Effects of sediment resuspension on phytoplankton production: teasing apart the influences of light, nutrients and algal entrainment

1. Wind‐induced sediment resuspension can affect planktonic primary productivity by influencing light penetration and nutrient availability, and by contributing meroplankton (algae resuspended from the lake bed) to the water column. We established relationships between sediment resuspension, light and nutrient availability to phytoplankton in a shallow lake on four occasions. 2. The effects of additions of surficial sediments and nutrients on the productivity of phytoplankton communities were measured in 300 mL gas‐tight bottles attached to rotating plankton wheels and exposed to a light gradient, in 24 h incubations at in situ temperatures. 3. While sediment resuspension always increased primary productivity, resuspension released phytoplankton from nutrient limitation in only two of the four experiments because the amount of available nitrogen and phosphorus entrained from the sediments was small compared with typical baseline levels in the water column. In contrast, chlorophyll a entrainment was substantial compared with baseline water column concentrations and the contribution of meroplankton to primary production was important at times, especially when seasonal irradiance in the lake was high. 4. Comparison of the in situ light climate with the threshold of light‐limitation of the phytoplankton indicated that phytoplankton in the lake were only likely to be light‐limited at times of extreme turbidity (e.g. >200 nephelometric turbidity units), particularly when these occur in winter. Therefore, resuspension influenced phytoplankton production mainly via effects on available nutrients and by entraining algae. The importance of each of these varied in time. 5. The partitioning of primary productivity between the water column and sediments in shallow lakes greatly influences the outcome of resuspension events for water column primary productivity.     

DO PHYSICAL FACTORS REGULATE PHYTOPLANKTON DISTRIBUTION PATTERNS IN LARGE, SHALLOW LAKES?

Factors that regulate phytoplankton dynamics in shallow, productive lakes are poorly understood, due to their predisposition for frequent algal blooms and sediment resuspension events. In Lake Apopka, greatest phytoplankton biomass reflects wind-induced resuspension of algae (meroplankton) that exists on the aphotic lake bottom in a layer approximately 5 cm thick; this assemblage is dominated by diatoms (>60% of total biomass) that can occur in resting stages. Once exposed to moderate light, meroplankton are capable of growth and photosynthetic rates comparable with surface populations. In Lake Okeechobee, remote sensing was used to assess the basin-wide distribution of suspended particles. Satellite reflectance values agreed well with in situ particle densities at 20 in-lake stations (average r²; LANDSAT = 0.81, AVHRR = 0.53), and maps of algal blooms (r² = 0.79, p ≤ 0.01). The greatest chlorophyll concentrations occurred in the vicinity of tributary nutrient inputs at the lake's perimeter, while turbidity increased towards the center of the lake reflecting predominant water circulation patterns. These results underscore the importance of physical-biological interactions in lakes.     

Impact of resuspended sediment on zooplankton feeding in Lake Waihola, New Zealand

1. Wind‐induced sediment resuspension in shallow lakes affects many physical and biological processes, including food gathering by zooplankton. The effects of suspended sediment on clearance rate were determined for a dominant cladoceran, Daphnia carinata, and calanoid copepod, Boeckella hamata, in Lake Waihola, New Zealand. 2. Animals were incubated at multiple densities for 4 days in lake water containing different amounts of suspended lake sediment. Rates of harvest of major food organisms were determined for each sediment level (turbidity) from changes in net growth rate with grazer density. 3. Daphnia cleared all food organisms 7–40 μm in length at similar rates, but was less efficient in its removal of free bacteria, phytoplankton <7 μm, and large cyanobacterial filaments. Elevation of sediment turbidity from 2 to 10 nephelometric turbidity units (NTU) (63 mg DW L^?1 added sediment) reduced Daphnia clearance of phytoplankton, heterotrophic flagellates and ciliates by 72–100%, and of amoebae and attached bacteria by 21–44%. Further inhibition occurred at higher turbidity. 4. Boeckella hamata removed microzooplankton primarily, rather than phytoplankton. The rate at which it cleared rotifers was reduced by 56% when turbidity was increased from 2.5 to 100 NTU. 5. In the absence of macrozooplankton, algal growth increased with sediment turbidity, suggesting that sediment also inhibits rotifer grazing. 6. As mid‐day turbidity in Lake Waihola is ≥10 NTU about 40% of the time, sediment resuspension may play a major role in moderating energy flow and structuring pelagic communities in this lake.     

Sedimentation modified by wind induced resuspension in a shallow tropical lagoon (Cote d'Ivoire)

In shallow environments, under certain conditions of fetch, wind velocity, bathymetry and bottom characteristics, resuspension can be generated by wind induced waves. In the tropical Ebrié lagoon, austral trade winds are dominant almost all year long, and their velocity shows a marked diel pattern with maximum speed between noon and midnight. Only austral trade winds with a speed >3 m s⁻¹ allow particle resuspension which is effective for depths<1.5 m. In these areas, significantly higher values of chlorophyll biomass and mineral seston are noted during the windy sequences. Granulometric and mineralogical analyses showed that only the surficial sediment (0–3 cm) was involved in resuspension. This process induces several effects: 1) an increase of the suspended matter concentration in the water and thus a light attenuation due to a higher turbidity, 2) a redistribution in the whole water column of nutrients from the pore water and 3) a removal of the finer fractions from the superficial sediment. On the contrary, for depths>1.5 m, particle sinking is permanent in depressions which are spontaneously transformed into anoxic systems. At the lagoon scale, sedimentation is significantly modified by wind induced resuspension. According to the bathymetry and the distance from a river, three sedimentary facies are recognized. Their grain size distributions are parabolic in areas where resuspension occurs, logarithmic in areas where no resuspension is possible and hyperbolic in the hollows and the main channels. Finally, a large part of the allochthonous inputs (from drainage and rivers) and autochthonous pelagic production is trapped into the Ebrié lagoon and less than 10% of the particles entering the lagoon are exported toward the Atlantic Ocean.     

Environmental control of phytoplankton productivity in turbulent turbid systems

Factors affecting phytoplankton productivity are analysed in turbid systems, such as shallow lakes and rivers. When resuspension from the sediment or loading from the catchment significantly increases inorganic (non-algal) turbidity and hence light attenuation potentials for high production are not realised. Energy available for phytoplankton growth is strongly regulated by underwater light availability which depends on the critical mixing depth, fluctuating light intensities and algal circulation patterns. Higher production rates in shallow waters are often compensated by greater algal respiration due to higher water temperatures when compared to deeper lakes.Total daily integral production of turbulent, turbid environments can be predicted from a combination of easily measured variables such as maximum photosynthetic rates, algal biomass, surface irradiance and some measure of underwater light attenuation.     

Effects of mixing on the pelagic food web in shallow lakes

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The importance of drawdown and sediment removal for the restoration of the eutrophied shallow Lake Kraenepoel (Belgium)

Lake Kraenepoel (Belgium) is a shallow lake (22 ha), divided in two basins since 1957 by a shallow dike. The lake was used for fish farming until World War II and was drawn down about every 5 years to harvest fish. Despite its dense historical carp population, it had clear water and a rich Littorelletea vegetation. During the course of the 20th century, the lake became eutrophic and the Littorelletea vegetation degraded. The northern basin, which was still drawn down about every decade after 1957, retained its clear water and had a dense submerged macrophyte vegetation. The southern basin, which was never drawn down after 1957 and which received direct surface water inputs, had become a turbid shallow lake with phytoplankton blooms in summer. In 2000, efforts were taken to restore the lake: the entire lake was drawn down, the fish community was biomanipulated, nutrient-rich surface water inputs were diverted from the southern basin and sediments were removed (only in the northern basin). Fish biomanipulation and sediment removal were successful in the northern basin, as nutrient levels declined and the Littorelletea vegetation recovered. In the southern basin, sediment analyses indicated that drawdown resulted in sediments with a lower water and organic matter content and water column turbidity decreased after the drawdown. But pH in the southern basin declined to <4, probably because sulphides in the sediment were oxidized during drawdown and sediment desiccation. In contrast, desiccated sediments were removed from the northern basin and pH did not decline below 6 after restoration. In spite of the still high dissolved nutrient concentrations, phytoplankton biomass declined significantly in the southern basin, probably due to acidification. However, no Littorelletea species colonised the lake bottom in the southern basin. Thus, lake drawdown may be a useful management technique to promote clear water conditions in shallow lakes. However, acidification due to sulphide oxidation may be an undesirable outcome and should be considered in drawdown and sediment desiccation manipulations.     

The effect of resuspension on algal production in a shallow lake

Waves cause erosion and resuspension of bottom sediments. In shallow lakes resuspension can take place over most of the lake area and the resuspended matter can stay in suspension for such a long time that the mean light intensity in the lake is reduced, causing reduced algal growth. The increase of suspended matter and light attenuation in the water of lake Tämnaren, Sweden, was found to be proportional to wind velocity to the third power. After each storm increased attenuation of light lasted for a week. The algal production was estimated to be reduced to only 15 % of what it would have been without increased turbidity due to resuspension.     

Resuspension of algal cells by benthivorous fish boosts phytoplankton biomass and alters community structure in shallow lakes

1. Positive effects of fish on algal biomass have variously been attributed to cascading top‐down effects and to nutrient enrichment by fish excretion. 2. Here, we used a combination of field and laboratory approaches to test an additional hypothesis, namely that the physical resuspension of settled algal cells by fish enhances algal biomass and alters community composition. 3. A multi‐lake survey showed that phytoplankton biomass and the fraction of motile algae increased with the concentration of inorganic suspended solids. This correlation could not be explained by wind‐induced resuspension because of the small size of the lakes. 4. In an enclosure experiment, chlorophyll‐a concentration, phytoplankton abundance and inorganic suspended solids increased significantly in the presence of Cyprinus carpio (common carp), but only if the fish had access to the sediment. No such effects were seen when a net prevented carp reaching the sediment. 5. The effects of enhanced nutrients and reduced zooplankton grazing as a result of fish feeding could not (fully) explain these observations, suggesting that the resuspension by carp of settled algae from a surface film on the sediment was the major factor in the outcome of the experiment. 6. An increase in diatoms and green algae (organisms with a relatively large sedimentation velocity) only in enclosures where carp could reach the sediment supported this view. 7. Several lines of evidence indicate that fish‐induced resuspension of algal cells from the sediment is an important mechanism that affects phytoplankton biomass and community composition in shallow lakes.     

Assessment of Components Controlling Phytoplankton Photosynthesis and Bacterioplankton Production in a Shallow,Alkaline, Turbid Lake (Neusiedlersee,Austria)

Various aspects of phytoplankton photosynthesis were investigated between 1968 and 1980 in Neusiedlersee, Austria, a shallow (z̄ = 1.3 m), well mixed lake with much turbidity resulting from frequent resuspension of inorganic sediment particles. The influences of the great environmental variability on biomass, photosynthesis and phytoplankton production were determined and discussed with respect to the increasing nutrient concentrations in the lake. Investigations of bacterioplankton biomass and production revealed their importance for nutrient regeneration and as food for large zooplankton populations. The results are compared with those from several other shallow lakes, and an attempt is made to identify characteristics to distinguish the different types.     

The effect of wave-reduction engineering on sediment resuspension in a large, shallow, eutrophic lake (Lake Taihu)

Frequent resuspension of sediments is recognized as an important process in large shallow lakes, impeding the recovery of eutrophic lakes. A large-scale project, including a wave barrier (3.3 km long) and a soft enclosure, was implemented to reduce wave energy and sediment resuspension in Lake Taihu, eastern China. The effects of the wave-reduction engineering on sediment resuspension and internal nutrient loading were investigated. Results showed that sediment resuspension rates as well as suspended solids (SS) in the areas protected by the wave barrier and the soft enclosure were significantly lower than in the unprotected areas. There was a positive relationship between total phosphorus (TP) and SS; thus internal loading of phosphorus was significantly reduced by the wave-reduction structure. High nutrient levels and phytoplankton biomass persisted during the experiment period, suggesting that additional measures, such as re-establishment of the macrophyte community, must be included to help restore the water quality in such a large, shallow and eutrophic lake.     

Phosphorus dynamics at multiple time scales in the pelagic zone of a large shallow lake in Florida,USA

Phosphorus (P) dynamics in large shallow lakes are greatly influenced by physical processes such as wind-driven sediment resuspension, at times scales from hours to years. Results from long-term (30 year) research on Lake Okeechobee, Florida (area 1,730 km², mean depth 2.7 m) illustrate key features of these P dynamics. Variations in wind velocity result in changes in water column transparency, suspended solids, and total P (TP). In summer there are diurnal changes in TP associated with afternoon winds, and in winter, when strong winds occur for multiple days, monthly average TP remains high compared to summer. The magnitude of daily and seasonal TP changes can exceed 100 μg l⁻¹. Hurricanes and tropical storms also cause extreme changes in TP that are superimposed on seasonal dynamics. When a hurricane passed 80 km south of the lake in October 1999, mean pelagic TP increased from 88 to 222 μg l⁻¹. During large resuspension events, light attenuation is substantially increased, and this influences the biomass and spatial extent of submerged plants, as well as water column TP. In Lake Okeechobee, TP concentrations typically are ∼20 μg l⁻¹ when submerged plants are dense, and soluble reactive P concentrations are reduced below detection, perhaps by the periphyton and plant uptake and by precipitation with calcium at high pH. In contrast, TP exceeds 50 μg l⁻¹ when submerged plants and periphyton are absent due to prolonged deep water, and phytoplankton biomass and algal bloom frequency both are increased. In Lake Okeechobee and other large shallow lakes, complex models that explicitly consider wind-wave energy, hydrodynamics, and sediment resuspension, transport, and key biological processes are needed to accurately predict how lake water TP will respond to different management options.     

The Influence of Macrophytes on Sediment Resuspension and the Effect of Associated Nutrients in a Shallow and Large Lake (Lake Taihu,China)

A yearlong campaign to examine sediment resuspension was conducted in large, shallow and eutrophic Lake Taihu, China, to investigate the influence of vegetation on sediment resuspension and its nutrient effects. The study was conducted at 6 sites located in both phytoplankton-dominated zone and macrophyte-dominated zone of the lake, lasting for a total of 13 months, with collections made at two-week intervals. Sediment resuspension in Taihu, with a two-week high average rate of 1771 g·m^-2·d^-1 and a yearly average rate of 377 g·m^-2·d^-1, is much stronger than in many other lakes worldwide, as Taihu is quite shallow and contains a long fetch. The occurrence of macrophytes, however, provided quite strong abatement of sediment resuspension, which may reduce the sediment resuspension rate up to 29-fold. The contribution of nitrogen and phosphorus to the water column from sediment resuspension was estimated as 0.34 mg·L^-1 and 0.051 mg·L^-1 in the phytoplankton-dominated zone. Sediment resuspension also largely reduced transparency and then stimulated phytoplankton growth. Therefore, sediment resuspension may be one of the most important factors delaying the recovery of eutrophic Lake Taihu, and the influence of sediment resuspension on water quality must also be taken into account by the lake managers when they determine the restoration target.     

Effects of resuspension and eutrophication level on summer phytoplankton dynamics in two hypertrophic areas of Lake Taihu,China

The composition of summer phytoplankton communities in two highly eutrophic areas of Lake Taihu, a large, shallow water body in China (area 2,338 km²; mean depth 2.0 m), was studied in relation to selected environmental variables. Both areas have multiple uses, including drinking water supply, fisheries, and recreation. The first area, Meiliang Bay, is much closer to central lake and subjected to wind-driven sediment resuspension, while the second, at Wuli Lake, is less affected by wave action but is considered more nutrient-rich due to its proximity to human inhabitation. By comparing two lake areas, we attempt to advance the understanding and management of different regions of shallow lakes, mainly differing in resuspension levels and eutrophication levels. A comparison of these two lake areas indicated a higher standing crop of phytoplankton in Wuli Lake than in Meiliang Bay. Canonic Correspondence Analysis indicated that nutrient status and sediment resuspension were the principal factors influencing phytoplankton composition in both areas. The dominant algal group in Meiliang Bay comprised immobile R-type strategists (Reynolds in Growth and reproductive strategies of freshwater phytoplankton, 1988), including filamentous diatoms and green algae (Aulacoseira granulate and Planctonema lauterbornii), and recruitment-beneficial RS-type strategists, mainly the colonial cyanobacteria (Microcystis spp.). The community of Wuli Lake was dominated by S-strategists, principally motile flagellates (Cryptomonas spp.) and C-strategists including small nonmotile centric diatoms and chlorococcal green algae.     

Clay-Turbid Interactions May Not Cascade—A Reminder for Lake Managers

Food web management is a frequently used lake restoration method, which aims to reduce phytoplankton biomass by strengthening herbivorous zooplankton through reduction of planktivorous fish. However, in clay‐turbid lakes several factors may reduce the effectivity of food web management. Increasing turbidity reduces the effectivity of fish predation and weakens the link between zooplankton and phytoplankton. Therefore, the effects of fish stock manipulations may not cascade to lower trophic levels as expected. Additionally, in clay‐turbid conditions invertebrate predators may coexist in high densities with planktivorous fish and negate the effects of fish reductions. For instance, in the stratifying regions of the clay‐turbid Lake Hiidenvesi, Chaoborus flavicans is the main regulator of cladocerans and occupies the water column throughout the day, although planktivorous Osmerus eperlanus is very abundant. The coexistence of chaoborids and fish is facilitated by a metalimnetic turbidity peak, which prevents efficient predation by fish. In the shallow parts of the lake, chaoborids are absent despite high water turbidity. We suggest that, generally, the importance of invertebrate predators in relation to vertebrate predators may change along turbidity and depth gradients. The importance of fish predation is highest in shallow waters with low turbidity. When water depth increases, the importance of fish in the top‐down regulation of zooplankton declines, whereas that of chaoborids increases, the change along the depth gradient being moderate in clear‐water lakes and steep in highly turbid lakes. Thus, especially deep clay‐turbid lakes may be problematic for implementing food web management as a restoration tool.     

Sedimentary losses of phosphorus in some natural and artificial Iowa lakes

Phosphorus sedimentation in four natural and four artificial Iowa lakes was measured by using sediment traps to determine if sedimentary phosphorus losses were greater in artificial lakes than in natural lakes and the limnological factors influencing phosphorus loss rates. Mean phosphorus sedimentation rates ranged from 13.3 to 218 mg · m^–2 day^–1. Although phosphorus sedimentation rates for the natural lakes as a group did not differ significantly from the rates for artificial lakes, there were significant differences among individual lakes. Phosphorus sedimentation rates also varied significantly during different seasons at different locations within a lake and at different depths within a location. Despite the variance, phosphorus sedimentation rates were strongly correlated with inorganic sediment concentrations and inorganic matter sedimentation rates, thus suggesting that inorganic sediments influence phosphorus sedimentation rates. When Iowa data were combined with data from published studies, mean sedimentation rates were directly correlated with mean chlorophyll a concentrations of the lakes. These data strongly suggest that sedimentation rates as measured by sediment traps are strongly influenced by the trophic status of a lake. Though sedimentation rates were higher in the more productive lakes, it is suggested that these rates represent only gross sedimentation rates rather than net sedimentation rates because of resuspension and resedimentation of bottom sediments.     

Does sediment resuspension have persistent effects on phytoplankton? Experimental studies in three shallow lakes

1. The persistence of effects of sediment resuspension on chlorophyll a , phytoplankton production and plant nutrients was examined by artificially resuspending sediment in enclosures in three shallow lakes and monitoring concentrations for 4–8 days. Realism was assessed by relating initial suspended solids concentrations to those observed during natural wind events.
2. Positive effects on the phytoplankton, persisting for at least 4–6 days were detected in eight of the nine experiments, and similar effects on nutrient concentrations were also common, although suspended solids had normally returned to control levels within 24 h
3. The phytoplankton during the periods of persistence was normally dominated by planktonic rather than benthic/meroplanktonic genera.
4. Several of the positive responses appeared to be associated with relief of nitrogen deficiency in the algae.
5. Persistent effects from previous resuspension events may cause baseline concentrations of phytoplankton and nutrients in shallow lakes to be overestimated and the effects of resuspension on phytoplankton and nutrients to be underestimated     

Why biomanipulation can be effective in peaty lakes

The effects of fish stock reduction (biomanipulation) was studied in an 85 ha shallow peaty turbid lake. The lake cleared in a 4-week period in April–May 2004, which demonstrated that biomanipulation can be effective in peaty lakes. We demonstrated that it is possible to reduce the fish stock to <25 kg ha⁻¹ benthivorous fish and <15 kg ha⁻¹ planktivorous fish, sufficiently low to switch the lake from a turbid to a clear state. Knowledge of lake morphology, fish stock, fish behaviour, and a variety of fishing methods was necessary to achieve this goal. It is expected that continuation of fisheries to remove young of the year planktivorous species is needed for several years, until macrophytes provide sufficient cover for zooplankton and can compete with phytoplankton. Cladocerans developed strongly after fish removal. The clearing of the lake coincided with a sudden decrease of filamentous cyanobacteria and suspended detritus, and a strong increase of Bosmina. We assume that Bosmina was able to reduce filamentous prokaryotes and detritus. After the disappearance of the cyanobacteria, Bosmina disappeared too. After the clearing of the lake Daphnia dominated in zooplankton and apparently was able to keep phytoplankton levels low. In our case, wind resuspension did not prevent biomanipulation from being successful. No correlation between windspeed and turbidity was found, neither in an 85 ha nor in a 230 ha shallow peaty lake. Regression analysis showed that on average 50% of the amount of suspended detritus can be explained by resuspension by fish and 50% by phytoplankton decomposition. The main goal of this biomanipulation experiment, clear water and increased submerged plant cover in a shallow peaty lake, was reached.     

Relative importance of periphyton and phytoplankton in turbid and clear vegetated shallow lakes from the Pampa Plain (Argentina): a comparative experimental study

We analyzed experimentally the relative contribution of phytoplankton and periphyton in two shallow lakes from the Pampa Plain (Argentina) that represent opposite scenarios according to the alternative states hypothesis for shallow lakes: a clear lake with submerged macrophytes, and a turbid lake with high phytoplankton biomass. To study the temporal changes of both microalgal communities under such contrasting conditions, we placed enclosures in the littoral zone of each lake, including natural phytoplankton and artificial substrata, half previously colonized by periphyton until a mature stage and half clean to analyze periphyton colonization. In the clear vegetated shallow lake, periphyton chlorophyll a concentrations were 3–6 times higher than those of the phytoplankton community. In contrast, phytoplankton chlorophyll a concentrations were 76–1,325 times higher than those of periphyton in the turbid lake. Here, under light limitation conditions, the colonization of the periphyton was significantly lower than in the clear lake. Our results indicate that in turbid shallow lakes, the light limitation caused by phytoplankton determines a low periphyton biomass dominated by heterotrophic components. In clear vegetated shallow lakes, where nitrogen limitation probably occurs, periphyton may develop higher biomass, most likely due to their higher efficiency in nutrient recycling.     

A comparison of the palaeolimnology of Peipsi and Võrtsjärv: connected shallow lakes in north-eastern Europe for the twentieth century, especially in relation to eutrophication progression and water-level fluctuations

We applied a multi-proxy palaeolimnological approach to provide insights into the natural variability and human-mediated trends of two interconnected temperate large shallow lakes, Peipsi and Võrtsjärv, during the twentieth century. The history of the lakes was assessed on the basis of age-related changes in the sediment main constituents (water, organic matter and carbonate), sub-fossil pigments, diatom assemblages and organic matter dissolved in pore water. The temporal changes in the palaeodata indicate an increase of the in-lake biological production in both lakes from about the 1960s, suggesting enhanced nutrient inputs. In subsequent decades, the gradual increase of autochthonous organic matter becomes more obvious, indicating progressive eutrophication of the lakes. Palaeolimnological indicators from the sediment record of Lake Peipsi indicate a slight recession of the lake’s eutrophication in the 1990s but not for Lake Võrtsjärv. The results of the study also suggest that after the lakes became eutrophied, the climatically induced water-level fluctuations ceased to be the main driver determining the abundance of phytoplankton. Responses of the lakes to human-induced impacts are better recorded in the sediments of Lake Peipsi than in those of Lake Võrtsjärv, which is shallower of the two and where the wave-induced resuspension of deposits markedly smooths or erases the signals of environmental changes. The results of the investigation expand the knowledge on how large shallow lakes respond to human-mediated and natural perturbations, including those in the lake catchment areas and the capability of the lakes to store the chronology and sequence of these changes.