Catastrophic transition in vertical distributions of phytoplankton: alternative equilibria in a water column

We showed a catastrophic transition between a surface maximum pattern and a subsurface maximum pattern of phytoplankton in a water column by a mathematical model considering the vertical distribution of phytoplankton and two resources, light and a limiting nutrient. In our model, we assumed that a water column consists of two layers: a complete mixing layer above a seasonal thermocline and an incomplete mixing layer below it. From numerical calculation of the model, we obtained that there are two stable vertical patterns of phytoplankton over a certain range of parameters of the model: a pattern having its maximum below the thermocline and another having its maximum above the thermocline. As other models having multiple stable equilibria, our model also exhibits a hysteresis effect and catastrophic transition when one of the parameters of the model changes continuously. These results indicate the possibility of the existence of alternative equilibria of vertical patterns of phytoplankton even if the trophic status and physical condition of the water column are similar. Moreover, the catastrophic transition between the steady states suggests one of the possible mechanisms of autumn algal blooms.     

Seasonal cycle of nitrogen and phytoplankton biomass in a well-mixed coastal system (Eastern English Channel)

The hydrological structure of the French coastal part of the eastern English Channel is strongly linked with tidal regimes and riverine input. Two distinct water masses are separated by a frontal area and drift along the coast in SW–NE direction. These two water masses are well-mixed during the entire year. We studied the seasonal dynamic of nitrogenous nutrients, chlorophyll a and organic particulate carbon and nitrogen at two stations, characteristic of these water masses, during the year 1994. Results show (i) a winter stock of nitrate and ammonium, (ii) a pre-bloom period corresponding to the use of ammonium, (iii) a high bloom period of short duration using nitrate, (iv) a post-bloom period with little phytoplanktonic activity probably limited by nutrients other than nitrogen and (v) an autumnal period of reconstitution of stock. The essential difference between the two stations is the importance of winter stock of nutrients and of bloom chlorophyll a concentration, with the coastal station richer than the offshore one. An assumption about the nitrogen available for new production in this area gives a value of 57% of the winter stock of inorganic nitrogen.     

The effect of water column mixing on phytoplankton succession, diversity and similarity

The lake number was used to describe the mixing condition forthree consecutive years (1992–1994) in Lake Tegel, Berlin,and compared to the successions of diatoms, dinoflagellatesand cyanobacteria, the main phytoplankton groups in the lake,as well as to diversity and similarity indices. Using both diversityand similarity indices in juxtaposition provides an indicationof the growth type of prevailing species (r- or K-strategists)and the degree of competition in the aqua-ecological system.A genera] pattern of these indices can be recognized as threephases: (i) high diversity—during spring, summer and autumn,interrupted by (ii) phases of low diversity during the latespring clear-water phase as the number of spring species plummeted,and (iii) during the late summer, climax populations of K-strategists.On a smaller time scale, similarity and diversity proved toreact sensitively to disturbances at frequencies intermediatein relation to the generation times of the phytoplankton. Thissupports the ‘intermediate disturbance hypothesis’,as proposed for phytoplankton by Padisak et al. [(eds) IntermediateDisturbance Hypothesis in Phytoplankton Ecology. Kluwer Academic,1993]. Diversity may remain quite high even for extended periodsduring summer climax situations, in conjunction with a highdegree of similarity, if deeper mixing of the epilimnion occursat time intervals of 2–3 weeks, as during the summer of1993. This enables the prevalence of ‘ruderal’ species,together with some motile K-strategists who actively seek optimaldepths for photosynthesis. During such summer situations describedby frequent occurrences of lower lake numbers, the epilimnionof Lake Tegel is mixed deeply enough to support ruderals, butnot too vigorously to counteract competitive advantages of motilespecies. Thus, vertical niche separation enhances diversity.     

The effect of elevated CO2 on growth and competition in experimental phytoplankton communities

We report an experiment designed to identify the effect of elevated CO₂ on species of phytoplankton in a simple laboratory system. Major taxa of phytoplankton differ in their ability to take up CO₂, which might lead to predictable changes in the growth rate of species and thereby shifts in the composition of phytoplankton communities in response to rising CO₂. Six species of phytoplankton belonging to three major taxa (cyanobacteria, diatoms and chlorophytes) were cultured in atmospheres whose CO₂ concentration was gradually increased from ambient levels to 1000 parts per million over about 100 generations and then maintained for a further 200 generations at elevated CO₂. The experimental design allowed us to trace a predictive sequence, from physiological features to the growth response of species to elevated CO₂ in pure culture, from the growth response in pure culture to competitive ability in pairwise mixtures and from pairwise competitive ability to shifts in the relative abundance of species in the full community of all six species. CO₂ altered the dynamics of growth in a fashion consistent with known differences among major taxa in their ability to take up and use CO₂. This pure‐culture response was partly successful in predicting the outcome of competition in pairwise mixtures, especially the enhanced competitive ability of chlorophytes relative to cyanobacteria, although generally statistical support was weak. The competitive response in pairwise mixtures was a good predictor of changes in competitive ability in the full community. Hence, there is a potential for forging a logical chain of inferences for predicting how phytoplankton communities will respond to elevated CO₂. Clearly further extensive experiments will be required to validate this approach in the greater complexity found in diverse communities and environments of natural systems.     

Nutrient competition and division synchrony in phytoplankton

In the Gulf Stream in October of 1968, division of at least nine species of dinoflagellates occurred between 03.00 h and 06.00h indicating a high level of developmental synchronization. Whereas light-dark and temperature cycles act independently on each species, the synchronizing agent in the Gulf Stream apparently couples the development of many species of the dinoflagellate community. It is proposed that diurnal fluctuation in nutrients, acting through interspecific competition, is the coupling and synchronizing agent in this environment. A mathematical model of interspecific competition shows that regular, short-term changes in nutrient levels are extremely sffective in synchronizing the development of several species. Calculations also suggest that division eynchrony is an indicator of the occurrence of competition for nutrients and that the degree of synchronization increases with the severity of nutrient limitation on the phytoplankton.     

Kinetics and bioenergetics of light-limited photoautotrophic growth of Spirulina platensis

Blue-green algae, Spirulina platensis, is cultivated under photoautotrophic growth conditions designed to have nearly uniform growth rate throughout the fermentor by illumination both sides of a rectangular vessel. The results show that growth rate and bioenergetic yield are a function of light intensity. Several kinetic models are considered to express the relationship between growth rate and light intensity.     

Response of Rhodopseudomonas capsulata to illumination and growth rate in a light-limited continuous culture.

Rhodopseudomonas capsulata was grown under anaerobic, photosynthetic conditions in a continuous culture device. Under light limitation, at a constant dilution rate, it was shown that cell composition, including photopigment (bacteriochlorophyll and carotenoids) and ribonucleic acid content, was not affected by incident light intensity; however, steady state culture density varied directly and linearly with light intensity. On the other hand, photopigment and ribonucleic acid levels were affected by growth rate regardless of light intensity. Additional experiments indicated a high apparent Ks for growth of R. capsulata with respect to light. These results were interpreted to mean that near the maximum growth rate (D = 0.45 h-1) some internal metabolic process became the limiting factor for growth, rather than the imposed energy limitation. A mathematical expression for the relation between steady-state culture density and dilution rate was derived and was found to adequately describe the data. A strong correlation was found between continuous cultures limited either by light or by a chemical energy source.     

Role of infochemical mediated zooplankton grazing in a phytoplankton competition model

Infochemicals released by marine phytoplankton play important roles in food web interactions by influencing the feeding behavior and selectivity of zooplanktonic predators. Recent modeling efforts have focused on the role of such chemicals as toxic grazing deterrents in phytoplankton competition. However, infochemicals may also be utilized as grazing cues, leading predators to profitable foraging patches. Here we investigate the role of infochemical mediated zooplankton grazing in a standard 3-species phytoplankton competition model, with the aim of further elucidating the ecological role of phytoplankton derived infochemicals. We then extend this to consider a more realistic 4-species model. The models produce a range of solutions depending on the strength of competition and microzooplankton grazing selectivity. Our key result is that infochemical chemoattractants, which increase the susceptibility of the producer to grazing, can provide a refuge for both competing phytoplankton species by attracting carnivorous copepods to consume microzooplankton grazers in a multi-trophic interaction. Our results indicate that infochemicals potentially have important consequences for the dynamics of marine food webs.     

Predicting the spatial dominance of phytoplankton in a light limited and incompletely mixed eutrophic water column using the PROTECH model

1. A phytoplankton community model [Phytoplankton RespOnses To Environmental CHange (PROTECH)] was used to examine the effect of a wide range of varied light intensities and mixed depths upon simulated phytoplankton populations. Two different column lengths of the simulated water body were examined (the upper 5 m and the whole 14.5 m water column) for each scenario.
2. The hypotheses tested were that: (i) under low light intensity and/or deep mixing the simulated community will be dominated by a phytoplankter with a low critical light intensity; (ii) at high light intensity and shallow mixing the simulated community will be dominated by small, fast-growing phytoplankters; (iii) under all conditions, except deep mixing, the largest proportion of phytoplankton biomass will be found near the surface.
3. It was found under most conditions that, although there was a bloom in the upper column (dominated by algae such as Chlorella , Ceratium or Rhodomonas ), the largest phytoplankton biomass in the water column was located 9 m below the surface and consisted of solely Asterionella . This bloom was missed by the 5-m samples. Thus, using the whole column sample lengths, hypothesis (i) was not rejected but hypotheses (ii) and (iii) were refuted.
4. The inclusion of specific movement characteristics of phytoplankton in the model allowed the possibility of the dominance of multiple spaces within the water column and should be included in any model-based investigation of this topic. Further, the results from the model suggest that a reduced depth of mixing creates greater environmental heterogeneity, allowing more species to persist.     

The interaction of components controlling net phytoplankton photosynthesis in a well-mixed lake (Lough Neagh, Northern Ireland)

The homogeneous distribution of the phytoplankton in a shallow (mean depth 8·6 m) unstratified lake, L. Neagh, Northern Ireland, facilitated the study of the interaction of components controlling gross photosynthesis per unit area. These included the photosynthetic capacity, the phytoplankton content of the euphotic zone, and a logarithmic function describing the effective radiation input. These factors were analysed for two sites, the open lake and Kinnego Bay, which respectively had standing crops of up to 90 and 300 mg chlorophyll a m^?3 and maximum daily rates of gross integral photosynthesis of 11·7 and 15·6 g O₂ m^?2 day^?1. Values are reduced by the high contribution to light attenuation by non-algal sources, which increases at low standing crops particularly in winter, when values of integral photosynthesis decrease to 0·5 g O₂ m^?2 day^?1. This relative change is the result of self-shading behaviour of the phytoplankton altering the crop content of the euphotic zone at different population densities. Changes in the irradiance function, incorporating day length, are largely responsible for the changes in daily rates of integral gross photosynthesis; as daily irradiance is also a determinant of water temperature, it exerts further influence through the photosynthetic capacity which was strongly correlated with temperature. Much of the gain in gross photosynthesis resulting from higher photosynthetic capacity may not be reflected in a higher net column photosynthesis, because of the greater proportional rise in respiration with temperature. The balance in the water column between respiration losses and photosynthetic input may frequently alter since the ratio of illuminated to dark zones is between 1/4 to 1/5 in the open lake, and small shifts in any of the controlling features may result in conditions unfavourable for growth. This is analysed especially for the increase of diatoms in spring, when small modifications of the underwater light field can delay growth.     

The relationship between phytoplankton distribution and water column characteristics in North West European shelf sea waters

Phytoplankton underpin the marine food web in shelf seas, with some species having properties that are harmful to human health and coastal aquaculture. Pressures such as climate change and anthropogenic nutrient input are hypothesized to influence phytoplankton community composition and distribution. Yet the primary environmental drivers in shelf seas are poorly understood. To begin to address this in North Western European waters, the phytoplankton community composition was assessed in light of measured physical and chemical drivers during the "Ellett Line" cruise of autumn 2001 across the Scottish Continental shelf and into adjacent open Atlantic waters. Spatial variability existed in both phytoplankton and environmental conditions, with clear differences not only between on and off shelf stations but also between different on shelf locations. Temperature/salinity plots demonstrated different water masses existed in the region. In turn, principal component analysis (PCA), of the measured environmental conditions (temperature, salinity, water density and inorganic nutrient concentrations) clearly discriminated between shelf and oceanic stations on the basis of DIN:DSi ratio that was correlated with both salinity and temperature. Discrimination between shelf stations was also related to this ratio, but also the concentration of DIN and DSi. The phytoplankton community was diatom dominated, with multidimensional scaling (MDS) demonstrating spatial variability in its composition. Redundancy analysis (RDA) was used to investigate the link between environment and the phytoplankton community. This demonstrated a significant relationship between community composition and water mass as indexed by salinity (whole community), and both salinity and DIN:DSi (diatoms alone). Diatoms of the Pseudo-nitzschia seriata group occurred at densities potentially harmful to shellfish aquaculture, with the potential for toxicity being elevated by the likelihood of DSi limitation of growth at most stations and depths.     

余氯对小球藻的影响以及损失评估

滨海电厂和液化天然气(LNG)工程排出的余氯水可能对周边海域浮游植物造成不利的影响,从而成为破坏海洋生态系统平衡的重要因素之一。在实验室条件下研究余氯对浮游植物的抑制作用和造成的损失。余氯抑制试验用小球藻(Chlorella sp.)进行,依据工程余氯排放工况,余氯实验浓度设计为0、0.02、0.05、0.10和0.20 mg/L 5个水平;实验温度为27℃,代表夏季;观察时间分别为0、24、48、72和96h。结果表明:高浓度(0.20mg/L)余氯对小球藻生长有抑制作用,最大值出现在48h时,48h之后抑制作用基本维持在同一水平;低浓度(0.02mg/L)对小球藻有促进作用;0.05mg/L组与0.10mg/L组在实验时间24、48h和72h时表现出余氯对小球藻生长抑制作用,在96h时与对照组藻类生长情况差异不显著。采用积分方程计算累计损失率,参考上海LNG工程2009年9月浮游植物细胞丰度值,计算得出工程余氯排放在72h内所造成的藻类理论损失量为1.97×1012个。     

长江口锋面附近咸淡水混合对浮游植物生长影响的现场培养

通过2009年6月调查航次,获得了营养盐等参数断面分布,表明咸淡水混合是控制营养盐分布的主要因素。为了解不同盐度梯度下浮游植物生长与营养盐吸收的关系,采集两个站位水样分别代表长江冲淡水(C1站)和外海水(I10站),按C1站水样比例,分100%、75%、50%、25%、0%不同比例混合进行现场模拟咸淡水混合培养,有以下认识:(1)平行结果表明培养过程中活体荧光极大值在100%混合组,且淡水比例越低,指数生长期0—48 h内生长速率越低,100%、75%、50%、25%组分别为1.18/d、1.12/d、1.14/d、0.77/d;(2)低于26盐度的水体中PO34-在48 h内可被迅速消耗而产生限制作用,是控制浮游植物生长的潜在限制因子;(3)除0%组外,各混合组DIN/P(DIN:溶解无机氮,Dissolved Inorganic Nitrogen,DIN=NO3-+NO-2+NH4+)比值在浮游植物指数生长期有升高趋势,100%组DIN/P比值增加了一倍。各组培养48 h后DIN/Si比值逐渐降低至原来的0.7左右,初始DIN/Si小于一定时间内硅藻吸收的(ΔDIN/ΔSi)比,是造成各组DIN/Si比值减小的原因。以上结果表明咸淡水混合过程中形成的营养盐梯度可造成浮游植物生长程度和速率差异,且可因局部浮游植物旺发而改变海水营养结构。     

Comparison of the light-limited growth of the nitrogen-fixing cyanobacteria Anabaena and Aphanizomenon

The effect of simultaneous N₂ fixation and light limitation on the growth of two strains of Anabaena sp. Bory de St. Vincent and Aphanizomenon flos-aquae (L.) Ralfs was investigated using continuous cultures. Under severely light-limited conditions, Aphanizomenon showed a broader absorption spectrum (due to the presence of phycoerythrin), a higher maximum efficiency of photosynthesis, a higher steady-state N₂ fixation activity and a higher growth affinity for light than did Anabaena . On the other hand, under light saturation, Anabaena showed a higher maximum rate of O₂ production and a higher maximum specific growth rate than Aphanizomenon . These monoculture results characterize Anabaena and Aphanizomenon , in relative terms, as a 'sun' and a 'shade' species respectively, and are in accordance with field observations. The difference between the two species in their acclimatory response is discussed in terms of a species-specific alteration of the PSI:PSII stoichiometry. Besides the species-specific modulation of the accessory pigments, such an acclimation would provide a biochemical basis for the observed physiological differences. The monoculture results were used to differentiate the niches of the two species and suggested that Aphanizomenon would competitively displace Anabaena under N₂-fixing, light-limited conditions. However, when both species were grown together, Anabaena became dominant and seemed to be the superior competitor for light. In order to explain this finding, the possible effects of release of allelopathic compounds, or dynamic aspects of light supply, are discussed.