A comprehensive appraisal of the mycorrhizal literature provides data for 336 plant families representing 99% of flowering plants, with regard to mycorrhizas and other nutritional adaptations. In total, arbuscular (AM), orchid, ectomycorrhizas (EM) and ericoid mycorrhizas and nonmycorrhizal (NM) roots occur in 74%, 9%, 2%, 1% and 6% of Angiosperm species respectively. Many families of NM plants have alternative nutritional strategies such as parasitism, carnivory, or cluster roots. The remaining angiosperms (8%) belong to families reported to have both AM and NM species. These are designated as NM-AM families here and tend to occur in habitats considered non-conducive to mycorrhizal fungi, such as epiphytic, aquatic, extremely cold, dry, disturbed, or saline habitats. Estimated numbers of species in each category of mycorrhizas is presented with lists of NM and EM families. Evolutionary trends are also summarised by providing data on all clades and orders of flowering and non-flowering vascular plants on a global scale. A case study of Western Australian plants revealed that plants with specialised nutritional modes such as carnivory, cluster roots, or EM were much more diverse in this ancient landscape with infertile soils than elsewhere. Detailed information on the mycorrhizal diversity of plants presented here is linked to a website (mycorrhizas.info) to allow data to remain current. Over a century of research effort has resulted in data on mycorrhizal associations of >10,000 plant species that are of great value, but also somewhat of a liability due to conflicting information about some families and genera. It is likely that these conflicts result in part from misdiagnosis of mycorrhizal associations resulting from a lack of standardisation in criteria used to define them. Families that contain both NM and AM species provide a second major source of inconsistency, but even when these are excluded there is a ∼10% apparent error rate in published lists of mycorrhizal plants. Arbuscules are linked to AM misdiagnosis since they are used less often than vesicles to recognise AM associations in roots and apparently occur sporadically in NM plants. Key issues with the diagnosis of mycorrhizal plants are discussed using the Cyperaceae as a case study. Detailed protocols designed to consistently distinguish AM from endophytic Glomeromycotan Fungus Colonisation (GFC) are provided. This review aims to stimulate debate and provide advice to researchers delving into root biology.

The extent to which terrestrial ecosystems can sequester carbon to mitigate climate change is a matter of debate. The stimulation of arbuscular mycorrhizal fungi (AMF) by elevated atmospheric carbon dioxide (CO(2)) has been assumed to be a major mechanism facilitating soil carbon sequestration by increasing carbon inputs to soil and by protecting organic carbon from decomposition via aggregation. We present evidence from four independent microcosm and field experiments demonstrating that CO(2) enhancement of AMF results in considerable soil carbon losses. Our findings challenge the assumption that AMF protect against degradation of organic carbon in soil and raise questions about the current prediction of terrestrial ecosystem carbon balance under future climate-change scenarios.

Ephemeral roots have essential roles in plant and ecosystem functioning. In forests, roots account for a major component of carbon cycling, yet few studies have examined ranges of root trait variation and how different species vary in root form and function in these communities. Root branching intensity, specific root length (SRL; root length per unit dry mass), root diameter, tissue density, phenolic concentration and nitrogen concentration were determined for the finest two root orders of 25 co-existing North American woody species sampled from mature plants in a single forest community. Trait correlations and multivariate patterns were examined to evaluate the most important trait differences among species. Branching intensity, SRL, and phenolic concentration varied most widely among species (coefficient of variation (CV) = 0.42, 0.57 and 0.58, respectively). Species predominately forming ectomycorrhiza (EM) had a higher branching intensity than those forming arbuscular mycorrhiza (AM) with mycorrhizal types correctly predicted in c. 70% of individual observations by branching intensity alone. There was notably no correlation between SRL and nitrogen. Variation in SRL among species mapped partially along phylogenetic lines (consistency index (CI) = 0.44), with remaining variation attributable to differences in species' ecological specialization. Variation found in root traits suggests different nutrient acquisition strategies within this community, which could have potential species-level effects on carbon and mineral nutrient cycling.

1. Stem xylem characteristics have a great impact on growth and adult stature of trees because of their role in mechanical support, long-distance water transport and whole-plant carbon allocation. Yet, despite the potential causal link between xylem traits and plant growth/adult stature, most studies have tried to link wood density, an indirect but easy to measure proxy for wood properties, to tree growth and size.
2. To determine whether xylem traits outperform wood density as predictors of tree growth and stature, we evaluated the covariation among wood density, xylem anatomical traits, tree diameter growth rate and adult stature in 40 Asian tropical tree species through principal component analyses and through bivariate correlation, both across species and across phylogenetically independent contrasts.
3. Vessel diameter exhibited a tight negative relationship with vessel frequency. Wood density showed a significant correlation with vessel diameter and density, but not with vessel fraction. Most correlations between functional traits indicate adaptive coordination, demonstrated by significant correlations between phylogenetically independent contrasts.
4. Across species, diameter growth rate and adult stature were positively correlated with vessel lumen diameter and potential hydraulic conductivity, but not with wood density. Thus, our results suggest that xylem anatomical traits that are linked to hydraulic conductivity are better predictors of tree growth rate and adult stature than wood density.
5. Synthesis. We found that xylem anatomical traits have a more significant influence on whole-plant performance due to their direct association with stem hydraulic conductivity, whereas wood density is decoupled from hydraulic function due to complex variations in xylem components.

Ectomycorrhizal fungal tissues comprise a significant forest-litter pool. Ectomycorrhizal (EM) fungi may also influence the decomposition of other forest-litter components via competitive interactions with decomposer fungi and by ensheathing fine roots. Because of these direct and indirect effects of ectomycorrhizal fungi, the factors that control the decomposition of EM fungi will strongly control forest-litter decomposition as a whole and, thus, ecosystem nutrient and carbon cycling. Some have suggested that chitin, a component of fungal cell walls, reduces fungal tissue decomposition because it is relatively recalcitrant. We therefore examined the change in chitin concentrations of EM fungal tissues during decomposition. Our results show that chitin is not recalcitrant relative to other compounds in fungal tissues and that its concentration is positively related to the decomposition of fungal tissues. Variation existing among EM fungal isolates in chitin concentration suggests that EM fungal community structure influences C and nutrient cycling.

Fine root survivorship is an important aspect of root ecology and is known to be influenced by a suite of covariates. However, the relative importance of each covariate on root survivorship is not clear. Here, we used minirhizotron-based data from 18 woody species to evaluate the relative strength of influence on root survivorship by root diameter, branch order, soil depth, and season of root birth, and to examine how the relationship between each covariate and root survivorship differed across species. We extracted hazard ratio estimates for 16 species from published studies that performed Cox proportional hazards regression analysis, and from our own unpublished data for two Chinese temperate tree species. The mean change in hazard ratio (CHR) and corresponding coefficient of variation for each factor were calculated across species. On average, root diameter and season of root birth had stronger effects on root survivorship than branch order and soil depth. However, the effects of season varied with species and were stronger in temperate forests, whereas the influence of diameter and order were relatively consistent across species. These results suggest that root structures such as diameter and branch order should be carefully considered in root classification and sampling, and adding season of birth (particularly in temperate forests) as a covariate in root survivorship analysis should further enhance our ability in achieving a better understanding of root demography and more accurate estimates of root turnover in forests of different climate zones.

1. Below-ground plant functional traits regulate plant soil interactions and may therefore strongly influence ecosystem responses to global change. Despite this, knowledge of how fine-root functional traits vary among plant species and along environmental gradients has lagged far behind our understanding of above-ground traits.
2. We measured species- and community-level root and leaf trait responses for 50 temperate rain forest species from 28 families of ferns, woody and herbaceous angiosperms and conifers, along a soil chronosequence in New Zealand that exhibits a strong gradient in soil nutrient availability. Relationships among species traits (both above- and below-ground) and their distribution along the chronosequence were tested using phylogenetic generalized least-squares regression to account for plant relatedness.
3. Distinctive root trait syndromes were observed; they were closely linked to species' distribution along the chronosequence. Species growing in the strongly P-limited late stages of the chronosequence had relatively high specific root length (SRL), thin root diameter, high root tissue density, high levels of root branching and low root nutrient concentrations compared to intermediate stages. Species on the youngest site also had high SRL, but had low root tissue density, thick root diameter and high root nutrient concentrations.
4. Species root and leaf nutrient concentrations were positively correlated, reflecting the strong underlying gradient in soil fertility. In contrast, the relationship between SRL and SLA was more complex; there was a weak positive correlation between SRL and SLA, but this conflicted with stronger patterns of increasing SRL and declining SLA with increasing site age.
5. Community-averaged trait values calculated using presence/absence data showed similar trends to the species-level patterns. In contrast, community averages calculated using species abundance-weighted data showed weaker relationships with site age, particularly for morphological traits. This suggests that much of the variation in morphological traits between sites was driven by shifts in the presence of subordinate or 'rare' species rather than by changes in the dominant species.
6. Synthesis. Our study demonstrates co-ordinated species- and community-level changes in root traits along a soil chronosequence. These results highlight the influence of soil nutrition on plant functional traits and contribute to our understanding of the drivers of community assembly in a changing environment.

Plants and their arbuscular mycorrhizal fungal symbionts interact in complex underground networks involving multiple partners. This increases the potential for exploitation and defection by individuals, raising the question of how partners maintain a fair, two-way transfer of resources. We manipulated cooperation in plants and fungal partners to show that plants can detect, discriminate, and reward the best fungal partners with more carbohydrates. In turn, their fungal partners enforce cooperation by increasing nutrient transfer only to those roots providing more carbohydrates. On the basis of these observations we conclude that, unlike many other mutualisms, the symbiont cannot be

Determining the variation in roots traits within a grass root system is important for understanding the role of fine roots in carbon and nutrient cycling in grassland ecosystems, where the majority of biomass and litter accumulation occur belowground. However, few studies have been conducted in this regard. In this study, the structural and chemical traits of shoot-derived and root-derived roots were examined in three perennial grasses—Cleistogenes squarrosa, Achnatherum sibiricum and Stipa grandis—aiming to explore structural differences, responses to nitrogen and water addition in different types of roots and their correlations with aboveground plant nitrogen. Our results showed significant differences between these two root types, with root-derived roots having higher N concentration, tissue density, and specific root length, but lower C: N and diameter than shoot-derived roots. Trait relationships between root N concentration and tissue density for the two root types differed from that reported among species. These traits in different types of roots were insensitive to resource addition. Furthermore, N concentration in shoot-derived roots was more strongly linked to aboveground plant N concentration than root-derived roots. The results of this study demonstrate structural differences within the root system that may reflect functional heterogeneity in grass roots.

Mounting evidence has shown strong linkage of root function with root branch order. However, it is not known whether this linkage is consistent in different species. Here, root anatomic traits of the first five branch order were examined in five species differing in plant phylogeny and growth form in tropical and subtropical forests of south China. In Paramichelia baillonii, one tree species in Magnoliaceae, the intact cortex as well as mycorrhizal colonization existed even in the fifth-order root suggesting the preservation of absorption function in the higher-order roots. In contrast, dramatic decreases of cortex thickness and mycorrhizal colonization were observed from lower- to higher-order roots in three other tree species, Cunninghamia lanceolata, Acacia auriculiformis and Gordonia axillaries, which indicate the loss of absorption function. In a fern, Dicranopteris dichotoma, there were several cortex layers with prominently thickened cell wall and no mycorrhizal colonization in the third- and fourth-order roots, also demonstrating the loss of absorptive function in higher-order roots. Cluster analysis using these anatomic traits showed a different classification of root branch order in P. baillonii from other four species. As for the conduit diameter-density relationship in higher-order roots, the mechanism underpinning this relationship in P. baillonii was different from that in other species. In lower-order roots, different patterns of coefficient of variance for conduit diameter and density provided further evidence for the two types of linkage of root function with root branch order. These linkages corresponding to two types of ephemeral root modules have important implication in the prediction of terrestrial carbon cycling, although we caution that this study was pseudo-replicated. Future studies by sampling more species can test the generality of these two types of linkage.

Litter decomposition is one of the most important and dynamic components of nutrient cycling in ecosystems. In addition to the well-known actions of plant roots on litter, roots also constitute an important biotic factor regulating litter decomposition. However, relatively little attention has been given to this. Relative to the upper litter layer, the middle and lower layers are enriched in water and nutrients, which provide favorable conditions for root growth. By proliferating in litter, roots potentially influence litter decomposition through a priming effect, mycorrhizal fungi and N uptake. We give an overview of root foraging in litter, including factors influencing it. We then review its influence and possible mechanisms of root growth on litter decomposition. We also propose a conceptual module that enhances understanding of the relationship between root and litter decomposition.

Litter decomposition is one of the most important and dynamic components of nutrient cycling in ecosystems. In addition to the well-known actions of plant roots on litter, roots also constitute an important biotic factor regulating litter decomposition. However, relatively little attention has been given to this. Relative to the upper litter layer, the middle and lower layers are enriched in water and nutrients, which provide favorable conditions for root growth. By proliferating in litter, roots potentially influence litter decomposition through a priming effect, mycorrhizal fungi and N uptake. We give an overview of root foraging in litter, including factors influencing it. We then review its influence and possible mechanisms of root growth on litter decomposition. We also propose a conceptual module that enhances understanding of the relationship between root and litter decomposition.

Although linkages of leaf and whole-plant traits to leaf lifespan have been rigorously investigated, there is a limited understanding of similar linkages of whole-plant and fine root traits to root lifespan. In comparisons across species, do suites of traits found in leaves also exist for roots, and can these traits be used to predict root lifespan? We observed the fine root lifespan of 12 temperate tree species using minirhizotrons in a common garden and compared their median lifespans with fine-root and whole-plant traits. We then determined which set of combined traits would be most useful in predicting patterns of root lifespan. Median root lifespan ranged widely among species (95-336 d). Root diameter, calcium content, and tree wood density were positively related to root lifespan, whereas specific root length, nitrogen (N) : carbon (C) ratio, and plant growth rate were negatively related to root lifespan. Root diameter and plant growth rate, together (R(2) = 0.62) or in combination with root N : C ratio (R(2) = 0.76), were useful predictors of root lifespan across the 12 species. Our results highlight linkages between fine root lifespan in temperate trees and plant functional traits that may reduce uncertainty in predictions of root lifespan or turnover across species at broader spatial scales.

Mycorrhizal-induced growth depression of plants in high-P soil has been reported in many species. The carbon costs of factors contributing to this growth depression were analyzed in Volkamer lemon (Citrus volkameriana Tan. & Pasq.) colonized by the mycorrhizal (M) fungus Glomus intraradices Schenck and Smith. M and nonmycorrhizal (NM) plants were each grown at two P-supply rates. Carbon budgets of M and NM plants were determined by measuring whole-plant carbon assimilation and respiration rates using gas-exchange techniques. Biomass, M colonization, tissue-P concentration, and total fatty acid concentration in the fibrous roots were determined. Construction costs of the fibrous roots were estimated from heat of combustion, N, and ash content. Root-growth respiration was derived from daily root growth and root-construction cost. M and NM plants grown in high-P soil were similar in P concentration, daily shoot carbon assimilation, and daily shoot dark respiration. At 52 d after transplanting (DAT), however, combined daily root plus soil respiration was 37% higher for M than for NM plants, resulting in a 20% higher daily specific carbon gain (mmol CO2 [mmol carbon]-1 d-1) in NM than M plants. Estimates of specific carbon gain from specific growth rates indicated about a 10% difference between M and NM plants. Absolute values of specific carbon gain estimated by whole-plant gas exchange and by growth analysis were in general agreement. At 52 DAT, M and NM plants at high P had nearly identical whole-plant growth rates, but M plants had 19% higher root dry weight with 10% higher daily rates of root growth. These allocation differences at high P accounted for about 51% of the differences in root/soil respiration between M and NM plants. Significantly higher fatty acid concentrations in M than NM fibrous roots were correlated with differences in construction costs of the fibrous roots. Of the 37% difference in daily total root/soil respiration observed between high-P M and NM plants at 52 DAT, estimated daily growth respiration accounted for only about 16%, two-thirds of which was associated with construction of lipid-rich roots, and the remaining one-third with greater M root growth rates. Thus, of the 37% more root/soil respiration associated with M colonization of high-P plants, 10% was directly attributable to building lipid-rich roots, 51% to greater M root biomass allocation, and the remaining 39% could have been used for maintenance of the fungal tissue in the root and growth and maintenance of the extramatrical hyphae.

Roots, as organs distinguishable developmentally and anatomically from shoots (other than by occurrence of stomata and sporangia on above-ground organs), evolved in the sporophytes of at least two distinct lineages of early vascular plants during their initial major radiation on land in Early Devonian times (c. 410-395 million years ago). This was some 15 million years after the appearance of tracheophytes and c. 50 million years after the earliest embryophytes of presumed bryophyte affinity. Both groups are known initially only from spores, but from comparative anatomy of extant bryophytes and later Lower Devonian fossils it is assumed that, during these times, below-ground structures (if any) other than true roots fulfilled the functions of anchorage and of water and nutrient acquisition, despite lacking an endodermis (as do the roots of extant Lycopodium spp.). By 375 million years ago root-like structures penetrated almost a metre into the substratum, greatly increasing the volume of mineral matter subject to weathering by the higher than atmospheric CO(2) levels generated by plant and microbial respiration in material with restricted diffusive contact with the atmosphere. Chemical weathering consumes CO(2) in converting silicates into bicarbonate and Si(OH)(4). The CO(2) consumed in weathering ultimately came from atmospheric CO(2) via photosynthesis and respiration; this use of CO(2) probably accounts for most of the postulated 10-fold decrease in atmospheric CO(2) from 400-350 million years ago, with significant effects on shoot evolution. Subsequent evolution of roots has yielded much-branched axes down to 40 microm diameter, a lower limit set by long-distance transport constraints. Finer structures involved in the uptake of nutrients of low diffusivity in soil evolved at least 400 million years ago as arbuscular mycorrhizas or as evaginations of

Introduced species inevitably experience novel selection pressures in their new environments as a result of changes in mutualist and antagonist relationships. While most previous work has examined how escape from specialist enemies has influenced herbivore or pathogen resistance of exotic species, post-introduction shifts in exotic dependence on mutualists have not been considered. In a common environment, we compared dependence on AM fungi of North American and European populations of Hypericum perforatum (St. John's Wort), a forb native to Europe. Introduced North American populations responded less to inoculation with AM fungi than did European populations. Root architecture was strongly correlated with mycorrhizal response, and introduced populations had finer root architecture than native populations. Finally, introduced populations exhibited decreased root and increased reproductive allocation relative to European populations, consistent with a transition to a weedier life history; however, biomass allocation patterns were uncorrelated with mycorrhizal response. These findings are the first demonstration of a genetically based reduction of mycorrhizal dependence and shift in root architecture in an introduced species.

* The ectomycorrhizal (ECM) symbiosis was historically considered restricted to the temperate zones, but recent studies have shown the importance of this symbiosis across the tropics. We examined ECM fungal diversity, host plant phylogeny and ECM host preferences in a rainforest dominated by the leguminous host plants Dicymbe corymbosa, Dicymbe altsonii and Aldina insignis. * Ectomycorrhizal fungi were identified by internal transcribed spacer rDNA sequencing and host species were verified with chloroplast trnL sequencing. To test whether Dicymbe and Aldina represent independent gains of the ECM symbiosis, we constructed a Fabaceae phylogeny using MatK and trnL. We identified four independent ECM lineages within the Fabaceae. * We detected a diverse community of 118 ECM species dominated by the /clavulina, /russula-lactarius, /boletus, and /tomentella-thelephora lineages. Ectomycorrhizal species in Agaricales, Atheliales and Polyporales may represent previously unrecognized tropical-endemic ECM lineages. Previous studies suggested that ECM fungi did not diversify in the tropics, but the /clavulina lineage appears to have a center of diversity in tropical South America. * Dicymbe and Aldina represent independent gains of the ECM symbiosis in Fabaceae but their fungal symbionts showed no host preferences. Spatial factors are more important than hosts in structuring the ECM fungal community in this ecosystem.

Tree hydraulic architecture exhibits patterns that propagate from tissue to tree scales. A challenge is to make sense of these patterns in terms of trade-offs and adaptations. The universal trend for conduits per area to decrease with increasing conduit diameter below the theoretical packing limit may reflect the compromise between maximizing the area for conduction versus mechanical support and storage. Variation in conduit diameter may have two complementary influences: one being compromises between efficiency and safety and the other being that conduit tapering within a tree maximizes conductance per growth investment. Area-preserving branching may be a mechanical constraint, preventing otherwise more efficient top-heavy trees. In combination, these trends beget another: trees have more, narrower conduits moving from trunks to terminal branches. This pattern: (1) increases the efficiency of tree water conduction; (2) minimizes (but does not eliminate) any hydraulic limitation on the productivity or tissue growth with tree height; and (3) is consistent with the scaling of tree conductance and sap flow with size. We find no hydraulic reason why tree height should scale with a basal diameter to the two-thirds power as recently claimed; it is probably another mechanical constraint as originally proposed. The buffering effect of capacitance on the magnitude of transpiration-induced xylem tension appears to be coupled to cavitation resistance, possibly alleviating safety versus efficiency trade-offs.

Here, we tested hypothesized relationships among leaf and fine root traits of grass, forb, legume, and woody plant species of a savannah community. CO2 exchange rates, structural traits, chemistry, and longevity were measured in tissues of 39 species grown in long-term monocultures. Across species, respiration rates of leaves and fine roots exhibited a common regression relationship with tissue nitrogen (N) concentration, although legumes had lower rates at comparable N concentrations. Respiration rates and N concentration declined with increasing longevity of leaves and roots. Species rankings of leaf and fine-root N and longevity were correlated, but not specific leaf area and specific root length. The C3 and C4 grasses had lower N concentrations than forbs and legumes, but higher photosynthesis rates across a similar range of leaf N. Despite contrasting photosynthetic pathways and N2-fixing ability among these species, concordance in above- and below-ground traits was evident in comparable rankings in leaf and root longevity, N and respiration rates, which is evidence of a common leaf and root trait syndrome linking traits to effects on plant and ecosystem processes.

A survey of 659 papers mostly published since 1987 was conducted to compile a checklist of mycorrhizal occurrence among 3,617 species (263 families) of land plants. A plant phylogeny was then used to map the mycorrhizal information to examine evolutionary patterns. Several findings from this survey enhance our understanding of the roles of mycorrhizas in the origin and subsequent diversification of land plants. First, 80 and 92% of surveyed land plant species and families are mycorrhizal. Second, arbuscular mycorrhiza (AM) is the predominant and ancestral type of mycorrhiza in land plants. Its occurrence in a vast majority of land plants and early-diverging lineages of liverworts suggests that the origin of AM probably coincided with the origin of land plants. Third, ectomycorrhiza (ECM) and its derived types independently evolved from AM many times through parallel evolution. Coevolution between plant and fungal partners in ECM and its derived types has probably contributed to diversification of both plant hosts and fungal symbionts. Fourth, mycoheterotrophy and loss of the mycorrhizal condition also evolved many times independently in land plants through parallel evolution.

Fine roots are critical components for plant mercury (Hg) uptake and removal, but the patterns of Hg distribution and turnover within the heterogeneous fine root components and their potential limiting factors are poorly understood. Based on root branching structure, we studied the total Hg (THg) and its cellular partitioning in fine roots in 6 Chinese subtropical trees species and the impacts of root morphological and stoichiometric traits on Hg partitioning. The THg concentration generally decreased with increasing root order, and was higher in cortex than in stele. This concentration significantly correlated with root length, diameter, specific root length, specific root area, and nitrogen concentration, whereas its cytosolic fraction (accounting for <10% of THg) correlated with root carbon and sulfur concentrations. The estimated Hg return flux from dead fine roots outweighed that from leaf litter, and ephemeral first-order roots that constituted 7.2-22.3% of total fine root biomass may have contributed most to this flux (39-71%, depending on tree species and environmental substrate). Our results highlight the high capacity of Hg stabilization and Hg return by lower-order roots and demonstrate that turnover of lower-order roots may be an effective strategy of detoxification in perennial tree species.

Bringing together leaf trait data spanning 2,548 species and 175 sites we describe, for the first time at global scale, a universal spectrum of leaf economics consisting of key chemical, structural and physiological properties. The spectrum runs from quick to slow return on investments of nutrients and dry mass in leaves, and operates largely independently of growth form, plant functional type or biome. Categories along the spectrum would, in general, describe leaf economic variation at the global scale better than plant functional types, because functional types overlap substantially in their leaf traits. Overall, modulation of leaf traits and trait relationships by climate is surprisingly modest, although some striking and significant patterns can be seen. Reliable quantification of the leaf economics spectrum and its interaction with climate will prove valuable for modelling nutrient fluxes and vegetation boundaries under changing land-use and climate.

Historically, ephemeral roots have been equated with 'fine roots' (i.e. all roots of less than an arbitrary diameter, such as 2 mm), but evidence shows that 'fine roots' in woody species are complex branching systems with both rapid-cycling and slow-cycling components. A precise definition of ephemeral roots is therefore needed. Using a branch-order classification, a rhizotron method and sequential sampling of a root cohort, we tested the hypothesis that ephemeral root modules exist within the branching Fraxinus mandshurica (Manchurian ash) root system as distal nonwoody lateral branches, which show anatomical, nutritional and physiological patterns distinct from their woody mother roots. Our results showed that in F. mandshurica, distal nonwoody root branch orders die rapidly as intact lateral branches (or modules). These nonwoody branch orders exhibited highly synchronous changes in tissue nitrogen concentrations and respiration, dominated root turnover, nutrient flux and root respiration, and never underwent secondary development. The ephemeral root modules proposed here may provide a functional basis for differentiating and sampling short-lived absorptive roots in woody plants, and represent a conceptual leap over the traditional coarse-fine root dichotomies based on arbitrary size classes.

Aims Roots play an important role in the acquisition of soil resources and ecosystem processes. In this study, roots of nine subtropical tree species were studied to: 1) examine root diameter, specific root length and tissue N concentrations of different root branch orders in various tree species; 2) identify seasonal patterns of root N concentration; and 3) test the hypothesis that roots of lower orders have more marked seasonal patterns of N as they belong to a foundational module that is functionally different from higher order roots.
Methods Nine tree species in Dinghushan, Guangdong were studied. Roots were sampled from at least three trees of each species in the same plot at six different times during one year, and individual roots were separated according to branch orders. We measured root diameter, specific root length and tissue N concentrations of each order of all nine species. Variations of root N concentrations among orders in each species were analyzed by one-way ANOVA.
Important findings There were significant differences in root N concentration among different branch orders, with first and second orders having consistently higher N concentrations than higher orders, as expected. However, contrary to our hypothesis, the first two orders, which should belong to the rapid-cycling fraction of tree root systems, did not demonstrate the most marked seasonal changes in N concentration. Thus, the marked seasonal patterns in N concentrations in lower order roots found in temperate trees may not occur in subtropical trees. We suggest this contrast between temperate and subtropical trees may be related to N storage patterns in roots in temperate trees due to distinct phenology and the lack of distinct seasonality in subtropical trees. More study is needed to identify specific mechanisms that regulate N concentrations in roots and other plant tissues in subtropical vs. temperate trees.

Aims Roots play an important role in the acquisition of soil resources and ecosystem processes. In this study, roots of nine subtropical tree species were studied to: 1) examine root diameter, specific root length and tissue N concentrations of different root branch orders in various tree species; 2) identify seasonal patterns of root N concentration; and 3) test the hypothesis that roots of lower orders have more marked seasonal patterns of N as they belong to a foundational module that is functionally different from higher order roots.
Methods Nine tree species in Dinghushan, Guangdong were studied. Roots were sampled from at least three trees of each species in the same plot at six different times during one year, and individual roots were separated according to branch orders. We measured root diameter, specific root length and tissue N concentrations of each order of all nine species. Variations of root N concentrations among orders in each species were analyzed by one-way ANOVA.
Important findings There were significant differences in root N concentration among different branch orders, with first and second orders having consistently higher N concentrations than higher orders, as expected. However, contrary to our hypothesis, the first two orders, which should belong to the rapid-cycling fraction of tree root systems, did not demonstrate the most marked seasonal changes in N concentration. Thus, the marked seasonal patterns in N concentrations in lower order roots found in temperate trees may not occur in subtropical trees. We suggest this contrast between temperate and subtropical trees may be related to N storage patterns in roots in temperate trees due to distinct phenology and the lack of distinct seasonality in subtropical trees. More study is needed to identify specific mechanisms that regulate N concentrations in roots and other plant tissues in subtropical vs. temperate trees.