Phosphorus and nitrogen relationships of Cladophora glomerata in two lake basins of different trophic status

• 1 Cladophora has increased in abundance and cover in the South Basin of Windermere, English Lake District, in recent years. During the growing seasons of 1992 and 1993, the maximum biomass of Cladophora in the South Basin, at nearly 200 g dry weight m^-2, was nearly six times greater and the percentage cover, at 95%, was nearly ten times greater than in the North Basin.
• 2 Nutrient analysis showed that Cladophora from the South Basin had significantly higher average contents of N than that from the North Basin. The suggested rate-limiting tissue N content, 1.3% of dry weight, was never reached in the South Basin but was reached in 20% of samples from the North Basin. In contrast, the average P content was not significantly different, at about 0.24% in both basins. The suggested rate-limiting P content, 0.16% of dry weight, was reached in 31% of the samples from the South Basin and 37% from the North Basin.
• 3 On average, the standing stock of P in the South Basin was 0.12 g m^-2, twelvefold that in the North Basin at 0.01 g m^-2 and the standing stock of N in the South Basin was 1.81 g m^-2, elevenfold that in the North Basin at 0.16 g m^-2. Maximum standing stocks of P were 0.38 g m^-2 in the South and 0.05 g m^-2 in the North Basin. Maximum standing stocks of N were 5.02 g m^-2 in the South and 0.60 g m^-2 in the North Basin.
• 4 The maximal rate of PO₄–P uptake ranged between 298 and 2949 μg P g^-1 DW h^-1 and the specific affinity varied between 2.9 and 24.3 μg P g^-1 DW h^-1 (mg PO₄–P m^-3)^-1. The higher values of both characteristics are within the range which is believed to indicate severe P limitation.
• 5 Using the nutrient uptake characteristics, rates of uptake at pelagic and littoral concentrations of PO₄–P were calculated to be between 0.1 and 1.7% of the maximal rates apart from one site where rates were up to 5.6% of V_maxi and 40% of V_maxi based on pelagic and littoral concentrations of PO₄–P, respectively. Despite these generally low rates of uptake, calculations suggest they were just sufficient to account for the measured rate of increase of P standing stock, assuming no biomass loss. Similar calculations suggested that pelagic concentrations of NO₃–N could support between 21 and 30% of maximal rates and these were between four and six times (North Basin) and twenty-five times (South Basin) the observed rates of increase in N standing stock. Littoral concentrations of NO₃–N were similar and so would have allowed similar rates.
• 6 Evidence from nutrient tissue contents, standing stocks and rates of uptake suggests that the growth of Cladophora, in both basins of Windermere, is controlled primarily by the availability of P.

     

Interannual variability of algal populations and their influence on lake metabolism

• 1 An input-output phosphorus budget is given for Windermere and its two basins based on data available for the late 1980s. The annual areal total phosphorus loading for the whole lake was 1.04 g P m^-2 yr^-1 and for the North and South Basins were 1.08 and 1.70 g P m^-2 yr^-1, respectively. For the whole lake and its South Basin the values were similar to the upper range of critical loads calculated according to the equation of Vollenweider (1976) for the transition between oligotrophy and eutrophy while that for the North Basin (1.08 g P m^-2 yr^-1) was within this range of critical loadings but towards its lower end.
• 2 Changes in the quality of summer phytoplankton are described for Windermere, particularly its South Basin, between 1978 and 1989 in relation to the utilization of nitrate-nitrogen (NO³-N) in the epilimnion, deoxygenarion of the hypolimnion and the ratio of epilimnetic volume to hypolimnetic volume, E_v/H_v The two basins of Windermere with values of E_v/H_v of 0.79 (South Basin) and 0.50 (North Basin) have contrasting conditions of summer deoxygenarion. The shallower South Basin shows marked interannual variability in the development of hypolimnetic anoxia. Years with large hypolimnetic anoxia during autumn are correlated with the production during summer of large populations of the poorly grazed blue-green alga Oscillatoria bourrellyi and exhaustion of NO₃-N in the upper layers. During years when anoxia does not develop the summer phytoplankton consists of small easily grazed algae or larger ones subject to parasitic epidemics. The deeper North Basin never becomes anoxic even though it can contain similar sized populations of O. bourrellyi to the South Basin.
• 3 A possible explanation of the between basin and, for the South Basin, between year variation of utilization of NO₃-N and level of hypolimnetic deoxygenarion is that algal quality can determine lake metabolism dependent upon lake or basin morphology. Poorly grazed large forms such as O. bourrellyi act as sinks for NO₃-N. On sedimentation such populations act as a ‘short circuit’ mechanism descending into deeper layers in sufficient quantities to cause anoxia. Other species subject to crustacean or microbial grazing are mineralized in the epilimnion with little sedimentation to the deeper waters. Subsequent recycling of nitrogen as NH₄-N takes place in the upper layers or thermocline which is more readily taken up by subsequent production. The influence of such ‘short circuit’ mechanisms is reduced in deep lakes and exacerbated in shallow ones.
• 4 The success of species such as O. bourrellyi is dependent upon a sufficient inoculum, an adequate supply of nutrients and the depth of intermittent mixing. The importance of these factors in regulating presence and timing of summer populations is illustrated and discussed.

     

ALKALINE PHOSPHATASE IN FRESHWATER CLADOPHORA‐EPIPHYTE ASSEMBLAGES: REGULATION IN RESPONSE TO PHOSPHORUS SUPPLY AND LOCALIZATION1

Extracellular alkaline phosphatase enzyme activity (APA) is important for algal phosphorus (P) acquisition in P‐limited freshwater ecosystems and is often used as an indicator of P deficiency. APA allows access to organic P (monophosphate esters), but the regulation of APA in response to availability of both PO₄³⁻ and organic P is poorly characterized. This study aimed to examine the regulation of APA in freshwater Cladophora‐epiphyte assemblages in response to PO₄³⁻ and a hydrolyzable organic P source, and for the first time to apply enzyme linked fluorescence (ELF) to localize APA within freshwater macroalgal‐epiphyte assemblages. In response to elevated PO₄³⁻ concentrations, a component of net APA was suppressed, but there was also a constitutive APA, which was maintained even after prolonged exposure to nearly 1,000 μM PO₄³⁻ and saturation of internal P pools. When supplied with organic glycerol P as the sole P source, the algae maintained APA in excess of needs for supplying PO₄³⁻ for uptake, resulting in PO₄³⁻ release into the medium. Constitutive APA may be adaptive to growth under chronic P limitation in oligotrophic freshwater habitats. Excess APA and release of PO₄³⁻ could benefit different algal and bacterial partners within assemblages. APA in both Cladophora sp. and epiphytic algae was localized with ELF only when ethanol fixation was omitted. In algal subsamples exposed to different P treatments, there was no correlation between bulk APA (using 4‐methylumbelliferyl phosphate [MUP] substrate) and % cell labeling with ELF, suggesting that ELF labeling of APA was at best semiquantitative in the algal assemblages.     

Soil fertility and fine root dynamics in response to 4 years of nutrient (N,P, K) fertilization in a lowland tropical moist forest,Panama

The question of how tropical trees cope with infertile soils has been challenging to address, in part, because fine root dynamics must be studied in situ. We used annual fertilization with nitrogen (N as urea, 12.5 g N m^?2 year^?1), phosphorus (P as superphosphate, 5 g P m^?2 year^?1) and potassium (K as KCl, 5 g K m^?2 year^?1) within 38 ha of old‐growth lowland tropical moist forest in Panama and examined fine root dynamics with minirhizotron images. We expected that added P, above all, would (i) decrease fine root biomass but, (ii) have no impact on fine root turnover. Soil in the study area was moderately acidic (pH = 5.28), had moderate concentrations of exchangeable base cations (13.4 cmol kg^?1), low concentrations of Bray‐extractable phosphate (PO₄ = 2.2 mg kg^?1), and modest concentrations of KCl‐extractable nitrate (NO₃ = 5.0 mg kg^?1) and KCl‐extractable ammonium (NH₄ = 15.5 mg kg^?1). Added N increased concentrations of KCl‐extractable NO₃ and acidified the soil by one pH unit. Added P increased concentrations of Bray‐extractable PO₄ and P in the labile fraction. Concentrations of exchangeable K were elevated in K addition plots but reduced by N additions. Fine root dynamics responded to added K rather than added P. After 2 years, added K decreased fine root biomass from 330 to 275 g m^?2. The turnover coefficient of fine roots <1 mm diameter ranged from 2.6 to 4.4 per year, and the largest values occurred in plots with added K. This study supported the view that biomass and dynamics of fine roots respond to soil nutrient availability in species‐rich, lowland tropical moist forest. However, K rather than P elicited root responses. Fine roots smaller than 1 mm have a short lifetime (<140 days), and control of fine root production by nutrient availability in tropical forests deserves more study.     

Phytoplankton community structure and environment in the Kenyan waters of Lake Victoria

• 1 Phytoplankton species composition, numerical abundance, spatial distribution and total biomass measured as chlorophyll a concentration were studied in relation to environmental factors in September 1994 (dry season) and March 1995 (rainy season), respectively, in the Kenyan waters of Lake Victoria; 103 species were recorded.
• 2 Blue‐green algae (Cyanophyceae) were most diverse, followed by diatoms (Bacillariophyceae), green algae (Chlorophyceae) and dinoflagellates (Dinophyceae).
• 3 Twinspan separated the phytoplankton communities in the Nyanza Gulf and those in the open lake during both seasons. During the dry season, the Nyanza Gulf was strongly dominated by blue‐greens, while diatoms dominated in the open lake. During the rainy season, blue‐greens remained dominant in the Nyanza Gulf although the number of species found was lower than during the dry season; in the open lake, blue‐greens replaced diatoms as the dominant group and there were more species than in the dry season.
• 4 Canonical correspondence analysis indicated that the phytoplankton species distribution was significantly correlated with turbidity during the dry season and with SiO₂ during the rainy season. Chlorophyll a concentrations ranging from 2.0 to 71.5 mg m^‐3 in the dry season and 2.0–17.2 mg m^‐3 in the rainy season confirm earlier reports of increasing phytoplankton biomass in Lake Victoria since the 1960s.

     

Filamentous algae in fish ponds of the Třeboň Biosphere Reserve-ecophysiological study

Filamentous algae in eutrophic carp ponds in South Bohemia (Central Europe) were studied from 1988 to 1990. High biomass (490 g DW m^-2) was attained by Cladophora fracta (O. F. Müll. ex Vahl) Kütz. after two months of growth. This marked growth depleted inorganic carbon in the water, but did not decrease the concentration of tissue nutrients. Laboratory measurements of final pH indicate that all the filamentous algae studied, except for Tribonema, are very efficient HCO₃ ^- users. An extremely high pH of 11.6 and oxygen concentration of 32 mg l^-1 were measured in the algal mats. High pH resulted in CaCO₃ precipitation, visible as white incrustations on algal filaments. The amount of precipitated CaCO₃ reached 134 kg ha^-1. After reaching peak biomass, 90% of the Cladophora decomposed over the next 95 days.The highest net photosynthetic rate in C. fracta was measured between pH range 8.5–10.0 and oxygen concentrations of 7–12 mg l^-1. Optimum temperature for photosynthesis was between 17–22°C.     

In situ measurements confirm the seasonal dominance of benthic algae over phytoplankton in nearshore primary production of a large lake

1. Despite the recognition of its importance, benthic primary production is seldom reported, especially for large lakes. We measured in situ benthic net primary production by monitoring flux in dissolved inorganic carbon (DIC) concentration in benthic incubation chambers, based on continuous measurements of CO_2(aq) flux, alkalinity, and the temperature‐dependent dissociation constants of carbonic acid (K₁ and K₂). This methodology has the advantages of monitoring net primary production directly as change in carbon, maintaining continuous water recirculation, and having sufficient precision to detect change in DIC over short (i.e. 15 min) incubations, even in alkaline waters. 2. Benthic primary production on Cladophora‐dominated rocky substrata in western Lake Ontario was measured biweekly. Maximum biomass‐specific net photosynthetic rates were highest in the spring (2.39 mgC g Dry Mass^?1 h^?1), decreased to negative rates by early summer (?0.76 mgC g DM^?1 h^?1), and exhibited a regrowth in late summer (1.98 mgC g DM^?1 h^?1). 3. A Cladophora growth model (CGM), previously validated to predict Cladophora biomass accrual in Lake Ontario, successfully simulated the seasonality and magnitude of biomass‐specific primary production during the first cohort of Cladophora growth. Averaged over this growing season (May–Aug), mean areal net benthic production at the estimated depth of peak biomass (2 m) was 405 mg C m^?2 d^?1. 4. We measured planktonic primary production in proximity to the benthic study and constructed a depth‐resolved model of planktonic production. Using the CGM, benthic primary production was compared with planktonic primary production for the period May–Aug. Net benthic production from the shoreline to the 12 m contour (1–2 km offshore) equalled planktonic production. Closer to shore, benthic primary production exceeded planktonic primary production. Failure to account for benthic primary production, at least during abundant Cladophora growth, will lead to large underestimates in carbon and nutrient flows in the nearshore zone of this Great Lake.     

Biofiltering efficiency, uptake and assimilation rates of Ulva clathrata (Roth) J. Agardh (Clorophyceae) cultivated in shrimp aquaculture waste water

The growth, biofiltering efficiency and uptake rates of Ulva clathrata were studied in a series of outdoor tanks, receiving waste water directly from a shrimp (Litopenaeus vannamei) aquaculture pond, under constant aeration and two different water regimes: (1) continuous flow, with 1 volume exchange a day (VE day^-1) and (2) static regime, with 1 VE after 4 days. Water temperature, salinity, pH, dissolved inorganic nitrogen (DIN), phosphate (PO₄), chlorophyll-a (chl-a), total suspended solids (TSS), macroalgal biomass (fresh weight) and tissue nutrient assimilation were monitored over 12 days. Ulva clathrata was highly efficient in removing the main inorganic nutrients from effluent water, stripping 70–82% of the total ammonium nitrogen (TAN) and 50% PO₄ within 15 h. Reductions in control tanks were much lower (Tukey HSD, P < 0.05). After 3 days, the mean uptake rates by the seaweed biomass under continuous flow were 3.09 mg DIN g DW day⁻¹ (383 mg DIN m⁻² day⁻¹) and 0.13 mg PO₄ g DW day⁻¹ (99 mg PO₄ m⁻² day⁻¹), being significantly higher than in the static regime (Tukey HSD, P < 0.05). The chl-a decreased in seaweed tanks, suggesting that U. clathrata inhibited phytoplankton growth. Correlations between the cumulative values of DIN removed from the water and total nitrogen assimilated into the seaweed biomass (r = 0.7 and 0.8, P < 0.05), suggest that nutrient removal by U. clathrata dominated over other processes such as phytoplankton and bacterial assimilation, ammonia volatilization and nutrient precipitation.     

Improvement of both lipid and biomass productivities of Qatar Chlorocystis isolate for biodiesel production and food security

Microalgae are considered a very promising alternative for biofuel production. Several strategies were developed to modulate and improve algae metabolites production to meet the requirements for biodiesel production. Most previous research evidenced that the increase of the lipid content is accompanied by a decrease of the biomass production, which increases the cost of the downstream processing. Hence, the challenge is to find special culture conditions that increase the lipid and the biomass productivities simultaneously. In the present work, we developed a strategy for the improvement of biomass and lipid productivities in a novel local microalga isolate, Chlorocystis sp. QUCCCM14, which was not previously known as a promising strain. Indeed, culturing QUCCCM14 using f/2 medium with 10× NaH₂PO₄ (0.15 g L^?1 NaNO₃ and 5.6 mg L^?1 NaH₂PO₄) resulted in an improvement of 3.178 folds the lipid productivity reaching 56.121 mg L^?1 day^?1 and enhanced the biomass productivity reaching 141.363 mg L^?1 day^?1, simultaneously. Comparative analyses of the FAME profiles demonstrated that fed‐batch culture with phosphate or nitrate separately leads to a high production of the omega 3 fatty acids (Linolenic acid), whereas fed‐batch culture with phosphate and nitrate simultaneously increased the production of fatty acids suitable for biodiesel production.     

Contrasting effects of low‐level phosphorus and nitrogen enrichment on growth of the mat‐forming alga Didymosphenia geminata in an oligotrophic river

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Nitrogen and phosphorus dynamics in a large river estimated by an in situ Lagrangian tracking approach

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Chlorophyll‐a concentrations and macroinvertebrate declines coincide with the collapse of overwintering diving duck populations in a large eutrophic lake

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Regulation of phosphate uptake reveals cyanobacterial bloom resilience to shifting N:P ratios

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Water quality of the Modder River,South Africa

The Modder River is a relatively small river in the central region of the Free State Province, South Africa and has a mean annual runoff of 184 × 106m³. Botshabelo is a city, which has developed in the catchment area of the river, and its sewage outflows are discharged into the Klein Modder, a tributary of the Modder River. This study was conducted in order to determine the influence of Botshabelo's sewage outflow on the water quality of the river. It was determined that the Modder and Klein Modder Rivers do not generally follow distinctive seasonal patterns in terms of chemical parameters, although NO₃-N and PO₄-P concentrations usually increase with increasing flow and conductivity decreases with increasing flow. Physical parameters such as turbidity, flow and temperature did however follow distinctive seasonal patterns from February 1996 to December 1997, as did phytoplankton growth. Low chlorophyll-a concentrations were exhibited in the winter and higher concentrations during spring. In the Klein Modder River, algal blooms occurred more frequently, and the algal biomass was higher than in the Modder River. This could be ascribed to the higher nutrient concentrations and lower flow velocities in the former. The inflow of the Klein Modder River into the Modder River caused on average, 112% increase in PO₄-P, 171% increase in NO₃-N, 50% increase in chlorophyll-a concentration, and 230% increase in E. coli counts.     

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.     

Clipping defoliation and nitrogen addition shift competition between a C3 grass (Leymus chinensis) and a C4 grass (Hemarthria altissima)

• Human‐induced disturbances, including grazing and clipping, that cause defoliation are common in natural grasslands. Plant functional type differences in the ability to compensate for this tissue loss may influence interspecific competition.
• To explore the effects of different intensities of clipping and nitrogen (N) addition on compensatory growth and interspecific competition, we measured accumulated aboveground biomass (AGB), belowground biomass (BGB), tiller number, non‐structural carbohydrates concentrations and leaf gas exchange parameters in two locally co‐occurring species (the C₃ grass Leymus chinensis and the C₄ grass Hemarthria altissima) growing in monoculture and in mixture.
• For both grasses, the clipping treatment had significant impacts on the accumulated AGB, and the 40% clipping treatment had the largest effect. BGB gradually decreased with increasing defoliation intensity. Severe defoliation caused a significant increase in tiller number. Stored carbohydrates in the belowground biomass were mobilised and transported aboveground for the growth of new leaves to compensate for clipping‐induced injury. The net CO₂ assimilation rate (A) of the remaining leaves increased with clipping intensity and peaked under clipping intensities of 20% or 40%. Nitrogen addition, at a rate of 10 g·N·m^?2·year^?1, enhanced A of the remaining leaves and non‐structural carbohydrate concentrations, which benefited plant compensatory growth, especially for the C₃ grass. Under the mixed planting conditions, the clipping and N addition treatments lowered the competitive advantage of the C₄ grass.
• The results suggest that a combination of defoliation and N deposition have the potential to benefit the coexistence of C₃ and C₄ grasses.

     

Preliminary observations on Gibbula umbilicalis (da Costa, 1778) on the Portuguese coast

Physiological indicators of nitrogen and phosphorus deficiency were used to determine the role of these nutrients in the development and proliferation of Cladophora in the Manawatu River, New Zealand. Nitrogen indicator tests showed that surplus nitrogen was available at all times. During a sustained low flow period when the dissolved reactive phosphorus was between 4–5 mg m^–3, phosphorus indicator tests revealed that the Cladophora proliferations were phosphorus deficient. A downstream trend of reduced nitrogen and phosphorus availability was also observed during this period. Sudden high river flows were frequent and had a major influence on the Cladophora biomass.     

GROWTH AND PHOSPHORUS UPTAKE BY THE TOXIC DINOFLAGELLATE ALEXANDRIUM CATENELLA (DINOPHYCEAE) IN RESPONSE TO PHOSPHATE LIMITATION1

Alexandrium catenella (Whedon et Kof.) Balech has exhibited seasonal recurrent blooms in the Thau lagoon (South of France) since first reported in 1995. Its appearance followed a strong decrease (90%) in phosphate (PO₄^3?) concentrations in this environment over the 1970–1995 period. To determine if this dinoflagellate species has a competitive advantage in PO₄^3?‐limited conditions in terms of nutrient acquisition, semicontinuous cultures were carried out to characterize phosphorus (P) uptake by A. catenella cells along a P‐limitation gradient using different dilution rates (DRs). Use of both inorganic and organic P was investigated from measurements of ³³PO₄^3? uptake and alkaline phosphatase activity (APA), respectively. P status was estimated from cellular P and carbon contents (Q_P and Q_C). Shifts in trends of Q_P/Q_C and Q_P per cell (Q_P·cell?1) along the DR gradient allowed the definition of successive P‐stress thresholds for A. catenella cells. The maximal uptake rate of ³³PO₄^3? increased strongly with the decrease in DR and the decrease in Q_P/Q_C, displaying physiological acclimations to PO₄^3? limitation. Concerning maximal APA per cell, the observation of an all‐or‐nothing pattern along the dilution gradient suggests that synthesis of AP was induced and maximized at the cellular scale as soon as PO₄^3? limitation set in. APA variations revealed that the synthesis of AP was repressed over a PO₄^3? threshold between 0.4 and 1 μM. As lower PO₄^3? concentrations are regularly observed during A. catenella blooms in Thau lagoon, a significant portion of P uptake by A. catenella cells in the field may come from organic compounds.     

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.

     

Spatial heterogeneity and short‐term oxygen dynamics in the rhizosphere of Vallisneria spiralis: Implications for nutrient cycling

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