The effect of herbivorous crustacean zooplankton on epilimnetic carbon and phosphorus cycling

Plastic limnocorrals (LCs; volume 70 m³) in oligo-mesotrophic Lake Lucerne, Switzerland were used to manipulate planktonic communities by (1) removing large zooplankton at the start of the experiment, and (2) adding phosphate during experiments of two weeks duration.Primary production (¹⁴C-assimilation), chlorophyll, standing crops and sedimentation of particulate organic carbon (POC) and of particulate phosphorus, as well as plankton composition were assessed simultaneously in two to four differently treated LCs. Carbon and phosphorus mass balances were calculated from assimilation, temporal change in standing crop of particulate matter, and sedimentation.A quick elimination of crustaceans by screening hardly increased primary production, but decreased sedimentation and mineralisation of particulate organic carbon, and thus significantly enhanced the standing crop of POC, but decreased POC turnover. The exclusion of crustaceans increased the mean residence time of total phosphorus by a reduced P loss by sedimentation.Increased grazing pressure during the experiment showed little grazing induced effect on plankton composition and primary production except at the very end.We conclude that in an oligo-mesotrophic lake, buffering mechanisms attenuate the impact of changing grazing pressures on primary production.     

Can results from limnocorral experiments be transferred to in situ conditions?

In 1982–1984 eight limnocorral (LC) experiments, each lasting for two weeks, were performed in mesotrophic Lake Lucerne, Switzerland, to study the effects of biomanipulation (removal of crustaceans by 95 µm filter nets) on zooplankton — phytoplankton relationships and epilimnetic carbon and phosphorus fluxes. Seston concentrations and to a lesser extent primary production rates were reduced in control LCs through zooplankton grazing, and settling flux increased through fecal pellet production. But C and P regeneration were not significantly affected.We found several indications that the LCs, despite of their large size ( 70 m³, 3 m in diameter and 11 m long) were artificial systems when compared to the surrounding lake: The eddy diffusion was diminished by about one order of magnitude, nutrients depleted (but phosphorus was supplied to the LCs), the phyto- and zooplankton showed lower standing crops in the control LCs than in the lake, the phytoplankton showed a shift from nannoplankton to netplankton, the crustacean zooplankton was mostly limited in their vertical migration, and the POC and PP sedimentation rates were increased. Moreover, in our set of experiments we always found outliers, which may have been caused by the different in situ conditions at the beginning of the experiments, and further enhanced by the complexity of the enclosed system.The problems of the LC-technique, such as replicability, scaling (size and time) and data extrapolation are discussed. The impact of crustacean zooplankton on particulate matter, i.e. seston reduction and sedimentation enhancement, can be extrapolated on a qualitative rather than on a quantitative basis for the Lake Lucerne ecosystem.     

Variation in the C:P ratio of suspended and settling seston and its significance for P uptake calculations

SUMMARY. 1. In 1981–84 limnocorral (LC) experiments were performed in Lake Lucerne. Switzerland, to manipulate the planktonic community by varying P fertilization and by removing large zooplankton (with a 95 μm screen).
2. The C:P ratios in both suspended and entrapped seston exceeded the 'ideal' C:P ratio of 106 proposed by Redfield, Ketchum & Richards (1963) when P was limiting algal growth.
3. P fertilization could decrease the sestonic C:P ratio to 106 only when P did not limit algal growth; P additions far exceeding the P loading of eutrophic lakes were necessary to obtain this situation.
4. Changes in epilimnetic C:P ratios were usually related to short- term changes in primary production, caused by variable in situ light conditions and turbulence, and subsequent variation in POC concentrations.
5. Entrapped seston in the 95 μm-filtered LCs showed C:P ratios slightly higher than those of suspended seston, indicating fast P release and slower C mineralization in the epilimnetic nutrient cycle.
6. Removing large crustacean zooplankton enhanced epilimnetic P mineralization, and C:P ratios of entrapped seston in the 95 μm-filtered LCs were increased.
7 Detritus formed a relatively high proportion of the seston and amounted to more than two-thirds of the measured POC concentration.
8. Calculations of algal P uptake using information on sestonic C:P ratios and ¹⁴C uptake rates are questionable, as long as detritus cannot be separated from algae and net carbon uptake cannot be accurately measured.     

Effects of a filter-feeding fish [silver carp, Hypophthalmichthys molitrix (Val.)] on phyto- and zooplankton in a mesotrophic reservoir: results from an enclosure experiment

SUMMARY 1. Silver carp, Hypophthalmichthys molitrix (Val.), feeds on both phyto- and zooplankton and has been used in lake biomanipulation studies to suppress algal biomass. Because reports on the effects of silver carp on lake food webs have been contradictory, we conducted an enclosure experiment to test how a moderate biomass of the fish (10 g wet weight m⁻³) affects phytoplankton and crustacean zooplankton in a mesotrophic temperate reservoir.
2. Phytoplankton biomass <30 μm and particulate organic carbon (POC) <30 μm were significantly higher in enclosures with silver carp than in enclosures without fish, whereas Secchi depth was lower. Total copepod biomass declined strongly in both treatments during the experiment, but it was significantly higher in fish-free enclosures. Daphnid biomass was also consistently higher in enclosures without fish, although this effect was not significant. However, the presence of fish led to a fast and significant decrease in the size at maturity of Daphnia galeata Sars. Thus, the moderate biomass of silver carp had a stronger negative effect on cladoceran zooplankton than on phytoplankton.
3. Based on these results and those of previous studies, we conclude that silver carp should be used for biomanipulation only if the primary aim is to reduce nuisance blooms of large phytoplankton species (e.g. cyanobacteria) that cannot be effectively controlled by large herbivorous zooplankton. Therefore, stocking of silver carp appears to be most appropriate in tropical lakes that are highly productive and naturally lack large cladoceran zooplankton.     

虾池生态系各有机碳库的储量

1997年6～8月于山东省海阳市黄海集团公司养虾场,采用围隔实验生态学方法研究了虾池生态系各有机碳库的储量,结果表明:溶解有机碳(DOC)、颗粒有机碳(POC)库储量(mgC/l)平均分别为9.95±2.04及3.10±1.60,DOC/POC比值平均为3.21.POC库中,腐质颗粒碳为总浮游生物碳的2倍.浮游植物碳、浮游动物碳、浮游细菌碳平均分别占总浮游生物碳的53%、13%及34%.腐质碳、总浮游生物碳平均分别占总有机碳(TOC)的92%及8%,腐质碳为总浮游生物碳的11.08倍.     

Characteristics of settling matter and its role in nutrient cycles in a deep oligotrophic lake

The settling flux of seston (dry weight, DW), chlorophyll a (Chl a), particulate organic carbon (POC), particulate organic nitrogen (PON), and particulate phosphorus (PP) was measured monthly in 1981–1983 at 10 different depths in Lake Chuzenji, Japan; an oligotrophic lake with a maximum depth of 163 m. The Ti concentration in entrapped matter was used to separate the sedimentation flux into allochthonous and autochthonous components. Inflow loads of dissolved nutrients (DN: 4.5, DP: 0.48 g m^-2a^-1) were almost sufficient to supply the autochthonous fluxes at 30 m (PON: 2.9, PP: 0.51 g m^-2a^-1 ), and this flux of POC (26.6 g m^-2a ^-1) was about one-third of primary production (84 g C M^-2a^-1). Sedimentation of particulate matter was the main path of losing nutrients from lake water, explaining more than 80% removal of inflow loads (TN, TP). Decomposition rates during sedimentation which were calculated from the vertical difference in the autochthonous flux agreed very closely with the results obtained by laboratory experiments of a 100-day incubation (content ratios from field observations were: POC 0.67, PON 0.65, PP 0.85; and from laboratory experiments they were: POC 0.68, PON 0.70, PP 0.94). These decomposition rates and those near the sediment interface were used to explain dissolved oxygen depletion and nitrate increase in the hypolimnion during stratification. The average sinking velocities were 1.82m d^-1 for seston and 1.16 m d^-1 for Chl a at 30m, they were influenced by Chl a content of seston.     

Biogeochemistry of organic matter in Lake Geneva: I — particulate hydrocarbons as biogenic and anthropogenic molecular markers

A study of the biogeochemistry of organic matter in Lake Geneva at a central station (SHL2), was carried out at key periods of the biological cycles from November 1985 to September 1986. The seasonal variability of particulate hydrocarbons was studied in the epilimnion and the hypolimnion, focusing on specific periods such as high phytoplankton productivity and rising herbivorous zooplankton activity (May 1986), clearness of waters (June 1986) and intense mixing of waters (February 1986).Analysis of non-aromatic hydrocarbons (NAH) indicated that their major source is the autochtonous phytoplankton. Year 1986 was characterized by a spring bloom of nanoplankton (chlorophyceae and cryptophyceae) which remained predominant during summer and fall. These biological events were correlated with an increase of biogenic NAH, particulary n-alkane and alkene concentrations; the latter increased by one order of magnitude in May. n-Alkane distributions maximized at n-C₁₇ or n-C₂₉, and the n-C₁₇/n-C₂₉ ratio varied in surface water from 2.8 in May to 0.06 in November of the preceding year, reflecting phytoplankton dominant species. Values of NAH/POC, n-alkanes/POC, alkenes/POC and pristane/phytane ratios underwent seasonal and vertical variations which could be related to grazing and/or settling of particles. NAH, n-alkane and alkene enrichment in hypolimnion particles was interpreted to be a result of zooplankton migrations.A series of 37 monoenes and polyenes having from 15 to 31 carbon atoms, originating from planktonic sources, were recognized in the particulate matter by gas chromatography/mass spectrometry (GC/MS). Their distribution was described in 3 sized particles (> 200 µm, 200-60 µm and < 60 µm, identified as adult zooplankton, young zooplankton and nanoplankton respectively) at two different periods of the biological cycle, May and June 1986. Low level inputs originating from terrestrial higher plants were evident in particles; the value of the Carbon Preference Index (CPI) in the carbon range 23–35 was less than 2.8.In all samples analyzed by GC, an unresolved complex mixture of hydrocarbons (UCM) was present and was interpreted, at least in May, as partly originating from the degradation of organic matter by microorganisms. Typical anthropogenic PAH, such as benzofluoranthene and dibenzo (a) anthracene, were found at a low level (9 and 3 ng l^–1 respectively). The absence of hopane-type hydrocarbons, except in November 1985 and in September 1986, and typical combustion PAH associated with particles indicated high temperature pyrolytic contaminants rather than petroleum-derived compounds.     

The role of zooplankton grazing in the formation of `clear water phase' in a shallow charophyte-dominated lake

In Chara-dominated shallow eutrophic Lake Prossa (Estonia), the collapse of spring phytoplankton community occurred in late May after which both primary production (PP) and phytoplankton biomass (B&pinf;) stayed at a very low level. By mid-June the Secchi depth had increased up to 2.6 m indicating the achievement of the `clear water phase', which persisted thoughout the rest of the vegetation period. The biomass of `edible' phytoplankton formed on average 53% of the total phytoplankton biomass, and the share of herbivorous zooplankton was on average 61% of the total zooplankton biomass. In spring zooplankton removed daily 27% of the total B&pinf; and 29% of PP by grazing while in summer these values rarely exceeded 5%. Zooplankton grazing was responsible for the decrease of `edible' (<31 μm) phytoplankton after its spring peak as well as for maintaining its biomass at a very low level during the whole vegetation period. Depletion of mineral forms of nitrogen and phosphorus that occurred most probably because of the development of charophytes by the end of May supported the collapse of the whole phytoplankton community and kept the water clear throughout the summer and autumn.

     

Benthic and pelagic food resources for zooplankton in shallow high-latitude lakes and ponds

1. Shallow lakes and ponds are a major component of the northern landscape and often contain a high zooplankton biomass despite clear waters that are poor in phytoplankton. 2. In this study we quantified zooplankton food sources and feeding rates in the shallow waters of two contrasting high‐latitude biomes: subarctic forest tundra (Kuujjuarapik, Quebec) and high arctic polar desert (Resolute, Nunavut). Five substrate types were tested (beads, bacteria, picophytoplankton, filamentous plankton and microbial mats). Special attention was given to the role of benthos, a component that is usually poorly integrated into models of aquatic foodwebs. 3. Consistent with observations elsewhere in the circumpolar region, high concentrations of adult macrozooplankton occurred in all sites (up to 17 100 crustaceans m^?3) while phytoplankton concentrations and primary productivity were low. The communities were composed of multiple species, including Daphnia middendorfiana, Hesperodiaptomus arcticus, Leptodiaptomus minutus, Artemiopsis stefanssoni and Branchinecta paludosa. 4. Detritus made 89–98% of the planktonic resource pool and bacteria contributed the highest biomass (up to 29 mg C m^?3) of the planktonic food particles available to zooplankton. Benthic resources were dominated by microbial mats that grew in nutrient‐rich conditions at the base of the ponds and which dominated overall ecosystem biomass and productivity. 5. All species were flexible in their feeding but there were large, order of magnitude differences in clearance rates among taxa. These differences likely resulted from different grazing strategies among cladocerans, copepods and fairy shrimps, and possibly also from adaptation to specific food types and size ranges that occur locally in these waters. 6. The subarctic cladocerans Ceriodaphnia quadrangula and D. middendorfiana, and the arctic fairy shrimp B. paludosa were observed to graze directly on the microbial mats and the feeding experiments confirmed their assimilation of benthic substrates. The other zooplankton species showed a more pelagic feeding mode but were capable of using microbial mat filaments, thus may be indirectly linked to benthic processes via resuspension. 7. Our study indicates that the classical aquatic food web in which phytoplankton provide the sole production base for grazers does not apply to northern shallow lakes and ponds. Instead, microbial mats increase the physical complexity of these high latitude ecosystems and likely play a role in sustaining their high zooplankton biomass.     

Effects of zooplankton on the sinking flux of organic carbon in Lake Biwa

To clarify the roles of zooplankton in the sedimentation of seston from the epilimnion, the sinking flux of particulate carbon was measured along with primary production rate and zooplankton biomass from July 1996 to October 1997 at a pelagic site in the north basin of Lake Biwa. During the study period, the flux varied seasonally from 66 to 510 mg C m⁻² day⁻¹ and was low in summer when zooplankton, composed mainly of Eodiaptomus japonicus and Daphnia galeata, were abundant. Simple correlation analysis revealed that the sinking flux correlated neither with the primary production rate nor with the amount of sestonic carbon above the sediment trap. However, the particle elimination rate, estimated as the difference between the primary production rate and the sinking flux, correlated positively with the zooplankton biomass. These results suggest that zooplankton play a substantial role in decreasing the sinking flux in Lake Biwa. Received: March 6, 2000 / Accepted: October 7, 2000     

The role of zooplankton in the ecological succession of plankton and benthic algae across a salinity gradient in the Shark Bay solar salt ponds

The relatively low biodiversity and simple hydrodynamics make solar salt ponds ideal sites for ecological studies. We have studied the ecological gradient of the primary ponds at the Shark Bay Resources solar salt ponds, Western Australia, using a coupled hydrodynamic ecological numerical model, DYRESM–CAEDYM. Seven ponds representative of the primary system were simulated with salinity ranging from 45 to 155 ppt. Five groups of organisms were simulated: three phytoplankton, one microbial mat plankton, and one zooplankton as well as dissolved inorganic and particulate organic nitrogen, phosphorus, and carbon. By extracting the various carbon fluxes from the model, we determined the role that the introduced zooplankton, Artemia sp., plays in grazing the particulate organic carbon (POC) from the water column in the high salinity ponds. We also examined the nutrient fluxes and stoichiometric ratios of the various organic components for each pond to establish the extent to which observed patterns in nutrient dynamics are mediated by the presence of Artemia sp. Model results indicated that Artemia sp. grazing was responsible for reduced water column POC in the higher salinity ponds. This resulted in an increase in photosynthetic available radiation (PAR) reaching the pond floor and consequent increase in microbial mat biomass, thus demonstrating the dual benefits of Artemia sp. to salt production in improved quality and quantity. In contrast, this study found no direct link between Artemia sp. and observed changes in planktonic algal species composition or nutrient limitation across the salinity gradient of the ponds. Guest Editors: J. John & B. Timms Salt Lake Research: Biodiversity and Conservation—Selected Papers from the 9th Conference of the International Society for Salt Lake Research     

Carbon pathways to zooplankton: insights from the combined use of stable isotope and fatty acid biomarkers

1. Numerous studies have quantified the relative contribution of terrestrial‐ and phytoplankton‐derived carbon sources to zooplankton secondary production in lakes. However, few investigated the pathways along which allochthonous and autochthonous carbon (C) was actually conveyed to consumers. 2. We suggest that the combined use of fatty acid and stable isotope biomarkers could solve this issue. We conducted a field study on two oligotrophic lakes, in which primary production increased significantly between 2002 and 2004. We used modelling to estimate the contribution of terrestrial‐ and phytoplankton‐derived C to particulate organic C (POC) and zooplankton production from their δ¹³C values in 2002 and 2004. 3. According to the isotope model, phytoplankton‐derived C accounted for a major part of the POC pool in both lakes and supported more Daphnia sp. production in 2004 than in 2002. Fatty acid data revealed that increased contribution of algal‐C to Daphnia production, although common between both lakes, was achieved through C pathways that were different. In one lake, Daphnia grazed more intensively on phytoplankton, whereas in the other there was greater grazing on bacteria. In the latter case, the increased primary production resulted in greater release of algal‐derived dissolved organic C (DOC), which may have supported extra bacterial and eventually Daphnia, production. 4. This is the first study illustrating that the combination of fatty acid and stable isotope biomarkers could further our understanding of the factors controlling the relative magnitude of food webs pathways conveying organic matter to zooplankton.     

Terrestrial subsidies to lake food webs: an experimental approach

Cross-ecosystem movements of material and energy are ubiquitous. Aquatic ecosystems typically receive material that also includes organic matter from the surrounding catchment. Terrestrial-derived (allochthonous) organic matter can enter aquatic ecosystems in dissolved or particulate form. Several studies have highlighted the importance of dissolved organic carbon to aquatic consumers, but less is known about allochthonous particulate organic carbon (POC). Similarly, most studies showing the effects of allochthonous organic carbon (OC) on aquatic consumers have investigated pelagic habitats; the effects of allochthonous OC on benthic communities are less well studied. Allochthonous inputs might further decrease primary production through light reduction, thereby potentially affecting autotrophic resource availability to consumers. Here, an enclosure experiment was carried out to test the importance of POC input and light availability on the resource use in a benthic food web of a clear-water lake. Corn starch (a C(4) plant) was used as a POC source due to its insoluble nature and its distinct carbon stable isotope value (δ(13)C). The starch carbon was closely dispersed over the bottom of the enclosures to study the fate of a POC source exclusively available to sediment biota. The addition of starch carbon resulted in a clear shift in the isotopic signature of surface-dwelling herbivorous and predatory invertebrates. Although the starch carbon was added solely to the sediment surface, the carbon originating from the starch reached zooplankton. We suggest that allochthonous POC can subsidize benthic food webs directly and can be further transferred to pelagic systems, thereby highlighting the importance of benthic pathways for pelagic habitats.     

Structural-functional relationships in the pelagic community of the eastern tropical Atlantic Ocean

Sampling was carried out in the eastern tropical Atlantic duringJuly–September, 1987 on four transects at latitudes 6,12, 18 and 24°W, from 6°S to 4°N. A total numberof 81 stations were occupied where physical, chemical and biologicalmeasurements were taken. Temperature, salinity, chlorophylla, phytoplankton, primary production (PP), bacterial production,micro-, meso-, macrozooplankton and mesopelagic fish and squidwere sampled in the upper 120 m layer. The biomass size spectrumof the plankton community was relatively flat within the divergencezone but the slope of the size spectrum increased towards theconvergence zone meaning an increased contribution of the small-sizedorganisms. Higher turnover rates (the ratio of PP to the totalcommunity biomass) were observed in regions where the size spectrasloped more steeply. PP exceeded the community metabolism bythree to five times in regions where the size spectra were sharplysloped. In regions with flat-type spectra (with slopes from-0.2 to 0) a balance between PP and community metabolism wasobserved. The variation coefficient and the dispersion index(the variance to mean ratio) indicated an increase of the spatialvariability of biomass in the sequence: ‘phytoplankton–mesozooplankton–macrozooplankton–micronekton’.At the higher trophic level of the zooplankton community, betweenthe biomass of squid and gelatinous zooplankton, a logarithmiclinear relationship was derived. Squid actively consumed gelatinousorganisms in regions of high concentrations of their prey, althoughlarge copepods, euphausiids, amphipods, flying fish, and myctophidfish also contributed to their diet.     

Role of bottom sediments in sustaining plankton production in a Lake ecosystem — Experimental demonstration using enclosed water bodies in a shallow eutrophic Lake

Using large enclosures set in Lake Suwa, experimental studies were conducted to examine the effects of bottom sediments on plankton production in a natural lake ecosystem. Successive changes in biotic and abiotic components in two types of enclosure with and without bottom sediments were surveyed throughout a one-month period in the early fall of 1985. Remarkable differences were found between the two types of enclosure with regard to the time courses of abundance of chlorophyll, PON, PP, TN, TP and zooplankton as well as those of POC/PON and POC/Chl ratios and primary production rate in terms of mg N m⁻² day⁻¹. Quantitative examination of the major causes of these differences revealed that the release of nitrogen and phosphorus from the bottom sediments was significantly effective for sustaining the active production of phytoplankton cells with a high nitrogen and phosphorus content, subsequently resulting in active zooplankton growth.     

The Significance of Zooplankton in the Cycling of Phosphorus in Lakes of Different Trophic Categories

The species composition, number, biomass and phosphorus excretion rate (PER) of zooplankton (Crustacea and Rotatoria) in 8 mesotrophic and eutrophic lakes were studied. PER were calculated using specific expenditures on metabolism estimated from the oxygen uptake, phosphorus content in zooplankton and field observations of number and biomass of planktonic Rotatoria and Crustacea. The zooplankton biomass and PER increased from mesotrophic to eutrophic lakes. The values of PER were compared with primary production and spring phosphorus concentration. The relative significance of zooplankton in phosphorus cycling declined with increasing trophic category of lakes.     

Impact of moderate silver carp biomass gradient on zooplankton communities in a eutrophic reservoir. Consequences for the use of silver carp in biomanipulation.

We examined the impacts of moderate gradient silver carp biomass (five levels from 0 to 36 g.m-3, i.e. about 0-792 kg.ha-1) on zooplankton communities of the eutrophic Villerest reservoir (France). During our mesocosm experiment changes in zooplankton assemblages were dependent on silver carp biomass. In the fishless and low fish biomass treatments, zooplankton abundance increased through time, owing to a peak in cladoceran density, but decreased (mainly cladocerans) at highest fish biomass. Copepods and rotifers were less affected at the highest fish biomass and dominated zooplankton communities. We highlighted that the presence of high silver carp biomass could lead to changes in phytoplankton assemblage via the impact on herbivorous zooplankton. Since silver carp efficiently graze on particles > 20 microns, the suppression of herbivorous cladocerans could result in an increase in small size algae (< 20 microns) abundance since these species would be released from grazers as well as competitors (large algae grazed by silver carp) and nutrients levels would be enhanced by fish internal loading. Our results showed that the use of low silver carp biomass (< 200 kg.ha-1) would allow us to minimize these negative effects.     

Size‐specific effects of bighead carp predation across the zooplankton size spectra

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Occurrence,Dynamics and Production of Picoplankton in Hungarian Shallow Lakes

Phototrophic picoplankton were detected in 10⁵-10⁶ cells/ml concentrations in seven water bodies of differing chemistry and trophic level. Dominant picoplankters were, at all sites, coccoid cyanobacteria of 0.8-1.2 μm dimensions, exhibiting red or yellow autofluorescence. Apart from the effects of water temperature their quantitative dynamics were significantly influenced by the nitrogen supply and the herbivorous zooplankton (Cladocera). Their maximum contribution to the total planktonic primary production was about 50%, both in mesotrophic and hypereutrophic environments. However during the bloom of filamentous nitrogen-fixing blue-greens their role became negligible. In phytoplankton communities the significance of picoplankton is overestimated several times, when based on the cell count, while it is underestimated on the basis of biomass. The most useful characteristics of phytoplankton size groups was the total surface area of their cells.     

Invasive trout increase the climatic sensitivity of zooplankton communities in naturally fishless lakes

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