Diffusional conductances to CO2 as a target for increasing photosynthesis and photosynthetic water-use efficiency

A key objective for sustainable agriculture and forestry is to breed plants with both high carbon gain and water-use efficiency (WUE). At the level of leaf physiology, this implies increasing net photosynthesis (A _N) relative to stomatal conductance (g _s). Here, we review evidence for CO₂ diffusional constraints on photosynthesis and WUE. Analyzing past observations for an extensive pool of crop and wild plant species that vary widely in mesophyll conductance to CO₂ (g _m), g _s, and foliage A _N, it was shown that both g _s and g _m limit A _N, although the relative importance of each of the two conductances depends on species and conditions. Based on Fick’s law of diffusion, intrinsic WUE (the ratio A _N/g _s) should correlate on the ratio g _m/g _s, and not g _m itself. Such a correlation is indeed often observed in the data. However, since besides diffusion A _N also depends on photosynthetic capacity (i.e., V _c,max), this relationship is not always sustained. It was shown that only in a very few cases, genotype selection has resulted in simultaneous increases of both A _N and WUE. In fact, such a response has never been observed in genetically modified plants specifically engineered for either reduced g _s or enhanced g _m. Although increasing g _m alone would result in increasing photosynthesis, and potentially increasing WUE, in practice, higher WUE seems to be only achieved when there are no parallel changes in g _s. We conclude that for simultaneous improvement of A _N and WUE, genetic manipulation of g _m should avoid parallel changes in g _s, and we suggest that the appropriate trait for selection for enhanced WUE is increased g _m/g _s.     

Seasonal Response of Grasslands to Climate Change on the Tibetan Plateau

Background

Monitoring vegetation dynamics and their responses to climate change has been the subject of considerable research. This paper aims to detect change trends in grassland activity on the Tibetan Plateau between 1982 and 2006 and relate these to changes in climate.

Methodology/Principal Findings

Grassland activity was analyzed by evaluating remotely sensed Normalized Difference Vegetation Index (NDVI) data collected at 15-day intervals between 1982 and 2006. The timings of vegetation stages (start of green-up, beginning of the growing season, plant maturity, start of senescence and end of the growing season) were assessed using the NDVI ratio method. Mean NDVI values were determined for major vegetation stages (green-up, fast growth, maturity and senescence). All vegetation variables were linked with datasets of monthly temperature and precipitation, and correlations between variables were established using Partial Least Squares regression. Most parts of the Tibetan Plateau showed significantly increasing temperatures, as well as clear advances in late season phenological stages by several weeks. Rainfall trends and significant long-term changes in early season phenology occurred on small parts of the plateau. Vegetation activity increased significantly for all vegetation stages. Most of these changes were related to increasing temperatures during the growing season and in some cases during the previous winter. Precipitation effects appeared less pronounced. Warming thus appears to have shortened the growing season, while increasing vegetation activity.

Conclusions/Significance

Shortening of the growing season despite a longer thermally favorable period implies that vegetation on the Tibetan Plateau is unable to exploit additional thermal resources availed by climate change. Ecosystem composition may no longer be well attuned to the local temperature regime, which has changed rapidly over the past three decades. This apparent lag of the vegetation assemblage behind changes in climate should be taken into account when projecting the impacts of climate change on ecosystem processes.     

Long-term relationships among atmospheric CO2, stomata, and intrinsic water use efficiency in individual trees

Leaf-level responses to increases in atmospheric carbon dioxide (CO(2)) concentrations could have large implications for water and carbon cycles. We investigated whether stomatal density, guard cell length, and intrinsic water use efficiency (iWUE) of 27 individual trees growing at the Arnold Arboretum in Boston, Massachusetts have responded to changing environmental conditions over the last 100 years. We examined leaves from 74 herbarium specimens collected from three genera-Acer (maples), Quercus (oaks), and Carpinus (hornbeams)-from 1893 to 2006. During this period, global average atmospheric CO(2) concentrations increased by approximately 29% (86 ppm), and temperatures in Boston increased by 1.8°C. Stomatal density and guard cell length were negatively correlated in oaks and hornbeams. Although stomatal density declined and guard cell length increased over time, the changes were not dependent on the magnitude of changes in CO(2) concentrations. Intrinsic WUE did not change significantly over time. Our findings suggest that iWUE may not respond to changes in CO(2) concentrations over the lifetimes of individual trees, possibly because of compensating changes in stomatal density and guard cell size. We provide an example of a method that can enable researchers to differentiate between genetic and plastic responses to global change in long-lived trees.     

Tree-ring analysis and conifer growth responses to increased atmospheric CO2 levels

Summary Tree-ring data of naturally grown connifers were analyzed to evaluate the possibility of enhanced tree growth due to increased atmospheric CO₂. Tree cores were obtained from 34 sites in four different climatic regions in the northern hemisphere. In each of the four regions, the sampling sites were located along ecological gradients between the subalpine treeline and low elevations and, sometimes, the arid forest border. Growth trends after 1950, when the atmospheric CO₂ concentration increased by more than 30 l·l^-1 indicate an increase in ring-widths at eight of the 34 sites. These chronologies were from sites which moderate temperature or water stress. In four cases the growth increase in the post-1950 period coincided with favorable climatic conditions. In the remaining four cases, the growth increase exceeded the upper bound response expected from CO₂ enrichment experiments with seedling conifer species. Therefore, increased growth in any of the tree-ring chronologies examined could not be solely attributed to higher atmospheric CO₂ concentrations.Major financial supporters: Swiss National Science Foundation (application no. 1.869-0.83); Swiss Federal Institute of Forestry Research, 8903 Birmensdorf, Switzerland; other financial supporters: Carbon Dioxide Research Division, U.S. Department of Energy under subcontract no. 11X-57507V with Martin Marietta Energy Systems, IncOperated by Martin Marietta Energy Systems, Inc., under contract DE-AC05-840R21400 with U.S. Department of Energy     

Interaction of ocean and biosphere in their transient responses to increasing atmospheric CO2

Increasing atmospheric CO₂ induces a net uptake of carbon in the ocean by a shift in chemical equilibrium in seawater, and in the terrestrial biosphere by a stimulated photosynthesis and productivity. The fractions absorbed in both biosphere and ocean decline with increasing dynamics of the release rate of CO₂ into the atmosphere. However, the relative portion of ocean absorption descends much faster with annual growth rate of CO₂ release than biospheric absorption does, due to a difference in dynamics. The equilibrium absorption capacity of the biosphere is estimated to be only one quarter of that of the ocean, but the current sink size of the biosphere is about half of that of the ocean.Apart from CO₂-stimulated carbon fixation, the biosphere releases CO₂ as a result of land use changes, in particular after deforestation. Both of these fluxes are of the order of 1–1.5 Pg of carbon per year. The CO₂-fertilization effect and regrowth together have turned the terrestrial biosphere as a whole from a source into a sink.     

The combined effects of warming and drying suppress CO2 and N2O emission rates in an alpine meadow of the eastern Tibetan Plateau

The eastern Tibetan Plateau has become increasingly warmer and drier since the 1990s. Such warming and drying has a great impact on ecosystem processes on the eastern Tibetan Plateau. To determine their combined effects on CO₂ and N₂O emission rates, we conducted a field manipulative experiment in an alpine meadow of the eastern Tibetan Plateau during the growing season of 2009. The experiment showed that warming manipulation increased soil temperature by 1?°C, and drying manipulation decreased soil water content by 6.8?%. We found that by counteracting the effect of low temperature in the area, experimental warming significantly increased soil microbial biomass, the number of bacteria, fungi, actinomycetes, ammonifying bacteria, nitrobacteria and denitrifying bacteria, and facilitated the emission rates of CO₂ and N₂O by 33.4 and 31.5?%, respectively. However, decreased precipitation further aggravated soil water stress and inhibited the numbers of these organisms, and reduced the emission rates of CO₂ and N₂O by 47.4 and 37.9?%, respectively. So decreased soil water content tended to offset the positive effect of warming. Compared to the positive effects of warming, decreased soil water content was shown in our study to have even greater impact on the eastern Tibetan Plateau during the growing season. Therefore, inhibition of CO₂ and N₂O emission rates (32.3 and 29.3?%, respectively) by warming and drying will intensify if the combined effects of these climatic trends persist in the region.     

CO2 emissions from biomass combustion for bioenergy: atmospheric decay and contribution to global warming

Carbon dioxide (CO₂) emissions from biomass combustion are traditionally assumed climate neutral if the bioenergy system is carbon (C) flux neutral, i.e. the CO₂ released from biofuel combustion approximately equals the amount of CO₂ sequestered in biomass. This convention, widely adopted in life cycle assessment (LCA) studies of bioenergy systems, underestimates the climate impact of bioenergy. Besides CO₂ emissions from permanent C losses, CO₂ emissions from C flux neutral systems (that is from temporary C losses) also contribute to climate change: before being captured by biomass regrowth, CO₂ molecules spend time in the atmosphere and contribute to global warming. In this paper, a method to estimate the climate impact of CO₂ emissions from biomass combustion is proposed. Our method uses CO₂ impulse response functions (IRF) from C cycle models in the elaboration of atmospheric decay functions for biomass‐derived CO₂ emissions. Their contributions to global warming are then quantified with a unit‐based index, the GWP_bio. Since this index is expressed as a function of the rotation period of the biomass, our results can be applied to CO₂ emissions from combustion of all the different biomass species, from annual row crops to slower growing boreal forest.     

Initial responses of soil CO2 efflux and C, N pools to experimental warming in two contrasting forest ecosystems, Eastern Tibetan Plateau, China

Alpine ecosystems are harsh environments where low temperatures are generally a limiting factor. Predicted global warming is thus expected to have a profound impact on alpine ecosystems in the future. This study was conducted to compare the effect of experimental warming on soils in two contrasting forest ecosystems (a dragon spruce plantation and a natural forest) using the open top chamber (OTC) method in the Eastern Tibetan Plateau of China. The OTC enhanced average daily mean soil temperatures by 0.61°C (plantation) and 0.55°C (natural forest), respectively, throughout the growing season. Conversely, soil volumetric moisture declined by 4.10% in the plantation and by 2.55% in the natural forest. Across all measuring dates, warming increased average soil CO₂ efflux by 10.6% in the plantation and by 15.4% in the natural forest. However, elevated temperatures did not affect the respiration quotient in either forest. Two-stage sulfuric acid hydrolysis was used to quantify labile and recalcitrant C and N fractions in the two contrasting soils. Warming significantly reduced labile C and N fractions in both ecosystems but did not influence the total, recalcitrant and microbial biomass C and N pools. Labile C, N and microbial biomass C showed significant interactions in warming × forest type × season. Irrespective of warming treatments, all measured pools were significantly larger in the natural forest compared to the plantation. Taken together, our results indicate that the lowered soil labile C and N pools may be induced by the increased soil CO₂ efflux. The responses of the natural forest soil were more sensitive to experimental warming than those of the plantation. We conclude that reforestation dramatically lowers soil C and N pools, further affecting the responses of forest soils to future global warming.     

Methanotrophic community structure and activity under warming and grazing of alpine meadow on the Tibetan Plateau

Knowledge about methanotrophs and their activities is important to understand the microbial mediation of the greenhouse gas CH₄ under climate change and human activities in terrestrial ecosystems. The effects of simulated warming and sheep grazing on methanotrophic abundance, community composition, and activity were studied in an alpine meadow soil on the Tibetan Plateau. There was high abundance of methanotrophs (1.2–3.4 × 10⁸ pmoA gene copies per gram of dry weight soil) assessed by real-time PCR, and warming significantly increased the abundance regardless of grazing. A total of 64 methanotrophic operational taxonomic units (OTUs) were obtained from 1,439 clone sequences, of these OTUs; 63 OTUs (98.4%) belonged to type I methanotrophs, and only one OTU was Methylocystis of type II methanotrophs. The methanotroph community composition and diversity were not apparently affected by the treatments. Warming and grazing significantly enhanced the potential CH₄ oxidation activity. There were significantly negative correlations between methanotrophic abundance and soil moisture and between methanotrophic abundance and NH₄–N content. The study suggests that type I methanotrophs, as the dominance, may play a key role in CH₄ oxidation, and the alpine meadow has great potential to consume more CH₄ under future warmer and grazing conditions on the Tibetan Plateau.     

Radial Growth of Qilian Juniper on the Northeast Tibetan Plateau and Potential Climate Associations

There is controversy regarding the limiting climatic factor for tree radial growth at the alpine treeline on the northeastern Tibetan Plateau. In this study, we collected 594 increment cores from 331 trees, grouped within four altitude belts spanning the range 3550 to 4020 m.a.s.l. on a single hillside. We have developed four equivalent ring-width chronologies and shown that there are no significant differences in their growth-climate responses during 1956 to 2011 or in their longer-term growth patterns during the period AD 1110–2011. The main climate influence on radial growth is shown to be precipitation variability. Missing ring analysis shows that tree radial growth at the uppermost treeline location is more sensitive to climate variation than that at other elevations, and poor tree radial growth is particularly linked to the occurrence of serious drought events. Hence water limitation, rather than temperature stress, plays the pivotal role in controlling the radial growth of Sabina przewalskii Kom. at the treeline in this region. This finding contradicts any generalisation that tree-ring chronologies from high-elevation treeline environments are mostly indicators of temperature changes.     

CO2浓度和温度升高对红桦根际微生物的影响

应用自控、封闭、独立的生长室系统,研究升高的大气CO2浓度(环境CO2浓度 350(±25)μmol.mol-1,EC)和温度(环境温度 2.0(±0.5)℃,ET)及其交互作用(ECT)对不同栽植密度条件下红桦根际土壤可培养微生物数量的影响。结果表明:(1)EC显著增加了高密度条件下根际细菌数量;在整个生长季中,最大的根际细菌数量增加出现在7月份;而EC对低密度处理的根际细菌数量影响不显著。除了5月和6月份,ET在其余月份均显著增加了根际细菌数量,但是与密度处理没有有意义的相关;ECT对高低密度处理的根际细菌数量均未产生有统计意义的影响。(2)EC对低密度条件下的根际放线菌数量有显著增加,而对高密度条件下的根际放线菌数量无显著影响;ET和ECT对高低密度条件下的根际放线菌数量均未产生有统计意义的影响。(3)EC和ET对高低密度条件下的根际真菌数量无显著增加,而ECT显著增加了根际真菌数量。     

Responses of CO2 efflux from an alpine meadow soil on the Qinghai Tibetan Plateau to multi-form and low-level N addition

Aims

To assess the effects of atmospheric N deposition on the C budget of an alpine meadow ecosystem on the Qinghai–Tibetan Plateau, it is necessary to explore the responses of soil-atmosphere carbon dioxide (CO₂) exchange to N addition.

Methods

Based on a multi-form, low-level N addition experiment, soil CO₂ effluxes were monitored weekly using the static chamber and gas chromatograph technique. Soil variables and aboveground biomass were measured monthly to examine the key driving factors of soil CO₂ efflux.

Results

The results showed that low-level N input tended to decrease soil moisture, whereas medium-level N input maintained soil moisture. Three-year N additions slightly increased soil inorganic N pools, especially the soil NH ₄ ⁺ -N pool. N applications significantly increased aboveground biomass and soil CO₂ efflux; moreover, this effect was more significant from NH ₄ ⁺ -N than from NO ₃ ^? -N fertilizer. In addition, the soil CO₂ efflux was mainly driven by soil temperature, followed by aboveground biomass and NH ₄ ⁺ -N pool.

Conclusions

These results suggest that chronic atmospheric N deposition will stimulate soil CO₂ efflux in the alpine meadow on the Qinghai–Tibetan Plateau by increasing available N content and promoting plant growth.     

Temperature controls ecosystem CO2 exchange of an alpine meadow on the northeastern Tibetan Plateau

Alpine ecosystems are extremely vulnerable to climate change. To address the potential variability of the responses of alpine ecosystems to climate change, we examined daily CO₂ exchange in relation to major environmental variables. A dataset was obtained from an alpine meadow on the Qinghai‐Tibetan Plateau from eddy covariance measurements taken over 3 years (2002–2004). Path analysis showed that soil temperature at 5 cm depth (T_s5) had the greatest effect on daily variation in ecosystem CO₂ exchange all year around, whereas photosynthetic photon flux density (PPFD) had a high direct effect on daily variation in CO₂ flux during the growing season. The combined effects of temperature and light regimes on net ecosystem CO₂ exchange (NEE) could be clearly categorized into three areas depending on the change in T_s5: (1) almost no NEE change irrespective of variations in light and temperature when T_s5 was below 0 °C; (2) an NEE increase (i.e. CO₂ released from the ecosystem) with increasing T_s5, but little response to variation in light regime when 0 °C≤T_s5≤8 °C; and (3) an NEE decrease with increase in T_s5 and PPFD when T_s5 was approximately >8 °C. The highest daily net ecosystem CO₂ uptake was observed under the conditions of daily mean T_s5 of about 15 °C and daily mean PPFD of about 50 mol m⁻² day⁻¹. The results suggested that temperature is the most critical determinant of CO₂ exchange in this alpine meadow ecosystem and may play an important role in the ecosystem carbon budget under future global warming conditions.     

No temperature acclimation of soil extracellular enzymes to experimental warming in an alpine grassland ecosystem on the Tibetan Plateau

Alpine grassland soils store large amounts of soil organic carbon (SOC) and are susceptible to rising air temperature. Soil extracellular enzymes catalyze the rate-limiting step in SOC decomposition and their catalysis, production and degradation rates are regulated by temperature. Therefore, the responses of these enzymes to warming could have a profound impact on carbon cycling in the alpine grassland ecosystems. This study was conducted to measure the responses of soil extracellular enzyme activity and temperature sensitivity (Q₁₀) to experimental warming in samples from an alpine grassland ecosystem on the Tibetan Plateau. A free air-temperature enhancement system was set up in May 2006. We measured soil microbial biomass, nutrient availability and the activity of five extracellular enzymes in 2009 and 2010. The Q₁₀ of each enzyme was calculated using a simple first-order exponential equation. We found that warming had no significant effects on soil microbial biomass C, the labile C or N content, or nutrient availability. Significant differences in the activity of most extracellular enzymes among sampling dates were found, with typically higher enzyme activity during the warm period of the year. The effects of warming on the activity of the five extracellular enzymes at 20 °C were not significant. Enzyme activity in vitro strongly increased with temperature up to 27 °C or over 30 °C (optimum temperature; T_opt). Seasonal variations in the Q₁₀ were found, but the effects of warming on Q₁₀ were not significant. We conclude that soil extracellular enzymes adapted to seasonal temperature variations, but did not acclimate to the field experimental warming.     

Climate change and the phytoplankton spring bloom: warming and overwintering zooplankton have similar effects on phytoplankton

Indoor mesocosms were used to study the combined effect of warming and of different densities of overwintering mesozooplankton (mainly copepods) on the spring development of phytoplankton in shallow, coastal waters. Similar to previous studies, warming accelerated the spring phytoplankton peak by ca. 1 day °C^?1 whereas zooplankton did not significantly influence timing. Phytoplankton biomass during the experimental period decreased with warming and with higher densities of overwintering zooplankton. Similarly, average cell size and average effective particle size (here: colony size) decreased both with zooplankton density and warming. A decrease in phytoplankton particle size is generally considered at typical footprint of copepod grazing. We conclude that warming induced changes in the magnitude and structure of the phytoplankton spring bloom cannot be understood without considering grazing by overwintering zooplankton.     

Effects of increasing UV-B radiation and atmospheric CO2 on photosynthesis and growth: implications for terrestrial ecosystems

Increases in UV-B radiation reaching the earth as a result of stratospheric ozone depletion will most likely accompany increases in atmospheric CO₂ concentrations. Many studies have examined the effects of each factor independently, but few have evaluated the combined effects of both UV-B radiation and elevated CO₂. In general the results of such studies have shown independent effects on growth or seed yield. Although interspecific variation is large, high levels of UV-B radiation tends to reduce plant growth in sensitive species, while CO₂ enrichment tends to promote growth in most C₃ species. However, most previous studies have not looked at temporal effects or at the relationship between photosynthetic acclimation to CO₂ and possible photosynthetic limitations imposed by UV-B radiation. Elevated CO₂ may provide some protection against UV-B for some species. In contrast, UV-B radiation may limit the ability to exploit elevated CO₂ in other species. Interactions between the effects of CO₂ enrichment and UV-B radiation exposure have also been shown for biomass allocation. Effects on both biomass allocation and photosynthetic acclimation may be important to ecosystem structure in terms of seedling establishment, competition and reproductive output. Few studies have evaluated ecosystem processes such as decomposition or nutrient cycling. Interactive effects may be subtle and species specific but should not be ignored in the assessment of the potential impacts of increases in CO₂ and UV-B radiation on plants.     

Responses of CO2, CH4 and N2O fluxes to livestock exclosure in an alpine steppe on the Tibetan Plateau, China

Background and aims

The photochemical reflectance index (PRI) is correlated to photosynthetic efficiency and has been successfully applied at multiple scales for remote estimation of physiological functioning. However, interpretation of the PRI signal can be confounded by many different variables including declines in photochemical pigments. Our study was aimed at investigating PRI in response to salinity stress, and evaluating physiological and pigment responses of two co-occurring shrubs, Baccharis halimifolia and Myrica cerifera in laboratory studies.

Methods

Photosynthesis, water relations, chlorophyll fluorescence, hyperspectral reflectance and leaf pigment contents were measured following salinity treatment.

Results

Physiological measurements indicated that both species exhibit adaptations which protect PSII during periods of stress. Chlorophyll fluorescence parameters were affected in both species, but indicated that other photochemical reactions (e.g. photorespiration) were important for energy dissipation in absence of chlorophyll changes. After many days of reduced photosynthesis, photochemical changes were detectable using PRI indicating chronic stress.

Conclusions

Variations in PRI were not related to changes in pigments but strongly related to tissue chlorides indicating salinity effects on the PRI signal. Thus, PRI is an indicator of salinity stress in these coastal species and may be as an early signal for increasing salt exposure associated with rising sea-level and climate change.     

Effect of increased atmospheric CO2 on the performance of an aquatic detritivore through changes in water temperature and litter quality

Cold water woodland streams, where terrestrially derived organic matter fuels aquatic food webs, can be affected by increases in atmospheric CO₂ concentrations, as these are predicted to lead to increases in water temperature and decreases in organic matter quality. In fact, elevated CO₂ (580 ppm) decreased the initial phosphorus concentration of birch litter by 30% compared with litter grown under ambient conditions (380 ppm). Here, we first assessed the effect of differences in litter quality on mass loss, microbial colonization and conditioned litter quality after submersion in a mountain stream for 2 weeks. Leaching did not change the relative differences between litter types, while fungal biomass was two fold higher in elevated litter. We then offered this litter (conditioned ambient and elevated) to a stream detritivore that was kept at 10 and 15 °C to assess the individual and interactive effects of increased temperature and decreased litter quality on invertebrate performance. When given a choice, the detritivore preferred elevated litter, but only at 10 °C. When fed litter types singularly, there was no effect of litter quality on consumption rates; however, the effect of temperature depended on individual size and time of collection. Growth rates were higher in individuals fed ambient litter at 10 °C when compared with individuals fed elevated litter at 15 °C. Mortality did not differ between litter types, but was higher at 15 °C than at 10 °C. Increases in temperature led to alterations in the individual body elemental composition and interacted with litter type. The performance of the detritivore was therefore more affected by increases in temperature than by small decreases in litter quality. However, it seems conceivable that in a future global warming scenario the simultaneous increases in water temperature and decreases in litter quality might affect detritivores performance more than predicted from the effects of both factors considered individually.     

Modifications of the carbon and nitrogen allocations in the plant (Triticum aestivum L.) soil system in response to increased atmospheric CO2 concentration

The aim of this work was to examine the response of wheat plants to a doubling of the atmospheric CO₂ concentration on: (1) carbon and nitrogen partitioning in the plant; (2) carbon release by the roots; and (3) the subsequent N uptake by the plants. The experiment was performed in controlled laboratory conditions by exposing fast-growing spring wheat plants, during 28 days, to a ¹⁴CO₂ concentration of 350 or 700 L L^–1 at two levels of soil nitrogen fertilization. Doubling CO₂ availability increased total plant production by 34% for both N treatment. In the N-fertilized soil, the CO₂ enrichment resulted in an increase in dry mass production of 41% in the shoots and 23% in the roots; without N fertilization this figure was 33% and 37%, respectively. In the N-fertilized soil, the CO₂ increase enhanced the total N uptake by 14% and lowered the N concentration in the shoots by 23%. The N concentration in the roots was unchanged. In the N-fertilized soil, doubling CO₂ availability increased N uptake by 32% but did not change the N concentrations, in either shoots or roots. The CO₂ enrichment increased total root-derived carbon by 12% with N fertilization, and by 24% without N fertilization. Between 85 and 90% of the total root derived-¹⁴C came from respiration, leaving only 10 to 15% in the soil as organic ¹⁴C. However, when total root-derived ¹⁴C was expressed as a function of root dry weight, these differences were only slightly significant. Thus, it appears that the enhanced carbon release from the living roots in response to increased atmospheric CO₂, is not due to a modification of the activity of the roots, but is a result of the increased size of the root system. The increase of root dry mass also resulted in a stimulation of the soil N mineralization related to the doubling atmospheric CO₂ concentration. The discussion is focused on the interactions between the carbon and nitrogen allocation, especially to the root system, and the implications for the acquisition of nutrients by plants in response to CO₂ increase.Abbreviations N soil fertilization without nitrogen - N soil fertilization with nitrogen