Elevated atmospheric CO2 affects decomposition of Festuca vivipara (L.) Sm. litter and roots in experiments simulating environmental change in two contrasting arctic ecosystems

Mass loss, together with nitrogen and carbon loss, from above-ground material and roots of Festuca vivipara were followed for 13 months in a high Arctic polar semi-desert and a low Arctic tree-line dwarf shrub heath. Festuca vivipara for the study was obtained from plants cultivated at two different CO₂ concentrations (350 and 500 μL L^–1) in controlled environment chambers in the UK. Each of the four resource types (shoots or roots from plants grown in elevated or ambient CO₂ concentrations) was subsequently placed in an experiment simulating aspects of environmental change in each Arctic ecosystem. Air, litter and soil temperatures were increased using open-topped polythene tents at both sites, and a 58% increase in summer precipitation was simulated at the high Arctic site. Mass loss was greatest at the low Arctic site, and from the shoot material, rather than the roots. Shoots grown under an elevated CO₂ concentration decomposed more slowly at the high Arctic site, and more quickly at the low Arctic one, than shoots grown at ambient CO₂. After 13 months, greater amounts of C and N remained in above-ground litter from plants grown under elevated, rather than ambient, CO₂ at the polar semi-desert site, although lower amounts of C remained in elevated CO₂ litter at the low Arctic ecosystem. In the high Arctic, roots grown in the 500 μL L^–1 CO₂ concentration decomposed significantly more slowly than below-ground material derived from the ambient CO₂ chambers. Elevated CO₂ concentrations significantly increased the inital C:N ratio, % soluble carbohydrates and α-cellulose content, and significantly decreased the inital N content, of the above-ground material compared to that derived from the ambient treatment. Initially, the C:N ratio and percentage N were similar in both sets of roots derived from the two different CO₂ treatments, but soluble carbohydrate and α-cellulose concentrations were higher, and percentage lignin lower, in the elevated CO₂ treatments.The tent treatments significantly retarded shoot decomposition in both ecosystems, probably because of lower litter bag moisture contents, although the additional precipitation treatment had no effect on mass loss from the above-ground material. The results suggest that neither additional summer precipitation (up to 58%), nor soil temperature increase of 1 °C, which may occur by the end of the next century as an effect of a predicted 4 °C rise in air temperature, had an appreciable effect on root decomposition in the short term in a high Arctic soil. However, at the low Arctic site, greater root decomposition, and a lower pool of root N remaining, were observed where soil temperature was increased by 2 °C in response to a 4 °C rise in air temperature. These results suggest that decomposition below-ground in this ecosystem would increase as an effect of predicted climate change. These data also show that there is a difference in the initial results of decomposition processes between the two Arctic ecosystems in response to simulated environmental change.     

The effects of elevated CO2 on symbiotic N2 fixation: a link between the carbon and nitrogen cycles in grassland ecosystems

The response of plants to elevated CO₂ is dependent on the availability of nutrients, especially nitrogen. It is generally accepted that an increase in the atmospheric CO₂ concentration increases the C:N ratio of plant residues and exudates. This promotes temporary N-immobilization which might, in turn, reduce the availability of soil nitrogen. In addition, both a CO₂ stimulated increase in plant growth (thus requiring more nitrogen) and an increased N demand for the decomposition of soil residues with a large C:N will result under elevated CO₂ in a larger N-sink of the whole grassland ecosystem. One way to maintain the balance between the C and N cycles in elevated CO₂ would be to increase N-import to the grassland ecosystem through symbiotic N₂ fixation. Whether this might happen in the context of temperate ecosystems is discussed, by assessing the following hypothesis: i) symbiotic N₂ fixation in legumes will be enhanced under elevated CO₂, ii) this enhancement of N₂ fixation will result in a larger N-input to the grassland ecosystem, and iii) a larger N-input will allow the sequestration of additional carbon, either above or below-ground, into the ecosystem. Data from long-term experiments with model grassland ecosystems, consisting of monocultures or mixtures of perennial ryegrass and white clover, grown under elevated CO₂ under free-air or field-like conditions, supports the first two hypothesis, since: i) both the percentage and the amount of fixed N increases in white clover grown under elevated CO₂, ii) the contribution of fixed N to the nitrogen nutrition of the mixed grass also increases in elevated CO₂. Concerning the third hypothesis, an increased nitrogen input to the grassland ecosystem from N₂ fixation usually promotes shoot growth (above-ground C storage) in elevated CO₂. However, the consequences of this larger N input under elevated CO₂ on the below-ground carbon fluxes are not fully understood. On one hand, the positive effect of elevated CO₂ on the quantity of plant residues might be overwhelming and lead to an increased long-term below-ground C storage; on the other hand, the enhancement of the decomposition process by the N-rich legume material might favour carbon turn-over and, hence, limit the storage of below-ground carbon.     

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.     

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 effects of free-air CO2 enrichment and soil water availability on spatial and seasonal patterns of wheat root growth

Spring wheat [ Triticum aestivum (L). cv. Yecora Rojo] was grown from December 1992 to May 1993 under two atmospheric CO₂ concentrations, 550 μmol mol^–1 for high-CO₂ plots, and 370 μmol mol^–1 for control plots, using a Free-Air CO₂ Enrichment (FACE) apparatus. In addition to the two levels of atmospheric CO₂, there were ample and limiting levels of water supply through a subsurface trip irrigation system in a strip, split-plot design. In order to examine the temporal and spatial root distribution, root cores were extracted at six growth stages during the season at in-row and inter-row positions using a soil core device (86 mm ID, 1.0 m length). Such information would help determine whether and to what extent root morphology is changed by alteration of two important factors, atmospheric CO₂ and soil water, in this agricultural ecosystem. Wheat root growth increased under elevated CO₂ conditions during all observed developmental stages. A maximum of 37% increase in total root dry mass in the FACE vs. Control plots was observed during the period of stem elongation. Greater root growth rates were calculated due to CO₂ enhancement until anthesis. During the early vegetative growth, root dry mass of the inter-row space was significantly higher for FACE than for Control treatments suggesting that elevated CO₂ promoted the production of first-order lateral roots per main axis. Then, during the reproductive period of growth, more branching of lateral roots in the FACE treatment occurred due to water stress. Significant higher root dry mass was measured in the inter-row space of the FACE plots where soil water supply was limiting. These sequential responses in root growth and morphology to elevated CO₂ and reduced soil water supports the hypothesis that plants grown in a high-CO₂ environment may better compensate soil-water-stress conditions.     

The Earthtron facility for below-ground manipulation study

A controlled environmental facility is necessary for investigation of the interaction between above-ground and below-ground components in microcosm experiments. The Earthtron is a simple computer-controlled chamber simulating natural environments: diurnal light/dark cycles, and separately controlled soil and air temperature, humidity and CO₂ concentration. In soil core incubation experiments, the Earthtron was able to simulate the dynamics of soil temperature field conditions. Environmental control also affects the dynamics of soil water and distribution patterns of nutrients in the microcosm, linking to the distribution of plant roots and soil biota. The Earthtron can not only reproduce field conditions but also predict the effects of global climatic change on terrestrial ecosystems.     

Nitrogen Fertilization Has a Stronger Effect on Soil Nitrogen-Fixing Bacterial Communities than Elevated Atmospheric CO2

Biological nitrogen fixation is the primary supply of N to most ecosystems, yet there is considerable uncertainty about how N-fixing bacteria will respond to global change factors such as increasing atmospheric CO₂ and N deposition. Using the nifH gene as a molecular marker, we studied how the community structure of N-fixing soil bacteria from temperate pine, aspen, and sweet gum stands and a brackish tidal marsh responded to multiyear elevated CO₂ conditions. We also examined how N availability, specifically, N fertilization, interacted with elevated CO₂ to affect these communities in the temperate pine forest. Based on data from Sanger sequencing and quantitative PCR, the soil nifH composition in the three forest systems was dominated by species in the Geobacteraceae and, to a lesser extent, Alphaproteobacteria. The N-fixing-bacterial-community structure was subtly altered after 10 or more years of elevated atmospheric CO₂, and the observed shifts differed in each biome. In the pine forest, N fertilization had a stronger effect on nifH community structure than elevated CO₂ and suppressed the diversity and abundance of N-fixing bacteria under elevated atmospheric CO₂ conditions. These results indicate that N-fixing bacteria have complex, interacting responses that will be important for understanding ecosystem productivity in a changing climate.     

Effects of elevated atmospheric CO2 on fine root length and distribution in an oak-palmetto scrub ecosystem in central Florida

Atmospheric CO₂ concentration is rising and it has been suggested that a portion of the additional carbon is being sequestered in terrestrial vegetation and much of that in below-ground structures. The objective of the present study was to quantify the effects of elevated atmospheric CO₂ on fine root length and distribution with depth with minirhizotrons in an open-top chamber experiment in an oak-palmetto scrub ecosystem at Kennedy Space Centre, Florida, USA. Observations were made five times over a period of one and a half years in three ambient chambers (350 p.p.m. CO₂), three CO₂ enriched chambers (700 p.p.m. CO₂), and three unchambered plots. Greater root length densities were produced in the elevated CO₂ chambers (14.2 mm cm^?2) compared to the ambient chambers (8.7 mm cm^?2). More roots may presumably lead to more efficient acquisition of resources. Fine root abundance varied significantly with soil depth, and there appeared to be enhanced proliferation of fine roots near the surface (0–12 cm) and at greater depth (49–61 cm) in the elevated CO₂ chambers. The vertical root distribution pattern may be a response to availability of nutrients and water. More studies are needed to determine if increased root length under CO₂ enriched conditions actually results in greater sequestering of carbon below ground.     

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.     

Stability of above-ground and below-ground processes to extreme drought in model grassland ecosystems: Interactions with plant species diversity and soil nitrogen availability

Extreme drought events have the potential to cause dramatic changes in ecosystem structure and function, but the controls upon ecosystem stability to drought remain poorly understood. Here we used model systems of two commonly occurring, temperate grassland communities to investigate the short-term interactive effects of a simulated 100-year summer drought event, soil nitrogen (N) availability and plant species diversity (low/high) on key ecosystem processes related to carbon (C) and N cycling. Whole ecosystem CO₂ fluxes and leaching losses were recorded during drought and post-rewetting. Litter decomposition and C/N stocks in vegetation, soil and soil microbes were assessed 4 weeks after the end of drought. Experimental drought caused strong reductions in ecosystem respiration and net ecosystem CO₂ exchange, but ecosystem fluxes recovered rapidly following rewetting irrespective of N and species diversity. As expected, root C stocks and litter decomposition were adversely affected by drought across all N and plant diversity treatments. In contrast, drought increased soil water retention, organic nutrient leaching losses and soil fertility. Drought responses of above-ground vegetation C stocks varied depending on plant diversity, with greater stability of above-ground vegetation C to drought in the high versus low diversity treatment. This positive effect of high plant diversity on above-ground vegetation C stability coincided with a decrease in the stability of microbial biomass C. Unlike species diversity, soil N availability had limited effects on the stability of ecosystem processes to extreme drought. Overall, our findings indicate that extreme drought events promote post-drought soil nutrient retention and soil fertility, with cascading effects on ecosystem C fixation rates. Data on above-ground ecosystem processes underline the importance of species diversity for grassland function in a changing environment. Furthermore, our results suggest that plant–soil interactions play a key role for the short-term stability of above-ground vegetation C storage to extreme drought events.     

Responses of shoot and root gas exchange, leaf blade expansion and biomass production to pulses of elevated C02 in hydroponic wheat

Short-term effects of elevated CO₂ during the early life phaseof plants may have long lasting consequences for growth andbiomass in later periods. We exposed hydroponically grown wheatseedlings to 5 d pulses of elevated CO₂ while leaf expansiongrowth as well as shoot and root gas exchange were measuredsimultaneously and continuously. Shoot photosynthesis, night-timeshoot respiration and below-ground respiration (largely by roots)roughly doubled when atmospheric CO₂ concentration was doubled.An interruption of CO₂ enrichment caused CO₂ assimilation andrespiration to return to control levels. However, while theresponse of photosynthesis was immediate, that of respirationshowed a hysteresis of about 3 d. Since shoot biomass increasedat elevated CO₂ (with no change in allocation pattern) equalfluxes per shoot or root system after a return to control CO₂concentrations indicate substantial downward adjustment of thecapacity for CO₂ fixation and release in high-CO₂ grown plants.Leaf expansion growth was completely unaffected by CO₂ enrichment,whereas tiller initiation was significantly increased (doubledin 18 d). We conclude that leaf growth in these wheat plantswas already carbon-saturated at ambient CO₂ concentration atoptimum mineral nutrient supply. The stimulation of growth ofwhole plants was exclusively due to enhanced tillering duringthis very early part of the life of these wheat plants. Key words: Allocation, atmospheric carbon dioxide enrichment, growth, photosynthesis, respiration, tillering, Triticum aestivum     

Exposure to warming and CO2 enrichment promotes greater above-ground biomass, nitrogen, phosphorus and arbuscular mycorrhizal colonization in newly established grasslands

Aims

In view of the projected increase in global air temperature and CO₂ concentration, the effects of climatic changes on biomass production, CO₂ fluxes and arbuscular mycorrhizal fungi (AMF) colonization in newly established grassland communities were investigated. We hypothesized that above- and below-ground biomass, gross primary productivity (GPP), AMF root colonization and nutrient acquisition would increase in response to the future climate conditions. Furthermore, we expected that increased below-ground C allocation would enhance soil respiration (R_soil).

Methods

Grassland communities were grown either at ambient temperatures with 375?ppm CO₂ (Amb) or at ambient temperatures +3°C with 620?ppm CO₂ (T+CO₂).

Results

Total biomass production and GPP were stimulated under T+CO₂. Above-ground biomass was increased under T+CO₂ while belowground biomass was similar under both climates. The significant increase in root colonization intensity under T+CO₂, and therefore the better contact between roots and AMF, probably determined the higher above-ground P and N content. R_soil was not significantly affected by the future climate conditions, only showing a tendency to increase under future climate at the end of the season.

Conclusions

Newly established grasslands benefited from the exposure to elevated CO₂ and temperature in terms of total biomass production; higher root AMF colonization may partly provide the nutrients required to sustain this growth response.     

Carbon balance of a tropical savanna of northern Australia

Through estimations of above- and below-ground standing biomass, annual biomass increment, fine root production and turnover, litterfall, canopy respiration and total soil CO₂ efflux, a carbon balance on seasonal and yearly time-scales is developed for a Eucalypt open-forest savanna in northern Australia. This carbon balance is compared to estimates of carbon fluxes derived from eddy covariance measurements conducted at the same site. The total carbon (C) stock of the savanna was 204±53 ton C ha^–1, with approximately 84% below-ground and 16% above-ground. Soil organic carbon content (0–1 m) was 151±33 ton C ha^–1, accounting for about 74% of the total carbon content in the ecosystem. Vegetation biomass was 53±20 ton C ha^–1, 39% of which was found in the root component and 61% in above-ground components (trees, shrubs, grasses). Annual gross primary production was 20.8 ton C ha^–1, of which 27% occurred in above-ground components and 73% below-ground components. Net primary production was 11 ton C ha^–1 year^–1, of which 8.0 ton C ha^–1 (73%) was contributed by below-ground net primary production and 3.0 ton C ha^–1 (27%) by above-ground net primary production. Annual soil carbon efflux was 14.3 ton C ha^–1 year^–1. Approximately three-quarters of the carbon flux (above-ground, below-ground and total ecosystem) occur during the 5–6 months of the wet season. This savanna site is a carbon sink during the wet season, but becomes a weak source during the dry season. Annual net ecosystem production was 3.8 ton C ha^–1 year^–1.     

Belowground positive and negative feedbacks on CO2 growth enhancement

In this paper we present a conceptual model of integrated plant-soil interactions which illustrates the importance of identifying the primary belowground feedbacks, both positive and negative, which can simultaneously affect plant growth responses to elevated CO₂. The primary negative feedbacks share the common feature of reducing the amount of nutrients available to plants. These negative feedbacks include increased litter C/N ratios, and therefore reduced mineralization rates, increased immobilization of available nutrients by a larger soil microbial pool, and increased storage of nutrients in plant biomass and detritus due to increases in net primary productivity (NPP). Most of the primary positive feedbacks share the common feature of being plant mediated feedbacks, the only exception being Zak et al.'s hypothesis that increased microbial biomass will be accompanied by increased mineralization rates. Plant nutrient uptake may be increased through alterations in root architecture, physiology, or mycorrhizal symbioses. Further, the increased C/N ratios of plant tissue mean that a given level of NPP can be achieved with a smaller supply of nitrogen.Identification of the net plant-soil feedbacks to enhanced productivity with elevated CO₂ are a critical first step for any ecosystem. It is necessary, however, that we first identify how universally applicable the results are from one study of one ecosystem before ecosystem models incorporate this information. The effect of elevated CO₂ on plant growth (including NPP, tissue quality, root architecture, mycorrhizal symbioses) can vary greatly for different species and environmental conditions. Therefore it is reasonable to expect that different ecosystems will show different patterns of interacting positive and negative feedbacks within the plant-soil system. This inter-ecosystem variability in the potential for long-term growth responses to rising CO₂ levels implies that we need to parameterize mechanistic models of the impact of elevated CO₂ on ecosystem productivity using a detailed understanding of each ecosystem of interest.     

Structural equation modelling reveals plant-community drivers of carbon storage in boreal forest ecosystems

Boreal forest ecosystems are important drivers of the global carbon (C) cycle by acting as both sinks and sources of atmospheric CO₂. While several factors have been proposed as determining the ability of boreal forest to function as C sinks, little is known about their relative importance. In this study, we applied structural equation modelling to a previously published dataset involving 30 boreal-forested islands that vary greatly in their historic fire regime, in order to explore the simultaneous influence of several factors believed to be important in influencing above-ground, below-ground and total ecosystem C accumulation. We found that wildfire was a major driver of ecosystem C sequestration, and exerted direct effects on below-ground C storage (presumably through humus combustion) and indirect effects on both above-ground and below-ground C storage through altering plant-community composition. By contrast, plant diversity influenced only below-ground C storage (and even then only weakly), while net primary productivity and decomposition had no detectable effect. Our results suggest that while boreal forests have great potential for storing significant amounts of C, traits of dominant plant species that promote below-ground C accumulation and the absence of wildfire are the most important drivers of C sequestration in these ecosystems.     

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.     

Interactive effects of elevated carbon dioxide and environmental stresses on root mass fraction in plants: a meta-analytical synthesis using pairwise techniques

Rising atmospheric CO₂ greatly enhances plant production, but its effect on biomass allocation, particularly in the presence of environmental stresses, is not well understood. Here, we used meta-analysis combined with pairwise techniques to examine root mass fraction (RMF; i.e., the fraction of root to total biomass) as affected by elevated CO₂ and environmental stresses. Our results showed that lower soil fertility increased RMF and the magnitude was similar for ambient and elevated CO₂-grown plants. Lower soil water also increased RMF, but to a greater extent at elevated than at ambient CO₂. While CO₂ enrichment had little effect on the magnitude of O₃-caused reduction in RMF in herbaceous species, it alleviated the adverse effect of higher O₃ on root production in woody species. These results demonstrate that CO₂ has less pronounced effects on RMF than other environmental factors. Under abiotic stresses, e.g., drought and higher O₃, elevated CO₂-grown plants will likely increase biomass allocation below-ground. Because of the non-uniform changes in drought and O₃ projected for different parts of the world, we conclude that elevated CO₂ will have regional, but not global, effects on biomass allocation under various global change scenarios.     

Root growth and functioning under atmospheric CO2 enrichment

This paper examines the extent to which atmospheric CO₂ enrichment may influence growth of plant roots and function in terms of uptake of water and nutrients, and carbon allocation towards symbionts. It is concluded that changes in dry matter allocation greatly depend on the experimental conditions during the experiment, the growth phase of the plant, and its morphological characteristics. Under non-limiting conditions of water and nutrients for growth, dry matter partitioning to the root is not changed by CO₂ enrichment. The increase in root/shoot ratio, frequently observed under limiting conditions of water and/or nutrients, enables the plant to explore a greater soil volume, and hence acquire more water and nutrients. However, more data on changes in dry matter allocation within the root due to atmospheric CO₂ are needed. It is concluded that nitrogen fixation is favored by CO₂ enrichment since nodule mass is increased, concomitant with an increase in root length. The papers available so far on the influence of CO₂ enrichment on mycorrhizal functioning suggest that carbon allocation to the roots might be increased, but also here more experiments are needed.Abbreviations LAR leaf area ratio - LWR leaf weight ratio - SWR stem weight ratio - RGR relative growth rate - R/S root/shoot - RWR root weight ratio     

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     

Rapid root closure after fire limits fine root responses to elevated atmospheric CO2 in a scrub oak ecosystem in central Florida, USA

Elevated atmospheric carbon dioxide (CO₂) often stimulates the growth of fine roots, yet there are few reports of responses of intact root systems to long‐term CO₂ exposure. We investigated the effects of elevated CO₂ on fine root growth using open top chambers in a scrub oak ecosystem at Kennedy Space Center, Florida for more than 7 years. CO₂ enrichment began immediately after a controlled burn, which simulated the natural disturbance that occurs in this system every 10–15 years. We hypothesized that (1) root abundance would increase in both treatments as the system recovered from fire; (2) elevated CO₂ would stimulate root growth; and (3) elevated CO₂ would alter root distribution. Minirhizotron tubes were used to measure fine root length density (mm cm^?2) every three months. During the first 2 years after fire recovery, fine root abundance increased in all treatments and elevated CO₂ significantly enhanced root abundance, causing a maximum stimulation of 181% after 20 months. The CO₂ stimulation was initially more pronounced in the top 10 cm and 38–49 cm below the soil surface. However, these responses completely disappeared during the third year of experimental treatment: elevated CO₂ had no effect on root abundance or on the depth distribution of fine roots during years 3–7. The results suggest that, within a few years following fire, fine roots in this scrub oak ecosystem reach closure, defined here as a dynamic equilibrium between production and mortality. These results further suggest that elevated CO₂ hastens root closure but does not affect maximum root abundance. Limitation of fine root growth by belowground resources – particularly nutrients in this nutrient‐poor soil – may explain the transient response to elevated CO₂.