The generation of phytoplankton patchiness by mesoscale current patterns

 Elken et al. (1994) suggested that phytoplankton patchiness can be generated by mesoscale eddies in light-limited, nutrient-replete environments. This hypothesis is explored using two ecological models of different physical complexity. The model results support the idea that the coupling of mesoscale eddy circulation and phytoplankton growth leads to differential growth rates and thus generates variability in phytoplankton distributions. The specific circulation of a cyclonic eddy isolates a phytoplankton population in its core. Due to the reduced vertical mixing, a higher growth rate is supported in the core, and phytoplankton concentrations increase compared to the surrounding environment. A one-dimensional model is used to explore the hypothesis in general and to perform sensitivity studies. A more realistic simulation uses a coupled three- dimensional model for the western Baltic Sea. Starting from vertically and horizontally homogeneous distributions for nutrients and plankton, the models generate patchiness due to the proposed mechanism. The described mechanism may apply for other mesoscale variable environments during light-limited growth periods as well, e.g., the frontal region of the Southern Ocean. Received: 31 March 2001 / Accepted: 31 August 2001     

Modeling spring-summer phytoplankton bloom in Lake Michigan with and without riverine nutrient loading

There were two phytoplankton blooms captured by remote sensing in Lake Michigan in 1998, one from March to May, and one during June. In this paper, those phytoplankton blooms were simulated by a coupled physical–biological model, driven by observed meteorological forcing in 1998. The model reasonably reproduced the lake currents. The biological model results, with and without riverine nutrient loading, were compared with the remote sensing data. A 3-month-long donut-like phytoplankton bloom that appeared in southern Lake Michigan was reasonably well simulated only when riverine input was included, indicating the importance of riverine nutrient input for supporting the growth of phytoplankton in Lake Michigan. The model with riverine input also captured a second event-driven phytoplankton bloom during June with weaker magnitude that occurred in mid-south Lake Michigan, which lasted for about 20 days. The major reason for the weaker bloom in June was that vertical mixing in the hydrodynamic model was too weak (leading to a mixed-layer depth of 20 m) to bring the bottom nutrient-rich water up to the epilimnion. High chlorophyll concentration that persisted in Green Bay for almost a year was simulated with less intensity.     

The nonlinear effects of the eddy viscosity and the bottom friction on the Lagrangian residual velocity in a narrow model bay

The nonlinear effects of the eddy viscosity and the bottom friction on the Lagrangian residual velocity (LRV) are studied numerically in a narrow model bay. Three groups of the experiments with different eddy viscosity and different forms of the bottom friction are designed and are carried out in the three kinds of the topography. When the eddy viscosity is obtained from a two-equation turbulence closure model, the pattern of the LRV is more complex than that of the time invariant eddy viscosity case and the intensity is from more than 1.3 times to one order smaller than that of the linear eddy viscosity condition. The LRV are also acquired when the eddy viscosity varies from the flood-averaged one to the ebb-averaged one. It is found that when the flood-averaged eddy viscosity is bigger than the ebb-averaged eddy viscosity (flood-dominated asymmetry), the direction of the breadth-averaged LRV and the 3D LRV is nearly opposite to that when the eddy viscosity asymmetry is reverse (ebb-dominated asymmetry). However, the intensity of the LRV for the ebb-dominated case decreases toward the flood-dominated case as the ratio of the maximum depth in the deep channel and the minimum depth on the shoal increases. The different forms of the bottom friction also play a role in the LRV. The structures of the 3D LRV and the depth-integrated LRV are simpler, and the intensity of the LRV is two times smaller when the linear bottom friction is used than those when the quadratic bottom friction is used.     

Initial growth of phytoplankton in turbid estuaries: A simple model

An idealised model is presented and analysed to gain more fundamental understanding about the dynamics of phytoplankton blooms in well-mixed, suspended sediment dominated estuaries. The model describes the behaviour of subtidal currents, suspended sediments, nutrients and phytoplankton in a channel geometry. The initial growth of phytoplankton and its spatial distribution is calculated by solving an eigenvalue problem. The growth rates depend on the position in the estuary due to along-estuary variations in nutrient concentration and suspended sediment concentration. The model yields an insight into how the onset of blooms in the model depends on physical and biological processes (turbulent mixing, fresh water discharge, light attenuation, imposed nutrient concentrations at the river and sea side). In particular, the model demonstrates that the joint action of spatial variations in turbidity and in nutrients causes the maximum phytoplankton concentrations to occur seaward of the estuarine turbidity maximum.     

新安江水库(千岛湖)湖泊区夏季热分层期间垂向理化及浮游植物特征

2010年7月对亚热带特大型水库——新安江水库湖泊区水体的垂向物理、化学参数以及浮游植物群落进行了观测研究,并应用Water-PAM对水体浮游植物垂向光合作用参数进行了测定.研究结果表明:夏季该水库湖泊区在水下10～20 m处形成明显的温跃层,垂向pH值、溶解氧及浊度的变化同叶绿素a浓度呈现高度一致;夏季浮游植物群落以硅藻占绝对优势,水体表层以梅尼小环藻(Cyclotella meneghiniana)为主,表层以下其它各层均以巴豆叶脆杆藻(Fragilaria cro-tonensis)为绝对优势种,垂向分布表现为5～10 m区间为浮游植物高密度区域,温跃层以下浮游植物密度显著下降,水温分层可能是决定浮游植物垂向分布的重要因素之一.浮游植物最大光合效率从表层向下层逐步降低,实际光合效率最大值出现在垂向10 m区域.     

鄱阳湖浮游植物时空变化特征及其对极端洪枯事件的响应

鄱阳湖作为中国最大的淡水湖泊,其水生态健康状态始终是人们关注的热点。近些年,鄱阳湖极端洪旱灾害频发,浮游植物生长受极端洪旱的影响发生了明显变化。为分析浮游植物时空变化特征、探究环境因子对鄱阳湖浮游植物影响机制以及极端洪枯事件对浮游植物的影响,利用结构方程模型(SEM)构建浮游植物与环境因子的影响路径模型,定量分析环境因子对浮游植物的影响程度。结果表明,鄱阳湖浮游植物以蓝藻、绿藻为主且有明显的季节特征,在7月丰水期浮游植物密度达到最高;由结构方程模型(SEM)可知,影响浮游植物密度最关键的因子为物理因子(水温>pH>透明度>溶解氧),其次为营养物质(总氮>硝态氮>总磷>磷酸盐),浮游植物对高温、高营养和高pH较偏好。2020年极端洪水和2022年极端高温干旱,浮游植物密度主要受水温、溶解氧、透明度等物理因子的影响;在影响较小的营养物质中,主要的限制性因素分别为磷和氮。与正常年份相比,极端洪水年鄱阳湖受入湖来水及降雨的增多,湖区水量急剧增加,“稀释”作用超过水温和透明度对浮游植物生长的促进作用造成浮游植物密度和生物量有所下降;在营养物质中,磷成为主要限制性因素。而极端高温干旱年受入湖来水的减少及湖水的快速蒸发,“浓缩”作用超过水温和透明度对浮游植物生长的抑制作用造成浮游植物密度和生物量显著增加,同时,营养物质对浮游植物的作用更加凸显。研究结果表明极端洪枯事件导致鄱阳湖浮游植物变化明显,确定其对浮游植物的影响机制,可以为极端洪枯事件下浮游植物监测、管理提供一定的理论基础。     

Ammonium,nitrate and phytoplankton interactions in a freshwater tidal estuarine zone: potential effects of cultural eutrophication

Nitrate and ammonium are the most important nitrogen sources for phytoplankton growth. Differential utilization of inorganic nitrogenous compounds by phytoplankton has been observed and may have significant impacts on primary productivity at local scales. We used enrichment experiments with natural phytoplankton populations from the freshwater tidal zone of the Guadiana estuary, a coastal ecosystem increasingly subjected to anthropogenic influences, to study the effects of nitrate and ammonium on N-consumption and phytoplankton growth. In addition, we used combined additions of nitrate and ammonium to understand the inhibitory effect of ammonium over nitrate uptake. Ammonium concentrations in the freshwater tidal reaches of the Guadiana estuary throughout the sampling period were too low to exert an inhibitory effect on nitrate uptake or a toxic effect on phytoplankton growth. Nitrate was clearly the main nitrogen source for phytoplankton at the study site. Overall, nitrate seemed to become limiting at concentrations lower than 20 μM and N-limitation was particularly significant during summer. A trend of decreasing nitrate uptake with increasing ammonium concentrations and uptake suggested an overall preference for ammonium. However, preference for ammonium was group-specific, and it was observed mainly in green algae and cyanobacteria. In fact, cyanobacteria relied only on ammonium as their N-source. On the contrary, diatoms preferred nitrate, and did not respond to ammonium additions. The increasing eutrophication in the Guadiana estuary and particularly increased inputs of nitrogen as ammonium due to urban waste effluents may result in a shift in phytoplankton community composition, towards a dominance of cyanobacteria and green algae.     

Hydrodynamic-phytoplankton model for short-term forecasts of phytoplankton in Lake Taihu, China

Phytoplankton biomass is an important factor for short-term forecasts of algal blooms. Our new hydrodynamic-phytoplankton model is primarily intended for simulating the spatial and temporal distribution of phytoplankton in Lake Taihu within a time frame of 1-5 days. The model combines two modules: a simple phytoplankton kinetics module for growth and loss; and a mass-transport module, which defines phytoplankton transport horizontally with a two dimensional hydrodynamic model. To adapt field data for model input and calibration, we introduce two simplifications: (a) exclusion of some processes related to phytoplankton dynamics like nutrient dynamics, sediment resuspension, mineralization and nitrification, and (b) use of monthly measured data of the nutrient state. Chlorophyll-α concentration, representing phytoplankton biomass, is the only state variable in the model. A sensitivity analysis was carried out to identify the most sensitive parameter set in the phytoplankton kinetics module. The model was calibrated with field data collected in 2008 and validated with additional data obtained in 2009. A comparison of simulated and observed chlorophyll-α concentration for 33 grid cells achieved an accuracy of 78.7%. However, mean percent error and mean absolute percent error were 13.4% and 58.2%, respectively, which implies that further improvement is necessary, e.g. by reducing uncertainty of the model input and by an improved parameter calibration.     

Multi-nest high-resolution model of submesoscale circulation features in the Gulf of Taranto

Recent oceanographic field measurements and high-resolution numerical modelling studies have revealed intense, transient, submesoscale motions characterised by a horizontal length scale of 100–10,000 m. This submesoscale activity increases in the fall and winter when the mixed layer (ML) depth is at its maximum. In this study, the submesoscale motions associated with a large-scale anticyclonic gyre in the central Gulf of Taranto were examined using realistic submesoscale-permitting simulations. We used realistic flow field initial conditions and multiple nesting techniques to perform realistic simulations, with very-high horizontal resolutions (> 200 m) in areas with submesoscale variability. Multiple downscaling was used to increase resolution in areas where instability was active enough to develop multi-scale interactions and produce 5-km-diameter eddies. To generate a submesoscale eddy, a 200-m resolution was required. The submesoscale eddy was formed through small-scale baroclinic instability in the rim of a large-scale anticyclonic gyre leading to large vertical velocities and rapid restratification of the ML in a time-scale of days. The submesoscale eddy was confirmed by observational data from the area and we can say that for the first time we have a proof that the model reproduces a realistic submesoscale vortex, similar in shape and location to the observed one.     

Hydrodynamic conditions associated with the formation,maintenance and dissipation of a phytoplankton thin layer in a coastal upwelling system

During May–June 2005, a 17-d cruise was carried out in Ría de Pontevedra (Galician Rías Baixas) to study the physical–biological interactions that may lead to subsurface aggregations of phytoplankton organisms in thin layers (TLs). Physical processes governed the initiation and development, maintenance, and decline of a diatom (toxin producing Pseudo-nitzschia spp. and Chaetoceros socialis) TL during an upwelling relaxation-upwelling–downwelling sequence. Differences in shear profiles appeared to lead to the formation of a TL during upwelling events. These results reveal that the coupling between maximum values of shear and buoyancy frequency can shape a subsurface chlorophyll maximum (SCM) into a TL. The effect of shear upon phytoplankton patches, which has been predicted on the basis of theoretical studies, has been corroborated in this study in which the vertical distribution of an observed TL was controlled by physical processes.     

Light climate as the key factor controlling the spring dynamics of phytoplankton in Lake Zürich

During the spring seasons of 1983, 1986 and 1987 the development of phytoplankton in Lake Zürich was investigated (from February to May) using samples taken at short term intervals. The aim was to describe the effects of the short term dynamics of environmental factors on the algal growth. The results could then be used to discuss the existing theories to assess the start of phytoplankton growth pulses in spring. Only 7 to 10 days without wind driven vertical mixing were required in spring to start the first growth pulse, despite of a still very unstable water column (sometimes inverse thermal stratification). Mainly flagellates andStephanodiscus hantzschii increased their biomass and achieved net growth rates of 0.1 and up to 0.65 d⁻¹ respectively. During such a phase the mixing depth was always smaller than the euphotic depth. Later on, at the start of the spring bloom (=last growth pulse in spring before the clear water stage), the intensity of vertical mixing as well as the mixing depth were markedly reduced due to an increase in heat input and low wind. Then flagellates dominated (contribution up to 75.5% of the areal biomass reaching 60 g fresh weight m⁻²) and the growth rate rose to a maximum of 0.65 d⁻¹. Standard models of critical depth considers that there is only a biomass increase if the mixing depth is smaller than the depth of a water layer positive balanced between production and respiration. This model for determining the beginning of a phytoplankton growth pulse in spring takes no account of the favorable light conditions for phytoplankton cells at calm and sunny days in February and March. The newly developed threshold value model takes these situations into account: It assumes that the phytoplankton biomass increases when the calculated effective light climate is equal or greater than a previously fixed threshold. The calculations are based on the mean light intensity within the mixed layer at windy days or within the euphotic depth (z _eu) at calm days. In Lake Zürich a minimum of 0.2 10⁶ J m⁻²d⁻¹ (=0.9 mol quanta m⁻²d⁻¹) has to be reached or surpassed in at least 3 days before an exponential increase of algal biomass can occur. The value does not depend on short term fluctuations in neither radiation nor mixing depth. It seems that this value is rather low comparing with those of investigations in other water bodies (up to 0.8 10⁶ J m⁻² d⁻¹) but high related to values from algal cultures (0.02 10⁶ J m⁻²d⁻¹). As the weather can only be forecasted a few days ahead with any certainty the period for a more or less accurate prediction of an algal bloom is restricted to about 1 to 5 days.     

Long-term changes in physical and chemical conditions of nutrient-poor lakes along a latitudinal gradient: is there a coherent phytoplankton community response?

To evaluate climate and atmospheric deposition induced physical and water chemical changes and their effects on phytoplankton communities, we used complete time series (14 years, monthly measurements during the growing season) of 18 physical and chemical variables and phytoplankton data from 13 nutrient-poor Swedish reference lakes along a latitudinal gradient. We found numerous strong significant changes over time that were most coherent among lakes for sulfate concentrations, conductivity, calcium, magnesium, chloride, potassium, water color, surface water temperature and the intensity of thermal stratification. Despite these pronounced coherent physical and water chemical changes over Sweden, the phytoplankton biomass and species richness of six phytoplankton groups, measured at the same time as the water chemical variables, showed only few and weak significant changes over time. The only coherent significant change over Sweden, occurring in seven lakes, was observed in the species richness of chlorophytes. The number of chlorophyte taxa significantly declined over Sweden. Using a partial least square model for each lake, we attributed the decline primarily to an increase in water temperatures and water color, which were among the most important variables for the model performance of each lake. All other taxonomic groups were driven primarily by non-coherent changes in nutrient concentrations, pH and probably also non-coherent grazing pressure. We concluded that coherent phytoplankton responses can only be achieved for taxonomic groups that are driven primarily by coherent physical/chemical changes. According to our study, chlorophytes belong to such a group, making them possible global change indicators. Our findings give new insights into global change effects on different phytoplankton taxonomic groups in nutrient-poor lakes.     

Cosmic ray modulation during the solar activity growth phase of cycle 24

Recent years allowed us to study long-term variations in the cosmic ray (CR) intensity at an unusually deep solar activity (SA) minimum between cycles 23 and 24 and during the SA growth phase in cycle 24, which was the cycle when SA was the lowest for the epoch of regular ground-based CR observations since 1951. The intensity maximum, the value of which depends on the particle energy, was observed in CR variations during the period of an unusually prolonged SA minimum: the CR density during the aformentioned period (2009) is higher than this density at previous CR maxima in cycles 19–23 for low-energy particles (observed on spacecraft and in the stratosphere) and medium-energy particles (observed with neutron monitors). After 2009 CR modulation at the SA growth phase was much weaker over three years (2010–2012) than during the corresponding SA growth periods in the previous cycles. The possible causes of this anomaly in CR variations, which are related to the CR residual modulation value at a minimum between cycles 23 and 24 and to variations in SA characteristics during this period, were examined. The contribution of different solar magnetic field characteristics and indices, taking into account sporadic solar activity, has been estimated.     

Abiotic controls of potentially harmful algal blooms in Santa Monica Bay,California

Despite the increasing occurrence of harmful phytoplankton blooms along the North American west coast, records of phytoplankton populations and related environmental conditions are uncommon. In this study, twice monthly measurements in the upper 50 m are used to assess physico-chemical conditions contributing to the growth of potentially harmful bloom taxa over two annual cycles (2004–2005) in the Santa Monica Bay, California. Results were compared to the predictions of the Intaglio model [Smayda, T.J., Reynolds, C.S., 2001. Community assembly in marine phytoplankton: application of recent models to harmful dinoflagellate blooms. Journal of Plankton Research 23, 447–461.] of phytoplankton community assembly. Potentially harmful taxa were present in every surface sample and were numerically dominant during the largest observed blooms, contributing up to 92% of the total phytoplankton abundance >5 μm. Large interannual variation was observed in the dominant taxa and bloom seasonality; Pseudo-nitzschia sp. dominated blooms in early 2004 (February and April), whereas Prorocentrum micans and Lingulodinium polyedrum blooms occurred in May and September of 2005, respectively. The Pseudo-nitzschia sp. blooms were associated with elevated nitrate, dissolved silicon and phosphate concentrations throughout the euphotic zone; the first bloom followed a strong upwelling and the second occurred during the onset of seasonal stratification. In contrast, the blooms of P. micans were associated with highly stratified, low nutrient waters. Multivariate analysis supports the roles of temperature, mixed-layer depth and nutrient concentrations as primary controls of bloom growth, following the conceptual Intaglio model. The strong presence of potentially harmful bloom species in the Santa Monica Bay during this study appears unusual in comparison to limited studies over the last several decades.     

砷污染治理背景下阳宗海浮游植物生物量的时空分布模式及驱动因子

在社会经济发展和流域开发持续的背景下,砷污染已成为我国部分水体面临的重要环境问题,目前对砷污染防控的生态效应与修复效果评价仍缺乏系统识别。湖泊生态系统中浮游植物是重要的生产者,砷等重金属污染可以直接影响浮游植物生长、物种演替和初级生产力水平,浮游植物已成为指示砷污染水平及其生态效应的敏感指标。本研究以长期受到砷污染胁迫并经历污染治理的阳宗海为研究对象,设置南、中、北3个调查位点,于2015年4月-2019年12月对浮游植物和水质因子开展季节调查和空间分析,通过识别浮游植物生物量的时空分布模式与驱动因子,评价了砷污染与治理下浮游植物生物量的变化机制和生态修复效果。调查结果显示,采样期间阳宗海浮游植物以蓝藻门为主,浮游植物的生物量范围为0.7~30.4 mg/L,平均生物量在2016年最低((3.0±1.8) mg/L)、在2017年最高((10.5±8.9) mg/L)。ANOVA分析结果显示,浮游植物生物量存在显著的季节差异而空间差异不明显。相关分析结果显示,阳宗海浮游植物生物量与砷浓度和透明度呈显著负相关,而与水体温度和pH呈显著正相关。多元线性回归分析进一步显示,砷和水温是驱动阳宗海浮游植物生物量变化的显著环境因子。由此可见,在重金属污染湖泊经过修复后,水体砷遗留物的毒性效应仍然对浮游植物生长产生了明显的抑制作用,表明了水体重金属污染物可能具有长期的沉积物释放作用与持久的生态毒理效应。     

Primary production in a turbid estuary, the Ems-Dollard: field and model studies

The Ems-Dollard Estuary has a steep axial gradient in turbidity which strongly regulates primary production of phytoplankton and microphytobenthos. A combination of the turbidity and nutrient gradients causes a high pelagic primary production in the outer, seaward part of the estuary, and a high benthic production on high-level mudflats (with long effective photoperiods) in the inner part. Model simulations were used to study phytoplankton succession and the factors influencing this succession. Light limitation was evaluated in a sensitivity run which used a lower compensation light intensity.     

The use of satellite data to characterize phytoplankton in Sea of Okhotsk water

For the identification of natural processes taking place in surface water layer in the Sea of Okhotsk and nearby water areas, satellite data were used to determine seasonal variations of chlorophyll “a” concentration and the problem of water typification by satellite data in the visible light range was solved. Analysis of the annual cycle of phytoplankton concentration variations confirmed the presence of a short explosion in phytoplankton population in the southern Sea of Okhotsk during ice cover formation, which had been predicted before by model calculations. Classes of water reflecting different geoenvironmental situations were identified for characteristic regions of the Sea of Okhotsk. The dependences of chlorophyll “a” concentration growth on temperature were calculated, and corrections were introduced in the standard relationships used in biooptic algorithms for different parts of the Sea of Okhotsk.     

基于15N稳定同位素示踪技术的太湖梅梁湾浮游植物群落氮吸收动力学研究

浮游植物对氮的吸收与其生长繁殖密切相关,太湖梅梁湾湖区蓝藻水华频频暴发,对该水域浮游植物氮吸收进行研究具有重要意义.本文分别在冬、春、夏、秋4个季节于梅梁湾采样,对水体常规理化指标和浮游植物群落结构进行分析,并利用¹⁵N稳定同位素示踪技术研究了浮游植物对铵态氮（NH₄⁺-N）、硝态氮（NO₃^--N）和尿素态氮（Urea-N）吸收的动力学特征.结果表明,太湖梅梁湾浮游植物群落除了秋季对NH₄⁺-N的吸收不符合米氏方程外,其余均符合.冬季和春季3种形态氮最大吸收速率（V_max）的大小依次为：NH₄⁺-N > NO₃^--N > Urea-N,而夏季为：NH₄⁺-N > Urea-N > NO₃^--N.3种形态氮V_max的季节变化规律为夏季 > 秋季 > 春季 > 冬季.V_max在不同季节以及不同形态氮之间的差异性可能与浮游植物群落组成以及水体中NH₄⁺-N浓度不同有关.浮游植物对NH₄⁺-N吸收的K_S值在冬、春季高于夏季,对Urea-N吸收的K_s值则在夏、秋季高于冬、春季,而对NO₃^--N吸收的K_s值则在夏季显著高于其他3个季节.冬季和春季梅梁湾浮游植物群落最容易受到NO₃^--N限制,而最不容易受到Urea-N的限制;而夏季,则最容易受到NO₃^--N限制,而最不容易受到NH₄⁺-N的限制,且浮游植物群落对NH₄⁺-N的亲和力最高.与NO₃^--N相比,秋季浮游植物更容易受到Urea-N的限制.不同季节,容易对浮游植物产生限制作用的氮的形态不同.     

Decadal variability of the Kuroshio Extension: mesoscale eddies and recirculations

An eddy-resolving multidecadal ocean model hindcast simulation is analyzed to investigate time-varying signals of the two recirculation gyres present respectively to the north and south of the Kuroshio Extension (KE) jet. The northern recirculation gyre (NRG), which has been detected at middepth recently by profiling float and moored current meter observations, is a major focus of the present study. Low-frequency variations in the intensity of the recirculation gyres are overall highly correlated with decadal variations of the KE jet induced by the basin-wide wind change. Modulation of the simulated mesoscale eddies and its relationship with the time-varying recirculation gyres are also evaluated. The simulated eddy kinetic energy in the upstream KE region is inversely correlated with the intensity of the NRG, consistent with previous observational studies. Eddy influence on the low-frequency modulation of the NRG intensity at middepth is further examined by a composite analysis of turbulent Sverdrup balance, assuming a potential vorticity balance between the mean advection and the convergent eddy fluxes during the different states of the recirculation gyre. The change in the NRG intensity is adequately explained by that inferred by the turbulent Sverdrup balance, suggesting that the eddy feedback triggers the low-frequency modulation of the NRG intensity at middepth.