Recent Land Use Change to Agriculture in the U.S. Lake States: Impacts on Cellulosic Biomass Potential and Natural Lands.

Perennial cellulosic feedstocks may have potential to reduce life-cycle greenhouse gas (GHG) emissions by offsetting fossil fuels. However, this potential depends on meeting a number of important criteria involving land cover change, including avoiding displacement of agricultural production, not reducing uncultivated natural lands that provide biodiversity habitat and other valued ecosystem services, and avoiding the carbon debt (the amount of time needed to repay the initial carbon loss) that accompanies displacing natural lands. It is unclear whether recent agricultural expansion in the United States competes with lands potentially suited for bioenergy feedstocks. Here, we evaluate how recent land cover change (2008-2013) has affected the availability of lands potentially suited for bioenergy feedstock production in the U.S. Lake States (Minnesota, Wisconsin, Michigan) and its impact on other natural ecosystems. The region is potentially well suited for a diversity of bioenergy production systems, both grasses and woody biomass, due to the widespread forest economy in the north and agricultural economy in the south. Based on remotely-sensed data, our results show that between 2008 and 2013, 836,000 ha of non-agricultural open lands were already converted to agricultural uses in the Lake States, a loss of nearly 37%. The greatest relative changes occurred in the southern half that includes some of the most diverse cultivable lands in the country. We use transition diagrams to reveal gross changes that can be obscured if only net change is considered. Our results indicate that expansion of row crops (corn, soybean) was responsible for the majority of open land loss. Even if recently lost open lands were brought into perennial feedstock production, there would a substantial carbon debt. This reduction in open land availability for biomass production is closing the window of opportunity to establish a sustainable cellulosic feedstock economy in the Lake States as mandated by current Federal policy, incurring a substantial GHG debt, and displacing a range of other natural ecosystems and their services.

Ectomycorrhizal fungal succession coincides with shifts in organic nitrogen availability and canopy closure in post-wildfire jack pine forests.

Successional changes in belowground ectomycorrhizal fungal (EMF) communities have been observed with increasing forest stand age; however, mechanisms behind this change remain unclear. It has been hypothesized that declines of inorganic nitrogen (N) and increases of organic N influence changes in EMF taxa over forest development. In a post-wildfire chronosequence of six jack pine (Pinus banksiana) stands ranging in age from 5 to 56 years, we investigated EMF community composition and compared shifts in taxa with detailed soluble inorganic and organic N data. Taxa were identified by internal transcribed spacer rDNA sequencing, and changes in community composition evaluated with non-metric multi-dimensional scaling (NMDS). Dissimilarities in the community data were tested for correlations with N variables. We observed a successional shift along NMDS axis 1 from such taxa as Suillus brevipes and Thelephora terrestris in sites age 5 and 11 to species of Cortinarius and Russula, among others, in the four older sites. This change was positively correlated with soluble organic N (SON) (r(2) = 0.902, P = 0.033) and free amino-acid N (r(2) = 0.945, P = 0.021), but not inorganic N. Overall, our results show a successional shift of EMF communities occurring between stand initiation and canopy closure without a change in species of the dominant plant-host, and associated with SON and free amino-acid N in soil. It is uncertain whether EMF taxa are responding to these organic N forms directly, affecting their availability, or are ultimately responding to changes in other site variables, such as belowground productivity.

Amino acid uptake by temperate tree species characteristic of low- and high-fertility habitats.

The relationship between inorganic nitrogen (N) cycling and plant productivity is well established. However, recent research has demonstrated the ability of plants to take up low molecular weight organic N compounds (i.e., amino acids) at rates that often rival those of inorganic N forms. In this study, we hypothesize that temperate forest tree species characteristic of low-fertility habitats will prefer amino acids over species characteristic of high-fertility habitats. We measured the uptake of (15)N-labeled amino acids (glycine, glutamine, arginine, serine), ammonium (NH(4)(+)), and nitrate (NO(3)(-)) by four tree species that commonly occur in eastern North America, where their abundances have been correlated with inorganic N availability. Specific uptake rates of amino acids were largely similar for all tree species; however, high-fertility species took up NH(4)(+) at rates more than double those of low-fertility species, rendering amino acid N relatively more important to the N nutrition of low-fertility species. Low-fertility species acquired over four times more total N from arginine compared to NH(4)(+) and NO(3)(-); high-fertility species acquired the most N from NH(4)(+). Arginine had the highest uptake rates of any amino acid by all species; there were no significant differences in uptake rates of the remaining amino acids. Our results support the idea that the dominant species in a particular habitat are those best able to utilize the most available N resources.

Plant-available organic and mineral nitrogen shift in dominance with forest stand age.

Studies of soil nitrogen (N) availability over stand development have almost exclusively focused on mineral N, yet we increasingly recognize that plants can take up organic N in the form of free amino acids at biologically important rates. We investigated amino-acid and mineral N availability along a 10-site chronosequence of jack pine stands, varying in age from 4 to 60 yr following wildfire. We measured free amino-acid N and mineral N in soil extracts; native proteolytic rates; net N mineralization rates; and microbial amino-acid consumption via a 15N leucine tracer assay in 6 of the 10 sites (4, 10, 18, 22, 46, and 55-yr-old). Amino-acid N was consistently low in the youngest sites (4-10 yr), increased rapidly in mid-aged sites (15-22 yr), and was highest in stand age 46. In contrast, mineral N exhibited a parabolic shape (R2 0.499; P < 0.0001), with the youngest site and the four oldest sites containing the highest amounts of mineral N. As a result, amino-acid N as a percentage of amino-acid N + mineral N was greatest in mid-aged stands (e.g., 67% in the 22-yr-old stand). We observed no trend in proteolytic rates across the chronosequence (P = 0.632). Percentage 15N tracer recovery was lowest in the extractable organic N pool for the 4, 10, and 18-yr-old sites, though only site age 10 was significantly different from the older sites. Percentage of recovery in the organic N pool was significantly positively related (R2 = 0.798; P < 0.05) to standing pools of amino-acid N. Overall, our results suggest that heterotrophic consumption, not production via proteolysis, controls soil free amino-acid availability. Higher microbial demand for free amino acids in younger vs. older sites likely results from greater microbial C and N limitation early in stand development due to the lack of fresh litter inputs. Since aminoacid N exceeds mineral N in a time period of stand development where jack pine growth rates and N demand are highest, we speculate that amino-acid N may be important to the N economy of these forests.

Changes in soil carbon and nitrogen cycling along a 72-year wildfire chronosequence in Michigan jack pine forests.

We investigated the changes in soil processes following wildfire in Michigan jack pine (Pinus banksiana) forests using a chronosequence of 11 wildfire-regenerated stands spanning 72 years. The objective of this study was to characterize patterns of soil nutrients, soil respiration and N mineralization with stand development, as well as to determine the mechanisms driving those patterns. We measured in situ N mineralization and soil respiration monthly during the 2002 growing season and used multiple regression analysis to determine the important factors controlling these processes. Growing-season soil respiration rates ranged from a low of 156 g C/m2 in the 7-year-old stand to a high of 254 g C/m2 in the 22-year-old stand, but exhibited no clear pattern with stand age. In general, soil respiration rates peaked during the months of July and August when soil temperatures were highest. We used a modified gamma function to model a temporal trend in total N mineralization (total N mineralization=1.853-0.276xagexe-0.814xage; R2=0.381; P=0.002). Total N mineralization decreased from 2.8 g N/m2 in the 1-year-old stand to a minimum value of 0.5 g N/m2 in the 14-year-old stand, and then increased to about 1.5 g N/m2 in mature stands. Changes in total N mineralization were driven by a transient spike in N turnover in the mineral soil immediately after wildfire, followed by a gradual accrual of a slow-cycling pool of N in surface organic horizons as stands matured. Thus, in Michigan jack pine forests, the accumulation of surface organic matter appears to regulate N availability following stand-replacing wildfire.

Kinetics of nitrogen uptake by Populus tremuloides in relation to atmospheric CO(2) and soil nitrogen availability.

Sustained increases in plant production in response to elevated atmospheric carbon dioxide (CO(2)) concentration may be constrained by the availability of soil nitrogen (N). However, it is possible that plants will respond to N limitation at elevated CO(2) concentration by increasing the specific N uptake capacity of their roots. To explore this possibility, we examined the kinetics of (15)NH(4) (+) and (15)NO(3) (-) uptake by excised roots of Populus tremuloides Michx. grown in ambient and twice-ambient CO(2) concentrations, and in soils of low- and high-N availability. Elevated CO(2) concentration had no effect on either NH(4) (+) or NO(3) (-) uptake, whereas high-N availability decreased the capacity of roots to take up both NH(4) (+) and NO(3) (-). The maximal rate of NH(4) (+) uptake decreased from 12 to 8 &mgr;mol g(-1) h(-1), and K(m) increased from 49 to 162 &mgr;mol l(-1), from low to high soil N availability.Because NO(3) (-) uptake exhibited mixedkinetics over the concentration range we used (10-500 &mgr;mol l( -1)), it was not possible to calculate V(max) and K(m). Instead, we used an uptake rate of 100 &mgr;mol g(-1) h(-1) as our metric of NO(3) (-) uptake capacity, which averaged 0.45 and 0.23 &mgr;mol g(-1) h(-1) at low- and high-N availability, respectively. The proximal mechanisms for decreased N uptake capacity at high-N availability appeared to be an increase in fine-root carbohydrate status and a decrease in fine-root N concentration. Both NH(4) (+) and NO(3) (-) uptake were inversely related to fine-root N concentration, and positively related to fine-root total nonstructural carbohydrate concentration. We conclude that soil N availability, through its effects on fine-root N and carbohydrate status, has a much greater influence on the specific uptake capacity of P. tremuloides fine roots than elevated atmospheric CO(2). In elevated atmospheric CO(2), changes in N acquisition by P. tremuloides appeared to be driven by changes in root architecture and biomass, rather than by changes in the amount or activity of N-uptake enzymes.

Nitrate deposition in northern hardwood forests and the nitrogen metabolism of Acer saccharum marsh.

It is generally assumed that plant assimilation constitutes the major sink for anthropogenic Nitrate NO 3- deposited in temperate forests because plant growth is usually limited by nitrogen (N) availability. Nevertheless, plants are known to vary widely in their capacity for NO 3- uptake and assimilation, and few studies have directly measured these parameters for overstory trees. Using a combination of field and greenhouse experiments, we studied the N nutrition of Acer saccharum Marsh. in four northern hardwood forests receiving experimental NO 3- additions equivalent to 30 kg N ha-1 year-1. We measured leaf and fine-root nitrate reductase activity (NRA) of overstory trees using an in vivo assay and used 15N to determine the kinetic parameters of NO 3- uptake by excised fine roots. In two greenhouse experiments, we measured leaf and root NRA in A. saccharum seedlings fertilized with 0-3.5 g NO 3- -N m-2 and determined the kinetic parameters of NO 3- and NH 4+ uptake in excised roots of seedlings. In both overstory trees and seedlings, rates of leaf and fine root NRA were substantially lower than previously reported rates for most woody plants and showed no response to NO 3- fertilization (range = non-detectable to 33 nmol NO 2- g-1 h-1). Maximal rates of NO 3- uptake in overstory trees also were low, ranging from 0.2 to 1.0 µmol g-1 h-1. In seedlings, the mean V max for NO 3- uptake in fine roots (1 µmol g-1 h-1) was approximately 30 times lower than the V max for NH 4+ uptake (33 µmol g-1 h-1). Our results suggest that A. saccharum satisfies its N demand through rapid NH 4+ uptake and may have a limited capacity to serve as a direct sink for atmospheric additions of NO 3- .