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Evidence for the role and fate of water-insoluble condensed tannins in the short-term reduction of carbon loss during litter decay


Warmer temperatures associated with climate change have the potential to accelerate litter decay and subsequently release large amounts of carbon stored in soils. Condensed tannins are widespread secondary metabolites, which accumulate to high concentrations in many woody plants and play key roles in forest soil nutrient cycles. Future elevated atmospheric CO2 concentrations are predicted to reduce nitrogen content and increase tannin concentrations in plant tissues, thus reducing litter quality for microbial communities and slowing decomposition rates. How the distinct condensed tannin fractions (water-soluble, acetone:MeOH-soluble and solvent-insoluble) impact soil processes, has not been investigated. We tested the impact of condensed tannin and nitrogen concentrations on decay rates of poplar and Douglas-fir litter at sites spanning temperature and moisture gradients in coastal rainshadow forests in British Columbia, Canada. The three condensed tannin fractions were quantified using recent improvements on the butanol-HCl assay. Decay was assessed based on carbon remaining, while changes in litter chemistry were primarily observed using two methods for proximate chemical analyses. After 0.6 and 1 year of decay, more carbon remained in poplar litter with high, compared to low, condensed tannin concentrations. By contrast, more carbon remained in Douglas fir litter than poplar litter during this period, despite lower condensed tannin concentrations. Rapid early decay was especially attributed to loss of soluble compounds, including water-soluble condensed tannins. Water-insoluble condensed tannin fractions, which were transformed to acid-unhydrolyzable residues over time, were associated with reduced carbon loss in high condensed tannin litter.

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Acid-determined cellulose


Acid-determined “lignin”


Acid-hydrolyzable fraction


Acid-unhydrolyzable residue




Condensed tannins


Low-condensed tannin treatment


Low-nitrogen treatment


High-condensed tannin treatment


High-nitrogen treatment




Non-polarizable extractables


Water-soluble extractables


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We thank Nicholas von Wittgenstein for preparing litter bags, David Dunn and Rebecca Dixon (Chemical Services Laboratory, Pacific Forestry Centre, NRCan) for proximate chemical analyses, and Dr. Caroline M. Preston (Pacific Forestry Centre, NRCan) for feedback and insightful conversations. Special thanks to Professor Thomas G. Whitham and the cottonwood research group at Northern Arizona University for the poplar litter samples and associated genetic and chemical information. This work was supported by the Natural Sciences and Engineering Research Council (NSERC) of Canada in the form of Discovery Grants to CPC and the NSERC CREATE Program in Forests and Climate Change.

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Correspondence to C. Peter Constabel.

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Responsible Editor: Jack Brookshire.

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Shay, PE., Peter Constabel, C. & Trofymow, J.A. Evidence for the role and fate of water-insoluble condensed tannins in the short-term reduction of carbon loss during litter decay. Biogeochemistry 137, 127–141 (2018).

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  • Poplar
  • Douglas-fir
  • Proanthocyanidin
  • Proximate chemistry
  • Butanol-HCl
  • Carbon sequestration