Effect of competition on the demography of planktonic cladocerans — Daphnia and Diaphanosoma

Summary Isolated and mixed continuous cultures of Daphnia hyalina and Diaphanosoma brachyurum in lake water were maintained under laboratory conditions to elucidate demographic effects of competition. Population dynamics curves were obtained. Interspecific competition was revealed by the decrease in the average density of animals in the mixed versus isolated cultures and by the extinction of one species in the presence of the other. Within the first 50 days either Diaphanosoma (4 cases) or Daphnia (1 case) was the superior competitor, depending on the initial conditions. Further cultivation resulted in the extinction of Diaphanosoma in the mixed cultures. There were no statistically significant differences between the maximum rates of population increase (r _m ) in Daphnia and Diaphanosoma at the concentration of edible algae about 2·10⁵ m³ml^-1(0.293 and 0.286 days^-1, respectively). Time lags for density-dependent parameters of the populations were evaluated by means of rank cross-correlations. Regardless of the species identity the time lags of fecundity, birth rate, and the rate of population growth were significantly higher in the superior competitor. The initial conditions of culturing affected the time lags which in their turn influenced the outcome of the interaction. Enhanced competitive ability due to the maximized time lags in Daphnia was not associated with the loss of population stability. Conversely, it brought about destabilization of Diaphanosoma populations which seemed to be the ultimate cause of its extinction observed in the end of the experiment. Time lag of the population growth rate was well predicted based on the half-sum of time lags in birth and death rates (r²=0.80, P<0.001). Daphnia responded to competition with a sharp shortening of the time lags of fecundity, birth rate, and the population growth rate. It increased clutch size and showed inverse relationship between the fecundity time lag and average fecundity even though it was strongly suppressed by Diaphanosoma. The competitive ability was not related to the percentage of adults in the populations. In contrast to the current belief the major result of interspecific competition in the experiment was not a decrease in the rate of population growth but was a reduction in population time lags.     

Characterization of the bacterial flora of mass cultivated Brachionus plicatilis

Bacterial density and composition in association of mass cultivated rotifers (Brachionus plicatilis, SINTEF-strain) was investigated, during experimental conditions identical to the procedures used for preparing rotifers as live food for marine cold water fish larvae. These procedures include cultivation, enrichment with squid meal and acclimation to low temperature by storage of the rotifer culture at 6 °C. Large variations were observed in the number of rotifer associated (1.8–7.6 · 10³ colony forming units per rotifer^–1) and free-living (0.6–25 10⁷ cells·ml^–1) bacteria. An increase of 50–150% in the bacterial number was normally observed after feeding the rotifer with squid meal, but after three days of acclimation at 6 °C, the bacterial numbers decreased to the initial level.After enrichment of the cultures with squid meal, the similarity in the composition of the bacterial flora between the rotifers and water was reduced. However, acclimation of the culture at 6 °C resulted in better agreement of the rotifer associated flora and that in water. Enrichment of the cultures induced a shift in the bacterial composition from Cytophaga/Flavobacterium dominance to Pseudomonas/Alcaligenes dominance. The bacterial flora of the rotifer cultures are dominated by presumably opportunistic species after enrichment, which may have detrimental effects when rotifers are fed as live food to marine fish larvae.     

Use of swimming speed and egg ratio as predictors of the status of rotifer cultures in aquaculture

This study evaluated the use of egg ratio (eggs rotifer^–1) and swimming speed (mm min^–1) as prediction criteria for production and culture quality in mass cultures of the rotifer Brachionus plicatilis. Egg ratio was determined to be a suitable predictor of rotifer growth and production in the cultures. Low egg ratios (i.e., 0–0.17 eggs rotifer^–1) indicate reduced rotifer population over time (i.e., negative net population growth rates). However, at this time egg ratio dynamics are not suitably understood to predict in advance a sudden population collapse.Swimming speed of reproductive, egg-carrying females in the exponential growth phase was 40–45 mm min^–1. During exponential growth swimming speed was independent of the food used. Lower swimming speeds were obtained in late stationary phase (10–25 mm min^–1) when yeast was used as a food source. Both environmental factors (e.g., accumulating metabolites) and changes in nutritional state of the rotifers may have affected the swimming speed, but environmental factors appear to be the most important. We believe that swimming speed has the potential of becoming an accurate predictor of culture quality in mass cultures of rotifers.     

High density culture of the freshwater rotifer,Brachionus calyciflorus

The freshwater rotifer, Brachionus calyciflorus is one of the live food organisms used for the mass production of larval fish. In this study possibility of obtaining high density cultures of the freshwater rotifer B. calyciflorus were investigated. The two culture systems used differed in their air and dissolved oxygen supplies using three temperatures in each case: 24, 28 and 32 °C. Rotifers were batch-cultured using 5 l-vessels and fed with the freshwater Chlorella. The growth rate of rotifers significantly increased with an increase in temperature. The maximum density of the rotifers with air-supply at 24 °C, 6500 ind. ml^–1, was significantly lower than those cultured at 28 and 32 °C, i.e. 8600 and 8100 ind. ml^–1, respectively. Dissolved oxygen levels decreased with time and ranged from 0.8 to 1.4 mg l^–1 when the density of freshwater rotifer was the highest at each temperature. The highest density (19200 ind. ml^–1) of freshwater rotifer was obtained in cultures with a supply of oxygen at 28 °C. Densities of 13500 and 17200 ind. ml^–1 were found at 24 and 32 °C, respectively. Levels of NH₃-N increased with time and a dramatic increase of NH₃-N was observed at high temperatures. Levels of NH₃-N at 24, 28 and 32 °C were 13.2, 18.5 and 24.5 mg l^–1, respectively. These levels coincided with the highest rotifer density at each of the three temperatures. When rotifers were cultured with an oxygen-supply and pH was adjusted to 7, the maximum density of rotifer reached 33500 ind. ml^–1 at 32 °C . These results suggested that high density culture of freshwater rotifer, B. calyciflorus could be achieved under optimal conditions with DO value of exceeding 5 mg l^–1 and NH₃-N values of lower than 12.0 mg l^–1.     

The dynamics of a model of a plankton-nutrient interaction

A stability analysis is given for a model of plankton dynamics introduced by Wroblewskiet al. (Global Biogeochem. Cycles 2, 199–218, 1988). The detailed dependence of the steady-states and their stability on the various model parameters is explicitly presented and analysed. It is shown that under certain conditions the coexistence of phytoplankton and zooplankton occurs in an orbitally stable oscillatory mode. A distinguished parameter is varied and the steady-states computed. The significance of the lack of stable steady-states leading to periodic population levels is investigated and related to certain oceanographic data. Partially supported by NSF Grant DMS-8902712. Supported by New Zealand University Grants Committee Postdoctoral Fellowship.     

Competitive interactions between the rotifer Synchaeta oblonga and the cladoceran Scapholeberis kingi Sars

Competition experiments showed that the small cladoceran Scapholeberis kingi rapidly excluded the rotifer Synchaeta oblonga from mixed-species cultures, but was itself unaffected by the presence of S. oblonga. Short-term experiments testing the effect of S. kingi on the survivorship and reproduction of S. oblonga showed that the former imposed a high mortality on the latter, even though shared food resources were abundant. These results indicate that adult S. kingi mechanically interferes with S. oblonga either by ingesting, or by rejecting in a damaged condition, individuals swept into its branchial chamber. In contrast to many other small species of cladocerans, and like large species of Daphnia, S. kingi has the potential to markedly suppress populations of some rotifer species through a combination of interference and exploitative competition.     

Production and nutritional quality of the rotifer Brachionus plicatilis fed marine Chlorella sp. at different cell densities

The effect of different cell densities of marine Chlorella sp. on the growth rate, doubling time and production of the rotifer Brachionus plicatilis was investigated. A significant increase in rotifer production was achieved at a density of 50 × 10⁶ Chlorella cells ml^–1. The nutritional quality of rotifers grown at different concentrations of Chlorella is discussed.     

The biochemical composition and calorific content of a rotifer and its algal food: comparison of a two stage chemostat and batch culture

It is often assumed that the use of a two-stage chemostat yields algal food with a well-defined nutritional composition that can maintain herbivores in a steady state of growth. In this study I investigated two bacteriafree culture techniques, continuous flow chemostats and batch cultures, to determine whether the biochemical composition of the rotifer Encentrum linnhei differed in the two cultures. Changes in the biochemical composition and calorific content of the algal food were also examined. In the rotifer reaction vessel only the lipid content of the algal food increased significantly with dilution rates, while significant decreases in protein and carbohydrates were detected at increasing algal densities. A different pattern was observed in the response of the unused algal cells to variables such as dilution, algal input and algal densities in the sump of the rotifer chemostat. In the chemostat the biochemical composition of the rotifers varied as expected with dilution rates, algal input and food availability but significant differences were found in the biochemical composition of the animals growing in the reaction vessel and those collected from the sump. In contrast, the biochemical content of batch-grown E. linnhei varied with time in a way that depended upon food availability and also on the biochemical state of the algal food. However, at the end of the exponential phase of growth, when maximum densities had been achieved, batch-grown rotifers were more biochemically nutritious than chemostat-grown animals in their steady-state phase.     

Differentiation during asexual reproduction and regeneration in a microturbellarian

Affordable biological technology for the reclamation of wastes and water of the waste streams from intensive livestock units is important in a country short of water. This study tested the concept of reclamation of waste by Streptocephalus macrourus (Crustacea: Anostraca) from the effluent of a high rate algal pond processing livestock wastes. S. macrourus showed a growth efficiency of 39% to 74% when fed optimal rations and cultured at densities between 10 and 400 1^–1. The maximum daily growth rates (0.15–0.21) approximate the growth rates of cladoceran or rotifer cultures managed for maximal biomass production. S. macrourus' ability to withstand crowding enabled the production from S. macrourus cultures (up to 91.8 mg dry mass l^–1 d^–1, or 1241 mg wet mass l^–1 d^–1) to exceed production recorded from cladoceran or rotifer cultures. Temperature influenced growth rate, with the highest growth rate occurring at 24 °C. The dilution rate of continuously fed cultures influenced growth rate, with the optimum dilution rate tested being 10 ml organism ^–1 d^–1. Mass mortality occurred when organisms were held at a density of 4000 l^–1. S. macrourus is able to convert algae grown on livestock waste efficiently into anostracan biomass, and is able to give a very high daily production.     

Induction of diapausing amictic eggs in Synchaeta pectinata

Amictic females of a clone of S. pectinata from Star Lake (Norwich, Vermont) may produce diapausing as well as non-diapausing (subitaneous) eggs. The proportion of diapausing eggs produced in cultures was unaffected by temperature (12 vs 19 °C) or rotifer population density (minima of 0.33 vs 3 ind. ml^–1) at 19 °C. However, at 19 °C this proportion was higher in cultures maintained at a low food level suppressing reproduction (5 × 10³ cells ml^–1 Cryptomonas erosa) than in those maintained at a high food level (2 × 10⁴ cells ml^–1); the treatment effect was marginally significant (p=0.067). Consistent with the effect of low food availability, a period of starvation was very effective in inducing the development of diapausing eggs. None of 19 females cultured individually from hatching at 19 °C on C. erosa (2 × 10⁴ cells ml^–1) in 1-ml volumes produced any diapausing eggs in 4 days (0 out of 349 eggs), while 13 out of 16 females subjected to a 15-hour starvation period 6 hours after birth produced one or more diapausing eggs during that time (34% of the 158 eggs produced by the 16 females were diapausing). Diapausing eggs produced and left at 19 °C hatched after 4 to 13 days. Those produced in cultures with a low food level took significantly longer to hatch (9.7 days) than those produced in cultures with a high food level (8.1 days) (p=0.022). In natural communities, S. pectinata should be able to respond directly and rapidly to poor food conditions by producing eggs that undergo an obligatory dormant period before resuming development.     

Axenic culture of <Emphasis Type="Italic">Brachionus plicatilis</Emphasis> using antibiotics

The rotifer Brachionus plicatilis culture is composed of complex microcosms including bacteria, protozoans, algae, and fungi. Previous studies reported methods to establish axenic rotifer cultures, but further refinement of these techniques is needed, for molecular biological research which requires pure culture to isolate nucleic acids from rotifers only. In order to render rotifer culture axenic, we tested five antibiotics: ampicillin (Amp), chloramphenicol (Cp), kanamycin (Km), nalidixic acid (Na), and streptomycin (Sm) at 30–100 μg/ml. Except for Cp, which reduces rotifer reproduction, all other antibiotics at the tested concentrations did not affect rotifer reproduction or show any toxic effects. A rotifer disinfection method was finally established by treating the resting eggs with 0.25% (w/v) sodium hypochlorite (NaOCl) for 3 min, washing with sterilized sea water, and then exposing the neonates to an Amp, Km, Na, and Sm mixture. Using four nutrient media, we confirmed that this protocol renders the rotifer culture bacterial and fungus free. The axenic rotifer culture generated here is useful not only for genetic analysis of Brachionus plicatilis, but for studying the rotifer life cycle without bacterial influence.     

Comparison of population dynamics between slow- and fast-growing strains of the rotifer Brachionus calyciflorus pallas in continuous culture

Summary The population dynamics of a slow- and a fast-growing strain of the rotifer Brachionus calyciflorus are compared. Rotifers were grown in steady-states, at various specific growth rates (), in both two-stage chemostat and turbidostat cultures on the green alga Chlorella pyrenoidosa. Population variables, including specific ingestion (I), loss (L) and filtration (F) rates, yield (Y), production (P) and half-saturation coefficient of growth (K _g), were calculated using a growth model based on saturation kinetics. I, L, F and K _g were shown to be higher and Y and P lower for the fast-growing strain. Differences between the two strains with regard to these variables may represent tradeoffs associated with the faster potential growth rate. Steady-state relationships between these values and for the fast-growing strain, however, deviated from model predictions which suggest a possible shift from carbon to non-carbon growth limitation.     

Seasonal changes in zooplankton abundance in the lower Rhine during 1987–1991

Zooplankton in the River Rhine was surveyed for five years at the Dutch sampling stations, Lobith (German/Dutch border) and Maassluis (at the point of discharge of the river into the North Sea). The zooplankton abundance showed an apparent seasonal pattern at both stations, characterized by low densities during the winter period, and higher densities during the summer period, with a spring peak. Zooplankton was dominated by rotifers at both stations, although during the winter periods the contribution of copepods was considerable. The rotifers were dominated byBrachionus angularis, B. calyciflorus, Keratella cochlearis andK. quadrata; the copepods by cyclopoid nauplii; the cladocerans by small-sized species mainly belonging toBosmina. At Maassluis the relative contribution of copepods was higher than at Lobith. Furthermore, the zooplankton at Maassluis included the speciesEurytemora affinis, characteristic for estuarine conditions. In spring, the rotifer density and water temperature and rotifer density and chlorophylla concentration were positively correlated. Furthermore, both rotifer density and chlorophylla were inversely correlated with discharge. The possible role of environmental factors (water temperature, chlorophyll content, discharge and biotic factors) controlling the river zooplankton dynamics is indicated.     

Zooplankton densities in a Hydrilla infested lake

The number of individuals and species of zooplankton were sampled concurrently with Hydrilla biomass and water quality for one year in a small, eutrophic central Florida lake. Throughout the study, rotifer species and individuals dominated the zooplankton. The abundance of the zooplankton tended to remain high when Hydrilla biomass was at its seasonal low during late winter and early spring. When hydrilla growth increased in the late spring and summer months causing a decrease in total alkalinity, specific conductivity, water color, turbidity, orthophosphate and chlorophyll a concentrations; the abundance of the zooplankton declined. During this time, there was a shift from limnetic to littoral species, principally rotifers. Hydrilla growth did not affect the mean number of cladoceran or copepod species, but may have led to an increase in rotifer species.     

Susceptibility of Ephemeral Pool <Emphasis Type="Italic">Hexarthra</Emphasis> to Predation by the Fairy Shrimp <Emphasis Type="Italic">Branchinecta mackini</Emphasis>: can Predation Drive Local Extinction?

Isolated rock pools in the US desert southwest often develop dense populations of a lithophilic species of the rotifer Hexarthra. I hypothesized that rotifers persist in these isolated ponds due to the absence of either competition with or predation by potentially sympatric crustaceans, especially anostracans. I tested the latter idea with laboratory and field experiments, in each case exposing rotifers to adult fairy shrimp (Branchinecta mackini, the dominant anostracan in this region in winter ponds) in 200 ml microcosms. In␣most cases individual female fairy shrimp had distinct negative effects on rotifer suspensions due to direct predatory consumption of the smaller animals. Estimated effective water processing rates ranged from ca. 50 to over 300 ml ind⁻¹ h⁻¹ and rotifer consumption rates by female B. mackini were between 280 and > 600 Hexarthra consumed per individual predator per hour. Male B. mackini never significantly reduced rotifer numbers in either laboratory or field microcosms. The results indicate that, while perhaps not the sole determinant of rotifer distribution in these ephemeral pools, fairy shrimp predation can have a strong negative influence on natural Hexarthra populations.     

Effect of two microalgae concentrations on body size and egg size of the rotifer <Emphasis Type="Italic">Brachionus calyciflorus</Emphasis>

The rotifer, Brachionus calyciflorus, was grown with two algae species (Chlorella sp. and Scenedesmus obliquus) at different concentrations (0.1, 1 and 10 × 10⁶ cells ml⁻¹). The body size (lorica biovolume) of individual rotifer and their egg size were measured when the populations were roughly in the exponential phase of population growth. The body size of the rotifers differed significantly (P < 0.05) among the two algae species used, however this effect was not observed for egg size. The body size of rotifers fed on higher densities of Chlorella sp. (10 × 10⁶ cells ml⁻¹) was significantly larger than for those fed on lower and medium densities (0.1 and 1 × 10⁶ cells ml⁻¹). Body size and egg size of rotifers fed with different amounts of Scenedesmus did not differ significantly. The egg size was significantly larger at higher food level of Chlorella. A significantly positive correlation was observed between the adult rotifer body size and their egg size.     

The relationship between Trichocerca pusilla (Jennings), Aulacoseira spp. and water temperature in Loch Leven,Scotland, U.K.

Loch Leven is a shallow, eutrophic lake in the Scottish lowlands that is famous for its brown trout (Salmo trutta L.) fishery. Studies of planktonic rotifer populations began here in January 1977. Since then, samples have been collected and analysed at more or less weekly intervals. Additional information on the composition and abundance of phytoplankton and crustacean zooplankton species, and on a variety of physical and chemical determinants, has been recorded on each sampling occasion.Long-term datasets, such as that described above, are invaluable for identifying interactions between components of the plankton that only appear for short periods each year, as these interactions would probably be overlooked in data spanning a shorter period of time. This study uses the long-term data from Loch Leven to examine the food and temperature requirements of the summer rotifer species Trichocerca pusilla (Lauterborn). The results suggest that T. pusilla prefers water temperatures above 12 °C and that it feeds, primarily, on the filamentous diatom Aulacoseira spp. During the summer months, its abundance was closely related to the availability of this diatom. When filaments of Aulacoseira spp. were abundant, rotifer densities reached 1000–3000 ind. l^–1 and when they were scarce (e.g. 1980, 1997 and 1998) T. pusilla densities also remained low (i.e. less than 100 ind. l^–1). The reason for the success or failure of Aulacoseira during the summer months each year is unclear but, in general, its abundance was related to the availability of dissolved silica in the water.     

Differentiated pellicle organization and lipopeptide production in standing culture of <Emphasis Type="Italic">Bacillus subtilis</Emphasis> strains

Pellicle formation and lipopeptide production was analysed in standing cultures of different Bacillus subtilis strains producing two or three families of lipopeptides. Despite its ability to produce surfactin, B. Subtilis ATCC 6633 was unable to form stable pellicle at air–water interface. For the ATTC 21332 and ATCC 9943 strains, it was shown for the first time that the lipopeptides were also produced in standing cultures at productivities similar or lower than those obtained when the culture medium is agitated. A differentiated behaviour was observed between these strains in repetitive batch cultures. B. subtilis 9943 formed a wrinkled, thinner and more resistant pellicle than B. subtilis 21332. The structure of the pellicle determined by electron microscopy observations showed that cells of B. subtilis 9943 formed microcolonies whereas those of B. subtilis 21332 rapidly died. Under these conditions, surfactin production by strain 21332 decreased after 2 days whereas it remained stable for B. subtilis 9943 during the 6 days of the cultures. These data indicate that cells of B. subtilis strains growing in pellicle can produce lipopeptides differently depending on their cellular organisation. M. Chollet-Imbert and F. Gancel have contributed equally to the scientific work.     

Experimental study on the impact of dinoflagellate Alexandrium species on populations of the rotifer Brachionus plicatilis

To investigate harmful effects of the dinoflagellate Alexandrium species on microzooplankton, the rotifer Brachionus plicatilis was chosen as an assay species, and tested with 10 strains of Alexandrium including one known non-PSP-producer (Alexandrium tamarense, AT-6). HPLC analysis confirmed the PSP-content of the various strains: Alexandrium lusitanicum, Alexandrium minutum and Alexandrium tamarense (ATHK, AT5-1, AT5-3, ATCI02, ATCI03) used in the experiment were PSP-producers. No PSP toxins were detected in the strains Alexandrium sp1, Alexandrium sp2.Exposing rotifer populations to the densities of 2000 cells ml⁻¹ of each of these 10 Alexandrium strains revealed that the (non-PSP) A. tamarense (AT-6) and two other PSP-producing algae: A. lusitanicum, A. minutum, did not appear to adversely impact rotifer populations. Rotifers exposed to these three strains were able to maintain their population numbers, and in some cases, increase them. Although some increases in rotifer population growth following exposures to these three algal species were noted, the rate was less than for the non-exposed control rotifer groups.In contrast, the remaining seven algal strains (A. tamarense ATHK, AT5-1, AT5-3, ATCI02, ATCI03; also Alexandrium sp1 and Alexandrium sp2) all have adverse effects on the rotifers. Dosing rotifers with respective algal cell densities of 2000 cells ml⁻¹ each, for Alexandrium sp1, Alexandrium sp2, and A. tamarense strains ATHK and ATCI03 showed mean lethal time (LT₅₀) on rotifer populations of 21, 28, 29, and 36h, respectively. The remaining three species (A. tamarense strains AT5-1, AT5-3, ATCI02) caused respective mean rotifer LT₅₀s of 56, 56, and 71 h, compared to 160 h for the unexposed “starved control” rotifers. Experiments to determine ingestion rates for the rotifers, based on changes in their Chlorophyll a content, showed that the rotifers could feed on A. lusitanicum, A. minutum and A. tamarense strain AT-6, but could graze to little or no extent upon algal cells of the other seven strains. The effects on rotifers exposed to different cell densities, fractions, and growth phases of A. tamarense algal culture were respectively compared. It was found that only the whole algal cells had lethal effects, with strongest impact being shown by the early exponential growth phase of A. tamarense. The results indicate that some toxic mechanism(s), other than PSP and present in whole algal cells, might be responsible for the adverse effects on the exposed rotifers.     

A field and laboratory study on factors affecting polymorphism in the rotifer Keratella tropica

Summary We present data on polymorphism of the rotifer Keratella tropica from an outdoor, mesoscale experiment and a series of laboratory induction assays. In the outdoor experiment the biomass of planktonic crustaceans was noticeably depressed by larval fish predation, and associated with this depression K. tropica underwent a striking reduction of caudal spines. Subsequent laboratory studies showed that the positive association between crustacean biomass and caudal spine length was a cause and effect relationship. This is the first record of rotifer morphological change as an indirect effect of fish predation. In laboratory experiments filtrates of monospecific cultures of a cyclopoid copepod, a calanoid copepod and 5 cladocerans induced a remarkable spine development. Morphological induction showed a direct relationship with the concentration of crustaceans, both under field and laboratory conditions. Long spines were found to be strong deterrents against small predators (Acanthocyclops robustus copepodites), but were useless against large ones (females of the same species). The morphotypes of K. tropica obtained by experimental induction from a single clone encompass much of the worldwide variation of the species.