Effects of elevated [CO2] at the community level mediated by root symbionts

This review examines the effects of elevated [CO₂] on plant symbioses with mycorrhizal fungi and root nodule bacteria, with emphasis on community and ecosystem processes. The effects of elevated [CO₂] on the relationships between single plant species and root symbionts are considered first. There is some evidence that plant infection by and/or biomass of root symbionts are stimulated by elevated [CO₂], but growth enhancement of the host seemingly depends on its degree of dependence on symbiosis and on soil nutrient availability. Second, the effects of elevated [CO₂] on the relationships between plant multispecies assemblages and soil, and likely impacts on above-ground and belowground diversity, are analysed. Experimental and modelling work have suggested the existence of complex feedbacks in the responses of plants and the rhizosphere to CO₂ enrichment. By modifying C inputs from plants to soil, elevated [CO₂] may affect the biomass, the infectivity, and the species/isolate composition of root symbionts. This has the potential to alter community structure and ecosystem functioning. Finally, the incorporation of type and degree of symbiotic dependence into the definition of plant functional types, and into experimental work within the context of global change research, are discussed. More experimental work on the effects of elevated [CO₂] at the community/ecosystem level, explicitly considering the role of root symbioses, is urgently needed.     

The nitrogen budget of a pine forest under free air CO2 enrichment

Elevated concentrations of atmospheric CO₂ increase plant biomass, net primary production (NPP) and plant demand for nitrogen (N). The demand for N set by rapid plant growth under elevated CO₂ could be met by increasing soil N availability or by greater efficiency of N uptake. Alternatively, plants could increase their nitrogen-use efficiency (NUE), thereby maintaining high rates of growth and NPP in the face of nutrient limitation. We quantified dry matter and N budgets for a young pine forest exposed to 4 years of elevated CO₂ using free-air CO₂ enrichment technology. We addressed three questions: Does elevated CO₂ increase forest NPP and the demand for N by vegetation? Is demand for N met by greater uptake from soils, a shift in the distribution of N between plants, microbes, and soils, or increases in NUE under elevated CO₂? Will soil N availability constrain the NPP response of this forest as CO₂ fumigation continues? A step-function increase in atmospheric CO₂ significantly increased NPP during the first 4 years of this study. Significant increases in NUE under elevated CO₂ modulated the average annual requirement for N by vegetation in the first and third growing seasons under elevated CO₂; the average stimulation of NPP in these years was 21% whereas the average annual stimulation of the N requirement was only 6%. In the second and fourth growing seasons, increases in NPP increased the annual requirement for N by 27-33%. Increases in the annual requirement for N were largely met by increases in N uptake from soils. Retranslocation of nutrients prior to senescence played only a minor role in supplying the additional N required by trees growing under elevated CO₂. NPP was highly correlated with between-plot variation in the annual rate of net N mineralization and CO₂ treatment. This demonstrates that NPP is co-limited by C availability, as CO₂ from the atmosphere, and N availability from soils. There is no evidence that soil N mineralization rates have increased under elevated CO₂. The correlation between NPP and N mineralization rates and the increase in the annual requirement for N in certain years imply that soil N availability may control the long-term productivity response of this ecosystem to elevated CO₂. Although we have no evidence suggesting that NPP is declining in response to >4 years of CO₂ fumigation, if the annual requirement of N continues to be stimulated by elevated CO₂, we predict that the productivity response of this forest ecosystem will decline over time.     

Modeling the belowground response of plants and soil biota to edaphic and climatic change—What can we expect to gain?

As atmospheric CO₂ concentrations continue to increase, so too will the emphasis placed on understanding the belowground response of plants to edaphic and climatic change. Controlled-exposure studies that address the significance of an increased supply of carbon to roots and soil biota, and the consequences of this to nutrient cycling will play a prominent role in this process. Models will also contribute to understanding the response of plants and ecosystems to changes in the earth's climate by incorporating experimental results into conceptual or quantitative frameworks from which potential feedbacks within the plant-soil system can be identified. Here we present five examples of how models can be used in this analysis and how they can contribute to the development of new hypotheses in the areas of root biology, soil biota, and ecosystem processes. Two examples illustrate the role of coarse and fine roots in nitrogen and phosphorus uptake from soils, the respiratory costs associated with this acquisition of nutrients, and the significance of root architecture in these relationships. Another example focuses on a conceptual model that has helped raise new ideas about the effects of elevated CO₂ on root and microbial biomass, and on nutrient dynamics in the rhizosphere. Difficulties associated with modeling the contribution of mycorrhizal fungi to whole-plant growth are also discussed. Finally, several broad-scale models are used to illustrate the importance of root turnover, litter decomposition, and nitrogen mineralization in determining an ecosystem's response to atmospheric CO₂ enrichment. We conclude that models are appropriate tools for use both in guiding existing studies and in identifying new hypotheses for future research. Development of models that address the complexities of belowground processes and their role in determining plant and ecosystem function within the context of rising CO₂ concentrations and associated climate change should be encouraged.     

Nitrogen cycling in microcosms of yellow birch exposed to elevated CO2: simultaneous positive and negative below-ground feedbacks

This study investigated simultaneous plant and soil feedbacks on growth enhancement with elevated [CO₂] within microcosms of yellow birch (Betula alleghaniensis Britt.) in the second year of growth. Understanding the integrated responses of model ecosystems may provide key insight into the potential net nutrient feedbacks on [CO₂] growth enhancements in temperate forests. We measured the net biomass production, C:N ratios, root architecture, and mycorrhizal responses of yellow birch, in situ rates gross nitrogen mineralization and the partitioning of available NH₄⁺ between yellow birch and soil microbes. Elevated atmospheric [CO₂] resulted in significant alterations in the cycling of N within the microcosms. Plant C/N ratios were significantly increased, gross mineralization and NH₄⁺ consumption rates were decreased, and relative microbial uptake of NH₄⁺ was increased, representing a suite of N cycling negative feedbacks on N availability. However, increased C/N ratios may also be a mechanism which allows plants to maintain higher growth with a constant or reduced N supply. Total plant N content was increased with elevated [CO₂], suggesting that yellow birch had successfully increased their ability to acquire nutrients during the first year of growth. However, plant uptake rates of NH₄⁺ had decreased in the second year. This discrepancy implies that, in this study, nitrogen uptake showed a trend through ontogeny of decreasing enhancement under elevated [CO₂]. The reduced N mineralization and relatively increased N immobilization are a potential feedback which may drive this ontogenetic trend. This study has demonstrated the importance of using an integrated approach to exploring potential nutrient-cycling feedbacks in elevated [CO₂].     

Influence of rhizodeposition under elevated CO2 on plant nutrition and soil organic matter

Atmospheric CO₂ concentrations can influence ecosystem carbon storage through net primary production (NPP), soil carbon storage, or both. In assessing the potential for carbon storage in terrestrial ecosystems under elevated CO₂, both NPP and processing of soil organic matter (SOM), as well as the multiple links between them, must be examined. Within this context, both the quantity and quality of carbon flux from roots to soil are important, since roots produce specialized compounds that enhance nutrient acquisition (affecting NPP), and since the flux of organic compounds from roots to soil fuels soil microbial activity (affecting processing of SOM).From the perspective of root physiology, a technique is described which uses genetically engineered bacteria to detect the distribution and amount of flux of particular compounds from single roots to non-sterile soils. Other experiments from several labs are noted which explore effects of elevated CO₂ on root acid phosphatase, phosphomonoesterase, and citrate production, all associated with phosphorus nutrition. From a soil perspective, effects of elevated CO₂ on the processing of SOM developed under a C4 grassland but planted with C3 California grassland species were examined under low (unamended) and high (amended with 20 g m^–2 NPK) nutrients; measurements of soil atmosphere 13C combined with soil respiration rates show that during vegetative growth in February, elevated CO₂ decreased respiration of carbon from C4 SOM in high nutrient soils but not in unamended soils.This emphasis on the impacts of carbon loss from roots on both NPP and SOM processing will be essential to understanding terrestrial ecosystem carbon storage under changing atmospheric CO₂ concentrations.Abbreviations SOM soil organic matter - NPP net primary productivity - NEP net ecosystem productivity - PNPP p-nitrophenyl phosphate     

Elevated carbon dioxide and ozone alter productivity and ecosystem carbon content in northern temperate forests

Three young northern temperate forest communities in the north‐central United States were exposed to factorial combinations of elevated carbon dioxide (CO₂) and tropospheric ozone (O₃) for 11 years. Here, we report results from an extensive sampling of plant biomass and soil conducted at the conclusion of the experiment that enabled us to estimate ecosystem carbon (C) content and cumulative net primary productivity (NPP). Elevated CO₂ enhanced ecosystem C content by 11%, whereas elevated O₃ decreased ecosystem C content by 9%. There was little variation in treatment effects on C content across communities and no meaningful interactions between CO₂ and O₃. Treatment effects on ecosystem C content resulted primarily from changes in the near‐surface mineral soil and tree C, particularly differences in woody tissues. Excluding the mineral soil, cumulative NPP was a strong predictor of ecosystem C content (r² = 0.96). Elevated CO₂ enhanced cumulative NPP by 39%, a consequence of a 28% increase in canopy nitrogen (N) content (g N m^?2) and a 28% increase in N productivity (NPP/canopy N). In contrast, elevated O₃ lowered NPP by 10% because of a 21% decrease in canopy N, but did not impact N productivity. Consequently, as the marginal impact of canopy N on NPP (?NPP/?N) decreased through time with further canopy development, the O₃ effect on NPP dissipated. Within the mineral soil, there was less C in the top 0.1 m of soil under elevated O₃ and less soil C from 0.1 to 0.2 m in depth under elevated CO₂. Overall, these results suggest that elevated CO₂ may create a sustained increase in NPP, whereas the long‐term effect of elevated O₃ on NPP will be smaller than expected. However, changes in soil C are not well‐understood and limit our ability to predict changes in ecosystem C content.     

甘南地区二氧化碳施肥效应对生态系统的影响

陆地生态系统是全球第二大碳库,其碳收支一直是气候变化研究的热点领域,而研究二氧化碳（CO₂）施肥效应又是全球变化碳循环领域较为关注的前沿部分。CO₂与生态系统关系复杂,当前仍无法厘清CO₂对陆地生态系统碳循环的影响作用。基于太阳辐射数据、气温数据及归一化植被指数数据等,利用光能利用率遥感模型,模拟2019年甘南地区的碳循环,选取三个指标,即GPP （陆地生态系统总初级生产力）、NPP （净初级生产力）和NEP （净生态系统生产力）来分析甘南地区植被固碳的时空变化特征及CO₂施肥效应。结果表明：（1）甘南地区2019年植被固碳总量约为2611 tC。甘南地区生态系统GPP、NPP和NEP季节性特征明显,其值均在夏季达到最高;而在空间上,GPP、NPP表现为东高西低的特征,NEP呈现出北高南低的分布特征。（2） CO₂对GPP、NPP存在正向的施肥效应,分别增加了14.4%和14.3%;而对NEP具有负向反馈效应,使其减少了0.3%,并且CO₂对NEP的影响整体也表现为北高南低的特征。研究揭示出：虽然CO₂在提升GPP和NPP时,正向的施肥效应明显,但是对甘南地区的NEP,即固碳量来说,CO₂的影响却很有限。因此在研究CO₂施肥效应时不应一概而论,生态地理环境对其的影响不可忽视。研究可以为揭示陆地生态系统碳循环的动态机制提供一定的理论依据。     

Long‐term dynamics of mycorrhizal root tips in a loblolly pine forest grown with free‐air CO2 enrichment and soil N fertilization for 6 years

Large‐scale, long‐term FACE (Free‐Air CO₂ enrichment) experiments indicate that increases in atmospheric CO₂ concentrations will influence forest C cycling in unpredictable ways. It has been recently suggested that responses of mycorrhizal fungi could determine whether forest net primary productivity (NPP) is increased by elevated CO₂ over long time periods and if forests soils will function as sources or sinks of C in the future. We studied the dynamic responses of ectomycorrhizae to N fertilization and atmospheric CO₂ enrichment at the Duke FACE experiment using minirhizotrons over a 6 year period (2005–2010). Stimulation of mycorrhizal production by elevated CO₂ was observed during only 1 (2007) of 6 years. This increased the standing crop of mycorrhizal tips during 2007 and 2008; during 2008, significantly higher mortality returned standing crop to ambient levels for the remainder of the experiment. It is therefore unlikely that increased production of mycorrhizal tips can explain the lack of progressive nitrogen limitations and associated increases in N uptake observed in CO₂‐enriched plots at this site. Fertilization generally decreased tree reliance on mycorrhizae as tip production declined with the addition of nitrogen as has been shown in many other studies. Annual NPP of mycorrhizal tips was greatest during years with warm January temperatures and during years with cool spring temperatures. A 2 °C increase in average late spring temperatures (May and June) decreased annual production of mycorrhizal root tip length by 50%. This has important implications for ecosystem function in a warmer world in addition to potential for forest soils to sequester atmospheric C.     

Progressive N limitation of plant response to elevated CO2: a microbiological perspective

A major uncertainty in predicting long-term ecosystem C balance is whether stimulation of net primary production will be sustained in future atmospheric CO₂ scenarios. Immobilization of nutrients (N in particular) in plant biomass and soil organic matter (SOM) provides negative feedbacks to plant growth and may lead to progressive N limitation (PNL) of plant response to CO₂ enrichment. Soil microbes mediate N availability to plants by controlling litter decomposition and N transformations as well as dominating biological N fixation. CO₂-induced changes in C inputs, plant nutrient demand and water use efficiency often have interactive and contrasting effects on microbes and microbially mediated N processes. One critical question is whether CO₂-induced N accumulation in plant biomass and SOM will result in N limitation of microbes and subsequently cause them to obtain N from alternative sources or to alter the ecosystem N balance. We reviewed the experimental results that examined elevated CO₂ effects on microbial parameters, focusing on those published since 2000. These results in general show that increased C inputs dominate the CO₂ impact on microbes, microbial activities and their subsequent controls over ecosystem N dynamics, potentially enhancing microbial N acquisition and ecosystem N retention. We reason that microbial mediation of N availability for plants under future CO₂ scenarios will strongly depend on the initial ecosystem N status, and the nature and magnitude of external N inputs. Consequently, microbial processes that exert critical controls over long-term N availability for plants would be ecosystem-specific. The challenge remains to quantify CO₂-induced changes in these processes, and to extrapolate the results from short-term studies with step-up CO₂ increases to native ecosystems that are already experiencing gradual changes in the CO₂ concentration.     

Effects of plant species diversity, atmospheric [CO2], and N addition on gross rates of inorganic N release from soil organic matter

A significant challenge in predicting terrestrial ecosystem response to global changes comes from the relatively poor understanding of the processes that control pools and fluxes of plant nutrients in soil. In addition, individual global changes are often studied in isolation, despite the potential for interactive effects among them on ecosystem processes. We studied the response of gross N mineralization and microbial respiration after 6 years of application of three global change factors in a grassland field experiment in central Minnesota (the BioCON experiment). BioCON is a factorial manipulation of plant species diversity (1, 4, 9 and 16 prairie species), atmospheric [CO₂] (ambient and elevated: 560 μmol mol^?1), and N inputs (ambient and ambient +4 g N m^?2 yr^?1). We hypothesized that gross N mineralization would increase with increasing levels of all factors because of stimulated plant productivity and thus greater organic inputs to soils. However, we also hypothesized that N addition would enhance, while elevated [CO₂] and greater diversity would temper, gross N mineralization responses because of increased and reduced plant tissue N concentrations, respectively. In partial support of our hypothesis, gross N mineralization increased with greater diversity and N addition, but not with elevated [CO₂]. The ratio of gross N mineralization to microbial respiration (i.e. the ‘yield’ of inorganic N mineralized per unit C respired) declined with greater diversity and [CO₂] suggesting increasing limitation of microbial processes by N relative to C in these treatments. Based on these results, we conclude that the plant supply of organic matter primarily controls gross N mineralization and microbial respiration, but that the concentration of N in organic matter input secondarily influences these processes. Thus, in systems where N limits plant productivity these global change factors could cause different long‐term ecosystem trajectories because of divergent effects on soil N and C cycling.     

A synthesis: The role of nutrients as constraints on carbon balances in boreal and arctic regions

As in many ecosystems, carbon (C) cycling in arctic and boreal regions is tightly linked to the cycling of nutrients: nutrients (particularly nitrogen) are mineralized through the process of organic matter decomposition (C mineralization), and nutrient availability strongly constrains ecosystem C gain through primary production. This link between C and nutrient cycles has implications for how northern systems will respond to future climate warming and whether feedbacks to rising concentrations of atmospheric CO₂ from these regions will be positive or negative. Warming is expected to cause a substantial release of C to the atmosphere because of increased decomposition of the large amounts of organic C present in high-latitude soils (a positive feedback to climate warming). However, increased nutrient mineralization associated with this decomposition is expected to stimulate primary production and ecosystem C gain, offsetting or even exceeding C lost through decomposition (a negative feedback to climate warming). Increased primary production with warming is consistent with results of numerous experiments showing increased plant growth with nutrient enrichment. Here we examine key assumptions behind this scenario: (1) temperature is a primary control of decomposition in northern regions, (2) increased decomposition and associated nutrient release are tightly coupled to plant nutrient uptake, and (3) short-term manipulations of temperature and nutrient availability accurately predict long-term responses to climate change.     

Modeled responses of terrestrial ecosystems to elevated atmospheric CO2: a comparison of simulations by the biogeochemistry models of the Vegetation/Ecosystem Modeling and Analysis Project (VEMAP)

Although there is a great deal of information concerning responses to increases in atmospheric CO₂ at the tissue and plant levels, there are substantially fewer studies that have investigated ecosystem-level responses in the context of integrated carbon, water, and nutrient cycles. Because our understanding of ecosystem responses to elevated CO₂ is incomplete, modeling is a tool that can be used to investigate the role of plant and soil interactions in the response of terrestrial ecosystems to elevated CO₂. In this study, we analyze the responses of net primary production (NPP) to doubled CO₂ from 355 to 710 ppmv among three biogeochemistry models in the Vegetation/Ecosystem Modeling and Analysis Project (VEMAP): BIOME-BGC (BioGeochemical Cycles), Century, and the Terrestrial Ecosystem Model (TEM). For the conterminous United States, doubled atmospheric CO₂ causes NPP to increase by 5% in Century, 8% in TEM, and 11% in BIOME-BGC. Multiple regression analyses between the NPP response to doubled CO₂ and the mean annual temperature and annual precipitation of biomes or grid cells indicate that there are negative relationships between precipitation and the response of NPP to doubled CO₂ for all three models. In contrast, there are different relationships between temperature and the response of NPP to doubled CO₂ for the three models: there is a negative relationship in the responses of BIOME-BGC, no relationship in the responses of Century, and a positive relationship in the responses of TEM. In BIOME-BGC, the NPP response to doubled CO₂ is controlled by the change in transpiration associated with reduced leaf conductance to water vapor. This change affects soil water, then leaf area development and, finally, NPP. In Century, the response of NPP to doubled CO₂ is controlled by changes in decomposition rates associated with increased soil moisture that results from reduced evapotranspiration. This change affects nitrogen availability for plants, which influences NPP. In TEM, the NPP response to doubled CO₂ is controlled by increased carboxylation which is modified by canopy conductance and the degree to which nitrogen constraints cause down-regulation of photosynthesis. The implementation of these different mechanisms has consequences for the spatial pattern of NPP responses, and represents, in part, conceptual uncertainty about controls over NPP responses. Progress in reducing these uncertainties requires research focused at the ecosystem level to understand how interactions between the carbon, nitrogen, and water cycles influence the response of NPP to elevated atmospheric CO₂. Received: 13 December 1996 / Accepted: 20 November 1997     

The impact of elevated CO2, increased nitrogen availability and biodiversity on plant tissue quality and decomposition

Elevated CO₂, increased nitrogen (N) deposition and increasing species richness can increase net primary productivity (NPP). However, unless there are comparable changes in decomposition, increases in productivity will most likely be unsustainable. Without comparable increases in decomposition nutrients would accumulate in dead organic matter leading to nutrient limitations that could eventually prohibit additional increases in productivity. To address this issue, we measured aboveground plant and litter quality and belowground root quality, as well as decomposition of aboveground litter for one and 2‐year periods using in situ litterbags in response to a three‐way factorial manipulation of CO₂ (ambient vs. 560 ppm), N deposition (ambient vs. the addition of 4 g N m⁻² yr⁻¹) and plant species richness (one, four, nine and 16 species) in experimental grassland plots. Litter chemistry responded to the CO₂, N and plant diversity treatments, but decomposition was much less responsive. Elevated CO₂ induced decreases in % N and % lignin in plant tissues. N addition led to increases in % N and decreases in % lignin. Increasing plant diversity led to decreases in % N and % lignin and an increase in % cellulose. In contrast to the litter chemistry changes, elevated CO₂ had a much lower impact on decomposition and resulted in only a 2.5% decrease in carbon (C) loss. Detectable responses were not observed either to N addition or to species richness. These results suggest that global change factors such as biodiversity loss, elevated CO₂ and N deposition lead to significant changes in tissue quality; however, the response of decomposition is modest. Thus, the observed increases in productivity at higher diversity levels and with elevated CO₂ and N fertilization are not matched by an increase in decomposition rates. This lack of coupled responses between production and decomposition is likely to result in an accumulation of nutrients in the litter pool which will dampen the response of NPP to these factors over time.     

Elevated CO2 increases nitrogen fixation and decreases soil nitrogen mineralization in Florida scrub oak

We report changes in nitrogen cycling in Florida scrub oak in response to elevated atmospheric CO₂ during the first 14 months of experimental treatment. Elevated CO₂ stimulated above-ground growth, nitrogen mass, and root nodule production of the nitrogen-fixing vine, Galactia elliottii Nuttall. During this period, elevated CO₂ reduced rates of gross nitrogen mineralization in soil, and resulted in lower recovery of nitrate on resin lysimeters. Elevated CO₂ did not alter nitrogen in the soil microbial biomass, but increased the specific rate of ammonium immobilization (NH₄⁺ immobilized per unit microbial N) measured over a 24-h period. Increased carbon input to soil through greater root growth combined with a decrease in the quality of that carbon in elevated CO₂ best explains these changes. These results demonstrate that atmospheric CO₂ concentration influences both the internal cycling of nitrogen (mineralization, immobilization, and nitrification) as well as the processes that regulate total ecosystem nitrogen mass (nitrogen fixation and nitrate leaching) in Florida coastal scrub oak. If these changes in nitrogen cycling are sustained, they could cause long-term feedbacks to the growth responses of plants to elevated CO₂. Greater nitrogen fixation and reduced leaching could stimulate nitrogen-limited plant growth by increasing the mass of labile nitrogen in the ecosystem. By contrast, reduced nitrogen mineralization and increased immobilization will restrict the supply rate of plant-available nitrogen, potentially reducing plant growth. Thus, the net feedback to plant growth will depend on the balance of these effects through time.     

Species control variation in litter decomposition in a pine forest exposed to elevated CO2

Net primary production and the flux of dry matter and nutrients from vegetation to soils has increased following four years of exposure to elevated CO₂ in a southern pine forest in NC, USA. This has increased the demand for nutrients to support enhanced rates of NPP and altered the conditions for litter decomposition on the forest floor. We quantified the chemistry and decomposition dynamics of leaf litter produced by five of the most abundant tree species in this ecosystem during the third and fourth growing seasons under elevated CO₂. The objectives of this study were to determine (i) if there were systemic or species‐specific changes in leaf litter chemistry associated with a sustained enhancement of plant growth under elevated CO₂; and (ii) whether the process of litter decomposition was altered by increased inputs of energy and nutrients to the forest floor in the plots under elevated CO₂. Leaf litter chemistry, including various C fractions and N concentration, was virtually unchanged by elevated CO₂. With few exceptions, plant litter produced under elevated CO₂ lost mass or N at the same relative rate as that produced under ambient CO₂. The relationship between initial litter chemistry and decomposition was not altered by elevated CO₂. The greater forest floor mass and nutrient content in the plots under elevated CO₂ had no consistent or long‐term effect on litter decomposition. Thus, we found no evidence that plant and microbial processes under elevated CO₂ resulted in systemic changes in mass loss or N dynamics during decomposition. In contrast to the limited effects of elevated CO₂ on litter chemistry and decomposition, there were large differences among species in initial litter chemistry, mass loss and N dynamics during decomposition. If the species composition of this forest community is altered by elevated CO₂, the indirect effect of a change in species composition will exert greater control over the long‐term rate of nutrient cycling than the direct effect of elevated CO₂ on litter chemistry and decomposition dynamics alone.     

Interactions among mycorrhizae, atmospheric CO2 and soil N impact plant community composition

We examined plant community responses to interactions between arbuscular mycorrhizal (AM) fungi and availability of atmospheric CO₂ and soil N. Communities of 14 plant species were grown in mesocosms containing living or killed AM fungal inoculum, ambient or elevated atmospheric CO₂ and low or enriched soil N. After one growing season, significantly different plant communities existed in the different treatments. Plant species richness was lowest in +N mesocosms and highest in +AM + CO₂ mesocosms. At ambient CO₂, AM fungi reduced richness but at elevated CO₂ they increased it. This was caused by changes in mortality rates of several C₃ forbs and may suggest that CO₂ enrichment ameliorates the carbon cost of some AM symbioses. Soil moisture was higher in +CO₂ mesocosms but +AM counteracted this effect. These results suggest that AM symbioses may be important mediators of plant community responses to anthropogenic CO₂ and N enrichment.     

Soil-related Constraints to the Carbon Dioxide Fertilization Effect

The significance of CO₂ fertilization in increasing net primary production (NPP) and of the underpinning mechanisms is widely recognized. This article focuses on soil resources, particularly availability of plant nutrients (e.g., nitrogen, phosphorus) and water affecting the NPP under enhanced levels of atmospheric CO₂. The review of the literature indicates the critical role of N in enhancing NPP at higher than ambient CO₂ concentration. Elevated CO₂ increases the total N uptake by 3–33% and induces a negative feedback in soil N dynamics. Decrease in tissue N concentration by 10–33%, and increase in the recalcitrant C- fractions in plant biomass results in progressive decline in soil N availability over time and necessitates supplemental application of N. In addition, higher quantity of soil available P is required. Despite a reduction in stomatal conductance and transpiration by about 20–27% at leaf level, these benefits are offset at the ecosystem level by higher water losses through soil evaporation, runoff and a rise in leaf temperature due to physiological forcing. A three-way interaction between atmospheric CO₂, soil nutrients and water availability may be the decisive factor to harnessing the benefits of CO₂ fertilization effect.     

Effects of inoculation of EPS-producing Pantoea agglomerans on wheat rhizosphere aggregation

We investigated plant and soil nitrogen pools and soil processes in monospecific stands of the C₃ sedge Scirpus olneyi and the C₄ grass Spartina patens grown in the field in open top chambers in a brackish marsh on the Chesapeake Bay. Stands of S. olneyi responded to eight years of elevated CO₂, by increased rates of net ecosystem gas exchange and a large stimulation of net ecosystem production. We conducted our study in the summer of 1994 and 1995 when soil cores were collected and aboveground biomass was estimated. Nitrogen concentration in elevated CO₂ treatments was reduced 15% in stems of S. olneyi and 8% in the upper 10 cm of the soil profile. While total plant nitrogen per unit of land area remained the same between treatments, total soil nitrogen showed a non-significant tendency to decrease in the upper 10 cm of the soil profile in elevated CO₂ both years of study. A significant decrease in soil bulk density largely contributed to the observed decrease in soil nitrogen. Exchangeable nitrogen and potential denitrification rates were also reduced in elevated CO₂, but net nitrogen mineralization was unchanged by elevated CO₂ treatment in S. olneyi both years. Plants and soils in a pure stand of the C₄ grass, S. patens, showed none of these effects of elevated CO₂ treatment. Our data provides evidence of changes in nitrogen dynamics of an ecosystem exposed to elevated CO₂ for eight years; however due to the variability in these data, we cannot say if or how these changes are likely to impact the effect of rising CO₂ on primary production or carbon accumulation in this ecosystem in the future.     

Diets of giants: the nutritional value of sauropod diet during the Mesozoic

A major uncertainty in estimating energy budgets and population densities of extinct animals is the carrying capacity of their ecosystems, constrained by net primary productivity (NPP) and its digestible energy content. The hypothesis that increases in NPP due to elevated atmospheric CO₂ contributed to the unparalleled size of the sauropods has recently been rejected, based on modern studies on herbivorous insects that imply a general, negative correlation of diet quality and increasing CO₂. However, the nutritional value of plants grown under elevated CO₂ levels might be very different for vertebrate megaherbivores than for insects. Here we show plant species‐specific responses in metabolizable energy and nitrogen content, equivalent to a two‐fold variation in daily food intake estimates for a typical sauropod, for dinosaur food plant analogues grown under CO₂ concentrations spanning estimates for Mesozoic atmospheric concentrations. Our results potentially rebut the hypothesis that constraints on sauropod diet quality were driven by Mesozoic CO₂ concentration.     

Effects of elevated [CO2] on forest growth and carbon storage: a modelling analysis of the consequences of changes in litter quality/quantity and root exudation

Many researchers have proposed that the stimulus of plant growth under elevated [CO₂] observed in short-term experiments will be moderated in the longer term by a reduction in soil nitrogen (N) availability linked to decreased litter quality and/or increased litter production. However, these negative feedbacks may be offset to some extent by a stimulus in N fixation linked to increased root exudation. The aim of this modelling study is to examine how changes in litter quality/quantity and root exudation –- if they occur –- will affect the CO₂ responses of net primary productivity and ecosystem carbon (C) storage on different timescales. We apply a model of C and N cycling in forest ecosystems (G’DAY) to stands of Norway spruce (Picea abies, L. Cast) growing at a N-limited experimental site at Flakaliden, Sweden, and draw the following conclusions: (1) in the absence of changes in litter quality and root exudation, the short-term CO₂ stimulus of litter quantity leads to only a minimal CO₂ stimulus of productivity or C storage in the medium term (≈ 20 years) and long term (≈ 200 years), because of constraints on soil N availability; (2) increasing plant nitrogen use efficiency (via a decrease in the N:C ratio of new litter) makes little impact on these results; (3) a significant CO₂ response in the medium term requires a substantial decrease in the N:C ratio of older litter, when it is approaching stabilisation as soil organic matter, although the long-term CO₂ response remains small; and (4) an increase in N fixation leads to a small effect on productivity in the short term, but a very large effect on both productivity and C storage in the long term. These results suggest that soil N constraints on the long-term CO₂-fertilisation effect can be overcome to a significant extent only by increases in N acquisition, although only modest increases may be required. This revised version was published online in June 2006 with corrections to the Cover Date.