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Karenia mikimotoi

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estimated to be 0.1 to 0.5 that of a <strong>Karenia</strong> brevis cell (Bridgers et al., 2004). It was<br />

suggested by these workers that blooms of raphidophytes would have to be of<br />

correspondingly higher densities to have the same effect as that of a K. brevis bloom.<br />

Also, as discussed by Smayda (2006), there is a higher probability that blooms will<br />

occur in chemically modified habitats and at fish-farming sites.<br />

It is interesting to note that two of the fatty acids from Fibrocapsa reported by Fu et al.<br />

(2004) as having strong haemolytic activity are widely promoted as human dietary<br />

supplements. Eicosapentaenoic acid (EPA) is a component of ‘omega-3’-rich fish oils,<br />

and non-toxic microalgae such as Porphyridium cruentum have been investigated as a<br />

possible source of arachidonic acid for use in infant feeding formulations. This<br />

highlights the fact that the toxic effect of such compounds probably operates at a<br />

microsopic level where there may be microzones of very high concentrations around<br />

individual cells (see sect. 3.4.3).<br />

3.9 Heterosigma akashiwo<br />

This alga is cosmopolitan and is widely distributed in the Atlantic and Pacific oceans<br />

and is found in European waters from Norway in the north to Spain and Portugal in the<br />

south (Smayda, 2006). Its toxic properties have been reviewed by Landsberg, (2002)<br />

and Smayda (2006). To quote the latter: “Heterosigma akashiwo is probably the most<br />

versatile harmful algal species known: it is antagonistic to organisms ranging in size<br />

from bacteria to invertebrates to fish. Its multiple modes of antagonism range from<br />

nutritional inadequacy, to suppression of feeding and growth, and to toxicity causing<br />

mortality.” These antagonistic properties towards various organisms have been<br />

tabulated in detail by Landsberg (2002).<br />

The exact nature of the toxins and mechanisms of toxicity in H. akashiwo and other<br />

raphidophytes remains unclear. Some mechanisms of ichthyotoxicity, which may also<br />

be involved with allelopathy, that have been postulated are production of reactive<br />

oxygen species and the production of toxins that have yet to be fully characterized<br />

(Twiner et al., 2004; Smayda, 2006; Kempton et al., 2008).<br />

Khan et al. (1997) reported that they had isolated from H. akashiwo four toxic<br />

components which were, on the basis of the analytical methods used, tentatively<br />

identified as brevetoxins, neurotoxins that have been well characterized and are also<br />

produced by the dinoflagellate <strong>Karenia</strong> brevis (see sect. 5.).<br />

Uncharacterised extracellular compounds excreted by H. akashiwo were found to have<br />

an irreversible effect on respiratory activity in human and mammalian cells in vitro<br />

(Twiner et al., 2004). These compounds were also found to affect cytosolic calcium<br />

concentrations in cultured sf19 insect cells leading to apoptosis (cell death). The effect<br />

was related to the concentration of extracellular Ca 2+ . It was concluded that the<br />

extracellular compounds were inhibiting the plasma membrane Ca 2+ -ATPase<br />

transporter (Twiner et al., 2005). It was suggested that the compounds may play a role<br />

in the ichthyotoxicity and allelopathy of the alga.<br />

In laboratory studies Keppler et al. (2005) demonstrated sublethal effects (significantly<br />

increased oyster hepatopancreas lysosomal destabilization rates) of cells of H.<br />

akashiwo on the southeastern oyster, Crassostrea virginica.<br />

The most notable impact of H. akashiwo has been on aquaculture where it has caused<br />

massive kills of farmed fish at various locations round the world and has also affected<br />

shellfish (Landsberg, 2002; Smayda, 2006; Kempton et al., 2008).<br />

24 A Literature review of the potential health effects of marine microalgae and macroalgae

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