Macroalgal Secondary Metabolites

The tuft-forming green alga Cladophora glomerata synthesizes a wide variety of toxic fatty acids, including capric and palmitoleic acids, which have been demonstrated to be insecticidal and allelopathic (Dodds and Gudder 1992). 

To date, no clear evidence of inducible defenses among freshwater macroalgae has been reported, in contrast to their marine algal counterparts. For example, certain species of marine brown algae increase phlorotannin production in response to damage by mesograzers (Amsler and Fairhead 2006 see Chaps. 3 and 7). Whether Chara and Cladophora, two species of freshwater chlorophytes with putative allelopathic activity, increase allelochemical concentration in response to competitors remains to be seen (see Sect. 5.7.3). 

Significant concentrations of cyanotoxins have been found to accumulate in the tissues of macroinvertebrates such as mollusks and crustaceans, presenting an indirect route of exposure for invertebrates, fish, and aquatic mammals at higher trophic levels (Negri and Jones 1995). In natural systems, mortality among benthic invertebrate herbivores is probably low because most bloom-forming bacteria are planktonic and only periodically come into contact with the benthos. Nevertheless, Kotak et al. (1996) determined that enhanced mortality of snails at the end of a bloom cycle in Canadian lakes was due to consumption of Microcystis cells that had formed a scum on the surface of macrophytes. Oberemm et al. (1999) found that aqueous microcystins, saxitoxins, and anatoxin-a all resulted in developmental delays in fish and salamander embryos. Interestingly, more severe malformations and enhanced mortality were observed when larvae were exposed to crude cyanobacterial extracts than to pure toxins applied at natural concentrations (Oberemm et al. 1999). 

There are numerous reports on herbivorous waterfowl and mammals that have died following consumption of cyanobacterial mats (e.g., Krienitz et al. 2003 WHO 2003). Most deaths are probably due to incidental ingestion of contaminated water during drinking or while feeding on aquatic macrophytes. 

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Evolution of chemical defenses according to the optimal defense theory presumes, in addition to costly defenses, that there is genetic variation for the defensive metabolites, that herbivory is the major selective agent for such metabolites, and that the chemical trait in question is efficient in reducing herbivory (Stamp 2003). Research on macroalgal chemical defenses has strongly emphasized the last precondition, which has mainly been studied by testing the deterrence effects of secondary metabolites in bioassays. The defensive role for the trait has been assumed on the basis of deterrence it provides. Veiy little research on the first two... 

Table 7.1 Published studies explicitly testing hypotheses derived from the Induced Defense Model, a corollary model to the Optimal Defense Model, concerning variation in the concentration of secondary metabolites or resistance to herbivores in different macroalgal species... 

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