The Absorption of Light in Lakes: Negative Impact of Dissolved Organic Carbon on Primary Productivity

Colored dissolved organic matter (CDOM) absorbs a substantial fraction of photosynthetically active radiation (PAR) in boreal lakes. However, few studies have systematically estimated how this light absorption influences pelagic primary productivity. In this study, 75 boreal lakes spanning wide and orthogonal gradients in dissolved organic carbon (DOC) and total phosphorus (TP) were sampled during a synoptic survey. We measured absorption spectra of phytoplankton pigments, CDOM, and non-algal particles to quantify the vertical fate of photons in the PAR region. Area-specific rates of gross primary productivity (PP_A) were estimated using a bio-optical approach based on phytoplankton in vivo light absorption and the light-dependent quantum yield of photochemistry in PSII measured by a PAM fluorometer. Subsequently, we calculated the effects of CDOM, DOC, and TP concentration on PP_A. CDOM absorbed the largest fraction of PAR in the majority of lakes (mean 56.3%, range 36.9–76.2%), phytoplankton pigments captured a comparatively minor fraction (mean 6.6%, range 2.2–28.2%). PP_A estimates spanned from 45 to 993 mg C m^?2 day^?1 (median 286 mg C m^?2 day^?1). We found contrasting effects of CDOM (negative) and TP (positive) on PP_A. The use of DOC or CDOM as predictors gave very similar results and the negative effect of these variables on PP_A can probably be attributed to shading. A future scenario of increased DOC, which is highly correlated with CDOM in these lakes, might impose negative effects on areal primary productivity in boreal lakes.     

Growth rate of alpine phytoplankton assemblages from contrasting watersheds and N‐deposition regimes exposed to nitrogen and phosphorus enrichments

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Light limitation of phytoplankton development in an oligotrophic lake - Loch Ness, Scotland

• 1 The underwater light climate in Loch Ness is described in terms of mixing depth (Z_m) and depth of the euphoric zone (Z_eu). During periods of complete mixing, Z_m equates with the mean depth of the loch (132 m), but even during summer stratification the morphometry of the loch and the strong prevailing winds produce a deep thermocline and an epilimnetic mixed layer of about 30 m or greater. Hence, throughout the year the quotient Z_m/Z_eu is exceptionally high and the underwater light climate particularly unfavourable for phytoplankton production and growth.
• 2 Phytoplankton biomass expressed as chlorophyll a is very low in Loch Ness, with a late summer maximum of less than 1.5 mg chlorophyll a m^-3 in the upper 30 m of the water column. This low biomass and the resulting very low photosynthetic carbon fixation within the water column are evidence that a severe restraint is imposed on the rate at which phytoplankton can grow in the loch.
• 3 The chlorophyll a content per unit of phytoplankton biovolume and the maximum, light-saturated specific rate of photosynthesis are both parameters which might be influenced by the light climate under which the phytoplankton have grown. However, values obtained from Loch Ness for both chlorophyll a content (mean 0.0045 mg mm^-3) and maximum photosynthetic rate (1–4 mg C mg Chla^-1 h^-1) are within the range reported from other lakes.
• 4 Laboratory bioassays with the natural phytoplankton community from Loch Ness on two occasions in late summer when the light climate in the loch is at its most favourable, suggest that even then limitation of phytoplankton growth is finely balanced between light and phosphorus limitation. Hence, for most of the year, when the light climate is less favourable, phytoplankton growth will be light limited.
• 5 Quotients relating mean annual algal biomass as chlorophyll a (c. 0.5 mg Chla m^-3) and the probable annual specific areal loading of total phosphorus (0.4–1.7 g TP m^-2 yr^-1) suggest that the efficiency with which phytoplankton is produced in Loch Ness per unit of TP loading is extremely low when compared with values from other Scottish lochs for which such an index has been calculated. This apparent inefficiency can be attributed to suppression of photosynthetic productivity in the water column due to the unfavourable underwater light climate.
• 6 These several independent sources of evidence lead to the conclusion that phytoplankton development in Loch Ness is constrained by light rather than by nutrients. Loch Ness thus appears to provide an exception to the generally accepted paradigm that phytoplankton development in lakes of an oligotrophic character is constrained by nutrient availability.

     

Primary productivity,phytoplankton and limiting nutrient factors in labrador lakes

The primary productivity of some lakes and reservoirs in western Labrador was measured by the ¹⁴C method in order to determine the range of productivities and the effects of impoundment. No primary productivity data previously existed for this part of Canada. Both the primary productivity and standing crops of phytoplankton were found to be low in a newly impounded lake but later rose to levels greater than in surrounding natural lakes. In nutrient enrichment experiments, carbon was never found to be limiting but phosphorus stimulated primary productivity when added alone or in combination with nitrogen.     

Nutrient enrichment experiments in three central Florida lakes of different trophic states

Seasonal nutrient enrichment experiments (short-term bioassays) were conducted in three Florida lakes of different trophic states to determine the effects of addition of various nutrient combinations upon chlorophyll a and phytoplankton standing crops. Nutrient enriched surface water samples with crustacean zooplankton removed were incubated in situ in clear polyethylene bags for 3 to 6 days. The 2⁵ factorial design employed two levels (ambient and enriched) of each of five nutrients [NH₄ ⁺, PO _inf4 ^sup3− , Fe⁻ -EDTA, SiO _inf3 ^sup2− and a cation (Ca²⁺ or K⁺) or trace elements]. Ammonium produced significant increases in chlorophyll a and phytoplankton standing crops in all experiments. Phosphate produced similar results in the mesotrophic lake, but the eutrophic lakes had both positive and nonsignificant responses which varied seasonally between lakes. Iron increased chlorophyll a in most experiments but affected total phytoplankton standing crop only during the summer and fall. Silicon had negative effects in some experiments. Cations and trace elements produced marked differences between lakes for chlorophyll a, but total phytoplankton standing crop showed few significant responses. Synergistic responses to two- and three-factor interactions were observed in all lakes. Differences in the responses of phytoplankton taxonomic divisions to enrichment may be responsible for much of the between lake variation in chlorophyll a and total phytoplankton volume responses. Nutrient limitations in these lakes are discussed and related to limnological factors and predictive models.     

Increase of atmospheric CO2 promotes phytoplankton productivity

It is usually thought that unlike terrestrial plants, phytoplankton will not show a significant response to an increase of atmospheric CO₂. Here we suggest that this view may be biased by a neglect of the effects of carbon (C) assimilation on the pH and the dissociation of the C species. We show that under eutrophic conditions, productivity may double as a result of doubling of the atmospheric CO₂ concentration. Although in practice productivity increase will usually be less, we still predict a productivity increase of up to 40% in marine species with a low affinity for bicarbonate. In eutrophic freshwater systems doubling of atmospheric CO₂ may result in an increase of the productivity of more than 50%. Freshwaters with low alkalinity appeared to be very sensitive to atmospheric CO₂ elevation. Our results suggest that the aquatic C sink may increase more than expected, and that nuisance phytoplankton blooms may be aggravated at elevated atmospheric CO₂ concentrations.     

Carbon-14 Uptake by Picoplankton and Total Phytoplankton in Eight New Zealand Lakes

Eight New Zealand lakes were surveyed for ¹⁴C uptake by phytoplankton as a function of light intensity. The results support the view that the photosynthetic picoplankton is an important contributor to primary productivity in oligotrophic lakes but is relatively unimportant in more eutrophic lakes. A comparison of carbon uptake vs. light intensity characteristics (P vs. I) of the picoplankton size class vs. that of the total phytoplankton community supports the view that the picoplankton size class may be adapted to utilization of dimmer light.     

Photosynthetic parameters of the entire euphotic phytoplankton of the Bransfield Strait,Summer 1985

Summary The photosynthetic parameters ^B, P^Bm, and I_k were derived from the relationship between primary productivity (P) and irradiance (I) for mixed samples, each mixture consisting of seven samples of phytoplankton collected from different depths, between the surface and 35 m at eight different sampling stations in the Bransfield Strait in January–February 1985. The ^B values obtained in this study were considerably higher than those generally reported for phytoplankton from a single depth in polar waters. These higher values may be attributed to the fact that most of the phytoplankton in the mixed samples came from depths with limiting irradiances, which had a greater yield than surface phytoplankton. The P^Bm value we obtained was lower than that reported by other authors, possibly because the mixed sample phytoplankton have saturation points at low light intensity. Our I_K values confirmed the ability of phytoplankton in these latitudes to reach maximum photosynthesis rates with low densities of photonic flux. When treated as independent parameters, the ^B and P _m ^B values showed a significant correlation for each sampling station. The efficiency of the entire euphotic phytoplankton of the water column, estimated as ₀ ^eu Pdz/ ₀ ^eu Bdz or Integral Assimilation Number (IAN), showed a predictable significant correlation with the ^B values calculated from the entire euphotic phytoplankton. Areas with high surface biomass were as productive as those with low surface phytoplankton biomass. Areas with low surface phytoplankton biomass showed higher ^B values than areas with high surface biomass, which suggests, that, the ^B values for the entire photosynthesizing phytoplankton in the water column may have greater predictive value than the ^B values for the phytoplankton from a single depth. This finding should be taken into account when satellites are used for studying marine production processes, especially in areas with very different euphotic depths and surface phytoplankton concentrations.This research was made possible by grants from Chilean Antarctic Institute (INACH); Chilean National Commission for Science and Technology (FONDECYT 0854-1989); and University of Chile (DTI N-2449-88)     

Carbon fixation and carbon availability in marine phytoplankton

It is widely believed that inorganic C does not limit the rate of short-term photosynthesis, the net productivity, or the maximum biomass, of marine phytoplankton. This lack of inorganic C restriction is less widely believed to hold for phytoplankton in many low alkalinity freshwaters or for seaweed in nutrient-enriched rock pools. These views are examined in the context of the physical chemistry of the inorganic C system in natural waters and of the ways in which various taxa of phytoplankton deal with inorganic C and discriminate between ¹²C and ¹³C. Using this information to interpret data obtained in the ocean or in freshwater suggests that short-term photosynthesis, production rate, and achieved biomass, of phytoplankton are rarely limited by inorganic C supply but, rather, that the widely suggested factors of limited light, nitrogen or phosphorus supply are the resource inputs which restrict productivity. Global change, by increasing atmospheric CO₂ partial pressure and global mean temperatures, is likely to increase the mean CO₂ concentration in the atmosphere, but the corresponding change in the oceans will be much less. There are, however, genotypic differences in the handling of inorganic C among the diversity of marine phytoplankton, and in impact on use of limiting nutrients, so increases in the mean CO₂ and HCO₃ ^- concentrations in surface ocean waters could cause changes in species composition. However, the rarity of inorganic C limitation of marine phytoplankton short-term photosynthesis, net productivity, or the maximum biomass, in today's ocean means that global change is unlikely to increase these three values in the ocean.     

Seasonality in Nutrients and Phytoplankton Production in Two Shallow Lakes: Waigani Lake,Papua New Guinea,and Barton Broad,Norfolk, England

Waigani Lake, near Port Moresby, Papua New Guinea and Barton Broad, Norfolk, England are both shallow lakes nutrient-enriched from sewage effluent disposal. In Waigani Lake phytoplankton biomass varied seasonally with lower levels (100-200 mg chlorophyll α m⁻³) during the wet season increasing to over 400 mg chlorophyll α m⁻³ at the end of the dry season. Secchi disc depths varied between 0. 11 and 0. 34 m. Phytoplankton productivity in Waigani Lake was very high throughout the year (range: A_max 4,370-21,000 mg O₂ m⁻³ h⁻¹) but production was lower during the wet season (range: A_max 4,370-12,700 mg O₂ m⁻³ h⁻¹). High surface productivity was recorded from August to December except on sampling days when the weather was overcast. Productivity throughout the year declined rapidly with depth. Algal biomass in Barton Broad varied from 3-10 mg chlorophyll α m⁻³ in winter but increased in spring and was very high in summer (200-500 mg chlorophyll α m⁻³). Secchi disc depth varied from 0.21 m in August 1976 to 1.76 m in December. Phytoplankton production in Barton Broad was low in winter (range: A_max 247-1,250 mg O₂ m⁻³ h⁻¹) but increased markedly in spring and summer with the highest rate (A_max 6,850 mg O₂ m⁻³ h⁻¹) being recorded in August. Surface inhibition was observed during summer except when the weather was overcast. Seasonality in nutrients and phytoplankton in Waigani Lake appear to be related to rainfall. Nutrient concentrations in Barton Broad are more closely related to phytoplankton activity which, in turn, correlates with seasonality in solar radiation.     

Spatial and temporal variation in nitrogen fixation and its importance to phytoplankton in phosphorus‐rich lakes

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Plant community structure determines primary productivity in shallow,eutrophic lakes

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Effects of glaciers on nutrient concentrations and phytoplankton in lakes within the Northern Cascades Mountains (USA)

We compared nitrate concentrations, phytoplankton biomass, and phytoplankton community structure in lakes fed by glacier melt and snowmelt (GSF lakes) and by snowmelt only (SF lakes) within North Cascades National Park (NOCA) in Washington State, USA. In the U.S. Rocky Mountains, glacier melting has greatly increased nitrate concentrations in GSF lakes (52–236 µg NO₃–N L^?1) relative to SF lakes (1–14 µg NO₃–N L^?1) and thereby stimulated phytoplankton changes in GSF lakes. Considering NOCA contains approximately one-third of the glaciers in the continental U.S., and many mountain lakes that receive glacier meltwater inputs, we hypothesized that NOCA GSF lakes would have greater nitrate concentrations, greater phytoplankton biomass, and greater abundance of nitrogen-sensitive diatom species than NOCA SF lakes. However, at NOCA nitrate concentrations were much lower and differences between lake types were small compared to the Rockies. At NOCA, nitrate concentrations averaged 13 and 5 µg NO₃–N L^?1 in GSF and SF lakes, respectively, and a nitrate difference was not detectable in several individual years. There also was no difference in phytoplankton biomass or abundance of nitrogen-sensitive diatoms between lake types at NOCA. In contrast to the Rockies, there also was not a significant positive relationship between watershed percent glacier area and lake nitrate at NOCA. Results demonstrate that biogeochemical responses to global change in Western U.S. mountain lake watersheds may vary regionally. Regional differences may be affected by differing nitrogen deposition, climate, geology, or microbial processes within glacier environments, and merit further investigation.     

Changes in phytoplankton,benthic invertebrate and fish assemblages of boreal lakes following invasion by Gonyostomum semen

1. During recent decades, Gonyostomum semen populations have spread in northern temperate regions forming dense blooms that may dominate the phytoplankton assemblage for extended periods. In this study, we investigate the effects of G. semen blooms in boreal brown water lakes with special emphasis on phytoplankton, fish and benthic invertebrate assemblages using data from 10 boreal lakes sampled annually over a 10‐year period. 2. Significant differences in phytoplankton and benthic invertebrate assemblages were found between lakes with high (3.01 mm³ L^?1; >80% phytoplankton biomass) and lakes with low G. semen biomass (0.03 mm³ L^?1; <5% phytoplankton biomass). In particular, high G. semen lakes had lower biomass of smaller, edible phytoplankton and a higher abundance and biomass of benthic invertebrates, especially Chaoborus flavicans, and perch than low G. semen lakes. 3. The length distribution of fish also suggested a tendency towards large and older fish and a lower recruitment success in high G. semen lakes, as denoted by lower abundances of fish shorter than 10 cm and higher biomass and abundance of fish longer than 15 cm in high G. semen lakes. 4. This study shows that high G. semen lakes are characterised by less edible phytoplankton, dominance by a few species and enhanced benthic secondary and fish production. Hence, the conjecture that high biomasses of G. semen create a bottleneck in the energy transfer to higher trophic levels seems less likely in boreal lakes.     

Phytoplankton and primary production in clear-vegetated,inorganic-turbid,and algal-turbid shallow lakes from the pampa plain (Argentina)

Shallow lakes often alternate between two possible states: one clear with submerged macrophytes, and another one turbid, dominated by phytoplankton. A third type of shallow lakes, the inorganic turbid, result from high contents of suspended inorganic material, and is characterized by low phytoplankton biomass and macrophytes absence. In our survey, the structure and photosynthetic properties (based on ¹⁴C method) of phytoplankton were related to environmental conditions in these three types of lakes in the Pampa Plain. The underwater light climate was characterized. Clear-vegetated lakes were more transparent (K _d 4.5–7.7 m⁻¹), had high DOC concentrations (>45 mg l⁻¹), low phytoplankton Chl a (1.6–2.7 μg l⁻¹) dominated by nanoflagellates. Phytoplankton productivity and photosynthetic efficiency (α ~ 0.03 mgC mgChla ⁻¹ h⁻¹ W⁻¹ m²) were relatively low. Inorganic-turbid lakes showed highest K _d values (59.8–61.4 m⁻¹), lowest phytoplankton densities (dominated by Bacillariophyta), and Chl a ranged from 14.6 to 18.3 μg l⁻¹. Phytoplankton-turbid lakes showed, in general, high K _d (4.9–58.5 m⁻¹) due to their high phytoplankton abundances. These lakes exhibited the highest Chl a values (14.2–125.7 μg l⁻¹), and the highest productivities and efficiencies (maximum 0.56 mgC mgChla ⁻¹ h⁻¹ W⁻¹ m²). Autotrophic picoplankton abundance, dominated by ficocianine-rich picocyanobacteria, differed among the shallow lakes independently of their type (0.086 × 10⁵–41.7 × 10⁵ cells ml⁻¹). This article provides a complete characterization of phytoplankton structure (all size fractions), and primary production of the three types of lakes from the Pampa Plain, one of the richest areas in shallow lakes from South America. Handling editor: J. Padisak     

Primary and new production in the deep Canada Basin during summer 2002

The NOAA Ocean Exploration program provided the opportunity to measure the carbon and nitrogen productivity across the Canada Basin. This research examined the major environmental factors limiting the levels of primary production and possible future climate change on the ecosystems. The vertical distributions of the carbon and nitrogen uptakes of phytoplankton had similar patterns as their respective biomass concentrations which were low at the surface and highest in the chlorophyll-maximum layer. The annual carbon and new production rates of phytoplankton in the Canada Basin were about 5 and 1 g C m^–2, respectively. Nutrients were determined to be a main limiting factor at the surface, whereas light may be a major factor limiting phytoplankton productivity in the chlorophyll-maximum layer for open waters. The bottom surface of the ice has a low specific uptake and productivity of phytoplankton, indicating that photosynthetic activity might be controlled by both light and nutrients.     

Effects on phytoplankton of nutrients added in conjunction with acidification

1. The effects of nitric acidification on phytoplankton were studied in a small, eperimentally manipulated, oligotrophic lake (L302N) in the Eperimental Lakes Area of Canada. The focus was altered after 9 years of acidification to investigate the possibility of using nutrient additions to stimulate recovery, followed by a controlled incremental recovery, in which the pH was increased to a predetermined target level. 2. Five years of additions of HNO₃ to L302N reduced its pH from 6.5 to 6.1. Nitrate concentration increased because the algal community was severely P deficient. The phytoplankton community structure and productivity were not significantly affected by these additions. 3. The phytoplankton community was significantly affected when pH was subsequently decreased over three successive years from 6.1 to 5.1 by the addition of HCl. Dominance shifted from chrysophytes to a co-dominance of chlorophytes and dinoflagellates, which altered the size structure of the community. Species diversity significantly decreased, although phytoplankton productivity remained unchanged. 4. At pH 5.1 nitrate and sulphate additions were made, creating conditions like those in lakes in eastern North America, which receive high loadings of nitrogen from the atmosphere. The phytoplankton assemblage shifted to dominance by small coccoidal chlorophytes. However, biomass and productivity were unaffected. 5. Finally, phosphate, as phosphoric acid, was added, along with nitrate and sulphate, to the epilimnion, which stimulated internal alkalinity generation and productivity. It is concluded that CO₂ concentrations and the form of N (nitrate vs. ammonia) affect algal composition but that P determines algal biomass and productivity. Chlorophytes were found to be good competitors for P when N and CO₂ were high; it is epected that cyanobacteria would be more competitive for P in low CO₂ systems. Conversely, dinoflagellates are most competitive in systems with low pH and high P, such as that which occurred in L302N. Although the P additions reduced N concentrations and created alkalinity, this is not a recommended remedial procedure in acidified lakes because it enhanced dinoflagellate abundance, which has been associated with fish kills. 6. When all additions ceased, the pH of L302N recovered from 5.1 to 5.8, chrysophytes and chlorophytes became more abundant and dinoflagellates decreased in abundance. Phytoplankton biomass decreased and species diversity increased. Phytoplankton productivity remained unchanged     

Limnology of naturally acidic,oligotrophic dune lakes in subtropical Australia,including chlorophyll — phosphorus relationships

This study provides the most comprehensive physico-chemical and phytoplankton data yet available for Australian dune lakes, which are among the world's most naturally acidic and oligotrophic freshwaters. Seasonal and spatial variations were examined in Blue Lagoon and Lake Freshwater, two ‘water-table window’ lakes in south-east Queensland. Like other dune lakes, they are acidic (minimum pH 4.20 and 4.55, respectively), polymictic water bodies with low concentrations of marine-derived major ions and almost undetectable levels of trace metals. While linmologically similar in winter, during spring-summer Lake Freshwater has significantly higher levels of chlorophyll-a, total phosphorus (TP) and turbidity than Blue Lagoon and other dune lakes, indicating seasonal mesotrophy. The key nutrient is TP, which has recently increased to a maximum of 17 μ· l⁻¹, due either to inputs from recreational sources, or to the death and decomposition of littoral vegetation resulting from falling water levels over the last decade. Inorganic nitrogen, though present only in small amounts, does not appear to limit the eutrophication process because of a shift in phytoplankton dominance from the usual desmids and dinoflagellates to N₂-fixing blue-green algae. A chlorophyll - TP linear regression derived for dune lakes indicates that at TP < 20 μg · l⁻¹ chlorophyll ‘yield’ is higher than in other lake types represented by regressions from the literature. This may be due to a more efficient utilization of the limited available phosphorus by dune lake algae which have adapted to the naturally oligotrophic environment. The implications of these findings for lake management are discussed.     

The zooplankton-phytoplankton interface in lakes of contrasting trophic status: an experimental comparison

We report here the results of an experimental study designed to compare algal responses to short-term manipulations of zooplankton in three California lakes which encompass a broad range of productivity (ultra-oligotrophic Lake Tahoe, mesotrophic Castle Lake, and strongly eutrophic Clear Lake). To assess the potential strength of grazing in each lake, we evaluated algal responses to a 16-fold range of zooplankton biomass. To better compare algal responses among lakes, we determined algal responses to grazing by a common grazer (Daphnia sp.) over a range ofDaphnia densities from 1 to 16 animals per liter. Effects of both ambient grazers andDaphnia were strong in Castle Lake. However, neither ambient zooplankton norDaphnia had much impact on phytoplankton in Clear Lake. In Lake Tahoe, no grazing impacts could be demonstrated for the ambient zooplankton butDaphnia grazing had dramatic effects. These results indicate weak coupling between phytoplankton and zooplankton in Clear Lake and Lake Tahoe, two lakes which lie near opposite extremes of lake trophic status for most lakes. These observations, along with work reported by other researchers, suggest that linkages between zooplankton and phytoplankton may be weak in lakes with either extremely low or high productivity. Biomanipulation approaches to recover hypereutrophic lakes which aim only to alter zooplankton size structure may be less effective if algal communities are dominated by large, inedible phytoplankton taxa.     

Impact of eutrophication on the carbon stable‐isotopic baseline of benthic invertebrates in two deep soft‐water lakes

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