A Review of Nontraditional Biomanipulation

The aim of this review is to identify problems, find general patterns, and extract recommendations for successful management using nontraditional biomanipulation to improve water quality. There are many obstacles that prevent traditional biomanipulation from achieving expectations: expending largely to remove planktivorous fish, reduction of external and internal phosphorus, and macrophyte re-establishment. Grazing pressure from large zooplankton is decoupled in hypereutrophic waters where cyanobacterial blooms flourish. The original idea of biomanipulation (increased zooplankton grazing rate as a tool for controlling nuisance algae) is not the only means of controlling nuisance algae via biotic manipulations. Stocking phytoplanktivorous fish may be considered to be a nontraditional method; however, it can be an effective management tool to control nuisance algal blooms in tropical lakes that are highly productive and unmanageable to reduce nutrient concentrations to low levels. Although small enclosures increase spatial overlap between predators and prey, leading to overestimates of the impact of predation, microcosm and whole-lake experiments have revealed similar community responses to major factors that regulate lake communities, such as nutrients and planktivorous fish. Both enclosure experiments and large-scale observations revealed that the initial phytoplankton community composition greatly impacted the success of biomanipulation. Long-term observations in Lake Donghu and Lake Qiandaohu have documented that silver carp (Hypophthalmichthys molitrix) and bighead carp (H. nobilis) (two filter-feeding planktivorous species commonly used in management) can suppress Microcystis blooms efficiently. The introduction of silver and bighead carp could be an effective management technique in eutrophic systems that lack macrozooplankton. We confirmed that nontraditional biomanipulation is only appropriate if the primary aim is to reduce nuisance blooms of large algal species, which cannot be controlled effectively by large herbivorous zooplankton. Alternatively, this type of biomanipulation did not work efficiently in less eutrophic systems where nanophytoplankton dominated.

Present status and changes of the phytoplankton community after invasion of Neosalanx taihuensis since 1982 in a deep oligotrophic plateau lake, Lake Fuxian in the subtropical China

Phytoplankton assemblages in the subtrophical oligotrophic Lake Fuxian, the second deepest lake in China, were investigated monthly from September 2002 to August 2003. A total of 113 species belonging to seven phyla were identified, among them, a filamentous green alga, Mougeotia sp., dominated almost throughout the study period and comprised most of the total phytoplankton biomass. Mougeotia sp. has made a substantial development during the past decades: it was absent in 1957, only occasionally present in 1983, increased substantially in 1993, and became predominant in 2002-2003. It is likely that natural invasion of the Taihu Lake noodlefish (Neosalanx taihuensis) has led to a change of dominant herbivorous zooplankton from small to large calanoid, which has increased grazing pressure on small edible algae, and thus has indirectly favored the development of the inedible filamentous Mougeotia sp.

Enhancement of planktonic rotifers by Microcystis aeruginosa blooms: An enclosure experiment in a shallow eutrophic lake

Enclosure experiments with three treatments (sediment addition, sediment + nitrogen enrichment, sediment + phosphorus enrichment) and unfertilized controls were performed in shallow hypereutrophic Lake Donghu during the summer of 2000. Dense Microcystis aeruginosa blooms occurred in all the enclosures during the experimental period but not in the surrounding lake water. Generally, the dominant rotifers were Polyarthra vulgalis, Filinia longiseta, Proales sp. and Asplanchna sp. at the beginning of the experiment, followed by a shift to Brachionus calyciflorus, Trichocerca similis, Cephalodella catellina and Anuraeopsis fissa, and finally to F. longiseta, Proales sp. and Keratella cochleris. M. aeruginosa blooms strongly suppressed the larger Diaphanosoma brachyurum but enhanced the development of the smaller cladocerans and rotifers that probably efficiently utilized organic matter from M. aeruginosa through the detritus food chain. The smaller cladoceran and rotifers coexisted successfully throughout the experimental period.

Summertime ciliate and copepod nauplii distributions and micro-zooplankton herbivorous activity in the Laizhou Bay, Bohai Sea, China

The abundance and biomass of ciliated protozoa and copepod nauplii were investigated at 21 grid stations and two anchored stations in the Laizhou Bay, Bohai Sea, China in June 1998. Dilution incubations were carried out to investigate micro-zooplankton grazing pressure at the anchored stations during spring tide and neap tide. The dominant species were Tintinnopsis amoyensis, T. chinglanensis, T. pallida and aloricate ciliates. A total of 13 species of tintinnids were found. The total abundance of ciliates and nauplii ranged from 30 to 2390 ind l(-1) at grid stations. Tintinnopsis amoyensis was the only ciliate found at the anchored stations and in concentrations which varied from 0 to 6700 ind l(-1). The spatial distribution of ciliates was patchy. Tintinnopsis amoyensis and T. pallida were distributed in the Weihe River mouth and Xiaoqinghe River mouth respectively. The aloricate ciliates, T. chinglanensis and Codonellopsis ostenfeldi dominated offshore in sequence. The water mixing process may affect the spatial pattern of the dominant ciliate species. The abundance and biomass of copepod nauplii were in the range of 0-140 ind l(-1) and 0-7 mu g C l(-1) respectively, with the peak appearing at grid station 15. The total biomass of ciliates and copepod nauplii was in the range of 1(.)5-25 mu g C l(-1). Water column biomass of ciliates and nauplii varied from 2(.)37 to 52(.)3 mg C m(-2). At the anchored stations, the phytoplankton growth rates ranged from undetectable to 0 21 d(-1) and micro-zooplankton grazing rates from 0 13 to 0(.)57 d(-1). The grazing pressure of micro-zooplankton were 12 to 43% of the chlorophyll standing stock and 84 to 267% of the chlorophyll (C) 2000 Academic Press.

Impacts of a small dam on riverine zooplankton

In order to explore the temporal impacts of a small dam on riverine zooplankton, monthly samples were conducted from November 2005 to June 2006 in a reach of Xiangxi River, China, which is affected by a small hydropower plant. A total of 56 taxa of zooplankton were recorded during the study and rotifers were the most abundant group, accounting for 97% of total taxa, while the others were copepod nauplii and copepod adults. This study indicated that: (1) the small dam in the Xiangxi River study area created distinct physical and ecological conditions relative to free-flowing lotic reaches despite the constrained channel and small size of the dam; (2) the existence of the plant's small dam had a significant effect on the zooplankton community. In long periods of drought or dry seasons the effect of the dam on potamoplankton was more pronounced (e.g., November, February, March, and May). But the downfall or the connectivity of channel appeared to decrease the effect of small hydropower plants on riverine zooplankton (e.g., April). The present observation underscores the need for additional studies that provide more basic data on riverine zooplankton communities and quantify ecological responses to dam construction over longer time spans.

Short-term effects of drawing water for connectivity of rivers and lakes on zooplankton community structure

During 28-29, September 2005, water was drawn from Hanjiang River and Houguan Lake to the Yangzi River via Sanjiao Lake and Nantaizi Lake in Wuhan in order to provide favorable conditions for ecosystem restoration. To evaluate the feasibility and validity of drawing water as a means of ecosystem restoration, zooplankton populations were studied 3 times (before, immediately after finishing and a month after drawing water) at seven locations from 27 Sept. 2005 to 2 Nov. 2005. Water quality in the lakes was mostly improved and zooplankton species richness decreased as soon as drawing water had finished but increased a month after drawing water. Zooplankton density and biomass was reduced in the lakes by drawing water but was increased at the entrance to Sanjiao Lake because of landform geometry change. Before drawing water, most species in Sanjiao Lake e.g., Brachionus sp. and Keratella sp. were tolerant of contamination. After drawing water oligotrophic-prone species such as Lecane ludwigii and Gastropus stylifer emerged. We conclude that drawing water could be important for improving water quality and favour ecosystem restoration. Dilution of nutrient concentrations may be an important role in the effect.

Field and experimental studies on the combined impacts of cyanobacterial blooms and small algae on crustacean zooplankton in a large, eutrophic, subtropical, Chinese lake

Field and experimental studies were conducted to evaluate the combined impacts of cyanobacterial blooms and small algae on seasonal and long-term changes in the abundance and community structure of crustacean zooplankton in a large, eutrophic, Chinese lake, Lake Chaohu. Seasonal changes of the crustacean zooplankton from 22 sampling stations were investigated during September 2002 and August 2003, and 23 species belonging to 20 genera were recorded. Daphnia spp. dominated in spring but disappeared in mid-summer, while Bosmina coregoni and Ceriodaphnia cornuta dominated in summer and autumn. Both maximum cladoceran density (310 ind. l(-1)) and biomass (5.2 mg l(-1)) appeared in autumn. Limnoithona sinensis, Sinocalanus dorrii and Schmackeria inopinus were the main species of copepods. Microcystis spp. were the dominant phytoplankton species and formed dense blooms in the warm seasons. In the laboratory, inhibitory effects of small colonial Microcystis on growth and reproduction of Daphnia carinata were more remarkable than those of large ones, and population size of D. carinata was negatively correlated with density of fresh large colonial Microcystis within a density range of 0-100 mg l(-1) (r = -0.82, P < 0.05). Both field and experimental results suggested that seasonal and long-term changes in the community structure of crustacean zooplankton in the lake were shaped by cyanobacterial blooms and biomass of the small algae, respectively, i.e., colonial and filamentous cyanobacteria contributed to the summer replacement of dominant crustacean zooplankton from large Daphnia spp. to small B. coregoni and C. cornuta, while increased small algae might be responsible for the increased abundance of crustacean zooplankton during the past decades.

Crustacean zooplankton distribution patterns and their biomass as related to trophic indicators of 29 shallow subtropical lakes

A comparative limnological study was carried out to present a snapshot of crustacean zooplankton communities and their relations to environmental factors to test whether there is a consistent relationship between crustacean biomass and trophic indicators among lake groups with similar trophic conditions. The study lakes showed a wide range of trophic status, with total phosphorus (TP) ranging from 0.008 to 1.448mgL(-1), and chlorophyll a from 0.7 to 146.1 mu g L-1, respectively. About 38 species of Crustacea were found, of which Cladocera were represented by 25 taxa (20 genera), and Copepoda by 13 taxa (I I genera). The most common and dominant species were Bosmina coregoni, Moina micrura, Diaphanosoma brachyurum, Cyclops vicinus, Thermocyclops taihokuensis, Mesocyclops notius and Sinocalanus dorrii. Daphnia was rare in abundance. Canonical correspondence analysis showed that except for four species (D. hyalina, S. dorrii, C. vicinus and M. micrura), almost all the dominant species had the same preference for environmental factors. Temperature, predatory cyclopoids and planktivorous fishes seem to be the key factors determining species distribution. TP was a relatively better trophic indicator than chlorophyll a to predict crustacean biomass. Within the three groups of lakes, however, there was no consistent relationship between crustacean biomass and trophic indicators. The possible reason might be that top-down and bottom-up control on crustaceans vary with lake trophic state. The lack of significant negative correlation between crustacean biomass and chlorophyll a suggests that there was little control of phytoplankton biomass by macrozooplankton in these shallow subtropical lakes. (c) 2007 Elsevier GmbH. All rights reserved.

Phytoplankton biomass in relation to nutrients and zooplankton in thirty subtropical lakes adjacent to the Yangtze River, China

A total of 30 shallow lakes, located along the middle and lower reaches of the Yangtze River, were studied to assess the relative importance of nutrients and zooplankton biomass in determining the phytoplankton biomass in subtropical China. Zooplankton biomass and nutrients both varied greatly in these lakes. Factor analysis and multiple linear regression showed that phytoplankton biomass was positively correlated with TN, NH4+, NO3- and TP, while it did not show any negative relationship to zooplankton biomass. Meanwhile, the phytoplankton biomass showed contrary relationships to the mass ratio of TN/TP in spring and summer, suggesting that in nutrient-richer lakes the dominant phytoplankton species have different preferences for TN/TP ratio. The insignificant top-down control of phytoplankton biomass may be attributed to the dominance of small-sized crustaceans and low crustacean biomass resulting from cyanobacterial dominance and planktivorous fish predation as well as other factors. Thus, it is likely that nutrients were more important than zooplankton biomass in explaining the total variance of phytoplaDkton biomass in these subtropical lakes.

Crustacean zooplankton size structure in aquaculture lakes: is larger size structure always associated with higher grazing pressure?

Crustacean zooplankton size structure in 27 aquaculture lakes was studied to test the hypothesis that larger size structure is associated with higher grazing pressure. Mean body length of crustaceans was positively correlated with increasing Chl a (r(2) = 0.40, P = 0.000) and TP (r(2) = 0.38, P = 0.000), contrary to the empirical studies. However, the ratio of zooplankton to phytoplankton biomass decreased significantly with increasing TP (r(2) = 0.27, P = 0.005) and mean body length (r(2) = 0.46, P = 0.000). Meanwhile, size structure showed no significant effect in explaining residual variations of phosphorus-chlorophyll relationship (P = 0.231). These results indicate that larger size structure was not always associated with higher zooplankton grazing pressure. It is likely that in aquaculture lakes crustacean zooplankton size structure was of minor importance in control of phytoplankton biomass, and it was mainly regulated by fish predation. The results showed in our study and the empirical studies might be a reflection of two different stages of lake eutrophication and fish predation intensity.

Effects of fish stocking on the zooplankton community structure in a shallow lake in China

Changes in the zooplankton community structure in relation to fishery practices in Lake Donghu, Wuhan, China were examined. The number of Protozoa species increased slightly, whereas the number of rotifers and crustaceans decreased from the 1960s to the 1990s. The total annual average densities of zooplankton increased 15-20 times in the 1990s compared with the 1960s. This increase was largely attributed to Protozoa, which contributed 93.4% by number of the total zooplankton density in 1991. Cladoceran densities decreased markedly from 1987. Changes in densities of rotifers and copepods were not evident. Trends in zooplankton biomass were similar to density. Large changes in zooplankton community structure coincided with markedly changes in concentration of chlorophyll a and transparency in Lake Donghu in 1987. The year 1987 seems to be the threshold year when the zooplankton community structure changed considerably. These changes were related to continuously increasing fish stock biomass in the lake. It was suggested that fish stocking and fish biomass should be a better managed for improvement of the quality of the lake's environment.

The rotifer assemblage in a shallow, eutrophic Chinese lake and its relationships with cyanobacterial blooms and crustacean zooplankton

Rotifer assemblage in the subtropical eutrophic Lake Chaohu was investigated monthly from September 2002 to August 2003. Forty-nine species belonging to 18 genera and 14 families were recorded. The highest densities of rotifer were observed during summer when there were heavy cyanobacterial blooms. There was a significant positive correlation between total rotifer density and the biomass of cyanobacteria. However, no correlations were found between the densities of rotifer and crustacean zooplankton, possibly owing to the paucity of large-bodied planktonic crustaceans. It is likely that the occurrence of cyanobacterial blooms not only caused the shift of dominant crustacean zooplankton from large species to smaller ones but also weakened the negative interaction between crustaceans and rotifers.

Impacts of filter-feeding fishes on the long-term changes of crustacean zooplankton in a eutrophic subtropical Chinese lake

Long-term changes In the crustacean zooplankton community (calanoid and cyclopoid copepods and cladocerans) were studied in Lake Donghu, a shallow and eutrophic Chinese lake. This lake had been earlier stocked with two pump Alter-feeding Ashes, silver carp (Hypopthalmichthys molitrix) and bighead carp (Aristichthys nobilis). During the 1950s and the mid-1980s, the ratio of copepods to cladocerans was relatively stable but showed a general increase thereafter. From the early-1980s to the 1990s, calanoid/cyclopoid ratios decreased obviously. In the 1990s, Cyclops vicinus, Diaphanosoma brachyurum, and Moina micrura were dominant the abundance of C. vicinus and M. micrura increased significantly; and D, brachyurum showed a substantial decrease. The study shows that under extremely high pressure of Ash predation, the species which could recover rapidly from fish predation would be the most likely to survive and increase their numbers.

Long-term changes of Copepoda community (1957-1996) in a subtropical Chinese lake stocked densely with planktivorous filter-feeding silver and bighead carp

In contrast to the relatively well documented impact of particulate-feeding fish on zooplankton communities, little attention has been devoted to the impact of filter-feeding fish. Filter-feeding silver and bighead carp are the most intensively cultured fish species in Asia and comprise much of the production of Chinese aquaculture. However, little information is known about the impact of either fish on the zooplankton community. Long-term changes in the Copepoda community (1957-1996) were studied at two sampling stations of a subtropical Chinese lake (Lake Donghu) dominated by silver and bighead carp. For both calanoids and cyclopoids, the littoral station (I) was much more resource profitable than the pelagic station (II). There has been a tremendous increase in the annual fish catch over the past 30 years due to the increased stocking with fingerlings of the two carp species. There was a notably higher fish density at Station I than at Station II. Cyclopoid abundance was notably higher at Station I than at Station II during the 1950s to the 1980s, while the reverse became true in the 1990s. This is probably because when fish abundance increased to an extremely high level, the impact of fish predation on the cyclopoids became more important than that of food resources at the littoral station. At both stations, cyclopoid abundance was relatively low in spite of the presence of abundant prey. Similarly, calanoid density did not differ significantly between the two stations in the 1950s and 1960s, but was significantly lower at Station I than at Station II during the 1980s and 1990s. Such changes are attributed to the gradient of fish predation between the stations and an increasing predation pressure by the fish. The increased fish predation also correlated with a shift in summer-dominant calanoids from larger species to smaller ones. In conclusion, the predaceous cyclopoids are affected by fish predation to a much lesser extent than the herbivorous calanoids, and therefore increased predation by filter-feeding fish results in a definite increase in the cyclopoid/calanoid ratio. Predation by filter-feeding fish has been a driving force in shaping the copepod community structure of Lake Donghu during the past decades.

Changes in the plankton community following introduction of filter-feeding planktivorous fish

1. We conducted enclosure experiments in a shallow eutrophic lake, in which a biomass gradient of the filter-feeding planktivore, silver carp, Hypophthalmichthys molitrix Valenciennes, was created, and subsequent community changes in both zooplankton and phytoplankton were examined. 2. During a summer experiment, a bloom of Anabaena flos-aquae developed (approximate to 8000 cells mL(-1)) solely in an enclosure without silver carp. Concurrent with, or slightly preceding the Anabaena bloom, the number of rotifer species and their abundance increased from seven to twelve species (1700-14 400 organisms L-1) after the bloom in this fish-free enclosure. Protozoans and bacteria were generally insensitive to the gradient of silver carp biomass. 3. During an autumn experiment, on the other hand, large herbivorous crustaceans were more efficient than silver carp in suppressing the algae, partly because the lower water temperature (approximate to 24 degrees C) inhibited active feeding of this warm-water fish and also formation of algal colonies. Heterotrophic nanoflagellate and bacterial densities were also influenced negatively by the crustaceans. 4. Correspondence analysis (CA) was applied to the weekly community data of zooplankton and phytoplankton. A major effect detected in the zooplankton community was the presence/absence of silver carp rather than the biomass of silver carp, whereas that in the phytoplankton community was the fish biomass before the Anabaena bloom, but shifted to the presence/absence of the fish after the bloom.