Regulation of Fatty Acid Production and Release in Benthic Algae: Could Parallel Allelopathy Be Explained with Plant Defence Theories?

Many organisms produce chemical compounds, generally referred as secondary metabolites, to defend against predators and competitors (allelopathic compounds). Several hypotheses have been proposed to explain the interaction between environmental factors and secondary metabolites production. However, microalgae commonly use simple metabolites having a role in primary metabolism as allelopathic compounds. The aim of this study was to determine whether classical theories of plant chemical defences could be applied to microalgae producing allelochemicals derived from the primary metabolism. Our study was designed to investigate how growth phase, algal population density, nutrient limitation and carbon assimilation affect the production and release of allelopathic free fatty acids (FFAs) among other FFAs. The model species used was Uronema confervicolum, a benthic filamentous green alga that produces two allelopathic FFAs (linoleic and α-linolenic acids) inhibiting diatom growth. FFAs have been quantified in algal biomass and in culture medium. Our results were analysed according to two classical plant defence theories: the growth-differentiation balance hypothesis (GDBH) and the optimal defence theory (ODT), based on the metabolic capacities for defence production and on the need for defence, respectively. While a higher production of allelopathic compounds under increased light conditions supports the use of GDBH with this microalga, the observation of a negative feedback mechanism mostly supports ODT. Therefore, both theories were insufficient to explain all the observed effects of environmental factors on the production of these allelochemicals. This highlights the needs of new theories and models to better describe chemical interactions of microalgae.

Allelopathic interactions involving benthic phototrophic microorganisms.

As a way to prevent resource depletion by other species, many phototrophic aquatic microorganisms produce inhibitory compounds. This process, known as allelopathy, has been widely studied in planktonic environments, where it is recognized as being a driving force of planktonic communities. However, in benthic environments, biofilms provide very particular micro-environments. The present review focuses on allelopathic interactions involving benthic phototrophic prokaryotes and micro-eukaryotes ('microalgae'), which generally form biofilms, and includes any interaction involving benthic microalgae either as the emitter or as the target in both marine and freshwater habitats. To support our hypothesis on the importance of allelopathy in biofilms due to the particularities of biofilms, we show that (i) reported allelopathic species and compounds are diverse and numerous in the three major groups of benthic phototrophic microorganisms, (ii) allelopathic benthic species could affect community composition, (iii) allelopathy in biofilms is currently underestimated because of the lack of suitable methods. As benthic primary producers represent an important source of organic carbon in some streams and littoral areas, these interactions could impact the whole ecosystem in these areas, probably more than in areas dominated by planktonic communities.

Impairment of benthic diatom adhesion and photosynthetic activity by allelopathic compounds from a green alga: involvement of free fatty acids?

The role of chemical interactions in shaping microbial communities has raised increasing interest over the last decade. Many benthic microorganisms are known to develop chemical strategies to overcome competitors, but the real importance of chemical interactions within freshwater biofilm remains unknown. This study focused on the biological and chemical mechanisms of an interaction involving two benthic microorganisms, an allelopathic filamentous green alga, Uronema confervicolum, and a common diatom, Fistulifera saprophila. Our results showed that functions critical for benthic phototrophic microorganisms were inhibited by U. confervicolum extracts. Growth, cell motility, adhesion, and photosynthetic activity were impaired at extract concentrations ranging between 5 and 20 µg ml(-1). The adhesion inhibition was mediated by intracellular nitric oxide (NO) induction. A bioassay-guided fractionation of the extract with HPLC helped to identify two C18 fatty acids present in the growth-inhibiting fractions: linoleic (LA) and α-linolenic (LNA) acids. These compounds represented 77% of the total free fatty acids of U. confervicolum and were present in the culture medium (1.45 µg l(-1) in total). Both could inhibit the diatom growth at concentrations higher than 0.25 µg ml(-1), but had no effect on cell adhesion. The discrepancy between the effective concentrations of fatty acids and the concentration found in culture medium may be explained by the presence of high-concentration microenvironments. The compounds involved in adhesion inhibition remain to be identified. Though further experiments with complex biofilms are needed, our results suggest that U. confervicolum may participate to the control of biofilm composition by inhibiting diatom adhesion.