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.     

Progressive N limitation in forests: review and implications for long-term responses to elevated CO2

Field studies have shown that elevated CO2 can cause increased forest growth over the short term (<6 years) even in the face of N limitation. This is facilitated to some degree by greater biomass production per unit N uptake (lower tissue N concentrations), but more often than not, N uptake is increased with elevated CO2 as well. Some studies also show that N sequestration in the forest floor is increased with elevated CO2. These findings raise the questions of where the "extra" N comes from and how long such growth increases can continue without being truncated by progressive N limitation (PNL). This paper reviews some of the early nutrient cycling literature that describes PNL during forest stand development and attempts to use this information, along with recent developments in soil N research, to put the issue of PNL with elevated CO2 into perspective. Some of the early studies indicated that trees can effectively "mine" N from soils over the long term, and more recent developments in soil N cycling research suggest mechanisms by which this might have occurred. However, both the early nutrient cycling literature and more recent simulation modeling suggest that PNL will at some point truncate the observed increases in growth and nutrient uptake with elevated CO2, unless external inputs of N are increased by either N fixation or atmospheric deposition.     

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.     

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.     

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.     

Long-term CO2 enrichment of a forest ecosystem: implications for forest regeneration and succession.

The composition and successional status of a forest affect carbon storage and net ecosystem productivity, yet it remains unclear whether elevated atmospheric carbon dioxide (CO2) will impact rates and trajectories of forest succession. We examined how CO2 enrichment (+200 microL CO2/L air differential) affects forest succession through growth and survivorship of tree seedlings, as part of the Duke Forest free-air CO2 enrichment (FACE) experiment in North Carolina, USA. We planted 2352 seedlings of 14 species in the low light forest understory and determined effects of elevated CO2 on individual plant growth, survival, and total sample biomass accumulation, an integrator of plant growth and survivorship over time, for six years. We used a hierarchical Bayes framework to accommodate the uncertainty associated with the availability of light and the variability in growth among individual plants. We found that most species did not exhibit strong responses to CO2. Ulmus alata (+21%), Quercus alba (+9.5%), and nitrogen-fixing Robinia pseudoacacia (+230%) exhibited greater mean annual relative growth rates under elevated CO2 than under ambient conditions. The effects of CO2 were small relative to variability within populations; however, some species grew better under low light conditions when exposed to elevated CO2 than they did under ambient conditions. These species include shade-intolerant Liriodendron tulipifera and Liquidambar styraciflua, intermediate-tolerant Quercus velutina, and shade-tolerant Acer barbatum, A. rubrum, Prunus serotina, Ulmus alata, and Cercis canadensis. Contrary to our expectation, shade-intolerant trees did not survive better with CO2 enrichment, and population-scale responses to CO2 were influenced by survival probabilities in low light. CO2 enrichment did not increase rates of sample biomass accumulation for most species, but it did stimulate biomass growth of shade-tolerant taxa, particularly Acer barbatum and Ulmus alata. Our data suggest a small CO2 fertilization effect on tree productivity, and the possibility of reduced carbon accumulation rates relative to today's forests due to changes in species composition.     

Litterfall 15N abundance indicates declining soil nitrogen availability in a free-air CO2 enrichment experiment

Forest productivity increases in response to carbon dioxide (CO2) enrichment of the atmosphere. However, in nitrogen-limited ecosystems, increased productivity may cause a decline in soil nitrogen (N) availability and induce a negative feedback on further enhancement of forest production. In a free-air CO2 enrichment (FACE) experiment, the response of sweetgum (Liquidambar styraciflua L.) productivity to elevated CO2 concentrations [CO2] has declined over time, but documenting an associated change in soil N availability has been difficult. Here we assess the time history of soil N availability through analysis of natural 15N abundance in archived samples of freshly fallen leaf litterfall. Litterfall delta15N declined from 1998 to 2005, and the rate of decline was significantly faster in elevated [CO2]. Declining leaf litterfall delta15N is indicative of a tighter ecosystem N cycle and more limited soil N availability. By integrating N availability over time and throughout the soil profile, temporal dynamics in leaf litterfall delta15N provide a powerful tool for documenting changes in N availability and the critical feedbacks between C and N cycles that will control forest response to elevated atmospheric CO2 concentrations.     

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.     

Nitrogen controls on ecosystem carbon sequestration: a model implementation and application to Saskatchewan,Canada

A plant–soil nitrogen (N) cycling model was developed and incorporated into the Integrated BIosphere Simulator (IBIS) of Foley et al. [Foley, J.A., Prentice, I.C., Ramankutty, N., Levis, S., Pollard, D., Sitch, S., Haxeltine, A., 1996. An integrated biosphere model of land surface process, terrestrial carbon balance and vegetation dynamics. Global Biogeochem. Cycles 10, 603–628]. In the N-model, soil mineral N regulates ecosystem carbon (C) fluxes and ecosystem C:N ratios. Net primary productivity (NPP) is controlled by feedbacks from both leaf C:N and soil mineral N. Leaf C:N determines the foliar and canopy photosynthesis rates, while soil mineral N determines the N availability for plant growth and the efficiency of biomass construction. Nitrogen controls on the decomposition of soil organic matter (SOM) are implemented through N immobilization and mineralization separately. The model allows greater SOM mineralization at lower mineral N, and conversely, allows greater N immobilization at higher mineral N. The model's seasonal and inter-annual behaviours are demonstrated. A regional simulation for Saskatchewan, Canada, was performed for the period 1851–2000 at a 10 km × 10 km resolution. Simulated NPP was compared with high-resolution (1 km × 1 km) NPP estimated from remote sensing data using the boreal ecosystem productivity simulator (BEPS) [Liu, J., Chen, J.M., Cihlar, J., Park, W.M., 1997. A process-based boreal ecosystem productivity simulator using remote sensing inputs. Remote Sens. Environ. 44, 81–87]. The agreement between IBIS and BEPS, particularly in NPP spatial variation, was considerably improved when the N controls were introduced into IBIS.     

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.     

大气CO2体积分数升高对植物N素吸收的影响

从影响植物N素吸收的因素来看，大气CO2体积分数升高条件下植物净光合作用增强，碳同化产物增多，利于改善N素吸收的能量和物质基础：植物根系生长增强，生物量增多且空间分布加大，有利于N素吸收；但土壤有效N供应能力的变化存在增强和减弱两种观点。从植物N素吸收的实际情况来看，大气CO2体积分数升高条件下植物N吸收总量并末增加，植物体内N质量分数普遍降低，某些种类植物N吸收形态也发生了改变。因此要阐明大气CO2体积分数升高对植物N素吸收的影响机制，必须探明土壤有效N供应能力的变化：CO2体积分数升高条件下N矿化作用是否增强，微生物和植物间是否存在对有效N的竞争，此外，CO2体积分数升高条件下植物根系形态特征变化和N素吸收(包括主动和被动吸收)的生理机制及其与环境因素的关系也值得进一步研究。     

Habitat type determines herbivory controls over CO2 fluxes in a warmer Arctic

High-latitude ecosystems store large amounts of carbon (C); however, the C storage of these ecosystems is under threat from both climate warming and increased levels of herbivory. In this study we examined the combined role of herbivores and climate warming as drivers of CO2 fluxes in two typical high-latitude habitats (mesic heath and wet meadow). We hypothesized that both herbivory and climate warming would reduce the C sink strength of Arctic tundra through their combined effects on plant biomass and gross ecosystem photosynthesis and on decomposition rates and the abiotic environment. To test this hypothesis we employed experimental warming (via International Tundra Experiment [ITEX] chambers) and grazing (via captive Barnacle Geese) in a three-year factorial field experiment. Ecosystem CO2 fluxes (net ecosystem exchange of CO2, ecosystem respiration, and gross ecosystem photosynthesis) were measured in all treatments at varying intensity over the three growing seasons to capture the impact of the treatments on a range of temporal scales (diurnal, seasonal, and interannual). Grazing and warming treatments had markedly different effects on CO2 fluxes in the two tundra habitats. Grazing caused a strong reduction in CO2 assimilation in the wet meadow, while warming reduced CO2 efflux from the mesic heath. Treatment effects on net ecosystem exchange largely derived from the modification of gross ecosystem photosynthesis rather than ecosystem respiration. In this study we have demonstrated that on the habitat scale, grazing by geese is a strong driver of net ecosystem exchange of CO2, with the potential to reduce the CO2 sink strength of Arctic ecosystems. Our results highlight that the large reduction in plant biomass due to goose grazing in the Arctic noted in several studies can alter the C balance of wet tundra ecosystems. We conclude that herbivory will modulate direct climate warming responses of Arctic tundra with implications for the ecosystem C balance; however, the magnitude and direction of the response will be habitat-specific.     

Calcium constrains plant control over forest ecosystem nitrogen cycling

Forest ecosystem nitrogen (N) cycling is a critical controller of the ability of forests to prevent the movement of reactive N to receiving waters and the atmosphere and to sequester elevated levels of atmospheric carbon dioxide (CO2). Here we show that calcium (Ca) constrains the ability of northern hardwood forest trees to control the availability and loss of nitrogen. We evaluated soil N-cycling response to Ca additions in the presence and absence of plants and observed that when plants were present, Ca additions "tightened" the ecosystem N cycle, with decreases in inorganic N levels, potential net N mineralization rates, microbial biomass N content, and denitrification potential. In the absence of plants, Ca additions induced marked increases in nitrification (the key process controlling ecosystem N losses) and inorganic N levels. The observed "tightening" of the N cycle when Ca was added in the presence of plants suggests that the capacity of forests to absorb elevated levels of atmospheric N and CO2 is fundamentally constrained by base cations, which have been depleted in many areas of the globe by acid rain and forest harvesting.     

Potential nitrogen constraints on soil carbon sequestration under low and elevated atmospheric CO2

The interaction between nitrogen cycling and carbon sequestration is critical in predicting the consequences of anthropogenic increases in atmospheric CO2 (hereafter, Ca). The progressive N limitation (PNL) theory predicts that carbon sequestration in plants and soils with rising Ca may be constrained by the availability of nitrogen in many ecosystems. Here we report on the interaction between C and N dynamics during a four-year field experiment in which an intact C3/C4 grassland was exposed to a gradient in Ca from 200 to 560 micromol/mol. There were strong species effects on decomposition dynamics, with C loss positively correlated and N mineralization negatively correlated with Ca for litter of the C3 forb Solanum dimidiatum, whereas decomposition of litter from the C4 grass Bothriochloa ischaemum was unresponsive to Ca. Both soil microbial biomass and soil respiration rates exhibited a nonlinear response to Ca, reaching a maximum at approximately 440 micromol/mol Ca. We found a general movement of N out of soil organic matter and into aboveground plant biomass with increased Ca. Within soils we found evidence of C loss from recalcitrant soil C fractions with narrow C:N ratios to more labile soil fractions with broader C:N ratios, potentially due to decreases in N availability. The observed reallocation of N from soil to plants over the last three years of the experiment supports the PNL theory that reductions in N availability with rising Ca could initially be overcome by a transfer of N from low C:N ratio fractions to those with higher C:N ratios. Although the transfer of N allowed plant production to increase with increasing Ca, there was no net soil C sequestration at elevated Ca, presumably because relatively stable C is being decomposed to meet microbial and plant N requirements. Ultimately, if the C gained by increased plant production is rapidly lost through decomposition, the shift in N from older soil organic matter to rapidly decomposing plant tissue may limit net C sequestration with increased plant production.     

大气CO2升高对土壤碳循环影响的研究进展

土壤有机碳是陆地碳库的重要组成部分,其积累和分解的变化直接影响全球的碳平衡。据估计,全球土壤（表层1m）有机碳积累总量相当于大气中碳总量的2～3倍。土壤是温室气体的源或汇,土壤碳库的变化将影响大气C02的浓度,因此,土壤碳库对人类活动的响应也是全球碳循环和全球变化研究的热点。在全球变化的大背景下,大气CO2升高导致植被生态系统碳平衡的改变进而对土壤碳循环产生影响。总结了陆地生态系统碳循环对大气C02浓度升高响应的主要生物学机制及过程,简述了大气C02浓度升高对影响土壤碳输入和输出的各因素的研究进展,并指出未来研究的主要方向。在大气C02浓度升高条件下,陆地生态系统碳循环的变化主要反映在以下几个方面：1）不同类型植物群落的净初级生产力（NPP）显著增加,但湿地植物的净初级生产力也有可能降低;2）光合产物向根系分配的数量增加,地上／地下生物量降低,根系形态发生变化,根系周转速率和根系分泌等过程的碳流量提高;3）植物含氮量降低,C／N提高,次生代谢产物增加,微生物生长受到抑制,植物残体分解速率降低;4）土壤呼吸速率显著增加,提高幅度受植物类型与土壤状况的影响;5）进入土壤的植物残体及分泌物的数量和性质影响土壤酶的活性,脱氢酶和转化酶活性增加,酚氧化酶和纤维素酶受植物类型与环境条件的影响;6）土壤中真菌的数量的增加幅度要高于细菌;7）CH4释放量增加,在植物的生长期表现更为明显。由于陆地生态系统碳循环的复杂性,研究结果仍有很大的不确定性。大气C02浓度升高与全球变化的其它表现间的交互作用将是今后研究的重点,同时由于土壤碳循环是一个由微生物介导的生物地球化学循环过程,因此,加强陆地生态系统碳循环的微生物机制研究也将为全面理解碳循环的过程提供更加准确的研究理论基础。     

Effect of elevated CO2 on coarse-root biomass in Florida scrub detected by ground-penetrating radar

Growth and distribution of coarse roots in time and space represent a gap in our understanding of belowground ecology. Large roots may play a critical role in carbon sequestration belowground. Using ground-penetrating radar (GPR), we quantified coarse-root biomass from an open-top chamber experiment in a scrub-oak ecosystem at Kennedy Space Center, Florida, USA. GPR propagates electromagnetic waves directly into the soil and reflects a portion of the energy when a buried object is contacted. In our study, we utilized a 1500 MHz antenna to establish correlations between GPR signals and root biomass. A significant relationship was found between GPR signal reflectance and biomass (R2 = 0.68). This correlation was applied to multiple GPR scans taken from each open-top chamber (elevated and ambient CO2). Our results showed that plots receiving elevated CO2 had significantly (P = 0.049) greater coarse-root biomass compared to ambient plots, suggesting that coarse roots may play a large role in carbon sequestration in scrub-oak ecosystems. This nondestructive method holds much promise for rapid and repeatable quantification of coarse roots, which are currently the most elusive aspect of long-term belowground studies.     

Growth, yield and photosynthesis of Panicum maximum and Stylosanthes hamata under elevated CO2

Plant height, biomass production, assimilatory functions and chlorophyll accumulation of Panicum maximum and Stylosanthes hamata in intercropping systems was influenced significantly under elevated CO2 (600 +/- 50 ppm) in open top chambers (OTCs). The plant height increased by 32.0 and 49.0% over the control in P. maximum and S. hamata respectively in intercropping system under elevated CO2 over open field grown crops (Ca). P. maximum and S. hamata produced 67 and 85% higher fresh and dry biomass respectively under elevated CO2. Rates of photosynthesis and stomatal conductance increased in both the crop species in intercropping systems under elevated CO2. The canopy photosynthesis (photosynthesis x leaf area index) of these crop species increased significantly under elevated CO2 over the open grown crops. The chlorophyll a and b accumulation were also higher in the leaves of both the crop species as grown in OTC with elevated CO2. The increased chlorophyll content, leaf area index and canopy photosynthesis led to higher growth and biomass production in these crop species under elevated CO2. The total carbon sequestration in crop biomass and soils during the three years was 21.53 Mg C/ha under elevated CO2. The data revealed that P. maximum and S. hamata intercropping system is the potential as a sink for the increasing level of CO2 in the atmosphere in the semi-arid tropics.     

Gastropod herbivory in response to elevated CO2 and N addition impacts plant community composition

In this study, the influence of elevated carbon dioxide (CO2) and nitrogen (N) deposition on gastropod herbivory was investigated for six annual species in a California annual grassland community. These experimentally simulated global changes increased availability of important resources for plant growth, leading to the hypothesis that species with the most positive growth and foliar nutrient responses would experience the greatest increase in herbivory. Counter to the expectations, shifts in tissue N and growth rates caused by N deposition did not predict shifts in herbivore consumption rates. N deposition increased seedling N concentrations and growth rates but did not increase herbivore consumption overall, or for any individual species. Elevated CO2 did not influence growth rates nor have a statistically significant influence on seedling N concentrations. Elevated CO2 at ambient N levels caused a decline in the number of seedlings consumed, but the interaction between CO2 and N addition differed among species. The results of this study indicate that shifting patterns of herbivory will likely influence species composition as environmental conditions change in the future; however, a simple trade-off between shifting growth rates and palatability is not evident.     

Temperature influences carbon accumulation in moist tropical forests

Evergreen broad-leaved tropical forests can have high rates of productivity and large accumulations of carbon in plant biomass and soils. They can therefore play an important role in the global carbon cycle, influencing atmospheric CO2 concentrations if climate warms. We applied meta-analyses to published data to evaluate the apparent effects of temperature on carbon fluxes and storages in mature, moist tropical evergreen forest ecosystems. Among forests, litter production, tree growth, and belowground carbon allocation all increased significantly with site mean annual temperature (MAT); total net primary productivity (NPP) increased by an estimated 0.2-0.7 Mg C x ha(-1) x yr(-1) x degrees C(-1). Temperature had no discernible effect on the turnover rate of aboveground forest biomass, which averaged 0.014 yr(-1) among sites. Consistent with these findings, forest biomass increased with site MAT at a rate of 5-13 Mg C x ha(-1) x degrees C(-1). Despite greater productivity in warmer forests, soil organic matter accumulations decreased with site MAT, with a slope of -8 Mg C x ha(-1) x degrees C(-1), indicating that decomposition rates of soil organic matter increased with MAT faster than did rates of NPP. Turnover rates of surface litter also increased with temperature among forests. We found no detectable effect of temperature on total carbon storage among moist-tropical evergreen forests, but rather a shift in ecosystem structure, from low-biomass forests with relatively large accumulations of detritus in cooler sites, to large-biomass forests with relatively smaller detrital stocks in warmer locations. These results imply that, in a warmer climate, conservation of forest biomass will be critical to the maintenance of carbon stocks in moist tropical forests.     

Modeling patterns of nonlinearity in ecosystem responses to temperature, CO2, and precipitation changes.

It is commonly acknowledged that ecosystem responses to global climate change are nonlinear. However, patterns of the nonlinearity have not been well characterized on ecosystem carbon and water processes. We used a terrestrial ecosystem (TECO) model to examine nonlinear patterns of ecosystem responses to changes in temperature, CO2, and precipitation individually or in combination. The TECO model was calibrated against experimental data obtained from a grassland ecosystem in the central United States and ran for 100 years with gradual change at 252 different scenarios. We primarily used the 100th-year results to explore nonlinearity of ecosystem responses. Variables examined in this study are net primary production (NPP), heterotrophic respiration (R(h)), net ecosystem carbon exchange (NEE), runoff, and evapotranspiration (ET). Our modeling results show that nonlinear patterns were parabolic, asymptotic, and threshold-like in response to temperature, CO2, and precipitation anomalies, respectively, for NPP, NEE, and R(h). Runoff and ET exhibited threshold-like pattern in response to both temperature and precipitation anomalies but were less sensitive to CO2 changes. Ecosystem responses to combined temperature, CO2, and precipitation anomalies differed considerably from the responses to individual factors in terms of response patterns and/or critical points of nonlinearity. Our results suggest that nonlinear patterns in response to multiple global-change factors were diverse and were considerably affected by combined climate anomalies on ecosystem carbon and water processes. The diverse response patterns in nonlinearity have profound implications for both experimental design and theoretical development.