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1.
The phosphorus cycle in the ecosystem of the shallow, hypertrophic Loosdrecht lakes (The Netherlands) was simulated by means of the dynamic eutrophication model PCLOOS. The model comprises three algal groups, zooplankton, fish, detritus, zoobenthos, sediment detritus and some inorganic phosphorus fractions. All organic compartments are modelled in two elements, carbon and phosphorus. Within the model system, the phosphorus cycle is considered as completely closed. Carbon and phosphorus are described independently, so that the dynamics of the P/C ratios can be modelled. The model has been partly calibrated by a method based on Bayesian statistics combined with a Range Check procedure.Simulations were carried out for Lake Loosdrecht for the periods before and after the restoration measures in 1984, which reduced the external phosphorus loading to the lake from ca. 2 mgP m–2 d–1 to 1 mgP m–2 d–1. The model outcome was largely comparable withthe measured data. Total phosphorus has slowly decreased from an average 130 µgP l–1 to ca. 80 µgP l–1, but chlorophyll-a (ca. 150 µg 1–1, summer-averaged) and seston concentrations (8–15 mgC 1–1) hardly changed since the restoration measures. About two-thirds of the seston consisted of detritus, while the phytoplankton remained dominated by filamentous cyanobacteria. The P/C ratio of the seston decreased from ca. 1.0% to 0.7%, while the P/C ratios of zooplankton, zoobenthos and fish have remained constant and are much higher. The system showed a delayed response to the decreased phosphorus loading until a new equilibrium was reached in ca. five years. Major reasons for the observed resilience of the lake in responding to the load reduction are the high phosphorus assimilation efficiency of the cyanobacteria and the high internal recycling of phosphorus. A further reduction of nutrient loading, perhaps in combination with additional measures like biomanipulation, will be the most fruitful additional restoration measure.  相似文献   

2.
In the nature reserve Botshol, the process of eutrophication is very successfully stopped and reversed by hydrological separation of the agricultural area from the nature reserve and by dephosphorizing the inlet-water. The external phosphate loading of about 0.6 g Pm–2y–1 decreased 80% to about 0.12 g Pm–2y–1. The Dutch governmental program in relation to eutrophication is focussed on the introduction of detergents without phosphates, dephosphorizing the water at sewage purification plants, reduction of fertilization of farming lands and international measures in relation to the discharge of phosphates in the rivers Rhine, Meuse and Scheldt. The effects of the governmental program on the external phosphate loading of Botshol will be a reduction of only 4%.For the average Dutch fresh waters the effect of the governmental program is a decrease of the external loading of about 6 g Pm–2y–1 to about 3 g Pm–2y–1, a phosphate loading still much to high to push back eutrophication noticeably.As an example also for a number of Dutch fresh waters, sensitive to eutrophication, the effects of the governmental program are calculated. For none of these waters the program will lead to an external phosphate loading that even comes close to the recent phosphate loading of Botshol.  相似文献   

3.
Since 1960 the water quality of the nature reserve Botshol has been deteriorated. An increase in the nutrient load caused an escalation in phytoplankton biomass and turbidity problems. This may have been caused by nutrient-rich water flowing into the reserve from the agricultural areas and from the polder of Nellestein. From 1980 to 1985 much research has been undertaken and participation of different disciplines gave the opportunity to analyse the cause of the changes. An integrated restoration plan has been drawn up to reduce the eutrophication. The plan was implemented in 1989 and in order to reverse the external nutrient load the following measures were taken. 1. Execution of a water management plan with the intention to isolate the agricultural areas from the reserve area. 2. Dephosphorization of the inlet water. These restoration measures resulted in a reduction of the external load from 0.6 to 0.1 g P m–2 year–1.  相似文献   

4.
Dokulil  M. T.  Jagsch  A. 《Hydrobiologia》1992,243(1):389-394
Following restoration measures (ring canalization, treatment plant with phosphorus precipitation), phosphorus loading declined step-wise from 26.21 t year–1 to 9.18 t year–1 during the period 1979–1984 while P-retention increased from 48% to 78%. Phosphorus loading was poorely correlated with precipitation. Inorganic nitrogen load, largely NO3– N, did not decline but was significantly correlated with precipitation (r = 0.95) throughout the investigation period (1978–1989).Total phosphorus loading reached acceptable levels in 1984 when compared to critical loading calculated according to Vollenweider (1976). Phosphorus input to the lake has remained at these levels in recent years.Average annual chlorophyll-a concentrations and biovolume of phytoplankton in the top 20 m layer of the lake decreased, in correspondence with the respective phosphorus concentrations, from eutrophic to mesotrophic levels. The decline was accompanied by a drastic reduction of blue-green algal populations, and especially of Oscillatoria rubescens D.C..  相似文献   

5.
As a result of high nutrient loading Lake Veluwe suffered from an almost permanent bloom of the blue-green algaOscillatoria agardhii Gomont. In 1979, the phosphorus loading of the lake was reduced from approx. 3 to 1 g P.m–2.a–1. Moreover, since then the lake has been flushed during winter periods with water low in phosphorus. This measure aimed primarily at interrupting the continuous algal bloom. The results of these measures show a sharp decline of total-phosphorus values from 0.40–0.60 mg P.l–1 (before 1980) to 0.10–0.20 mg P.l–1 (after 1980). Summer values for chlorophylla dropped from 200–400 mg.m–3 to 50–150 mg.m–3.The increase in transparency of the lake water was relatively small, from summer values of 15–25 cm before the implementation of the measures to 25–45 cm afterwards. The disappointing transparency values may be explained by the decreasing chlorophylla and phosphorus content of the algae per unit biovolume. Blue-green algae are gradually loosing ground. In the summer of 1985 green algae and diatoms dominated the phytoplankton for the first time since almost 20 years. To achieve the ultimate water quality objectives (transparency values of more than 100 cm in summer), the phosphorus loading has to be reduced further.  相似文献   

6.
The formerly rich characean community in Botshol with six species of which the rareNitellopsis obtusa andChara hispida dominated at many sites, decreased to only two species,Chara globularis andC. connivens, in the period 1980–1988. The macrophyteNajas marina also remained at some sites, and the aquatic mossFontinalis antipyretica and the filamentous algaVaucheria dichotoma predominated at many sites. These phenomena may have been due to eutrophication by the inlet of polluted water. This process of eutrophication was stopped by restoration measures in 1989, resulting in a lower phosphorus concentration (ca. 0.024 mg l–1) and a higher water transparency. Immediately after these measures the Characeae community increased strongly in abundance and number of species. During the summer of 1990, and especially of 1991, a spectacular growth occurred ofChara connivens. Chara connivens was often accompanied byChara hispida. Other species with scattered occurrence wereChara aculeolata, C. aspera, C. contraria andC. Globularis. The reasons for the shift in dominance fromNitellopsis obtusa toChara connivens are discussed. One of the reasons may be the recent higher chloride content which is one of the consequences of the restoration measures.  相似文献   

7.
Simons  Jan  Ohm  Marieke  Daalder  Remco  Boers  Peter  Rip  Winnie 《Hydrobiologia》1994,(1):243-253
Till about 1965 a rich characean community occurred in the shallow peat lake Botshol with six species of which the rare Nitellopsis obtusa and Chara major dominated at many sites. In the period 1980–1988 characean biomass strongly decreased and only two species, Chara globularis and C. connivens, remained in small populations at a few localities. Of the macrophyte Najas marina also some small populations remained, while the aquatic moss Fontinalis antipyretica and the filamentous alga Vaucheria dichotoma predominated at many sites. These phenomena may have been due to eutrophication by the input of polluted water. This process of impoverishment was stopped by restoration measures in 1989, resulting in a lower phosphorus concentration (ca 25 µg l-1) and a higher water transparency. Immediately after these measures the Characeae community increased in abundance and number of species. During the summer of 1990, and especially 1991, a spectacular growth occurred of Chara connivens. C. connivens was often accompanied by C. major. Other species with scattered occurrence were C. aculeolata, C. aspera, C. contraria and C. globularis. The reasons for the shift in dominance from Nitellopsis obtusa to Chara connivens are discussed. From growth experiments evidence was obtained that neither the recent higher chloride level, nor the lowered phosphate concentration were the main factors for the domination of Chara connivens.  相似文献   

8.
Horppila  Jukka  Kairesalo  Timo 《Hydrobiologia》1992,(1):323-331
Lake Vesijärvi, southern Finland, suffered sewere eutrophication by sewage effluent from the city of Lahti during the 1960's and the early 1970's. The municipal sewage loading was diverted from the lake in 1976 and the lake started to recover. However, in the 1980's blue-green algal blooms increased again and the recovery of the lake faded. Enclosure experiments demonstrated that high roach (Rutilus rutilus) biomass is one of the key factors in the fading recovery of the lake. In this study, the influence of roach and another cyprinid fish species (bleak, Alburnus alburnus) to planktonic algal productivity and biomass in Lake Vesijärvi was examined. Enclosure experiments in the field showed the impacts of planktivorous bleak on water quality; in an enclosure with a density of 1 fish m–2 average daily algal production (1370 mg C m–2) and chlorophyll-a concentration (50–90 µg 1–1) were more than twice that in an enclosure without fish. Laboratory experiments showed that the availability of planktonic food affects the foraging behaviour of roach and consequently the internal nutrient loading from the sediment into the water. Roach caused the highest phosphorus loading and turbidity when there was no zooplanktonic food available in the water. The possible interactions between planktivorous and omnivorous fish species are discussed.  相似文献   

9.
In 1984 the external phosphorus load of the shallow eutrophic Loosdrecht lakes was reduced from 3.3 to 1.0 mg m–2 d–1. The effect of phosphorus release from the sediment on lake restoration was investigated. Diffusive release under aerobic conditions (20 °C) decreased from 1 mg m–2 d–1 in 1984 to 0.3 mg m–2 d–1 in 1990. The generation of inorganic phosphorus due to mineralization during summer equals 3 mg m–2 d–1, which is much higher than the measured rate of diffusive release. Despite that, the phosphorus release is hardly stimulated by anaerobic conditions, which indicates that only a small amount of phosphorus is adsorbed by ferric iron in the top sediment layer. This apparent discrepancy is probably caused by the uptake of inorganic phosphorus uptake during resuspension and the loss of inorganic phosphorus with downward seepage.The estimated removal of phosphorus due to downward seepage of 0.8 mg m–2 d–1 agrees well with the average phosphorus retention in the lake. This indicates that sediment burial and diagenesis are unimportant mechanisms for withdrawing phosphorus from the nutrient cycle.Between 1982 and 1991 the total phosphorus content of the upper 2 cm of the sediment decreased from 0.94 to 0.60 g kg–1 DW. At present, about 20% of total phosphorus in this layer is potentially bioavailable, but largely incorporated in easily degradable organic matter. This pool is much smaller in deeper layers. Based on the estimated and measured rates and pool sizes, the annual average phosphorus cycle in the lakes was modelled to evaluate the effects of various restoration measures. The main predictions of the model are: 1) further reduction of the external load may cause a gradual decrease of the total phosphorus concentration in the lake water; 2) dredging and iron addition, without reduction of the external load, may give a rapid improvement followed by a slow return to the present situation; and 3) reduction of the external load, combined with a cut off of downward seepage will not improve the water quality.  相似文献   

10.
Turbid water, high phosphorus (P)-loading and disappearing Chara communities forced local water authorities to carry out restoration measures in the lakes and marshes of the Botshol The Netherlands. The reduction of the external-P input could be reached by chemical treatment of the brackish suppletion water and by separating the area from nearby agricultural areas. A side effect of these measures was an increase of chloride from 400 mg l−1 to 1000 mg l−1 in the surface water of Botshol. Internal biogeochemical processes were investigated with phytometers and direct measurements of soil nutrient availability in greenhouse experiments. The increased chloride levels were assumed to increase soil pore water P. The first experiment showed higher P in the peat-soils treated with the highest Cl-concentration and an increased leaching of PO4 from the lake-bottom peat-soils. No reaction of the phytometer Epilobium hirsutum was found. In the second experiment the 800 mg l−1 Cl-treatment resulted in significantly higher biomass of Carex acutiformis grown on treated bank soil. N-uptake by the phytometer Carex acutiformis was significantly higher. The available-P and total-P in the bank soil did not show a treatment effect. The two studies showed under similar ‘standardized’ conditions a treatment effect of chloride on the P-availability, resulting in higher PO4 leaching and increased plant nutrient concentrations and biomass. The field study showed higher available-P concentrations in the shore zone than in remote areas. The high chloride levels after restoration impact internal nutrient availability in the Botshol wetlands, on soil loaded with P in recent and historic times.  相似文献   

11.
Lake Sempach, located in the central part of Switzerland, has a surface area of 14 km2, a maximum depth of 87 m and a water residence time of 15 years. Restoration measures to correct historic eutrophication, including artificial mixing and oxygenation of the hypolimnion, were implemented in 1984. By means of the combination of external and internal load reductions, total phosphorus concentrations decreased in the period 1984–2000 from 160 to 42 mg P m–3. Starting from 1997, hypolimnion oxygenation with pure oxygen was replaced by aeration with fine air bubbles. The reaction of the plankton has been investigated as part of a long-term monitoring program. Taxa numbers, evenness and biodiversity of phytoplankton increased significantly during the last 15 years, concomitant with a marked decline of phosphorus concentration in the lake. Seasonal development of phytoplankton seems to be strongly influenced by the artificial mixing during winter and spring and by changes of the trophic state. Dominance of nitrogen fixing cyanobacteria (Aphanizomenon sp.), causing a severe fish kill in 1984, has been correlated with lower N/P-ratio in the epilimnion. Buoyant algae such as Planktothrix rubescens (syn. Oscillatoria rubescens) increased in abundance due to enlargement of the trophogenic layer and extended mixing depth during winter. The interactions between zoo- and phytoplankton seemed to be depressed as a result of restoration measures. Zooplankton composition changed to more carnivorous and less herbivorous species. Oxygenation of the hypolimnion induced bioturbation of sediments, mainly by oligochaetae worms, and stimulated germination of spores and cysts and hatching of resting eggs.  相似文献   

12.
The abundance of Gloeotrichia echinulata colonies in the sediments of Lake Erken and their phosphorus content were investigated to determine the contribution of Gloeotrichia colonies to total sediment phosphorus. Moreover, the potential size of the algal inoculum and the migration to the water during summer were estimated.The surplus phosphorus content of the resting colonies in the sediment was about 45% of total phosphorus, which maximized at 8.5 µg P (mg dw)–1 or 81 ng P colony–1. The C:P ratio (by weight) in the early colonies appearing in the lake water was 50:1, while the ratio stabilized at 150 during the major migration period. The internal supply of surplus phosphorus was used during the pelagic growth of the colonies.The internal phosphorus loading to the epilimnion of Lake Erken due to Gloeotrichia migration could, from the measurements of the increase in particulate epilimnetic phosphorus, be estimated at 40 mg P m –2 or 2.5 mg P m–2 d–1 in late July and early August. Determination of the number of colonies in the sediment before and during the migration verified this value to be a conservative estimate of the internal phosphorus loading due to Gloeotrichia migration to the epilimnion in Lake Erken.The sediment P content calculated from the P concentration in early epilimnion colonies resulted in a value of 35 µg P (g dw)–1 as a maximum. This corresponds to only 3% of the total phosphorus content in Lake Erken sediment.  相似文献   

13.
Kitaka  Nzula  Harper  David M.  Mavuti  Kenneth M. 《Hydrobiologia》2002,488(1-3):73-80
The main river supplying Lake Naivasha, Kenya, the Malewa, drains a catchment given over to largely subsistence cultivation and animal husbandry. The lake itself is the focus for an intensive horticultural industry based upon irrigation from the lake. The Malewa, however, is relatively independent of the impact of industry, and so its contribution to eutrophication of the lake was evaluated. Two periods of study, a very wet-dry and a `normal' wet-dry season showed that the river contribution of phosphorus led to a total phosphorus loading of 1.4 g m–2 lake surface ann–1 in the very wet period compared to 0.2 in the `normal'. Chlorophyll `a' in the open water of the lake was significantly related to soluble reactive phosphorus. The lake is now eutrophic by normal limnological criteria.  相似文献   

14.
The paper summarizes the results of a ten-year (1981–1991) zooplankton research on the Lake Loosdrecht, a highly eutrophic lake. The main cause of the lake's eutrophication and deteriorating water quality was supply up to mid 1984 of water from the River Vecht. This supply was replaced by dephosphorized water from the Amsterdam-Rhine Canal in 1984. The effects of this and other restoration measures on the lake's ecosystem were studied. Despite a reduction in the external P-load from ca. 1.0 g P m–2 y–1 to ca. 0.35 g m–2 y–1 now, the filamentous prokaryotes, including cyanobacteria and Prochlorothrix, continue to dominate the phytoplankton.Among the crustacean plankton Bosmina spp, Chydorus sp. and three species of cyclopoid copepods and their nauplii are quite common. Though there was no major change in the composition of abundant species, Daphnia cucullata, which is the only daphnid in these lakes, became virtually extinct since 1989. Among about 20 genera and 40 species of rotifers the important ones are: Anuraeopsis fissa, Keratella cochlearis, Filinia longiseta and Polyarthra. The rotifers usually peak in mid-summer following the crustacean peak in spring. The mean annual densities of crustaceans decreased during 1988–1991. Whereas seston (< 150 µm) mean mass in the lake increased since 1983 by 20–60%, zooplankton (> 150 µm) mass decreased by 15–35%.The grazing by crustacean community, which was attributable mainly to Bosmina, had mean rates between 10 and 25% d–1. Between 42 and 47% of the food ingested was assimilated. In spring and early summer when both rotifers and crustaceans have their maximal densities the clearance rates of the rotifers were much higher. Based on C/P ratios, the zooplankton (> 150 µm) mass contained 2.5 times more phosphorus than seston (< 150 µm) mass so that the zooplankton comprised 12.5 % of the total-P in total particulate matter in the open water, compared with only 4.5% of the total particulate C. The mean excretion rates of P by zooplankton varied narrowly between 1.5 and 1.8 µg P 1 d–1, which equalled between 14 and 28% d–1 of the P needed for phytoplankton production.The lack of response to restoration measures cannot be ascribed to one single factor. Apparently, the external P-loading is still not low enough and internal P-loading, though low, may be still high enough to sustain high seston levels. Intensive predation by bream is perhaps more important than food quality (high concentrations of filamentous cyanobacteria) in depressing the development of large-bodied zooplankton grazers, e.g. Daphnia. This may also contribute to resistance of the lake's ecosystem to respond to rehabilitation measures.  相似文献   

15.
Human activity has been the cause of continuing decline of water quality in most Dutch lakes. Development of lake restoration programmes must take into account the lake functions. Major reduction of the nutrient and pollutant loading is the primary step in lake restoration. Still, the recovery of eutrophic lakes is retarded frequently because of internal phosphorus loading by the lakes' sediments. Sediment dredging, as an additional tool for water quality management to accelerate accomplishing the desired water quality, is studied. In this paper we evaluate the preliminary results of eight lake restoration projects in the Netherlands. The lakes are compared in order to estimate the magnitude of the internal phosphorus loading. Dredging as an additional measure was carried out twice in the peatlake Geerplas. In the Nieuwkoop Lakes only the external phosphorus loading was substantially reduced from 0.9 to 0.2 g P m–2 y–1. Provisional results of these two shallow peatlake restoration projects focussed on eutrophication abatement with and without dredging, are presented. Both show a decrease in phosphorus concentration in the lakes. The necessity to dredge the lakes is discussed.  相似文献   

16.
Synopsis Predation and contaminants are two possible factors in the poor recruitment of young lake charr Salvelinus namaycush in the Great Lakes. We measured the feeding rate of slimy sculpins Cottus cognatus and burbot Lota lota on young lake charr (uncontaminated young from eggs of a hatchery brood stock and contaminated young from eggs of Lake Michigan lake charr) in laboratory test chambers with a cobble substrate. The median daily consumption rate of sculpins for all tests was 2 lake charr eggs (N = 22 tests; 95% confidence interval, 0–13) and 2 lake charr free embryos (N = 31 tests; 95% confidence interval, 0–10). Feeding rate did not differ between hatchery and contaminated prey. Slimy sculpins continued to feed on lake charr when another prey organism, the deepwater amphipod Pontoporeia hoyi, was present. Feeding by burbot on free embryos (4–36 d–1) increased as the mobility of young increased, but burbot consumed about 10% of their body weight weekly in free-swimming young (140–380 d–1). Predation on lake charr eggs by sculpins could be considerable over the 100 to 140 d incubation period, and burbot could eat large numbers of free-swimming lake charr as the young fish left the reef. Predation pressure on young lake charr may inhibit rehabilitation of self-sustaining populations of lake charr on some reefs unless a critical egg density has been reached.  相似文献   

17.
The aims of this study were to document the mainly chemical behaviour of two linked artificial lakes used for both stormwater management and recreation in the new town of Craigavon. Further, the understanding of their behaviour should help in their management and the design of other similar lakes.The lake mean total phosphorus (73 µg P l–1), nitrate (0.50 mg N l–1) and chlorophyll a (25 µg l–1) concentrations, Secchi depth (1.2 m) and the estimated total phosphorus loading (1.98 g m–2 a–1) all classify the main lake as eutrophic. An important source of the phosphorus load on the lakes is the urban area of Craigavon (52% of the total load). The interrelationships between total phosphorus, chlorophyll a and Secchi depth in the main lake are similar to those in natural ones. In addition, the lake follows the total phosphorus load — trophic state relationships (lake total phosphorus and chlorophyll a concentrations and Secchi depth) found to apply elsewhere. These two points indicate that the artificial lakes in Craigavon behave similarly to natural ones.  相似文献   

18.
Relationships between chlorophyll a content of the water, the shoreline-length: water area ratio and the annual total fish yield as catch per unit effort (CUE: kg ha–1 100 h–1 as annual mean values) have been calculated by multivariable regression. The determination coefficient (r 2 = 0.913) showed a significant dependence of fish yield on morphometry of different lake areas. Accordingly, fish carrying capacity of the open water areas, calculated from chlorophyll a content and S/A, ranged from 12 to 34%, but that of the littoral zone between 66 and 88%. These findings have also been supported by echo-sounding records of the horizontal distribution of fish.Bream (Abramis brama L.) contributes the majority (70–80%) of fish stock and yield. Its food mainly consists of zooplankton and benthic invertebrates in ratios that are widely variable with season and depend on the age of fish. Average daily food consumption of individuals (age group 3 + and over) varies between 2 and 5 g. Bream consumes two- to three-times more food in the SW basin than in the NE one. This means that the present stocks inhabiting areas from NE to SW consume annually 13249–20085 t yr–1 of food. According to estimated calorific values, the annual energy consumption of local populations along the longitudinal axis of the lake varies between 93 and 141 kJ m–2 yr–1. The efficiency of energy transfer from primary producers to fish is low and varies from 0.04 to 0.1%.  相似文献   

19.
The relationship between total phosphorus and chlorophyll a concentration was determined for Skinner Lake, Indiana over an annual cycle in 1978–79. Total nitrogen:total phosphorus ratios in the epilimnion ranged from 19 to 220 suggesting a phosphorus-dependent algal yield in the epilimnion. Approximately 90% of annual TP loading reached the lake via streamflow, and 93% of this entered during snowmelt and spring-overturn periods. At that time incoming water flushed the lake 2.4 times. Atmospheric loading accounted for 1.4% of annual TP load. Internal hypolimnetic TP loading occurred during summer stratification. Mean [chl a] for the ice-free period was 15.15 mg m–3, within the range expected for eutrophic lakes.The 1978–79 data were used in conjuction with the Vollenweider & Kerekes (1980) model to produce a model specific for the Skinner Lake system. The model predicted mean epilimnetic total phosphorus and chlorophyll a concentrations from mean total phosphorus concentration in inlet streams and from lake water residence time during the period of spring overturn and summer stratification. The Skinner-specific model was tested in 1982 and it closely predicted observed mean epilimnetic [TP] and [chl a] during the ice-free period. This study shows that variability in lake models which average data over an annual period can be reduced by considering lake-specific seasonal variation in hydrology and external TP loading.  相似文献   

20.
S. mossambicus was exposed to toxic and sublethal concentrations of the fertilizer diammonium phosphate (0.2 to 1.0 g l–1). Mortality, food utilization and growth were studied. At a concentration of 0.6 g l–1 DAP, 100% mortality was observed within 96 h; no mortality occurred at 0.5 g l–1; LC50 was 0.55 g l–1. Rearing the fish in increasing sublethal concentrations of DAP, it was found that the feeding rate decreased from 25.4 mg g–1 fish–1 d–1 (fish reared in DAP-free water) to 10.1 mg g–1 d–1 at the highest sublethal concentration (0.5 g l–1). Growth rate was drastically reduced. At high sublethal concentrations of DAP, the fish lost reserve energy, in addition to the energy obtained from food intake for survival, as a result of increased swimming activity and opercular beats.  相似文献   

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