Analysis of the self-shading effect on algal vertical distribution in natural waters

Self-shading of light by algae growing in a column of water plays an important role in the dynamics of algal blooms. Thus without self-shading the algal concentration would increase more rapidly, making the nutrient limitation too strong. Apart from the practical importance of self-shading, its inherent nonlinearity in the growth dynamics leads to an interesting mathematical problem, which warrants detailed analytical investigation. Our mathematical model for the self-shading effect includes vertical diffusion, algal settling, gross production, and collective losses of algae. Steady-state solutions of the model equation are investigated in detail by the phase plane method, and their stability examined. Finally we discuss the vertical profile of algal concentration.Contribution No. 291 of the Marine Sciences Research Center     

Promotion of harmful algal blooms by zooplankton predatory activity

The relationship between algae and their zooplanktonic predators typically involves consumption of nutrients by algae, grazing of the algae by zooplankton which in turn enhances predator biomass, controls algal growth and regenerates nutrients. Eutrophication raises nutrient levels, but does not simply increase normal predator-prey activity; rather, harmful algal bloom (HAB) events develop often with serious ecological and aesthetic implications. Generally, HAB species are outwardly poor competitors for nutrients, while their development of grazing deterrents during nutrient stress ostensibly occurs too late, after the nutrients have largely been consumed already by fast-growing non-HAB species. A new mechanism is presented to explain HAB dynamics under these circumstances. Using a multi-nutrient predator-prey model, it is demonstrated that these blooms can develop through the self-propagating failure of normal predator-prey activity, resulting in the transfer of nutrients into HAB growth at the expense of competing algal species. Rate limitation of this transfer provides a continual level of nutrient stress that results in HAB species exhibiting grazing deterrents protecting them from top-down control. This process is self-stabilizing as long as nutrient demand exceeds supply, maintaining the unpalatable status of HABs; such events are most likely under eutrophic conditions with skewed nutrient ratios.     

Algal Diet of Small-Bodied Crustacean Zooplankton in a Cyanobacteria-Dominated Eutrophic Lake

Small-bodied cladocerans and cyclopoid copepods are becoming increasingly dominant over large crustacean zooplankton in eutrophic waters where they often coexist with cyanobacterial blooms. However, relatively little is known about their algal diet preferences. We studied grazing selectivity of small crustaceans (the cyclopoid copepods Mesocyclops leuckarti, Thermocyclops oithonoides, Cyclops kolensis, and the cladocerans Daphnia cucullata, Chydorus sphaericus, Bosmina spp.) by liquid chromatographic analyses of phytoplankton marker pigments in the shallow, highly eutrophic Lake Võrtsjärv (Estonia) during a seasonal cycle. Copepods (mainly C. kolensis) preferably consumed cryptophytes (identified by the marker pigment alloxanthin in gut contents) during colder periods, while they preferred small non-filamentous diatoms and green algae (identified mainly by diatoxanthin and lutein, respectively) from May to September. All studied cladoceran species showed highest selectivity towards colonial cyanobacteria (identified by canthaxanthin). For small C. sphaericus, commonly occuring in the pelagic zone of eutrophic lakes, colonial cyanobacteria can be their major food source, supporting their coexistence with cyanobacterial blooms. Pigments characteristic of filamentous cyanobacteria and diatoms (zeaxanthin and fucoxanthin, respectively), algae dominating in Võrtsjärv, were also found in the grazers’ diet but were generally avoided by the crustaceans commonly dominating the zooplankton assemblage. Together these results suggest that the co-occurring small-bodied cyclopoid and cladoceran species have markedly different algal diets and that the cladocera represent the main trophic link transferring cyanobacterial carbon to the food web in a highly eutrophic lake.     

The importance of Protozoa in controlling the abundance of planktonic algae in lakes

Apart from ciliates, there are other protozoans which can form an important element in the herbivorous zooplankton of lakes.
The protozoans discussed in this paper feed on colonial algae. They can greatly reduce the numbers of certain Chlorophyta in Windermere, Esthwaite Water and Blelham Tarn, lakes in the English Lake District.
A very short period, often 7–14 days, can suffice for the destruction of more than 99% of an algal population.
Experiments with a species of Pseudospora showed that it would only ingest one out of six green algal species, although, in nature, Pseudosporae which are morphologically similar to this species have been observed in all these algae.
The importance of such protozoans, and of parasitic fungi, has been underrated or neglected in studies of the quantitative relationships between primary and secondary production in the plankton.     

Positive feedbacks between bottom-up and top-down controls promote the formation and toxicity of ecosystem disruptive algal blooms: A modeling study

Harmful algal blooms that disrupt and degrade ecosystems (ecosystem disruptive algal blooms, EDABs) are occurring with greater frequency and severity with eutrophication and other adverse anthropogenic alterations of coastal systems. EDAB events have been hypothesized to be caused by positive feedback interactions involving differential growth of competing algal species, low grazing mortality rates on EDAB species, and resulting decreases in nutrient inputs from grazer-mediated nutrient cycling as the EDAB event progresses. Here we develop a stoichiometric nutrient–phytoplankton–zooplankton (NPZ) model to test a conceptual positive feedback mechanism linked to increased cell toxicity and resultant decreases in grazing mortality rates in EDAB species under nutrient limitation of growth rate. As our model EDAB alga, we chose the slow-growing, toxic dinoflagellate Karenia brevis, whose toxin levels have been shown to increase with nutrient (nitrogen) limitation of specific growth rate. This species was competed with two high-nutrient adapted, faster-growing diatoms (Thalassiosira pseudonana and Thalassiosira weissflogii) using recently published data for relationships among nutrient (ammonium) concentration, carbon normalized ammonium uptake rates, cellular nitrogen:carbon (N:C) ratios, and specific growth rate. The model results support the proposed positive feedback mechanism for EDAB formation and toxicity. In all cases the toxic bloom was preceded by one or more pre-blooms of fast-growing diatoms, which drew dissolved nutrients to low growth rate-limiting levels, and stimulated the population growth of zooplankton grazers. Low specific grazing rates on the toxic, nutrient-limited EDAB species then promoted the population growth of this species, which further decreased grazing rates, grazing-linked nutrient recycling, nutrient concentrations, and algal specific growth rates. The nutrient limitation of growth rate further increased toxin concentrations in the EDAB algae, which further decreased grazing-linked nutrient recycling rates and nutrient concentrations, and caused an even greater nutrient limitation of growth rate and even higher toxin levels in the EDAB algae. This chain of interactions represented a positive feedback that resulted in the formation of a high-biomass toxic bloom, with low, nutrient-limited specific growth rates and associated high cellular C:N and toxin:C ratios. Together the elevated C:N and toxin:C ratios in the EDAB algae resulted in very high bloom toxicity. The positive feedbacks and resulting bloom formation and toxicity were increased by long water residence times, which increased the relative importance of grazing-linked nutrient recycling to the overall supply of limiting nutrient (N).     

Evaluation of eutrophication control in Lake Chivero,Zimbabwe, by multivariate analysis of zooplankton

Zooplankton was a useful tool in evaluation of eutrophication control in Lake Chivero, a tropical man made impoundment. Principal component analysis revealed both spatial and temporal changes in zooplankton community structure which were related to nutrient inflows as well as changes in nutrient content of the lake. Studies on filtration rates of different algae by Bosmina longirostris and Daphnia lumholtzi demonstrated that the response of zooplankton to eutrophication was related to their inability to utilize colonial algal species that develop in a nutrient rich-environment.     

浮游动物诱发藻类群体的形成

从研究蓝藻水华形成机理的需要出发,综述了浮游动物的牧食压力对藻类群体形成的诱发作用。指出诱发藻类群体形成的化合物来自牧食性浮游动物对藻类的有效牧食,是藻类群体形成的重要原因之一,而这些诱发性的化合物并不是有关生物体的组成成分,是种间相互作用的结果。藻类群体的形成方式有源于一个母细胞的分裂和业已存在的单细胞的聚合两种方式,栅藻的诱发性群体可能是来自一个母细胞的分裂,而在其它藻类的诱发性群体形成如铜绿微囊藻则可能是业已存在的单细胞的聚合。由于藻类形成群体后能显著降低浮游动物对其牧食速率,因此,这种诱发性群体形成的现象,可以解释为藻类对变化的牧食压力的一种有效的反牧食防御策略,也是两者协同进化的结果。浮游动物对藻类群体形成的重要作用,在研究模拟蓝藻群体及水华形成值得借鉴应用。作者还提出推测,水华蓝藻的群体形成,可能就是在富营养化条件下藻类快速生长,加上浮游动物的牧食压力共同作用下联合驱动的结果,而这种群体形成很可能在积累到一定程度后,结合特定的气象水文等理化因子,就会聚集于水表“爆发”出肉眼可见的水华。因此,开展浮游动物牧食作用对水华蓝藻早期群体形成诱发效应的研究不仅能加深对水华形成的全面认识,而且对于进一步认识藻类的诱发性反牧食防御适应机制、揭示生态系统中生物之间的复杂关系也具有重要的理论意义。     

Effects of the blue-green alga Microcystis aeruginosa on zooplankton competitive relations

Summary Field distribution patterns and laboratory feeding experiments have suggested that blooms of colonial blue-green algae strongly inhibit relatively large-bodied daphnid cladocerans. We conducted laboratory experiments to test the hypothesis that blooms of the colonial blue-green alga Microcystis aeruginosa would shift competitive dominance away from large-bodied daphnid cladocerans toward smaller-bodied cladocerans, copepods, and rotifers. In laboratory competition experiments, increasing the proportion of M. aeruginosa in the algal food supply resulted in a shift from dominance by the relatively largebodied cladoceran Daphnia ambigua to dominace by the copepod Diaptomus reighardi. The small-bodied cladoceran Bosmina longirostris was always numerically heavily dominant over D. ambigua, but its estimated population biomasses were only slightly higher than those of D. ambigua. Daphnia ambigua consistently outcompeted the rotifer Brachionus calyciflorus. Our results demonstrate that blooms of M. aeruginosa can alter zooplankton competitive relations in laboratory experiments, favoring small-bodied cladocerans and copepods at the expense of large-bodied cladocerans. However, contrary to predictions, blooms of M. aeruginosa did not improve the competitive ability of rotifers.     

Spatial patterns and relationships between phytoplankton, zooplankton and water quality in the Saimaa lake system, Finland

The interactions between phytoplankton and zooplankton were studied in two large lakes in the Saimaa lake system, Finland. Both are subjected to substantial waste water loading, and exhibit a clear trophic gradient between the loaded and unloaded areas. The phytoplankton and zooplankton were compared in terms of composition, abundance and biomass at 34–39 stations located in different parts of the lakes. At least four mechanisms were thought to affect the composition of plankton communities: (1) the amount of nutrients (trophic gradient), (2) grazing of algae by herbivores, (3) the effect of the algal species composition on feeding by zooplankters (large, colonial algae in the more loaded parts of the lakes) and (4) the regeneration and reorganization of nutrients.     

Influence of metazooplankton on interactions of bacteria and phytoplankton in an oligotrophic lake

Commensalism based on organic carbon supplied by phytoplanktonand competition for mineral nutrients are important interactionsbetween bacteria and phytoplankton in oligotrophic clear-watersystems. Both interactions are influenced by zooplankton activity.To examine the relation ship between algae and bacteria in LakeLa Caldera, we studied: the correlations among phyto plankton,bacteria and phosphorus (P) dynamics; the ratio of organic carbonsupplied by algae to organic carbon demand by bacteria; andthe importance of P remineralized by metazooplankton for bothcommunities. Phytoplankton and bacteria had a similar seasonaldynamics, and there was a sig nificant and positive relationshipbetween bacterial abundance and algal biomass (P<0.01). However,the release of organic carbon from phytoplankton was usuallyhigher than the bacterioplankton carbon requirement. P availablevia zooplankton remineralization satisfied between 74 and 316%of the minimum P demands of algae and bacteria. To elucidatewhether zooplankton operate similarly on algae and bacterialgrowth or indirectly influence bacterial growth through phytoplanktonmetab olism, we performed zooplankton manipulation experiments.High zooplankton biomass in these experiments stimulated bothprimary and bacterial production, but release of organic carbonfrom phytoplankton declined. These results suggest a directstimulus of bacterial growth, so algae and bac teria can balancegrazing losses by compensatory growth. Further, the algal decreaseof the organic carbon supply for bacteria could, over time,lead to a change in the algae-bacteria interaction from competitionto commensalism. This reduction in organic carbon excretioncould affect the balance of the competitive interaction.     

Keystone predator effects and grazer control of planktonic primary production

If prey species exhibit trade-offs in their ability to utilize resources versus their ability to avoid predation, predators can facilitate prey turnover along gradients of productivity, shifting dominance from edible to inedible prey (the keystone predator effect). I tested this model under controlled, laboratory conditions, using a model aquatic system composed of zooplankton as the top consumer, a diverse community of algae as prey, and nutrients as basal resources. Nutrient manipulations (low and high) were crossed with presence–absence of zooplankton. Results supported theoretical predictions. Algal biomass increased in response to enrichment regardless of predator presence/absence. However, predators and nutrients had an interactive effect on algal biomass and size structure. At the low nutrient level, algal-prey were dominated by edible forms and attained similar biomass regardless of zooplankton presence/absence. At the high level of enrichment, presence of zooplankton favored higher levels of algal biomass and shifted dominance to large, inedible taxa. At the termination of the experiment, I performed a series of lab-based assays on the resultant algal community in order to quantify trade-offs among algal size classes in maximal population growth rates (as a measure of competitive ability for nutrients) and susceptibility to zooplankton grazing. Assays provided support for a size-based keystone trade-off. Small size classes of algae displayed higher maximal growth rates but were more susceptible to grazing effects. Large size classes were protected from grazing but showed low rates of population growth in response to enrichment.     

在不同蓄水位下三峡库区春季水华特征及趋势分析

基于2004—2015年三峡库区支流春季水华的监测数据,开展了三峡库区不同蓄水位下春季水华特征及趋势分析。结果表明:总共有26条支流发生春季水华,香溪河为水华发生概率最高的支流。水华藻类优势种主要是硅藻和甲藻,甲藻水华出现的概率随着蓄水位的增加而增加,但是在持续几年175 m蓄水之后,甲藻水华出现的频率下降,硅藻水华出现的频率增加。在175 m蓄水之前,水华发生期藻类密度、叶绿素a含量及水体营养盐水平相对较低,此后都有了较高的提升,但是在持续几年175 m蓄水之后,藻类密度、叶绿素a浓度以及营养盐浓度都有下降趋势。对藻密度及叶绿素a含量与主要环境因子进行主成分分析(Principal components analysis)发现,藻密度和叶绿素a与总磷、总氮、高锰酸盐指数聚类一簇,藻密度、叶绿素a浓度与总磷、溶解氧、高锰酸盐指数显著正相关。在蓄水之后,三峡库区支流营养水平有所增加,随着蓄水位的增加,三峡库区支流断面的平均流速降低,在合适的温度下,三峡库区支流水华易发生。     

Responses to fish predation and nutrients by plankton at different levels of taxonomic resolution

1. To improve mechanistic understanding of plankton responses to eutrophication, a mesocosm experiment was performed in the shallow littoral zone of a south Swedish lake, in which nutrient and fish gradients were crossed in a fully factorial design. 2. Food chain theory accurately predicted total biomass development of both phyto‐ and zooplankton. However, separating zooplankton and algae into finer taxonomic groups revealed a variety of responses to both nutrient and fish gradients. 3. That both nutrients and fish are important for phytoplankton dynamics was seen more clearly when viewing each algal group separately, than drawing conclusions only from broad system variables such as chlorophyll a concentration or total phytoplankton biovolume. 4. In some taxa, physiological constraints (e.g. sensitivity to high pH and low concentrations of free CO₂) and differences in competitive ability may be more important for the biomass development than fish predation, grazing by herbivorous zooplankton, and nutrient availability. 5. We conclude that food chain theory accurately predicted responses in system variables, such as total zooplankton or algal biomass, which are shaped by the dynamics of certain strong interactors (‘keystone species’), such as large cladocerans, cyanobacteria and edible algae (<50 μm), whereas responses at finer taxonomic levels cannot be predicted from current theory.     

赤潮过程中“藻-菌”关系研究进展

微生物对促进海洋物质循环,维持水生环境的生态平衡具有重要作用。在赤潮事件中,基于微生物(尤其是细菌)的多样性和重要性,它们与藻类之间的相互关系成为了研究的热点。过去20年里,人们从不同角度对"藻-菌"间的关系进行了探索,包括物理学过程、生物学过程、环境过程以及化学过程。就化学过程而言,它作为一种较早出现的技术,在以往的研究中带给人们许多认识藻菌关系的方法。随着学科的渗透,化学法有了拓展与延伸,为人们认识藻菌关系带来了新的契机。从化学生态学领域来梳理"藻-菌"关系中涉及的现象和行为,包括菌对藻的有益面、菌对藻的有害面、以及藻类应答细菌行为的化学途径;并从信号语言(群体感应、化感作用)的角度来阐释两者之间的互生或克生关系。通过文献综述的方式来解读藻菌关系的互作过程和机理,为认识赤潮的发生和防控方法提供借鉴。     

Using Daphnia physiology to drive food web dynamics: A theoretical revisit of Lotka-Volterra models

The Lotka–Volterra model is the most commonly used framework to describe the dynamics of ecological systems in which two species interact, one as a predator and the other as prey. Theoretical ecologists have since built on variants of these equations, frequently applying them to model the dynamics of algal-herbivore interactions in aquatic systems. In this study, we augment a Lotka–Volterra system by introducing a bioenergetically-explicit, ecophysiological model to examine how variations in resource allocation affect zooplankton growth and subsequently phytoplankton dynamics. Ingested material within a zooplankter's gut is separated into distinct internal congener pools that are used to support physiological processes occurring in a hierarchical direction: neurological functions, energetics, osmoregulatory maintenance, waste management, and finally growth. Consistent with the predictions of the “stoichiometric knife edge” theory, our analysis suggests that a balanced algal congener composition is required to optimize zooplankton internal congener saturations, resulting in a maximal allocation of energy to growth. In examining the advantages rendered by different strategies of minimum and optimum somatic quotas when experiencing phosphorus-enrichment conditions, we show that herbivores with narrow homeostatic bounds and animals with low minimum quotas (or depletion specialists) achieve optimal performance first. Our analysis also predicts patterns of multiple stable equilibria in which the same environmental conditions can be characterized by dramatically different prey-to-predator ratios. Importantly, abrupt shifts from one state to another can be induced not only by short-term variations in food abundance but also by variations in the nutritional quality of the prey. Our predictions have profound implications for connecting microscopic processes with macroscopic patterns and offer new insights into the multitude of factors that modulate food web dynamics.     

Multiple dynamics in a single predator-prey system: experimental effects of food quality.

Recent work with the freshwater zooplankton Daphnia has suggested that the quality of its algal prey can have a significant effect on its demographic rates and life-history patterns. Predator-prey theory linking food quantity and food quality predicts that a single system should be able to display two distinct patterns of population dynamics. One pattern is predicted to have high herbivore and low algal biomass dynamics (high HBD), whereas the other is predicted to have low herbivore and high algal biomass dynamics (low HBD). Despite these predictions and the stoichiometric evidence that many phytoplankton communities may have poor access to food of quality, there have been few tests of whether a dynamic predator-prey system can display both of these distinct patterns. Here we report, to the authors' knowledge, the first evidence for two dynamical patterns, as predicted by theory, in a single predator-prey system. We show that the high HBD is a result of food quantity effects and that the low HBD is a result of food quality effects, which are maintained by phosphorus limitation in the predator. These results provide an important link between the known effects of nutrient limitation in herbivores and the significance of prey quality in predator-prey population dynamics in natural zooplankton communities.     

Field and experimental studies on the combined impacts of cyanobacterial blooms and small algae on crustacean zooplankton in a large,eutrophic, subtropical,Chinese lake

Field and experimental studies were conducted to evaluate the combined impacts of cyanobacterial blooms and small algae on seasonal and long-term changes in the abundance and community structure of crustacean zooplankton in a large, eutrophic, Chinese lake, Lake Chaohu. Seasonal changes of the crustacean zooplankton from 22 sampling stations were investigated during September 2002 and August 2003, and 23 species belonging to 20 genera were recorded. Daphnia spp. dominated in spring but disappeared in mid-summer, while Bosmina coregoni and Ceriodaphnia cornuta dominated in summer and autumn. Both maximum cladoceran density (310 ind. l⁻¹) and biomass (5.2 mg l⁻¹) appeared in autumn. Limnoithona sinensis, Sinocalanus dorrii and Schmackeria inopinus were the main species of copepods. Microcystis spp. were the dominant phytoplankton species and formed dense blooms in the warm seasons. In the laboratory, inhibitory effects of small colonial Microcystis on growth and reproduction of Daphnia carinata were more remarkable than those of large ones, and population size of D. carinata was negatively correlated with density of fresh large colonial Microcystis within a density range of 0–100 mg l⁻¹ (r = −0.82, P< 0.05). Both field and experimental results suggested that seasonal and long-term changes in the community structure of crustacean zooplankton in the lake were shaped by cyanobacterial blooms and biomass of the small algae, respectively, i.e., colonial and filamentous cyanobacteria contributed to the summer replacement of dominant crustacean zooplankton from large Daphnia spp. to small B. coregoni and C. cornuta, while increased small algae might be responsible for the increased abundance of crustacean zooplankton during the past decades.     

典型红树林生态系统藻类多样性及其在生态过程中的作用

藻类是红树林生态系统重要的生物类群, 根据生态习性可分为浮游植物、底栖微藻和大型藻类三个生态类群, 它们在红树林生态系统生物多样性、初级生产、元素循环等方面起着重要作用。但在红树林生态系统中, 关注重点多集中在红树植物和动物, 对其中的藻类重视不够, 且多数研究集中在近20年以及亚洲的红树林区。事实上, 红树林生态系统藻类非常丰富, 其多样性研究有助于深入揭示红树林生态系统的结构与功能。本文介绍了红树林生态系统藻类的组成类群及其重要性, 重点对红树林区浮游植物、底栖硅藻和大型海藻的种类组成、地理分布及其与初级生产力、水质污染、元素循环、碳库形成等生态过程中的作用的研究动态和进展等进行了总结。根据已有研究, 红树林区浮游植物和底栖硅藻的种类数一般为几十到上百种, 其中硅藻在种类和数量上都占绝对优势, 它们是重要的初级生产者、饵料生物和水质污染指示生物; 红树林区底栖大型藻类主要由红藻、绿藻、褐藻、蓝藻组成, 绿藻的种类较多, 红藻在数量上占优势; 藻类是红树林湿地碳库的重要贡献者, 在红树林湿地生态系统碳汇和碳循环中起重要作用。红树林生态系统是个高度动态和异质的系统, 今后应加强红树林藻类多样性的长周期、大尺度变化及不同生境藻类的综合研究, 关注大陆径流和潮汐对藻类多样性和蓝碳的影响, 借助沉积物藻类记录, 探明红树林区藻类的长周期变化, 反演气候变化和人类活动对红树林生态系统的影响过程和机制。     

POSITIVE FEEDBACK AND THE DEVELOPMENT AND PERSISTENCE OF ECOSYSTEM DISRUPTIVE ALGAL BLOOMS1

Harmful algal blooms (HABs) have occurred with increasing frequency in recent years with eutrophication and other anthropogenic alterations of coastal ecosystems. Many of these blooms severely alter or degrade ecosystem function, and are referred to here as ecosystem disruptive algal blooms (EDABs). These blooms are often caused by toxic or unpalatable species that decrease grazing rates by planktonic and benthic herbivores, and thereby disrupt the transfer of nutrients and energy to higher trophic levels, and decrease nutrient recycling. Many factors, such as nutrient availability and herbivore grazing have been proposed to separately influence EDAB dynamics, but interactions among these factors have rarely been considered. Here we discuss positive feedback interactions among nutrient availability, herbivore grazing, and nutrient regeneration, which have the potential to substantially influence the dynamics of EDAB events. The positive feedbacks result from a reduction of grazing rates on EDAB species caused by toxicity or unpalatability of these algae, which promotes the proliferation of the EDAB species. The decreased rates also lower grazer‐mediated recycling of nutrients and thereby decrease nutrient availability. Since many EDAB species are well‐adapted to nutrient‐stressed environments and many exhibit increased toxin production and toxicity under nutrient limitation, positive feedbacks are established which can greatly increase the rate of bloom development and the adverse effects on the ecosystem. An understanding of how these feedbacks interact with other regulating factors, such as benthic/pelagic nutrient coupling, physical forcing, and life cycles of EDAB species provides a substantial future challenge.