Shaver, G.R., Johnson, L.C., Cades, D.H., Murray, G., Laundre, J.A., Rastetter, E.B., Nadelhoffer, K.J., and Giblin, A.E. "Biomass and CO2 flux in wet sedge tundras: Responses to nutrients, temperature, and light." Ecological Monographs 1998 68(1): 75-97.

The aim of this research was to analyze the effects of increased N or P availability, increased air temperature, and decreased light intensity on wet sedge tundra in northern Alaska. Nutrient availability was increased for 69 growing seasons, using N and P fertilizers in factorial experiments at three separate field sites. Air temperature was increased for six growing seasons, using plastic greenhouses at two sites, both with and without N + P fertilizer. Light intensity (photosynthetically active photon flux) was reduced by 50% for six growing seasons at the same two sites, using optically neutral shade cloth. Responses of wet sedge tundra to these treatments were documented as changes in vegetation biomass, N mass, and P mass, changes in whole-system CO2 fluxes, and changes in species composition and leaf-level photosynthesis.

Biomass, N mass, and P mass accumulation were all strongly P limited, and biomass and N mass accumulation also responded significantly to N addition with a small N P interaction. Greenhouse warming alone had no significant effect on biomass, N mass, or P mass, although there was a consistent trend toward increased mass in the greenhouse treatments. There was a significant negative interaction between the greenhouse treatment and the N + P fertilizer treatment, i.e., the effect of the two treatments combined was to reduce biomass and N mass significantly below that of the fertilizer treatment only. Six years of shading had no significant effect on biomass, N mass, or P mass.

Ecosystem CO2 fluxes included net ecosystem production (NEP; net CO2 flux), ecosystem respiration (RE, including both plant and soil respiration), and gross ecosystem production (GEP; gross ecosystem photosynthesis). All three fluxes responded to the fertilizer treatments in a pattern similar to the responses of biomass, N mass, and P mass, i.e., with a strong P response and a small, but significant, N response and N P interaction. The greenhouse treatment also increased all three fluxes, but the greenhouse plus N + P treatment caused a significant decrease in NEP because RE increased more than GEP in this treatment. The shade treatment increased both GEP and RE, but had no effect on NEP. Most of the changes in CO2 fluxes per unit area of ground were due to changes in plant biomass, although there were additional, smaller treatment effects on CO2 fluxes per unit biomass, per unit N mass, and per unit P mass.

The vegetation was composed mainly of rhizomatous sedges and rushes, but changes in species composition may have contributed to the changes in vegetation nutrient content and ecosystem-level CO2 fluxes. Carex cordorrhiza, the species with the highest nutrient concentrations in its tissues in control plots, was also the species with the greatest increase in abundance in the fertilized plots. In comparison with Eriophorum angustifolium, another species that was abundant in control plots, C. cordorrhiza had higher photosynthetic rates per unit leaf mass. Leaf photosynthesis and respiration of C. cordorrhiza also increased with fertilizer treatment, whereas they decreased or remained constant in E. angustifolium.

The responses of these wet sedge tundras were similar to those of a nearby moist tussock tundra site that received an identical series of experiments. The main difference was the dominant P limitation in wet sedge tundra vs. N limitation in moist tussock tundra. Both tundras were relatively unresponsive to the increased air temperatures in the greenhouses but showed a strong negative interaction between the greenhouse and fertilizer treatments. New data from this study suggest that the negative interaction may be driven by a large increase in respiration in warmed fertilized plots, perhaps in relation to large increases in P concentration.