Differential sensitivity of green algae to allelopathic substances from Chara

Three short-term laboratory experiments were conducted to investigate allelopathic effects of a mixture of Chara globularis var. globularis Thuillier and Chara contraria var. contraria A. Braun ex Kützing on three different green algae. Single phytoplankton species were exposed to filtered water originating from charophyte cultures. Phytoplankton growth was monitored by determination of chlorophyll concentrations in batch cultures. The change in chlorophyll concentration during the experiments was analysed with a logistic growth model, resulting in an estimate of the exponential growth rate and the duration of the lag phase of the single green algae. The results indicate allelopathic effects of Chara on the growth of the green algae Selenastrum capricornutum Printz and Chlorella minutissima Fott et Nováková, whereas Scenedesmus obliquus (Turpin) Kützing did not seem to be affected. The exponential growth rate of S. capricornutum decreased 7% in the presence of water from a charophyte culture, while the growth rate of C. minutissima decreased with 3%. The allelopathic effect of Chara did not increase when the green alga C. minutissima was P-limited. The effect of Chara was different when young sprouts were used. With young sprouts the duration of the lag phase of C. minutissima was extended (25%), whilst for old plants the growth rate of this green alga decreased. Although the inhibiting effect of charophytes on specific phytoplankton species is rather small, the differential sensitivity of the species to Chara might influence the composition and biomass of phytoplankton communities in the field.     

Interaction between the macrophyte <Emphasis Type="Italic">Stratiotes aloides</Emphasis> and filamentous algae: does it indicate allelopathy?

The aquatic macrophyte Stratiotes aloides Linnaeus, which has recently received attention in studies on allelopathy, has been shown to suppress phytoplankton growth. In the Netherlands, S. aloides often co-occurs with floating filamentous algae. However, filamentous algae are generally absent in close proximity to S. aloides, resulting in gaps in filamentous algae mats. We analyzed whether those gaps may be caused by allelopathic substances excreted by S. aloides or by nutrient depletion. We studied in a field survey the colonization of natural S. aloides by filamentous algae and determined in situ nutrient concentrations in natural S. aloides stands. To analyze the relative importance of allelopathy and nutrient competition in the interaction between S. aloides and filamentous algae, we carried out field experiments. Introduction of artificial (non-allelopathic) plants in natural S. aloides stands enabled us to compare the colonization by filamentous algae of both Stratiotes sp. and artificial plants. The filamentous algae were absent in close vicinity to S. aloides. Significantly lower concentrations of ortho-phosphate and potassium were observed close to S. aloides as compared with the filamentous algae. In the field experiments the artificial plants were rapidly colonized by filamentous algae, mainly Cladophera Kützing and Spirogyra Link, while all natural plants remained free of such algae. Additionally, most nutrient concentrations did not significantly differ in the proximity of artificial or natural stands of S. aloides. The concentrations of the major growth-limiting nutrients, phosphate and nitrate, were significantly higher and nonlimiting in natural Stratiotes stands. Our main conclusion is that, although allelopathic interactions between S. aloides and filamentous algae do occur under natural conditions, nutrient competition between the two can also be an important factor.     

Allelopathic activity of Stratiotes aloides on phytoplankton—towards identification of allelopathic substances

The allelopathic activity of the aquatic macrophyte, Stratiotes aloides, was determined with laboratory experiments. Active compounds exuded in the medium or present in plant tissue were extracted using standard procedures and solid phase extraction (SPE). The activity towards various cyanobacteria and chlorophytes was tested in two different bioassay systems using agar plates and liquid cultures of phytoplankton. Extracts and exudates of S. aloides affected phytoplankton growth. SPE-enriched exudates and enriched water from a natural Stratiotes stand caused inhibition of target species, however, also some controls were active. Phytoplankton species exhibited differential sensitivity to extracts of S. aloides. We observed inhibitory and stimulatory effects on phytoplankton. In general, more cyanobacteria than other phytoplankton species were inhibited, and the inhibition of cyanobacteria was stronger. In most cases, nutrient (P or K) limitation of Synechococcus elongatus and Scenedesmus obliquus decreased the sensitivity of these species towards allelochemicals from Stratiotes aloides, except for P-limited cultures of Scenedesmus. The allelopathically active compound(s) from Stratiotes are moderately lipophilic and most likely no phenolic compounds. Our results indicate that allelopathy (besides nutrient interference and shading) might also account for the low phytoplankton and filamentous algae densities in the vicinity of Stratiotes plants, at least during certain phases of the life-cycle of Stratiotes.     

Winter ecology of shallow lakes: strongest effect of fish on water clarity at high nutrient levels

While the structuring role of fish in lakes is well studied for the summer season in North temperate lakes, little is known about their role in winter when fish activity and light irradiance potentially are lower. This is unfortunate as the progressing climate change may have strong effects on lake winter temperature and possibly on trophic dynamics too. We conducted an enclosure experiment with and without the presence of fish throughout winter in two shallow lakes with contrasting phosphorus concentrations. In hypertrophic Lake Søbygård, absence of fish led to higher biomass of zooplankton, higher grazing potential (zooplankton:phytoplankton ratio) and, accordingly, lower biomass of phytoplankton and chlorophyll a (Chl a), while the concentrations of total nitrogen (TN), total phosphorus (TP), oxygen and pH decreased. The average size of egg-bearing Daphnia and Bosmina and the minimum size of egg-bearing specimens of the two genera rose. In the less eutrophic Lake Stigsholm, zooplankton and their grazing potential were also markedly affected by fish. However, the decrease in Chl a was slight, and phytoplankton biovolume, pH and the oxygen concentration were not affected. TN was higher when fish were absent. Our results indicate that: (i) there is a notable effect of fish on zooplankton community structure and size during winter in both eutrophic and hypertrophic North temperate lakes, (ii) Chl a can be high in winter in such lakes, despite low light irradiance, if fish are abundant, and (iii) the cascading effects on phytoplankton and nutrients in winter may be more pronounced in hypertrophic lakes. Climate warming supposedly leading to reduced winter mortality and dominance of small fish may enhance the risk of turbid state conditions in nutrient-enriched shallow lakes, not only during the summer season, but also during winter.     

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.