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geosmin natural bicyclic terpene earthy odor volatile sesquiterpene irregular precursor farnesyl pyrophosphate methylisoborneol musty smell odor detection beetroots scent filamentous bacteria blue-green algae actinomyces prokaryotes eukaryotes anabaena phormidium planktothrix streptomyces biosynthesis geosmin synthase recirculating aquaculture systems saltwater fish depuration rendering substance matter material compound component molecule molecular structural structure formula skeletal geometry ball stick