A nonautonomous model for the effects of refuge and additional food on the dynamics of phytoplankton-zooplankton system

In this paper, a mathematical model for the interacting dynamics of phytoplankton-zooplankton is proposed. The phytoplankton have the ability to take refuge and release toxins to avoid over predation by zooplankton. The zooplankton are provided some additional food to persist in the system. The phytoplankton are assumed to be affected directly by external toxic substances whereas zooplankton are affected indirectly by feeding on the affected phytoplankton. We incorporate seasonal variations in the model, assuming the level of nutrients, refuge and the rate of toxins released by phytoplankton as functions of time. Our results show that when high toxicity and refuge cause extinction of zooplankton, providing additional food supports the survival of zooplankton population and controls the phytoplankton population. Prey refuge and additional food have stabilizing effects on the system; higher values of the former results in extinction of zooplankton whereas phytoplankton disappear for larger values of the latter. Seasonality in nutrients level and toxins released by phytoplankton generate higher periodic solutions while time-dependent refuge of phytoplankton causes the occurrence of a period-three solution. The possibility of finding additional food for zooplankton may push back the ecosystem to a simple stable state from a complex dynamics.     

Do phytoplankton communities evolve through a self-regulatory abundance–diversity relationship?

A small group of phytoplankton species that produce toxic or allelopathic chemicals has a significant effect on plankton dynamics in marine ecosystems. The species of non-toxic phytoplankton, which are large in number, are affected by the toxin-allelopathy of those species. By analysis of the abundance data of marine phytoplankton collected from the North-West coast of the Bay of Bengal, an empirical relationship between the abundance of the potential toxin-producing species and the species diversity of the non-toxic phytoplankton is formulated. A change-point analysis demonstrates that the diversity of non-toxic phytoplankton increases with the increase of toxic species up to a certain level. However, for a massive increase of the toxin-producing species the diversity of phytoplankton at species level reduces gradually. Following the results, a deterministic relationship between the abundance of toxic phytoplankton and the diversity of non-toxic phytoplankton is developed. The abundance-diversity relationship develops a unimodal pathway through which the abundance of toxic species regulates the diversity of phytoplankton. These results contribute to the current understanding of the coexistence and biodiversity of phytoplankton, the top-down vs. bottom-up debate, and to that of abundance-diversity relationship in marine ecosystems.     

A method for the determination of glycollic acid in the extracellular products of cultured and natural phytoplankton populations

A method for accurately isolating and characterizing glycollic acid excreted by phytoplankton has been developed; it is applicable to both laboratory and field conditions. Results are given for phytoplankton in culture and natural phytoplankton communities.     

Omnivory by Planktivores Stabilizes Plankton Dynamics, but May Either Promote or Reduce Algal Biomass

Classical models of phytoplankton–zooplankton interaction show that with nutrient enrichment such systems may abruptly shift from limit cycles to stable phytoplankton domination due to zooplankton predation by planktivorous fish. Such models assume that planktivorous fish eat only zooplankton, but there are various species of filter-feeding fish that may also feed on phytoplankton. Here, we extend these classical models to systematically explore the effects of omnivory by planktivorous fish. Our analysis indicates that if fish forage on phytoplankton in addition to zooplankton, the alternative attractors predicted by the classical models disappear for all realistic parameter settings, even if omnivorous fish have a strong preference for zooplankton. Our model also shows that the level of fish biomass above which zooplankton collapse should be higher when fish are omnivorous than when fish are zooplanktivorous. We also used the model to explore the potential effects of the now increasingly common practice of stocking lakes with filter-feeding fish to control cyanobacteria. Because omnivorous filter-feeding fish forage on phytoplankton as well as on the main grazers of phytoplankton, the net effect of such fish on the phytoplankton biomass is not obvious. Our model suggests that there may be a unimodal relationship between the biomass of omnivorous filter-feeding fish and the biomass of phytoplankton. This implies that to manage for reductions in phytoplankton biomass, heavy stocking or strong reduction of such fish is best.     

An Examination of Cell Volume in Dominant Phytoplankton Species of the Central and Southern Adriatic Sea

The phytoplankton biomass expressed as total cell volume is often of doubtful value due to cell size and volume variations in the organisms present. Cell volume calculations for 95 phytoplankton species and volume-frequency diagrams for 25 dominant species were analyzed for the Adriatic Sea. Recommendations for more precise work dealing with phytoplankton cell volume and biomass are given. Temporal variations in the average microplankton cell volume are presented and discussed.     

RELATING PHYTOPLANKTON BIOMASS AND PRODUCTION TO EPISODIC PHYSICAL FORCING IN SOUTHEASTERN LAKE MICHIGAN

Spatial and temporal dynamics in phytoplankton reflect of the combined effects of the physical and chemical environments and associated biological responses. Although alterations in phytoplankton are well-documented for a variety of lentic waters, the exact linkages between environmental forcing and phytoplankton assemblages remain poorly understood (particularly for coastal systems). A recurrent sediment resuspension event occurs every late winter/early spring in southeastern Lake Michigan, often extending greater than ten km in width and 300 km in length. Inherently, such a large-scale and dramatic physical process would be thought to dramatically influence phytoplankton assemblages; however, linkages between the turbidity plume and phytoplankton assemblages have been postulated, but never verified. As such, the episodic nature of the plume provided an opportunity to examine the effects of a short-term physical forcing event on coastal phytoplankton in relation to more persistent, seasonal meteorological forcing. Lake phytoplankton assemblages within and outside of the RCP were examined during the spring isothermal period from 1998 to 2000. Here, we describe results from the 1998 and 1999 field seasons characterizing the distribution of phytoplankton biomass and composition within and adjacent to the RCP and their relationship to particulate and dissolved constituents. In addition, the spatial and temporal patterns in production and photosynthetic characteristics of the phytoplankton community are examined.     

海洋浮游植物与生物碳汇

系统描述了浮游植物与海洋碳汇相关的几个过程:初级生产、浮游植物沉降、浮游动物粪球打包沉降、经典食物链碳汇、溶解有机碳生产和转化、透明胞外聚合颗粒物(TEP)凝聚网,和CO₂分压升高(海水酸化)影响下浮游植物功能群转变及中国海可能的生物碳汇前景展望。提出海洋初级生产过程和TEP凝聚网过程是中国海生物碳汇的关键过程,而中国海的黄海中部及长江口区域是生物碳汇研究的重点区域,建议将硅藻及其碳汇过程作为今后研究的重点。     

Phytoplankton response to a changing climate

Phytoplankton are at the base of aquatic food webs and of global importance for ecosystem functioning and services. The dynamics of these photosynthetic cells are linked to annual fluctuations of temperature, water column mixing, resource availability, and consumption. Climate can modify these environmental factors and alter phytoplankton structure, seasonal dynamics, and taxonomic composition. Here, we review mechanistic links between climate alterations and factors limiting primary production, and highlight studies where climate change has had a clear impact on phytoplankton processes. Climate affects phytoplankton both directly through physiology and indirectly by changing water column stratification and resource availability, mainly nutrients and light, or intensified grazing by heterotrophs. These modifications affect various phytoplankton processes, and a widespread advance in phytoplankton spring bloom timing and changing bloom magnitudes have both been observed. Climate warming also affects phytoplankton species composition and size structure, and favors species traits best adapted to changing conditions associated with climate change. Shifts in phytoplankton can have far-reaching consequences for ecosystem structure and functioning. An improved understanding of the mechanistic links between climate and phytoplankton dynamics is important for predicting climate change impacts on aquatic ecosystems.     

Dynamics of phytoplankton in Finnish lakes

The studies on lake phytoplankton in Finland are reviewed and the major aspects of the phytoplankton dynamics are discussed. Special attention has been paid to the factors limiting productivity and species succession in different communities. After the early mainly taxonomical and floristic publications on phytoplankton at the end of last century, phytoplankton studies in lakes have proceeded along two different lines: 1) the species composition of communities and taxonomy, and 2) their production ecology or dynamics. Recently, both approaches have been combined, resulting in some profound ecological studies. In many lakes, phosphorus has been shown to be a limiting factor for phytoplankton productivity. However, it has also been shown that the irradiance and water temperature may effectively regulate the seasonal trend of phytoplankton productivity. This is the case especially in polyhumic forest lakes, where allochthonous material seems to play a major role also in primary production ecology.     

Implications of seasonal mixing for phytoplankton production and bloom development

Based on a 1D model considering phytoplankton and nutrients in a vertical water column, we investigate the consequences of temporal and spatial variations in turbulent mixing for phytoplankton production and biomass. We show that in seasonally mixed systems, the processes controlling phytoplankton production and the sensitivity of phytoplankton abundance to ambient light, trophic state and mixed-layer depth differ substantially from those at steady state in systems with time-constant diffusivities. In seasonally mixed systems, the annually replenished nutrient pool in the euphotic zone is an important factor for phytoplankton production supporting bloom development, whereas without winter mixing, production mainly depends on the diffusive nutrient flux during stratified conditions. Seasonal changes in water column production are predominantly determined by seasonal changes in phytoplankton abundance, but also by seasonal changes in specific production resulting from the transport of nutrients, the exploitation of the nutrient pool and the increase in light shading associated with phytoplankton growth. The interplay between seasonal mixing and the vertical distribution of mixing intensities is a key factor determining the relative importance of the processes controlling phytoplankton production and the sensitivity of the size and timing of the annual maximum phytoplankton abundance to the abiotic conditions.     

Patchy agglomeration as a transition from monospecies to recurrent plankton blooms

We propose a model for explaining both red tides and recurring phytoplankton blooms. Three assumptions are made, namely the presence of toxin producing phytoplankton, the satiation phenomenon in zooplankton's feeding, modelled by a Holling type II response, and phytoplankton aggregation leading to formation of patches. The dynamics of the plankton population is shown to depend on the fraction of the phytoplankton population that aggregates to form colonies and on the number of the latter.     

A PHYLOGENETIC APPROACH TO THE ESTIMATION OF PHYTOPLANKTON TRAITS1

The quantitative characterization of the ecology of individual phytoplankton taxa is essential for model resolution of many aspects of aquatic ecosystems. Existing literature cannot directly parameterize all phytoplankton taxa of interest, as many traits and taxa have not been sampled. However, valuable clues on the value of traits are found in the evolutionary history of species and in common correlations between traits. These two resources were exploited with an existing, statistically consistent method built upon evolutionary concepts. From a new data set with >700 observations on freshwater phytoplankton traits and a qualitative phytoplankton phylogeny, estimates were derived for the size, growth rate, phosphate affinity, and susceptibility to predation of 277 phytoplankton types, from evolutionary ancestors to present‐day species. These estimates account simultaneously for phylogenetic relationships between types, as imposed by the phylogeny, and approximate power‐law relationships (e.g., allometric scaling laws) between traits, as reconstructed from the data set. Results suggest that most phytoplankton traits are to some extent conserved in evolution: cross‐validation demonstrated that the use of phylogenetic information significantly improves trait value estimates. By providing trait value estimates as well as uncertainties, these results could benefit most quantitative studies involving phytoplankton.     

Dynamics of nutrient-phytoplankton interaction in the presence of viral infection

The present paper deals with the problem of a nutrient-phytoplankton (N-P) populations where phytoplankton population is divided into two groups, namely susceptible phytoplankton and infected phytoplankton. Conditions for coexistence or extinction of populations are derived taking into account general nutrient uptake functions and Holling type-II functional response as an example. It is observed that the three component systems persist when the infected phytoplankton population is not able to consume nutrient.     

Some aspects of the seasonal distribution of flagellates in mountain lakes

In a larger regional survey in Tyrol, phytoplankton species composition and biovolume of mid-altitude and high-mountain lakes was studied. Results from eight lakes showed that flagellates (mainly Chrysophyceae, Dinophyceae, and Cryptophyceae) are important components of the phytoplankton.In the mid-altitude lakes a spring and an autumn maximum of Chrysophyceae as well as a summer maximum of large dinoflagellates are observed, whereas Cryptophyceae and Dinophyceae show irregular distributions. In the high-mountain lakes the seasonal variations of phytoplankton, including flagellates, are limited by the long duration of the winter situation. However similar sequences of phytoplankton assemblages as in the midaltitude lakes can be observed. Flagellates in high-mountain lakes are important to sustain phytoplankton standing crop under the winter snow and ice cover.In order to show similarities and differences of high-mountain and mid-altitude lakes, vertical profiles of phytoplankton from three lakes and seasonal patterns of Gymnodinium uberrimum from two lakes are compared. In addition the patterns of cryptomonads differing in their ecological requirements (Cryptomonas spp. and Rhodomonas minuta) are shown for a meromictic mid-altitude lake.     

How do sinking phytoplankton species manage to persist?

Phytoplankton require light for photosynthesis. Yet, most phytoplankton species are heavier than water and therefore sink. How can these sinking species persist? Somehow, the answer should lie in the turbulent motion that redisperses sinking phytoplankton over the vertical water column. Here, we show, using a reaction-advection-diffusion equation of light-limited phytoplankton, that there is a turbulence window sustaining sinking phytoplankton species in deep waters. If turbulent diffusion is too high, phytoplankton are mixed to great depths, and the depth-averaged light conditions are too low to allow net positive population growth. Conversely, if turbulent diffusion is too low, sinking phytoplankton populations end up at the ocean floor and succumb in the dark. At intermediate levels of turbulent diffusion, however, phytoplankton populations can outgrow both mixing rates and sinking rates. In this way, the reproducing population as a whole can maintain a position in the well-lit zone near the top of the water column, even if all individuals within the population have a tendency to sink. This theory unites earlier classic results by Sverdrup and Riley as well as our own recent findings and provides a new conceptual framework for the understanding of phytoplankton dynamics under the influence of mixing processes.     

The impact of diel vertical migration of <Emphasis Type="Italic">Daphnia</Emphasis> on phytoplankton dynamics

Diel vertical migration (DVM) of large zooplankton is a very common phenomenon in the pelagic zone of lakes and oceans. Although the underlying mechanisms of DVM are well understood, we lack experimental studies on the consequences of this behaviour for the zooplankton’s food resource—the phytoplankton. As large zooplankton species or individuals migrate downwards into lower and darker water strata by day and upwards into surface layers by night, a huge amount of herbivorous biomass moves through the water column twice a day. This migration must have profound consequences for the phytoplankton. It is generally assumed that migration supports an enhanced phytoplankton biomass and a change in the composition of the phytoplankton community towards smaller, edible algae in the epilimnion of a lake. We tested this assumption for the first time in field experiments by comparing phytoplankton biomass and community assemblage in mesocosms with and without artificially migrating natural stocks of Daphnia hyalina. We show that DVM can enhance phytoplankton biomass in the epilimnion and that it has a strong impact on the composition of a phytoplankton community leading to an advantage for small, edible algae. Our results support the idea that DVM of Daphnia can have strong effects on phytoplankton dynamics in a lake.     

Iron bioavailability to phytoplankton: an empirical approach

Phytoplankton are often limited by iron in aquatic environments. Here we examine Fe bioavailability to phytoplankton by analyzing iron uptake from various Fe substrates by several species of phytoplankton grown under conditions of Fe limitation and comparing the measured uptake rate constants (Fe uptake rate/ substrate concentration). When unchelated iron, Fe′, buffered by an excess of the chelating agent EDTA is used as the Fe substrate, the uptake rate constants of all the eukaryotic phytoplankton species are tightly correlated and proportional to their respective surface areas (S.A.). The same is true when FeDFB is the substrate, but the corresponding uptake constants are one thousand times smaller than for Fe′. The uptake rate constants for the other substrates we examined fall mostly between the values for Fe′ and FeDFB for the same S.A. These two model substrates thus empirically define a bioavailability envelope with Fe′ at the upper and FeDFB at the lower limit of iron bioavailability. This envelope provides a convenient framework to compare the relative bioavailabilities of various Fe substrates to eukaryotic phytoplankton and the Fe uptake abilities of different phytoplankton species. Compared with eukaryotic species, cyanobacteria have similar uptake constants for Fe′ but lower ones for FeDFB. The unique relationship between the uptake rate constants and the S.A. of phytoplankton species suggests that the uptake rate constant of Fe-limited phytoplankton has reached a universal upper limit and provides insight into the underlying uptake mechanism.     

全球气候变化下的海洋浮游植物多样性

理解全球气候变化对地球生态系统的影响是全世界广泛关注的问题, 而相比于陆地生态系统, 海洋生态系统对全球气候变化更为敏感。全球气候变化对海洋的影响主要表现在海洋暖化、海洋酸化、大洋环流系统的改变、海平面上升、紫外线辐射增强等方面。浮游植物是海洋生态系统最重要的初级生产者, 同时对海洋碳循环起到举足轻重的作用, 其对全球气候变化的响应主要体现在物种分布、初级生产力、群落演替、生物气候学等方面。具体表现在以下方面: 暖水种的分布范围在扩大, 冷水种分布范围在缩小; 浮游植物全球初级生产力降低; 浮游植物群落会向细胞体积更小的物种占优势的方向转变; 浮游植物水华发生的时间提前、强度增强; 一些有害物种水华的发生频率也会增加; 海洋表层海水的酸化会影响浮游植物特别是钙化类群的生长和群落多样性; 紫外辐射增强对浮游植物的生长起到抑制作用; 厄尔尼诺、拉尼娜、降水量的增加通常抑制浮游植物生长。浮游植物生长和分布的变化会体现在多样性的各个层面上。对于浮游植物在全球变化各种驱动因子下的生理生态学和长周期变动观测等是今后研究的重要方向, 也将为理解全球变化下的浮游植物-多样性-生态系统响应与反馈机制提供基本信息。     

Trophic relationships in the pelagic zone of Mondsee,Austria

Data are presented on nutrient concentrations, phytoplankton biovolume development, zooplankton composition and population dynamics, and fish from a deep, stratifying, alpine lake (Mondsee, Austria) during a three-year period between 1982 and 1984. Development of the phytoplankton is closely related to structuring events of the physico-chemical environment. Dissolved silicate and phosphorus concentrations are critical for the summer situation. During summer algal abundance is largely affected by grazing of zooplankton, but no clear-water phase was observed at the end of the spring peak of phytoplankton.Temperature and food are factors responsible for the timing and growth of the zooplankton populations. Because of close overlap in the epilimnion, exploitative and mechanical interference competition and predation by invertebrate and vertebrate predators are the main structuring forces acting on the zooplankton community, and hence influence phytoplankton indirectly.     

The role of highly unsaturated fatty acids in aquatic foodweb processes

1. Polyunsaturated fatty acids (PUFA) are almost exclusively synthesized by plants. Animals can convert from one form of PUFA to another through elongation and desaturation, but very few can synthesize PUFA de novo. PUFA play an important role in regulating cell membrane properties, serve as precursors for important animal hormones and are essential for animals. 2. In aquaculture studies, highly unsaturated fatty acids (HUFA), a subset of PUFA, have been found to be critical for maintaining high growth, survival and reproductive rates and high food conversion efficiencies for a wide variety of marine and freshwater organisms. 3. The plankton literature suggests high food-quality algae species are rich in HUFA and low food-quality algae are poor in HUFA. Adding semi-pure emulsions of HUFA to algae monocultures can markedly increase the growth rates of zooplankton feeding on these mixtures. 4. A study measuring zooplankton biomass accrual when feeding on natural phytoplankton found a strong correlation between phytoplankton HUFA (specifically eicosapentaenoic acid) content and herbivorous zooplankton production. 5. The aquatic ecology literature suggests that planktonic foodwebs with high HUFA content phytoplankton have high zooplankton to phytoplankton biomass ratios, while systems with low HUFA phytoplankton have low zooplankton biomass. Also, the seasonal succession of plankton in many temperate lakes follows patterns tied to phytoplankton HUFA content, with intense zooplankton grazing and ‘clear-water-phases’ characteristic of periods when the phytoplankton is dominated by HUFA-rich species. 6. Herbivorous zooplankton production is constrained by the zooplankton’s ability to ingest and digest phytoplankton. It is becoming increasingly clear, however, that much of the phytoplankton which is assimilated may be nutritionally inadequate. HUFA may be key nutritional constituents of zooplankton diets, and may determine energetic efficiency across the plant–animal interface, secondary production and the strength of trophic coupling in aquatic pelagic foodwebs.