NUTRIENT LIMITATION OF BENTHIC ALGAE IN LAKE MICHIGAN: THE ROLE OF SILICA1

In the Laurentian Great Lakes, phytoplankton growth and biomass are secondarily limited by silica (Si), as a result of phosphorus (P) enrichment. Even modest levels of P enrichment can induce secondary Silimitation, which, in turn, promotes a shift from the native diatom phytoplankton flora to chlorophyte and cyanobacteria species. However, very little is known about the nutritional status of benthic populations and their response to nutrient enrichment. Two experiments were performed in the littoral zone of Lake Michigan where nutrients were delivered to in situ benthic algal (episammic and epilithic) assemblages using nutrient‐diffusing substrata. In order to test the hypothesis that benthic algae in Lake Michigan are Si limited, a 2 × 3 factorial experiment was used to deliver all combinations of Si, N, and P to resident assemblages growing on artificial substrata composed of natural (Si rich) versus calcium carbonate (Si poor) sand. A second experiment utilized a serial enrichment to evaluate the role of Si in mediating changes in taxonomic composition. These findings indicate that benthic algae in Lake Michigan exhibit signs of secondary Si limitation, and that their response to enrichment is similar to the phytoplankton. Moreover, natural sand substrata may provide a source of Si to resident benthic algae.     

Local disturbance history affects patchiness of benthic river algae

1. Recent research has shown that high‐flow events in streams leave a small‐scale mosaic of bed patches that have experienced scouring, sediment deposition (fill), or remained stable. Few studies have investigated if this ‘local disturbance history’ contributes to the patchy distribution of benthic organisms in streams and rivers. 2. In the present research, we demonstrate that local disturbance history in a mid‐sized river can have both short‐ and long‐term effects on epilithic algae. Chains buried vertically in the substratum of the river bed (236 in a 800‐m reach) indicated that two floods (return periods ≤1 year) caused a mosaic of bed patches with different disturbance histories. Once after the first and twice after the second flood, we sampled epilithic algae (mainly diatoms) in replicate patches that had been scoured, filled, or remained stable during the respective event. Algal biomass and cell density per substratum area were determined. 3. Three months after the first flood, algal biomass, total diatom density, diatom taxon richness, and densities of six of nine most common taxa were highest in fill patches. Six days after the second flood, biomass was highest in stable patches, indicating a refugium function of these patches. The refugium patches consisted of average‐sized stones, in contrast to previous studies of flood refugia for benthic algae in which these refugia were always large and/or immobile substrata. Four weeks after the second flood, diatoms tended to be most abundant in scour patches. With one exception, these differences between patch types could not be attributed to differences in local near‐bed current velocity or water depth. 4. The effects of disturbance history were more complex than a simple refugium function of stable patches because algal patterns changed with time since the last disturbance, possibly depending on the successional state of the algal mats.     

A manipulative study of macroinvertebrate grazers in Hong Kong streams: do snails compete with insects?

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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.     

Production stability and biomass quality in microalgal cultivation – Contribution of community dynamics

The prospect of using constructed communities of microalgae in algal cultivation was confirmed in this study. Three different algal communities, constructed of diatoms (Diatom), green algae (Green), and cyanobacteria (Cyano), each mixed with a natural community of microalgae were cultivated in batch and semi‐continuous mode and fed CO₂ or cement flue gas (12–15% CO₂). Diatom had the highest growth rate but Green had the highest yield. Changes in the community composition occurred throughout the experiment. Green algae were the most competitive group, while filamentous cyanobacteria were outcompeted. Euglenoids, recruited from scarce species in the natural community became a large part of the biomass in semi‐steady state in all communities. High temporal and yield stability were demonstrated in all communities during semi‐steady state. Valuable products (lipids, proteins, and carbohydrates) comprised 61.5 ± 5% of ash‐free biomass and were similar for the three communities with lipids ranging 14–26% of dry mass (DM), proteins (15–28% DM) and carbohydrates (9–23% DM). Our results indicate that culture functions (stability, biomass quality) were maintained while dynamic changes occurred in community composition. We propose that a multispecies community approach can aid sustainability in microalgal cultivation, through complementary use of resources and higher culture stability.     

Benthic algal biomass in an unshaded first-order lowland stream: distribution and regulation

The vertical distribution of algal biomass in the bed sediment and the seasonal development of benthic algae on stones and fine-grained sediments were studied in a small unshaded stream. In addition, field experiments were conducted on the role of irradiance and phosphorus in regulating algal biomass. We found that algal biomass was high at a sediment depth of ten centimetres. Comparison of studies on algal biomass where different depths of the sediment are used should therefore be made with caution. Substrata-dependent differences in algal biomass development were substantial. While algal biomass development on stones was controlled by macroinvertebrate grazing, that on the fine-grained sediment followed the dynamics of incident irradiance, but was attenuated by sediment rebedding. Because of the high grazing pressure on algal biomass on stony substrata, no significant response to phosphorus enrichment was attained. In contrast, algal biomass development on fine-grained sediments was phosphorus-limited. Heavy shading of the fine-grained sediments did not significantly affect algal biomass development, thus suggesting that phosphorus limitation prevents algae from fully utilizing the light resource in this stream. This revised version was published online in July 2006 with corrections to the Cover Date.     

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

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

How protective are refuges? Quantifying algal protection in rock crevices

1. Refuges can be functionally important if they harbour sufficient organisms during disturbance to augment population recovery. I quantified the protection of stream algae in crevice refuges using the applied, severe disturbance of scrubbing. Scrubbing effectively removed visible surface algae, and algae remaining on stones were considered protected. 2. In a field experiment, substrata with different quantities of crevice (glass bottles; greywacke; schist; pumice) were incubated in a channelised stream. The possible influence of growth conditions was investigated concurrently; half of the substrata were suspended in the water column, the rest were placed on the bottom (providing differential access to grazers and exposing them to different flow conditions). 3. Rougher substrata had greater total algal biomass than smoother substrata; this pattern resulted from more algal biomass in crevices of the rougher substrata. Protection from scrubbing ranged from about 5% of total algal biomass on glass and greywacke to 80% on pumice. In contrast, algal biomass removed by scrubbing was similar among the experimental substrata. Suspended substrata had more chironomid grazers than those on the bottom, and also greater algal biomass, possibly because of high algal concentrations in the chironomid retreats. 4. A survey of stones from three rivers supported the experimental results; namely, rough pumice protected more algae from scrubbing than did smoother greywacke. 5. In a separate experimental assay, there was no difference in algal growth on agar plates with and without added powdered rock substrata, suggesting that crevice characteristics and not substratum chemistry produced the differences in algal assemblages between rough and smooth surfaces. 6. Results indicate that rough stream stones may protect sufficient algae to augment their recovery in streams following disturbance.     

Nitrogen limitation of the phytobenthos in Alpine lakes: results from nutrient‐diffusing substrata

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LINKING BENTHIC ALGAL BIOMASS TO STREAM SUBSTRATUM TOPOGRAPHY1

The physical properties of substrata significantly influence benthic algal development. We explored the relationships among substratum surface texture and orientation with epilithic microphytobenthic biomass accumulation at the whole‐substratum and micrometer scales. Unglazed clay tiles set at three orientations (horizontal, vertical, and 45°), and six substrata of varying surface roughness were deployed in a prairie stream for 3 weeks. Substrata were analyzed for loosely attached, adnate, and total benthic algal biomass as chl a, and confocal laser scanning microscopy was used to measure substrata microtopography (i.e., roughness, microscale slope angles, and three‐dimensional surface area). At the whole‐substratum level, vertical substrata collected significantly (P < 0.05) less algal biomass, averaging 34% and 36% less than horizontal and 45° substrata, respectively. Benthic algal biomass was also significantly less on smoother surfaces; glass averaged 29% less biomass than stream rocks. At the microscale level, benthic algal biomass was the greatest at intermediate values, peaking at a mean roughness of approximately 17 μm, a mean microscale slope of 50°, and a projected/areal surface area ratio of 2:1. The proportion of adnate algae increased with surface roughness (26% and 67% for glass and brick, respectively), suggesting that substratum type changes the efficiency of algal removal by brushing. Individual substrata and microsubstrata characteristics can have a strong effect on benthic algae development and potentially affect reach scale algal variability as mediated by geomorphology.     

Response of an algal assemblage to nutrient enrichment and shading in a Hawaiian stream

To investigate the effects of nitrate enrichment, phosphate enrichment, and light availability on benthic algae, nutrient-diffusing clay flowerpots were colonized with algae at two sites in a Hawaiian stream during spring and autumn 2002 using a randomized factorial design. The algal assemblage that developed under the experimental conditions was investigated by determining biomass (ash-free dry mass and chlorophyll a concentrations) and composition of the diatom assemblage. In situ pulse amplitude-modulated fluorometry was also used to model photosynthetic rate of the algal assemblage. Algal biomass and maximum photosynthetic rate were significantly higher at the unshaded site than at the shaded site. These parameters were higher at the unshaded site with either nitrate, or to a lesser degree, nitrate plus phosphate enrichment. Analysis of similarity of diatom assemblages showed significant differences between shaded and unshaded sites, as well as between spring and autumn experiments, but not between nutrient treatments. However, several individual species of diatoms responded significantly to nitrate enrichment. These results demonstrate that light availability (shaded vs. unshaded) is the primary limiting factor to algal growth in this stream, with nitrogen as a secondary limiting factor.     

Periphyton distribution and abundance on substrata of different size along a gradient of stream trophy de Montréal

In stream, substrata of different size present different degree ofstability, current, erosion, and deposition to colonizingorganisms. In this study, we tested the importance of substratumsize ranging from sand to small boulders for periphytondistribution and abundance. Because trophy strongly affect streamorganisms, we sampled at nine sites chosen to represent the rangeof nutrients typical of Eastern Ontario and Western Québec. Alarge part of the variability in algal biomass (as chlorophyll)among sites was explained by trophy (as seston or totalphosphorus). However, there was also an effect of substratum size.Cobbles had the highest biomass, and gravel the lowest; sand andboulders were intermediate. Assemblages on different substrata weredifferent in taxonomy and life forms. Cyanobacterial colonies andmotile diatoms dominated the finer substrata while adnate andfilamentous algae were more developed on the larger ones.Consequently periphyton on fine sediments was more loosely attachedthan on rocks. Average algal size was not related to substratumsize but increased significantly with trophy confirmingobservations in benthic and planktonic assemblages inlakes.     

Exclusion size and material have minimal effects on stream benthic algae and macroinvertebrate colonization within submerged cages

Despite their widespread use in grazer–biofilm studies, stream exclusion cages have inherent physical properties that may alter benthic organism colonization and growth. We used laboratory studies and a field experiment to determine how exclusion cage design (size and material) alters light availability, water velocity, and benthic organism colonization. We measured light reduction by various plastic cage materials and flow boundary layer thickness across a range of exclusion cage sizes in the laboratory. We also deployed multiple exclusion cage designs based on commonly available materials into a second-order stream to assess algae and macroinvertebrate colonization differences among exclusion cages. All plastics reduced some light (190–700 nm wavelengths) and blocked more ultraviolet light than photosynthetically active radiation. Exclusion cage size did not influence flow boundary layer thickness, but larger exclusions tended to have higher velocity at the substrata surface. Despite light and water velocity differences, algal biomass, macroinvertebrate density, and community composition were similar between exclusion cage types. However, algal assemblages outside exclusion cages differed in composition and had higher biomass compared to inside exclusion cages. In terms of algal and macroinvertebrate colonization, plastic exclusion cage size and material appear to be flexible within the sizes tested, but differences can still exist between exclusion cage communities and those within the stream. Overall, artifacts of screened exclusion cages do not appear to introduce large bias in results of grazer–biofilm studies, but efforts to design exclusion cages that better mimic the natural system should continue.     

Interactive effects of three pervasive marine stressors in a post-disturbance coral reef

Ecosystems are commonly affected by natural, episodic disturbances that can abruptly and drastically alter communities. Although it has been shown that resilient ecosystems can eventually recover to pre-disturbed states, the extent to which communities in early stages of recovery could be affected by multiple anthropogenic stressors is poorly understood. Pervasive and rising anthropogenic stressors in coastal marine systems that could interactively affect the recovery of these systems following natural disturbances include high sedimentation, nutrient enrichment, and overfishing. Using a 6-month field experiment, we examined the effects of all combinations of these three stressors on key functional groups in the benthic community growing on simulated, post-disturbance reef patches within a system recovering from large-scale natural disturbances (corallivorous seastar outbreak and cyclone). Our study revealed that sedimentation, nutrient enrichment, and overfishing (simulated using exclusion cages) interactively affected coral survival and algal growth, with taxon-specific effects at multiple scales. First, our treatments affected corals and algae differently, with sedimentation being more detrimental to macroalgal growth but less detrimental to coral (Porites rus) survival in caged plots, driving significant interactions between sedimentation and caging for both taxa. We also observed distinct responses between coral species and between algal functional groups, with the most extensive responses from algal turf biomass, for which sedimentation suppressed the synergistic (positive) combined effect of nutrient enrichment and caging. Our findings suggest that different combinations of ubiquitous anthropogenic stressors, related to either sea- or land-based activities, interactively influence community recovery from disturbance and may alter species compositions in the resulting community. Our findings further suggest that anthropogenic stressors could promote further degradation of coral reefs following natural disturbances by inhibiting recovery to coral-dominated states that provide vital ecosystem services to coastal populations worldwide.     

Spatial and seasonal variations in benthic algal assemblages in streams in monsoonal Hong Kong

Samples from stone surfaces were collected in pools within four unpolluted hillstreams (two shaded and two unshaded) in monsoonal Hong Kong (lat. 23°N) to elucidate the extent of spatial (within and among streams) and temporal (seasonal) variations in algal biomass and assemblage composition. Sampling continued for over 12 months, incorporating the dry season when streams were at baseflow, and the wet season when spates were frequent. We anticipated that algal biomass would be lower in shaded streams and during the wet season, with associated seasonal differences in assemblage composition or relative abundance of different growth forms (e.g. erect versus prostrate). Benthic chlorophyll a (a proxy for algal biomass) varied among streams from an annual mean of 11.0–22.3 mg m⁻². Dry-season standing stocks were 18% higher than during the wet season when spate-induced disturbance reduced algal standing stocks. Algal biomass varied significantly at the stream scale, but not at the pool scale, and was lower in unshaded streams, where standing stocks may have been limited by high densities of algivorous balitorid loaches (mainly Pseudogastromyzon myersi). An overriding effect of grazers on algal biomass could also have reduced variations resulting from spate-induced disturbance. Significant differences in assemblage composition among streams, which were dominated by diatoms and cyanobacteria (totally 82 taxa) were not systematically related to shading conditions. Seasonal variations in algal assemblages were statistically significant but rather minor, and did not involve major shifts in composition or growth form caused by spate-induced disturbance. The abundance of filamentous cyanobacteria in all the streams may have been due to ‘gardening’ by balitorid loaches that removed erect or stalked diatoms and favoured cyanobacteria that persist through basal regeneration of filaments. This explanation requires validation through manipulative experiments. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: Luis Mauricio Bini     

Responses of benthic algae to nutrient enrichment in a shallow lake: Linking community production,biomass, and composition

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Flood disturbance effects on benthic diatom assemblage structure in a semiarid river network

Disturbances such as floods and droughts play a central role in determining the structure of riverine benthic biological assemblages. Extreme disturbances from flash floods are often restricted to part of the river network and the magnitude of the flood disturbance may lessen as floods propagate downstream. The present study aimed to characterize the impact of summer monsoonal floods on the resistance and resilience of the benthic diatom assemblage structure in nine river reaches of increasing drainage size within the Gila River in the southwestern United States. Monsoonal floods had a profound effect on the diatom assemblage in the Gila River, but the effects were not related to drainage size except for the response of algal biomass. During monsoons, algal biomass was effectively reduced in smaller and larger systems, but minor changes were observed in medium systems. Resistance and resilience of the diatom assemblage to floods were related to specific species traits, mainly to growth forms. Tightly adhered, adnate and prostrate species (Achnanthidium spp., Cocconeis spp.) exhibited high resistance to repeated scour disturbance. Loosely attached diatoms, such as Nitzschia spp. and Navicula spp., were most susceptible to drift and scour. However, recovery of the diatom assemblage was very quick indicating a high resilience, especially in terms of biomass and diversity. Regional hydroclimatic models predict greater precipitation variability, which will select for diatoms resilient to bed‐mobilizing disturbances. The results of this study may help anticipate future benthic diatom assemblage patterns in the southwestern United States resulting from a more variable climate.     

Organic matter availability structures microbial biomass and activity in a Mediterranean stream

1. We compared microbial biomass (bacteria, fungi, algae) and the activity of extracellular enzymes used in the decomposition of organic matter (OM) among different benthic substrata (leaves, coarse and fine substrata) over one hydrological year in a Mediterranean stream.
2. Microbial heterotrophic biomass (bacteria plus fungi) was generally higher than autotrophic biomass (algae), except during short periods of high light availability in the spring and winter. During these periods, sources of OM shifted towards autochthonous sources derived mainly from algae, which was demonstrated by high algal biomass and peptidase activity in benthic communities.
3. Heterotrophic activity peaked in the autumn. Bacterial and fungal biomass increased with the decomposition of cellulose and hemicellulose compounds from leaf material. Later, lignin decomposition was stimulated in fine (sand, gravel) and coarse (rocks, boulders and cobbles) substrata by the accumulation of fine detritus.
4. The Mediterranean summer drought provoked an earlier leaf fall. The resumption of the water flow caused the weathering of riparian soils and subsequently a large increase in dissolved organic carbon and nitrate, which led to growth of bacteria and fungi.     

Climate-induced shifts in an experimental phytoplankton community: a mechanistic approach

Climate change is likely to have far-reaching effects on biotic interactions in aquatic ecosystems. We investigated the effect of different spring warming scenarios on the succession of three algal groups (cyanobacteria, diatoms and green algae) in 10-l microcosms. We fitted these microcosm data to a simple mechanistic model to estimate the effect of different climate warming scenarios on the population dynamics of these algal functional groups. Experimental and model results indicate that the different algal functional groups respond differently to climate warming under phosphorus-limited conditions. Whereas the successional sequence, from diatoms to green algae to cyanobacteria, was not affected by the different climate warming scenarios, cyanobacteria showed a stronger response to the different climate warming scenarios than diatoms or green algae. Both the growth rates and peak abundances of cyanobacteria were significantly higher in the average and warm spring scenarios than in the cold spring scenario. Our findings illustrate that integration of models and microcosm experiments are a useful approach in predicting the impacts of rising temperatures on the dynamics of phytoplankton communities.     

Three-year study of benthic algal spring bloom development in a small, Danish lowland stream

Seasonal development of benthic algae was studied over a three-year period in a small, nutrient-rich lowland stream to investigate inter-annual variation in the algal spring bloom and differences in algal biomass regulation on two different substrata: fine-grained sediments and stones. The algal spring bloom was initiated when irradiance at the sediment surface exceeded 7 mol photons m^-2 d^-1 and mean water velocity was concomitantly below the threshold for bed load transport in the stream. Large inter-annual and substratum-dependent differences in peak algal biomass were observed, thus suggesting that different parameters regulate algal biomass development on the two substrata. On fine-grained sediments algal biomass development was predominantly coupled to light availability, while on stony substrata algal composition and peak biomass might be affected by invertebrate grazing.