Biota can be described in terms of elemental composition, expressed as an atomic ratio of carbon:nitrogen:phosphorus (refs 1-3). The elemental stoichiometry of microoorganisms is fundamental for understanding the production dynamics and biogeochemical cycles of ecosystems because microbial biomass is the trophic base of detrital food webs. Here we show that heterotrophic microbial communities of diverse composition from terrestrial soils and freshwater sediments share a common functional stoichiometry in relation to organic nutrient acquisition. The activities of four enzymes that catalyse the hydrolysis of assimilable products from the principal environmental sources of C, N and P show similar scaling relationships over several orders of magnitude, with a mean ratio for C:N:P activities near 1:1:1 in all habitats. We suggest that these ecoenzymatic ratios reflect the equilibria between the elemental composition of microbial biomass and detrital organic matter and the efficiencies of microbial nutrient assimilation and growth. Because ecoenzymatic activities intersect the stoichiometric and metabolic theories of ecology, they provide a functional measure of the threshold at which control of community metabolism shifts from nutrient to energy flow.

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Periphyton assemblage data collected from 233 stream site-visits (49 in 1993, 56 in 1994, and 128 in 1995) throughout the Mid-Appalachian region were used to develop a periphyton index of biotic integrity (PIBI) based on 1) algal genera richness; 2) the relative abundances of diatoms, Cyanobacteria, dominant diatom genus, acidophilic diatoms, eutraphentic diatoms, and motile diatoms; 3) chlorophyll and biomass (ash-free dry mass) standing crops; and 4) alkaline phosphatase activity. Thirty-seven diatom genera and 38 non-diatom genera were collected. The relative richness and relative abundance (RA) of these genera were used to calculate the RA metrics of the PIBI. PIBI scores ranged from 48.0 to 85.1 among the 233 site-visits with an overall regional mean (Ϯ1 SE) of 66.1 Ϯ 0.5. The 10 metrics and the PIBI were correlated with 27 chemical, 12 physical habitat, and 3 landscape variables. Overall, PIBI was inversely correlated with stream depth, stream water color, and Fe. Component metrics were significantly correlated with several chemical (Al, acid neutralizing capacity, Cl, Fe, Mn, N, Na, P, pH, Si, SO 4 , total suspended solids), physical habitat (channel embeddedness, riparian disturbances, stream depth, stream width, substrate composition), and landscape (% of the watershed in forest, agriculture, and urban land uses) variables. Canonical correlation analysis revealed significant correlations between the 10 PIBI metrics and 4 significant environmental gradients related to general human disturbances (stream acidity, stream substrate composition, and stream and riparian habitat). Analysis of variance revealed significant differences in PIBI scores for lowland vs highland streams, and among stream orders. Annual differences were explained by differences in the proportions of sampling sites in lowland streams in each year. The univariate distribution of PIBI scores was used to set threshold PIBI values for the assessment of ecological condition in Mid-Appalachian streams.

Human alteration of the nitrogen (N) cycle has produced benefits for health and well-being, but excess N has altered many ecosystems and degraded air and water quality. US regulations mandate protection of the environment in terms that directly connect to ecosystem services. Here, we review the science quantifying effects of N on key ecosystem services, and compare the costs of N-related impacts or mitigation using the metric of cost per unit of N. Damage costs to the provision of clean air, reflected by impaired human respiratory health, are well characterized and fairly high (e.g. costs of ozone and particulate damages of $28 per kg NO x -N). Damage to services associated with productivity, biodiversity, recreation and clean water are less certain and although generally lower, these costs are quite variable (< $2.2-56 per kg N). In the current Chesapeake Bay restoration effort, for example, the collection of available damage costs clearly exceeds the projected abatement costs to reduce N loads to the Bay ($8-15 per kg N). Explicit consideration and accounting of effects on multiple ecosystem services provides decision-makers an integrated view of N sources, damages and abatement costs to address the significant challenges associated with reducing N pollution.

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