Polyphenols as regulators of plant-litter-soil interactions in northern California's pygmy forest: A positive feedback?

The convergent evolution of polyphenol-rich plant communities has occurred on highly acidic and infertile soils throughout the world. The pygmy forest in coastal northern California is an example of an ecosystem on an extremely infertile soil that has exceptionally high concentrations of polyphenols. Many 'negative feedbacks' have been identified whereby plants degrade fertile soils through production of polyphenol-rich litter, sequestering soil nutrients into unavailable form and creating unfavorable conditions for seed germination, root growth, and nutrient uptake. But in the context of plant-litter-soil interactions in ecosystems adapted to soils that are inherently acidic and infertile (such as the pygmy forest), there are also many 'positive feedbacks' that result from polyphenol production. By inhibiting decomposition, polyphenols regulate the formation of a mor-humus litter layer, conserving nutrients and creating a more favorable medium for root growth. Polyphenols shift the dominant pathway of nitrogen cycling from mineral to organic forms to minimize potential N losses from the ecosystem and maximize litter-N recovery by mycorrhizal symbionts. Polyphenol complexation of Al, Mn and Fe reduce potential Al toxicity and P fixation in soil. Polyphenols regulate organic matter dynamics, leading to the accumulation of organic matter with cation exchange capacity to minimize leaching of nutrient cations. Humic substances derived from polyphenolic precursors coat rhizosphere soil surfaces, improving physical and chemical conditions for root growth and nutrient cycling. Although their long-accepted adaptive value for antiherbivore defense is now in doubt, polyphenol alteration of soil conditions and regulation of nutrient cycling illustrate how fitness can be influenced by the 'extended' phenotype in plant-litter-soil interactions.