high-N Soil Research Articles

Soil organic carbon increase via microbial assimilation or soil protection against the priming effect is mediated by the availability of soil N relative to input C

Labile C inputs into soils will be partially transformed into soil organic carbon (SOC) through microbial assimilation or physicochemical protection as such mineral-organic interactions and soil aggregation. The C inputs may stimulate the decomposition of native SOC, inducing a phenomenon known as the priming effect. Increasing C inputs may increase SOC content, yet the relative role of these mechanisms in controlling the magnitudes of SOC increase among soils remains unclear. Four soils differing in microbiology and N content were incubated with 13C-labeled glucose at the amounts of 0.5 (G0.5), 1.0 (G1.0), and 2.0 (G2.0) g C kg−1 soil for 48 days. The objectives of this study were 1) to quantify the fates of added glucose-C, the priming effect, and the changes in SOC and mineral N contents after the incubation, and 2) to identify the main mechanisms for SOC increase with the increased amount of C input and the effects of initial soil N availability and microbial composition. 13C isotope probing showed that 19 to 26 % and 37 to 96 % of added glucose were utilized for microbial respiration and growth, respectively, after the incubation in all soils, with the glucose-C use efficiency ranging from 30 to 60 %. The remaining accounted for 4 to 37 % in all cases except for UltisolHN at G1.0 and G2.0 (–23 % and −2%), which was confirmed as the fraction protected in soil or re-utilized by microbes via acid extraction. The soil-protected glucose-C was higher, while glucose-C use efficiency was lower with the increased amount of added glucose only in the soils with low N. Glucose addition induced a positive priming effect, a SOC increase, and a mineral N loss irrespective of soil type and addition amount. Glucose addition increased the cumulative fungal biomass in all cases and the cumulative bacterial biomass only at G2.0 in all soils and at G1.0 in the high-N soils. It increased the final fungal necromass only at G0.5 in MollisolLN and UltisolHN and at G2.0 in MollisolHN and the final bacterial necromass in all cases except for the low-N soils and UltisolHN at G2.0. With the increased amount of added glucose, the SOC content increased linearly and the priming effect and fungal biomass increased non-linearly, with larger magnitudes in the soils with lower N. These results indicated that glucose was quickly protected after addition and remained in the soil until soil N became a limiting factor for microbial utilization. The availability of soil N relative to input C mediated microbial growth and its controls over SOC change via the soil protection and priming effects. The carbon sequestration potential with increased input of dissolved substrates as such glucose e. g. through root exudation was likely larger in soils with lower fertility due to reduced substrate-C use efficiency and increased effect of soil protection despite an increased priming effect. Further studies are needed on more substrate types as they affect substrate-C use efficiency and soil protection mechanisms.

Biological nitrogen fixation by irrigated dry bean (Phaseolus vulgaris L.) genotypes

Dry bean (Phaseolus vulgaris L.) is usually considered to be poor at biological nitrogen fixation (BNF), but large variations in this trait have been observed among bean genotypes. We evaluated 16 bean genotypes for N2 fixation ability in four N treatments: (i) uninoculated in low-N soil (30 kg N ha−1), (ii) inoculated with commercial Rhizobium leguminosarum bv. phaseoli inoculant Nitrastik-D® in low-N soil, (iii) inoculated with commercial R. leguminosarum bv. phaseoli inoculant Nodulator® in low-N soil, and (iv) uninoculated in high-N soil (100 kg N ha−1). There were differences between genotypes in all the plant parameters that were measured, but only nodulation was affected by N treatment. The 100 kg N ha−1 treatment suppressed nodulation. Seven genotypes nodulated well with either inoculant, two genotypes nodulated better with Nitrastik-D than with Nodulator, three nodulated better with Nodulator than with Nitrastik-D, and four nodulated poorly with either inoculant. Cultivars AC Redbond, Island, and Resolute, all currently commercially grown, did not fix much N2 at flowering (4–8 kg N ha−1) or maturity (19–34 kg N ha−1). By contrast, germplasm lines PI 136692 (red bean), GH-196 (pinto bean), and LEF2RB (carioca bean) had high BNF capability at flowering (10–11 kg N ha−1) and especially at maturity (60–72 kg N ha−1), in addition to high seed yield (2778–2897 kg ha−1), indicating their superior ability to support both of these economically important traits throughout plant growth. These three genotypes would be valuable to breeders for the genetic improvement of BNF in dry bean cultivars.

Influence of Nitrogen Rate, Seeding Rate, and Weed Removal Timing on Weed Interference in Barley and Effect of Nitrogen on Weed Response to Herbicides

Field experiments were conducted at the Montana State University Southern Agricultural Research Center, Huntley, MT, in 2011 through 2013 to determine the effect of nitrogen (N) rate, seeding rate, and weed removal timing on weed interference in barley. A delay in weed removal timing from the 3- to 4-leaf (LF) stage to the 8- to 10-LF stage of barley resulted in up to 3.5-fold increase in total weed biomass and 10% reduction in barley biomass, and this was unaffected by a N rate that ranged from 56 (low) to 168 (high) kg ha−1. The effect of N rate on barley biomass was more pronounced when weed removal was delayed from the 3- to 4-LF stage to the 8- to 10-LF stage of barley and in nontreated plots. Increasing the barley seeding rate from 38 to 152 kg ha−1increased the barley plant density by 50%, biomass by 13%, and grain yield by 29%, averaged over N rates and weed removal timing. On the basis of 5 and 10% levels of acceptable yield loss, the addition of ≥112 kg N ha−1delayed the critical timing of weed removal by at least 1.3 wk in barley, compared with the 56 kg N ha−1rate. A medium or high N rate prevented reduction in barley grain quality (plumpness and test weight) observed when the seeding rate was increased from 38 to 76 or 152 kg ha−1at the low N rate. In a separate greenhouse study, the effect of N rate on the effectiveness of various herbicides for controlling wild oat, green foxtail, kochia, or Russian thistle was investigated. Results highlighted that wild oat or green foxtail grown under 56 kg N ha−1(low N) soil required 1.4 to 2.6 times higher doses of clodinafop, fenoxaprop, flucarbazone, glyphosate, glufosinate, pinoxaden, or tralkoxydim for 50% reduction in shoot dry weights (GR50) compared with plants grown under 168 kg N ha−1(high N). Similarly, a reduced efficacy of thifensulfuron methyl + tribenuron methyl, metsulfuron methyl, or bromoxynil+pyrasulfotole was observed (evident from the GR50values) for kochia or Russian thistle grown under low- vs. high-N soil. Information gained from this research will aid in developing cost-effective, integrated weed management (IWM) strategies in cereals and in educating growers on the importance of fertilizer N management as a component of IWM programs.

Managing soil nitrogen to restore annual grass-infested plant communities: effective strategy or incomplete framework?

Theoretical and empirical work has established a positive relationship between resource availability and habitat invasibility. For nonnative invasive annual grasses, similar to other invasive species, invader success has been tied most often to increased nitrogen (N) availability. These observations have led to the logical assumption that managing soils for low N availability will facilitate restoration of invasive plant-dominated systems. Although invasive annual grasses pose a serious threat to a number of perennial-dominated ecosystems worldwide, there has been no quantitative synthesis evaluating the degree to which soil N management may facilitate restoration efforts. We used meta-analysis to evaluate the degree to which soil N management impacts growth and competitive ability of annual and perennial grass seedlings. We then link our analysis to current theories of plant ecological strategies and community assembly to improve our ability to understand how soil N management may be used to restore annual grass-dominated communities. Across studies, annual grasses maintained higher growth rates and greater biomass and tiller production than perennials under low and high N availability. We found no evidence that lowering N availability fundamentally alters competitive interactions between annual and perennial grass seedlings. Competitive effects of annual neighbors on perennial targets were similar under low and high N availability. Moreover, in most cases perennials grown under competition in high-N soils produced more biomass than perennials grown under competition in low-N soils. While these findings counter current restoration and soil N management assumptions, these results are consistent with current plant ecological strategy and community assembly theory. Based on our results and these theories we argue that, in restoration scenarios in which the native plant community is being reassembled from seed, soil N management will have no direct positive effect on native plant establishment unless invasive plant propagule pools and priority effects are controlled the first growing season.

Managing soil nitrogen to restore annual grass infested plant communities: An effective strategy or incomplete framework?

Theoretical and empirical work has established a positive relationship between resource availability and habitat invasibility. For nonnative invasive annual grasses, similar to other invasive species, invader success has been tied most often to increased nitrogen (N) availability. These observations have led to the logical assumption that managing soils for low N availability will facilitate restoration of invasive plant-dominated systems. Although invasive annual grasses pose a serious threat to a number of perennial-dominated ecosystems worldwide, there has been no quantitative synthesis evaluating the degree to which soil N management may facilitate restoration efforts. We used meta-analysis to evaluate the degree to which soil N management impacts growth and competitive ability of annual and perennial grass seedlings. We then link our analysis to current theories of plant ecological strategies and community assembly to improve our ability to understand how soil N management may be used to restore annual grass-dominated communities. Across studies, annual grasses maintained higher growth rates and greater biomass and tiller production than perennials under low and high N availability. We found no evidence that lowering N availability fundamentally alters competitive interactions between annual and perennial grass seedlings. Competitive effects of annual neighbors on perennial targets were similar under low and high N availability. Moreover, in most cases perennials grown under competition in high-N soils produced more biomass than perennials grown under competition in low-N soils. While these findings counter current restoration and soil N management assumptions, these results are consistent with current plant ecological strategy and community assembly theory. Based on our results and these theories we argue that, in restoration scenarios in which the native plant community is being reassembled from seed, soil N management will have no direct positive effect on native plant establishment unless invasive plant propagule pools and priority effects are controlled the first growing season.

Plant species, atmospheric CO2 and soil N interactively or additively control C allocation within plant-soil systems

Two plant species, Medicago truncatula (legume) and Avena sativa (non-legume), were grown in low- or high-N soils under two CO2 concentrations to test the hypothesis whether C allocation within plant-soil system is interactively or additively controlled by soil N and atmospheric CO2 is dependent upon plant species. The results showed the interaction between plant species and soil N had a significant impact on microbial activity and plant growth. The interaction between CO2 and soil N had a significant impact on soil soluble C and soil microbial biomass C under Madicago but not under Avena. Although both CO2 and soil N affected plant growth significantly, there was no interaction between CO2 and soil N on plant growth. In other words, the effects of CO2 and soil N on plant growth were additive. We considered that the interaction between N2 fixation trait of legume plant and elevated CO2 might have obscured the interaction between soil N and elevated CO2 on the growth of legume plant. In low-N soil, the shoot-to-root ratio of Avena dropped from 2.63 +/- 0.20 in the early growth stage to 1.47 +/- 0.03 in the late growth stage, indicating that Avena plant allocated more energy to roots to optimize nutrient uptake (i.e. N) when soil N was limiting. In high-N soil, the shoot-to-root ratio of Medicago increased significantly over time (from 2.45 +/- 0.30 to 5.43 +/- 0.10), suggesting that Medicago plants allocated more energy to shoots to optimize photosynthesis when N was not limiting. The shoot-to-root ratios were not significantly different between two CO2 levels.

Feedbacks of Soil Inoculum of Mycorrhizal Fungi Altered by N Deposition on the Growth of a Native Shrub and an Invasive Annual Grass

Anthropogenic nitrogen (N) deposition causes shifts in vegetation types as well as species composition of arbuscular mycorrhizal (AM) fungi and other soil microorganisms. A greenhouse experiment was done to determine whether there are feedbacks between N-altered soil inoculum and growth of a dominant native shrub and an invasive grass species in southern California. The region is experiencing large-scale loss of Artemisia californica shrublands and replacement by invasive annual grasses under N deposition. Artemisia californica and Bromus madritensis ssp. rubens were grown with soil inoculum from experimental plots in a low N deposition site that had (1) N-fertilized and (2) unfertilized soil used for inoculum, as well as (3) high-N soil inoculum from a site exposed to atmospheric N deposition for four decades. All treatments plus a nonmycorrhizal control were given two levels of N fertilizer solution. A. californica biomass was reduced by each of the three inocula compared to uninoculated controls under at least one of the two N fertilizer solutions. The inoculum from the N-deposition site caused the greatest growth depressions. By contrast, B. madritensis biomass increased with each of the three inocula under at least one, or both, of the N solutions. The different growth responses of the two plant species may be related to the types of AM fungal colonization. B. madritensis was mainly colonized by a fine mycorrhizal endophyte, while A. californica had primarily coarse endophytes. Furthermore, A. californica had a high level of septate, nonmycorrhizal root endophytes, while B. madritensis overall had low levels of these endophytes. The negative biomass response of A. californica seedlings to high N-deposition inoculum may in part explain its decline; a microbially-mediated negative feedback may occur in this system that causes poor seedling growth and establishment of A. californica in sites subject to N deposition and B. madritensis invasion.

Photosynthetic acclimation of overstory Populus tremuloides and understory Acer saccharum to elevated atmospheric CO2 concentration: interactions with shade and soil nitrogen.

We exposed Populus tremuloides Michx. and Acer saccharum Marsh. to a factorial combination of ambient and elevated atmospheric CO2 concentrations ([CO2]) and high-nitrogen (N) and low-N soil treatments in open-top chambers for 3 years. Our objective was to compare photosynthetic acclimation to elevated [CO2] between species of contrasting shade tolerance, and to determine if soil N or shading modify the acclimation response. Sun and shade leaf responses to elevated [CO2] and soil N were compared between upper and lower canopy leaves of P. tremuloides and between A. saccharum seedlings grown with and without shading by P. tremuloides. Both species had higher leaf N concentrations and photosynthetic rates in high-N soil than in low-N soil, and these characteristics were higher for P. tremuloides than for A. saccharum. Electron transport capacity (Jmax) and carboxylation capacity (Vcmax) generally decreased with atmospheric CO2 enrichment in all 3 years of the experiment, but there was no evidence that elevated [CO2] altered the relationship between them. On a leaf area basis, both Jmax and Vcmax acclimated to elevated [CO2] more strongly in shade leaves than in sun leaves of P. tremuloides. However, the apparent [CO2] x shade interaction was largely driven by differences in specific leaf area (m2 g-1) between sun and shade leaves. In A. saccharum, photosynthesis acclimated more strongly to elevated [CO2] in sun leaves than in shade leaves on both leaf area and mass bases. We conclude that trees rooted freely in the ground can exhibit photosynthetic acclimation to elevated [CO2], and the response may be modified by light environment. The hypothesis that photosynthesis acclimates more completely to elevated [CO2] in shade-tolerant species than in shade-intolerant species was not supported.

Genotypic variation in physiological and growth responses of Populus tremuloides to elevated atmospheric CO2 concentration.

Physiological and biomass responses of six genotypes of Populus tremuloides Michx., grown in ambient t (357 micromol mol(-1)) or twice ambient (707 micromol mol(-1)) CO2 concentration ([CO2]) and in low-N or high-N soils, were studied in 1995 and 1996 in northern Lower Michigan, USA. There was a significant CO2 x genotype interaction in photosynthetic responses. Net CO2 assimilation (A) was significantly enhanced by elevated [CO2] for five genotypes in high-N soil and for four genotypes in low-N soil. Enhancement of A by elevated [CO2] ranged from 14 to 68%. Genotypes also differed in their biomass responses to elevated [CO2], but biomass responses were poorly correlated with A responses. There was a correlation between magnitude of A enhancement by elevated [CO2] and stomatal sensitivity to CO2. Genotypes with low stomatal sensitivity to CO2 had a significantly higher A at elevated [CO2] than at ambient [CO2], but elevated [CO2] did not affect the ratio of intercellular [CO2] to leaf surface [CO2]. Stomatal conductance and A of different genotypes responded differentially to recovery from drought stress. Photosynthetic quantum yield and light compensation point were unaffected by elevated [CO2]. We conclude that P. tremuloides genotypes will respond differentially to rising atmospheric [CO2], with the degree of response dependent on other abiotic factors, such as soil N and water availability. The observed genotypic variation in growth could result in altered genotypic representation within natural populations and could affect the composition and structure of plant communities in a higher [CO2] environment in the future.

ATMOSPHERIC CO2, SOIL-N AVAILABILITY, AND ALLOCATION OF BIOMASS AND NITROGEN BYPOPULUS TREMULOIDES

Our ability to predict whether elevated atmospheric CO2 will alter the cycling of C and N in terrestrial ecosystems requires understanding a complex set of feedback mechanisms initiated by changes in C and N acquisition by plants and the degree to which changes in resource acquisition (C and N) alter plant growth and allocation. To gain further insight into these dynamics, we grew six genotypes of Populus tremuloides Michx. that differ in autumnal senescence (early vs. late) under experimental atmospheric CO2 (35.7 and 70.7 Pa) and soil-N availability (low and high) treatments. Atmospheric CO2 concentrations were manipulated with open-top chambers, and soil-N availability was modified in open-bottom root boxes by mixing different proportions of native A and C horizon soil. Net N mineralization rates averaged 61 ng N·g−1·d−1 in low-N soil and 319 ng N·g−1·d−1 in high-N soil. After 2.5 growing seasons, we harvested above- and belowground plant components in each chamber and determined total biomass, N concentration, N content, and the relative allocation of biomass and N to leaves, stems, and roots. Elevated CO2 increased total plant biomass 16% in low-N soil and 38% in high-N soil, indicating that the growth response of P. tremuloides to elevated CO2 was constrained by soil-N availability. Greater growth under elevated CO2 did not substantially alter the allocation of biomass to above- or belowground plant components. At both levels of soil-N availability, elevated CO2 decreased the N concentration of all plant tissues. Despite declines in tissue N concentration, elevated CO2 significantly increased whole-plant N content in high-N soil (ambient = 137 g N/chamber; elevated = 155 g N/chamber), but it did not influence whole-plant N content in low-N soil (36 g N/chamber). Our results indicate that plants in high-N soil obtained greater amounts of soil N under elevated CO2 by producing a proportionately larger fine-root system that more thoroughly exploited the soil. The significant positive relationship between fine-root biomass and total-plant N content we observed in high-N soil further supports this contention. In low-N soil, elevated CO2 did not increase fine-root biomass or production, and plants under ambient and elevated CO2 obtained equivalent amounts of N from soil. In high-N soil, it appears that greater acquisition of soil N under elevated CO2 fed forward within the plant to increase rates of C acquisition, which further enhanced plant growth response to elevated CO2.

ATMOSPHERIC CO2AND THE COMPOSITION AND FUNCTION OF SOIL MICROBIAL COMMUNITIES

Elevated atmospheric CO2 has the potential to increase the production and alter the chemistry of organic substrates entering soil from plant production, the magnitude of which is constrained by soil-N availability. Because microbial growth in soil is limited by substrate inputs from plant production, we reasoned that changes in the amount and chemistry of these organic substrates could affect the composition of soil microbial com- munities and the cycling of N in soil. We studied microbial community composition and soil-N transformations beneath Populus tremuloides Michx. growing under experimental atmospheric CO2 (35.7 and 70.7 Pa) and soil-N-availability (low N 5 61 ng N·g 21 ·d 21 and high N 5 319 ng N·g 21 ·d 21 ) treatments. Atmospheric CO2 concentration was modified in large, open-top chambers, and we altered soil-N availability in open-bottom root boxes by mixing different proportions of A and C horizon material. We used phospholipid fatty-acid analysis to gain insight into microbial community composition and coupled this analysis to measurements of soil-N transformations using 15 N-pool dilution techniques. The infor- mation presented here is part of an integrated experiment designed to elucidate the phys- iological mechanisms controlling the flow of C and N in the plant-soil system. Our ob- jectives were (1) to determine whether changes in plant growth and tissue chemistry alter microbial community composition and soil-N cycling in response to increasing atmospheric CO2 and soil-N availability and (2) to integrate the results of our experiment into a synthesis of elevated atmospheric CO2 and the cycling of C and N in terrestrial ecosystems. After 2.5 growing seasons, microbial biomass, gross N mineralization, microbial im- mobilization, and nitrification (gross and net) were equivalent at ambient and elevated CO2, suggesting that increases in fine-root production and declines in fine-root N concentration were insufficient to alter the influence of native soil organic matter on microbial physiology; this was the case in both low- and high-N soil. Similarly, elevated CO2 did not alter the proportion of bacterial, actinomycetal, or fungal phospholipid fatty acids in low-N or high-N soil, indicating that changes in substrate input from greater plant growth under elevated CO2 did not alter microbial community composition. Our results differ from a substantial number of studies reporting increases and decreases in soil-N cycling under elevated CO 2. From our analysis, it appears that soil-N cycling responds to elevated atmospheric CO 2 in experimental situations where plant roots have fully colonized the soil and root-associated C inputs are sufficient to modify the influence of native soil organic matter on microbial physiology. In young developing ecosystems where plant roots have not fully exploited the soil, microbial metabolism appears to be regulated by relatively large pools of soil organic matter, rather than by the additional input of organic substrates under elevated CO 2.

INTERACTIVE EFFECTS OF ATMOSPHERIC CO2AND SOIL-N AVAILABILITY ON FINE ROOTS OFPOPULUS TREMULOIDES

The objective of this experiment was to understand how atmospheric carbon dioxide (CO2) and soil-nitrogen (N) availability influence Populus tremuloides fine-root growth and morphology. Soil-N availability may limit the growth response of forests to elevated CO2 and interact with atmospheric CO2 to alter litter quality and ecosystem carbon (C) and N cycling. We established a CO2 × N factorial field experiment and grew six genotypes of P. tremuloides for 2.5 growing seasons in 20 large open-top chamber/root-box experimental units at the University of Michigan Biological Station in northern lower Michigan (USA). In this paper we describe an integrated examination of how atmospheric CO2 and soil-N availability influence fine-root morphology, growth, mortality, and biomass. We also studied the relationship between root biomass and total soil respiration. Over 80% of the absorbing root length of P. tremuloides was accounted for by roots <0.4 mm in diameter, and specific root length (100–250 m/g) was much greater than reports for other temperate and boreal deciduous trees. Elevated atmospheric CO2 increased the diameter and length of individual roots. In contrast, soil N had no effect on root morphology. Fine-root length production and mortality, measured with minirhizotrons, was controlled by the interaction between atmospheric CO2 and soil N. Rates of root production and mortality were significantly greater at elevated CO2 when trees grew in high-N soil, but there were no CO2 effects at low soil N. Fine-root biomass increased 137–194% in high-N compared to low-N soil, and elevated atmospheric CO2 increased fine-root biomass (52%) in high soil N, but differences in low soil N were not significant. Across all treatments, dynamic estimates of net fine-root production were highly correlated with fine-root biomass (soil cores; r = 0.975). Mean rates of soil respiration were more than double in high-N compared to low-N soil, and elevated atmospheric CO2, when compared to ambient atmospheric CO2, increased mean rates of soil respiration 19% in 1995 and 25% in 1996. Across all treatments, total root biomass was linearly related to mean rates of soil respiration (r2 = 0.96). Our results indicate that atmospheric CO2 and soil-N availability strongly interact to influence P. tremuloides fine-root morphology, growth, and C turnover. Aspen-dominated ecosystems of the future are likely to have greater productivity fueled by greater nutrient uptake due to greater root length production. Further, it appears that elevated atmospheric CO2 will result in greater C inputs to soil through greater rates of fine-root production and turnover, especially in high-fertility soils. Increased C inputs to soil result in greater rates of soil respiration. At this time, it is not clear what effects increased rates of root turnover will have on C storage in the soil.

GAS EXCHANGE, LEAF NITROGEN, AND GROWTH EFFICIENCY OFPOPULUS TREMULOIDESIN A CO2-ENRICHED ATMOSPHERE

Predicting forest responses to rising atmospheric CO2 will require an understanding of key feedbacks in the cycling of carbon and nitrogen between plants and soil microorganisms. We conducted a study for 2.5 growing seasons with Populus tremuloides grown under experimental atmospheric CO2 and soil-N-availability treatments. Our objective was to integrate the combined influence of atmospheric CO2 and soil-N availability on the flow of C and N in the plant–soil system and to relate these processes to the performance of this widespread and economically important tree species. Here we consider treatment effects on photosynthesis and canopy development and the efficiency with which this productive capacity is translated into aboveground, harvestable yield. We grew six P. tremuloides genotypes at ambient (35 Pa) or elevated (70 Pa) CO2 and in soil of low or high N mineralization rate at the University of Michigan Biological Station, Pellston, Michigan, USA (45°35′ N, 84°42′ W). In the second year of growth, net CO2 assimilation rate was significantly higher in elevated-CO2 compared to ambient-CO2 plants in both soil-N treatments, and we found little evidence for photosynthetic acclimation to high CO2. In the third year, however, elevated-CO2 plants in low-N soil had reduced photosynthetic capacity compared to ambient-CO2, low-N plants. Plants in high-N soil showed the opposite response, with elevated-CO2 plants having higher photosynthetic capacity than ambient-CO2 plants. Net CO2 assimilation rate was linearly related to leaf N concentration (log:log scale), with identical slopes but different intercepts in the two CO2 treatments, indicating differences in photosynthetic N-use efficiency. Elevated CO2 increased tissue dark respiration in high-N soil (+22%) but had no significant effect in low-N soil (+9%). There were no CO2 effects on stomatal conductance. At the final harvest, stem biomass and total leaf area increased significantly due to CO2 enrichment in high-N but not in low-N soil. Treatment effects on wood production were largely attributable to changes in leaf area, with no significant effects on growth efficiency. We conclude that harvest intervals for P. tremuloides on fertile sites will shorten with rising atmospheric CO2, but that tree size at canopy closure may be unaffected.

Soil nematodes indicate food web responses to elevated atmospheric CO 2

Summary To understand the impact of rising levels of atmospheric CO 2 on ecosystems, we need to understand plant responses to elevated CO 2 , as well as how those plant responses in turn affect their environment. An important component of the environment of a plant is the soil biota living near plant roots. Soil nematodes are representative of a large portion of this biota, since they are abundant and trophically diverse in most soils. In a three-year field experiment, we studied the responses of soil nematodes to increased root growth of trees growing in high and low nitrogen soils under ambient and twice-ambient atmospheric CO 2 , a two-by-two factorial experimental design. Our hypothesis was that in the high-N soil, increased root growth resulting from twice-ambient atmospheric CO 2 would positively affect nematode density, supporting a more abundant and trophically complex nematode community. Trembling aspen ( Populus tremuloides ) were grown in twenty open-top chambers under the four treatments, replicated five times. In low-N soil, twice-ambient CO 2 was associated with higher density of the most abundant plant-feeding taxon (Trichodoridae), lower density of one bacteriafeeding taxon (Rhabditidae), and lower evenness of the community, compared to ambient CO 2 . In high-N soil, twice-ambient CO 2 was associated with higher density of predator/omnivores, lower diversity, and a larger value of Bonger's Maturity Index, compared to ambient CO 2 . In soils under young deciduous trees, such as the aspens in this experiment, increased root growth under elevated CO 2 may result in significant changes in soil food web community structure that may provide clues about the fate of carbon under elevated CO 2 .

Ionic balance and organic acids in western redcedar, western hemlock, and Douglas-fir seedlings grown in low- and high-N soils

Seedlings of western redcedar (Thuja plicata Donn ex. D. Don), western hemlock (Tsuga heterophylla (Raf.) Sarg.), and Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) were transplanted into soils with low and high levels of available NO3-(and total N). Current-year foliage was sampled after 10 weeks to determine the effect of N availability on foliar cation-anion balance (C-A) and the concentrations of low molecular weight organic acids of the three species. Carboxylate concentrations were estimated by using the difference between sums of cations and anions (C-A): 750 mequiv.·kg-1for western redcedar, 351 mequiv.·kg-1for western hemlock, and 266 mequiv.·kg-1for Douglas-fir. Quinic acid was a primary constituent, accounting for 40% of the total for western redcedar and 75% for western hemlock and Douglas-fir. Oxalic acid was present in greatest concentration in the foliage of western redcedar (65 mequiv.·kg-1) but was a minor constituent in western hemlock and Douglas-fir. The quantified acids accounted for only 15% of the C-A of western redcedar but &gt;80% of the C-A of western hemlock and Douglas-fir. A considerable portion of the C-A balance not accounted for in redcedar may be associated with the accumulation of CaCO3. Litterfall deposition of CaCO3may lead to the consumption of H+ions and enrichment of exchangeable soil Ca in the rooting zone of long-lived western redcedar trees. No statistically significant differences among the soils were detected with regard to C-A or the concentration of organic acids.

Dry matter and nitrogen allocation in western redcedar, western hemlock, and Douglas fir seedlings grown in low- and high-N soils

Des semis de Thuja plicata Don ex. D. Don, de Tsuga heterophylla (Raf.) Sarg et de Douglas (Pseudotsuga menziesii (Mirb.) Franco) ont ete replantes dans 48 pots contenant du sol avec des quantites faibles ou elevees de nitrate NO 3 - (et d'azote total) selon 4 traitements : temoin sans apport, fertilise avec 60 mg kg -1 de (NH 4 )2SO 4 , un apport de 15 mg kg -1 de dicyandiamide (DCD) un inhibiteur de la nitrification, un traitement combine avec 60 mg kg -1 de (NH 4 )2SO 4 et le DCD. Le poids sec et le contenu en azote des tissus des semis ont ete determines apres 8 semaines. Les semis du traitement a faibles reserves en azote assimilable ont accumule 65 % de matiere seche et 40 % de l'azote relativement au traitement a haut niveau d'azote. Le transfert interne d'azote depuis les aiguilles d'un an et les compartiments du tronc et des branches chez le Thuja et le Douglas, mais pas pour le Tsuga, est une importante source d'azote pour la croissance des aiguilles et racines de l'annee dans le traitement le plus pauvre en azote. Le Tsuga presente l'accroissement relatif le plus eleve en matiere seche (Log e (DM final ) - Log e (DM initial ) = RDMI) et en azote (Log e (N final ) - Log e (N intial ) = RNI) pour chacun des traitements et a accumule 35 % d'azote en plus pour le traitement faible et 10 % en plus pour le traitement eleve que les autres especes. Le RDMI du Thuja est intermediaire entre celui du Tsuga et du Douglas, alors que son RNI est le plus faible quel que soit le sol. Les resultats suggerent que le Tsuga est plus efficient que le Thuja ou le Douglas pour l'absorption d'azote inorganique, specialement dans les sols pauvres en azote.

Ionic balance and organic acids in western redcedar, western hemlock, and Douglas-fir seedlings grown in low- and high-N soils

Ionic balance and organic acids in western redcedar, western hemlock, and Douglas-fir seedlings grown in low- and high-N soils

Response of soil biota to elevated atmospheric CO2 in poplar model systems.

We tested the hypotheses that increased belowground allocation of carbon by hybrid poplar saplings grown under elevated atmospheric CO2 would increase mass or turnover of soil biota in bulk but not in rhizosphere soil. Hybrid poplar saplings (Populus×euramericana cv. Eugenei) were grown for 5 months in open-bottom root boxes at the University of Michigan Biological Station in northern, lower Michigan. The experimental design was a randomized-block design with factorial combinations of high or low soil N and ambient (34 Pa) or elevated (69 Pa) CO2 in five blocks. Rhizosphere microbial biomass carbon was 1.7 times greater in high-than in low-N soil, and did not respond to elevated CO2. The density of protozoa did not respond to soil N but increased marginally (P < 0.06) under elevated CO2. Only in high-N soil did arbuscular mycorrhizal fungi and microarthropods respond to CO2. In high-N soil, arbuscular mycorrhizal root mass was twice as great, and extramatrical hyphae were 11% longer in elevated than in ambient CO2 treatments. Microarthropod density and activity were determined in situ using minirhizotrons. Microarthropod density did not change in response to elevated CO2, but in high-N soil, microarthropods were more strongly associated with fine roots under elevated than ambient treatments. Overall, in contrast to the hypotheses, the strongest response to elevated atmospheric CO2 was in the rhizosphere where (1) unchanged microbial biomass and greater numbers of protozoa (P < 0.06) suggested faster bacterial turnover, (2) arbuscular mycorrhizal root length increased, and (3) the number of microarthropods observed on fine roots rose.