Synchronization of cell death in a dinoflagellate population is mediated by an excreted thiol protease

Regulated programmed cell death (PCD) processes have been documented in several phytoplankton species and are hypothesized to play a role in population dynamics. However, the mechanisms leading to the coordinated collapse of phytoplankton blooms are poorly understood. We showed that the collapse of the annual bloom of Peridinium gatunense, an abundant dinoflagellate in Lake Kinneret, Israel, is initiated by CO2 limitation followed by oxidative stress that triggers a PCD-like cascade. We provide evidences that a protease excreted by senescing P. gatunense cells sensitizes younger cells to oxidative stress and may consequently trigger synchronized cell death of the population. Ageing of the P. gatunense cultures was characterized by a remarkable rise in DNA fragmentation and enhanced sensitivity to H2O2. Exposure of logarithmic phase (young) cultures to conditioning media from stationary phase (old) cells sensitized them to H2O2 and led to premature massive cell death. We detected the induction of specific extracellular protease activity, leupeptin-sensitive, in ageing cultures and in lake waters during the succession of the P. gatunense bloom. Partial purification of the conditioned media revealed that this protease activity is responsible for the higher susceptibility of young cells to oxidative stress. Inhibition of the protease activity lowered the sensitivity to oxidative stress, whereas application of papain to logarithmic phase P. gatunense cultures mimicked the effect of the spent media and enhanced cell death. We propose a novel mechanistic framework by which a population of unicellular phytoplankton orchestrates a coordinated response to stress, thereby determine the fate of its individuals.     

Allelopathic effects of the Baltic cyanobacteria Nodularia spumdigena, Aphanizomenon flos-aquae and Anabaena lemmermannii on algal monocultures

Allelopathy, the release of extracellular compounds that inhibit the growth of other microorganisms, may be one factor contributing to the formation and/or maintenance of cyanobacterial blooms. We investigated the allelopathic effects of three cyanobacterial species (Nodularia spumigena, Aphanizomenon flos-aquae and Anabaena lemmermannii) that frequently form mass-occurrences in the Baltic Sea. We exposed monocultures of three phytoplankton species (Thalassiosira weissflogii, Rhodomonas sp. and Prymnesium parvum) to cell-free filtrates of the three cyanobacteria, and quantified allelopathic effects with cell counts. We also investigated the role of the growth phase of cyanobacteria in their allelopathy, by comparing the effects of an exponential and a stationary phase culture of N. spumigena. All tested cyanobacteria inhibited the growth of Rhodomonas sp., but none of them affected P. parvum. The effects on T. weissflogii were more variable, and they were amplified by repeated filtrate additions compared to a single filtrate addition. N. spumigena was more allelopathic in exponential than in stationary growth phase, whereas the culture filtrate was more hepatotoxic in stationary phase. Hepatotoxins were thus probably not involved in the allelopathic effects, which is also indicated by the allelopathic properties of the non-toxic A. flos-aquae and A. lemmermannii. The results demonstrate that the common Baltic cyanobacteria affect some coexisting phytoplankton species negatively. Allelopathy may therefore play a role in interspecific competition and contribute to cyanobacterial bloom maintenance.     

Paradoxically, prior acquisition of antioxidant activity enhances oxidative stress-induced cell death

Oxidative stress has been implicated in the induction of programmed cell death in a wide variety of organisms. Acquiring antioxidant capacity is thought to enhance the viability of cells challenged by a subsequent oxidative stress. Counter-intuitively, we show that in two phytoplankton species, Chlamydomonas reinhardtii and Peridinium gatunense , representing the green and red plastid lineages, oxidative stress induced cell death in cultures that already possessed high antioxidant activity but not in cells that exhibited low activity. Cell death of low antioxidant possessing cultures was markedly enhanced by the addition of dehydroascorbate, a product of ascorbate peroxidase (APX), but not of ascorbate or reduced glutathione, and was preceded by increased metacaspase expression and activity. These data suggested that the level of APX and its products, strongly upregulated by oxidative stress, serves as a possible surveillance signal, reporting that the cells already experienced an earlier oxidative stress. Our data presents a novel role of APX in antioxidant activity and response to oxidative stress in photosynthetic microorganisms. Elimination of cysts production by phytoplankton cells that were already damaged by oxidative stress (indicated by the rise in oxidized proteins) as the inoculum for the following year's population may be the evolutionary trigger for this phenomenon.     

Allelopathic effects of Microcystis aeruginosa on green algae and a diatom: Evidence from exudates addition and co-culturing

Allelopathic interactions among phytoplankton species are regarded as one of the important factors contributing to phytoplankton species competition and succession. The role and extent of allelopathic effects of blooming freshwater cyanobacteria on other phytoplankton species in eutrophied waters, however, are still unknown. We examined the allelopathic effect of Microcystis aeruginosa on two common green algae (Scenedesmus quadricauda, Chlorella pyrenoidosa) and a diatom (Cyclotella meneghiniana) by adding exudates from different growth phases and in co-culture tests. Exudates of M. aeruginosa from the exponential growth phase and the stationary phase significantly inhibited the growth of S. quadricauda, C. pyrenoidosa and C. meneghiniana, whereas those from the decline phase increased their growth. The presence of M. aeruginosa extremely inhibited the growth of all tested species in co-cultures within 24 h. Our results indicate that under the tested environmental conditions (25 °C, light 80 μmol quanta m⁻² s⁻¹, manual shaking twice a day), allelopathic effects of M. aeruginosa on other phytoplankton species can significantly contribute to their competitive success.     

Temporary cyst formation in phytoplankton: a response to allelopathic competitors?

Competition among phytoplankton for limiting resources may involve direct or indirect interactions. A direct interaction of competitors is the release of chemicals that inhibit other species, a process known as allelopathy. Here, we investigated the allelopathic effect of three toxic microalgae species (Alexandrium tamarense, Karenia mikimotoi and Chrysochromulina polylepis) on a natural population of the dinoflagellate Scrippsiella trochoidea. Our major findings were that in addition to causing death of S. trochoidea cells, the allelopathic species also induced the formation of temporary cysts in S. trochoidea. Because cysts were not lysed, encystment may act as a defence mechanism for S. trochoidea to resist allelochemicals, especially when the allelopathic effect is moderate. By forming temporary cysts, S. trochoidea may be able to overcome the effect of allelochemicals, and thereby have an adaptive advantage over other organisms unable to do so.     

Competing phytoplankton undermines allelopathy of a bloom-forming dinoflagellate

Biotic interactions in the plankton can be both complex and dynamic. Competition among phytoplankton is often chemically mediated, but no studies have considered whether allelopathic compounds are modified by biotic interactions. Here, we show that compounds exuded during Karenia brevis blooms were allelopathic to the cosmopolitan diatom Skeletonema costatum, but that bloom allelopathy varied dramatically among collections and years. We investigated several possible causes of this variability and found that neither bloom density nor concentrations of water-borne brevetoxins correlated with allelopathic potency. However, when we directly tested whether the presence of competing phytoplankton influenced bloom allelopathy, we found that S. costatum reduced the growth-inhibiting effects of bloom exudates, suggesting that S. costatum has a mechanism for undermining K. brevis allelopathy. Additional laboratory experiments indicated that inducible changes to K. brevis allelopathy were restricted to two diatoms among five sensitive phytoplankton species, whereas five other species were constitutively resistant to K. brevis allelopathy. Our results suggest that competitors differ in their responses to phytoplankton allelopathy, with S. costatum exhibiting a previously undescribed method of resistance that may influence community structure and alter bloom dynamics.     

活性氧在植物抗病反应中的作用（综述）

介绍植物与病原体相互作用中的氧化爆发现象,总结了氧化爆发的几个特点。重点讨论在植物与病原体相互作用的活性氧产生机制,概述活性氧在植物抗病反应中的可能作用。     

Harmful algal blooms of allelopathic microalgal species: The role of eutrophication

The ability of certain harmful algal species to produce and release chemicals that inhibit the growth of co-occurring phytoplankton species, here considered as allelopathy, is closely associated with competition for limiting nutrient resources. Many phytoplankton cells are known to release elevated amounts of organic compounds under nutrient limitation. Eutrophication alters the nitrogen-to-phosphorus balance and, when nutrient availability is unbalanced, nutrient limitation may result. Algal species that can compete successfully for available growth-limiting nutrient(s) have the potential to become dominant and form blooms. The stress conditions imposed by the shifted nutrient supply ratios can, in some algae, stimulate production of allelochemicals that inhibit potential competitors. Thus, under cultural eutrophication, altered nutrient (N, P) ratios and limiting nutrient supplies can stimulate increased production of allelochemicals, including toxins, by some algal species and accentuate the adverse effects of these substances on other algae. Future investigation on the characterization of the chemical compounds involved in the allelopathic process are needed to advance the study of the mode of action of phytoplankton allelochemicals.     

Comparing grazing by Dreissena polymorpha on phytoplankton in the presence of toxic and non-toxic cyanobacteria

SUMMARY 1. The feeding behaviour of the zebra mussel ( Dreissena polymorpha ) was studied in the laboratory on different combinations of food, including a green alga ( Chlamydomonas reinhardtii ) and toxic and non-toxic strains of the cyanobacterium Microcystis aeruginosa .
2. The highest clearance rate of phytoplankton by zebra mussels was found when the mussels were feeding on a mixture of Chlamydomonas and non-toxic Microcystis , the lowest on a mixture of Chlamydomonas and toxic Microcystis .
3. The differences found in the clearance rates between food combinations can be partly explained by the production of pseudofaeces containing live phytoplankton cells. Zebra mussels expelled significantly more live phytoplankton cells in the presence of toxic Microcystis than in the presence of non-toxic Microcystis . The pseudofaeces contained predominantly live Chlamydomonas cells. Proportionately much less live Microcystis cells were encountered in the pseudofaeces.
4. Consequently, grazing of zebra mussels on a combination of Chlamydomonas and Microcystis may finally result in a dominance of Chlamydomonas over Microcystis . The presence of toxic Microcystis may even strengthen this shift.     

Allelopathic inhibitory effect of Myriophyllum aquaticum (Vell.) Verdc. on Microcystis aeruginosa and its physiological mechanism

Eutrophication and algal blooms are the most serious environmental problems in the world, and biological tools, especially the allelopathic inhibitory effects of aquatic macrophytes on phytoplankton growth have been receiving world-wide attention. In our experiments, the allelopathic inhibitory effect of Myriophyllum aquaticum culture water on Microcystis aeruginosa and its eco-physiological mechanism were investigated by initial addition assays and continuous addition assays. The results showed that the growth of M. aeruginosa was markedly inhibited by M. aquaticum culture water. Compared with initial addition assays, M. aquaticum exhibited stronger inhibitory potential on M. aeruginosa by continuous addition assays, indicating that allelopathic compounds might be excreted continuously by M. aquaticum, and the inhibitory effects would be cumulative. We also found that the relative content of chlorophyll a (Chl a), phycocyanin (PC) and allophycocyanin (APC) of M. aeruginosa decreased to 52.7%, 15.3% and 7.6% respectively after being treated by M. aquaticum culture water for 5 days, and phycobiliprotein (especially APC) decreased more than Chl a. These results suggest that the phycobiliprotein would be the target of allelopathic inhibition of M. aquaticum on M. aeruginosa, and a new macrophyte to control cyanobacterial blooms would be found.     

粉绿狐尾藻（Myriophyllum aquaticum）对铜绿微囊藻（Microcystis aeruginosa）的化感抑制效应及其生理机制

由于生物法,尤其是水生植物化感控藻方法在治理水体富营养化及水华现象中具有多方面的优势,近年来该领域的理论和技术研究备受世界各国关注.采用初始添加种植水和连续添加种植水的方法,研究挺水型粉绿狐尾藻对铜绿微囊藻的化感抑藻效应,并从光合作用的角度探讨其对铜绿微囊藻化感抑制的生理生态机制及作用靶点.研究结果表明:粉绿狐尾藻能够分泌某些化感物质有效抑制铜绿微囊藻的正常生长,其抑藻效应是通过连续释放某些化感物质作用于铜绿微囊藻而实现的,且具有累积性;粉绿狐尾藻分泌化感物质对铜绿微囊藻的Chl a、PC及APC的损伤程度存在差异(如处理5 d后,Chl a、PC、APC的相对含量分别降至52.7%、15.3%、7.6%),其中藻胆蛋白(尤其是APC)比Chl a 更为敏感,说明藻胆蛋白是粉绿狐尾藻化感物质抑制铜绿微囊藻的关键靶点.研究为水生植物化感控藻技术的发展提供了新的材料,并有助于深入了解水体生态系统的化学生态作用及其机制.     

Allelopathic inhibition of phytoplankton by exudates from Stratiotes aloides

The allelopathic potential of exudates from the aquatic macrophyte Stratiotes aloides on the growth of phytoplankton was investigated. A selection of phytoplankton species, occurring in habitats similar to that of Stratiotes, was used: two cyanobacterial strains (toxic and non-toxic Microcystis aeruginosa), one green alga (Scenedesmus obliquus) and one eustigmatophyte (Nannochloropsis limnetica). The results indicate allelopathic effects of Stratiotes on phytoplankton in six of the eight cases, expressed in an extended duration of the initial biovolume doubling time. The overall inhibitory effect (8–51%) was strain-specific for the two cyanobacteria. We also studied the effect of irradiance on the allelopathic potential of exudates from Stratiotes. Irradiance influenced the response of Scenedesmus only. The inhibitory effect of Stratiotes exudates on the growth of this green alga was stronger at 35 μmol m⁻² s⁻¹ than at 105 μmol m⁻² s⁻¹. We conclude that Stratiotes has allelopathic effects on phytoplankton, and that irradiance can, but does not always determine the extent of the allelopathic inhibition. In our experiments, the sensitivity of cyanobacteria to Stratiotes exudates was not higher than for other phytoplankton strains, but within cyanobacteria, the toxic strain was more sensitive than the non-toxic one.     

Responses of Phyto- and Zooplankton Communities to Prymnesium polylepis (Prymnesiales) Bloom in the Baltic Sea

A large bloom of Prymnesium polylepis occurred in the Baltic Sea during the winter 2007 – spring 2008. Based on numerous reports of strong allelopathic effects on phytoplankton exerted by P. polylepis and its toxicity to grazers, we hypothesized that during this period negative correlations will be observed between P. polylepis and (1) main phytoplankton groups contributing to the spring bloom (i.e., diatoms and dinoflagellates), and (2) zooplankton growth and abundance. To test these hypotheses, we analyzed inter-annual variability in phytoplankton and zooplankton dynamics as well as growth indices (RNA∶DNA ratio) in dominant zooplankton in relation to the Prymnesium abundance and biomass. Contrary to the hypothesized relationships, no measurable negative responses to P. polylepis were observed for either the total phytoplankton stocks or the zooplankton community. The only negative response, possibly associated with P. polylepis occurrence, was significantly lower abundance of dinoflagellates both during and after the bloom in 2008. Moreover, contrary to the expected negative effects, there were significantly higher total phytoplankton abundance as well as significantly higher winter abundance and winter-spring RNA∶DNA ratio in dominant zooplankton species in 2008, indicating that P. polylepis bloom coincided with favourable feeding conditions for zooplankton. Thus, primary consumers, and consequently also zooplanktivores (e.g., larval fish and mysids), may benefit from haptophyte blooms, particularly in winter, when phytoplankton is scarce.     

A 3D Individual-Based Model to Study Effects of Chemotaxis,Competition and Diffusion on the Motile-Phytoplankton Aggregation

In this paper, we develop a 3D-individual-based model (IBM) to understand effect of various small-scale mechanisms in phytoplankton cells, on the cellular aggregation process. These mechanisms are: spatial interactions between cells due to their chemosensory abilities (chemotaxis), a molecular diffusion and a demographical process. The latter is considered as a branching process with a density-dependent death rate to take into account the local competition on resources. We implement the IBM and simulate various scenarios under real parameter values for phytoplankton cells. To quantify the effects of the different processes quoted above on the spatial and temporal distribution of phytoplankton, we used two spatial statistics: the Clark–Evans index and the group belonging percentage. Our simulation study highlights the role of the branching process with a weak-to-medium competition in reinforcing the aggregating structure that forms from attraction mechanisms (under suitable conditions for diffusion and attraction forces), and shows by contrast that aggregations cannot form when competition is high.     

Can filter‐feeding fishes improve water quality in lakes?

1.  In this paper we examine the potential of a cichlid fish species ( Sarotherodon galilaeus ) to both maintain positive growth rates through filter-feeding on phytoplankton and improve water quality in Lake Kinneret through suppression of dinoflagellate ( Peridinium gatunense ) blooms.
2.  Seasonal plankton consumption by S. galilaeus from Lake Kinneret was examined experimentally by monitoring changes in plankton assemblages during 24 h in 5-m³ mesocosms containing varying densities of fish. Taxon-specific grazing rates ranged from 0 to 17 mg g_fish day^–1, with mean total consumption of 1.6% fish body weight per day. During the spring bloom of P. gatunense , S. galilaeus consumed mostly (94%) netphytoplankton (≥20 μm). The remaining 6% consisted mostly of nanophytoplankton (<20 μm). During the summer and fall, net- and nanophytoplankton accounted for 54 and 42%, respectively, of the diet of S. galilaeus . Zooplankton and flagellated and ciliated protozoans made up the remaining 4%.
3.  Simulations using a fish bioenergetics model indicated that consumption rates ( C ) were near maximum in spring (90% C _max), while consumption was reduced in summer-fall (59% C _max). Sarotherodon galilaeus obtains sufficient energy through filter-feeding year-round, although most growth (≥60%) occurs during the spring P. gatunense bloom.
4.  Despite efficient feeding on P. gatunense and nanophytoplankton by S. galilaeus , estimates of instantaneous plankton mortality caused by ingestion were two orders of magnitude lower than maximum potential plankton growth rates. Thus the potential for the S. galilaeus population in Lake Kinneret to positively affect water quality through algal suppression is low.     

Searching for allelopathic effects of submerged macrophytes on phytoplankton—state of the art and open questions

Allelopathy, here defined as biochemical interactions between aquatic primary producers, has always been intriguing as a process explaining the dominance of certain plant or algal species over others. Negative chemical interference has been invoked as one of the steering mechanisms behind mutual dominance of either submerged macrophytes or phytoplankton in shallow eutrophic lakes. Yet, despite much effort, convincing evidence for allelopathic interactions in situ is still missing. Also, laboratory approaches often lack reality. Inspired by a series of talks at the Shallow Lakes 2005 meeting in Dalfsen, the Netherlands, we argue that there is circumstantial but strong evidence that allelopathic interference between submerged macrophytes and phytoplankton may indeed exist in aquatic ecosystems despite the problems associated with research in this field. We first discuss experimental approaches combining laboratory and field studies, based on examples presented at this meeting. We then discuss the impact of nutrient status of both producing and target organism and biotic factors such as herbivory or pathogens that might affect allelopathy. Further topics are the potential seasonality of effects and the species-specificity of certain allelochemicals. We conclude with some thoughts why a final proof for allelopathy in situ might remain difficult or even inaccessible in some cases, and why we nevertheless should not abandon this idea.     

Use of phytoplankton-derived dissolved organic carbon by different types of bacterioplankton

Phytoplankton and heterotrophic prokaryotes are major components of the microbial food web and interact continuously: heterotrophic prokaryotes utilize the dissolved organic carbon derived from phytoplankton exudation or cell lysis (DOCp), and mineralization by heterotrophic prokaryotes provides inorganic nutrients for phytoplankton. For this reason, these communities are expected to be closely linked, although the study of the interactions between them is still a major challenge. Recent studies have presented interactions between phytoplankton and heterotrophic prokaryotes based on coexistence or covariation throughout a time-series. However, a real quantification of the carbon flow within these networks (defined as the interaction strength, IS) has not been achieved yet. This is critical to understand the selectivity degree of bacteria responding to specific algal DOCp. Here we used microautoradiography to quantify the preferences of the major heterotrophic prokaryote phylogenetic groups on DOC derived from several representative phytoplankton species, and expressed these preferences as an IS value. The distribution of the ISs was not random but rather skewed towards weak interactions, in a similar way as the distributions described for stable complex non-microbial ecosystems, indicating that there are some cases of high specificity on the use of specific algal DOCp by some bacterial groups, but weak interactions are more common and may be relevant as well. The variety of IS patterns observed supports the view that the vast range of different resources (different types of organic molecules) available in the sea selects and maintains the high levels of diversity described for marine bacterioplankton.     

A mannan binding lectin is involved in cell-cell attachment in a toxic strain of Microcystis aeruginosa

Microcystin, a hepatotoxin that represents a serious health risk for humans and livestock, is produced by the bloom-forming cyanobacterium Microcystis aeruginosa in freshwater bodies worldwide. Here we describe the discovery of a lectin, microvirin (MVN), in M. aeruginosa PCC7806 that shares 33% identity with the potent anti-HIV protein cyanovirin-N from Nostoc ellipsosporum. Carbohydrate microarrays were employed to demonstrate the high specificity of the protein for high-mannose structures containing alpha(1-->2) linked mannose residues. Lectin binding analyses and phenotypic characterizations of MVN-deficient mutants suggest that MVN is involved in cell-cell recognition and cell-cell attachment of Microcystis. A binding partner of MVN was identified in the lipopolysaccharide fraction of M. aeruginosa PCC7806. MVN is differentially expressed in mutants lacking the hepatotoxin microcystin. Additionally, MVN-deficient mutants contain much lower amounts of microcystin than the wild-type cells. We discuss a possible functional correlation between microcystin and the lectin and possible implications on Microcystis morphotype formation. This study provides the first experimental evidence that microcystins may have an impact on Microcystis colony formation that is highly important for the competitive advantage of Microcystis over other phytoplankton species.     

Seasonal dependence and functional implications of macrophyte–phytoplankton allelopathic interactions

Invasive plant species such as Ludwigia hexapetala might have a competitive advantage if they produce allelopathically active compounds against primary producers. Both phytoplankton and plant community structure may be affected due to different, species‐specific sensitivity to allelochemicals. Moreover, such allelopathic interactions could vary over the year depending on (i) the plant's phenological stage and (ii) the abilities of the native macrophytes to suppress—or the non‐native macrophytes to stimulate—the non‐native macrophyte population. We tested the allelopathic effects of aqueous leaf extracts of L. hexapetala on the photosynthetic activity of three target phytoplankton strains (Scenedesmus communis, a toxic Microcystis aeruginosa strain and a non‐toxic Microcystis aeruginosa strain) over three seasons of development (spring, summer and autumn). We also tested seasonal allelopathic effects of aqueous leaf extracts of both L. hexapetala (i.e. the non‐native invasive species) and the native Mentha aquatica on L. hexapetala seed germination. Finally, we identified three main secondary compounds present in the aqueous leaf extracts of L. hexapetala and we tested each individual compound on the phytoplankton's photosynthetic activity and on L. hexapetala seed germination. We observed marked seasonal and species‐specific patterns of L. hexapetala allelopathy on phytoplankton. The photosynthetic activities of S. communis and the toxic M. aeruginosa strain were stimulated by L. hexapetala aqueous leaf extracts in autumn and spring, respectively, whereas the non‐toxic M. aeruginosa strain was strongly inhibited in these two seasons. In summer, photosynthesis of all phytoplankton strains was inhibited. The germination rate of L. hexapetala seeds was stimulated by both L. hexapetala and M. aquatica aqueous leaf extracts, especially in summer, concomitant with the strong negative effects observed on the three phytoplankton strains. Three flavonoid glycosides (myricitrin, prunin and quercitrin) were identified as the main secondary compounds present in the L. hexapetala aqueous leaf extracts. The photosynthetic activity of S. communis was slightly stimulated by the three compounds. The photosynthetic activity of the toxic M. aeruginosa strain was stimulated by myricitrin and quercitrin, whereas that of the non‐toxic M. aeruginosa strain was inhibited by prunin. Finally, the germination rate and the germination velocity of L. hexapetala seeds were stimulated by myricitrin and prunin. These findings suggest that L. hexapetala could favour the photosynthetic activity of toxic cyanobacteria in spring and reduce their photosynthetic activity in summer, potentially leading to drastic changes in the phytoplankton communities and therewith ecological functioning of invaded ponds. Moreover, the stimulation of its seed germination could give a strong competitive advantage to L. hexapetala, thus promoting its invasiveness.     

Allelopathy of Aquatic Autotrophs

Allelopathy in aquatic environments may provide a competitive advantage to angiosperms, algae, or cyanobacteria in their interaction with other primary producers. Allelopathy can influence the competition between different photoautotrophs for resources and change the succession of species, for example, in phytoplankton communities. Field evidence and laboratory studies indicate that allelopathy occurs in all aquatic habitats (marine and freshwater), and that all primary producing organisms (cyanobacteria, micro- and macroalgae as well as angiosperms) are capable of producing and releasing allelopathically active compounds. Although allelopathy also includes positive (stimulating) interactions, the majority of studies describe the inhibitory activity of allelopathically active compounds. Different mechanisms operate depending on whether allelopathy takes place in the open water (pelagic zone) or is substrate associated (benthic habitats). Allelopathical interactions are especially common in fully aquatic species, such as submersed macrophytes or benthic algae and cyanobacteria. The prevention of shading by epiphytic and planktonic primary producers and the competition for space may be the ultimate cause for allelopathical interactions. Aquatic allelochemicals often target multiple physiological processes. The inhibition of photosynthesis of competing primary producers seems to be a frequent mode of action. Multiple biotic and abiotic factors determine the strength of allelopathic interactions. Bacteria associated with the donor or target organism can metabolize excreted allelochemicals. Frequently, the impact of surplus or limiting nutrients has been shown to affect the overall production of allelochemicals and their effect on target species. Similarities and differences of allelopathic interactions in marine and freshwater habitats as well as between the different types of producing organisms are discussed. Referee: Dr. Friedrich Jüttner, Universität Zürich-Limnologische Station, Institut für Pflanzen biologie, Universität Zürich, Seestrasse 187, Ch-8802 Klichberg ZH, Switzerland