Nutrient dynamics and plankton cycles in artificial ponds used in the production of oyster Ostrea edulis L. spat

The European flat oyster, Ostrea edulis L., has been successfully cultured in Cork Harbour on the south coast of Ireland, using non-tidal, man-made ponds for over 20 years. Broodstock placed in ponds spawn and the released larvae grow and settle on mussel shell cultch. This study investigates the nutrient dynamics and plankton cycles that occur from the time the ponds are filled. The incoming water to the ponds, when filled in May-June, is usually nutrient rich, with high numbers of diatoms. The nutrients become depleted quickly and the phytoplankton becomes dominated by smaller algal species, particularly flagellates and by dinoflagellates. Following mussel shell addition, orthophosphate levels increase and this pulse in nutrients is generally followed by an increase in microplankton, particularly Chlorella species. This increase often reaches bloom proportions and, if so, ponds are flushed. This addition of water increases the nutrient level and changes the phytoplankton composition, usually by introducing new dinoflagellate and diatom species.     

Suspended culture of the sea cucumber Apostichopus japonicus below a Pacific oyster raft – potential for integrated multi‐trophic aquaculture

To discuss the possibility of co‐culturing Pacific oyster with the sea cucumber Apostichopus japonicus, a field experiment was conducted in an oyster farm. Apostichopus japonicus juveniles (mean wet weight, 0.08 g) were cultured below an oyster raft and at a control station for 216 days, and the wet weight and stable isotope ratios (δ¹³C, δ¹⁵N) were analysed together with settling organic matter (OM) collected using sediment traps. All sea cucumbers cultured below the raft survived (survival rate, 100%), while at the control station one individual disappeared (96%). During 216 days, the juveniles at the oyster and control stations grew to a mean weight of 5.5 and 2.6 g, attaining respective specific growth rates of 2.0% and 1.6% (paired t‐test, P < 0.001). Settlement rates of carbon and nitrogen at the oyster station were ~5 times larger than those at the control station. The stable isotope analysis showed that settling OM at both stations originated from coastal phytoplankton and that phytoplankton represented the primary food source for A. japonicus. The rapid growth of A. japonicus at the oyster station was concluded to be due to the abundant supply of oyster biodeposits, which could be ingested by this species.     

Enclosure culture ban and its effects on water quality and phytoplankton community in a subtropical shallow lake

Enclosure culture, ubiquitous anthropogenic activity in shallow lakes, was prohibited in East Taihu Lake with all breeding infrastructure demolished at the end of 2018. However, the ecological responses of water quality change and phytoplankton community succession to the enclosure culture ban remain largely unknown. In this study, we conducted field investigations monthly in East Taihu Lake to present a comprehensive study of water quality and phytoplankton community variation before (2018) and after (2019) the enclosure culture ban. With the cessation of aquaculture activities, water quality significantly improved, TN and TP decreased by 54% and 34%, respectively, and eutrophic state shifted from light eutrophic (mean TSI = 59) to mesotrophic status (mean TSI = 48) in autumn. Chl-a decreased from 33.34 ± 16.87 to 24.84 ± 15.43 μg/L, and SDD increased from 0.76 ± 0.19 to 0.95 ± 0.36 m. Meanwhile, phytoplankton abundance significantly decreased from 3.6 × 10⁷ to 1.6 × 10⁷ cell/L, with the contribution of Cyanophyta markedly decreasing from 82% to 63%. In addition, the heterogeneity of phytoplankton community markedly increased. The changes in water quality and phytoplankton following the enclosure culture ban were largely due to the suspension of external nutrient input from breeding activity and improved water clarity. Redundancy analysis (RDA) analysis further elucidated that water temperature and nutrients were the main factors affecting phytoplankton dynamics. Our findings showed that the ban on enclosure culture had an extremely positive ecological impact.     

Water quality upstream and downstream of a commercial oyster aquaculture facility in Chesapeake Bay,USA

Oyster aquaculture is an expanding industry in the Chesapeake Bay. Oysters remove nitrogen (N) and phosphorus (P) from the water column through filtration and conversion of phytoplankton into shell and tissue, but also continuously excrete these same nutrients back into the water column as inorganic compounds readily available for plant or algal uptake. The objective of this study was to assess multiple water quality parameters upstream and downstream of a commercial oyster aquaculture facility in the mesohaline region of the Chesapeake Bay. Results of the study indicated a 78.4% average increase in total ammonia nitrogen (TAN) concentration and a 19.4% decrease in chlorophyll-a (Chl-a) concentration downstream of the facility. There was no significant change in the concentration of reactive phosphate (RP), nitrate–nitrogen (NO₃⁻–N), or nitrite–nitrogen (NO₂⁻–N) as water passed through the facility. It was determined that velocity of water through the facility had no influence on the change in TAN or Chl-a concentration from upstream to downstream of the facility. Increased reduction in Chl-a concentration from upstream to downstream was related to higher upstream concentrations of Chl-a. There was no correlation between increased rates of Chl-a removal and downstream TAN. Results of this study suggest that oyster aquaculture can significantly increase the amount of available inorganic nitrogen in the water column immediately downstream of a facility, independent of upstream availability of phytoplankton and flow velocity of water through the facility.     

Growth,morphometrics and nutrient content of farmed eastern oysters,Crassostrea virginica (Gmelin), in New Hampshire,USA

When harvested, oysters represent a removal from the ecosystem of nutrients such as nitrogen (N) and carbon (C). A number of factors potentially affect nutrient content, but a quantitative understanding across the geographical range of the eastern oysters is lacking. This study was designed to quantify the relationships among various metrics of farmed eastern oysters near its northern geographical range focusing on nutrient content. Hatchery‐reared oysters were deployed in polyethylene bags at six sites, and were measured on multiple occasions from 2010 to 2012. A quadratic polynomial fit to the combined datasets for shell height indicated that on average a ‘cocktail’ size oyster (63 mm shell height) would be reached after 2 year, and ‘regular’ size (76 mm) would require 3 year. There were significant differences in growth rates and oyster nutrient content among the sites; means for %N in soft tissue ranged from 6.9 to 8.6, and 0.07 to 0.18 in shell. Per cent N in soft tissue and shell were highest at two sites at the mouths of rivers with elevated dissolved inorganic N concentrations in the water. Grand means (all sites, seasons and years combined) of soft tissue N and C for regular size oysters were 7.3% and 38.5%, respectively; and for shell N and C were 0.13% and 12.0% respectively. Our study extends the range of data on nutrient content of the eastern oyster to northern New England, and indicates that oyster size, seasonality, and nutrient concentration in ambient water potentially affect %N and %C content of oysters.     

Suspended culture evaluation of Pacific oyster Crassostrea gigas in a tropical estuary

The growth and survival of Crassostrea gigas were evaluated in hanging culture in the Chone River estuary (Bahía de Caráquez), Manabí province, Ecuador. Oyster seeds (~10 mm) were confined in lantern nets at a density equal to the occupation of half the surface of the basket base, until reaching commercial size (80 mm). A sample for oyster biometric, parasitic and bacterial analysis was obtained monthly. Temperature, salinity, oxygen concentration, seston and phytoplankton biomass were determined. At the end of the study, the oysters were analysed for heavy metal concentration. The commercial oyster production was extrapolated to estimate the possible economic performance of a family production module (a 7 × 7 m bamboo raft). The results show a great biological feasibility of culture of C. gigas with harvests of commercial size oysters in only 5 months and acceptable survivals (accumulated >70%, after 3 weeks of crop seed selection). The heavy metal concentrations and the parasitological and bacteriological analyses did not indicate levels of contamination. The economic projections suggest that, even with a profitability of 57%, the initially invested capital could be recovered within the first harvest.     

The East River tidal strait,New York City,New York,a high-nutrient,low-chlorophyll coastal system

The East River tidal strait, located between New York Harbor and Western Long Island Sound, is characterized by high suspended silt concentrations with low organic content kept in suspension by intense tidal currents. Inorganic nutrients, including nitrate, nitrite, ammonia, and phosphate, were high even during the summer. Dissolved inorganic nitrogen (DIN) concentrations generally were above 20 µM and did not likely limit phytoplankton growth. Despite high nutrient concentrations, median chlorophyll a concentration was only 1.53 µg l^?1, making the East River tidal strait a high-nutrient, low-chlorophyll (HNLC) area, likely a result of suspended silt blocking light penetration into the surface water. There were times at which the ratio of mixed layer to depth of the euphotic zone was generally greater than what has been suggested for phytoplankton to produce net primary production. The high-nutrient East River tidal strait is likely one of the sources of nutrients fueling summer phytoplankton production and consequent hypoxia in the Western Long Island Sound as silt settles from surface water in the lower turbulence conditions of the western narrows of Long Island Sound, thereby allowing light penetration and subsequent consumption of dissolved nutrients by phytoplankton.

     

Modelling annual cycles of primary production in different regions of the Bohai Sea

The Bohai Sea is a typical shallow sea in which the average depth of water is about 18 m, and which has a rich biodiversity and abundance of marine life. The values of primary production differ among the four different regions of the Bohai Sea. A biological model consisting of nutrients, phytoplankton, zooplankton and detritus is used to simulate the annual cycles of plankton and primary production in these four regions. The simulation is forced by monthly observational data. Water transparency, nutrient concentrations, temperature and incident irradiance influence the amount and variation of plankton and primary production. The model simulations indicate that transparency and nutrient concentration are the principal causes of the differences in primary production among the four regions of the Bohai Sea.     

引水对尚湖浮游植物群落结构和营养盐浓度的影

摘 要：通过对引污补水作用下小型通江湖泊（以尚湖为例）中浮游植物群落和营养盐时空变化的研究发现：①研究期间,尚湖共检出浮游植物8门39属,从望虞河引水后,尚湖浮游植物群落组成中蓝藻门富营养化习见种的种类显著增加;②引水后尚湖的西湖区营养盐浓度上升2-3倍,东湖区基本维持不变;③经过较长时间的沉降和水体自净之后,浮游植物群落中蓝藻门富营养化习见种的种类减少,营养状态降至引水前水平。从望虞河引水补给尚湖,对其浮游植物群落和营养水平有显著影响。     

The impacts of aquacultured oysters, Crassostrea virginica (Gmelin, 1791) on water column nitrogen and sedimentation: results of a mesocosm study

To determine effects of aquacultured oysters Crassostrea virginica (Gmelin, 1791) on the overlying water column, a mesocosm study was performed at the Marine Ecosystem Research Laboratory (MERL) from June to October, 2000. The MERL facility is located adjacent to Narragansett Bay and consists of fourteen 13,000-l mesocosm tanks designed to simulate the Bay environmental conditions. Two hundred oysters (≈35 mm valve height; nominally filtering about 55 l/day/individual) were placed into three mesocosms, and three mesocosms were maintained without oysters as controls. Experiments were run with varying rates of water exchange in the tanks ranging from 0% to 100% per day (13,000 l/day). Parameters that were measured and compared between the two treatments included chlorophyll-a, particulate organic and inorganic matter, sedimentation, nitrate, ammonia, selected phytoplankton species and oyster growth rates. Oysters affected phytoplankton species composition and increased rates of sedimentation. Large diatoms were net sampled, and Nitzchia striata was predominant in mesocosms with oysters, while Skeletonema costatum dominated the control tanks. Ammonia excretion rates were determined for C. virginica using the salicylate–hypochlorite method. Ammonia excretion can be described by the allometric equation E=50.65w^0.699 when E is the ammonia excretion rate in μg/h, and w is the soft tissue dry weight in grams. Based on rates of ammonia excretion by oysters and observed steady states of ammonia and other forms of inorganic nitrogen in mesocosm tanks, it can be hypothesized that ammonia generated by oysters is taken up by rapidly regenerating phytoplankton in the water column.     

Relative importance of phytoplankton and periphyton on oyster-culture pens in a eutrophic tropical lagoon

The relative importance of biomass and productivity of phytoplankton and periphyton on oyster-culture pens was quantified from the tidal inlet to the inner region along a transect across a eutrophic tropical lagoon representing a gradient in flushing time over a complete seasonal cycle. Water flushing modified nutrient concentrations in the water column and the responses of phytoplankton and periphyton to nutrient enrichment. Nutrient concentrations were greater at sites in the inner region subject to poor flushing and lower at sites in the outer region subject to fast flushing. Phytoplankton chlorophyll a was greater at sites in the inner region. However, periphyton biomass decreased with increasing flushing times. Daily production rates of phytoplankton were also greater at sites in the inner region, but no significant differences in those of periphyton were observed among the study sites. On the basis of lagoon area, periphyton contributed <6% of the daily system gross production. The relative contribution of periphyton to net production was negligible when compared to that of phytoplankton. Nevertheless, there was a spatial shift in their relative contribution to the system biomass. Periphyton biomass exceeded that of phytoplankton at a well-flushed site with a large number of oyster farms available for periphyton colonization. Our results clearly demonstrate the modifying effects of tidal flushing as well as substratum area on the relative importance of periphyton biomass on oyster-culture pens in response to nutrient enrichment in coastal lagoons.     

A non‐classical biomanipulation experiment in Gonghu Bay of Lake Taihu: control of Microcystis blooms using silver and bighead carp

A non‐classical biomanipulation experiment was carried out in Gonghu Bay of Lake Taihu in 2009. Silver and bighead carp were stocked in a large fish enclosure to control cyanobacterial blooms. Water quality, plankton abundance, and the intracellular and extracellular microcystins (MCs) in lake water were investigated monthly in 2009. The concentrations of nitrogen nutrients were significantly lower in the fish enclosure than in the surrounding lake, while phosphorus (especially total phosphorus) concentration was higher in fish enclosure. During the blooming period, Cyanophyta contributed to more than 90% of the phytoplankton in the surrounding lake, whereas it represented only 40–80% in the fish enclosure. The phytoplankton and crustacean zooplankton biomasses and the zooplankton/phytoplankton ratios were all significantly lower in the fish enclosure than in the lake. This result suggested that silver and bighead carp can effectively suppress the phytoplankton biomass with the initial stocking density of 7.5 g m^?3 for silver carp and 1.1 g m^?3 for bighead carp, despite a simultaneous decrease in the grazing pressure of the zooplankton on the phytoplankton. During the blooming period, the intracellular and extracellular MCs in the fish enclosure were reduced by 93.8% and 69.8% compared with the surrounding lake. MCs content varied from 0.34 to 18.8 ng (mean 4.8 ng) MC‐LReqg^?1 wet weight in the muscle sample of silver and bighead carp in the experimental enclosure, which suggested that these fish were safe to consume for human. However, the long‐term effects of MCs on aquatic ecosystem and on public health cannot be overlooked.     

Determination of critical limits of soil nutrients for use in optimizing fertilizer rates for fish ponds in red, lateritic soil zones

Mesocosms containing lateritic soil from West Bengal, India and de-ionized water were treated with different amounts of nitrogen (N), phosphorus (P), and potassium (K) fertilizers. Concentrations of available N, P, and K in soils and gross primary productivity were measured for the mesocosms. Critical concentrations of the three nutrients were estimated by the graphical method of Cate and Nelson that has been used widely for establishing critical soil nutrient concentrations in fertilization research on terrestrial crop plants. Critical concentrations of the three nutrients were: available N, 200 mg kg⁻¹; available P, 13 mg kg⁻¹; available K, 80 mg kg⁻¹. Ponds sited in red, lateritic soil in West Bengal were treated monthly with 20, 15, and 3 kg ha⁻¹ application⁻¹ of N, P, and K, respectively. There was an increase in gross primary productivity when the critical concentrations of available N, P, and K established in the mesocosm study were attained in the pond soils. Fish production in fertilized ponds is positively correlated with gross primary productivity. Thus, the concept of critical nutrient concentrations in soils could be a valuable tool in pond fertilization research.     

The culture potential of Paracentrotus lividus (Lamarck 1816) in Greece: a preliminary report

Two hundred and forty adult and sub-adult Paracentrotus lividus were divided into two major groups (adults, sub-adults) based on their diameter differences (above or below 40 mm) and allocated to baskets suspended from the walkways of finfish cages, in South Evoikos Gulf, Central Greece. Each group were fed a commercial finfish diet, while their respective controls remained unfed during the whole duration of the experiment. Total weight and gonad weight of all treatment groups increased while the respective weights of the control groups decreased by the end of the experiment. In addition, gonad indices of the treatment groups increased by the end of the experiment, except for the sub-adult groups. No mortalities were observed in the treatment groups, while the low mortalities in the control groups (1–1.5%) can be attributed to the nutrient status of the surrounding waters and the increased plankton populations and sedentary organisms. It has therefore been shown that fish/sea urchin polyculture in Greece can be low risk and beneficial for the producer system, as long as prevailing water quality and environmental aquaculture standards are kept.     

The efficiency and condition of oysters and macroalgae used as biological filters of shrimp pond effluent

Current shrimp pond management practices generally result in elevated concentrations of nutrients, suspended solids, bacteria and phytoplankton compared with the influent water. Concerns about adverse environmental impacts caused by discharging pond effluent directly into adjacent waterways have prompted the search for cost‐effective methods of effluent treatment. One potential method of effluent treatment is the use of ponds or raceways stocked with plants or animals that act as natural biofilters by removing waste nutrients. In addition to improving effluent water quality prior to discharge, the use of natural biofilters provides a method for capturing otherwise wasted nutrients. This study examined the potential of the native oyster, Saccostrea commercialis (Iredale and Roughley) and macroalgae, Gracilaria edulis (Gmelin) Silva to improve effluent water quality from a commercial Penaeus japonicus (Bate) shrimp farm. A system of raceways was constructed to permit recirculation of the effluent through the oysters to maximize the filtration of bacteria, phytoplankton and total suspended solids. A series of experiments was conducted to test the ability of oysters and macroalgae to improve effluent water quality in a flow‐through system compared with a recirculating system. In the flow‐through system, oysters reduced the concentration of bacteria to 35% of the initial concentration, chlorophyll a to 39%, total particulates (2.28–35.2 µm) to 29%, total nitrogen to 66% and total phosphorus to 56%. Under the recirculating flow regime, the ability of the oysters to improve water quality was significantly enhanced. After four circuits, total bacterial numbers were reduced to 12%, chlorophyll a to 4%, and total suspended solids to 16%. Efforts to increase biofiltration by adding additional layers of oyster trays and macroalgae‐filled mesh bags resulted in fouling of the lower layers causing the death of oysters and senescence of macroalgae. Supplementary laboratory experiments were designed to examine the effects of high effluent concentrations of suspended particulates on the growth and condition of oysters and macroalgae. The results demonstrated that high concentrations of particulates inhibited growth and reduced the condition of oysters and macroalgae. Allowing the effluent to settle before biofiltration improved growth and reduced signs of stress in the oysters and macroalgae. A settling time of 6 h reduced particulates to a level that prevented fouling of the oysters and macroalgae.     

Projections of nutrient supply to a wakame (Undaria pinnatifida) seaweed farm on the Sanriku Coast of Japan

Wakame (Undaria pinnatifida) aquaculture is one of the major fishery industries on the Sanriku Coast of Japan. Low concentrations of nutrients in the autumn, when the germinated wakame sporophytes are installed at aquaculture facilities, induce defective growth, resulting in withering. To determine the characteristics of the nutrient variations in this region, we conducted hydrographic measurements off the coast of Iwate Prefecture. The measured results showed that in the autumn, vertical mixing pumped nutrients from the deeper layers to the mixed layer, where nutrients had been depleted during the summer. Based on these results, we developed a one‐dimensional model incorporating vertical mixing by atmospheric cooling to project the vertical distribution of the nutrient concentrations. By conducting ensemble calculations, we created a probability projection calendar for the nutrient supply up to 50 days ahead. This information was provided to wakame farmers before they installed the germinated sporophytes. The projected probability calculated from the initial conditions on 11 October 2016 was 0% until 25 October and gradually increased to 85% on 30 November. The nitrate concentration detected by an ultraviolet nitrate sensor moored at a wakame farm revealed that the nutrient concentrations reached a critical threshold on 18 November. The projected probability on that date was 75%; therefore, the projection was successful. The observed temporal evolution of the temperature, salinity, and nutrient concentrations was well reproduced by our model. The information obtained using this model is expected to reduce the risk of wakame withering and improve the efficiency of wakame production.     

Selective particle feeding by the Chilean oyster, Ostrea chilensis; implications for nursery culture and broodstock conditioning

Hatchery broodstock conditioning and nursery culture of the Chilean flat oyster Ostrea chilensis have been hampered by the poor performance of oysters fed typical microalgal hatchery diets. To determine the feeding capabilities of this species the selective removal and consumption of natural planktonic assemblages and artificial inert particles (polystyrene beads) by juvenile and adult oysters were examined experimentally. The arrangement of the eulaterofrontal cirri of the ctenidia was also examined to infer their potential efficiency of particle selection for feeding. Polystyrene beads of 45 and 15 μm in diameter had high rates of removal from suspensions by both juvenile (45 μm = 70%, 15 μm = 73%) and adult (45 μm = 88%, 15 μm = 87%) oysters. In contrast, beads of 6 μm diameter had lower rates of removal (adults = 68%, juveniles = 53%), while 1 μm beads were not removed at all. Both adult and juvenile oysters feeding upon natural plankton assemblages removed only microphytoplankton (20-75 μm in size) despite the presence of nanophytoplankton (2-20 μm), picophytoplankton (< 2 μm), cyanobacterium Synechoccocus spp. (< 2 μm), and bacterial cells (< 75 μm) in the experimental suspensions. Eulaterofrontal cirri of both juvenile (15.2 μm ± 0.9 SE) and adult oysters (18.9 μm ± 0.3 SE) are the shortest that have been reported for any ostreid species which helps to explain the inability of this species to retain small food particles. The clearance rates for oysters feeding on microphytoplankton only were 1.49 (± 0.05 SE) and 7.1 (± 1.2 SE) l h^− 1g^− 1 for juveniles and adults respectively. These values are much higher than previously reported for this species being fed smaller sized cultured microalgae. Our results strongly suggest that the difficulties in the nursery and broodstock hatchery culture of this oyster may be due to inappropriate phytoplankton diets. We recommend the provision of cultured microalgae of 20-75 μm in diameter for improving the performance of hatchery maintained juvenile and adult O. chilensis.     

北海营盘新珍珠贝养殖区浮游植物现状调查

于2010-2011年按季节在北海营盘新珍珠贝养殖区海域进行了4次调查采样,对该海域的浮游植物进行了初步研究。结果表明：该海域共有浮游植物36属82 种,春季以细柱藻属占优,夏季以海链藻属占优,秋季以角毛藻属占优,冬季以菱形藻属和海链藻属占优。各季节各站位浮游植物数量变化范围在2.8×104～9.6×104cells/ml,均位于同一数量级,多样性指数范围在2.54～2.94之间,均匀度在0.47～0.55之间,水质处于轻度污染状态,作为规划的珍珠贝养殖区,该海域饵料生物数量偏低,是规划时需要考虑的因素。     

太泊湖浮游生物初步调查及鲢鳙鱼产力评估

2013年5、7、9和11月,对太泊湖浮游生物和水体状况开展了调查,评估了鲢鳙鱼产力。总碱度变化范围为62.92~89.09 mg/L;总氮、总磷及氨氮含量均严重超过地标水环境质量标准的Ⅱ类标准限值。共监测到浮游植物7门51属,平均密度1 593×104个/L,平均生物量20.166 mg/L;绿藻门、蓝藻门、硅藻门以及裸藻门为优势种群,生物量分别占浮游植物总生物量的24.3%、8.9%、16.2%和25.84%。共采集到浮游动物25种,其中原生动物5种、轮虫13种、枝角类5种、桡足类2种。浮游动物平均数量1 771.85个/L,平均生物量2.508 mg/L;枝角类、轮虫、原生动物、桡足类的生物量分别占总生物量的15.9%、78.7%、0.1%和35.2%,浮游动物多样性指数为0.36~0.46,均匀度指数为0.34~0.41。综合判断太泊湖处于富营养化状态。太泊湖鲢鱼产力151.2 kg/hm2,全湖约322 t;鳙鱼产力56.4 kg/hm2,全湖约120 t。     

Effects of common carp Cyprinus carpio (L.) and feed addition in rohu Labeo rohita (Hamilton) ponds on nutrient partitioning among fish, plankton and benthos

The effects of introducing common carp (CC) and of adding artificial feed to fertilized rohu ponds on water quality and nutrient accumulation efficiency were studied. All ponds were stocked with 15 000 rohu ha^?1. Treatments included ponds with rohu alone, rohu plus 5000 common carp ha^?1 and rohu plus 10 000 CC ha^?1. A comparison was also made between supplementally fed and non‐fed ponds. The overall highest nitrogen (N) and phosphorus (P) concentrations were observed in ponds with 5000 CC ha^?1, followed by ponds with 10 000 and 0 CC ha^?1. The largest fractions of N and P inputs accumulating in fish, phytoplankton and zooplankton were observed in ponds with 5000 CC ha^?1, followed by ponds with 10 000 CC ha^?1 and subsequently ponds without CC. Relatively more nutrients accumulated in benthic organisms in ponds without than in ponds with CC. A smaller fraction of the nutrient input was retained in fish, plankton and benthic organisms in ponds without CC compared with ponds with CC. Compared with 5000 CC ha^?1, stocking 10 000 CC ha^?1 can be considered as overstocking, because this leads to lower fish production and relatively less nutrients retained in plankton and benthic organisms.