Nitrogen cycling during seven years of atmospheric CO2 enrichment in a scrub oak woodland

Experimentally increasing atmospheric CO2 often stimulates plant growth and ecosystem carbon (C) uptake. Biogeochemical theory predicts that these initial responses will immobilize nitrogen (N) in plant biomass and soil organic matter, causing N availability to plants to decline, and reducing the long-term CO2-stimulation of C storage in N limited ecosystems. While many experiments have examined changes in N cycling in response to elevated CO2, empirical tests of this theoretical prediction are scarce. During seven years of postfire recovery in a scrub oak ecosystem, elevated CO2 initially increased plant N accumulation and plant uptake of tracer 15N, peaking after four years of CO2 enrichment. Between years four and seven, these responses to CO2 declined. Elevated CO2 also increased N and tracer 15N accumulation in the O horizon, and reduced 15N recovery in underlying mineral soil. These responses are consistent with progressive N limitation: the initial CO2 stimulation of plant growth immobilized N in plant biomass and in the O horizon, progressively reducing N availability to plants. Litterfall production (one measure of aboveground primary productivity) increased initially in response to elevated CO2, but the CO2 stimulation declined during years five through seven, concurrent with the accumulation of N in the O horizon and the apparent restriction of plant N availability. Yet, at the level of aboveground plant biomass (estimated by allometry), progressive N limitation was less apparent, initially because of increased N acquisition from soil and later because of reduced N concentration in biomass as N availability declined. Over this seven-year period, elevated CO2 caused a redistribution of N within the ecosystem, from mineral soils, to plants, to surface organic matter. In N limited ecosystems, such changes in N cycling are likely to reduce the response of plant production to elevated CO2.     

Niche complementarity for nitrogen: an explanation for the biodiversity and ecosystem functioning relationship?

The relationship between plant diversity and productivity has largely been attributed to niche complementarity, assuming that plant species are complementary in their resource use. In this context, we conducted an 15N field study in three different grasslands, testing complementarity nitrogen (N) uptake patterns in terms of space, time, and chemical form as well as N strategies such as soil N use, symbiotic N fixation, or internal N recycling for different plant species. The relative contribution of different spatial, temporal, and chemical soil N pools to total soil N uptake of plants varied significantly among the investigated plant species, within and across functional groups. This suggests that plants occupy distinct niches with respect to their relative N uptake. However, when the absolute N uptake from the different soil N pools was analyzed, no spatial, temporal, or chemical variability was detected, but plants, and in particular functional groups, differed significantly with respect to their total soil N uptake irrespective of treatment. Consequently, our data suggest that absolute N exploitation on the ecosystem level is determined by species or functional group identity and thus by community composition rather than by complementary biodiversity effects. Across functional groups, total N uptake from the soil was negatively correlated with leaf N concentrations, suggesting that these functional groups follow different N use strategies to meet their N demands. While our findings give no evidence for a biodiversity effect on the quantitative exploitation of different soil N pools, there is evidence for different and complementary N strategies and thus a potentially beneficial effect of functional group diversity on ecosystem functioning.     

Atmospheric CO2 and O3 alter the flow of 15N in developing forest ecosystems

Anthropogenic O3 and CO2-induced declines in soil N availability could counteract greater plant growth in a CO2-enriched atmosphere, thereby reducing net primary productivity (NPP) and the potential of terrestrial ecosystems to sequester anthropogenic CO2. Presently, it is uncertain how increasing atmospheric CO2 and O3 will alter plant N demand and the acquisition of soil N by plants as well as the microbial supply of N from soil organic matter. To address this uncertainty, we initiated an ecosystem-level 15N tracer experiment at the Rhinelander (Wisconsin, USA) free air CO2-O3 enrichment (FACE) facility to understand how projected increases in atmospheric CO2 and 03 alter the distribution and flow of N in developing northern temperate forests. Tracer amounts of 15NH4+ were applied to the forest floor of developing Populus tremuloides and P. tremuloides-Betula papyrifera communities that have been exposed to factorial CO2 and O3 treatments for seven years. One year after isotope addition, both forest communities exposed to elevated CO2 obtained greater amounts of 15N (29%) and N (40%) from soil, despite no change in soil N availability or plant N-use efficiency. As such, elevated CO2 increased the ability of plants to exploit soil for N, through the development of a larger root system. Conversely, elevated O3 decreased the amount of 15N (-15%) and N (-29%) in both communities, a response resulting from lower rates of photosynthesis, decreases in growth, and smaller root systems that acquired less soil N. Neither CO2 nor 03 altered the amount of N or 15N recovery in the forest floor, microbial biomass, or soil organic matter. Moreover, we observed no interaction between CO2 and 03 on the amount of N or 15N in any ecosystem pool, suggesting that 03 could exert a negative effect regardless of CO2 concentration. In a CO2-enriched atmosphere, greater belowground growth and a more thorough exploitation of soil for growth-limiting N is an important mechanism sustaining the enhancement of NPP in developing forests (0-8 years following establishment). However, as CO2 accumulates in the Earth's atmosphere, future O3 concentrations threaten to diminish the enhancement of plant growth, decrease plant N acquisition, and lessen the storage of anthropogenic C in temperate forests.     

Grassland establishment under varying resource availability: a test of positive and negative feedback

The traditional logic of carbon (C) and nitrogen (N) interactions in ecosystems predicts further increases or decreases in productivity (positive feedback) in response to high and low fertility in the soil, respectively; but the potential for development of feedback in ecosystems recovering from disturbance is less well understood. Furthermore, this logic has been challenged in grassland ecosystems where frequent fires or grazing may reduce the contribution of aboveground litter inputs to soil organic matter pools and nutrient supply for plant growth, relative to forest ecosystems. Further, if increases in plant productivity increase soil C content more than soil N content, negative feedback may result from increased microbial demand for N making less available for plant growth. We used a field experiment to test for feedback in an establishing grassland by comparing aboveground net primary productivity (ANPP) and belowground pools and fluxes of C and N in soil with enriched, ambient, and reduced N availability. For eight years annual N enrichment increased ANPP, root N, and root tissue quality, but root C:N ratios remained well above the threshold for net mineralization of N. There was no evidence that N enrichment increased root biomass, soil C or N accrual rates, or storage of C in total, microbial, or mineralizable pools within this time frame. However, the net nitrogen mineralization potential (NMP) rate was greater following eight years of N enrichment, and we attributed this to N saturation of the microbial biomass. Grassland developing under experimentally imposed N limitation through C addition to the soil exhibited ANPP, root biomass and quality, and net NMP rate similar to the ambient soil. Similarity in productivity and roots in the reduced and ambient N treatments was attributed to the potentially high nitrogen-use efficiency (NUE) of the dominant C4 grasses, and increasing cover of legumes over time in the C-amended soil. Thus, in a developing ecosystem, positive feedback between soil N supply and plant productivity may promote enhanced long-term N availability and override progressive N limitation as C accrues in plant and soil pools. However, experimentally imposed reduction in N availability did not feed back to reduce ANPP, possibly due to shifts in NUE and functional group composition.     

Elevated CO2 stimulates net accumulations of carbon and nitrogen in land ecosystems: a meta-analysis

The capability of terrestrial ecosystems to sequester carbon (C) plays a critical role in regulating future climatic change yet depends on nitrogen (N) availability. To predict long-term ecosystem C storage, it is essential to examine whether soil N becomes progressively limiting as C and N are sequestered in long-lived plant biomass and soil organic matter. A critical parameter to indicate the long-term progressive N limitation (PNL) is net change in ecosystem N content in association with C accumulation in plant and soil pools under elevated CO2. We compiled data from 104 published papers that study C and N dynamics at ambient and elevated CO2. The compiled database contains C contents, N contents, and C:N ratio in various plant and soil pools, and root:shoot ratio. Averaged C and N pool sizes in plant and soil all significantly increase at elevated CO2 in comparison to those at ambient CO2, ranging from a 5% increase in shoot N content to a 32% increase in root C content. The C and N contents in litter pools are consistently higher in elevated than ambient CO2 among all the surveyed studies whereas C and N contents in the other pools increase in some studies and decrease in other studies. The high variability in CO2-induced changes in C and N pool sizes results from diverse responses of various C and N processes to elevated CO2. Averaged C:N ratios are higher by 3% in litter and soil pools and 11% in root and shoot pools at elevated relative to ambient CO2. Elevated CO2 slightly increases root:shoot ratio. The net N accumulation in plant and soil pools at least helps prevent complete down-regulation of, and likely supports, long-term CO2 stimulation of C sequestration. The concomitant C and N accumulations in response to rising atmospheric CO2 may reflect intrinsic nature of ecosystem development as revealed before by studies of succession over hundreds to millions of years.     

Urbanization increases grassland carbon pools: effects of landscaping in Colorado's front range.

During the past few decades, urban and suburban developments have grown at unprecedented rates and extents with unknown consequences for ecosystem function. Carbon pools of soil and vegetation on landscaped properties were examined in the Front Range of Colorado, USA, in order to characterize vegetation and soils found in urban green spaces; analyze their aboveground biomass, vegetative C storage, and soil C storage; and compare these suburban ecosystem properties to their counterparts in native grassland and cultivated fields. Anthropogenic activities leave clear signatures on all three C compartments measured. Management level dominates the response of grass production, biomass, and N tissue concentration. This, in turn, influences the amount of C and N both stored in and harvested from sites. The site age dominates the amount of woody biomass as well as soil C and N. Soil texture only secondarily affects total soil carbon and total bulk density. Established urban green spaces harbor larger C pools, more than double in some cases, than native grasslands or agricultural fields on a per-area basis. Lawn grass produces more biomass and stores more C than local prairie or agricultural fields. Introduced woody vegetation comprises a substantial C pool in urban green spaces and represents a new ecosystem feature. After an initial decrease with site development, soil organic carbon (SOC) pools surpass those in grasslands within two decades. In addition to the marked increase of C pools through time, a shift in storage from belowground to aboveground occurs. Whereas grasslands store approximately 90% of C belowground, urban green spaces store a decreasing proportion of the total C belowground in soils through time, reaching approximately 70% 30-40 years after construction. Despite the substantial increase in C pools in this urban area, it is important to recognize that this shift is distinct from C sequestration since it does not account for a total C budget, including increased anthropogenic C emissions from these sites.     

东北北部温带森林和干草地土壤养分分布及影响因素

土壤是陆地生态系统碳储存的重要场所,其养分变化与全球陆地碳循环密切相关。土壤养分是植物生长的重要保证,而土壤各养分之间是紧密联系的。理解土壤养分变化与环境因素的关系有助于更好地了解陆地生态系统碳、氮、磷循环。本研究以东北北部自东向西沿降水量梯度变化纬度带上的温带森林与干草地生态系统为研究对象,利用气象数据和野外土壤实测数据,分析了纬度带上不同植被类型土壤的有机碳、全氮、碳氮比、速效磷的空间分布格局及其与环境因子（年降水量、年均温、土壤pH值）的关系。研究纬度带上降水量自东向西逐渐减少,植被类型从温带森林过渡到干草原,与降水量和植被类型对应,植被生物量也自东向西呈现从高到低的分布梯度。研究结果表明：从整个研究带上来说,降水量与土壤pH值是土壤养分空间分布的决定因素,沿纬度带从东到西,随着降水量逐渐减少,土壤pH值逐渐增加,而土壤有机碳、全氮、碳氮比、速效磷含量逐渐减少。但如果将森林和草地分别讨论则发现,森林和草地生态系统的土壤养分环境控制因素有较大差别。对于草地生态系统而言,降水量和土壤pH值仍然是其土壤养分含量的控制因子,但森林生态系统由于所处区域降水量充足,降水量不再是其土壤养分的控制因子,降水量只与森林土壤碳氮比呈显著正相关。研究还发现森林土壤的速效磷含量与温度呈正相关,与土壤pH值呈负相关,说明温度对东北北部温带森林的土壤养分含量具有一定的控制作用。     

Ecosystem development in roadside grasslands: biotic control, plant-soil interactions, and dispersal limitations

Roadside grasslands undergoing secondary succession are abundant, and represent ecologically meaningful examples of novel, human-created ecosystems. Interactions between plant and soil communities (hereafter plant-soil interactions) are of major importance in understanding the role of biotic control in ecosystem functioning, but little is known about these links in the context of ecosystem restoration and succession. The assessment of the key biotic communities and interactions driving ecosystem development will help practitioners to better allocate the limited resources devoted to roadside grassland restoration. We surveyed roadside grasslands from three successional stages (0-2, 7-9, and >20 years) in two Mediterranean regions of Spain. Structural equation modeling was used to evaluate how interactions between plants, biological soil crusts (BSCs), and soil microbial functional diversity (soil microorganisms) affect indicators of ecosystem development and restoration: plant similarity to the reference ecosystem, erosion control, and soil C storage and N accumulation. Changes in plant community composition along the successional gradient exerted the strongest influence on these indicators. High BSC cover was associated with high soil stability, and high soil microbial functional diversity from late-successional stages was associated with high soil fertility. Contrary to our expectations, the indirect effects of plants, mediated by either BSCs or soil microorganisms, were very weak in both regions, suggesting a minor role for plant-soil interactions upon ecosystem development indicators over long periods. Our results suggest that natural vegetation dynamics effectively improved ecosystem development within a time frame of 20 years in the grasslands evaluated. They also indicate that this time could be shortened if management actions focus on: (1) maintaining well-conserved natural areas close to roadsides to enhance plant compositional changes towards late-successional stages, (2) increasing BSC cover in areas under strong erosion risk, to avoid soil loss, and (3) enhancing soil microbial functional diversity in resource-limited areas, to enhance soil C and N accumulation.     

Changes in microbial community characteristics and soil organic matter with nitrogen additions in two tropical forests

Microbial communities and their associated enzyme activities affect the amount and chemical quality of carbon (C) in soils. Increasing nitrogen (N) deposition, particularly in N-rich tropical forests, is likely to change the composition and behavior of microbial communities and feed back on ecosystem structure and function. This study presents a novel assessment of mechanistic links between microbial responses to N deposition and shifts in soil organic matter (SOM) quality and quantity. We used phospholipid fatty acid (PLFA) analysis and microbial enzyme assays in soils to assess microbial community responses to long-term N additions in two distinct tropical rain forests. We used soil density fractionation and 13C nuclear magnetic resonance (NMR) spectroscopy to measure related changes in SOM pool sizes and chemical quality. Microbial biomass increased in response to N fertilization in both tropical forests and corresponded to declines in pools of low-density SOM. The chemical quality of this soil C pool reflected ecosystem-specific changes in microbial community composition. In the lower-elevation forest, there was an increase in gram-negative bacteria PLFA biomass, and there were significant losses of labile C chemical groups (O-alkyls). In contrast, the upper-elevation tropical forest had an increase in fungal PLFAs with N additions and declines in C groups associated with increased soil C storage (alkyls). The dynamics of microbial enzymatic activities with N addition provided a functional link between changes in microbial community structure and SOM chemistry. Ecosystem-specific changes in microbial community composition are likely to have far-reaching effects on soil carbon storage and cycling. This study indicates that microbial communities in N-rich tropical forests can be sensitive to added N, but we can expect significant variability in how ecosystem structure and function respond to N deposition among tropical forest types.     

Nitrogen limitation of net primary productivity in terrestrial ecosystems is globally distributed

Our meta-analysis of 126 nitrogen addition experiments evaluated nitrogen (N) limitation of net primary production (NPP) in terrestrial ecosystems. We tested the hypothesis that N limitation is widespread among biomes and influenced by geography and climate. We used the response ratio (R approximately equal ANPP(N)/ANPP(ctrl)) of aboveground plant growth in fertilized to control plots and found that most ecosystems are nitrogen limited with an average 29% growth response to nitrogen (i.e., R = 1.29). The response ratio was significant within temperate forests (R = 1.19), tropical forests (R = 1.60), temperate grasslands (R = 1.53), tropical grasslands (R = 1.26), wetlands (R = 1.16), and tundra (R = 1.35), but not deserts. Eight tropical forest studies had been conducted on very young volcanic soils in Hawaii, and this subgroup was strongly N limited (R = 2.13), which resulted in a negative correlation between forest R and latitude. The degree of N limitation in the remainder of the tropical forest studies (R = 1.20) was comparable to that of temperate forests, and when the young Hawaiian subgroup was excluded, forest R did not vary with latitude. Grassland response increased with latitude, but was independent of temperature and precipitation. These results suggest that the global N and C cycles interact strongly and that geography can mediate ecosystem response to N within certain biome types.     

Nitrogen regulation of the climate-carbon feedback: evidence from a long-term global change experiment

Modeling studies have shown that nitrogen (N) strongly regulates ecosystem responses and feedback to climate warming. However, it remains unclear what mechanisms underlie N regulation of ecosystem-climate interactions. To examine N regulation of ecosystem feedback to climate change, we have conducted a warming and clipping experiment since November 1999 in a tallgrass prairie of the Great Plains, USA. Infrared heaters were used to elevate soil temperature by an average of 1.96 degrees C at a depth of 2.5 cm from 2000 to 2008. Yearly biomass clipping mimicked hay or biofuel feedstock harvest. We measured carbon (C) and N concentrations, estimated their content and C:N ratio in plant, root, litter, and soil pools. Warming significantly stimulated C storage in aboveground plant, root, and litter pools by 17%, 38%, and 29%, respectively, averaged over the nine years (all P < 0.05) but did not change soil C content or N content in any pool. Plant C:N ratio and nitrogen use efficiency increased in the warmed plots compared to the control plots, resulting primarily from increased dominance of C4 plants in the community. Clipping significantly decreased C and N storage in plant and litter pools (all P < 0.05) but did not have interactive effects with warming on either C or N pools over the nine years. Our results suggest that increased ecosystem nitrogen use efficiency via a shift in species composition toward C4 dominance rather than plant N uptake is a key mechanism underlying warming stimulation of plant biomass growth.     

Soil C and N changes with afforestation of grasslands across gradients of precipitation and plantation age

Afforestation, the conversion of unforested lands to forests, is a tool for sequestering anthropogenic carbon dioxide into plant biomass. However, in addition to altering biomass, afforestation can have substantial effects on soil organic carbon (SOC) pools, some of which have much longer turnover times than plant biomass. An increasing body of evidence suggests that the effect of afforestation on SOC may depend on mean annual precipitation (MAP). The goal of this study was to test how labile and bulk pools of SOC and total soil nitrogen (TN) change with afforestation across a rainfall gradient of 600-1500 mm in the Rio de la Plata grasslands of Argentina and Uruguay. The sites were all former grasslands planted with Eucalyptus spp. Overall, we found that afforestation increased (up to 1012 kg C x ha(-1) x yr(-1)) or decreased (as much as 1294 kg C x ha(-1) x yr(-1)) SOC pools in this region and that these changes were significantly related to MAP. Drier sites gained, and wetter sites lost, SOC and TN (r2 = 0.59, P = 0.003; and r2 = 0.57, P = 0.004, respectively). Labile C and N in microbial biomass and extractable soil pools followed similar patterns to bulk SOC and TN. Interestingly, drier sites gained more SOC and TN as plantations aged, while losses reversed as plantations aged in wet sites, suggesting that plantation age in addition to precipitation is a critical driver of changes in soil organic matter with afforestation. This new evidence implies that longer intervals between harvests for plantations could improve SOC storage, ameliorating the negative trends found in humid sites. Our results suggest that the value of afforestation as a carbon sequestration tool should be considered in the context of precipitation and age of the forest stand.     

Nitrogen retention in urban lawns and forests

Lawns are a dominant cover type in urban ecosystems, and there is concern about their impacts on water quality. However, recent watershed-level studies suggest that these pervious areas might be net sinks, rather than sources, for nitrogen (N) in the urban environment. A 15N pulse-labeling experiment was performed on lawn and forest plots in the Baltimore (Maryland, U.S.A.) metropolitan area to test the hypothesis that lawns are a net sink for atmospheric-N deposition and to compare and contrast mechanisms of N retention in these vegetation types. A pulse of 15N-NO3-, simulating a precipitation event, was followed through mineral soils, roots, Oi-layer/thatch, aboveground biomass, microbial biomass, inorganic N, and evolved N2 gas over a one-year period. The 15N label was undetectable in gaseous samples, but enrichment of other pools was high. Gross rates of production and consumption of NO3- and NH4+ were measured to assess differences in internal N cycling under lawns and forests. Rates of N retention were similar during the first five days of the experiment, with lawns showing higher N retention than forests after 10, 70, and 365 days. Lawns had larger pools of available NO3- and NH4+; however, gross rates of mineralization and nitrification were also higher, leading to no net differences in NO3- and NH4+ turnover times between the two systems. Levels of 15N remained steady in forest mineral soils from day 70 to 365 (at 23% of applied 15N), but continued to accumulate in lawn mineral soils over this same time period, increasing from 20% to 33% of applied 15N. The dominant sink for N in lawn plots changed over time. Immobilization in mineral soils dominated immediately (one day) after tracer application (42% of recovered 15N); plant biomass dominated the short term (10 days; 51%); thatch and mineral-soil pools together dominated the medium term (70 days; 28% and 36%, respectively); and the mineral-soil pool alone dominated long-term retention (one year; 70% of recovered 15N). These findings illustrate the mechanisms whereby urban and suburban lawns under low to moderate management intensities are an important sink for atmospheric-N deposition.     

三江源区不同建植年代人工草地群落演替与土壤养分变化

研究了了三源区不同建植期人工修复草地在不同演替阶段毒杂草[主要是甘肃马先蒿(Pedicularis kansuensis)]的入侵规律、数量特征,植物群落物种组成、生物苗和草地质最以及土壤养分、微生物活性的变化规律.结果表明,不同建植期人工修复草地植物群落的种类组成、植物功能群组成和群落数量特征存在显著差异.随着演替时间的推移,人工草地群落盖度、高度、物种数、生物最和多样性指数均表现出"V"字型变化规律,杂类草--甘肃马先蒿的数量特征变化尤为明显,在4 a的人工草地群落中开始局部入侵,在5～6 a的人工草地群落中大面积入侵,其入侵速度、入侵面积达到高峰期.土壤的含水量、容重、土壤中有机质、氮素和磷素在演替过程(7 a、9 a草地)中逐渐降低,到一定时期又逐步增加;随着演替的进行,不同建植期人工草地的土壤微牛物生物量碳和酶活性均呈"V"字型,变化.对于退化生态系统的恢复首先是植被恢复,其次是土壤肥力的恢复.土壤有机质等养分的积累、微生物活性的改善不仅能使土壤-植物复合系统的功能得以恢复,同时也能促进物种多样性的形成,有利于人工草地群落稳定性的提高.在试验区尽管植被恢复演替进行得比较缓慢,但从土壤发展的角度看,仍属进展演替.所以,在退化高寒草甸的恢复过程中,若降低和有效控制外界的干扰(如围栏封育),可为退化草地恢复提供繁殖体与土壤环境,实现人工草地逐步向恢复(正向)演替进行.图3表6参34     

Nitrogen uptake, distribution, turnover, and efficiency of use in a CO2-enriched sweetgum forest

The Progressive Nitrogen Limitation (PNL) hypothesis suggests that ecosystems in a CO2-enriched atmosphere will sequester C and N in long-lived biomass and soil organic pools, thereby limiting available N and constraining the continued response of net primary productivity to elevated [CO2]. Here, we present a six-year record of N dynamics of a sweetgum (Liquidambar styraciflua) stand exposed to elevated [CO2] in the free-air CO2 enrichment (FACE) experiment at Oak Ridge, Tennessee, USA. We also evaluate the concept of PNL for this ecosystem from the perspective of N uptake, content, distribution, and turnover, and N-use efficiency. Leaf N content was 11% lower on a leaf mass basis (NM) and 7% lower on a leaf area basis (NA) in CO2-enriched trees. However, there was no effect of [CO2] on total canopy N content. Resorption of N during senescence was not altered by [CO2], so NM of litter, but not total N content, was reduced. The NM of fine roots was not affected, but the total amount of N required for fine-root production increased significantly, reflecting the large stimulation of fine-root production in this stand. Hence, total N requirement of the trees was higher in elevated [CO2], and the increased requirement was met through an increase in N uptake rather than increased retranslocation of stored reserves. Increased N uptake was correlated with increased net primary productivity (NPP). N-use efficiency, however, did not change with CO2 enrichment because increased N productivity was offset by lower mean residence time of N in the trees. None of the measured responses of plant N dynamics in this ecosystem indicated the occurrence of PNL, and the stimulation of NPP by elevated [CO2] was sustained for the first six years of the experiment. Although there are some indications of developing changes in the N economy, the N supply in the soil at this site may be sufficient to meet an increasing demand for available N, especially as the roots of CO2-enriched trees explore deeper in the soil profile.     

Climatic/edaphic controls on soil carbon/nitrogen response to shrub encroachment in desert grassland.

The proliferation of woody plants in grasslands over the past 100+ years can alter carbon, nitrogen, and water cycles and influence land surface-atmosphere interactions. Although the majority of organic carbon in these ecosystems resides belowground, there is no consensus on how this change in land cover has affected soil organic carbon (SOC) and total nitrogen (TN) pools. The degree to which duration of woody plant occupation, climate, and edaphic conditions have mediated SOC and TN responses to changes in life-form composition are poorly understood. We addressed these issues at a desert grassland site in Arizona, USA, where the leguminous shrub velvet mesquite (Prosopis velutina) has proliferated along an elevation/precipitation/temperature gradient and on contrasting soil morphologic surfaces. On sandy loam complexes of mid-Holocene origin, mean SOC and TN of soils in the grassland matrix increased approximately 68% and approximately 45%, respectively, with increasing elevation. Soil organic carbon pools were comparable and TN pools were approximately 23% higher in Pleistocene-aged clay loam complexes co-occurring with Holocene-aged soils at the upper elevation/climatic zone. Across the site, belowground resources associated with large Prosopis plants were 21-154% (SOC) and 18-127% (TN) higher than those in the grassy matrix. The variance in SOC and TN pools accounted for by Prosopis stem size (a rough surrogate for time of site occupation) was highest at the low- and mid-elevation sites (69-74%) and lowest at the upper elevation site (32-38%). Soil delta15N values ranged from 5.5 per thousand to 6.7 per thousand across the soil/elevation zones but were comparable in herbaceous and shrub-impacted soils and exhibited a weak relationship with Prosopis basal stem diameter (r2 < 0.1) and TN (r2 < 0.08). The SOC delta13C values decreased linearly with increasing Prosopis basal diameter, suggesting that size and isotopic composition of the SOC pool is a function of time of Prosopis site occupation. Isotopic mixture models indicate that encroachment of C3 woody plants has also promoted SOC additions from C4 plant sources, indicative of long-term herbaceous facilitation. Grassy sites in contrasting soil/elevation combinations, initially highly distinctive in their SOC pool size and delta13C, appear to be converging on similar values following approximately 100 years of woody plant proliferation.     

Plants control the seasonal dynamics of microbial N cycling in a beech forest soil by belowground C allocation

Soil microbes in temperate forest ecosystems are able to cycle several hundreds of kilograms of N per hectare per year and are therefore of paramount importance for N retention. Belowground C allocation by trees is an important driver of seasonal microbial dynamics and may thus directly affect N transformation processes over the course of the year. Our study aimed at unraveling plant controls on soil N cycling in a temperate beech forest at a high temporal resolution over a time period of two years, by investigating the effects of tree girdling on microbial N turnover. In both years of the experiment, we discovered (1) a summer N mineralization phase (between July and August) and (2) a winter N immobilization phase (November-February). The summer mineralization phase was characterized by a high N mineralization activity, low microbial N uptake, and a subsequent high N availability in the soil. During the autumn/winter N immobilization phase, gross N mineralization rates were low, and microbial N uptake exceeded microbial N mineralization, which led to high levels of N in the microbial biomass and low N availability in the soil. The observed immobilization phase during the winter may play a crucial role for ecosystem functioning, since it could protect dissolved N that is produced by autumn litter degradation from being lost from the ecosystem during the phase when plants are mostly inactive. The difference between microbial biomass N levels in winter and spring equals 38 kg N/ha and may thus account for almost one-third of the annual plant N demand. Tree girdling strongly affected annual N cycling: the winter N immobilization phase disappeared in girdled plots (microbial N uptake and microbial biomass N were significantly reduced, while the amount of available N in the soil solution was enhanced). This was correlated to a reduced fungal abundance in autumn in girdled plots. By releasing recently fixed photosynthates to the soil, plants may thus actively control the annual microbial N cycle. Tree belowground C allocation increases N accumulation in microorganisms during the winter which may ultimately feed back on plant N availability in the following growing season.     

Interactive effects of plant species diversity and elevated CO2 on soil biota and nutrient cycling

Terrestrial ecosystems consist of mutually dependent producer and decomposer subsystems, but not much is known on how their interactions are modified by plant diversity and elevated atmospheric CO2 concentrations. Factorially manipulating grassland plant species diversity and atmospheric CO2 concentrations for five years, we tested whether high diversity or elevated CO2 sustain larger or more active soil communities, affect soil aggregation, water dynamics, or nutrient cycling, and whether plant diversity and elevated CO2 interact. Nitrogen (N) and phosphorus (P) pools, symbiotic N2 fixation, plant litter quality, soil moisture, soil physical structure, soil nematode, collembola and acari communities, soil microbial biomass and microflora community structure (phospholipid fatty acid [PLFA] profiles), soil enzyme activities, and rates of C fluxes to soils were measured. No increases in soil C fluxes or the biomass, number, or activity of soil organisms were detected at high plant diversity; soil H2O and aggregation remained unaltered. Elevated CO2 affected the ecosystem primarily by improving plant and soil water status by reducing leaf conductance, whereas changes in C cycling appeared to be of subordinate importance. Slowed-down soil drying cycles resulted in lower soil aggregation under elevated CO2. Collembola benefited from extra soil moisture under elevated CO2, whereas other faunal groups did not respond. Diversity effects and interactions with elevated CO2 may have been absent because soil responses were mainly driven by community-level processes such as rates of organic C input and water use; these drivers were not changed by plant diversity manipulations, possibly because our species diversity gradient did not extend below five species and because functional type composition remained unaltered. Our findings demonstrate that global change can affect soil aggregation, and we advocate that soil aggregation should be considered as a dynamic property that may respond to environmental changes and feed back on other ecosystem functions.     

Fate of nitrogen in riparian forest soils and trees: an 15N tracer study simulating salmon decay

We introduced an 15N-NH4+ tracer to the riparian forest of a salmon-bearing stream (Kennedy Creek, Washington, USA) to quantify the cycling and fate of a late-season pulse of salmon N and, ultimately, mechanisms regulating potential links between salmon abundance and tree growth. The 15N tracer simulated deposition of 7.25 kg of salmon (fresh) to four 50-m2 plots. We added NH4+ (the initial product of salmon carcass decay) and other important nutrients provided by carcasses (P, S, K, Mg, Ca) to soils in late October 2003, coincident with local salmon spawning. We followed the 15N tracer through soil and tree pools for one year. Biological uptake of the 15N tracer occurred quickly: 64% of the 15N tracer was bound in soil microbiota within 14 days, and roots of the dominant riparian tree, western red cedar (Thuja plicata), began to take up 15N tracer within seven days. Root uptake continued through the winter. The 15N tracer content of soil organic matter reached a maximum of approximately 52%, five weeks after the application, and a relative equilibrium of approximately 40% within five months. Six months after the addition, in spring 2004, at least 37% of the 15N tracer was found in tree tissues: approximately 23% in foliage, approximately 11% in roots, and approximately 3% in stems. Within the stems, xylem and phloem sap contained approximately 96% of the tracer N, and approximately 4% was in structural xylem N. After one year, at least 28% of the 15N tracer was still found in trees, and loss from the plots was only approximately 20%. The large portion of tracer N taken up in the fall and reallocated to leaves and stems the following spring provides mechanistic evidence for a one-year-lagged tree-growth response to salmon nutrients. Salmon nutrients have been deposited in the Kennedy Creek system each fall for centuries, but the system shows no evidence of nutrient saturation. Rates of N uptake and retention are a function of site history and disturbance and also may be the result of a legacy effect, in which annual salmon nutrient addition may lead to increased efficiency of nutrient uptake and use.     

Nutrient regulation of organic matter decomposition in a tropical rain forest

Terrestrial biosphere-atmosphere CO2 exchange is dominated by tropical forests, so understanding how nutrient availability affects carbon (C) decomposition in these ecosystems is central to predicting the global C cycle's response to environmental change. In tropical rain forests, phosphorus (P) limitation of primary production and decomposition is believed to be widespread, but direct evidence is rare. We assessed the effects of nitrogen (N) and P fertilization on litter-layer organic matter decomposition in two neighboring tropical rain forests in southwest Costa Rica that are similar in most ways, but that differ in soil P availability. The sites contain 100-200 tree species per hectare and between species foliar nutrient content is variable. To control for this heterogeneity, we decomposed leaves collected from a widespread neotropical species, Brosimum utile. Mass loss during decomposition was rapid in both forests, with B. utile leaves losing >80% of their initial mass in <300 days. High organic matter solubility throughout decomposition combined with high rainfall support a model of litter-layer decomposition in these rain forests in which rapid mass loss in the litter layer is dominated by leaching of dissolved organic matter (DOM) rather than direct CO2 mineralization. While P fertilization did not significantly affect mass loss in the litter layer, it did stimulate P immobilization in decomposing material, leading to increased P content and a lower C:P ratio in soluble DOM. In turn, increased P content of leached DOM stimulated significant increases in microbial mineralization of DOM in P-fertilized soil. These results show that, while nutrients may not affect mass loss during decomposition in nutrient-poor, wet ecosystems, they may ultimately regulate CO2 losses (and hence C storage) by limiting microbial mineralization of DOM leached from the litter layer to soil.