Tannins in nutrient dynamics of forest ecosystems - a review

Tannins make up a significant portion of forest carbon pools and foliage and bark may contain up to 40% tannin. Like many other plant secondary compounds, tannins were believed to function primarily as herbivore deterrents. However, recent evidence casts doubts on their universal effectiveness against herbivory. Alternatively, tannins may play an important role in plant - plant and plant - litter - soil interactions. The convergent evolution of tannin-rich plant communities on highly acidic and infertile soils throughout the world, and the intraspecific variation in tannin concentrations along edaphic gradients suggests that tannins can affect nutrient cycles. This paper reviews nutrient dynamics in forest ecosystems in relation to tannins. Tannins comprise a complex class of organic compounds whose concentration and chemistry differ greatly both among and within plant species. Because the function and reactivity of tannins are strongly controlled by their chemical structure, the effects of tannins on forest ecosystem processes are expected to vary widely. Tannins can affect nutrient cycling by hindering decomposition rates, complexing proteins, inducing toxicity to microbial populations and inhibiting enzyme activities. As a result, tannins may reduce nutrient losses in infertile ecosystems and may alter N cycling to enhance the level of organic versus mineral N forms. The ecological consequences of elevated tannin levels may include allelopathic responses, changes in soil quality and reduced ecosystem productivity. These effects may alter or control successional pathways. While a great deal of research has addressed tannins and their role in nutrient dynamics, there are many facets of tannin biogeochemistry that are not known. This lack of information hinders a complete synthesis of tannin effects on forest ecosystem processes and nutrient cycling. Areas of study that would help clarify the role of tannins in forest ecosystems include improved characterization and quantification techniques, enhanced understanding of structure-activity relationships, investigation of the fate of tannins in soil, further determination of the influence of environmental factors on plant tannin production and decomposition, and additional information on the effects of tannins on soil organisms.