Steady state phytoplankton assemblages during thermal stratification in deep alpine lakes. Do they occur?

Phytoplankton seasonal and long-term succession can be described and functionally classified by associations similar as in terrestrial vegetation studies. Such a concept has to include 'climax' into pelagic succession which in turn leads to the question to what extent steady state assemblages occur and if during periods of dynamic equilibrium can be identified. Here we explore the situation with respect to the above question for deep, nutrient poor, alpine lakes in Austria. We first track the long-term development of phytoplankton biomass, their taxonomic structure and their relation to total phosphorus and chlorophyll-a as predictors of trophic state over the past 35 years. We then analyse this data set for coherent algal associations which can be ascribed to trait separated functional groups according to Reynolds et al. (2002). A three year period of stable environmental conditions has then be extracted from the progression of trophic state indices, having similar dominating species each year. These years were finally analysed for steady state conditions according to definitions given in Sommer et al. (1993). During thermal stratification, achievement of an equilibrium could be ruled out although coexistence of several dominating species lasted for several weeks. Habitat templates were constructed from environmental variables prior to biomass peaks for two species important in summer assemblages, the dinoflagellate Ceratium hirundinella and the diatom Fragilaria crotonensis. In summary, functional groups proved to be a valid and useful concept to describe species succession of phytoplankton in deep alpine lakes while pelagic climax is much less clear and steady state conditions were never met.     

Macrophytes moderate the taxonomic and functional composition of phytoplankton assemblages during a nutrient loading experiment

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Steady-state assemblages of phytoplankton in four temperate lakes (NE U.S.A.)

For four temperate lakes (Northeast U.S.A.) we identify periods of persistent phytoplankton assemblages and investigate the ecological conditions that correlate to these persistent assemblages. Periods of persistent assemblages, here considered as steady-state phases, were defined according to equilibrium criteria (two or three coexisting species, contributing to 80% of the standing biomass, for at least 2 weeks) defined by Sommer et al. (1993, Hydrobiologia 249: 1–7). For all four lakes, samples were taken weekly during the ice-free season and phytoplankton attributes (biomass, assemblages, diversity, species richness, change rates) and abiotic variables (temperature, I^* – as light mean in the mixing zone – z_mix, and nutrients) were analysed. Chodikee (CH), an eutrophic and rapidly flushed lake, did not show any persistent phase. The remaining three lakes showed single steady-state phases that occurred at varying times during the ice-free season. Steady-state phases occurred during early stratification in late spring in the stably stratified oligotrophic Mohonk Lake (MO), in the late summer stratification in the meso-eutrophic Stissing Lake (ST), and during spring mixing in Wononscopomuc Lake (WO). MO showed a 3-week period with dominance of F assemblage (Botryococcus braunii, Willea wilhelmii and Eutetramorus planctonicus), characteristic for clear epilimnia, tolerant to low nutrient and sensitive to high turbidity. For three weeks, ST had a stable assemblage with dominance of Lo(Woronichinia sp.), common assemblage in summer epilimnion of mesotrophic lakes and sensitive to prolonged or deep mixing; and P, assemblage able to live in eutrophic epilimnia with mild light and sensitive to stratification and silica depletion. In contrast, the mesotrophic Wononscopomuc Lake (WO) showed persistent assemblages during a 4-week period of spring circulation, when a dinoflagellate (Lo) was co-dominant with Nitzschia acicularis (C). The latter species is characteristic for mesotrophic lakes, tolerant to low light and sensitive to stratification and silica depletion. Both Lo and P assemblages, among seven others, had before been quoted, in literature, as dominant in maturing stages. We could not find consistent statistical differences between the periods classified as steady-state and non-steady-state. However, the data demonstrated that prolonged period of both mixing and stratification can maintain dominant assemblages. Although, historically sensed as opposite mechanisms, both mixing and stratification, if persistent, were observed maintaining dominant assemblages because both scenarios are characterized by environmental constancy.     

Dominant species, functional assemblages and frequency of equilibrium phases in late summer phytoplankton assemblages in Hungarian small shallow lakes

Late summer phytoplankton associations were studied qualitatively and quantitatively in 80 Hungarian lakes altogether (mostly shallow salt lakes, reservoirs, oxbows, gravel pit lakes). Equilibrium phases sensu Sommer et al. (1993) were found only in 17 lakes. Most of them were under some kind (high salt content or very low level of nutrients) of stress factor. It is concluded that environmental stress forces phytoplankton communities towards equilibrium. No relationship between occurrence of equilibria and trophic state was found. Species number of non-equilibrated lakes was almost three times as high as those in equilibrium. Of the 31 recently described (Reynolds et al., 2002) phytoplankton assemblages most of those were recognized that are likely to occur in shallow lakes. Separation of a functional group W _S from W2 for Synura dominated lakes is suggested. It seemed also necessary to raise a group (Y _Ph) for lakes dominated by Phacotus. Sorting of Dinophyta species into different already described functional groups is desirable.     

Phytoplankton assemblage at equilibrium in large and deep subalpine lakes: a case study from Lago Maggiore (N. Italy)

In 1996, we studied the phytoplankton seasonal succession in Lago Maggiore (N. Italy) through weekly sampling. Such a frequency enabled us to evaluate the changes of the phytoplankton assemblage in the light of the equilibrium and non-equilibrium theories. The distinct phases of changing of the species composition were identified separating the samples by means of cluster analysis and non-metric multidimensional scaling (NMDS). We recognised well distinct phytoplankton associations, whose seasonal succession followed a clear cyclic path throughout the year, with spring and summer phases respectively characterised by a rapid turn-over of the assemblages and by a relative stability. Moreover, we observed an increase of species number and Shannon-Wiener diversity during the spring, followed by a summer decline of the diversity in spite of an unchanged species number. Because of the dominance of the same few species for about two months during summer, coupled with small fluctuations of the total biomass, we could identify the summer assemblage as a steady state assemblage. The aim of the present contribution, although describing the whole seasonal succession, is to draw the attention towards the species composition at the steady state, taking into account the functional properties of the species involved. The possible role of the metalimnetic niche in selecting a particular summer assemblage in deep and large lakes will be discussed.     

金河湾城市湿地浮游植物功能类群演替及驱动因子

于2015年春(5月)、夏(8月)、秋(10月)三季,在金河湾湿地4类水体共设置12个采样点对浮游植物功能类群演替及与水环境变量关系进行分析。研究期间共鉴定浮游植物376个种,隶属于8门10纲19目19科101属。浮游植物种类组成主要以硅藻门(39.62%)和绿藻门(35.64%)为主,其次裸藻门(10.11%)和蓝藻门(9.84%)的藻类所占比例较高,甲藻门、隐藻门、金藻门和黄藻门所占比例较低。调查期间金河湾湿地浮游植物细胞丰度季节间差异显著(P0.05),整体上呈现夏季秋季春季的规律。春、夏、秋三季共划分20个不同的功能类群,双因素方差分析(Two-way ANOVA)和单因子交叉相似性检验(One-way ANOSIM)表明:代表性功能类群在季节间演替明显(P0.05),群落构成差异显著(P0.01)。SIMPER分析指出,S2/H1/B/D/Lo/X1/MP是引起金河湾湿地各季节之间浮游植物群落结构差异的主要贡献类群。通过代表性功能类群和10个水环境变量的典范对应分析(CCA)探索环境变量与功能类群演替的关系。经分析,总氮(TN)是驱动金河湾湿地浮游植物功能类群演替的主要环境变量,电导率(SpCond)、pH与功能类群演替密切相关。     

Spatial and temporal dynamics of the steady-state phytoplankton assemblages in a temperate shallow hypertrophic lake (Lake Manyas,Turkey)

Spatial and temporal dynamics of phytoplankton biomass and species composition in the shallow hypertrophic Lake Manyas, Turkey, were studied biweekly from January 2003 to December 2004 to determine steady-state phases in phytoplankton assemblages. Steady-state phases were defined when one, two or three coexisting species contributed to at least 80% of the standing biomass for at least 2 weeks and during that time the total biomass did not change significantly. Ten steady-state phases were identified throughout the study peiod. During those periods, Achnanthes microcephala (Kützing) Cleve twice dominated the phytoplankton biomass alone and contributed to more than 50% of the total biomass in seven phases. Microcystis aeruginosa (Kützing) Kützing, Anabaena spiroides Klebahn, Cyclotella stylorum Brightwell, Pediastrum boryanum (Turpin) Meneghini and Phacus pusillus Lemmermann were also represented once in steady-state phytoplankton assemblages. A. microcephala was dominant usually during cold periods of the year, while M. aeruginosa and A. spiroides were usually dominant in warm seasons. The total number of species showed a clear decrease during steady-state phases at all stations. All stations were significantly different in terms of the measured physical and chemical parameters (P < 0.05) and phytoplankton biomass (F = 117, P < 0.05).     

Phytoplankton equilibrium phases during thermal stratification in a deep subtropical reservoir

1. Equilibrium and non-equilibrium hypotheses have often been used to explain observations in community ecology. Published case studies have demonstrated that steady state phytoplankton assemblages are more likely to occur in deep lakes than in shallow mixed ones.
2. Phytoplankton seasonal succession was studied by weekly sampling in Faxinal Reservoir (S Brazil), a subtropical deep, clear, warm monomictic and slightly eutrophic reservoir. This study demonstrated an alternation of steady and non-steady state phases of phytoplankton assemblages with different dominant species during the steady states.
3. During the studied period, three steady states were identified with different dominant algal species: Anabaena crassa (Cyanobacteria), Nephrocytium sp. (green algae) and Asterionella (diatoms).
4. Each steady state in Faxinal Reservoir developed under stratified conditions of the water column according to the predictions of the disturbance concepts. Apparently, the major forces driving the development and persistence of these steady-state phases were closely related to thermal stratification and its consequences.
5. This study is the first report on development of more than one steady state within a year in a stratified water body. The development of three steady states might be the result of the relatively long stratification period in the Faxinal Reservoir and to its unique geochemical features.     

Continuous culture algal bioassays for organic pollutants in aquatic ecosystems

Short-term responses of phytoplankton to organic pollutants are highly transitory. Time-course studies of non-steady state cells in continuous culture showed varying growth or photosynthetic responses such as enchancement, inhibition, adaptation (or development of resistance) or rebound, depending on the direction of changes in the intracellular toxicant concentration and the duration of exposure. However, steady-state cells in a two-stage chemostat system exhibited an increased tolerance to toxicants and subtle physiological effects such as photosynthetic enhancement which was accompanied by a considerable leakage of photosynthesates. It is important to understand such steady-state responses for the prediction and assessment of ecological impact by organic pollution on phytoplankton, since the time scale of changes in the toxicant/biomass ratio in most natural waters is long enough to approximate an equilibrium state.     

Taxa-specific eco-sensitivity in relation to phytoplankton bloom stability and ecologically relevant lake state

Phytoplankton (including plant-like, animal-like algae and Cyanobacteria) blooms have recently become a serious global threat to the sustenance of ecosystems, to human and animal health and to economy. This study focused on the composition and stability of blooms as well as their taxa-specific ecological sensitivity to the main causal factors (especially phosphorus and nitrogen) in degraded urban lakes. The analyzed lakes were assessed with respect to the trophic state as well as ecological status. Total phytoplankton biomass (ranging from 1.5 to 181.3 mg dm⁻³) was typical of blooms of different intensity, which can appear during a whole growing season but are the most severe in early or late summer. Our results suggested that steady-state and non-steady-state bloom assemblages including mono-, bi- and multi-species or heterogeneous blooms may occur in urban lakes. The most intense blooms were formed by the genera of Cyanobacteria: Microcystis, Limnothrix, Pseudanabaena, Planktothrix, Bacillariophyta: Cyclotella and Dinophyta mainly Ceratium and Peridinium. Considering the sensitivity of phytoplankton assemblages, a new eco-sensitivity factor was proposed (E-SF), based on the concept of Phytoplankton Trophic Index composed of trophic scores of phytoplankton taxa along the eutrophication gradient. The E-SF values of 0.5, 1.3, 6.7 and 15.1 were recognized in lakes having a high, good, moderate or poor ecological status, respectively. For lake restoration, each type of bloom should be considered separately because of different sensitivities of taxa and relationships with environmental variables. Proper recognition of the taxa-specific response to abiotic (especially to N and P enrichment) and biotic factors could have significant implications for further water protection and management.     

The structure and dynamics of phytoplankton assemblages from the inner part of the Thermaikos Gulf,Greece. I. Phytoplankton composition and biomass from May 1988 to April 1989

Qualitative and quantitative analyses of phytoplankton from the inner part of Thermaikos Gulf were carried out in the period May 1988–April 1989. A total of 154 taxa were identified in the phytoplankton. the monthly values of the phytoplankton biomass ranged between 614 and 6700 mg/m³ (wet weight). The seasonal development of the phytoplankton biomass exhibited three peaks. Diatoms, dinoflagellates and cryptophytes were the most important constituents of the gulf phytoplankton. Throughout the year, five different phytoplankton assemblages were distinguished.     

Sunlight Effects on the Osmotrophic Uptake of DMSP-Sulfur and Leucine by Polar Phytoplankton

Even though the uptake and assimilation of organic compounds by phytoplankton has been long recognized, very little is still known about its potential ecological role in natural marine communities and whether it varies depending on the light regimes the algae experience. We combined measurements of size-fractionated assimilation of trace additions of ³H-leucine and ³⁵S-dimethylsulfoniopropionate (DMSP) with microautoradiography to assess the extent and relevance of osmoheterotrophy in summer phytoplankton assemblages from Arctic and Antarctic waters, and the role of solar radiation on it was further investigated by exposing samples to different radiation spectra. Significant assimilation of both substrates occurred in the size fraction containing most phytoplankton (>5 µm), sunlight exposure generally increasing ³⁵S-DMSP-sulfur assimilation and decreasing ³H-leucine assimilation. Microautoradiography revealed that the capacity to take up both organic substrates seemed widespread among different polar algal phyla, particularly in pennate and centric diatoms, and photosynthetic dinoflagellates. Image analysis of the microautoradiograms showed for the first time interspecific variability in the uptakes of ³⁵S-DMSP and ³H-leucine by phytoplankton depending on the solar spectrum. Overall, these results suggest that the role of polar phytoplankton in the utilization of labile dissolved organic matter may be significant under certain conditions and further confirm the relevance of solar radiation in regulating heterotrophy in the pelagic ocean.     

Functional groups of phytoplankton shaping diversity of shallow lake ecosystems

Phytoplankton of eutrophic shallow lakes are frequently dominated by one species or species of the same functional group, resulting in species-pure algal assemblages. Knowledge of the structure of these assemblages is essential to understand their functioning; therefore, species and functional diversity were investigated in five sub-types of eutrophic shallow lake. Among the sub-types, astatic saline lakes and hypertrophic ponds had type-specific assemblages dominated by S_N and W0, W1 codons. The diversity of the phytoplankton in the sub-types was quite similar, except for the astatic saline lakes, which were characterised by lower values of both functional and species diversity. We found that both functional and species diversity were low when bloom-forming cyanobacteria (H1, S_N functional groups) became dominant. Dominance of other groups (J, Y, L_O and W1) did not coincide with decrease in species diversity. Analysis of the biovolume versus diversity relationships revealed that decrease in diversity might be expected at biovolume >20 mm³ l⁻¹ for shallow lakes.     

Phytoplankton strategies and diversity under different nutrient levels and planktivorous fish densities in a shallow Mediterranean lake

Two mesocosm experiments were carried out to investigate thedynamic ef fects of nutrients (nitrogen and phosphorus) andplanktivorous fish additions on phytoplankton strategies anddiversity. The phylogenetic and functional approaches were usedto understand phytoplankton ecology in shallow Mediterraneanlakes. The experimental approach is new for the study of algalfunctional groups. Nutrient loading and fish stocks enhancedbiomass of small algae but decreased phytoplankton diversityand species richness. Faster species replacement and fluctuationsin diversity occurred above loadings of 1 µM P and 21µM N. Mesotrophic conditions favoured a diverse pool ofspecies, including nostocales and unicellular flagellate algae(functional groups S_n, S₁, L₀, Y, Reynolds et al., 2002). C-strategistchlorophytes (small algae from functional group X₁) dominatedmid-successional assemblages with good light and accessiblenutrients. High nutrient concentrations, dim light, presenceof organic matter and of larger zooplankton favoured to functionalgroups S₁ of oligophotic filamentous cyanobacteria and J ofmixotrophic Scenedemus species. Intermediate nutrient levelswith total phosphorus (TP) 10 µM, water quiescence, transparencyand smaller zooplankton prompted dominance of chroococcal cyanobacteria(functional groups X₁ and K). Resulting patterns agree and reinforcethe validity of plankton functional groups associated with warm,shallow enriched systems, although some changes in the groupsare suggested in relation to the structuring role of nutrientsand grazing on the functional scheme for phytoplankton.     

Controlling factors of phytoplankton assemblages in wetlands: an experimental approach

The aim of this work is to answer some questions like: what factors control the phytoplankton assemblage? What factor or factors are perturbing the assemblage? What factors are driving or maintaining the stability? Are the different responses to the control factors dependent on the hierarchy level? For that, we tested experimentally the influence of herbivory, planktivory, nutrients and sediment on phytoplankton assemblages and its stability from a hypertrophic wetland (Las Tablas de Daimiel National Park, Spain) in three microcosm experiments. The study of the steady state phytoplankton assemblages in this perturbed system could point out some underlying processes instead of competition. The presence of planktivorous fish and the different composition of zooplankton have minor importance in phytoplankton composition. Conversely, sediment is of paramount importance, promoting a more diverse phytoplankton assemblage. When sediment or nutrient pulses are absent, phytoplankton become dominated by slow-growing algae, present but not dominant in the studied wetland community. We suggest that alternate states of phytoplankton assemblages in a eutrophic wetland occur as a sequence of substitutions persistently altered by perturbations, thanks to the close coupling with sediment, and that possibly trophic relationships are irrelevant.     

Coupling of phytoplankton and detritus in a shallow,eutrophic lake (Lake Loosdrecht,The Netherlands)

An oscillating steady state is described of phytoplankton, dominated by Prochlorothrix hollandica and Oscillatoria limnetica, and sestonic detritus in shallow, eutrophic Lake Loosdrecht (The Netherlands). A steady-state model for the coupling of the phytoplankton and detritus is discussed in relation to field and experimental data on phytoplankton growth and decomposition. According to model predictions, the phytoplankton to detritus ratio decreases hyperbolically at increasing phytoplankton growth rate and is independent of a lake's trophic state. The seston in L. Loosdrecht contains more detritus than phytoplankton as will apply to many other lakes. The model provides a basis for estimating the loss rate of the detritus, including decomposition, sedimentation and hydraulic loss. In a shallow lake like L. Loosdrecht detritus will continue to influence the water quality for years.     

Analysis of candidate gene expression patterns of adult male <Emphasis Type="Italic">Macrobrachium rosenbergii</Emphasis> morphotypes in response to a social dominance hierarchy

Because phytoplankton communities exhibit seasonal patterns driven by changes in physical factors, grazing pressure, and nutrient limitations, climate change, in combination with local phosphorus management policies are expected to impact phytoplankton annual dynamic. We used long-term monitoring data from Lake Geneva (from 1974 to 2010) to test if changes in phytoplankton seasonal succession across years is related to re-oligotrophication, inter-annual variability in thermal conditions, and Daphnia sp. density. We used a Bayesian method to identify species assemblages and wavelet analysis to detect transient dynamics in seasonal periodicity. A decrease in phosphorus concentrations appeared to play a major role in the inter-annual replacement of species assemblages. Furthermore, some species assemblages exhibited a change in their seasonal periodicity that was most likely induced by changes in Daphnia sp. density. Finally, we demonstrated that flexibility in the pattern of phytoplankton seasonal successions played a stabilizing role at the community level. The results suggest that phenology and inter-annual changes in seasonal dynamics of phytoplankton assemblages are important components to consider for explaining long-term variability in phytoplankton community.     

2012年夏季挪威海和格陵兰海浮游植物群落结构的色素表征

对2012年中国第5次北极科学考察期间的挪威海和格陵兰海两个断面的光合色素进行了高效液相色谱（HPLC）分级分析,通过藻类色素化学分类分析软件（CHEMTAX）获得了不同浮游植物类群对叶绿素a的贡献,进而得到该海域表层和次表层（30 m）的浮游植物群落结构。结果表明：表层总叶绿素a的浓度为23.59 ng/L,低于次表层的30.38 ng/L,其中浮游植物根据粒径划分对总叶绿素a的贡献由高到低依次是微型浮游植物、小型浮游植物和微微型浮游植物。该海域同时存在葱绿叶绿素（Prasino）、墨角藻黄素（Fuco）、别藻黄素（Allo）、多甲藻素（Perid）、玉米黄素（Zea）、19-丁墨甲藻黄素（19’BF）和19-六已墨甲藻黄素（19’HF）等色素,其浓度和分布与温盐和营养盐等环境因子存在一定的相关性。不同粒径浮游植物色素组成显示,微微型浮游植物群落中以S型定鞭藻（28%）、N型定鞭藻（21%）、硅藻（18%）和青绿藻（12%）占优;微型浮游植物群落的优势类群为S型定鞭藻（53%）、N型定鞭藻（20%）和硅藻（12%）;而小型浮游植物群落主要为硅藻（63%）和甲藻（17%）。     

Phytoplankton assemblages in twenty-one Sicilian reservoirs: relationships between species composition and environmental factors

Data collected in a limnological survey, carried out between 1987 and 1988 on 21 Sicilian reservoirs of varying trophic state, were ordinated using CANOCO 3.1 to generalise the way in which the structure of phytoplankton assemblage is conditioned by both physical and chemical variables. The results showed that in these man-made lakes, characterised by conspicuous water-level fluctuations, the annual and interannual variability in the abundance and composition of phytoplankton may be strongly influenced by their peculiar hydraulic regimes rather than by nutrient availability. In particular, it was highlighted that, from the early summer, water abstraction often leads to increased circulation and to the deepening of the mixed layer. In this way, an increase of the ratio of mixing depth to euphotic depth is forced, with the result that phytoplankton cells experience longer periods in darkness as they are carried through the mixed layer. Phytoplankton assemblages change in species composition in response to the environmental variation. Both the raising of the trophic state, with an increase in phytoplankton biomass and a decrease in transparency, and the intensified abstraction enhance the role of light availability in promoting the development of specific phytoplankton assemblages adapted to the modified physical environment. Light climate is an important influence on the species structure of the phytoplankton, especially in the higher part of the trophic gradient. In contrast, the influence of nutrients on the structure of the assemblages appears to be higher in the lower part of the trophic spectrum or in those environments characterised by a higher hydrological stability during the year.     

Changes in phytoplankton,benthic invertebrate and fish assemblages of boreal lakes following invasion by Gonyostomum semen

1. During recent decades, Gonyostomum semen populations have spread in northern temperate regions forming dense blooms that may dominate the phytoplankton assemblage for extended periods. In this study, we investigate the effects of G. semen blooms in boreal brown water lakes with special emphasis on phytoplankton, fish and benthic invertebrate assemblages using data from 10 boreal lakes sampled annually over a 10‐year period. 2. Significant differences in phytoplankton and benthic invertebrate assemblages were found between lakes with high (3.01 mm³ L^?1; >80% phytoplankton biomass) and lakes with low G. semen biomass (0.03 mm³ L^?1; <5% phytoplankton biomass). In particular, high G. semen lakes had lower biomass of smaller, edible phytoplankton and a higher abundance and biomass of benthic invertebrates, especially Chaoborus flavicans, and perch than low G. semen lakes. 3. The length distribution of fish also suggested a tendency towards large and older fish and a lower recruitment success in high G. semen lakes, as denoted by lower abundances of fish shorter than 10 cm and higher biomass and abundance of fish longer than 15 cm in high G. semen lakes. 4. This study shows that high G. semen lakes are characterised by less edible phytoplankton, dominance by a few species and enhanced benthic secondary and fish production. Hence, the conjecture that high biomasses of G. semen create a bottleneck in the energy transfer to higher trophic levels seems less likely in boreal lakes.