High resolution measurements of sediment resuspension above an accumulation bottom in a stratified lake

A detailed record of suspended particulate matter (SPM) concentrations in the benthic boundary layer (BBL) 1.5 m above an accumulation bottom and 13.5 m below the surface was obtained from frequent (30 min interval) beam attenuation measurements made with a Sea Tech transmissometer in the main basin of Lake Erken, a moderately deep (mean depth 9 m, maximum depth 21 m) dimictic lake in central Sweden. Concentrations of SPM (g m^–3) were not as strongly correlated to the beam attenuation coefficient (c, [m^–1]), as were concentrations of the inorganic SPM fraction. Apparently, this was caused by large optically inactive organic particles which significantly affected the measurements of SPM, but had little effect on the attenuation of light.When the water column was thermally stratified, SPM concentrations in the BBL showed a seasonal increase which was related to an increase in the thermocline depth. As the epilimnion deepened, there was also a marked increase in the occurrence of rapid and large changes in SPM concentration. After the loss of stratification, the amount of SPM and the temporal variability in its concentration was reduced. Since surface waves could not influence sediment resuspension at the depth of measurement, these data show the importance of internal waves in promoting sediment resuspension in areas of sediment accumulation. The relatively short period in each summer, when the thermocline reaches a sufficient depth to allow for resuspension over accumulation bottoms, can have important consequences for both the redistribution of lake sediments and the internal loading of phosphorus.     

Sediment resuspension and light attenuation in Peoria Lake: can macrophytes improve water quality in this shallow system?

We examined sediment resuspension and light attenuation in relation to the potential for macrophytes to improve water quality conditions in Peoria Lake, Illinois (U.S.A.). The lake exhibited high total suspended solids (TSS) loading and retention of predominantly fine-grained particles in 2000. Large fetches along prevailing wind rose, coupled with shallow morphometry and sediment particles composed of >90% silt and clay resulted in frequent periods of sediment resuspension. As calculated (wave theory) shear stress increased above the critical shear stress (measured experimentally), turbidity increased substantially at a resuspension monitoring station. Resuspension model explorations suggested that establishment of submersed aquatic macrophytes could substantially reduce sediment resuspension in Peoria Lake. However, K _d is currently very high, while Secchi transparency low, at in-lake stations. Thus, in order to establish a persistent macrophyte population in the lake to control resuspension, the underwater light regime will have to improve quite dramatically.     

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.     

Aspects of modelling sediment transport and night conditions in Lake Marken

In lake Marken (Markermeer), a shallow lake in the Netherlands, sediment transport and suspended solids concentration are dominated by wind induced resuspension of sediment. The suspended solids concentration in the lake determines the attenuation of light. A 2-dimensional sediment transport model, STRESS-2d, was used to estimate the special distribution of the silt content for periods with different wind conditions. The model was calibrated using measured values of the fall velocity distributions of sediment, freshly deposited material and suspended solids. The specific light attenuation coefficient of the various sediment classes was also measured and used in the light attenuation model, CLEAR, together with simulation results from the STRESS-2d model, to simulate the light attenuation variations in time and space. By comparison of the simulation results for different scenarios, the influence of regional planning features on the sediment balance and the light attenuation can quantified. A provisional example of this method is presented.     

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.     

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.     

Fisheries management as an additional lake restoration measure: biomanipulation scaling-up problems

Restoration of the highly eutrophic Reeuwijk lakes (ca. 700 ha) started in 1986 by reducing the external phosphorus loading. As an additional measure to improve the quality of the lake water, the structure of the fish population in Lake Klein Vogelenzang (18 ha) was altered in 1989 by the removal of ca. 100 kg ha^–1 bream from the lake in April and December. This constituted about 50% of the total bream biomass in the lake.The fish-stock reduction in April, 1989, was initially followed by high phosphorus concentrations, probably the result of considerable phosphorus release from the sediments. The resulting heavy algal blooms that occurred reduced the transparency to very low values. During the summer the zooplankton population increased markedly in numbers coinciding with reductions in total suspended matter including (blue-green) algae. A great improvement in Secchi-disc transparency was observed and by the end of December, 1989, the bottom of the lake (1.5–2.0 m) was visible. After heavy storms in January and February 1990, transparency dropped to < 1 m as a result of resuspension of high concentrations of suspended matter from the bottom sediments. Although transparency over the rest of 1990 was higher than in 1988, i.e. the year preceding the removal of fish (biomanipulation), it was lower than expected, based on the results of 1989. The study shows that technical and biological factors can cause serious management problems for the implementation of biomanipulation in larger water bodies.     

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.     

Quantifying measures to limit wind-driven resuspension of sediments for improvement of the ecological quality in some shallow Dutch lakes

Although phosphorus loadings are considered the main pressure for most shallow lakes, wind-driven resuspension can cause additional problems for these aquatic ecosystems. We quantified the potential effectiveness of measures to reduce the contribution of resuspended sediments, resulting from wind action, to the overall light attenuation for three comparable shallow peat lakes with poor ecological status in the Netherlands: Loosdrecht, Nieuwkoop, and Reeuwijk (1.8–2.7 m depth, 1.6–2.5 km fetch). These measures are: 1. wave reducing barriers, 2. water level fluctuations, 3. capping of the sediment with sand, and 4. combinations of above. Critical shear stress of the sediments for resuspension (V _crit), size distribution, and optical properties of the suspended material were quantified in the field (June 2009) and laboratory. Water quality monitoring data (2002–2009) showed that light attenuation by organic suspended matter in all lakes is high. Spatial modeling of the impact of these measures showed that in Lake Loosdrecht limiting wave action can have significant effects (reductions from 6% exceedance to 2% exceedance of V _crit), whereas in Lake Nieuwkoop and Lake Reeuwijk this is less effective. The depth distribution and shape of Lake Nieuwkoop and Lake Reeuwijk limit the role of wind-driven resuspension in the total suspended matter concentration. Although the lakes are similar in general appearance (origin, size, and depth range) measures suitable to improve their ecological status differ. This calls for care when defining the programme of measures to improve the ecological status of a specific lake based on experience from other lakes.     

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.     

Effects of Inorganic Turbidity on the Phytoplankton of an Amazonian Lake Impacted by Bauxite Tailings

The dumping of bauxite tailings into Batata Lake, an Amazonian clear-water lake, generated high levels of turbidity and caused a serious decrease in phytoplankton densities, which could possibly be the result of a photosynthetic limitation due to light attenuation together with an increase in algal sinking due to the adhesion of clay particles. This study aimed to investigate the sinking process through the addition of different suspended clay concentrations in columns containing Batata lake water. Since no effect of the suspended clays on Batata Lake phytoplankton sinking was observed, it was then evaluated, under laboratory conditions, whether the low conductivity of the Batata Lake water could interfere with the algae-clay aggregation process. Cultures of two algal species known to be capable to aggregate to Batata Lake suspended clays in algal culture medium: Staurodesmus convergens and Phormidium amoenum, were added to both the low conductivity Batata Lake water (14 μS cm^?1) and the high conductivity algal culture media (WC – 300 μS cm^?1 and Z8 – 560 μS cm^?1) together with Batata lake suspended clays. In both algal culture media and Batata lake water the two species had their sinking accelerated due to clay adhesion. It is thus suggested that the decrease in phytoplankton densities recorded in Batata Lake may not be related to an increase in phytoplankton loss rates due to algal-clay aggregation, but rather are a consequence of decreasing growth rates because of light attenuation.     

Scaling of Pelagic Metabolism to Size,Trophy and Forest Cover in Small Danish Lakes

ABSTRACT We tested whether pelagic light and nutrient availability, metabolism, organic pools and CO₂-supersaturation were related to lake size and surrounding forest cover in late summer–autumn measurements among 64 small (0.02–20 ha), shallow seepage lakes located in nutrient-rich, calcareous moraine soils in North Zealand, Denmark. We found a strong implicit scaling to lake size as light availability increased significantly with lake size while nutrient availability, phytoplankton biomass and dissolved organic matter declined. Forest lakes had significantly stronger net heterotrophic traits than open lakes as higher values were observed for light attenuation above and in the water, dissolved organic matter, pelagic community respiration (R) relative to maximum gross primary production (R/GPP) and CO₂-supersaturation. Total-phosphorus was the main predictor of phytoplankton biomass (Chl) despite a much weaker relationship than observed in previous studies of larger lakes. Maximum gross primary production increased with algal biomass and decreased with dissolved organic matter, whereas community respiration increased with dissolved organic matter and particularly with gross primary production. These results suggest that exogenous organic matter supplements primary production as an energy source to heterotrophs in these small lakes, and particularly so in forest lakes experiencing substantial shading from the forest and dissolved humic material. This suggestion is supported by 20–30-fold CO₂ supersaturation in the surface water of the smallest forest lakes and more than sixfold supersaturation in 75% of all measurements making these lakes among the most supersaturated temperate lakes examined so far.     

Summer sedimentation in six shallow arctic lakes

The movement of sediment between the lake bottom and water column of shallow lakes can be sizeable due to the large potential for resuspension in these systems. Resuspended sediments have been shown to alter phytoplankton community composition and elevate water column production and nutrient concentrations. We measured the summer sedimentation rates of two lakes in 2003 and six lakes in 2004. All lakes were shallow and located in the Alaskan Arctic. In 2004, turbidity, light attenuation, total sediment:chlorophyll a mass in the sediment traps, and thermal stratification were also measured in each of the lakes. The sediment:chlorophyll a mass was much greater than if the sediment was derived from phytoplankton production in all of the lakes, indicating that the source of the sedimenting material was resuspension and allochthonous inputs. Consistent with these findings, the temporal variation in sedimentation rate was synchronous between most lakes, and sedimentation rate was positively related to wind speed and rainfall suggesting that sedimentation rate was strongly influenced by landscape-scale factors (e.g., wind and rain events). Two of the lakes are located on deposits of loess that accumulated during past glacial periods. These two lakes had sedimentation rates that were significantly greater and more variable than any of the other lakes in the study, as well as high turbidity and light attenuation. Our results indicate that sedimentation in these shallow arctic lakes is supported primarily by allochthonous inputs and resuspension and that landscape-scale factors (e.g., weather and geology) impact on the transport of materials between the lake bottom and water column. Handling editor: J. Saros     

Structural and functional characteristics of epiphyton and epipelon in relation to their distribution in Lake Vechten

Epiphyton and epipelon were quantitatively collected, respectively, from the submerged macrophytes and the sandy lake bottom of Lake Vechten (The Netherlands). On a weight basis, epiphyton was maximal in autumn and epipelon in summer. In winter the chemical composition of epiphyton and epipelon was similar. In summer the epiphyton had on a unit weight basis more organic matter and carbonate, and had per unit organic matter a higher algal number, nitrogen and energy content than the epipelon. Algae predominating the epiphyton were filamentous greens and pennate diatoms; those in the epipelon were pennate diatoms and blue-green algae. In both cases, species known to frequent the phytoplankton were abundant. The diatoms were quantified using paper chromatographic pigment analyses. Both the epiphyton and the epipelon exhibited maximal photosynthesis in mid summer. That light was generally the limiting factor was evident from periphyton developed on artificial substrates. This periphyton differed widely in its composition from that on the natural substrates, mainly because the latter collected much more sedimenting matter.In dense Ceratophyllum stands light was severely attenuated and the significant gradients in oxygen and pH were caused by the differences with depth in the proportions of photosynthesis and respiration. The oxygen content and pH at the bottom decreased owing to epipelic respiration. The epiphytic composition depended greatly on the degree of light attenuation. The epiphytic and epipelic respiration, except during part of the early summer, exceeded photosynthesis on a 24 h basis; this included the macrophytic photosynthesis during the time the vegetation was maximally developed. During the growing season import of organic matter, i.e. deposited seston, greatly exceeded that due to the photosynthetic production. After the summer maximum, the epipelon decreased faster than predicted from its oxygen exchange. It was concluded that sedimentation and resuspension determined mainly the changes in epiphyton and epipelon. Especially when covered with vegetation, the lower littoral of Lake Vechten plays a large part in the aerobic decomposition of sestonic organic matter.     

Light-limited algal growth in Lake Loosdrecht: steady state studies in laboratory scale enclosures

Phytoplankton growth in the shallow, turbid Lake Loosdrecht (The Netherlands) is importantly influenced by light availability, and thus the concentrations of the various light-attenuating materials. The system is highly eutrophic and supports an algal biomass of ca. 160 mg Chl m^–3. A model is proposed here which predicts algal growth in the lake as a function of the light received and subsequent attenuation in the water column by phytoplankton, tripton and background colour. The model is based on an energy balance which relates growth rate to the true growth yield on light energy and the energy demand for cell maintenance. The coefficients for energy conversion (Y = 0.002 gDW kJ^–1) and cell maintenance (µ_e = 0.031 day^–1) were determined from steady state growth kinetics of Prochlorothrix hollandica in light-limited laboratory flow systems with the same depth as the lake and receiving summer average conditions of irradiance. Light attenuation by phytoplankton and tripton were quantified using specific attenuation coefficients: 0.011 m² mg^–1 Chl for the phytoplankton and 0.23 m² g^–1 DW for tripton.The growth studies demonstrated that Lake Loosdrecht can support a much higher algal biomass in the absence of non-algal particulate matter. The proposed model is used to predict chlorophyll a concentrations in dependence on growth rate and levels of tripton. Since approximately 75% of the sestonic dry weight in Lake Loosdrecht may be attributed to tripton, it is concluded that the algal biomass is markedly lowered by the abundance of tripton in the water column. A knowledge of the sources and fate of tripton in the lake is thus of fundamental importance in modelling phytoplankton dynamics.     

ESTIMATION OF CHLOROPHYLL a FROM TIME SERIES MEASUREMENTS OF HIGH SPECTRAL RESOLUTION REFLECTANCE IN AN EUTROPHIC LAKE

We acquired high spectral resolution reflectance data in Carter Lake, a eutrophic oxbow on the Iowa–Nebraska border, from April 1995 to April 1996. Chlorophyll a, total seston, sestonic organic matter, Secchi depth, and nephelometric turbidity were determined for each respective spectral measurement. Changes in algal taxonomic structure and abundance coincided with the development and senescence of a midsummer through autumn bloom of Anabaena. Taxonomic structure was more diverse in late winter and spring when Synedra sp. (diatom) and several chlorophytes and dinoflagellates were present. Overall, chlorophyll a varied from about 20 to 280 μg·L^?1, Secchi transparency from 18 to 74 cm, and seston dry weight from 11 to 48 mg·L^?1 in February and September, respectively. Particulate matter completely dominated lake water light attenuation. Dissolved organic matter had low optical activity. The most sensitive spectral feature to variation in chlorophyll a concentration was the magnitude of the scattering peak near 700 nm. The 700-nm peak correlated to chlorophyll concentration through the relationships between algal pigment absorption near 670 nm and the cell biomass and surface-related scattering signal in the near infrared. An algorithm relating the height of the 700-nm reflectance peak above a reference baseline between 670 and 850 nm to chlorophyll a was accurate and robust despite large variations in optical constituents caused by both strong seasonality in the algal system and short-term variations in seston from wind-induced sediment resuspension. The present algorithms were successfully used in other systems with different seasonality and productivity patterns. The coefficients of the models relating chlorophyll a and spectral reflectance variables appeared to be ecosystem specific: both the intercept and slope for the models in this study were moderately lower than for several other recently published results. We validated our algorithm coefficients with a second, independent dataset. The standard error for chlorophyll a prediction was ±28 μg·L^?1.     

Coupling dissolved organic carbon,CO2 and productivity in boreal lakes

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Phytoplankton biomass in Lake Xolotlán (Managua): Its seasonal and horizontal distribution

Some well-documented studies on restoring eutrophic lake systems in The Netherlands by fish stock management have been evaluated with the emphasis on the role of macrophytes. Furthermore, the factors determining the light climate for submerged macrophytes in a large shallow eutrophic lake (Lake Veluwe) have been assessed and the potential success of biomanipulation in large scale projects is discussed. Today relatively little attention has been paid to macrophyte management although the importance of macrophytes in lake restoration has been recognized regularly. The biomanipulation strategy was successful in small scale projects. In a large scale project, however, wind-induced resuspension may largely determine the underwater light climate through attenuation by the water column and periphytic layer. Therefore, restoration of relatively large waterbodies by fish stock management only is expected not to lead to any noteworthy improvement of the light climate for submerged macrophytes. Additional measures aimed at reducing wind-induced resuspension of sediment particles and reestablishing of the macrophyte stands are required for successful biomanipulation strategies. Water quality managers should pay more attention to macrophyte stands in biomanipulation projects because macrophytes enhance a more stable and diverse ecosystem. Restoration objectives and the methods of their achievement must be carefully planned since an abundant submerged macrophyte vegetation may have undesirable effects as well.     

Paleolimnology of Two Shallow Lakes in Central Alberta,Canada

An interpretation of postglacial change in water quality and productivity has been made for two shallow lakes in Central Alberta, Canada, namely Hastings Lake (longitude 113° 00′ W; latitude 53° 30′ N) and Lac Ste. Anne (longitude 114° 21′ W; latitude 53° 41′ N). Erosion rates around Lac Ste. Anne have remained constant. Similarly productivity has changed little although macrophytes contributed more to total production during the early stages. Hastings Lake has responded more sensitively to changes in the balance of precipitation and evaporation. From 5500 to 4000 year B. P. it was shallower than at present. Productivity during this early phase was less but macrophytes and allochthonous organic matter (i. e., leaf litter) probably contributed a greater proportion of the organic influx. After water levels rose productivity increased remaining steady until 2500 year B. P. when a slight decline occurred. Throughout the high water period oxygen depletion has not been serious. Any period of reducing conditions has been brief. Productivity has never been nutrient-limited. Elevation of the lake surface increased potential volume for production and reduced turbulent resuspension of bottom sediments permitting greater light penetration, and enhanced algal production. Difference between the sedimentary record of these two lakes, as well as the two basins of Hastings Lake, demonstrates the individualistic responses of basins and lakes to climatic events.     

The influence of light environment on depth of visual feeding by larvae and fry of Coregonus pollan (Thompson) in Lough Neagh

The depth limits for visual feeding by pollan larvae and fry, Coregonus pollan , were determined in Lough Neagh, Northern Ireland, by combining laboratory estimates of the minimum light threshold for feeding activity with field measurements of spectral, diurnal and seasonal changes in light penetration. Although feeding thresholds were lowest in the blue light (400–490 nm), the rapid attenuation of this region in the lake meant that the depth limits of feeding in situ were determined by response to the green region (490–595 nm). Effects of changes in irradiance and light penetration on the depth of feeding are considered during a seasonal study. Increased sensitivity, as the juvenile stages develop, may be offset by reduced light penetration due to algal growth. Over an annual period the maximum feeding depth for pollan varied between 2.6 and 4.1 m. In the juvenile stages of two other predatory species from the lake, perch, Perca fluviatilis , and pike, Esox lucius , the corresponding ranges were 1.1–1.9 m and 2.7–4.6 m respectively.