Stoichiometric Constraints on Food-Web Dynamics: A Whole-Lake Experiment on the Canadian Shield

A whole-lake manipulation of food-web structure (introduction of a top predator, northern pike, to a minnow-dominated lake) was performed in a Canadian Shield lake (L110) to examine the stoichiometric consequences of changes in planktonic community structure generated by altered food-web structure. Minnow abundance, zooplankton biomass and community composition, microconsumer abundance, and concentration and carbon–phosphorus (C:P) ratio of suspended particulate matter were monitored in L110 and unmanipulated L240 before (1992) and after (1993–95) pike introduction. Algal biomass in L110 determined from microscopic examination for postmanipulation and premanipulation periods was also compared with dynamics in a suite of unmanipulated reference lakes from long-term monitoring records. Pike were added in spring in 1993 and 1994 in sufficient quantity to raise pike biomass to levels of around 22 kg ha^{− 1} by 1994. Minnow populations in L110 responded dramatically, decreasing to levels 30% (1993), 10% (1994), and less than 1% (1995) of premanipulation values. However, most components lower in the food web did not respond in a manner consistent with predictions of existing food-web theory, such as the idea of cascading trophic interactions (CTI). While Daphnia biomass increased in L110 in the first year following manipulation, consistent with CTI, this effect was temporary and Daphnia collapsed in 1995, the year of lowest minnow abundance. Total zooplankton biomass in both lakes declined during the study period and, contrary to CTI, this decline appeared somewhat stronger in L110 than in L240. Dominant microconsumers (heterotrophic microflagellates) did not differ among years in either lake and did not appear to respond to food-web manipulation. At the bottom of the food web, no changes in bacterial biomass occurred in either lake. However, total concentrations of particulate matter appeared to increase in L110 after manipulation (contrary to expectations based on the theory of CTI) while algal biomass did not change in the manipulated lake relative to reference systems. Finally, particulate C:P increased in both L110 and L240 during the study period. The lack of strong response of Daphnia, the lack of response of the microbial food web, decreases in zooplankton biomass and increases in particulate biomass following reduction of minnow populations after piscivore introduction are at odds with expectations from existing food-web theory, such as the idea of CTI as currently formulated. However, the extremely high C:P ratios in particulate matter at the base of the food webs in these lakes, the coincidence of zooplankton declines and increases in particulate C:P ratios, and the results of small-scale mesocosm food-quality experiments are consistent with a hypothesis of a stoichiometric constraint operating on food-web dynamics in this and similar ecosystems. Received 22 April 1997; accepted 8 July 1997.     

Zooplankton contribution to particulate phosphorus and nitrogen in lakes

Based on the recognition of rather constant species-specificelement to dry weight ratios in freshwater zooplankton, poolsof metazoan zooplankton P and N were calculated for 45 Norwegianlakes of varying trophy. On the average zooplankton constituted20.4±12.3% and 4.6±4.3% of particulate P and Nrespectively. The fraction was considerably higher in many oligotrophiclakes, at most >50 and 20% respectively, but declined to15% (P) and 5% (N) in meso- to eutrophic lakes. In general,phytoplankton contribution to particulate P was <50%, leavinga large share of particulate P to bacteria and microzooplankton.The zooplankton proportion of particulate P was only weaklyinfluenced by the predation pressure in terms of fish communitystructure. Zooplankton P is important in the overall lake metabolism,and knowledge of this pool may be used to estimate zooplankton-mediatedloss and regeneration of P and improve total nutrient elementbudgets in lakes.     

Elemental stoichiometry of lower food web components in arctic and temperate lakes

Lakes were surveyed to assess the potential patterns of latitudinalvariation in carbon:nitrogen:phosphorus (C:N:P) stoichiometryof lower food web components. Thirty-four lakes were surveyedat an arctic latitude (68°38'N, 149°38'W) and 10 lakesat a temperate latitude (46°13'N, 89°32'W) during 1997.The temperate data set was augmented with earlier survey resultsemploying similar methods. It was hypothesized that differencesin environmental variables across latitude would cause differencesin community C:N:P ratios, leading to differences in trophicinteractions. Physical measurements (light, temperature), sestonand zooplankton were collected from each lake. Seston and zooplanktonwere analyzed for C, N and P content, and zooplankton were countedand measured for biomass estimates. The degree of trophic interactionbetween seston and zooplankton was assessed by (i) measuringelemental imbalances between seston and zooplankton and (ii)calculating the potential recycling ratio by the zooplanktoncommunity available for seston. Seston C:nutrient, but not N:P,ratios were higher in temperate than arctic lakes. Conversely,arctic zooplankton had higher C:nutrient, but not N:P, ratiosthan zooplankton in temperate lakes. Elemental imbalances weregreater in temperate than in arctic lakes, but N:P stoichiometryof potential zooplankton recycling was nearly identical betweenthe two latitudes. Zooplankton community C:N:P ratios were notrelated to either latitude or seston C:N:P. In accordance withstoichiometric theory, relative abundances of calanoid copepodswere positively correlated with seston C:N in temperate lakes.Additionally, relative abundances of Daphnia were negativelycorrelated with seston C:N ratios in temperate and arctic lakes,and positively correlated with N:P ratios in the arctic. Ingeneral, these results suggest that seston and zooplankton communitystoichiometry differ across latitude, and these differenceshave the potential to affect trophic interactions.     

Food quality for Daphnia in humic and clear water lakes

1. Growth and reproduction of Daphnia fed lake seston were measured in two categories of meso‐ to eutrophic lakes differing with respect to terrestrial organic matter influence (humic and clear water lakes). The content of highly unsaturated fatty acids (HUFA), P and N, as well as the taxonomical composition of seston were analysed. 2. Seston HUFA and C : P ratios were similar between lake categories, whereas C : N ratios were lower in the clear water lakes in both spring and summer. Despite the similarity in HUFA and P content of seston, Daphnia growth rate, clutch size and the proportion of gravid females were, respectively, about 1.5, 3 and 6 times higher in the clear water lakes. 3. Differences in growth and reproduction were related to a combination of higher N content and good fatty acid quality of the seston in the clear water lakes. Relatively high biomass of edible algae, such as Rhodomonas sp. and Cryptomonas sp., in the clear water lakes, and differences in water pH likely contributed to the observed differences in Daphnia growth and reproduction between lake categories. Additionally, it is possible that Daphnia was energy limited in the humic lakes despite high particulate organic carbon (POC) concentrations, as the contribution of non‐algal and detrital C to the POC pool was high. 4. Our results suggest that dietary HUFA content has the potential to improve herbivore growth and reproduction if N and P are not limiting. N merits more attention in studies of zooplankton nutrition.     

Nutrient stoichiometry of linked catchment‐lake systems along a gradient of land use

1. Catchments export nutrients to aquatic ecosystems at rates and ratios that are strongly influenced by land use practices, and within aquatic ecosystems nutrients can be processed, retained, lost to the atmosphere, or exported downstream. The stoichiometry of carbon and nutrients can influence ecosystem services such as water quality, nutrient limitation, biodiversity, eutrophication and the sequestration of nutrients and carbon in sediments. However, we know little about how nutrient stoichiometry varies along the pathway from terrestrial landscapes through aquatic systems. 2. We studied the stoichiometry of nitrogen and phosphorus exported by three catchments of contrasting land use (forest versus agriculture) and in the water column and sediments of downstream reservoirs. We also related stoichiometry to phytoplankton nutrient limitation and the abundance of heterocystous cyanobacteria. 3. The total N : P of stream exports varied greatly among catchments and was 18, 54 and 140 (molar) in the forested, mixed‐use and agricultural catchment, respectively. Total N : P in the mixed layers of the lakes was less variable but ordered similarly: 35, 52 132 in the forested, mixed‐use and agricultural lake, respectively. In contrast, there was little variation among systems in the C : N and C : P ratios of catchment exports or in reservoir seston. 4. Phytoplankton in the forested lake were consistently N limited, those in the agricultural lake were consistently P limited, and those in the mixed‐use lake shifted seasonally from P‐ to N limitation, reflecting N : P supply ratios. Total phytoplankton and cyanobacteria biomass were highest in the agricultural lake, but heterocystous (potentially N fixing) cyanobacteria were most abundant in the forested lake, corresponding to low N : P ratios. 5. Despite large differences in catchment export and water column N : P ratios, the N : P of sediment burial (integrated over several decades) was very low and remarkably similar (4.3–7.3) across reservoirs. N and P budgets constructed for the agricultural reservoir suggested that denitrification could be a major loss of N, and may help explain the relatively low N : P of buried sediment. 6. Our results show congruence between the catchment export N : P, reservoir N : P, phytoplankton N versus P limitation and the dominance of heterocystous cyanobacteria. However, the N : P stoichiometry of sediments retained in the lakes was relatively insensitive to catchment stoichiometry, suggesting that a common set of biogeochemical processes constrains sediment N : P across lakes of contrasting catchment land use.     

Detritivory and the stoichiometry of nutrient cycling by a dominant fish species in lakes of varying productivity

Little is known about the stoichiometry of nutrient cycling by detritivores. Therefore, we explored stoichiometric relationships in an omnivorous/detritivorous fish (gizzard shad, Dorosoma cepedianum) in three lakes that differed in productivity. Gizzard shad can feed on plankton and sediment detritus, but in all three lakes adult gizzard shad derived >98% of carbon (C) and phosphorus (P), and >90% of nitrogen (N) from sediment detritus, and the remainder from zooplankton.
Gizzard shad selectively consumed detritus with higher C, N and P concentrations than ambient lake sediments. Selective detritivory (i.e. the nutrient content of consumed detritus divided by the nutrient content of ambient detritus) was most pronounced in the lake with the lowest detrital nutrient concentrations. N and P cycling rates per fish were also consistently higher in this lake, in agreement with the prediction of stoichiometry theory that excretion rates should increase with food nutrient content. Among-lake differences in nutrient cycling rates were unrelated to inter-lake variation in fish body nutrient contents, which was minimal. The N:P ratio excreted was near Redfield (∼14:1) in all three lakes.
Stoichiometric analyses showed that the C:N and C:P ratios of sediment detritus were much higher (∼2.8×) than ratios of gizzard shad bodies, revealing substantial N and P imbalances between consumers and their food source. Gizzard shad alleviate N imbalance by selectively feeding on high N detritus (low C:N, high N:P), and apparently alleviate P imbalance by excreting nutrients at a higher N:P than that of their food or their bodies. Thus, this detritivore apparently regulates nutrient acquisition and allocation via both pre-absorption processes (selective feeding) and post-absorptive processes (differential N and P excretion).     

Essential fatty acids and phosphorus in seston from lakes with contrasting terrestrial dissolved organic carbon content

1. It is often assumed that lakes highly influenced by terrestrial organic matter (TOM) have low zooplankton food quality because of elemental and/or biochemical deficiencies of the major particulate organic carbon pools. We used the biochemical [polyunsaturated fatty acids, especially eicosapentaenoic acid (EPA) – 20:5ω3] and elemental (C : P ratio) composition of particulate matter (PM) as qualitative measures of potential zooplankton food in two categories of lakes of similar primary productivity, but with contrasting TOM influence (clear water versus humic lakes). 2. C : P ratios (atomic ratio) in PM were similar between lake categories and were above 400. The concentration (μg L⁻¹) and relative content (μg mg C⁻¹) of EPA, as well as the particulate organic carbon concentration, were higher in the humic lakes than in the clear‐water lakes. 3. Our results show high fatty acid quality of PM in the humic lakes. The differences in the biochemical quality of the potential zooplankton food between lake categories can be attributed to the differences in their phytoplankton communities. 4. High biochemical quality of the food can result in high efficiency of energy transfer in the food chain and stimulate production at higher trophic levels, assuming that zooplankton are able to ingest and digest the resource available.     

Determinants of seston C : P-ratio in lakes

1. The ratio of carbon to phosphorus (C : P) in seston is a major determinant of energy transfer in aquatic food webs and may vary more than an order of magnitude owing to various extrinsic and intrinsic factors. In this study, the determinants of C : P‐ratios in lake particulate matter (seston) was assessed in 112 Norwegian lakes, covering a C : P (atomic ratio) from 24 to 1842 (mean 250). 2. No overall effects of lake area, season or latitude on C : P was detected. Particulate P, but not particulate C, correlated with C : P. Multivariate analysis including a range of lake properties revealed total dissolved P, as the major determinant of sestonic C : P, with the fraction of detritus in total seston, chlorophyll or Secchi depth and lake colour as significant contributors. Together these parameters explained 30% of observed variance if using dissolved P and 81% if using total P as input variable to the multivariate model. 3. Chlorophyll and Secchi depth were highly correlated and substitutable in the analysis. Phytoplankton community composition did not affect seston C : P, probably reflecting the fact that live phytoplankton generally contributed <25% of the seston pool. 4. Total P correlated positively with C : P and is the key determinant of phytoplankton biomass and thus Secchi depth; the latter parameters contributed negatively to seston C : P, probably owing to increased light attenuation. These lake data thus support the light : nutrient ratio hypothesis, i.e. that high light and low P cause skewed uptake ratios of C to P. 5. Zooplankton biomass in general and Daphnia biomass in particular, was negatively correlated with C : P, probably reflecting a negative impact of poor seston quality at high C : P. Zooplankton grazing and nutrient recycling may also have contributed to a negative correlation between zooplankton biomass and sestonic C : P.     

How light and nutrients affect the relationship between autotrophic and heterotrophic biomass in a tropical black water periphyton community

This study examines how nutrients and light affect the relationship between autotrophic biomass and non-autotrophic periphyton organic matter in a tropical black water lake biofilm community. We hypothesized that there is no positive correlation between autotrophic and non-autotrophic organic matter in the periphytic community of a black water humic lake, where non-algal components of periphyton can rely on carbon sources external to the periphyton matrix and where nutrient availability is low. Second, we sought to test our hypothesis that non-autotrophic periphyton organic matter will benefit from nutrient enhancement in a lake where the availability of DOC is high. We performed a field experiment using in situ lake mesocosms to manipulate nutrient concentrations and light availability in a 2 × 2 factorial design. Control treatments (no nutrient added) and nutrient treatments (N + P) were compared in different light conditions: high light (near surface water) and low light (near bottom). No positive correlation was found between autotrophic biomass and non-autotrophic periphyton organic matter, but a negative correlation was observed in high nutrient and light conditions. The low C:P and N:P ratios revealed that the non-autotrophic organic matter mostly comprised a heterotrophic microbial biofilm. High levels of light and nutrients together caused significant changes in periphyton community properties. The non-autotrophic periphyton organic matter was negatively affected by nutrient addition, whereas autotrophic biomass was positively affected, especially in high light conditions. Our results strongly suggest that non-autotrophic periphyton organic matter in a humic lake is primarily comprised of a bacterial biofilm that directly competes for nutrients with autotrophs in the periphytic community. We also observed no effect of nutrient addition on periphyton growing in light-limited conditions. These results suggest that heterotrophic periphytic organisms might experience carbon limitation despite the high availability, but usually low quality, of dissolved carbon in the water column of humic lakes.     

Phosphorus in settling matter and bottom sediments in lakes loaded by agriculture

In shallow lakes, the cycling of P between water and bottom sediments is strongly influenced by wind-induced resuspension of particulate matter. The significance of this P flux as an algal nutrient source is unclear. We examined gross sedimentation in 3 open and shallow agriculturally loaded lakes. In addition, we estimated the potential P-release from settling and bottom matter by laboratory tests. The mean daily rate of gross sedimentation was 21–170 g m^–2 d^–1 of dry sediment, 0.04–0.18 g m^–2 d^–1 of P and 0.18–2.0 g m^–2 d^–1 of N; being the highest in the shallowest and most eutrophic lake. In Lake Karhijärvi, where the most intensive measurements were taken, wind explained the temporal variation in the gross sedimentation to some extent. The settling matter consisted of inorganic particles low in nutrients, especially during peak sedimentation periods. On average, 7.7 ± 3.1% (x ± 95% confidence interval) of the P in the settling matter in L. Karhijärvi was in an algal-available form according to 2–3 week bioassays. In the bottom matter of the three lakes, 3.0 ± 1.7% and 2.5 ± 3.6%, and 4.3 ± 3.7% of the P was utilized by the algae. In L. Karhijärvi, resuspension of the potentially available P exceeded 20 times the external loading during the open water season. According to sorption tests, P is released from the bottom matter only when the concentration of o-P is <2 g l^–1. Although such a low value cannot be determined with common analytical procedures, it seems probable that the P concentration allows P desorption during P-limited periods. However, the significance of resuspended matter as an algal nutrient source calls for further research.     

Chemical composition of modern and pre-acidification sediments in the Tatra Mountain lakes

Concentrations of major nutrients (C, N, P) and acid soluble metals (Ca, Mg, K, Al, Fe, Mn, Pb, and Zn) were determined in modern (0–1 cm) and pre-acidification (5–10 cm) sediment layers collected from 37 alpine and 3 forest lakes in the Tatra Mountains (Slovakia, Poland) in 1996–1998. Sediment composition reflected catchment characteristics and productivity of lakes. In the sediments of alpine lakes, C and N concentrations decreased and Mg increased with a decreasing proportion of vegetation and soil in the catchment. Decreasing Ca:Mg ratios in sediments along the vegetation gradient was inverse to that in water, and could be associated with different ratios of cations in water leachate from catchments and in solids which enter the lake due to soil erosion. Phosphorus concentrations increased with the proportion of moraine areas, with till soils rich in P. Concentrations of C, N, P, and Ca in sediments positively correlated to their concentrations in water. Sediment concentrations of Al and Al:Ca ratios increased with decreasing sediment and water pH. A negative correlation between water pH and concentrations of organic C in water and sediments indicated the important impact of organic acids on the acid status of the lakes exposed to higher terrestrial export of organic matter. Compared to the pre-acidification period, the modern sediments had significantly higher Fe, Mn, Zn, Pb, and K, but lower Mg concentrations. The Zn and Pb enrichment was more evident in oligotrophic alpine lakes than in more productive forest lakes and was independent of lake water or sediment pH. Fe and Mn concentrations in the modern sediments were higher than in ambient soils and bedrock, while those in pre-acidification sediments were similar to contemporary soils and to the rock layer. The enrichment of the modern sediments with Fe and Mn thus probably resulted from both their redox recycling and ecosystem acidification.     

西藏达则错盐湖沉积背景与有机沉积结构

以西藏拟溞(Daphniopsis tibetana Sars)为优势浮游动物物种的低盐度盐湖是西藏湖泊的一个重要类型,以达则错为代表,分析了其沉积背景及沉积物组成。结果如下:(1)湖泊敞水区无机沉积以内生化学沉积为主,可代表深水盐湖无机沉积物的自然沉积过程。(2)达则错盐湖浮游植物以蓝藻、硅藻、裸藻、绿藻为主,总生物量11.35 mg/L;浮游动物生物量为4.92 mg/L,其中西藏拟溞占 82.30%;浮游植物残体受盐梯度影响在盐梯度层之上聚集,而浮游动物残体及粪粒(Fecal pellets)因外表有碳酸盐附着可穿过盐梯度层沉积湖底,生物残体与浮游动物代谢产物构成了沉积有机物的物质基础。(3)表层沉积物平均含水量为66.70%,粒径0.004-0.02 mm范围内的颗粒物含量最大,占20.42%,其次为<0.004 mm的粘土,占4.53%。(4)表层沉积物总有机碳(TOC)平均含量为27.99 mg/g(干重),其中颗粒有机碳(POC)约为18.11 mg/g,占TOC的64.70%;在POC中,西藏拟溞粪粒贡献最大,约占POC的60.48%,占TOC的39.06%,占沉积物总量的1.12%,其次为西藏拟溞残体,占POC的38.85%。分析结果表明盐湖因其独特的水化学和生物学特征具有较强的沉积能力,以化学沉积为主的无机沉积及以西藏拟溞粪粒和残肢碎屑为主的有机沉积构成了该类型盐湖颗粒物沉降及沉积的主要过程。     

Effects of nitrogen enrichment on zooplankton biomass and N:P recycling ratios across a DOC gradient in northern-latitude lakes

We used data from whole-lake studies to assess how changes in food quantity (phytoplankton biomass) and quality (phytoplankton community composition, seston C:P and N:P) with N fertilization affect zooplankton biomass, community composition and C:N:P stoichiometry, and their N:P recycling ratio along a gradient in lake DOC concentrations. We found that despite major differences in phytoplankton biomass with DOC (unimodal distributions, especially with N fertilization), no major differences in zooplankton biomass were detectable. Instead, phytoplankton to zooplankton biomass ratios were high, especially at intermediate DOC and after N fertilization, implying low trophic transfer efficiencies. An explanation for the observed low phytoplankton resource use, and biomass responses in zooplankton, was dominance of colony forming chlorophytes of reduced edibility at intermediate lake DOC, combined with reduced phytoplankton mineral quality (enhanced seston N:P) with N fertilization. N fertilization, however, increased zooplankton N:P recycling ratios, with largest impact at low DOC where phytoplankton benefitted from light sufficiently to cause enhanced seston N:P. Our results suggest that although N enrichment and increased phytoplankton biomass do not necessarily increase zooplankton biomass, bottom-up effects may still impact zooplankton and their N:P recycling ratio through promotion of phytoplankton species of low edibility and altered mineral quality.

     

Benthic carbon is inefficiently transferred in the food webs of two eutrophic shallow lakes

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Does microbial biomass affect pelagic ecosystem efficiency? An experimental study

Bacteria and other microorganisms in the pelagic zone participate in the recycling of organic matter and nutrients within the water column. The microbial loop is thought to enhance ecosystem efficiency through rapid recycling and reduced sinking rates, thus reducing the loss of nutrients contained in organisms remaining within the photic zone. We conducted experiments with lake communities in 5400-liter mesocosms, and measured the flux of materials and nutrients out of the water column. A factorial design manipulated 8 nutrient treatments: 4 phosphorus levels × 2 nitrogen levels. Total sedimentation rates were greatest in high-N mesocosms; within N-surplus communities, 1 µM P resulted in 50% increase in total particulate losses. P additions without added N had small effects on nutrient losses from the photic zone; +2 µM P tanks received 334 mg P per tank, yet after 14 days lost only 69 mg more particulate-P than did control communities. Nutrient treatments resulted in marked differences in phytoplankton biomass (twofold N effect, fivefold P effect in +N mesocosms only), bacterioplankton densities (twofold N-effect, twofold P effects in -N and +N mesocosms), and the relative importance of autotrophic picoplankton (maximum in high NY mesocosms). Multiple regression analysis found that of 8 plankton and water chemistry variables, the ratio of autotrophic picoplankton to total phytoplankton (measured as chlorophyll ) explained the largest portion of the total variation in sedimentation loss rates (65% of P-flux, 57% of N-flux, 26% of total flux). In each case, systems with greater relative importance of autotrophic picoplankton had significantly reduced loss rates. In contrast, greater numbers of planktonic bacteria were associated with increased sedimentation rates and lower system efficiency. We suggest that different microbial components may have contrasting effects on the presumed enhanced efficiency provided by the microbial loop.Correspondence to: J. D. Wehr.     

Autotroph:herbivore biomass ratios; carbon deficits judged from plankton data

A survey on phytoplankton:zooplankton biomass ratios was performed in 342 Norwegian lakes, covering a wide range in lake size and productivity (total phosphorus: 3–246 g l^–1), but with most localities being oligo- to mesotrophic. Mean phytoplankton biomass was 88 g C l^–1, yet with the majority below 50 g C l^–1and a median of 25 g C l^–1. Total zooplankton biomass displayed a mean and median of 37 and 26 g C l^–1, respectively. Cladocerans were by far the dominant group, making up a median of almost 60% of total zooplankton biomass. Total zooplankton biomass as well as that of major aggregated metazoan taxa (cladocerans, calanoid copepods, cyclopoid copepods and rotifers) all showed a positive, but weak correlation with total phytoplankton biomass. These weak correlations suggest that algal biomass per se is a poor predictor of zooplankton biomass. An average phyto-:zooplankton biomass ratio (C:C) of 2.8 (SD±4.7) was found. 30% of the lakes had a phyto-:zooplankton biomass ratio below unity. While there was no correlation between the phyto-:zooplankton biomass ratio with increasing productivity in terms of P concentration, there was a higher biomass ratio in lakes with high fish predation pressure. The low ratio of phyto-:zooplankton biomass suggest major requirements from non-algal sources of C in the zooplankton diet. The need for dietary subsidizing is also supported by the fact that more than 75% of the lakes had algal biomass less than the estimated threshold for net positive growth of zooplankton, although it should be kept in mind that a high share of picoplankton would imply an underestimation of autotroph biomass in these lakes. Since the C-deficiency apparently is most pronounced in oligotrophic systems, it contradicts the view that the detritus pathways plays a predominant role in highly productive systems only, but while the source of detritus probably is mostly of autochthonous origin in eutrophic lakes, allochthonous detritus will be more important in oligotrophic systems.     

The sediment column as a record of trophic status: examples from Bosherston Lakes,SW Wales

Bosherston Lakes are a series of interconnected, mesotrophic to hypereutrophic, artificially-created coastal marl lakes in Dyfed, South West Wales. Progressive eutrophication of the lake system has been produced by a high external phosphorus loading which includes phosphorus-rich effluent from a sewage treatment works (STW) in the catchment of the Lakes.Cores were taken from four sites of varying eutrophic status within the Lakes. In the surface sediment layer, organic C, N and P concentrations generally correlate directly with trophic status and reflect distance from the source of P input. At one site, sediment stratigraphy records a clear transition at 20–15 cm depth, marked by a sharp upward increase in porosity, organic C, N, and P, and iron-associated-P; decreases in organic matter C/N, C/P and N/P ratios; a sharp decrease in carbonate, and a change in the subfossil diatom assemblage. Lead-210 dating indicates that this change occurred in the period 1919 to 1938.The diatom stratigraphy and sediment geochemistry suggest that this transition reflects an increase in trophic status at this site, probably as a result of the influx of nutrient-rich water. This took place when the management of the Stackpole estate surrounding the lake system, fell into decline during the period 1919–1938.     

Sources and fluxes of particulate organic matter in shallow coastal waters characterized by summer macroaggregate formation

The origin and temporal variation in composition of sedimented particulate organic matter (POM) in the Gulf of Trieste (northern Adriatic) was studied over the year with special reference to the composition and sedimentation of macroaggregates in summer of 1991 using sediment traps.Suspended and sedimented POM, comprising a minor part of the largely inorganic total particulate matter, was prevalently of marine origin and composed mostly of humic substances followed by carbohydrates and proteins. Seasonal variations of particulate proteins and carbohydrates were correlated with variations of phytoplankton biomass. New production, occurring in late spring as a consequence of massive riverine inputs of N, Si and P nutrients in the surface layer of the Gulf, produced high particulate carbohydrate and protein concentrations. Subsequent depletion of introduced nutrients caused the decrease of particulate protein concentration but not that of particulate carbohydrate. The prolonged plankton biosynthesis of carbohydrates successively produced marine snow and later macroaggregates. The macroaggregates were characterized by ¹³C value of –19.9. and their carbohydrates were mostly composed of glucose followed in decreasing order by mannose, fructose, galactose, arabinose, ribose, xylose and fucose, suggesting a prevalent origin from phytoplankton structural heteropolysaccharides. Sedimentation of particulate organic constituents in the summertime, characterized by the massive presence of macroaggregates in the surface layer above the pycnocline, was the highest at a depth of 10 m at the end of this phenomenon, about six weeks after its first appearance. Sedimented macroaggregates were clearly traced by a characteristic ¹³C signal and higher carbohydrate concentrations. The monosaccharide composition was influenced by selective degradation in the water column. Sedimented POM in the bottom layer was, on the other hand, more affected by sediment resuspension. The mean yearly decrease of particulate protein-C and carbohydrate-C by 40–50% in the water column between the depths of 10 and 20 m indicates the preferential utilization of these constituents by microorganisms. The decrease of particulate humic-C is probably more the result of the export of particulate matter from the Gulf. This study also indicates that the macroaggregate formation has little impact on the annual C and N budget in such coastal areas.     

Elemental ratios and the uptake and release of nutrients by phytoplankton and bacteria in three lakes of the Canadian shield

The dynamics of carbon (C), nitrogen (N), and phosphorus (P), elemental ratios, and dark uptake/release of N and P in bacterial and phytoplankton size fractions were studied during summer 1992 in three lakes of contrasting food web structure and trophic status (L240, L110, L227). We wished to determine if phytoplankton and bacteria differed in their elemental characteristics and to evaluate whether the functional role of bacteria in nutrient cycling (i.e., as sink or source) depended on bacterial elemental characteristics. Bacterial contributions to total suspended particulate material and to fluxes of nutrients in the dark were substantial and varied for different elements. This indicated that some techniques for assaying phytoplankton physiological condition are compromised by bacterial contributions. C/N ratios were generally less variable than C/P and N/P ratios. Both elemental ratios and biomass-normalized N and P flux indicated that phytoplankton growth in each lake was predominantly P-limited, although in L227 these data reflect the dominance of N-fixing cyanobacteria, and N was likely limiting early in the sampling season. In L227, phytoplankton N/P ratio and biomass-normalized N flux were negatively correlated, indicating that flux data were likely a reasonable measure of the N status of the phytoplankton. However, for L227 phytoplankton, P-flux per unit biomass was a hyperbolic function of N/P, suggesting that the dominant L227 cyanobacteria have a limited uptake and storage capacity and that P-flux per unit biomass may not be a good gauge of the P-limitation status of phytoplankton in this situation. Examination of N-flux data in the bacterial size fraction relative to the N/P ratio of the bacteria revealed a threshold N/P ratio (22:1 N/P, by atoms), below which, bacteria took up and sequestered added N, and above which, N was released. Thus, the functional role of bacteria in N cycling in these ecosystems depended on their N/P stoichiometry.