Effects of experimental freezing on soil nitrogen dynamics in soils from a net nitrification gradient in a nitrogen-saturated hardwood forest ecosystem

This study examined effects of soil freezing on N dynamics in soil along an N processing gradient within a mixed hardwood dominated watershed at Fernow Experimental Forest, West Virginia. Sites were designated as LN (low rates of N processing), ML (moderately low), MH (moderately high), and HN (high). Soils underwent three 7-day freezing treatments (0, –20, or –80 °C) in the laboratory. Responses varied between temperature treatments and along the gradient. Initial effects differed among freezing treatments for net N mineralization, but not nitrification, in soils across the gradient, generally maintained at LN < ML ≤ MH < HN for all treatments. Net N mineralization potential was higher following freezing at –20 and –80 °C than control; all were higher than at 0 °C. Net nitrification potential exhibited similar patterns. LN was an exception, with net nitrification low regardless of treatment. Freezing response of N mineralization differed greatly from that of nitrification, suggesting that soil freezing may decouple two processes of the soil N cycle that are otherwise tightly linked at our site. Results also suggest that soil freezing at temperatures commonly experienced at this site can further increase net nitrification in soils already exhibiting high nitrification from N saturation.

The Ecological Significance of the Herbaceous Layer in Temperate Forest Ecosystems

Despite a growing awareness that the herbaceous layer serves a special role in maintaining the structure and function of forests, this stratum remainsan underappreciated aspect of forest ecosystems. In this article I review and synthesize information concerning the herb layer’s structure,composition, and dynamics to emphasize its role as an integral component of forest ecosystems. Because species diversity is highest in the herb layeramong all forest strata, forest biodiversity is largely a function of the herb-layer community. Competitive interactions within the herb layer candetermine the initial success of plants occupying higher strata, including the regeneration of dominant overstory tree species. Furthermore, the herblayer and the overstory can become linked through parallel responses to similar environmental gradients. These relationships between strata varyboth spatially and temporally. Because the herb layer responds sensitively to disturbance across broad spatial and temporal scales, its dynamics canprovide important information regarding the site characteristics of forests, including patterns of past land-use practices. Thus, the herb layer has asignificance that belies its diminutive stature.

Effects of Silvicultural Practices on Soil Carbon and Nitrogen in a Nitrogen Saturated Central Appalachian (USA) Hardwood Forest Ecosystem

Silvicultural treatments represent disturbances to forest ecosystems often resulting in transient increases in net nitrification and leaching of nitrate and base cations from the soil. Response of soil carbon (C) is more complex, decreasing from enhanced soil respiration and increasing from enhanced postharvest inputs of detritus. Because nitrogen (N) saturation can have similar effects on cation mobility, timber harvesting in N-saturated forests may contribute to a decline in both soil C and base cation fertility, decreasing tree growth. Although studies have addressed effects of either forest harvesting or N saturation separately, few data exist on their combined effects. Our study examined the responses of soil C and N to several commercially used silvicultural treatments within the Fernow Experimental Forest, West Virginia, USA, a site with N-saturated soils. Soil analyses included soil organic matter (SOM), C, N, C/N ratios, pH, and net nitrification. We hypothesized the following gradient of disturbance intensity among silvicultural practices (from most to least intense): even-age with intensive harvesting (EA-I), even-age with extensive harvesting, even-age with commercial harvesting, diameter limit, and single-tree harvesting (ST). We anticipated that effects on soil C and N would be greatest for EA-I and least with ST. Tree species exhibited a response to the gradient of disturbance intensity, with early successional species more predominant in high-intensity treatments and late successional species more predominant in low-intensity treatments. Results for soil variables, however, generally did not support our predictions, with few significant differences among treatments and between treatments and their paired controls for any of the measured soil variables. Multiple regression indicated that the best predictors for net nitrification among samples were SOM (positive relationship) and pH (negative relationship). This finding confirms the challenge of sustainable management of N-saturated forests.

Acidification and Nutrient Cycling

Additions of acid anions can alter the cycling of other nutrients and elements within an ecosystem. As strong acid ions move through a forest, they may increase the concentrations of nitrogen (N) and sulfur (S) in the soil solution and stream water. Such treatments also may increase or decrease the availability of other anions, cations and metal ions in the soil. A number of studies in Europe and North America have documented increases in base cation concentrations such as calcium (Ca) and magnesium (Mg) with increased N and S deposition (Foster and Nicolson 1988, Feger 1992, Norton et al. 1994, Adams et al. 1997, Currie et al. 1999, Fernandez et al. 2003). Experiments in Europe also have evaluated the response of forested watersheds to decreased deposition (Tietema et al. 1998, Lamersdorf and Borken 2004). In this chapter, we evaluate the effects of the watershed acidification treatment on the cycling of N, S, Ca, Mg and potassium (K) on Fernow WS3.

Temporal and spatial variation of nitrogen transformations in nitrogen-saturated soils of a central Appalachian hardwood forest

We studied temporal and spatial patterns of soil nitrogen (N) dynamics from 1993 to 1995 in three watersheds of Fernow Experimental Forest, W.V.: WS7 (24-year-old, untreated); WS4 (mature, untreated); and WS3 (24-year-old, treated with (NH4)2SO since 1989 at the rate of 35 kg Nha–1year–1). Net nitrification was 141, 114, and115 kg Nha–1year–1, for WS3, WS4, and WS7, respectively, essentially 100% of net N mineralization for all watersheds. Temporal (seasonal) patterns of nitrification were significantly related to soil moisture and ambient temperaturein untreated watersheds only. Spatial patterns of soil water NO3–of WS4 suggest that microenvironmental variabilitylimits rates of N processing in some areas of this N-saturated watershed, in part by ericaceous species in the herbaceous layer. Spatial patterns of soil water NO3–in treated WS3 suggest that later stages of N saturation may result inhigher concentrations with less spatial variability. Spatial variability in soil N variables was lower in treated WS3 versus untreated watersheds. Nitrogen additions have altered the response of N-processing microbes to environmental factors, becoming less sensitive to seasonal changes in soil moisture and temperature. Biotic processes responsible forregulating N dynamics may be compromised in N-saturated forest ecosystems.

Effects of Atmospheric Nitrogen Deposition on the Herbaceous Layer of a Central Appalachian Hardwood Forest

Additions of nitrogen (N) have been shown to alter species diversity of plant communities, with most experimental studies having been carried out in communities dominated by herbaceous species. We examined seasonal and inter-annual patterns of change in the herbaceous layer of two watersheds of a central Appalachian hardwood forest that differed in experimental treatment. This study was carried out at the Fernow Experimental Forest, West Virginia, using two adjacent watersheds: WS4 (mature, second-growth hardwood stand, untreated reference), and WS3. Seven circular 0.04-ha sample plots were established in eachwatershed to represent its full range of elevation and slope aspect. The herbaceous layer was sampled by identifying and visually estimating cover (%) of all vascular plants. Sampling was carried out in mid-July of 1991 and repeated at approximately the same time in 1992. In 1994, these same plots were sampled each month fromMay to October. Seasonal patterns of herb layer dynamics were assessed for the complete 1994 data set, whereasinter-annual variability was based on plot data from 1991, 1992, and the July sample of 1994. There were nosignificant differences between watersheds for any sample year for any of the other herb layer characteristics measured, including herb layer cover, species richness, evenness, and diversity. Cover on WS4 decreased significantly from 1991 to 1992, followed by no change to 1994. By contrast, herb layer cover did not varysignificantly across years on WS3. Cover of the herbaceous layer of both watersheds increased from early in the growing season to the middle of the growing season, decreasing thereafter, with no significant differencesbetween WS3 and WS4 for any of the monthly cover means in 1994. Similar seasonal patterns found for herblayer cover—and lack of significant differences between watersheds—were also evident for species diversityand richness. By contrast, there was little seasonal change in herb layer species evenness, which was nearlyidentical between watersheds for all months except October. Seasonal patterns for individual species/speciesgroups were closely similar between watersheds, especially for Viola rotundifolia and Viola spp. Species richnessand species diversity were linearly related to herb layer cover for both WS3 and WS4, suggesting that spatialand temporal increases in cover were more related to recruitment of herb layer species than to growth of existingspecies. Results of this study indicate that there have been negligible responses of the herb layer to 6 yr of additions to WS3.

Effects of experimental nitrogen additions on plant diversity in an old-growth tropical forest

Response of plant biodiversity to increased availability of nitrogen (N) has been investigated in temperate and boreal forests, which are typically N-limited, but little is known in tropical forests. We examined the effects of artificial N additions on plant diversity (species richness, density and cover) of the understory layer in an N saturated old-growth tropical forest in southern China to test the following hypothesis: N additions decrease plant diversity in N saturated tropical forests primarily from N-mediated changes in soil properties. Experimental additions of N were administered at the following levels from July 2003 to July 2008: no addition (Control); 50 kg N ha−1 yr−1 (Low-N); 100 kg N ha−1 yr−1 (Medium-N), and 150 kg N ha−1 yr−1 (High-N). Results showed that no understory species exhibited positive growth response to any level of N addition during the study period. Although low-to-medium levels of N addition (≤100 kg N ha−1 yr−1) generally did not alter plant diversity through time, high levels of N addition significantly reduced species diversity. This decrease was most closely related to declines within tree seedling and fern functional groups, as well as to significant increases in soil acidity and Al mobility, and decreases in Ca availability and fine-root biomass. This mechanism for loss of biodiversity provides sharp contrast to competition-based mechanisms suggested in studies of understory communities in other forests. Our results suggest that high-N additions can decrease plant diversity in tropical forests, but that this response may vary with rate of N addition.