This Special Issue of New Phytologist contains the latest information and new ideas about how root dynamics might alter in the face of a globally changing environment. The importance of this topic is clear: changes in the production and turnover of roots in forests and grasslands in response to rising atmospheric CO2 concentrations, elevated temperatures, altered precipitation, or nitrogen deposition could be a key link between plant responses and longer-term changes in soil organic matter and ecosystem carbon balance.

The introductory review (Norby & Jackson, 2000), which draws together the different contributions to the volume, asks three central questions:

[bull ] Do elevated atmospheric CO2, nitrogen deposition, and climatic change alter the dynamics of root production and mortality?

[bull ] How do physiological responses of roots to global change factors impact whole-plant and ecosystem metabolism?

[bull ] What are the implications of root dynamics for soil microbial communities and the fate of carbon in soil?

Ecosystem-level observations of root production and mortality in response to global change factors are just starting to emerge. The challenge to root biologists is to overcome the profound methodological and analytical problems and assemble a more comprehensive data set from which ecosystem responses can be explained. The commissioned reviews and research papers in this volume attempt to meet that challenge. Following the introductory review, three papers provide a framework for subsequent analyses by presenting a global perspective on root turnover, a review of morphological and physiological attributes of roots, and a discussion of concepts of carbon allocation in plants. This is followed by a series of papers describing experimental studies on the effects of elevated CO2 and climatic change in various ecosystems. Three papers consider the physiological responses of roots to global change factors, followed by three papers reviewing mycorrhizal interactions and soil biology, and the implications for carbon sequestration in soil. The final paper returns to a global perspective with an analysis of how roots are handled in models of global change. Throughout these articles there is information on topics such as methodology for studying root dynamics, the major gaps in our knowledge, and the idea that leaves are a good analogy for roots.

Changes in the production and turnover of roots in forests and grasslands in response to rising atmospheric CO2 concentrations, elevated temperatures, altered precipitation, or nitrogen deposition could be a key link between plant responses and longer-term changes in soil organic matter and ecosystem carbon balance. Here we summarize the experimental observations, ideas, and new hypotheses developed in this area in the rest of this volume. Three central questions are posed. Do elevated atmospheric CO2, nitrogen deposition, and climatic change alter the dynamics of root production and mortality? What are the consequences of root responses to plant physiological processes? What are the implications of root dynamics to soil microbial communities and the fate of carbon in soil? Ecosystem-level observations of root production and mortality in response to global change parameters are just starting to emerge. The challenge to root biologists is to overcome the profound methodological and analytical problems and assemble a more comprehensive data set with sufficient ancillary data that differences between ecosystems can be explained. The assemblage of information reported herein on global patterns of root turnover, basic root biology that controls responses to environmental variables, and new observations of root and associated microbial responses to atmospheric and climatic change helps to sharpen our questions and stimulate new research approaches. New hypotheses have been developed to explain why responses of root turnover might differ in contrasting systems, how carbon allocation to roots is controlled, and how species differences in root chemistry might explain the ultimate fate of carbon in soil. These hypotheses and the enthusiasm for pursuing them are based on the firm belief that a deeper understanding of root dynamics is critical to describing the integrated response of ecosystems to global change.

Root turnover is a critical component of ecosystem nutrient dynamics and carbon sequestration and is also an important sink for plant primary productivity. We tested global controls on root turnover across climatic gradients and for plant functional groups by using a database of 190 published studies. Root turnover rates increased exponentially with mean annual temperature for fine roots of grasslands (r2 = 0.48) and forests (r2 = 0.17) and for total root biomass in shrublands (r2 = 0.55). On the basis of the best-fit exponential model, the Q10 for root turnover was 1.4 for forest small diameter roots (5 mm or less), 1.6 for grassland fine roots, and 1.9 for shrublands. Surprisingly, after accounting for temperature, there was no such global relationship between precipitation and root turnover. The slowest average turnover rates were observed for entire tree root systems (10% annually), followed by 34% for shrubland total roots, 53% for grassland fine roots, 55% for wetland fine roots, and 56% for forest fine roots. Root turnover decreased from tropical to high-latitude systems for all plant functional groups. To test whether global relationships can be used to predict interannual variability in root turnover, we evaluated 14 yr of published root turnover data from a shortgrass steppe site in northeastern Colorado, USA. At this site there was no correlation between interannual variability in mean annual temperature and root turnover. Rather, turnover was positively correlated with the ratio of growing season precipitation and maximum monthly temperature (r2 = 0.61). We conclude that there are global patterns in rates of root turnover between plant groups and across climatic gradients but that these patterns cannot always be used for the successful prediction of the relationship of root turnover to climate change at a particular site.

Root turnover is important to the global carbon budget as well as to nutrient cycling in ecosystems and to the success of individual plants. Our ability to predict the effects of environmental change on root turnover is limited by the difficulty of measuring root dynamics, but emerging evidence suggests that roots, like leaves, possess suites of interrelated traits that are linked to their life span. In graminoids, high tissue density has been linked to increased root longevity. Other studies have found root longevity to be positively correlated with mycorrhizal colonization and negatively correlated with nitrogen concentration, root maintenance respiration and specific root length. Among fruit trees, apple roots (which are of relatively small diameter, low tissue density and have little lignification of the exodermis) have much shorter life spans than the roots of citrus, which have opposite traits. Likewise, within the branched network of the fine root system, the finest roots with no daughter roots tend to have higher N concentrations, faster maintenance respiration, higher specific root length and shorter life spans than secondary and tertiary roots that bear daughter roots. Mycorrhizal colonization can enhance root longevity by diverse mechanisms, including enhanced tolerance of drying soil and enhanced defence against root pathogens. Many variables involved in building roots might affect root longevity, including root diameter, tissue density, N concentration, mycorrhizal fungal colonization and accumulation of secondary phenolic compounds. These root traits are highly plastic and are strongly affected by resource supply (CO2, N, P and water). Therefore the response of root longevity to altered resource availability associated with climate change can be estimated by considering how changes in resource availability affect root construction and physiology. A cost–benefit approach to predicting root longevity assumes that a plant maintains a root only until the efficiency of resource acquisition is maximized. Using an efficiency model, we show that reduced tissue Nconcentration and reduced root maintenance respiration, both of which are predicted to result from elevated CO2, should lead to slightly longer root life spans. Complex interactions with soil biota and shifts in plant defences against root herbivory and parasitism, which are not included in the present efficiency model, might alter the effects of future climate change on root longevity in unpredicted ways.

We review four hypotheses for the control of carbon acquisition by roots, and conclude that the functional equilibrium hypothesis can offer a good description of C acquisition by roots relative to shoots, but is deficient mechanistically. The hypothesis that import into roots is solely dependent on export from the shoot, itself determined by features of the shoot alone (the ‘push’ hypothesis), is supported by some but not all the evidence. Similarly, the idea that root demand, a function of the root alone, determines import into it (the ‘pull’ hypothesis), is consonant with some of the evidence. The fourth, general, hypothesis (the ‘shared control’ hypothesis) – that acquisition of C by roots is controlled by a range of variables distributed between root and shoot – accords with both experiment and theory. Top-down metabolic control analysis quantifies the control of C flux attributable to root relative to source leaf. We demonstrate that two levels of mechanistic control, short-term regulation of phloem transport and control of gene expression by compounds such as sugars, underlie distributed control. Implications for the impact of climate change variables are briefly discussed.

Growth of crops in CO2-enriched atmospheres typically results in significant changes in root growth and development. Increased root carbohydrates stimulate root growth either directly (functioning as substrates) or indirectly (functioning as signal molecules) by enhancing cell division or cell expansion, or both. Although highly variable, the literature suggests that, generally, initiation and stimulation of lateral roots is favored over the elongation of primary roots, leading to more highly branched, shallower root systems. Such architectural shifts can render root systems less efficient, perhaps contributing to the lower specific root activities often reported. Allocation of carbon (C) to roots fluctuates through the life of the plant; root functional and growth responses should therefore not be viewed as static. In annual crops, C allocation to belowground processes changes as vegetative growth switches to reproduction and maturation. Reductions in C allocation to roots over time might cause temporal shifts in root deployment, perhaps affecting root demography. However, significant changes in root turnover (defined here as root flux or mortality relative to total root pool size) as a result of decreased root longevities in crop plants are unlikely. Consideration of changing C allocation to roots, a more thorough understanding of the mechanistic controls on root longevity, and a better characterization of the rooting habits (life histories) of different crop species will further our understanding of how increasing atmospheric [CO2] will affect root demography. This knowledge will lead the way toward a more thorough understanding of the linkage of atmosphere with belowground plant function and also that of plant function with soil biology and structure. Ultimately, successful modeling of global C and nitrogen (N) cycles will require empirical data concerning spatial and temporal deployment of roots for a range of crop species grown under different agricultural management systems.

The objectives of this paper were to review the literature on the responses of root systems to elevated CO2 in intact, native grassland ecosystems, and to present the results from a 2-yr study of root production and mortality in an intact calcareous grassland in Switzerland. Previous work in intact native grassland systems has revealed that significant stimulation of the size of root systems (biomass, length density or root number) is not a universal response to elevated CO2. Of the 12 studies reviewed, seven showed little or no change in root-system size under elevated CO2, while five showed marked increases (average increase 38%). Insufficient data are available on the effects of elevated CO2 on root production, mortality and life span to allow generalization about effects. The diversity of experimental techniques employed in these native grassland studies also makes generalization difficult. In the present study, root production and mortality were monitored in situ in a species-rich calcareous grassland community using minirhizotrons in order to test the hypothesis that an increase in these two measures would help explain the increase in net ecosystem CO2 uptake (net ecosystem exchange) previously observed under elevated CO2 at this site (600 vs 350 μl CO2 l−1; eight 1.2-m2 experimental plots per CO2 level using the screen-aided CO2 control method). However, results from the first 2 yr showed no difference in overall root production or mortality in the top 18 cm of soil, where 80–90% of the roots occur. Elevated CO2 was associated with an upward shift in root length density: under elevated CO2 a greater proportion of roots were found in the upper 0–6-cm soil layer, and a lower proportion of roots in the lower 12–18 cm, than under ambient CO2. Elevated CO2 was also associated with an increase in root survival probability (RSP; e.g. for roots still alive 280 d after they were produced under ambient CO2, RSP = 0.30; elevated CO2, RSP = 0.56) and an increase (48%) in median root life span in the deepest (12–18 cm) soil layer. The factors driving changes in root distribution and longevity with depth under elevated CO2 were not clear, but might have been related to increases in soil moisture under elevated CO2 interacting with vertical patterns in soil temperatures. Thus extra CO2 taken up in this grassland ecosystem during the growing season under elevated CO2 could not be explained by changes in root production and mortality. However, C and nutrient cycling might be shifted closer to the soil surface, which could potentially have a substantial effect on the activities of soil heterotrophic organisms as CO2 levels rise.

Elevated CO2 increases root growth and fine (diam. [les ]2 mm) root growth across a range of species and experimental conditions. However, there is no clear evidence that elevated CO2 changes the proportion of C allocated to root biomass, measured as either the root[ratio ]shoot ratio or the fine root[ratio ]needle ratio. Elevated CO2 tends to increase mycorrhizal infection, colonization and the amount of extramatrical hyphae, supporting their key role in aiding the plant to more intensively exploit soil resources, providing a route for increased C sequestration. Only two studies have determined the effects of elevated CO2 on conifer fine-root life span, and there is no clear trend. Elevated CO2 increases the absolute fine-root turnover rates; however, the standing crop root biomass is also greater, and the effect of elevated CO2 on relative turnover rates (turnover[ratio ]biomass) ranges from an increase to a decrease. At the ecosystem level these changes could lead to increased C storage in roots. Increased fine-root production coupled with increased absolute turnover rates could also lead to increases in soil organic C as greater amounts of fine roots die and decompose. Although CO2 can stimulate fine-root growth, it is not known if this stimulation persists over time. Modeling studies suggest that a doubling of the atmospheric CO2 concentration initially increases biomass, but this stimulation declines with the response to elevated CO2 because increases in assimilation are not matched by increases in nutrient supply.

Soil temperature can influence the functioning of roots in many ways. If soil moisture and nutrient availability are adequate, rates of root length extension and root mortality increase with increasing soil temperature, at least up to an optimal temperature for root growth, which seems to vary among taxa. Root growth and root mortality are highly seasonal in perennial plants, with a flush of growth in spring and significant mortality in the fall. At present we do not understand whether root growth phenology responds to the same temperature cues that are known to control shoot growth. We also do not understand whether the flush of root growth in the spring depends on the utilization of stored nonstructural carbohydrates, or if it is fueled by current photosynthate. Root respiration increases exponentially with temperature, but Q10 values range widely from c. 1.5 to > 3.0. Significant questions yet to be resolved are: whether rates of root respiration acclimate to soil temperature, and what mechanisms control acclimation if it occurs. Limited data suggest that fine roots depend heavily on the import of new carbon (C) from the canopy during the growing season. We hypothesize that root growth and root respiration are tightly linked to whole-canopy assimilation through complex source–sink relationships within the plant. Our understanding of how the whole plant responds to dynamic changes in soil temperature, moisture and nutrient availability is poor, even though it is well known that multiple growth-limiting resources change simultaneously through time during a typical growing season. We review the interactions between soil temperature and other growth-limiting factors to illustrate how simple generalizations about temperature and root functioning can be misleading.

How ecosystems adapt to climate changes depends in part on how individual trees allocate resources to their components. A review of research using tree seedlings provides some support for the hypothesis that some tree species respond to exposure to drought with increases in root[ratio ]shoot ratios but little change in total root biomass. Limited research on mature trees over moderately long time periods (2–10 yr), has given mixed results with some studies also providing evidence for increases in root: shoot ratios. The Throughfall Displacement Experiment (TDE) was designed to simulate both an increase and a decrease of 33% in water inputs to a mature deciduous forest over a number of years. Belowground research on TDE was designed to examine four hypothesized responses to long-term decreases in water availability; (1) increases in fine-root biomass, (2) increases in fine root[ratio ]foliage ratio, (3) altered rates of fine-root turnover (FRT), and (4) depth of rooting. Minirhizotron root elongation data from 1994 to 1998 were examined to evaluate the first three hypotheses. Differences across treatments in net fine-root production (using minirhizotron root elongation observations as indices of biomass production) were small and not significant. Periods of lower root production in the dry treatment were compensated for by higher growth during favorable periods. Although not statistically significant, both the highest production (20 to 60% higher) and mortality (18 to 34% higher) rates were found in the wet treatment, resulting in the highest index of FRT. After 5 yr, a clear picture of stand fine-root-system response to drought exposure has yet to emerge in this forest ecosystem. Our results provide little support for either an increase in net fine-root production or a shift towards an increasing root[ratio ]shoot ratio with long-term drought exposure. One possible explanation for higher FRT rates in the wet treatment could be a positive relationship between FRT and nitrogen and other nutrient availability, as treatments have apparently resulted in increased immobilization of nutrients in the forest floor litter under drier conditions. Such hypotheses point to the continued need to study the interactions of water stress, nutrient availability and carbon-fixation efficiency in future long-term studies.

Temperate forests are recipients of anthropogenic nitrogen (N) deposition. Because growth in these ecosystems is often limited by N availability, elevated N inputs from the atmosphere can influence above- and belowground production in forests. Although fine-root production is the largest component of belowground production in forests, it is unclear whether or how increases in Navailability to forest trees accompanying increased N deposition might influence fine-root growth. Uncertainties as to how fine-root dynamics (i.e. production and turnover) vary in relation to soil N availability contribute to this problem. Although fine-root biomass typically decreases along soil N availability gradients in forests, it is unclear whether fine-root production and turnover also decrease along these gradients. Here, four possible relationships between fine-root turnover, fine-root production, and forest soil N availability are evaluated to develop a general hypothesis about changes in rooting dynamics that might accompany increases in N deposition. The four possible relationships are as follows. (1) Fine-root turnover rates do not systematically change with N availability in forest soils. If this is true, then fine-root production rates decrease with fine-root biomass in relation to soil N availability, and increased N deposition could lead to decreased fine-root production in forests. (2) Decreases in photosynthate allocation belowground along N availability gradients will function to slow fine-root turnover (or increase life span) as N availability increases with N deposition, thereby dramatically decreasing fine-root production. (3) Fine-root production might increase with N availability even though fine-root biomass typically decreases with N availability. This could occur if fine-root metabolism and turnover increase (life span decreases) with soil N supply. Increases in fine-root production accompanying increases in N availability, if large enough, could result in constant proportions of forest production being allocated to fine roots as soil N availability increases with N deposition. (4) Although fine-root turnover and production might both increase as N becomes more available to tree roots, the proportional allocation of total primary production to fine roots could decrease. Identifying the most likely of these four possibilities requires intersite comparisons of forest root dynamics along gradients of soil N availability and N deposition. Collective results of studies that use sequential sampling of fine-root biomass to estimate production suggest that fine-root turnover and production either; do not vary systematically, or that they decrease as N availability increases. By contrast, studies using ecosystem C or N budgets suggest that fine-root turnover and production both increase with N availability and that similar increases might be expected with elevated N deposition. It is argued here that assumptions underlying most biomass-based estimates of fine-root production are more suspect than are assumptions underlying element budget-based estimates. If so, it is likely that N deposition will function to decrease forest fine-root biomass but to stimulate fine-root turnover and production. However, increases in fine- root turnover and production could eventually decrease if chronically elevated N deposition leads to forest stand mortality.

Global warming over the next century is likely to be associated with a change in the extent to which atmospheric and soil temperatures fluctuate, on both a daily and a seasonal basis. The average annual temperature of the Earth's surface is expected to increase, as is the frequency of hot days. In this review, we explore what effects short-term and long-term changes in temperature are likely to have on root respiratory metabolism, and what impacts such changes will have on daily, seasonal and annual CO2 release by roots under field conditions. We demonstrate that Q10 values, and the degree of acclimation, differ between and within plant species. Changes in the temperature sensitivity of respiration with measuring temperature are highlighted. Temperature-dependent changes in adenylate control and substrate supply are likely to control the Q10 and degree of acclimation of root respiration. Limitations in respiration capacity are unlikely to control respiratory flux at most temperatures. The potential role of nonphosphorylating pathways such as the alternative oxidase in controlling Q10 values is highlighted. The possibility that potentially rapid changes in adenylate control might underlie the acclimation response (rather than slow changes in enzyme capacity) has implications for the total amount of CO2 respired by roots daily and annually. Our modelling suggests that rapid acclimation will result in near-perfect homeostasis of respiration rates and minimize annual CO2 release. However, annual CO2 release increases substantially if the speed of full acclimation is lower. Our modelling exercise also shows that high Q10 values have the potential to increase daily and annual CO2 release substantially, particularly if the frequency of hot days increases after global warming.

There is a growing recognition that accurate predictions of plant and ecosystem responses to global change require a better understanding of the mechanisms that control acquisition of growth-limiting resources. One such key mechanism is root physiological capacity to acquire nutrients. Changes in kinetics of root nitrogen (N) uptake might influence the extent to which terrestrial ecosystems will be able to sequester excesses in carbon (C) and N loads. Despite its significant role in determining plant and ecosystem cycling of C and N, there is little information on whether, or how, root nutrient uptake responds to global change. In this review various components of global change, namely increased CO2 concentration, increased soil temperature and increased atmospheric N deposition and their effects on kinetics of root nutrient uptake are examined. The response of root nutrient uptake kinetics to high CO2 is highly variable. Most of this variability might be attributable to differences in experimental protocols, but more recent evidence suggests that kinetic responses to high CO2 are also species-specific. This raises the possibility that elevated CO2 might alter community composition by shifting the competitive interaction of co-occurring species. Uptake of NH4+ and NO3− seem to be differentially sensitive to high CO2, which could influence ecosystem trajectory toward N saturation. Increased soil temperature might increase N and P uptake capacity to a greater extent in species from warm and fluctuating soil habitats than in species from cold and stable soil environments. The few available data also indicate that increased soil temperature elicits a differential effect on uptake of NH4+ versus NO3−. Root uptake kinetics are generally down-regulated in response to long-term exposure to atmospheric N deposition. The extent of this down-regulation might, however, vary among species, stages of succession, land-use history and plant demand. Nonetheless, it is suggested that root N uptake kinetics might be an accurate biological indicator of the ecosystem capacity to retain N. The results reviewed here clearly highlight the scanty nature of the literature in the area of root nutrient absorption responses to global change. It is also clear that effects of one component of global change on root nutrient absorption capacity might be counterbalanced by another. Therefore, the generalizations offered here must be viewed with caution and more effort should be directed to rigorously test these initial observations in future research.

Defining root death in studies of root dynamics is problematic because cell death occurs gradually and the resulting effects on root function are not well understood. In this study, metabolic activity of grape roots of different ages was assessed by excised root respiration and tetrazolium chloride reduction. We investigated changes in metabolic activity and patterns of cell death occurring with root age and changes in root pigmentation. Tetrazolium chloride reduction of roots of different ages was strongly correlated to respiration (R2 = 0.786). As roots aged, respiration and tetrazolium chloride reduction declined similarly, with minimum metabolic activity reached at six weeks. Tetrazolium chloride reduction indicated that the onset of root browning corresponded to a 77% reduction in metabolic activity (P < 0.001). Anatomical examination of roots at each pigmentation stage showed that even though some cells in brown roots were still alive, these roots were functionally dead. The effect of using different definitions of root death in relation to root survivorship was determined in a study of ‘Concord’ grapes with two pruning treatments, using three criteria for root death: browning, blackening or shriveling, and disappearance. There was no effect of vine pruning on root life span when life span was defined as the time from first appearance to the onset of browning. However, if death was judged as the point when roots either became black or shriveled or disappeared, vine pruning decreased root life span by 34% and 40%, respectively (P < 0.001), and also increased the decay constant for root decomposition by about 45% (P < 0.001). We conclude that the discrepancy among determinations of root life span assessed with different definitions of death might be partly caused by the latter evaluations of root life span incorporating a portion of root decomposition in definitions of root death.

Arbuscular mycorrhizal (AM) symbioses are a potentially important link in the chain of response of ecosystems to elevated atmospheric [CO2]. By promoting plant phosphorus uptake and acting as a sink for plant carbon, they can alleviate photosynthetic down-regulation. Because hyphal turnover is likely to be fast, especially in warmer soils, they can also act as a rapid pathway for the return of carbon to the atmosphere. However, most experiments on AM responses to [CO2] have failed to take into account the difference in growth of mycorrhizal and non- mycorrhizal plants; those that have done so suggest that AM colonization of roots is little altered by [CO2], although this issue remains to be resolved. Very little is known about the effects of other factors of global environmental change on mycorrhizas. These issues need urgent attention. It is also necessary to understand the potential for the various AM fungal taxa to respond differentially to environmental changes, including carbon supply and soil temperature and moisture, especially because of the differential abilities of plant and fungal species to migrate in response to changing environments. Indeed, there is a need for a new approach to the study of mycorrhizal associations, which has been too plant-centred. It is essential to regard the fungus as an organism itself, and to understand its biology both as an entity and as part of a symbiosis.

In this review, we discuss the potential for mycorrhizal fungi to act as a source or sink for carbon (C) under elevated CO2 and nitrogen deposition. Mycorrhizal tissue has been estimated to comprise a significant fraction of soil organic matter and below-ground biomass in a range of systems. The current body of literature indicates that in many systems exposed to elevated CO2, mycorrhizal fungi might sequester increased amounts of C in living, dead and residual hyphal biomass in the soil. Through this process, the fungi might serve as a negative feedback on the rise in atmospheric CO2 levels caused by fossil fuel burning and deforestation. By contrast, a few preliminary studies suggest that N deposition might increase turnover rates of fungal tissue and negate CO2 effects on hyphal biomass. If these latter responses are consistent among ecosystems, C storage in hyphae might decline in habitats surrounding agricultural and urban areas. When N additions occur without CO2 enrichment, effects on mycorrhizal growth are inconsistent. We note that analyses of hyphal decomposition under elevated CO2 and N additions are extremely sparse but are critical in our understanding of the impact of global change on the cycling of mycorrhizal C. Finally, shifts in the community composition of arbuscular and ectomycorrhizal fungi with increasing CO2 or N availability are frequently documented. Since mycorrhizal groups vary in growth rate and tissue quality, these changes in species assemblages could produce unforeseeable impacts on the productivity, survivorship, or decomposition of mycorrhizal biomass.

There is considerable uncertainty about how rates of soil carbon (C) and nitrogen (N) cycling will change as CO2 accumulates in the Earth's atmosphere. We summarized data from 47 published reports on soil C and N cycling under elevated CO2 in an attempt to generalize whether rates will increase, decrease, or not change. Our synthesis centres on changes in soil respiration, microbial respiration, microbial biomass, gross N mineralization, microbial immobilization and net N mineralization, because these pools and processes represent important control points for the below-ground flow of C and N. To determine whether differences in C allocation between plant life forms influence soil C and N cycling in a predictable manner, we summarized responses beneath graminoid, herbaceous and woody plants grown under ambient and elevated atmospheric CO2. The below-ground pools and processes that we summarized are characterized by a high degree of variability (coefficient of variation 80–800%), making generalizations within and between plant life forms difficult. With few exceptions, rates of soil and microbial respiration were more rapid under elevated CO2, indicating that (1) greater plant growth under elevated CO2 enhanced the amount of C entering the soil, and (2) additional substrate was being metabolized by soil microorganisms. However, microbial biomass, gross N mineralization, microbial immobilization and net N mineralization are characterized by large increases and declines under elevated CO2, contributing to a high degree of variability within and between plant life forms. From this analysis we conclude that there are insufficient data to predict how microbial activity and rates of soil C and N cycling will change as the atmospheric CO2 concentration continues to rise. We argue that current gaps in our understanding of fine-root biology limit our ability to predict the response of soil microorganisms to rising atmospheric CO2, and that understanding differences in fine-root longevity and biochemistry between plant species are necessary for developing a predictive model of soil C and N cycling under elevated CO2.

The representation of root activity in models is here confined to considerations of applications assessing the impacts of changes in climate or atmospheric [CO2]. Approaches to modelling roots can be classified into four major types: models in which roots are not considered, models in which there is an interplay between only selected above-ground and below-ground processes, models in which growth allocation to all parts of the plants depends on the availability and matching of the capture of external resources, and models with explicit treatments of root growth, architecture and resource capture. All models seem effective in describing the major root activities of water and nutrient uptake, because these processes are highly correlated, particularly at large scales and with slow or equilibrium dynamics. Allocation models can be effective in providing a deeper, perhaps contrary, understanding of the dynamic underpinning to observations made only above ground. The complex and explicit treatment of roots can be achieved only in small-scale highly studied systems because of the requirements for many initialized variables to run the models.