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.     

Responses of an old-field plant community to interacting factors of elevated [CO2], warming, and soil moisture

Aims: The direct effects of atmospheric and climatic change factors—atmospheric[CO₂], air temperature and changes in precipitation—canshape plant community composition and alter ecosystem function.It is essential to understand how these factors interact tomake better predictions about how ecosystems may respond tochange. We investigated the direct and interactive effects of[CO₂], warming and altered soil moisture in open-top chambers(OTCs) enclosing a constructed old-field community to test howthese factors shape plant communities. Materials and methods: The experimental facility in Oak Ridge, TN, USA, made use of4-m diameter OTCs and rain shelters to manipulate [CO₂] (ambient,ambient + 300 ppm), air temperature (ambient, ambient + 3.5°C)and soil moisture (wet, dry). The plant communities within thechambers comprised seven common old-field species, includinggrasses, forbs and legumes. We tracked foliar cover for eachspecies and calculated community richness, evenness and diversityfrom 2003 to 2005. Important findings: This work resulted in three main findings: (1) warming had species-specificeffects on foliar cover that varied through time and were alteredby soil moisture treatments; (2) [CO₂] had little effect onindividual species or the community; (3) diversity, evennessand richness were influenced most by soil moisture, primarilyreflecting the response of one dominant species. We concludethat individualistic species responses to atmospheric and climaticchange can alter community composition and that plant populationsand communities should be considered as part of analyses ofterrestrial ecosystem response to climate change. However, predictionof plant community responses may be difficult given interactionsbetween factors and changes in response through time.     

Genotypic variation in response of quaking aspen (Populus tremuloides) to atmospheric CO2 enrichment

Enriched atmospheric CO₂ alters the quantity and quality of plant production, but how such effects vary among plant genotypes is poorly known. We evaluated the independent and interactive effects of CO₂ and nutrient availability on growth, allocation and phytochemistry of six aspen (Populus tremuloides Michx.) genotypes. One-year-old trees, propagated from root cuttings, were grown in CO₂-controlled glasshouses for 64 days, then harvested. Foliage was analyzed for levels of water, nitrogen, starch, phenolic glycosides and condensed tannins. Of seven plant growth/allocation variables measured, four (biomass production, stem growth, relative growth rate and root:shoot ratio) exhibited marginally to highly significant CO₂ 2 genotype interactions. CO₂ enrichment stimulated growth of some genotypes more than others, and this interaction was itself influenced by soil nutrient availability. In addition, enriched CO₂ increased the magnitude of the among-genotype variance for four of the growth/allocation variables. Of six foliar chemical constituents analyzed, CO₂-mediated responses of two (the phenolic glycoside tremulacin and condensed tannins) varied among genotypes. Moreover, enriched CO₂ increased the magnitude of among-genotype variance for four of the chemical variables. Given the importance of these growth and chemical characteristics to the biological fitness of aspen, this research suggests that projected atmospheric CO₂ increases are likely to alter the genetic structures and evolutionary trajectories of aspen populations.     

Interaction of soil moisture and elevated CO2 on the above-ground growth rate, root length density and gas exchange of turves from temperate pasture

Interactions between water availability and elevated atmosphericCO₂ concentrations have the potential to be important factorsin determining future forage supply from temperate pastures.Using large turves from an established pasture, the responseof these communities at 350 or 700 l l^–1 CO₂ to a soilmoisture deficit and to recovery from the deficit in comparisonto turves that were well-watered throughout was measured. Priorto this experiment the turves had been exposed to the CO₂ treatmentsfor 324 d. Net CO₂ exchange continued at elevated CO₂ even when the volumetricsoil moisture content was less than 0.10 m³ m^–3 soil;at the same moisture deficit gas exchange at ambient CO₂ waszero. The additional carbon fixed by the elevated CO₂ turveswas primarily allocated below-ground as shown by the maintenanceof root length density at the same level as in well-wateredturves. When the dry turves were rewatered there was compensatorygrowth at ambient CO₂ so that the above-ground growth rate exceededthat of turves that had not experienced a moisture deficit.At the start of this experiment, the turves that were growingat 700 l I^–1 CO₂ had a greater proportion of legume (principallywhite clover, Trifolium repens L.) in the harvested herbage.There was a trend for the legume content at elevated CO₂ tobe reduced under a soil moisture deficit. The results indicate different strategies in response to soilmoisture deficits depending on the CO₂ concentration. At ambientCO₂, growth stopped, but plants were able to respond stronglyon rewatering; while at elevated CO₂ growth continued (particularlybelow-ground), but no additional growth was evident on rewatering.Ecosystem gas exchange measurements taken at the end of theexperiment (after 429 d of exposure to CO₂) showed 33% moreCO₂ was fixed at elevated CO₂ with only a small (12%) and nonsignificantdownward regulation. Key words: Carbon dioxide, climate change, grassland, gas exchange, soil moisture deficit     

Elevated [CO2] and nutrient status modified leaf phenology and growth rhythm of young Populus trichocarpa trees in a 3-year field study

Young individuals of a single clone of black cottonwood, in Iceland, were exposed for 3 years to elevated atmospheric CO₂ concentrations [CO₂] in whole-tree chambers at natural and high nutrient availability. No treatment effects were found at bud break or the start of shoot extension in spring. Autumn phenology was, however, affected both by elevated [CO₂] and changes in nutrient status. The time of annual growth cessation was linearly related to leaf nitrogen concentration, irrespective of CO₂ treatment. At low (natural) nutrient availability, elevated [CO₂] accelerated growth cessation and bud set, which reduced the period of active growth. An earlier and more pronounced leaf senescence and corresponding loss of photosynthetic capacity further decreased carbon acquisition in elevated [CO₂]. The negative [CO₂] effect on duration of shoot extension and leaf senescence existed, but was not as pronounced, when trees grew at higher nutrient availability. Improved nutrient availability extended the shoot extension period and delayed leaf senescence. It is suggested that trees grown in elevated [CO₂] altered their autumn phenology as an effect of a signal similar to that in trees growing at low nutrient availability, i.e. an imbalance between carbon and nitrogen sources. These alterations in autumn phenology may be important when predicting how trees will grow in a future CO₂ environment.     

Effects of CO2Enrichment on Trees and Forests: Lessons to be Learned in View of Future Ecosystem Studies

Because of their prominent role in global bioproductivity andbecause of their complex structure and function, forests andtree species deserve particular attention in studies on thelikely impact of elevated atmospheric CO₂on terrestrial vegetation.Besides a synoptic review of some of the most prominent above-groundresponse processes, particular attention is given to below-groundresponses of trees to elevated atmospheric CO₂, while some feedbackprocesses and interactions with various biotic and abiotic factorsare also briefly summarized. At the leaf level there is littleevidence of the long-term loss of sensitivity to CO₂that wassuggested by earlier experiments with tree seedlings in pots.Future studies on photosynthesis measurements will probablynot alter our conclusions about acclimation, but should focusmore on respiration under elevated CO₂, which is still poorlyunderstood. At the tree level, the increase in growth observedin elevated CO₂results from an increase in both leaf area andleaf photosynthetic rate (per unit leaf area). Tree growth enhancementis generally larger at high rates of nutrient supply; when nutrientsupply rates do not meet growth rates, tree nutrient statusdeclines and nutrients become limiting. In many studies at thecanopy level, a shift in whole-tree carbon allocation patterntowards below-ground parts has been associated with increasedatmospheric CO₂concentrations. At the ecosystem level, a largeramount of carbon being allocated below-ground could show upby either (1) more root growth and turnover, (2) enhanced activityof root-associated microorganisms, (3) larger microbial biomasspools and enhanced microbial activity, or (4) increased lossesof soil carbon through soil respiration. Fine root productionis generally enhanced, but it is not clear whether this responsewould persist in a forest. As elevated CO₂stimulates biomassproduction, litterfall and rhizodeposition also increase. Thisincreased delivery of labile organic matter to the soil couldinfluence soil microbial communities and subsequent decompositionrates, nutrient availability and carbon storage in soil. Thereare, however, contradictory hypothesis about the direction inwhich nutrient availability will be affected. Knowledge of theresponse of these and other ecophysiological processes to elevatedCO₂is the key to understanding the functioning of the wholeforest ecosystem. Our current knowledge is sufficiently largewith regard to how the carbon uptake process and individualtree growth respond under atmospheric changes, but more emphasisshould be put in future experiments on the interactions betweenvarious processes, such as the carbon and nitrogen cycles, andon below-ground responses. Copyright 1999 Annals of Botany Company Global climatic changes, elevated CO₂, forests, trees, below-ground processes, mycorrhizae, roots, decomposition.     

The growth response of plants to elevated CO2 under non-optimal environmental conditions

Under benign environmental conditions, plant growth is generally stimulated by elevated atmospheric CO₂ concentrations. When environmental conditions become sub- or supra-optimal for growth, changes in the biomass enhancement ratio (BER; total plant biomass at elevated CO₂ divided by plant biomass at the current CO₂ level) may occur. We analysed literature sources that studied CO₂2environment interactions on the growth of herbaceous species and tree seedlings during the vegetative phase. For each experiment we calculated the difference in BER for plants that were grown under 'optimal' and 'non-optimal' conditions. Assuming that interactions would be most apparent if the environmental stress strongly diminished growth, we scaled the difference in the BER values by the growth reduction due to the stress factor. In our compilation we found a large variability in CO₂2environment interactions between experiments. To test the impact of experimental design, we simulated a range of analyses with a plant-to-plant variation in size common in experimental plant populations, in combination with a number of replicates generally used in CO₂2environment studies. A similar variation in results was found as in the compilation of real experiments, showing the strong impact of stochasticity. We therefore caution against strong inferences derived from single experiments and suggest rather a reliance on average interactions across a range of experiments. Averaged over the literature data available, low soil nutrient supply or sub-optimal temperatures were found to reduce the proportional growth stimulation of elevated CO₂. In contrast, BER increased when plants were grown at low water supply, albeit relatively modestly. Reduced irradiance or high salinity caused BER to increase in some cases and decrease in others, resulting in an average interaction with elevated CO₂ that was not significant. Under high ozone concentrations, the relative growth enhancement by elevated CO₂ was strongly increased, to the extent that high CO₂ even compensated in an absolute way for the harmful effect of ozone on growth. No systematic difference in response was found between herbaceous and woody species for any of the environmental variables considered.     

Initial effects of experimental warming on above- and belowground components of net ecosystem CO2 exchange in arctic tundra

The Arctic contains extensive soil carbon reserves that could provide a substantial positive feedback to atmospheric CO₂ concentrations and global warming. Evaluation of this hypothesis requires a mechanistic understanding of the in situ responses of individual components of tundra net ecosystem CO₂ exchange (NEE) to warming. In this study, we measured NEE, total ecosystem respiration and respiration from below ground in experimentally warmed plots within Alaskan acidic tussock tundra. Soil warming of 2-4°C during a single growing season caused strong increases in total ecosystem respiration and belowground respiration from moss-dominated inter-tussock areas, and similar trends from sedge-dominated tussocks. Consequently, the overall effect of the manipulation was to substantially enhance net ecosystem carbon loss during mid-summer. Components of vascular plant biomass were closely correlated with total ecosystem respiration and belowground respiration in control plots of both microsites, but not in warmed plots. By contrast, in the warmed inter-tussock areas, belowground respiration was most closely correlated with organic-layer depth. Warming in tussock areas was associated with increased leaf nutrient pools, indicating enhanced rates of soil nutrient mineralisation. Together, these results suggest that warming enhanced net ecosystem CO₂ efflux primarily by stimulating decomposition of soil organic matter, rather than by increasing plant-associated respiration. Our short-term experiment provides field evidence to support previous growth chamber and modelling studies indicating that arctic soil C reserves are relatively sensitive to warming and could supply an initial positive feedback to rising atmospheric CO₂ concentrations/changing climate.     

Mechanisms of Phosphorus Acquisition for Ponderosa Pine Seedlings under High CO2and Temperature

To test the hypothesis that elevated atmospheric CO₂and elevatedtemperature, simulating current and predicted future growingseason conditions, act antagonistically on phosphorus acquisitionof ponderosa pine, seedlings were grown in controlled-environmentchambers in a two temperature (25/10 °C and 30/15 °C)xtwoCO₂(350 and 700 µl^-1) experimental design. Mycorrhizalseedlings were watered daily with a nutrient solution with Padded in organic form as inositol hexaphosphate (64 ppm P).Thus seedlings were challenged to use active forms of P acquisition.Elevated CO₂increased the relative growth rate by approx. 5%which resulted in an approx. 33% increase in biomass after 4months. There was no main effect of temperature on growth. Increasedgrowth under elevated CO₂and temperature was supported by increasesin specific absorption rate and the specific utilization rateof P. The contribution of mycorrhizae to P uptake may have beengreater under simulated future conditions, as elevated CO₂increasedthe number of mycorrhizal roots. There was no main effect oftemperature on root phosphatase activity, but elevated CO₂causeda decrease in activity. The inverse pattern of root phosphataseactivity and mycorrhizal infection across treatments suggestsa physiological coordination between these avenues of P acquisition.The concentration of oxalate in the soil increased under elevatedCO₂and decreased under elevated temperature. This small molecularweight acid solubilizes inorganic P making it available foruptake. Increased mycorrhizal infection and exudation of oxalateincreased P uptake in ponderosa pine seedlings under elevatedCO₂, and there was no net negative effect of increased temperature.The increased carbon status of pine under elevated CO₂may facilitateuptake of limiting P in native ecosystems. Atmospheric CO₂; climate change; growth analysis; oxalate; Pinus ponderosa ; ponderosa pine; phosphorus uptake; rhizosphere; root phosphatase; temperature     

Responses of soil microorganisms to resource availability in urban, desert soils

Terrestrial desert ecosystems are strongly structured by the distribution of plants, which concentrate resources and create islands of fertility relative to interplant spaces. Atmospheric nitrogen (N) deposition resulting from urbanization has the potential to change those spatial patterns via resource inputs, resulting in more homogeneous soil resource availability. We sampled soils at 12 desert remnant sites around Phoenix, Arizona along a model-predicted gradient in N deposition to determine the degree to which deposition has altered spatial patterns in soil resource availability and microbial activity. Soil microbial biomass and abundance were not influenced by atmospheric N deposition. Instead, plant islands remained strong organizers of soil microbial processes. These islands of fertility exhibited elevated pools of resources, microbial abundance, and activity relative to interspaces. In both plant islands and interspaces, soil moisture and soil N concentrations predicted microbial biomass and abundance. Following experimental wetting, carbon dioxide (CO₂) flux from soil of interspaces was positively correlated with N deposition, whereas in plant islands, soil CO₂ flux was positively correlated with soil moisture content and soil organic matter. Soil CO₂ flux in both patch types showed rapid and short-lived responses to precipitation, demonstrating the brief time scales during which soil biota may process deposited materials. Although we observed patterns consistent with N limitation of microbes in interspaces, we conclude that atmospheric N deposition likely accumulates in soils because microbes are primarily limited by water and secondarily by carbon or nitrogen. Soil microbial uptake of atmospherically deposited N likely occurs only during sparse and infrequent rainfall.     

Effects of climate on soil phosphorus cycle and availability in natural terrestrial ecosystems

Climate is predicted to change over the 21st century. However, little is known about how climate change can affect soil phosphorus (P) cycle and availability in global terrestrial ecosystems, where P is a key limiting nutrient. With a global database of Hedley P fractions and key‐associated physiochemical properties of 760 (seminatural) natural soils compiled from 96 published studies, this study evaluated how climate pattern affected soil P cycle and availability in global terrestrial ecosystems. Overall, soil available P, indexed by Hedley labile inorganic P fraction, significantly decreased with increasing mean annual temperature (MAT) and precipitation (MAP). Hypothesis‐oriented path model analysis suggests that MAT negatively affected soil available P mainly by decreasing soil organic P and primary mineral P and increasing soil sand content. MAP negatively affected soil available P both directly and indirectly through decreasing soil primary mineral P; however, these negative effects were offset by the positive effects of MAP on soil organic P and fine soil particles, resulting in a relatively minor total MAP effect on soil available P. As aridity degree was mainly determined by MAP, aridity also had a relatively minor total effect on soil available P. These global patterns generally hold true irrespective of soil depth (≤10 cm or >10 cm) or site aridity index (≤1.0 or >1.0), and were also true for the low‐sand (≤50%) soils. In contrast, available P of the high‐sand (>50%) soils was positively affected by MAT and aridity and negatively affected by MAP. Our results suggest that temperature and precipitation have contrasting effects on soil P availability and can interact with soil particle size to control soil P availability.     

大气CO2浓度升高对土壤碳库稳定性的影响

工业革命以来,大气CO₂浓度持续上升,升高的CO₂浓度会改变植物光合产物积累、土壤碳库的碳输入和碳输出过程,进而通过影响有机碳组成和周转特征来调控土壤碳库动态变化。土壤碳库是陆地生态系统碳库的重要组成部分,其碳储量的微小变化都会对大气CO₂浓度和气候变化产生巨大影响。但目前关于CO₂浓度升高对土壤碳库动态和稳定性的影响还不清楚,很大程度上限制了预测陆地生态系统碳循环对气候变化的反馈。系统综述国内外大气CO₂浓度升高对植被生产力、植被碳输入和土壤碳库影响的研究进展,旨在揭示土壤碳库物理、化学组成以及周转特征对CO₂浓度升高的响应过程和机理,探讨CO₂升高情境下土壤微生物特征对土壤碳库稳定性的影响和驱动机制,为深入理解全球变化下的土壤碳循环特征提供理论支撑。     

Effects of CO2 enrichment and intraspecific competition on biomass partitioning, nitrogen content and microbial biomass carbon in soil of perennial ryegrass and white clover

Seedlings of perennial ryegrass (Lolium perenne L. cv. Parcour)and white clover (Trifolium repens L. cv. Karina) grown at fivedifferent plant densities were exposed to ambient (390 ppm)and elevated (690 ppm) CO₂ concentrations. After 43 d the effectsof CO₂ enrichment and plant density on growth of shoot and root,nitrogen concentration of tissue, and microbial biomass carbon(C_mic) in soil were determined. CO₂ enrichment of Lolium perenneincreased shoot growth on average by 17% independent of plantdensity, while effects on root biomass ranged between -4% and+ 107% due to an interaction with plant density. Since tilernumber per plant was unaffected by elevated CO₂, the small responseof shoot growth to CO₂ enrichment was atributed to low sinkstrength. A significant correlation between nitrogen concentrationof total plant biomass and root fraction of total plant drymatter, which was not changed by CO₂ enrichment, indicates thatnitrogen status of the plant controls biomass partitioning andthe effect of CO₂ enrichment on root growth. Effects of elevatedCO₂ and plant density on shoot and root growth of Trifoliumrepens were not significantly interacting and mean CO₂-relatedincrease amounted to 29% and 66%, respectively. However, growthenhancement due to elevated CO₂ was strongest when leaf areaindex was lowest. Total amounts of nitrogen in shoots and rootswere bigger at 690 ppm than at 390 ppm CO₂. There was a significantincrease in C_mic in experiments with both species whereas plantdensity had no substantial effect. Key words: CO₂ enrichment, intraspecific competition, biomass partitioning, Lolium perenne, Trifolium repens, grassland     

Saturation response of enhanced vegetation productivity attributes to intricate interactions

Evidence for the multifaceted responses of terrestrial ecosystems has been shown by the weakening of CO₂ fertilization-induced and warming-controlled productivity gains. The intricate relationship between vegetation productivity and various environmental controls still remains elusive spatially. Here several inherent preponderances make China a natural experimental setting to investigate the interaction and relative contributions of five drivers to gross primary productivity for the period from 1982 to 2018 (i.e., elevated atmospheric CO₂ concentrations, climate change, nutrient availability, anthropogenic land use change, and soil moisture) by coupling multiple long-term datasets. Despite a strikingly prominent enhancement of vegetation productivity in China, it exhibits similar saturation responses to the aforementioned environmental drivers (elevated CO₂, climatic factors, and soil moisture). The CO₂ fertilization-dominated network explains the long-term variations in vegetation productivity in humid regions, but its effect is clearly attenuated or even absent in arid and alpine environments controlled by climate and soil moisture. Divergence in interactions also provides distinct evidence that water availability plays an essential role in limiting the potential effects of climate change and elevated CO₂ concentrations on vegetation productivity. Unprecedented industrialization and dramatic surface changes may have breached critical thresholds of terrestrial ecosystems under the diverse natural environment and thus forced a shift from a period dominated by CO₂ fertilization to a period with nonlinear interactions. These findings suggest that future benefits in terrestrial ecosystems are likely to be counteracted by uncertainties in the complicated network, implying an increasing reliance on human societies to combat potential risks. Our results therefore highlight the need to account for the intricate interactions globally and thus incorporate them into mitigation and adaptation policies.     

Experimental warming had little effect on carbon-based secondary compounds, carbon and nitrogen in selected alpine plants and lichens

Global warming is expected to change plant defence through its influence on plant primary resources. Increased temperature (T) will increase photosynthesis, and thus carbon (C) availability, but may also increase soil mineralization and availability of nitrogen (N). More access to C and N is expected to mainly increase plant growth, and, according to hypotheses on resource based defence, this could lower plant concentrations of carbon-based secondary compounds (CBSCs).We used two already established warming experiment with open top chambers (OTCs) and control plots in alpine south-western Norway, one on a ridge (8 years’ treatment) and a one in a leeside (3 years’ treatment), to study the effects of warming on plant and lichen defensive compound concentrations. The study included five vascular plant and six lichen species.One vascular plant species had lower concentration of CBSCs under elevated T, while the others did not respond to the treatment. In lichens there were no effects of warming on CBSCs, but a tendency to reduced total C concentrations. However, there were effects of warming on nitrogen, as the concentration decreased inside OTCs for three species, while it increased for one lichen species. Lichens generally had higher CBSC and total C concentrations on the ridge than in the leeside, but no such pattern were seen for vascular plants.No elevated temperature effect on CBCSs is most probably a result of high constitutive defence under the limiting alpine conditions, suggesting that chemical defence is little subject to change under climate warming, at least on a short-term basis. We suggest that the driving forces of plant defence in the arctic-alpine should be tested individually under controlled conditions, and suggest that competition from other plants may be a greater threat under climate warming than increased herbivory or disease attacks.     

Root system adjustments: regulation of plant nutrient uptake and growth responses to elevated CO2

Nutrients such as nitrogen (N) and phosphorus (P) often limit plant growth rate and production in natural and agricultural ecosystems. Limited availability of these nutrients is also a major factor influencing long-term plant and ecosystem responses to rising atmospheric CO₂ levels, i.e., the commonly observed short-term increase in plant biomass may not be sustained over the long-term. Therefore, it is critical to obtain a mechanistic understanding of whether elevated CO₂ can elicit compensatory adjustments such that acquisition capacity for minerals increases in concert with carbon (C) uptake. Compensatory adjustments such as increases in (a) root mycorrhizal infection, (b) root-to-shoot ratio and changes in root morphology and architecture, (c) root nutrient absorption capacity, and (d) nutrient-use efficiency can enable plants to meet an increased nutrient demand under high CO₂. Here we examine the literature to assess the extent to which these mechanisms have been shown to respond to high CO₂. The literature survey reveals no consistent pattern either in direction or magnitude of responses of these mechanisms to high CO₂. This apparent lack of a pattern may represent variations in experimental protocol and/or interspecific differences. We found that in addressing nutrient uptake responses to high CO₂ most investigators have examined these mechanisms in isolation. Because such mechanisms can potentially counterbalance one another, a more reliable prediction of elevated CO₂ responses requires experimental designs that integrate all mechanisms simultaneously. Finally, we present a functional balance (FB) model as an example of how root system adjustments and nitrogen-use efficiency can be integrated to assess growth responses to high CO₂. The FB model suggests that the mechanisms of increased N uptake highlighted here have different weights in determining overall plant responses to high CO₂. For example, while changes in root-to-shoot biomass allocation, r, have a small effect on growth, adjustments in uptake rate per unit root mass, [`(n)]\bar \nu , and photosynthetic N use efficiency, p*, have a significantly greater leverage on growth responses to elevated CO₂ except when relative growth rate (RGR) reaches its developmental limit, maximum RGR (RGR_max).     

气候变化对作物矿质元素利用率影响研究进展

作物矿质元素利用率对气候变化的响应是目前全球变化研究中既重要、又复杂,且认知最少的科学领域。这个科学问题的研究关系到解密或预测陆地植物及农作物矿质胁迫对全球气候变化响应的机理,为将来农业投入提供理论依据,是应对气候变化的当务之急。目前只有少数研究,通过模拟试验,探索性地开展了CO_2浓度或温度升高的环境条件下,矿质元素在土壤-植物系统迁移、分布和储存特征的研究。从相关的文献报道来看,CO_2浓度升高环境条件下,小麦和水稻作物籽粒中大量和痕量元素的富集水平一般呈下降趋势。但温度升高情况下,作物各器官对对矿质元素的吸收情况则更为复杂。正由于气候因素与植物矿质元素利用率之间关系的复杂性,在气候变化背景下,解密作物矿质胁迫对全球气候变化响应的科学问题,尚需改进试验方法、手段,从土壤性质、作物生态生理,以及农业生态系统中矿质元素在土壤-作物系统中迁移转化的过程,全面考察作物矿质元素利用率对气候变化的响应机理。     

Reduction of transpiration and altered nutrient allocation contribute to nutrient decline of crops grown in elevated CO2 concentrations

Plants grown in elevated [CO₂] have lower protein and mineral concentrations compared with plants grown in ambient [CO₂]. Dilution by enhanced production of carbohydrates is a likely cause, but it cannot explain all of the reductions. Two proposed, but untested, hypotheses are that (1) reduced canopy transpiration reduces mass flow of nutrients to the roots thus reducing nutrient uptake and (2) changes in metabolite or enzyme concentrations caused by physiological changes alter requirements for minerals as protein cofactors or in other organic complexes, shifting allocation between tissues and possibly altering uptake. Here, we use the meta‐analysis of previous studies in crops to test these hypotheses. Nutrients acquired mostly by mass flow were decreased significantly more by elevated [CO₂] than nutrients acquired by diffusion to the roots through the soil, supporting the first hypothesis. Similarly, Mg showed large concentration declines in leaves and wheat stems, but smaller decreases in other tissues. Because chlorophyll requires a large fraction of total plant Mg, and chlorophyll concentration is reduced by growth in elevated [CO₂], this supports the second hypothesis. Understanding these mechanisms may guide efforts to improve nutrient content, and allow modeling of nutrient changes and health impacts under future climate change scenarios.     

Temporal and Nutritional Influences on the Response to Elevated CO2 in Selected British Grasses

To investigate the duration of the CO₂ response and its interactionwith mineral nutrition, CO₂-enrichment experiments were performedon four British grasses of differing ecology and functionaltype: Arrhenatherum elatius (L.) Beauv., Festuca ovina L., Festucarubra L. and Poa annua L. Naturally-lit, glasshouse cabinetswere used, with a non-limiting water supply and a daytime meantemperature of 18 °C. Two CO₂ treatments were maintainedat nominal concentrations of 350 and 700 vpm and were combinedfactorially with two levels of balanced mineral nutrition atconductivities of 0·1 and 1 mS cm^-1. Harvests took placeat planting-out, and at 16, 37 and 58 d thereafter. Fitted curves were used to derive instantaneous values of totaldry weight, relative growth rate (RGR), shoot weight fraction(SWF) and unit shoot rate (USR) for all combinations of species,CO₂ level, nutrient level and time of harvesting. At the higher nutrient level there was a reasonably close agreementwith previous estimates of the CO₂ response in the four species.The response, if any, most often arose from an increase in USRbeing accompanied by a less than proportionate decline in SWF.Responses were sustained throughout the period studied. At thelower nutrient level, all species showed a CO₂ response initially,but this declined at a rate which was inversely related to theCO₂-responsiveness of the species at the higher nutrient level. The underlying ontogenetic drift appeared to be markedly towardsadjustment in SWF and away from that of USR. However, this driftwas retarded, suspended or even reversed by low-nutrient conditionsand/or by high CO₂ responsiveness in the species itself.Copyright1995, 1999 Academic Press Climate change, CO₂ enrichment, plant strategies, mineral nutrition, growth analysis, relative growth rate, shoot weight fraction, unit shoot rate, functional equilibria     

Links between soil microbial communities and plant traits in a species‐rich grassland under long‐term climate change

Climate change can influence soil microorganisms directly by altering their growth and activity but also indirectly via effects on the vegetation, which modifies the availability of resources. Direct impacts of climate change on soil microorganisms can occur rapidly, whereas indirect effects mediated by shifts in plant community composition are not immediately apparent and likely to increase over time. We used molecular fingerprinting of bacterial and fungal communities in the soil to investigate the effects of 17 years of temperature and rainfall manipulations in a species‐rich grassland near Buxton, UK. We compared shifts in microbial community structure to changes in plant species composition and key plant traits across 78 microsites within plots subjected to winter heating, rainfall supplementation, or summer drought. We observed marked shifts in soil fungal and bacterial community structure in response to chronic summer drought. Importantly, although dominant microbial taxa were largely unaffected by drought, there were substantial changes in the abundances of subordinate fungal and bacterial taxa. In contrast to short‐term studies that report high resistance of soil fungi to drought, we observed substantial losses of fungal taxa in the summer drought treatments. There was moderate concordance between soil microbial communities and plant species composition within microsites. Vector fitting of community‐weighted mean plant traits to ordinations of soil bacterial and fungal communities showed that shifts in soil microbial community structure were related to plant traits representing the quality of resources available to soil microorganisms: the construction cost of leaf material, foliar carbon‐to‐nitrogen ratios, and leaf dry matter content. Thus, our study provides evidence that climate change could affect soil microbial communities indirectly via changes in plant inputs and highlights the importance of considering long‐term climate change effects, especially in nutrient‐poor systems with slow‐growing vegetation.