Allelopathy prevents competitive exclusion and promotes phytoplankton biodiversity

It has been hypothesized that allelopathy can prevent competitive exclusion and promote phytoplankton diversity in aquatic ecosystems, where numerous species coexist on a limited number of resources. However, experimental proof‐of‐principle is not available to support this hypothesis. Here we present the first experimental evidence to support this hypothesis by demonstrating that allelopathy promotes the coexistence of two phytoplankton species, Ankistrodesmus falcatus and Oscillatoria sp., that compete for a single limiting nutrient. By performing long‐term competition experiments in nitrate‐limited continuous cultures, and by describing the population dynamics using a mechanistic model, we demonstrate that when allelopathy comes into play, one of the following outcomes is possible depending on the relative initial abundances of the species: dominance of the stronger competitor for nitrate (the non‐allelopathic species), oscillatory coexistence, or dominance of the weaker competitor (the allelopathic species). Our model analysis revealed that sustained oscillatory coexistence of the two species would be a common outcome of this experiment. Our study confirms for the first time, based on laboratory experiments combined with mechanistic models, that allelopathy can alter the predicted outcome of inter‐specific competition in a nutrient‐limited environment and increase the potential for the coexistence of more species than resources, thereby contributing to the identification of endogenous mechanisms that explain the extreme diversity of phytoplankton communities.     

Competing Effects of Toxin-Producing Phytoplankton on Overall Plankton Populations in the Bay of Bengal

The coexistence of a large number of phytoplankton species on a seemingly limited variety of resources is a classical problem in ecology, known as ‘the paradox of the plankton’. Strong fluctuations in species abundance due to the external factors or competitive interactions leading to oscillations, chaos and short-term equilibria have been cited so far to explain multi-species coexistence and biodiversity of phytoplankton. However, none of the explanations has been universally accepted. The qualitative view and statistical analysis of our field data establish two distinct roles of toxin-producing phytoplankton (TPP): toxin allelopathy weakens the interspecific competition among phytoplankton groups and the inhibition due to ingestion of toxic substances reduces the abundance of the grazer zooplankton. Structuring the overall plankton population as a combination of nontoxic phytoplankton (NTP), toxic phytoplankton, and zooplankton, here we offer a novel solution to the plankton paradox governed by the activity of TPP. We demonstrate our findings through qualitative analysis of our sample data followed by analysis of a mathematical model.     

Ingredients for protist coexistence: competition, endosymbiosis and a pinch of biochemical interactions

1. The interaction between mutualism, facilitation or interference and exploitation competition is of major interest as it may govern species coexistence. However, the interplay of these mechanisms has received little attention. This issue dates back to Gause, who experimentally explored competition using protists as a model [Gause, G.F. (1935) Vérifications expérimentales de la théorie mathématique de la lutte pour la vie. Actualités Scientifiques et Industrielles, 277]. He showed the coexistence of Paramecium caudatum with a potentially allelopathic species, Paramecium bursaria. 2. Paramecium bursaria hosts the green algae Chlorella vulgaris. Therefore, P. bursaria may benefit from carbohydrates synthesised by the algae. Studying endosymbiosis with P. bursaria is possible as it can be freed of its endosymbiont. In addition, C. vulgaris is known to produce allelochemicals, and P. bursaria may benefit also from allelopathic compounds. 3. We designed an experiment to separate the effects of resource exploitation, endosymbiosis and allelopathy and to assess their relative importance for the coexistence of P. bursaria with a competitor that exploits the same resource, bacteria. The experiment was repeated with two competitors, Colpidium striatum or Tetrahymena pyriformis. 4. Results show that the presence of the endosymbiont enables the coexistence of competitors, while its loss leads to competitive exclusion. These results are in agreement with predictions based on resource equilibrium density of monocultures (R*) supporting the idea that P. bursaria's endosymbiont is a resource provider for its host. When P. bursaria and T. pyriformis coexist, the density of the latter shows large variation that match the effects of culture medium of P. bursaria. Our experiment suggests these effects are because of biochemicals produced in P. bursaria culture. 5. Our results expose the hidden diversity of mechanisms that underlie competitive interactions. They thus support Gauses's speculation (1935) that allelopathic effects might have been involved in his competition experiments. We discuss how a species engaged both in competition for a resource and in costly interference such as allelopathy may counterbalance these costs with a resource-provider endosymbiont.     

Toxic phytoplankton as a keystone species in aquatic ecosystems: stable coexistence to biodiversity

The effect of allelochemicals released by toxic species in plankton community is often taken into account to reveal plankton biodiversity. Using a minimal chemostat model we show that the interaction between toxic and non‐toxic phytoplankton species with changing competitive effects among species due to allelopathy helps to promote the stable coexistence of many species on a single resource and hence can solve the paradox of plankton. We emphasize toxic phytoplankton as a keystone species that strongly uncovers its allelochemicals on other non‐toxic phytoplankton and enhances the species persistence and diversity in aquatic ecosystems. In addition, we analyze the consistency of ecosystem functioning and species diversity using a number of approaches, such as sampling hypothesis with selection and complementarity effects, cascading extinction–reinvasion, and examining system dynamics at different enrichment levels and toxicity. Our results suggest that chemostats with one toxic and one or more nontoxic phytoplankton species can be used for the experimental verification of the stable coexistence of many species on a single resource in aquatic ecology.     

Interactions between allelopathic properties and growth kynetics in four freshwater phytoplankton species studied by model simulations

We chose four species of freshwater phytoplankton: the chlorophyceans Ankistrodesmus falcatus, Chlamydomonas reinhardtii and Selenastrum capricornutum, and the cyanobacteria Oscillatoria sp. in order to study their competitive abilities for nitrate and their allelopathic properties. We parameterized models of nitrate uptake and growth with laboratory experiments. According to them, the species were ranked (from the best to the worst competitors): S. capricornutum, C. reinhardtii, A. falcatus and Oscillatoria sp. C. reinhardtii and Oscillatoria sp. were previously reported as allelopathic. In the present work, Oscillatoria sp. was allelopathic only against A. falcatus. However, none of our species was sensitive to C. reinhardtii. Additionally, we found an unknown allelopathic effect of A. falcatus against Oscillatoria sp. Our findings point out the high specificity of allelopathic interactions. With these data, we constructed a model of interspecific competition for nitrate, including allelopathic interactions. By performing model simulations, we studied how three factors influence the outcome of competition: relative abundance of competing species, resistance to allelopathy, and nitrate concentration. Our simulations showed that the initial ratio of species abundances will significantly determine the outcome of competition. If the worst competitor was the allelopathic species, the more it needs to outnumber the competing species, unless it is very sensitive to allelopathy (not defended). Nitrate has an important influence, showing a non-intuitive outcome of competition experiments at low nitrate concentrations, where the worst competitor (allelopathic species) wins competition in the majority of cases, whereas at intermediate concentrations, the better competitor dominates except for unfavorable ratios of abundances. With the increased amounts of nitrate, conditions again favor the worst competitor (the stronger allelopathic species). Despite the potential for two species coexistence showed by previous theoretical analysis of systems was similar to ours, our simulations did not detect this outcome. We hypothesized that this is due to the strong allelopathic effect of Oscillatoria sp.     

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.     

Resisting annihilation: relationships between functional trait dissimilarity,assemblage competitive power and allelopathy

Allelopathic species can alter biodiversity. Using simulated assemblages that are characterised by neutrality, lumpy coexistence and intransitivity, we explore relationships between within‐assemblage competitive dissimilarities and resistance to allelopathic species. An emergent behaviour from our models is that assemblages are more resistant to allelopathy when members strongly compete exploitatively (high competitive power). We found that neutral assemblages were the most vulnerable to allelopathic species, followed by lumpy and then by intransitive assemblages. We find support for our modeling in real‐world time‐series data from eight lakes of varied morphometry and trophic state. Our analysis of this data shows that a lake's history of allelopathic phytoplankton species biovolume density and dominance is related to the number of species clusters occurring in the plankton assemblages of those lakes, an emergent trend similar to that of our modeling. We suggest that an assemblage's competitive power determines its allelopathy resistance.     

Spatial polymorphism of bacteriocins and other allelopathic traits

Summary Many bacterial species carry plasmids that encode both the production of a highly specific toxin (bacteriocin) that kills competitors of the same species and immunity to that toxin. A great diversity of bacteriocins is produced within a single species. I present a model for the dynamics of competition between allelopathic and susceptible types. The model applies to most kinds of allelopathic competition. My primary goal is, however, to explain the widespread genetic polymorphism for bacteriocins. The model includes competition for scarce resources, competition through toxin production, spatial diffusion of individuals and toxins at varying rates and spatial variation in habitat quality. I draw three main conclusions from this reaction—diffusion model. (1) Polymorphism of toxin producers and susceptibles cannot be maintained within a single spatial location when individuals and the toxin mix randomly. (2) Susceptibles are generally favoured in poor habitats, where the rate of resource competition per interaction increases relative to the resource-independent rate of toxic killing. By contrast, toxic producers are generally favoured in good habitats, where the rate of resource competition is lower. (3) Spatial variation in habitat quality can lead to spatial polymorphism; susceptibles tend to win in poor habitats and producers tend to win in good habitats.     

Mutualistic relationships between phytoplankton and bacteria caused by carbon excretion from phytoplankton

For the competition system of phytoplankton and bacteria through inorganic phosphorus, our mathematical model showed that mutualistic relationships between them could occur due to production and consumption of extracellular organic carbon by phytoplankton and bacteria. In our model, phytoplankton are limited in their growth by light and phosphorus, and bacteria are limited in their growth by phosphorus and carbon released from phytoplankton. We adopted permanence as a criterion of the coexistence in mathematical analysis, and led necessary conditions of permanence in the model. Under these coexistence conditions, we estimated the strength of total effects of interactions between phytoplankton and bacteria at the steady state by press perturbation method. The results of this estimation indicated the mutualistic interactions between phytoplankton and bacteria. This suggests that mutualistic situation could occur due to the introduction of carbon flow from phytoplankton to bacteria, even if phytoplankton and bacteria compete with each other through common resource, inorganic phosphorus.     

Allelopathic adaptation can cause competitive coexistence

The maintenance of plant diversity is often explained by the ecological and evolutionary consequences of resource competition. Recently, the importance of allelopathy for competitive interactions has been recognized. In spite of such interest in allelopathy, we have few theories for understanding how the allelopathy influences the ecological and evolutionary dynamics of competing species. Here, I study the coevolutionary dynamics of two competing species with allelopathy in an interspecific competition system, and show that adaptive trait dynamics can cause cyclic coexistence. In addition, very fast adaptation such as phenotypic plasticity is likely to stabilize the population cycles. The results suggest that adaptive changes in allelopathy can lead to cyclic coexistence of plant species even when their ecological characters are very similar and interspecific competition is stronger than intraspecific competition, which should destroy competitive coexistence in the absence of adaptation.     

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.     

An ericoid shrub plays a dual role in recruiting both pines and their fungal symbionts along primary succession gradients

Relative importance of positive and negative interactions between plant species may change along disturbance and resource gradients. Positive interactions are suggested to prevail in low resource, low productivity (high stress) conditions and negative interactions in high resource availability. A dwarf shrub, mountain crowberry Empetrum nigrum ssp. hermaphroditum, is known to have allelopathic impacts on both Scots pine Pinus sylvestris and its ectomycorrhizal symbionts. We aimed to study if the outcome of Empetrum impacts on Scots pine changes along primary succession gradients on the dune shores of Bothnian Bay, in Finland, where abiotic stress gradually changes to biotic stress along the succession. We found that Empetrum may act as a facilitator despite its allelopathic effects, since the proportion of Scots pine seedlings established in Empetrum patches was higher than in patches without the shrub in early and mid succession stages, whereas patches without Empetrum were preferred in late succession. The amount of mycelial fungal biomass (ergosterol) in the soil in the vicinity of the seedling roots was higher in Empetrum patches than in patches without Empetrum and it increased along the succession gradient. Proportion of pine root tips colonised by suilloid morphotypes with abundant external mycelia and the diversity of ectomycorrhizal morphotypes were higher in mid successional stage in Empetrum patches compared to patches without Empetrum. Our results suggest that in the harsh physical conditions of the dune shore Empetrum facilitates pine seedling establishment in the early and mid stages of succession by providing mechanical and physical shelter whereas in late succession negative interactions (competition and allelopathy) between the shrub and the pine are dominating. To our knowledge we present the first finding that an ericoid mycorrhizal shrub could enhance both the performance of the ectomycorrhizal host tree and the tree's fungal symbionts.     

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     

Separation of allelopathy from resource competition using rice/barnyardgrass mixed-cultures

Plant-plant interference is the combined effect of allelopathy, resource competition, and many other factors. Separating allelopathy from resource competition is almost impossible in natural systems but it is important to evaluate the relative contribution of each of the two mechanisms on plant interference. Research on allelopathy in natural and cultivated plant communities has been hindered in the absence of a reliable method that can separate allelopathic effect from resource competition. In this paper, the interactions between allelopathic rice accession PI312777, non-allelopathic rice accession Lemont and barnyardgrass were explored respectively by using a target (rice)-neighbor (barnyardgrass) mixed-culture in hydroponic system. The relative competitive intensity (RCI), the relative neighbor effect (RNE) and the competitive ratio (CR) were used to quantify the intensity of competition between each of the two different potentially allelopathic rice accessions and barnyardgrass. Use of hydroponic culture system enabled us to exclude any uncontrolled factors that might operate in the soil and we were able to separate allelopathy from resource competition between each rice accession and barnyardgrass. The RCI and RNE values showed that the plant-plant interaction was positive (facilitation) for PI312777 but that was negative (competition) for Lemont and barnyardgrass in rice/barnyardgrass mixed-cultures. The CR values showed that one PI312777 plant was more competitive than 2 barnyardgrass plants. The allelopathic effects of PI312777 were much more intense than the resource competition in rice/barnyardgrass mixed cultures. The reverse was true for Lemont. These results demonstrate that the allelopathic effect of PI312777 was predominant in rice/barnyardgrass mixed-cultures. The most significant result of our study is the discovery of an experimental design, target-neighbor mixed-culture in combination with competition indices, can successfully separate allelopathic effects from competition.     

Functional correlates of allelopathic potential in a successional plant community

Allelopathy, plant–plant interactions mediated through chemical production, is an active area of ecological research. Despite this widespread interest, we still lack community scale information on the prevalence of this interaction and the types of species that may be expected to be allelopathic. To address this research need, the allelopathic potential of 65 plant species from all stages of succession in the Piedmont region of New Jersey, USA, was determined with laboratory bioassays. The strength of each species’ allelopathic activity was then related to life form, origin, and fundamental plant traits. The vast majority of species tested exhibited significant allelopathic effects in the bioassays, with many of these having fairly strong effects. Overall, the allelopathic potential of species decreased with life span, roughly following the successional transitions from short-lived to long-lived herbs and to woody species. Herbaceous species on average were more allelopathic than woody species, but there was no difference between native and non-native species once life form was accounted for. In a principal components analysis, allelopathy was associated with other plant traits, but these relationships differed between woody and herbaceous species. Allelopathic potential was positively associated with plant height in herbaceous species, but negatively associated with height, leaf mass, and seed mass in woody species. These results indicate that allelopathy may be a quite common ecological strategy in plants and is equally common in both native and non-native species. The linkage of allelopathy with other plant functional traits suggests that allelopathy can and should be integrated into the broader suite of plant strategies that are studied.     

Biological control of phytoplankton by the subtropical submerged macrophytes Egeria densa and Potamogeton illinoensis: a mesocosm study

1. In temperate regions, submerged macrophytes can hamper phytoplankton blooms. Such an effect could arise directly, for instance via allelopathy, or indirectly, via competition for nutrients or the positive interaction between submerged macrophytes and zooplankton grazing. However, there is some evidence that the positive interaction between submerged macrophytes and zooplankton grazing is less marked in warmer regions, where the interaction is less well studied, and that negative effects of higher water plants on phytoplankton biomass are weaker. 2. We carried out two consecutive mesocosm experiments in Uruguay (subtropical South America) to study the effects of two common submerged macrophytes from this region (Egeria densa and Potamogeton illinoensis) on phytoplankton biomass, in the absence of zooplankton grazing. We compared phytoplankton development between different macrophyte treatments (no macrophytes, artificial macrophytes, real Egeria and real Potamogeton). We used artificial macrophytes to differentiate between physical effects (i.e. shading, sedimentation and competition with periphyton) and biological effects (i.e. nutrient competition and allelopathy). 3. In Experiment 1, we found no evidence for physical effects of macrophytes on phytoplankton biomass, but both macrophyte species seemed to exert strong biological effects on phytoplankton biomass. Only Egeria affected phytoplankton community structure, particularly tempering the dominance of Scenedesmus. Nutrient addition assays revealed that only Egeria suppressed phytoplankton through nutrient competition. 4. We performed a second mesocosm experiment with the same design, but applying saturating nutrient conditions as a way of excluding the effects of competition for nutrients. This experiment showed that both macrophytes were still able to suppress phytoplankton through biological mechanisms, providing evidence for allelopathic effects. Our results indicate that both common macrophytes are able to keep phytoplankton biomass low, even in the absence of zooplankton grazing.     

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.     

Separation of allelopathy and resource competition by the boreal dwarf shrub Empetrum hermaphroditum Hagerup

An experimental technique was used to separate and evaluate the magnitude of allelopathic interference relative to resource competition by the boreal dwarf shrub Empetrum hermaphroditum Hagerup (Empetraceae). To test for resource competition and allelopathy, respectively, Scots pine (Pinus sylvestris L.) seedlings were grown in both the greenhouse and in the field over a 3 year period, in four different treatments within E. hermaphroditum vegetation: (1) PVC tubes were used to reduce effects of interspecific below-ground competition; (2) activated carbon was spread on the soil to adsorb toxins leached from E. hermaphroditum litter and green leaves, thus reducing effects of allelopathic interference; (3) E. hermaphroditum vegetation was left untreated to evaluate inhibiting effects when both allelopathy and resource competition were present; (4) PVC tubes, placed in E. hermaphroditum vegetation spread with activated carbon were used to determine growth of seedlings when both allelopathy and resource competition were reduced. Scots pine seedlings grown in untreated vegetation (with both root competition and allelopathy present) had the lowest shoot length and dry weight; seedlings with both allelopathy and root competition reduced (activated carbon in tube) were the largest. Reducing either root competition alone (tube treatment) or allelopathy alone (carbon treatment) produced seedlings of intermediate size, but reduced competition had a greater effect than reduced allelopathy (although, in the greenhouse, significantly so only for root biomass). In the greenhouse experiment, biomass production of seedlings grown free of both interactions (carbon in tube) was greater than the simple sum of the growth response to the individual interactions (tube treatment and carbon treatment, respectively). Larger shoot:root ratios were also found when pine seedlings were grown without tubes (i.e. when resource competition was occurring). In the field, the removal of allelopathy (carbon treatments) increased shoot:root ratio when compared to the removal of resource competition. The study showed that two different interference mechanisms of E. hermaphroditum can be separated and quantified, and that below-ground competition and allelopathy by E. hermaphroditum are both important factors retarding growth of Scots pine.     

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.     

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.