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Research seeking to explain the ecological role of polyphenolics (phlorotannins) in plants and brown algae has largely focused on 2 alternative concepts, the carbonhutrient (Cm) balance and the inducible defense models. We tested the hierarchy of effects of both models on phlorotannin production in the brown alga Fucus veslculosus (Fucales) by simultaneously manipulating the N environment and simulating herbivory for 2 oceanic (high and low intertidal) and estuarine populations. We measured phlorotannin levels in algae under control, grazed, N-enriched, and grazed + N-enriched treatments with time (0 to 14 d ) throughout the year to determine onset and duration of the response. We found greater support for the inducible defense model; generally, both grazed and grazed + Nenriched fronds had significantly higher phlorotannin concentrations than control thalli. When we found an inducible response, it was rapid (within 3 d) and relatively long term (>2 wk). However, the induced response was minimal for both oceanic populations during March, perhaps due to fixed-C limitation, and was absent for the estuarine and high intertidal populations during June, the period of peak phlorotannins at both sites. Although Nenrichment resulted in depressed concentrations of phlorotannins only for the estuarine population, we did measure a significant negative correlation between tissue N and phenolics for the oceanic population, a s predicted by the C/N balance model. Thus, while the induc~ble defense response takes preeminence over resource availability effects (C/N balance hypothesis), this study revealed that phlorotannin production is likely controlled by a complex interaction of environmental, developmental and defense-related factors, emphasizing the applicability of both models in marine systems.

Interactive effects of inducible defense and resource availability on phlorotannins in the North Atlantic brown alga Fucus vesiculosus