Spatial coexistence of phytoplankton species in ecological timescale

The species diversity of phytoplankton is usually very high in wild aquatic systems, as seen in the paradox of plankton. Coexistence of many competitive phytoplankton species is extremely common in nature. However, experiments and mathematical theories show that interspecific competition often leads to the extinction of most inferior species. Here, we present a lattice version of a multi-species Lotka–Volterra competition model to demonstrate the importance of local interaction. Its mathematical equilibrium is the exclusion of all but one superior species. However, temporal coexistence of many competitive species is possible in an ecological time scale if interactions are local instead of global. This implies that the time scale is elongated many orders when interactions are local. Extremely high species diversity of phytoplankton in aquatic systems may be maintained by spatial coexistence in an ecological time scale.Tatsuo Miyazaki, Kei-ichi Tainaka, and Jin Yoshimura contributed equally to this study.     

In situ microcosm experiments on the influence of nitrate and light on phytoplankton community composition

Phytoplankton blooms are fundamental features of coastal ecosystems, but the processes that select for blooms of certain species are not well understood. The aim of this work was to investigate experimentally the interaction of light and nutrients (nitrate) in structuring phytoplankton community composition in Pelorus Sound, New Zealand. Microcosm experiments were conducted in situ nine times throughout the year, providing controls and treatments for increased nutrients and decreased light. Nitrate availability was found to be limiting to phytoplankton growth during spring and summer. Small- to medium-sized, chain-forming diatom taxa such as Chaetoceros sp., Skeletonema sp., Pseudonitzschia sp. and Thalassiosira sp. responded most rapidly to nitrate enrichment, increasing their biovolume up to 32-fold during the 5-day experiments. A long-term phytoplankton monitoring database showed that these taxa have historically dominated the phytoplankton assemblage, suggesting that intense competition for nitrate is a key component in structuring the phytoplankton community. Many of the taxa that were able to withstand light reduction in the shaded treatments were rare historically in Pelorus Sound, suggesting that light is secondary to nitrate availability in structuring the phytoplankton community composition in this coastal embayment.     

Immersed in situ microcosms: A tool for the assessment of pollution impact on phytoplankton

In situ phytoplankton microcosms were developed and characterized for use in toxicity testing. The microcosms contained 225 μm filtered seawater maintained in 1 liter glass bottles attached to a plastic frame and immersed at 3 m under the sea surface. Synechococcus and picoeukaryote population dynamics in microcosms and the surrounding water were compared. A bloom-like behaviour observed for Synechococcus in these phytoplankton microcosms was avoided when 10% of the culture volume was replaced, every two days, by filtered seawater. After 2 weeks, no significant difference in Synechococcus and picoeukaryotes cell counts was observed in microcosms compared to the surrounding free seawater. Synechococcus fluorescence at 545 nm (phycoerythrobilin) fluctuated with a similar pattern in such microcosms and in free seawater and were shown to be correlated to light intensity fluctuations over a two week experiment. The in situ microcosms were used to study the impact of low copper additions. Synechococcus populations were dramatically decreased by copper addition, while picoeukaryote populations were increased simultaneously. Our data show that drastic changes in species composition can occur at copper concentrations encountered in polluted coastal areas.     

Seasonal succession of phytoplankton in a large subarctic river

The factors influencing the abundance of phytoplankton in the Yellowknife River, in the Canadian subarctic, were determined from collections made for 42 consecutive months from June 1975 to November 1978. The spring bloom of plankton occured during April of each year in response to changing light conditions. WhileChlamydomonas lapponica was dominant during this period, it was replaced during the early part of the summer by a rapid succession ofDinobryon species in whichD. cylindricum was followed byD. sociale and in turn byD. bavaricum andD. divergens. Although low nutrient levels permitted the development ofDinobryon during the summer, the abundance of diatoms was greatly limited by the concentrations of SiO₂ (< 0.1 g/m³). Algal densities began to decline in August and reached low overwintering levels by November. The absence of a fall bloom in densities was due to a combination of low temperatures and nutrient levels.P.O. Box 2310, Yellowknife, Northwest Territories, X1A 2P7, Canada     

Effects of nitrogen limitation on species replacement dynamics during early secondary succession on a semiarid sagebrush site

Summary A soil nitrogen (N) availability gradient was induced on a disturbed sagebrush site in northwestern Colorado by fertilizing with nitrogen (high available N), applying sucrose (low available N), and applying neither nitrogen nor sucrose (control). Species composition was studied for 3 years. At the end of the study, N concentration of aboveground tissue of 3 major species was determined. The rate of species replacement was most rapid on plots receiving the sucrose treatment and was slowest on plots receiving the N treatment. Early-seral dominats had greater tissue N concentrations when availability of the resource was high but lower tissue N concentrations when available soil N became limited. Midseral dominants displayed the opposite pattern. These results suggest that the supply of available soil N, and therefore the dynamics of N incorporation in perennial plant tissue, is a primary mechanism in controlling the rate of secondary succession within this semiarid ecosystem.     

有害赤潮藻对浮游动物影响的研究进展

有害赤潮藻对浮游动物的影响在赤潮危害的研究中越来越受到关注,本文综述了有害赤潮藻对浮游动物的毒害作用,主要表现为影响浮游动物的存活和生长,降低母体的产卵力、卵的受精率或受精卵的孵化率以及减慢胚胎发育或造成畸形,影响幼体的正常发育,抑制浮游动物的摄食和改变其正常行为等;这些毒害作用主要是由赤潮藻产生的藻毒素或其它一些胞外渗出物通过降低浮游动物对食物的利用率、造成生理损伤、破坏细胞膜、降低亲体产生的配子质量、抑制有丝分裂和胚胎发育等造成的．某些赤潮藻营养成分单一如缺乏多不饱和脂肪酸,也会对浮游动物的生长繁殖产生影响．     

Predation-driven dynamics of zooplankton and phytoplankton communities in a whole-lake experiment

Summary 1. Species compositions of zooplankton and phytoplankton were followed in Tuesday Lake before and after experimental manipulation of its fish populations (addition of piscivorous largemouth bass, removal of planktivorous minnows). Plankton dynamics were compared to those of adjacent, unmanipulated Paul Lake, where piscivorous fish have been dominant historically. 2. Indices of similarity for the zooplankton communities in the two lakes in 1984 prior to the manipulation were low; however, following the manipulation in spring, 1985, similarity of the zooplankton in the two lakes rose considerably and remained high throughout 1986. This was the result of an increase in Tuesday Lake of previously rare large-bodied cladocerans (Daphnia pulex, Holopedium gibberum) which were the dominants in Paul Lake, and the disappearance in Tuesday Lake of the dominant small-bodied copepod Tropocyclops prasinus, a minor component of the Paul Lake zooplankton. These observations are consistent with prior observations of the effects of size-selective predation on zooplankton communities. 3. Phytoplankton communities also responded strongly to the manipulation, with similarity indices for the two lakes rising from low levels in 1984 to high levels of similarity in 1985 and 1986, reflecting the decrease of formerly dominant Tuesday Lake taxa which were unimportant in Paul Lake and the appearance or increase in Tuesday Lake of several taxa characteristic of the Paul Lake phytoplankton assemblage. these results clearly show that food web structure can have pronounced effects on community composition at all levels of the food web, and that, just as zooplankton communities are structured by sizeselective predation, phytoplankton communities are structured by herbivory. These observations may provide some insight into factors governing the complex distributions of phytoplankton species among various lakes.A contribution from the University of Notre Dame Environmental Research Center, funded by NSF grants BSR-83-08918 and BSR-86-06271     

The species composition of Antarctic phytoplankton interpreted in terms of Tilman's competition theory

Summary An attempt was made, to test for the impact of resource competition on Antarctic marine phytoplankton. According to theory, species composition near competitive equilibrium should be determined by the ratios of limiting resources. Enrichment bioassays identified silicon and nitrogen as limiting nutrients for some of the most important phytoplankton species during early austral summer in the region near the Antarctic Peninsula. Together with the generally acknowledged limiting resource light, this gave three meaningful ratios of essential resources (Si:N, Si:light, N:light) and one ratio of substitutable resources (NO₃:NH₄). Phytoplankton species assemblages were found to be well separated by the ratios of the essential resources and by mixing depth. Nine out of 12 individual species were found to be separated along at least one of the gradients of resource ratios. Where comparison with competition experiments was available, predicted and realized distributions of species were compatible.     

Effects of leaf litter species on macroinvertebrate community properties and mosquito yield in Neotropical tree hole microcosms

Detritus quality and quantity affect macroinvertebrate productivity and distribution in many freshwater ecosystems. This study experimentally investigated the effects of leaf litter from Ceiba pentandra, Dipteryx panamensis, Ficus yoponensis, and Platypodium elegans on macroinvertebrate species composition, richness, and abundance in artificial water-filled tree holes in a lowland moist forest of Panama. Species composition was similar among treatments, but species richness and longevity differed among litter types and were consistently highest with Platypodium litter. Similar patterns were observed in natural tree holes of the focal tree species. The mosquito Culex mollis was the most abundant species in the field experiment. Average conductivity and dissolved oxygen concentration differed among leaf species, but pH did not. Leaf toughness was positively correlated with mean macroinvertebrate abundance and cumulative species richness. A laboratory experiment measured C. mollis yield and pupation time in tree hole microcosms containing the four litter species. Cumulative mosquito mass and time to pupation differed among leaf litter species, with Platypodium litter supporting the greatest yield. Pupation was slowest on Ceiba litter. Grazing by mosquito larvae facilitated leaf decomposition in all treatments. Results suggest that differences in macroinvertebrate species richness and mosquito yield can be attributed to differences in nutritional quality among litter species. Received: 14 October 1998 / Accepted: 21 February 1999     

Comparison of productivity and phytoplankton in a warm (Kongsfjorden) and a cold (Hornsund) Spitsbergen fjord in mid-summer 2002

Kongsfjorden and Hornsund are two glacial fjords without sills on the West Spitsbergen coast. Both sites are under the influence of relatively warm Atlantic-derived water, although Hornsund is more influenced by cold water from the Barents Sea. In this study, we compared the impacts of cold Arctic and warmer Atlantic waters on the pelagic ecosystems of Kongsfjorden and Hornsund. Both fjords were strongly influenced by Atlantic-derived waters during summer (2002). Diatoms were the most substantial contributors to phytoplankton biomass, especially in outer basins of both fjords, whereas the second most important contributors were autotrophic dinoflagellates in Kongsfjorden and nanoflagellates in Hornsund. Total phytoplankton biomass was highest in Hornsund. Primary production rates were an order of magnitude lower in Kongsfjorden than in Hornsund, and increased from inner to outer fjord (from 2.47 to 4.48 mg C m⁻² h⁻¹ in Kongsfjorden and from 14.00 to 86.65 mg C m⁻² h⁻¹ in Hornsund). Chlorophyll-a concentration was also substantially lower in Kongsfjorden. Zooplankton was dominated by omnivorous species in Kongsfjorden and herbivorous in Hornsund. Observed differences between the fjords may originate from (1) advection of different waters into the fjords; (2) differences in freshwater runoff and turbidity, and (3) timing of the phytoplankton bloom. Climate warming will likely increase the Atlantic water influence, and result in reduced production of diatoms and increase in flagellates.     

Competition among the pioneers in a seasonal soft-bottom benthic succession: field experiments and analysis of the Gilpin-Ayala competition model

Summary Controlled experiments, designed to assess the effects of pioneers on succession on an intertidal sandflat, provided evidence for interspecific competition between juvenile Hobsonia florida (Polychaeta, Ampharetidae) and oligochaetes. The field data were fitted to both the linear Volterra and non-linear Gilpin-Ayala competition equations. With its greater number of parameters, the Gilpin-Ayala model must provide a better fit to observed population abundances. The Gilpin-Ayala model is flawed as an explanation of the population trajectories of the H. florida and oligochaetes, because its non-linearity parameter affects only intraspecific competion. With either model our field data demonstrate a solution to Hutchinson's paradox. With competition coefficients near unity and similar carrying capacities, the predicted population trajectories are heavily dependent on initial conditions. The predicted times to competitive exclusion are long and can easily exceed the typical period of environmental constancy. Our study offers evidence for Neill's competitive bottleneck: competition acts primarily on the developmental stages of one of a pair of competing species. The permanent meiofauna may act as a competitive bottleneck for the population growth of benthic macrofauna. The mechanism of this competitive interaction probably involves exploitative interspecific competition for benthic diatoms.     

The response of zooplankton and phytoplankton from the North American Great Lakes to filtration

Filtration of ballast water was investigated as a means of minimizing the introduction of nonindigenous zooplankton and phytoplankton by ships visiting the North American Great Lakes-St. Lawrence Seaway system (GLSLSS). An automatic backwash screen filtration (ABSF) system with nominal filtration options of 25, 50 or 100 μm was mounted on the deck of an operating Seaway-sized dry bulk carrier, the MV Algonorth. Water was pumped through the ABSF with a deck mounted pump at 341 m³ hr⁻¹ during routine ship operations in the GLSLSS, and effectiveness of the various screen pore sizes at removing taxonomic categories of zooplankton and phytoplankton was measured using matched treatment and control ballast tanks. The smallest pore sizes (25 and 50 μm) performed better than the 100 μm pore size at removing biological material. There was no difference in the filtration efficiency of the 25 and 50 μm screens relative to macro- or microzooplankton in these tests, but this result was probably due to low densities of macrozooplankton, and soft-bodied (aloricate) characteristics of the microzooplankton present. The 25 and 50 μm pore sizes were subjected to more controlled tests on board a stationary barge platform equipped with triplicate 700 L catchment bins moored in Duluth Harbor of Lake Superior. In these tests, filter pore size, organism size and rigidity influenced zooplankton removal efficiency by the ABSF. The 25 μm screen reduced both macrozooplankton and microzooplankton significantly more than the 50 μm screen. Zooplankton width was more determinative of filtration performance than length, and both filters removed loricate species of rotifers significantly more efficiently than aloricate species of the same length and width size classes. The 25 and 50 μm ABSF also significantly reduced algal densities, with the exception of colonial and filamentous green algae (50 μm only). Filter efficiency relative to algal particles was influenced by filter pore size, organism morphology and structure, and intake density, while algal particle size was not determinative. This research provides compelling evidence that 25 or 50 μm filtration is a potentially powerful means of reducing densities of organisms discharged by ships operating in the Great Lakes but an additional treatment step would be necessary to effectively minimize risk and meet the International Maritime Organization's discharge standards associated with organisms of all sizes in the water column.     

Plankton composition and water quality in a pond of spring origin in Turkey

Phyto/zooplankton composition, chlorophyll a, and some water quality parameters were investigated in a spring-originated pond in Central Anatolia between February 2001 and January 2002. Water temperature, pH, dissolved oxygen, Secchi depth, total and calcium hardness, nitrate-nitrogen, nitrite-nitrogen, ammonia-nitrogen, total phosphorus, and soluble reactive phosphorus levels were analyzed. A total of 49 species belonging to Bacillariophyceae, Chlorophyceae, Cyanophyceae, Cryptophyceae, and Dinophyceae were identified. The highest phytoplankton abundance was found in August, whereas the lowest was determined in January. Phytoplankton abundance increased from February to August and declined in the following months. The Bacillariophyceae were dominant in the phytoplankton community. A total of 21 species of Rotifera, 2 species of Cladocera, and 1 genus of Copepoda were found. The zooplankton community was dominated by Rotifera. The highest abundance of zooplankton was recorded in July and the lowest value in November. The annual mean concentration of chlorophyll a was measured as 1.90 μg l⁻¹. In spite of these eutrophic levels (mean values of total phosphorus and nitrate-nitrogen: 0.069 mg P l⁻¹ and 0.68 mg N l⁻¹), phytoplankton cannot grow satisfactorily because of the short water retention time (0.6 day⁻¹). The shallowness of the pond together with the low phytoplankton biomass and the high concentrations of nutrients are discussed.     

Seasonal variations in upwelling and in the grazing impact of copepods on phytoplankton off A Coruña (Galicia, NW Spain)

The impact of grazing by copepods on phytoplankton was studied during a seasonal cycle on the Galician shelf off A Coruña (NW Spain). Grazing was estimated by measuring the chlorophyll gut content and the evacuation rates of copepods from three mesh-size classes: 200-500 (small), 500-1000 (medium), and 1000-2000 μm (large). Between February 1996 and June 1997, monthly measurements of water temperature, chlorophyll concentration, primary production rates, and copepod abundance, chlorophyll gut content, and evacuation rates were taken at an 80-m-deep, fixed shelf station. Additionally, the same measurements were collected daily during two bloom events in March and in July 1996. Small copepods were the most abundant through the seasonal cycle. The highest grazing impact, however, was due to the medium and large size classes. Grazing by small copepods exceeded grazing by medium and large copepods only during phytoplankton spring blooms. The impact of copepod grazing (considering all size fractions) was generally low. On average, 2% of the phytoplankton biomass and 6% of the primary production were removed daily by the copepod community. Maximum grazing impact values (9% of the phytoplankton biomass and 39% of the primary production) were found in mid-summer. These results suggest that most of the phytoplankton biomass would escape direct copepod grazing in this upwelling area.     

An ecosystem model of the phytoplankton competition in the East China Sea, as based on field experiments

An ecological dynamic model for the simulation of two pelagic phytoplankton groups is developed in this article. Model parameters were adjusted and validated based on the light-limited field culture experiments and the mesocosm experiments in the East China Sea (ECS). The calculation comparisons from the proposed model, along with field experiment observations, show that the model simulate the datasets very well, qualitatively and quantitatively. The parameters’ sensitivity analysis indicates that the competition between the diatoms and dinoflagellates is most sensitive to the photosynthetic process, followed by the exudation process of the phytoplankton, while the autolysis and respiration processes of phytoplankton and the grazing and exudation processes of zooplankton can also influence this competition to some extent. The sensitive parameters include: the photosynthetic optimal specific rate; the optimal irradiance and optimal temperature for phytoplankton growth; and the half-saturation constant for limiting nutrients, etc. Results of the sensitivity analysis also indicate that light, temperature and limiting nutrients are the controlling environmental factors for the competition between the diatoms and dinoflagellates in the ECS. In order to explore the effects of light and nutrients on the phytoplankton competition, simulations were carried out with varying light and nutrient conditions. Model simulations suggest that the diatoms favor higher irradiance, lower DIN/PO₄–P ratios, higher SiO₄–Si/DIN ratios and higher nutrient concentrations, as compared to the dinoflagellates. These results support the speculation that the increase in the DIN/PO₄−P ratio and the decrease in the SiO₄–Si/DIN ratio in the ECS may be responsible for the composition change in the functional Harmful Algal Bloom (HAB) groups from the diatom to the dinoflagellate communities over the last two decades. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: L. Naselli-Flores     

Phylogenetic diversity of bacterial communities associated with bioremediation of crude oil in microcosms

Bioremediation, mainly by indigenous bacteria, has been regarded as an effective way to clean up oil pollution after an oil spill. In order to obtain a systematic understanding of the succession of bacterial communities associated with oil bioremediation, sediments collected from the Penglai 19-3 oil platform were co-incubated with crude oil. Oil biodegradation was assessed on the basis of changes in oil composition monitored by GC–MS. Changes in the bacterial community structure were detected by two 16S rRNA gene based culture-independent methods, denaturing gradient gel electrophoresis (DGGE) and clone library. The results suggested that crude oil was rapidly degraded during the 30-day bioremediation period. Bacteria affiliated with the genus Pseudomonas dominated all three clone libraries. But dramatic changes were also detected in the process of biodegradation of crude oil. The “professional hydrocarbonocastic bacteria” (e.g., Alcanivorax) became abundant in the two samples during the bioremediation period. Meanwhile, δ-proteobacteria was only detected in the two samples. Information on the bacterial community revealed in this study will be useful in developing strategies for bioremediation of crude oil dispersed in the marine ecosystem.     

Seasonal succession of phytoplankton in Lake Yogo over 2 years: effect of artificial manipulation

We studied the seasonal variation in concentrations of nutrients and phytoplankton in Lake Yogo for 2 years, from May 2000 to May 2002, in order to clarify the seasonal succession of phytoplankton and the effect of various manipulations on it. It was revealed that in spite of the installation of aeration systems and the pumping of mesotrophic water from Lake Biwa during the summer season, the trophic state of Lake Yogo overall has not improved during the past few decades. However, the pumping of water from Lake Biwa did affect the concentrations of nutrients and the periods of cyanobacterial bloom during the summer. The pumping period was different in each year, and the cyanobacterial bloom occurred during the period without pumping in both years. The aeration destratification was not strong enough to prevent cyanobacterial blooms. Cyanobacteria and Bacillariophyceae contributed most to the phytoplankton biomass in both years. Aphanizomenon, Anabaena, and Microcystis were the main genera among cyanobacteria. The bloom of Aphanizomenon or Anabaena occurred early in the summer, and was then replaced by Microcystis. Aphanizomenon was almost always present, and often formed bloom even in winter. The seasonal succession of Bacillariophyceae was almost the same in both years and was well categorized: winter-growing species such as Aulacoseira pusilla (F. Meister) Tuji et Houki and species of Thalassiosiraceae, spring-growing species such as Asterionella formosa Hassall, Fragilaria crotonensis Kitton, and Synedra cf. acus, and fall-growing species such as Aulacoseira ambigua (Grunow) Simonsen, and Aulacoseira granulata (Ehrenb.) Simonsen.     

Zooplankton, phytoplankton and the microbial food web in two turbid and two clearwater shallow lakes in Belgium

Components of the pelagic food web in four eutrophic shallow lakes in two wetland reserves in Belgium (Blankaart and De Maten) were monitored during the course of 1998–1999. In each wetland reserve, a clearwater and a turbid lake were sampled. The two lakes in each wetland reserve had similar nutrient loadings and occurred in close proximity of each other. In accordance with the alternative stable states theory, food web structure differed strongly between the clearwater and turbid lakes. Phytoplankton biomass was higher in the turbid than the clearwater lakes. Whereas chlorophytes dominated the phytoplankton in the turbid lakes, cryptophytes were the most important phytoplankton group in the clearwater lakes. The biomass of microheterotrophs (bacteria, heterotrophic nanoflagellates and ciliates) was higher in the turbid than the clearwater lakes. Biomass and community composition of micro- and macrozooplankton was not clearly related to water clarity. The ratio of macrozooplankton to phytoplankton biomass – an indicator of zooplankton grazing pressure on phytoplankton – was higher in the clearwater when compared to the turbid lakes. The factors potentially regulating water clarity, phytoplankton, microheterotrophs and macrozooplankton are discussed. Implications for the management of these lakes are discussed.     

Comparison of species composition between different grassland management treatments after 25 years

Abstract. To identify management treatments suitable for the conservation of extensively managed grasslands, the ‘Fallow experiments in Baden‐Württemberg’ were set up in 1975. In this investigation, species composition of the grazing, mowing, mulching, controlled burning and unmanaged (succession) treatments were analysed after 25 yr of continuous management in Arrhenatherum elatius and Bromus erectus grasslands. Through ordination analyses it was found that species composition is strongly dependent on the management treatment. The first axis, identified by ordination analysis, essentially corresponded to a gradient of decreasing disturbance frequency. Controlled burning resulted in a unique species composition. Grazing, mowing and mulching twice a year were found to be most suitable for the conservation of unimproved, species‐rich grasslands.