A Coordination Model of Whole-plant Carbon Allocation in Relation to Water Stress

Although water is an important determinant of the allocationof material between roots and shoots during growth, and oftenparallels the effects of nitrogen, few models have explicitlyconsidered allocation in relation to water supply. We use coordinationtheory to develop a simple exponential model that considersallocation of dry matter between shoots and roots during growthin relation to carbon and watersupplies, and accounts for theeffects of water stress on growth. We compare coordinationvs.optimization(global and local) versions of the exponential model by examiningsimilarities and differences in model behaviour obtained underconstant and variable environmental conditions, and with drasticallychanging conditions (mild, moderate and severe water stress).The greatest differences between coordination and optimizationexist in the drastically changing conditions. In a second versionof the model, we remove the restriction of exponential growthand show how coordination principles can be extended to a morecomplicated structure. The non-exponential model is used toanalyse experimental data on the effects of different pot sizes(and hence water availability) on root restriction and plantgrowth as reported by Thomas and Strain (Plant Physiology96:627–634, 1991). With further refinements, the coordinationmodel has potential as a tool to model plant growth in relationto water supply under various environmental conditions. Optimization; functional balance; root: shoot ratio; root restriction     

Continental‐scale nitrogen pollution is shifting forest mycorrhizal associations and soil carbon stocks

Most tree roots on Earth form a symbiosis with either ecto‐ or arbuscular mycorrhizal fungi. Nitrogen fertilization is hypothesized to favor arbuscular mycorrhizal tree species at the expense of ectomycorrhizal species due to differences in fungal nitrogen acquisition strategies, and this may alter soil carbon balance, as differences in forest mycorrhizal associations are linked to differences in soil carbon pools. Combining nitrogen deposition data with continental‐scale US forest data, we show that nitrogen pollution is spatially associated with a decline in ectomycorrhizal vs. arbuscular mycorrhizal trees. Furthermore, nitrogen deposition has contrasting effects on arbuscular vs. ectomycorrhizal demographic processes, favoring arbuscular mycorrhizal trees at the expense of ectomycorrhizal trees, and is spatially correlated with reduced soil carbon stocks. This implies future changes in nitrogen deposition may alter the capacity of forests to sequester carbon and offset climate change via interactions with the forest microbiome.     

Response of tree biomass and wood litter to disturbance in a Central Amazon forest

We developed an individual-based stochastic-empirical model to simulate the carbon dynamics of live and dead trees in a Central Amazon forest near Manaus, Brazil. The model is based on analyses of extensive field studies carried out on permanent forest inventory plots, and syntheses of published studies. New analyses included: (1) growth suppression of small trees, (2) maximum size (trunk base diameter) for 220 tree species, (3) the relationship between growth rate and wood density, and (4) the growth response of surviving trees to catastrophic mortality (from logging). The model simulates a forest inventory plot, and tracks recruitment, growth, and mortality of live trees, decomposition of dead trees (coarse litter), and how these processes vary with changing environmental conditions. Model predictions were tested against aggregated field data, and also compared with independent measurements including maximum tree age and coarse litter standing stocks. Spatial analyses demonstrated that a plot size of ~10 ha was required to accurately measure wood (live and dead) carbon balance. With the model accurately predicting relevant pools and fluxes, a number of model experiments were performed to predict forest carbon balance response to perturbations including: (1) increased productivity due to CO₂ fertilization, (2) a single semi-catastrophic (10%) mortality event, (3) increased recruitment and mortality (turnover) rates, and (4) the combined effects of increased turnover, increased tree growth rates, and decreased mean wood density of new recruits. Results demonstrated that carbon accumulation over the past few decades observed on tropical forest inventory plots (~0.5 Mg C ha^–1 year^–1) is not likely caused by CO₂ fertilization. A maximum 25% increase in woody tissue productivity with a doubling of atmospheric CO₂ only resulted in an accumulation rate of 0.05 Mg C ha^–1 year^–1 for the period 1980–2020 for a Central Amazon forest, or an order of magnitude less than observed on the inventory plots. In contrast, model parameterization based on extensive data from a logging experiment demonstrated a rapid increase in tree growth following disturbance, which could be misinterpreted as carbon sequestration if changes in coarse litter stocks were not considered. Combined results demonstrated that predictions of changes in forest carbon balance during the twenty-first century are highly dependent on assumptions of tree response to various perturbations, and underscores the importance of a close coupling of model and field investigations.     

Repeated insect defoliation effects on growth, nitrogen acquisition, carbohydrates, and root demography of poplars

Large-scale outbreaks of defoliating insects are common in temperate forests. The effects of defoliation on tree physiology are expected to cascade through the entire forest ecosystem, altering carbon, nitrogen, and water fluxes, and subsequently affecting nitrogen cycling and plant-herbivore interactions. If these post-defoliation changes are largely driven by N deficiency, tree root system responses to defoliation should be central to regulating the long-term effects of defoliation; N fertilization should reverse the effects. We examined these phenomena in a 3-year large-scale replicated manipulative field experiment in a hybrid poplar plantation, where we regulated defoliation by gypsy moths as well as nitrogen availability. To our knowledge, this is the first manipulative field experiment at this scale to examine the effects of severe insect defoliation on whole-tree physiology. Defoliation decreased tree growth and increased the rate of top dieback in the stand. Defoliation led to transient declines in carbon allocation to starch in fine roots, trunk, and twigs in the year of heaviest defoliation. Root production and root mortality were unaffected by the heaviest defoliation, but nitrate and ammonium uptake were strongly depressed. N fertilization increased tree growth, but did not alter defoliation effects on starch accumulation or top dieback. Defoliation and fertilization treatments did not interact. In this system, defoliation effects on tree recovery of leaf nitrogen lost to herbivory were primarily driven by effects on nitrogen uptake, rather than effects on root production or mortality.     

高浓度CO2对树木生理生态的影响研究进展

大气CO2浓度升高已成为世界范围内的重要环境问题。CO2浓度升高势必会对植物的生理生态变化产生重要影响。综述了国内外有关高浓度CO2对树木生理生态影响研究的最新进展,具体包括高浓度CO2对树木生长发育、光合和呼吸作用、抗氧化系统、树木代谢物质、挥发性有机化合物以及树木凋落物等方面的影响。高浓度CO2一般会促进树木地上植株的生长和发育,但也因树种差异而有所不同。最新研究表明,高浓度CO2促进了树木细根周转,树木根系生长在大气CO2浓度升高条件下表现为促进作用,这种作用加快了全球森林生态系统的C循环。高浓度CO2虽然在一定程度上促进树木光合速率的增加,但长期熏蒸也往往会发生光合驯化,这种现象产生的生理学机制目前仍无定论。高浓度CO2对树木呼吸作用尤其是根系呼吸的影响将是未来研究的重点和难点。高浓度CO2一般会提高树木抗氧化酶活性与抗氧化剂含量,但不同树种响应高浓度CO2的过程和机理也有所差异。研究表明,高浓度CO2一般对树木凋落物的分解产生不利影响,但也因树种而异。需要强调的是,目前关于树木地下部分、树木对高浓度CO2的适应机理和重要过程(碳氮水耦合及基因调控等)以及多个树种包括不同类型树种及不同品种之间比较研究较少;关于某一重要生理生态机制(如根系生理代谢)尤其是多个生态因子复合条件下缺乏长期深入的研究。在此基础上给出了大气CO2浓度升高下树木生理生态学研究的未来发展方向,包括高CO2浓度条件下树木根系生理代谢及机制、树木碳氮水耦合的生理过程及机制、不同生态因子复合作用对树木生理影响机制以及树木分子作用机理等方面的研究。这些研究不仅将丰富森林树木应对未来气候变化的有关科学理论,也为全球气候变化背景下实现森林树种生态功能的优化选择及森林生态系统的可持续发展与经营提供重要的生理生态学理论依据和参考。     

Climate change in forest ecosystems: A field experiment addressing the effects of raising temperature and reduced rainfall on early life cycle stages of oaks

Higher temperatures and reduced rainfalls that are expected with the advance of climate change can impair the emergence and establishment of tree seedlings in forest ecosystems. These climatic changes can also decrease the availability of soil resources and reduce the performance of seedlings. We evaluated these effects in a temperate forest from Mexico with two native oak species (Quercus crassifolia and Quercus eduardii). As recently emerged oak seedlings are highly sensitive to changing environmental conditions, our field experiment was conducted across the season in which seedling emergence occurs (October–February). In the field, we used open-top chambers to increase temperature and rainout shelters to reduce rainfall, while controls were exposed to the current climate. Experimental plots of both treatments were established beneath the forest canopy because most oaks recruit in understory habitats. In these plots, we sowed acorns of both species in October 2015 and recorded seedling emergence and survival until February 2016, also monitoring temperature, precipitation and contents of water and nitrogen in the soil. On seedlings that survived until the end of the experiment we measured their growth, photosynthetic efficiency and foliar contents of water, carbon and nitrogen. Both the emergence and survival of Q. crassifolia seedlings were lower in climate change plots than in controls, but no differences were found for Q. eduardii. However, seedlings of both species had lower growth rates, photosynthetic efficiencies and contents of water, nitrogen and carbon in climate change simulation plots. These results indicate that climate change can impair tree seedling establishment in oak forest, also suggesting that their development will be constrained by reduced water and nitrogen availability.     

Ectomycorrhizal fungi slow soil carbon cycling

Respiration of soil organic carbon is one of the largest fluxes of CO₂ on earth. Understanding the processes that regulate soil respiration is critical for predicting future climate. Recent work has suggested that soil carbon respiration may be reduced by competition for nitrogen between symbiotic ectomycorrhizal fungi that associate with plant roots and free‐living microbial decomposers, which is consistent with increased soil carbon storage in ectomycorrhizal ecosystems globally. However, experimental tests of the mycorrhizal competition hypothesis are lacking. Here we show that ectomycorrhizal roots and hyphae decrease soil carbon respiration rates by up to 67% under field conditions in two separate field exclusion experiments, and this likely occurs via competition for soil nitrogen, an effect larger than 2 °C soil warming. These findings support mycorrhizal competition for nitrogen as an independent driver of soil carbon balance and demonstrate the need to understand microbial community interactions to predict ecosystem feedbacks to global climate.     

Bioengineering ground improvement considering root water uptake model

Bioengineering features of native vegetation are currently being evolved to enhance soil stiffness, slope stabilisation and erosion control. The effects of tree roots on soil moisture content and ground settlement are discussed in this paper. Matric suction induced by tree roots is a key factor, governing the properties of unsaturated soils, directly imparting stability to slopes and resistance for yielding behaviour. A mathematical model for the rate of root water uptake that considers ground conditions, type of vegetation and climatic parameters has been developed. This study highlights the inter-related parameters contributing to the development of a conceptual evapo-transpiration and root moisture uptake equilibrium model that is then incorporated in a comprehensive numerical finite element model. The developed model considers fully coupled-flow-deformation behaviour of soil. Field measurements obtained by the Authors from a site in Victoria, South of Australia, are used to validate the model. In this study, the active tree root distribution has been predicted by measuring soil organic content distribution. The predicted results show acceptable agreement with the field data in spite of the assumptions made for simplifying the effects of soil heterogeneity and anisotropy. The results prove that the proposed root water uptake model can reliably predict the region of the maximum matric suction away from the tree axis.     

Toward an allocation scheme for global terrestrial carbon models

The distribution of assimilated carbon among the plant parts has a profound effect on plant growth, and at a larger scale, on terrestrial biogeochemistry. Although important progress has been made in modelling photosynthesis, less effort has been spent on understanding the carbon allocation, especially at large spatial scales. Whereas several individual-level models of plant growth include an allocation scheme, most global terrestrial models still assume constant allocation of net primary production (NPP) among plant parts, without any environmental coupling. Here, we use the CASA biosphere model as a platform for exploring a new global allocation scheme that estimates allocation of photosynthesis products among leaves, stems, and roots depending on resource availability. The philosophy underlying the model is that allocation patterns result from evolved responses that adjust carbon investments to facilitate capture of the most limiting resources, i.e. light, water, and mineral nitrogen. In addition, we allow allocation of NPP to vary in response to changes in atmospheric CO₂. The relative magnitudes of changes in NPP and resource-use efficiency control the response of root:shoot allocation. For ambient CO₂, the model produces realistic changes in above-ground allocation along productivity gradients. In comparison to the CASA standard estimate using fixed allocation ratios, the new allocation scheme tends to favour root allocation, leading to a 10% lower global biomass. Elevated CO₂, which alters the balance between growth and available resources, generally leads to reduced water stress and consequently, decreased root:shoot ratio. The major exception is forest ecosystems, where increased nitrogen stress induces a larger root allocation.     

树木根系衰老研究的意义与现状

树木根系是树木重要的组成部分,具有养分和水分的吸收、传输和储存、树体的固定与支撑等重要的生理功能．在树木根系形成以后,常常遭遇到养分和水分胁迫,因此,其养分和水分的吸收功能尤其重要,在森林土壤中,养分和水分具有很大的时间和空间异质性,随着养分和水分在时间和空间上的变化,树木及时地主动调整其碳在根系中的分配,从而导致部分根系衰老或死亡,在林学上,树木根系衰老与养分和水分吸收关系密切,因而与树木生产力有直接的关系．在生态系统乃至全球尺度上,树木根系衰老影响碳循环和养分循环,因为根系对碳的消耗占树木通过光合作用所固定的碳的比例相当大,且含有丰富的养分．树木根系衰老受许多环境因子的影响,生物因子有真菌、细菌、病毒、土壤小型动物等,非生物因子有水分、温度、土壤养分、重金属等,这些因子对树木根系衰老的影响机制并不相同,尽管在树木根系衰老研究领域取得了长足的进步,提出了许多不同的假设,但仍有许多问题尚未解决,这些假设也需要更多的实验来验证,运用细胞学、生物化学、土壤科学、遗传学等多学科的交叉研究可进一步揭示根系衰老的本质。     

Root water uptake and profile soil water as affected by vertical root distribution

Water uptake by plant roots is a main process controlling water balance in field profiles and vital for agro-ecosystem management. Based on the sap flow measurements for maize plants (Zea mays L.) in a field under natural wet- and dry-soil conditions, we studied the effect of vertical root distribution on root water uptake and the resulted changes of profile soil water. The observations indicate that depth of the most densely rooted soil layer was more important than the maximum rooting depth for increasing the ability of plants to cope with the shortage of water. Occurrence of the most densely rooted layer at or below 30-cm soil depth was very conducive to maintaining plant water supply under the dry-soil conditions. In the soil layers colonized most densely by roots, daytime effective soil water saturation (S _e) always dropped dramatically due to the high-efficient local water depletion. Restriction of the rooting depth markedly increased the difference of S _e between the individual soil layers particularly under the dry-soil conditions due likely to the physical non-equilibrium of water flow between the layers. This study highlights the importance of root distribution and pattern in regulating soil water use and thereby improving endurance of plants to seasonal droughts for sustainable agricultural productivity.     

Fine-root growth in a forested bog is seasonally dynamic,but shallowly distributed in nutrient-poor peat

Background and aims

Fine roots contribute to ecosystem carbon, water, and nutrient fluxes through resource acquisition, respiration, exudation, and turnover, but are understudied in peatlands. We aimed to determine how the amount and timing of fine-root growth in a forested, ombrotrophic bog varied across gradients of vegetation density, peat microtopography, and changes in environmental conditions across the growing season and throughout the peat profile.

Methods

We quantified fine-root peak standing crop and growth using non-destructive minirhizotron technology over a two-year period, focusing on the dominant woody species in the bog: Picea mariana, Larix laricina, Rhododendron groenlandicum, and Chamaedaphne calyculata.

Results

The fine roots of trees and shrubs were concentrated in raised hummock microtopography, with more tree roots associated with greater tree densities and a unimodal peak in shrub roots at intermediate tree densities. Fine-root growth tended to be seasonally dynamic, but shallowly distributed, in a thin layer of nutrient-poor, aerobic peat above the growing season water table level.

Conclusions

The dynamics and distribution of fine roots in this forested ombrotrophic bog varied across space and time in response to biological, edaphic, and climatic conditions, and we expect these relationships to be sensitive to projected environmental changes in northern peatlands.

     

Contingent effects of water balance variation on tree cover density in semiarid woodlands

Aim The local distribution of woody vegetation affects most functional aspects of semiarid landscapes, from soil erosion to nutrient cycling. With growing concern about anthropogenic climate change, it has become critical to understand the ecological determinants of woody plant distribution in semiarid landscapes. However, relatively little work has examined the determinants of local variation in woody cover. Here we examine water balance controls associated with patterns of tree cover density in a topographically complex semiarid woodland. Location Los Pinos Mountains, Sevilleta National Wildlife Refuge LTER, New Mexico, USA. Methods To explore the relationship between local water balance variation and tree cover density, we used a combination of high‐resolution (1 × 1 m), remotely sensed imagery and quantitative estimates of water balance variation in space and time. Regression tree analysis (RTA) was used to identify the environmental parameters that best predict variation in tree cover density. Results Using six predictor variables, the RTA explains 39% of the deviance in tree cover density over the landscape. The relationship between water balance conditions and tree cover density is highly contingent; that is, similar tree cover densities occur under very different combinations of water balance parameters. Thus, the effect of one environmental parameter on tree cover density depends on the values of other parameters. After tree cover density is adjusted for water balance conditions, residual variation is related to tree cover density in the neighbourhood of a particular location. Conclusion In semiarid landscapes, vegetation structure is largely controlled by water supply and demand. Results presented here indicate that localized feedbacks and site‐specific historical processes are critical for understanding the responses of semiarid vegetation to climate change.     

Increased forest carbon storage with increased atmospheric CO2 despite nitrogen limitation: a game‐theoretic allocation model for trees in competition for nitrogen and light

Changes in resource availability often cause competitively driven changes in tree allocation to foliage, wood, and fine roots, either via plastic changes within individuals or through turnover of individuals with differing strategies. Here, we investigate how optimally competitive tree allocation should change in response to elevated atmospheric CO₂ along a gradient of nitrogen and light availability, together with how those changes should affect carbon storage in living biomass. We present a physiologically‐based forest model that includes the primary functions of wood and nitrogen. From a tree's perspective, wood is an offensive and defensive weapon used against neighbors in competition for light. From a biogeochemical perspective, wood is the primary living reservoir of stored carbon. Nitrogen constitutes a tree's photosynthetic machinery and the support systems for that machinery, and its limited availability thus reduces a tree's ability to fix carbon. This model has been previously successful in predicting allocation to foliage, wood, and fine roots along natural productivity gradients. Using game theory, we solve the model for competitively optimal foliage, wood, and fine root allocation strategies for trees in competition for nitrogen and light as a function of CO₂ and nitrogen mineralization rate. Instead of down‐regulating under nitrogen limitation, carbon storage under elevated CO₂ relative to carbon storage at ambient CO₂ is approximately independent of the nitrogen mineralization rate. This surprising prediction is a consequence of both increased competition for nitrogen driving increased fine root biomass and increased competition for light driving increased allocation to wood under elevated CO₂.     

Water shortage affects the water and nitrogen balance in Central European beech forests

Whilst forest policy promotes cultivation and regeneration of beech dominated forest ecosystems, beech itself is a highly drought sensitive tree species likely to suffer from the climatic conditions prognosticated for the current century. Taking advantage of model ecosystems with cool-moist and warm-dry local climate, the latter assumed to be representative for future climatic conditions, the effects of climate and silvicultural treatment (different thinning regimes) on water status, nitrogen balance and growth parameters of adult beech trees and beech regeneration in the understorey were assessed. In addition, validation experiments with beech seedlings were carried out under controlled conditions, mainly in order to assess the effect of drought on the competitive abilities of beech. As measures of water availability xylem flow, shoot water potential, stomatal conductance as well as delta (13)C and delta (18)O in different tissues (leaves, phloem, wood) were analysed. For the assessment of nitrogen balance we determined the uptake of inorganic nitrogen by the roots as well as total N content and soluble N compounds in different tissues of adult and young trees. Retrospective and current analysis of delta (13)C, growth and meteorological parameters revealed that beech growing under warm-dry climatic conditions were impaired in growth and water balance during periods with low rain-fall. Thinning affected water, N balance and growth mostly of young beech, but in a different way under different local climatic conditions. Under cool, moist conditions, representative for the current climatic and edaphic conditions in beech forests of Central Europe, thinning improves nutrient and water status consistent to published literature and long-term experience of forest practitioners. However, beech regeneration was impaired as a result of thinning at higher temperatures and under reduced water availability, as expected in future climate.     

Modeling long-term crop response to fertilizer and soil nitrogen

A simple nitrogen balance model to calculate long-term changes in soil organic nitrogen, nitrogen uptake by the crop and recovery of applied nitrogen, is presented. It functions with time intervals of one year or one growing season. In the model a labile and a stable pool of soil organic nitrogen are distinguished. Transfer coefficients for the various inputs of nitrogen are established that specify the fractions taken up by the crop, lost from the system, and incorporated in soil organic nitrogen. It is shown how input data, model parameters and initial pool sizes can be derived and how the model can be used for calculating long-term changes in total soil organic nitrogen and uptake by the crop. For nitrogen applied annually as fertilizer or organic material the time course of nitrogen uptake and recovery of applied nitrogen is calculated. To test the sensitivity of the model, calculations have been performed for different environmental conditions with higher or lower risks for losses. The model has also been applied to establish fertilizer recommendations for a certain target nitrogen uptake by the crop. Finally, for agricultural systems where periods of cropping alternate with peroids of green fallow the time course of nitrogen uptake by the crop is calculated.     

植物光合产物分配及其影响因子研究进展

植物光合产物分配受环境因子和生物因子的共同影响。为增进对植物对全球变化响应的理解, 从植物个体水平与群落/生态系统水平综述了植物光合产物分配的影响因子与影响机理的最新研究进展。植物个体在光照增强及受水分和养分胁迫时, 会将光合产物更多地分配到根系; CO₂浓度升高对植物光合产物分配的影响受土壤氮素的制约, 植物受氮素胁迫时, CO₂浓度升高会促进光合产物更多地分配到根系; 反之, 对植物光合产物分配没有影响。植物群落/生态系统的光合产物分配对环境因子的响应不敏感; 光合产物向根系的分配比例随其生长阶段逐渐降低。功能平衡假说、源汇关系假说和相关生长关系假说分别从环境因子、个体发育和环境因子与个体发育协同作用方面阐述了植物光合产物分配的影响机理。在此基础上,指出了未来拟重点加强的研究方向: 1)生态系统尺度的光合产物向呼吸部分的分配研究; 2)地下净初级生产力(belowground net primary productivity, BNPP)研究; 3)温室和野外条件下及幼苗和成熟林光合产物分配对环境因子响应的比较研究; 4)生态系统尺度的多因子控制试验; 5)整合环境因子和个体发育对植物光合产物分配格局的影响研究。     

Impacts of climate change on the vegetation of Africa: an adaptive dynamic vegetation modelling approach

Recent IPCC projections suggest that Africa will be subject to particularly severe changes in atmospheric conditions. How the vegetation of Africa and particularly the grassland–savanna–forest complex will respond to these changes has rarely been investigated. Most studies on global carbon cycles use vegetation models that do not adequately account for the complexity of the interactions that shape the distribution of tropical grasslands, savannas and forests. This casts doubt on their ability to reliably simulate the future vegetation of Africa. We present a new vegetation model, the adaptive dynamic global vegetation model (aDGVM) that was specifically developed for tropical vegetation. The aDGVM combines established components from existing DGVMs with novel process‐based and adaptive modules for phenology, carbon allocation and fire within an individual‐based framework. Thus, the model allows vegetation to adapt phenology, allocation and physiology to changing environmental conditions and disturbances in a way not possible in models based on fixed functional types. We used the model to simulate the current vegetation patterns of Africa and found good agreement between model projections and vegetation maps. We simulated vegetation in absence of fire and found that fire suppression strongly influences tree dominance at the regional scale while at a continental scale fire suppression increases biomass in vegetation by a more modest 13%. Simulations under elevated temperature and atmospheric CO₂ concentrations predicted longer growing periods, higher allocation to roots, higher fecundity, more biomass and a dramatic shift toward tree dominated biomes. Our analyses suggest that the CO₂ fertilization effect is not saturated at ambient CO₂ levels and will strongly increase in response to further increases in CO₂ levels. The model provides a general and flexible framework for describing vegetation response to the interactive effects of climate and disturbances.     

Fine roots and ectomycorrhizas as indicators of environmental change

Abstract

Human-induced and natural stress factors can affect fine roots and ectomycorrhizas. Therefore they have potential utility as indicators of environmental change. We evaluated, through meta-analysis, the magnitude of the effects of acidic deposition, nitrogen deposition, increased ozone levels, elevated atmospheric carbon dioxide, and drought on fine roots and ectomycorrhizal (ECM) characteristics. Ectomycorrhizal colonization was an unsuitable parameter for environmental change, but fine root length and biomass could be useful. Acidic deposition had a significantly negative impact on fine roots, root length being more sensitive than root biomass. There were no significant effects of nitrogen deposition or elevated tropospheric ozone on the quantitative root parameters. Elevated CO₂ had a significant positive effect. Drought had a significantly negative effect on fine root biomass. The negative effect of acidic deposition and the positive effect of elevated CO₂ increased over time, indicating that effects were persistent contrary the other factors. The meta-analysis also showed that experimental conditions, including both laboratory and field experiments, were a major source of variation. In addition to quantitative changes, environmental changes affect the species composition of the ectomycorrhizal fungal community.