Toxigenic effects of diatoms on grazers, phytoplankton and other microbes: a review.

Traditionally, diatoms have been regarded as providing the bulk of the food that sustains the marine food chain and important fisheries. However, this view was challenged almost two decades ago on the basis of laboratory and field studies showing that when copepods, the principal predators of diatoms, feed on certain diatom diets, they produce abnormal eggs that either fail to develop to hatching or hatch into malformed (i.e. teratogenic) nauplii that die soon afterwards. Over the years, many explanations have been advanced to explain the causes for reproductive failure in copepods and other marine and freshwater invertebrates including diatom toxicity, or nutritional deficiency and poor assimilation of essential compounds in the animal gut. Here we review the literature concerning the first possibility, that diatoms produce cytotoxic compounds responsible for growth inhibition and teratogenic activity, potentially sabotaging future generations of grazers by inducing poor recruitment. The cytotoxic compounds responsible for these effects are short chain polyunsaturated aldehydes (PUAs) and other oxygenated fatty acid degradation products such as hydroxides, oxo-acids, and epoxyalcohols (collectively termed oxylipins) that are cleaved from fatty acid precursors by enzymes activated within seconds after crushing of cells. Such toxins are suggested to have multiple simultaneous functions in that they not only deter herbivore feeding but some also act as allelopathic agents against other phytoplankton cells, thereby affecting the growth of competitors, and also signalling population-level cell death and termination of blooms, with possible consequences for food web structure and community composition. Some oxylipins also play a role in driving marine bacterial community diversity, with neutral, positive or negative interactions depending on the species, thereby shaping the structure of bacterial communities during diatom blooms. Several reviews have already been published on diatom-grazer interactions so this paper does not attempt to provide a comprehensive overview, but rather to consider some of the more recent findings in this field. We also consider the role of diatom oxylipins in mediating physiological and ecological processes in the plankton and the multiple simultaneous functions of these secondary metabolites.