carbon, the algal C : P ratio, and the C : P ratio of non‐algal organic matter in periphyton. 2. The mean C : Chl ratio of periphyton (405) was relatively high with values in >25% of the samples exceeding 500. On average, 8.4% of total periphyton C was accounted for by C in algal cells. Only 15% of samples were found to have more than 15% periphyton C in cellular algal carbon. Our model showed a nonlinear relationship between periphytic C : P ratios and the C : P ratio of algal cells in the periphyton when non‐algal organic matter was present. However, even at relatively low cellular algal C (<10% of total C), algal C : P ratios can strongly affect the C : P ratio of periphyton as a whole (i.e. algal cells plus other organic matter). 3. The high C : Chl ratios and the low biovolume‐derived algal C of periphyton samples in our data set indicate that algal cells are typically a minor component of organic carbon in periphyton, However, this minor contribution would not preclude algal cellular stoichiometry from notably influencing periphyton C : P ratios.     

Interactions between nutrient availability and hydroperiod shape macroinvertebrate communities in Florida Everglades marshes

Hydroperiod and nutrient status are known to influence aquatic communities in wetlands, but their joint effects are not well explored. I sampled floating periphyton mat and flocculent detritus (floc) infaunal communities using 6-cm diameter cores at short- and long-hydroperiod and constantly inundated sites across a range of phosphorus (P) availability (total phosphorus in soil, floc and periphyton). Differences in community structure between periphyton and floc microhabitats were greater than any variation attributable to hydroperiod, P availability, or other spatial factors. Multivariate analyses indicated community structure of benthic-floc infauna was driven by hydroperiod, although crowding (no. g⁻¹ AFDM) of individual taxa showed no consistent responses to hydroperiod or P availability. In contrast, community structure of periphyton mat infauna was driven by P availability, while densities of mat infauna (no. m⁻²) were most influenced by hydroperiod (+correlations). Crowding of mat infauna increased significantly with P availability in short-hydroperiod marshes, but was constant across the P gradient in long-hydroperiod marshes. Increased abundance of floating-periphyton mat infauna with P availability at short-hydroperiod sites may result from a release from predation by small fish. Community structure and density were not different between long-hydroperiod and constantly inundated sites. These results have implications for the use of macroinvertebrates as indicators of water quality in wetlands and suggest the substrate sampled can influence interpretation of ecological responses observed in these communities.     

Effects of a macroalgal mat (Ulva lactuca) on estuarine sand flat copepods: an experimental study

The effects of a macroalgal mat (Ulva lactuca) on intertidal sand flat copepods were studied in Jamaica Bay, NY. In 1999, Ulva was removed from paired quadrats in the algal mat. Density and species richness were estimated from sediment cores over seven biweekly periods and compared with Ulva-removal quadrats and with an unvegetated reference site. In a separate analysis at the same site, species composition and density of all copepods within the algal mat were determined from replicated quadrat samples over seven dates. Emergence traps were deployed in all treatments in 1999 to quantify copepod movements from sediments into the water column during flooding tide. Dissolved oxygen (DO) was continuously monitored over several 96-h periods in and above the algal mat. In 2000, three biomass levels of Ulva (0, 100, 200 g DW per m²) were added to paired quadrats in the unvegetated zone. Density and species richness were estimated over eight time periods. Ulva removal resulted in significant increases in density and species richness, but levels remained lower than at the unvegetated site. Likewise, addition of Ulva biomass produced significant declines in copepod density. However, the differences between low- and high-Ulva-addition treatments were insignificant. Increases in phytal copepods after Ulva addition did not compensate for losses of epibenthic and infaunal species. The overall effect of the Ulva mat was a net loss of copepod density, including the loss of nearly all infaunal species. Anoxia in sediments and within the algal mat are correlated with declines in infaunal species and with escape by some infaunal species from the sediment into the water column.     

The role of canopy cover on the recovery of periphyton and macroinvertebrate communities after a month-long flood

We studied the recovery of periphyton and macroinvertebrate communities in a second order stream after a month-long spate that began as increased discharge due to snowmelt in April 2000 but continued through May as a result of frequent rainstorms. We sampled macroinvertebrates and periphyton in June 2000 at three different sites—an upstream site in a conifer tree plantation, a second site ca. 800 m downstream in a mixed hardwood forest, and a third site ca. 1.4 km downstream where there was an open canopy. Periphyton abundance was low on 6 June at all sites but increased 10-fold at the open canopy site the following week (13 June), appearing as a thick mat of Ulothrix. By 26 June, periphyton biomass (as both chlorophyll a and ash free dry mass) had decreased by 80% whereas densities of Chironomidae increased 50-fold and Baetidae increased 4-fold at the open canopy site; little change had occurred in periphyton and macroinvertebrate communities at the forested sites from early to late June. Our results suggest that baetids and chironomids rapidly responded to the increased algal resource at the open canopy site due to their life history characteristics and high growth rates. Chironomids decreased the green mat of Ulothrix by using this alga in tube construction and both chironomids and baetids likely consumed this alga. The varied responses of the periphyton and macroinvertebrate communities at the forested versus open sites demonstrate a resistance to impacts by floods at the forested sites, whereas at the open canopy site periphyton and macroinvertebrate communities were more resilient to the flood disturbance. Handling editor: R. Bailey     

Removal of settled sediments and periphyton from macrophytes by grazing invertebrates in the littoral zone of a large oligotrophic lake

• 1 The resistance and resilience of littoral zone communities to sedimentation will depend both on the extent to which sediment deposition affects productivity, and on interactions within the communities. A series of hypotheses were set up and tested to examine interactions and feedback mechanisms among deposited sediments, periphyton, macrophytes and grazers in a large oligotrophic lake subject to fluctuating sediment loadings.
• 2 Although sediments incorporated into periphyton reduced light availability to macrophytes, periphytic algae were generally the dominant light absorbing component under natural conditions. When grazers were absent, both sediments incorporated in the periphyton and periphytic algal densities increased, and both were then important in reducing light available to macrophytes.
• 3 Grazing rate and assimilation efficiency for the dominant grazer, the prosobranch gastropod Potamopyrgus antipodarum, increased with increasing sediment content under natural lake conditions to reach a maximum at 10 mg sediment cm^?2.
• 4 An increase in sediment incorporation into periphyton films resulted in an increased grazing rate and hence grooming of sediments from macrophytes.
• 5 Grazing invertebrates can play a major role in maintenance of littoral communities by continuously grooming macrophyte hosts of periphytic algae and settled sediments.

     

Spatial distribution of denitrifying activity in a stream draining an agricultural catchment

SUMMARY 1. A field study was made of the spatial distribution of denitrification activity in the sediment of the River Dorn, Oxfordshire, England.
2. An assay of denitrifying enzyme activity was used to examine the distribution of denitrification with depth in cores of sediment representative of the types found in the stream. The maximum activity recorded in a predominantly silt sediment core was 5 times greater than that recorded in a sandy gravel core. In both fine sand and silt cores, peaks in denitrifier enzyme activity were shown to correspond to the limit of the nitrate diffusion front. At this depth the redox potential dropped rapidly from + 300 mV to 0 or less. Denitrifying enzyme activity in the stream water was negligible.
3. In situ denitrification activity (I DA) measurements were carried out in an 800 m reach of the Dorn using the acetylene inhibition technique on small sediment cores. Concurrent measurements were also made of stream depth and velocity, nitrate concentration in the interstitial water, and the wet bulk density, loss on ignition, mineraliz- able carbon and total nitrogen contents of the sediment. Mineralizable carbon was the variable which showed the best correlation with I DA. Highest IDAs were associated with accumulations of fine-grained sediment at meander bends. Mean IDAs measured under flood conditions were significantly higher ( P <0.05) than those measured under baseflow. It was estimated that denitrification reduced the nitrate load in the River Dorn by 15% under summer baseflow conditions     

Uncoupling of omnivore-mediated positive and negative effects on periphyton mats

The riverine grass shrimp (Palaemonetes paludosus) and eastern mosquitofish (Gambusia holbrooki) consume periphyton and small invertebrates, potentially affecting periphyton through negative effects (i.e., consumption) and/or positive effects such as nutrient regeneration, physical stimulation, and trophic cascades. We performed field experiments in the Everglades in which omnivores and periphyton were maintained in cages, with a fraction of the periphyton held in omnivore-exclusion bags that allowed passage of nutrients but prevented its consumption or physical disturbance. In some instances, periphyton growth rate increased with increasing omnivore biomass. Omnivores probably stimulated periphyton growth through nutrient regeneration, possibly subsidizing periphyton with nutrients derived from ingested animal prey. The net balance of omnivore-mediated negative and positive effects varied among experiments because of seasonal and spatial differences in periphyton characteristics. Consumption of periphyton mats might have been reduced by the arrangement of palatable algae (green algae and diatoms) within a matrix of unpalatable ones (CaCO(3)-encrusting filamentous cyanobacteria). In a laboratory feeding experiment, mosquitofish consumed more green algae and diatoms in treatments with disrupted mat structure than in those with intact mats. No difference in diet was observed for shrimp. Our study underscores the complexity of consumer-periphyton interactions in which periphyton edibility affects herbivory and consumers influence periphyton through multiple routes that cannot be fully appreciated in experiments that only investigate net effects.     

Evaluating the multiple mechanisms underlying herbivore-algal interactions in streams

Stream algal responses to herbivory were investigated under different environmental conditions. Snail densities and nutrient concentrations were manipulated in experimental enclosures to document the influence of nutrient availability on the magnitude of algal responses to herbivory. Periphyton mats in other enclosures were subjected to physical disruption by artificial means to evaluate the influence of disturbance on algal abundance. The impact of herbivory on algal abundance decreased substantially with increase in water column nutrient concentrations. This result was explained by findings that: (1) algal accumulation was constrained by nutrient availability under ambient water quality conditions. (2) accumulation of most algal populations was stimulated by nutrient enrichment only under grazed conditions. Thus, snail grazing simultaneously exerted a negative impact on algal abundance, by removal and consumption of a portion of the periphyton mat, and a stimulative effect, by increasing the availability of nutrients to remaining cells. Algal responses to artificial disturbance indicated that stimulative effects of herbivory were caused by the physical disruption of thr assemblage rather than by other processes (e. g., nutrient regeneration). However, consumptive losses far outweighed stimulative effects on algal abundance under ambient nutrient conditions. The magnitude of these two antagonistic effects was comparable under enriched conditions because grazing had only slight effects on algal abundance. Thus, the importance of different mechanisms of algal-herbivore interactions is strongly influenced by ambient environmental conditions, a finding that has important implications for predicting the outcome of herbivore-algal interactions in ecosystems with pronounced temporal and spatial variation in biotic and abiotic conditions.     

Efficacy of algal metrics for assessing nutrient and organic enrichment in flowing waters

1. Algal-community metrics were calculated for periphyton samples collected from 976 streams and rivers by the U.S. Geological Survey's National Water-Quality Assessment (NAWQA) Programme during 1993–2001 to evaluate national and regional relations with water chemistry and to compare whether algal-metric values differ significantly among undeveloped and developed land-use classifications.
2. Algal metrics with significant positive correlations with nutrient concentrations included indicators of trophic condition, organic enrichment, salinity, motility and taxa richness. The relative abundance of nitrogen-fixing algae was negatively correlated with nitrogen concentrations, and the abundance of diatom species associated with high dissolved oxygen concentrations was negatively correlated with both nitrogen and phosphorus concentrations. Median algal-metric values and nutrient concentrations were significantly lower at undeveloped sites than those draining agricultural or urban catchments.
3. Total algal biovolume did not differ significantly among major river catchments or land-use classifications, and was only weakly correlated with nitrate (positive) and suspended-sediment (negative) concentrations. Estimates of periphyton chlorophyll a indicated an oligotrophic–mesotrophic boundary of about 21 mg m⁻² and a mesotrophic–eutrophic boundary of about 55 mg m⁻² based on upper and lower quartiles of the biovolume data distribution.
4. Although algal species tolerance to nutrient and organic enrichment is well documented, additional taxonomic and autecological research on sensitive, endemic algal species would further enhance water-quality assessments.     

High nutrient availability leads to weaker top‐down control of stream periphyton: Compensatory feeding in Ancylus fluviatilis

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