Randy A. Dahlgren

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.

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.

The Nature, Properties and Management of Volcanic Soils

Abstract Soils formed in volcanic ejecta have many distinctive physical, chemical, and mineralogical properties that are rarely found in soils derived from other parent materials. These distinctive properties are largely attributable to the formation of noncrystalline materials (e.g., allophane, imogolite, ferrihydrite) containing variable charge surfaces, and the accumulation of organic matter. Formation of noncrystalline materials is directly related to the properties of volcanic ejecta as a parent material, namely the rapid weathering of glassy particles. The composition of the colloidal fraction forms a continuum between pure Al–humus complexes and pure allophane/imogolite, depending on the pH and organic matter characteristics of the weathering environment. For soil management purposes, volcanic soils are often divided into two groups based on the colloidal composition of the surface horizons: allophanic soils dominated by allophane and imogolite, and nonallophanic soils dominated by Al–humus complexes and 2:1 layer silicates. Volcanic soils exhibit a wide range of agricultural productivity, depending on the degree or intensity of pedogenic development and the colloidal composition of the rooting zone. The different charge characteristics of allophanic and nonallophanic soils is the most important factor regulating chemical fertility attributes. Phosphorus fixation, strong acidity, and aluminum toxicity are the primary chemical limitations to agricultural productivity. Volcanic soils generally have high physical fertility (tilth) and mature soils are relatively resilient to erosion and compaction. To maximize the productivity of volcanic soils, proper management based on an understanding of the unique physical, chemical, and mineralogical properties of these soils must be practiced.

Contribution of bedrock nitrogen to high nitrate concentrations in stream water

Concentrations of nitrate in stream water throughout the world are reported to be elevated relative to natural background levels. This enrichment is commonly attributed to anthropogenic activities such as atmospheric emissions, livestock feeding, agricultural runoff,, timber harvesting practices and domestic/industrial effluent discharge,. Here we show that bedrock containing appreciable concentrations of fixed nitrogen contribute a surprisingly large amount of nitrate to surface waters in certain California watersheds,o an extent that even small areas of these rocks have a profound influence on water quality. As 75% of the rocks now exposed at the Earths surface are sedimentary in origin, and as these rocks contain about 20% of the global nitrogen inventory, ‘geological’ nitrogen may be a large and hitherto unappreciated source of nitrate to surface waters. Such a natural nitrate source may be especially significant given that nitrate contamination at very low levels can contribute to surface water eutrophication, may cause infant methaemoglobinaemia (‘blue baby’ syndrome) and has been implicated in certain cancers. In addition, geological nitrogen may be a source of the ‘missing’ nitrogen noted in several biogeochemical studies of ecosystem nitrogen budgets.

The effects of whole-tree clear-cutting on soil processes at the Hubbard Brook Experimental Forest, New Hampshire, USA

The effects of whole-tree clear-cutting on soil processes and streamwater chemistry were examined in a northern hardwood forest at the Hubbard Brook Experimental Forest, New Hampshire. Soil processes were examined by monitoring soil solution chemistry collected using zero-tension lysimeters from the Oa, Bh and Bs horizons at three sites along an elevational/vegetation gradient. Whole-tree clear-cutting created a severe ecosystem disturbance leading to leaching losses of nutrients from the soil profile, increased acidification, and elevated concentrations of Al-ions in soil solutions and streamwater. The response was driven by the process of nitrification that led to production of nitric acid in both the forest floor and mineral soil horizons. This acidity was largely neutralized by release and leaching of basic cations and inorganic monomeric Al-ions leaching with the NO3-ions. The major source of nutrient loss was from the forest floor. The chemical response to the clear-cut was most intense during the second year following the treatment and declined to near reference concentrations in 4–5 years. High elevation sites showed the greatest response to disturbance and the slowest recovery of soil solution concentrations to pre-cut concentrations. Shallow soils and a slower recovery of vegetation at the upper elevation sites were the primary factors contributing to the enhanced disturbance and delayed recovery (and enhanced response to disturbance in the upper elevation sites).

Soil development along an elevational transect in the western Sierra Nevada, California

Soil development along an elevational transect on the western slopes of the the central Sierra Nevada was investigated to assess the effects of climate on soil properties and processes. The transect of seven soils formed in granitic residuum spans elevations from 198 to 2865 m with mean annual temperature and precipitation differences of 13°C (3.9–16.7) and 94 cm (33–127), respectively. Soil pH decreased by about two units and base saturation decreased from 90 to 10% with increasing elevation. Concentrations of organic C in the solum increased with elevation, with the largest single increase occurring between the oak woodland (5–6 kg C/m2) and mixed-conifer sites (10–15 kg C/m2). Clay mineralogy showed a general trend of desilication and hydroxy-Al interlayering of 2:1 layer silicates with increasing elevation. The degree of chemical weathering, based on clay and secondary Fe oxide concentration in the solum, showed a maximum (clay = 536 kg/m2 and Fe oxides = 24 kg/m2) at mid-elevations having intermediate levels of precipitation and temperature. While some soil properties show a continuous progression (e.g., organic carbon, base saturation, clay mineralogy) with elevation, other properties (e.g., pH, soil color, clay and secondary Fe oxide concentrations) show a pronounced change (threshold-type step over a short distance at about 1600 m. The explanation for the abrupt nature of this shift is not known; however, it coincides with the approximate elevation of the present-day average effective winter snow-line.

Contribution of amino compounds to dissolved organic nitrogen in forest soils

Dissolved organic nitrogen (DON) may play an important role in plantnutrition and nitrogen fluxes in forest ecosystems. In spite of the apparentimportance of DON, there is a paucity of information concerning its chemicalcomposition. However, it is exactly this chemical characterization that isrequired to understand the importance of DON in ecosystem processes. Theprimaryobjective of this study was to characterize the distribution of free aminoacidsand hydrolyzable peptides/proteins in the DON fraction of Oa horizon leachatesalong an extreme edaphic gradient in northern California. Insitu soil solutions were extracted by centrifugation from Oahorizonscollected beneath Pinus muricata (Bishop pine) andCupressus pygmaea (pygmy cypress) on slightlyacidic/fertile and highly acidic/infertile sites. DON accounted for 77 to99% of the total dissolved nitrogen in Oa horizon leachates. Nitrogen infree amino acids and alkyl amines ranged from 0.04–0.07 mgN/L on the low fertility site to 0.45–0.49 mg N/L onthe high fertility site, and accounted for 1.5 to 10.6% of the DON fraction.Serine, glutamic acid, leucine, ornithine, alanine, aspartic acid andmethylamine were generally the most abundant free amino compounds. Combinedamino acids released by acid hydrolysis accounted for 48 to 74% of theDON, suggesting that proteins and peptides were the main contributor to DON inOa horizon leachates. Together, nitrogen from free andcombined amino compounds accounted for 59 to 78% of the DON. Most of theDON was found in the hydrophobic fraction, which suggests the presence ofprotein/peptide-polyphenol complexes or amino compounds associated withhumic substances. Because free and combined amino acids can be an importantnitrogen source for some plants, soil DON may play an important role in plantnutrition and ecosystem function.

Linking chemical reactivity and protein precipitation to structural characteristics of foliar tannins

Tannins influence ecosystem function by affecting decomposition rates, nutrient cycling, and herbivory. To determine the role of tannins in ecological processes, it is important to quantify their abundance and understand how structural properties affect reactivity. In this study, purified tannins from the foliage of nine species growing in the pygmy forest of the northern California coast were examined for chemical reactivity, protein precipitation capacity (PPC), and structural characteristics (13C NMR). Reactivity of purified tannins varied among species 1.5-fold for the Folin total phenol assay, and 7-fold and 3-fold, respectively, for the acid butanol and vanillin condensed tannin assays. There was about a 5-fold difference in PPC. Variation in chemical reactivity and PPC can be largely explained by differences in structural characteristics of the tannins determined by 13C NMR. In particular, the condensed versus hydrolyzable tannin content, as well as the hydroxylation pattern of the B-ring and stereochemistry at the C-2–C-3 position appear to influence reactivity. Due to the large differences in chemical reactivity among species, it is necessary to use a well-characterized purified tannin from the species of interest to convert assay values to concentrations. Our results suggest that structural characteristics of tannins play an important role in regulating their reactivity in ecological processes.

A review of vegetated buffers and a meta-analysis of their mitigation efficacy in reducing nonpoint source pollution.

Vegetated buffers are a well-studied and widely used agricultural management practice for reducing nonpoint-source pollution. A wealth of literature provides experimental data on their mitigation efficacy. This paper aggregated many of these results and performed a meta-analysis to quantify the relationships between pollutant removal efficacy and buffer width, buffer slope, soil type, and vegetation type. Theoretical models for removal efficacy (Y) vs. buffer width (w) were derived and tested against data from the surveyed literature using statistical analyses. A model of the form Y = K x (1-e(-bxw)), (0 < K < or = 100) successfully captured the relationship between buffer width and pollutant removal, where K reflects the maximum removal efficacy of the buffer and b reflects its probability to remove any single particle of pollutant in a unit distance. Buffer width alone explains 37, 60, 44, and 35% of the total variance in removal efficacy for sediment, pesticides, N, and P, respectively. Buffer slope was linearly associated with sediment removal efficacy either positively (when slope < or = 10%) or negatively (when slope > 10%). Buffers composed of trees have higher N and P removal efficacy than buffers composed of grasses or mixtures of grasses and trees. Soil drainage type did not show a significant effect on pollutant removal efficacy. Based on our analysis, a 30-m buffer under favorable slope conditions (approximately 10%) removes more than 85% of all the studied pollutants. These models predicting optimal buffer width/slope can be instrumental in the design, implementation, and modeling of vegetated buffers for treating agricultural runoff.

Simultaneous Sorption of Cd, Cu, Ni, Zn, Pb, and Cr on Soils Treated with Sewage Sludge Supernatant

Disposal of sewage sludge creates the potential for heavy metal accumulation in theenvironment. This study assessed nine soils currently used as Dedicated Land Disposal units(DLDs) for treatment and disposal of municipal sewage sludge in the vicinity of Sacramento,California. Adsorption characteristics of these soils for Cd, Cu, Ni, Zn, Pb, and Cr were studiedby simultaneously mixing these elements in the range of 0-50 µmol L-1 with sludgesupernatant and reacting with the soil using a soil:supernatant ratio of 1:30, pH = 4.5 or 6.5, andconstant ionic strength (0.01 M Na-acetate). The concentration of metals in the supernatant wasdetermined after a 24 hr equilibration period. Adsorption isotherms showed that metal sorptionwas linearly related to its concentration in the supernatant solution. The distribution coefficientKd (Kd = concentration on solid phase/concentration in solution phase) was computed as theslope of the sorption isotherm. The distribution coefficients were significantly correlated to soilorganic matter content for Ni, Cu, Cd, and Pb at pH 4.5 and for Ni, Cu, Zn, and Cd at pH 6.5.There was also a correlation between Kd and soil specific surface area but no relationship to othersoil properties such as CEC, clay content, and noncrystalline Fe and Al materials. Therefore, soilorganic carbon and surface area appear to be the most important soil properties influencing metaladsorption through formation of organo-metal complexes. The Kd values for all elements werehigher at pH 6.5 than at 4.5. Selectivity between metals resulted in the following metal affinitiesbased on their Kd values: Pb>Cu>Zn>Ni>Cd≈Cr at pH 4.5 andPb>Cu≈Zn>Cd>Ni>Cr at pH 6.5.