Resource exploitation of patches is influenced not simply by the rate of root production in the patches but also by the lifespan of the roots inhabiting the patches. We examined the effect of sustained localized nitrogen (N) fertilization on root lifespan in four tree species that varied widely in root morphology and presumed foraging strategy. The study was conducted in a 12-year-old common garden in central Pennsylvania using a combination of data from minirhizotron and root in-growth cores. The two fine-root tree species, Acer negundo L. and Populus tremuloides Michx., exhibited significant increases in root lifespan with local N fertilization; no significant responses were observed in the two coarse-root tree species, Sassafras albidum Nutt. and Liriodendron tulipifera L. Across species, coarse-root tree species had longer median root lifespan than fine-root tree species. Localized N fertilization did not significantly increase the N concentration or the respiration of the roots growing in the N-rich patch. Our results suggest that some plant species appear to regulate the lifespan of different portions of their root system to improve resource acquisition while other species do not. Our results are discussed in the context of different strategies of foraging of nutrient patches in species of different root morphology.

Our understanding of plant responses to biotic and abiotic drivers is largely based on above-ground plant traits, with little focus on below-ground traits despite their key role in water and nutrient uptake. Here, we aimed to understand the extent to which above- and below-ground traits are co-ordinated, and how these traits respond to soil moisture gradients and plant intraspecific competition.

Leaf dark respiration (Rdark ) is an important yet poorly quantified component of the global carbon cycle. Given this, we analyzed a new global database of Rdark and associated leaf traits. Data for 899 species were compiled from 100 sites (from the Arctic to the tropics). Several woody and nonwoody plant functional types (PFTs) were represented. Mixed-effects models were used to disentangle sources of variation in Rdark. Area-based Rdark at the prevailing average daily growth temperature (T) of each site increased only twofold from the Arctic to the tropics, despite a 20°C increase in growing T (8-28°C). By contrast, Rdark at a standard T (25°C, Rdark (25) ) was threefold higher in the Arctic than in the tropics, and twofold higher at arid than at mesic sites. Species and PFTs at cold sites exhibited higher Rdark (25) at a given photosynthetic capacity (Vcmax (25) ) or leaf nitrogen concentration ([N]) than species at warmer sites. Rdark (25) values at any given Vcmax (25) or [N] were higher in herbs than in woody plants. The results highlight variation in Rdark among species and across global gradients in T and aridity. In addition to their ecological significance, the results provide a framework for improving representation of Rdark in terrestrial biosphere models (TBMs) and associated land-surface components of Earth system models (ESMs). © 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.

Stomata regulate rates of carbon assimilation and water loss. Arbuscular mycorrhizal (AM) symbioses often modify stomatal behavior and therefore play pivotal roles in plant productivity. The size of the AM effect on stomatal conductance to water vapor (g s ) has varied widely, has not always been apparent, and is unpredictable. We conducted a meta-analysis of 460 studies to determine the size of the AM effect under ample watering and drought and to examine how experimental conditions have influenced the AM effect. Across all host and symbiont combinations under all soil moisture conditions, AM plants have shown 24 % higher g s than nonmycorrhizal (NM) controls. The promotion of g s has been over twice as great during moderate drought than under amply watered conditions. The AM influence on g s has been even more pronounced under severe drought, with over four times the promotion observed with ample water. Members of the Claroideoglomeraceae, Glomeraceae, and other AM families stimulated g s by about the same average amount. Colonization by native AM fungi has produced the largest promotion. Among single-AM symbionts, Glomus deserticola, Claroideoglomus etunicatum, and Funneliformis mosseae have had the largest average effects on g s across studies. Dicotyledonous hosts, especially legumes, have been slightly more responsive to AM symbiosis than monocotyledonous hosts, and C3 plants have shown over twice the AM-induced promotion of C4 plants. The extent of root colonization is important, with heavily colonized plants showing ×10 the g s promotion of lightly colonized plants. AM promotion of g s has been larger in growth chambers and in the field than in greenhouse studies, almost ×3 as large when plants were grown under high light than low light, and ×2.5 as large in purely mineral soils than in soils having an organic component. When AM plants have been compared with NM controls given NM pot culture, they have shown only half the promotion of g s as NM plants not given anything at inoculation to control for associated soil organisms. The AM effect has been much greater when AM plants were larger or had more phosphorus than NM controls. These findings should assist in further investigations of predictions and mechanisms of the AM influence on host g s.

Here, the coevolution of mycorrhizal fungi and roots is assessed in the light of evidence now available, from palaeobotanical and morphological studies and the analysis of DNA-based phylogenies. The first bryophyte-like land plants, in the early Devonian (400 million years ago), had endophytic associations resembling vesicular-arbuscular mycorrhizas (VAM) even before roots evolved. Mycorrhizal evolution would have progressed from endophytic hyphae towards balanced associations where partners were interdependent due to the exchange of limiting energy and nutrient resources. Most mycorrhizas are mutualistic, but in some cases the trend for increasing plant control of fungi culminates in the exploitative mycorrhizas of achlorophyllous, mycoheterotrophic plants. Ectomycorrhizal, ericoid and orchid mycorrhizas, as well as nonmycorrhizal roots, evolved during the period of rapid angiosperm radiation in the Cretaceous. It is hypothesised that roots gradually evolved from rhizomes to provide more suitable habitats for mycorrhizal fungi and provide plants with complex branching and leaves with water and nutrients. Selection pressures have caused the morphological divergence of roots with different types of mycorrizas. Root cortex thickness and exodermis suberization are greatest in obllgately mycorrhizal plants, while nonmycorrhizal plants tend to have fine roots, with more roots hairs and relatively advanced chemical defences. Major coevolutionary trends and the relative success of plants with different root types are discussed. Contents Summary 275 I. Introduction 276 II. Mycorrhizal Fungi 276 III. The Dawn of Mycorrhizas 279 IV. Mycorrhizal Associations of Living and Extinct Plants 282 V. Evolution of Roots 288 VI. The Root as a Habitat for Fungi 290 VII. Mycorrhizal Evolution Trends 295 Acknowledgements 298 References 298.

Minirhizotrons were used to observe fine root (≤1 mm) production, mortality, and longevity over 2 years in four sugar-maple-dominated northern hardwood forests located along a latitudinal temperature gradient. The sites also differed in N availability, allowing us to assess the relative influences of soil temperature and N availability in controlling fine root lifespans. Root production and mortality occurred throughout the year, with most production occurring in the early portion of the growing season (by mid-July). Mortality was distributed much more evenly throughout the year. For surface fine roots (0-10 cm deep), significant differences in root longevity existed among the sites, with median root lifespans for root cohorts produced in 1994 ranging from 405 to 540 days. Estimates of fine root turnover, based on the average of annual root production and mortality as a proportion of standing crop, ranged from 0.50 to 0.68 year for roots in the upper 30 cm of soil. The patterns across sites in root longevity and turnover did not follow the north to south temperature gradient, but rather corresponded to site differences in N availability, with longer average root lifespans and lower root turnover occurring where N availability was greater. This suggests the possibility that roots are maintained as long as the benefit (nutrients) they provide outweighs the C cost of keeping them alive. Root N concentrations and respiration rates (at a given temperature) were also higher at sites where N availability was greater. It is proposed that greater metabolic activity for roots in nitrogen-rich zones leads to greater carbohydrate allocation to those roots, and that a reduction in root C sink strength when local nutrients are depleted provides a mechanism through which root lifespan is regulated in these forests.

Temperate deciduous forest understoreys are experiencing widespread changes in community composition, concurrent with increases in rates of nitrogen supply. These shifts in plant abundance may be driven by interspecific differences in nutrient foraging (i.e. conservative vs. acquisitive strategies) and, thus, adaptation to contemporary nutrient loading conditions. This study sought to determine if interspecific differences in nutrient foraging could help explain patterns of shrub success and decline in eastern North American forests.Using plants grown in a common garden, fine root traits associated with nutrient foraging were measured for six shrub species. Traits included the mean and skewness of the root diameter distribution, specific root length (SRL), C:N ratio, root tissue density, arbuscular mycorrhizal colonization and foraging precision. Above- and below-ground productivity were also determined for the same plants, and population growth rates were estimated using data from a long-term study of community dynamics. Root traits were compared among species and associations among root traits, measures of productivity and rates of population growth were evaluated.Species fell into groups having thick or thin root forms, which correspond to conservative vs. acquisitive nutrient foraging strategies. Interspecific variation in root morphology and tissue construction correlated with measures of productivity and rates of cover expansion. Of the four species with acquisitive traits, three were introduced species that have become invasive in recent decades, and the fourth was a weedy native. In contrast, the two species with conservative traits were historically dominant shrubs that have declined in abundance in eastern North American forests.In forest understoreys of eastern North America, elevated nutrient availability may impose a filter on species success in addition to above-ground processes such as herbivory and overstorey canopy conditions. Shrubs that have root traits associated with rapid uptake of soil nutrients may be more likely to increase in abundance, while species without such traits may be less likely to keep pace with more productive species.© The Author 2017. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For Permissions, please email: journals.permissions@oup.com

The roots of the majority of tree species are associated with either arbuscular mycorrhizal (AM) or ectomycorrhizal (EM) fungi. The absorptive roots of tree species also vary widely in their diameter. The linkages between root thickness, mycorrhiza type and nutrient foraging are poorly understood. We conducted a large root ingrowth experiment in the field to investigate how absorptive roots of varying thickness and their associated fungi (AM vs. EM) exploit different nutrient patches (inorganic and organic) in a common garden. In nutrient-rich patches, thin-root tree species more effectively proliferated absorptive roots than thick-root tree species, whereas thick-root tree species proliferated more mycorrhizal fungal biomass than thin-root tree species. Moreover, nutrient patches enriched with organic materials resulted in greater root and mycorrhizal fungal proliferation compared to those enriched with inorganic nutrients. Irrespective of root morphology, AM tree species had higher root foraging precision than mycorrhizal hyphae foraging precision within organic patches, whereas EM tree species exhibited the opposite. Our findings that roots and mycorrhizal fungi are complementary in foraging within nutrient patches provide new insights into species coexistence and element cycling in terrestrial ecosystems.© 2016 by the Ecological Society of America.

There is growing recognition that classifying terrestrial plant species on the basis of their function (into 'functional types') rather than their higher taxonomic identity, is a promising way forward for tackling important ecological questions at the scale of ecosystems, landscapes or biomes. These questions include those on vegetation responses to and vegetation effects on, environmental changes (e.g. changes in climate, atmospheric chemistry, land use or other disturbances). There is also growing consensus about a shortlist of plant traits that should underlie such functional plant classifications, because they have strong predictive power of important ecosystem responses to environmental change and/or they themselves have strong impacts on ecosystem processes. The most favoured traits are those that are also relatively easy and inexpensive to measure for large numbers of plant species. Large international research efforts, promoted by the IGBP–GCTE Programme, are underway to screen predominant plant species in various ecosystems and biomes worldwide for such traits. This paper provides an international methodological protocol aimed at standardising this research effort, based on consensus among a broad group of scientists in this field. It features a practical handbook with step-by-step recipes, with relatively brief information about the ecological context, for 28 functional traits recognised as critical for tackling large-scale ecological questions.

For 76 annual, biennial, and perennial species common in the grasslands of central Minnesota, USA, we determined the patterns of correlations among seven organ‐level traits (specific leaf area, leaf thickness, leaf tissue density, leaf angle, specific root length, average fine root diameter, and fine root tissue density) and their relationships with two traits relating to growth form (whether species existed for part of the growing season in basal, non‐caulescent form and whether species were rhizomatous or not). The first correlation of traits showed that grasses had thin, dense leaves and thin roots while forbs had thick, low‐density leaves and thick roots without any significant differences in growth form or life history. The second correlation of traits showed a gradient of species from those with high‐density roots and high‐density erect leaves to species with low‐density roots and low‐density leaves that were held parallel to the ground. High tissue density species were more likely to exist as a basal rosette for part of the season, were less likely to be rhizomatous, and less likely to be annuals. We examined the relationships between the two axes that represent the correlations of traits and previously collected data on the relative abundance of species across gradients of nitrogen addition and disturbance. Grasses were generally more abundant than forbs and the relative abundance of grasses and forbs did not change with increasing nitrogen addition or soil disturbance. High tissue density species became less common as fertility and disturbance increased.

The relative competitive abilities of Agropyron desertorum and Agropyron spicatum under rangeland conditions were compared using Artemisia tridentata ssp. wyomingensis transplants as indicator plants. We found A. desertorum to have substantially greater competitive ability than A. spicatum as manifested by the responses of Artemisia shrubs that were transplanted into nearly monospecific stands of these grass species. The Artemisia indicator plants had lower survival, growth, reproduction, and late-season water potential in the neighborhoods dominated by A. desertorum than in those dominated by A. spicatum. In similar, essentially monospecific grass stands, neutron probe soil moisture measurements showed that stands of A. desertorum extracted water more rapidly from the soil profile than did those of A. spicatum. These differences in extraction rates correlate clearly with the differences in indicator plant success in the respective grass stands. Nitrogen and phosphorus concentrations in Artemisia tissues suggested these nutrients were not limiting indicator plant growth and survival in the A. desertorum plots.

The identification of plant functional traits that can be linked to ecosystem processes is of wide interest, especially for predicting vegetational responses to climate change. Root diameter of the finest absorptive roots may be one plant trait that has wide significance. Do species with relatively thick absorptive roots forage in nutrient-rich patches differently from species with relatively fine absorptive roots? We measured traits related to nutrient foraging (root morphology and architecture, root proliferation, and mycorrhizal colonization) across six coexisting arbuscular mycorrhizal (AM) temperate tree species with and without nutrient addition. Root traits such as root diameter and specific root length were highly correlated with root branching intensity, with thin-root species having higher branching intensity than thick-root species. In both fertilized and unfertilized soil, species with thin absorptive roots and high branching intensity showed much greater root length and mass proliferation but lower mycorrhizal colonization than species with thick absorptive roots. Across all species, fertilization led to increased root proliferation and reduced mycorrhizal colonization. These results suggest that thin-root species forage more by root proliferation, whereas thick-root species forage more by mycorrhizal fungi. In mineral nutrient-rich patches, AM trees seem to forage more by proliferating roots than by mycorrhizal fungi. © 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.

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 \nfor total root biomass in shrublands (r2 = 0.55). On the basis of the best‐fit exponential model, the Q10 for root \nturnover was 1.4 for forest small diameter roots (5 mm or less), 1.6 for grassland fine roots, and 1.9 for shrublands. \nSurprisingly, after accounting for temperature, there was no such global relationship between precipitation and \nroot turnover. The slowest average turnover rates were observed for entire tree root systems (10% annually), \nfollowed by 34% for shrubland total roots, 53% for grassland fine roots, 55% for wetland fine roots, and 56% for \nforest fine roots. Root turnover decreased from tropical to high‐latitude systems for all plant functional groups. \nTo test whether global relationships can be used to predict interannual variability in root turnover, we evaluated \n14 yr of published root turnover data from a shortgrass steppe site in northeastern Colorado, USA. At this site \nthere was no correlation between interannual variability in mean annual temperature and root turnover. Rather, \nturnover was positively correlated with the ratio of growing season precipitation and maximum monthly \ntemperature (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.

Volumetric soil water content and soil water potential were measured beneath a winter wheat crop during the 1975 growing season. Almost no rain fell between mid-May and mid-July and the soil dried continuously until the potential was less than – 20 bars to a depth of 80 cm. Evaporation was separated from drainage by denning an ‘effective rooting depth’ at which the hydraulic gradient was zero.

* Different portions of tree root systems play distinct functional roles, yet precisely how to distinguish roots of different functions within the branching fine-root system is unclear. * Here, anatomy and mycorrhizal colonization was examined by branch order in 23 Chinese temperate tree species of both angiosperms and gymnosperms forming ectomycorrhizal and arbuscular-mycorrhizal associations. * Different branch orders showed marked differences in anatomy. First-order roots exhibited primary development with an intact cortex, a high mycorrhizal colonization rate and a low stele proportion, thus serving absorptive functions. Second and third orders had both primary and secondary development. Fourth and higher orders showed mostly secondary development with no cortex or mycorrhizal colonization, and thus have limited role in absorption. Based on anatomical traits, it was estimated that c. 75% of the fine-root length was absorptive, and 68% was mycorrhizal, averaged across species. * These results showed that: order predicted differences in root anatomy in a relatively consistent manner across species; anatomical traits associated with absorption and mycorrhizal colonization occurred mainly in the first three orders; the single diameter class approach may have overestimated absorptive root length by 25% in temperate forests.

Leaf-level determinants of species environmental stress tolerance are still poorly understood. Here, we explored dependencies of species shade (T(shade)) and drought (T(drought)) tolerance scores on key leaf structural and functional traits in 339 Northern Hemisphere temperate woody species. In general, T(shade) was positively associated with leaf life-span (L(L)), and negatively with leaf dry mass (M(A)), nitrogen content (N(A)), and photosynthetic capacity (A(A)) per area, while opposite relationships were observed with drought tolerance. Different trait combinations responsible for T(shade) and T(drought) were observed among the key plant functional types: deciduous and evergreen broadleaves and evergreen conifers. According to principal component analysis, resource-conserving species with low N content and photosynthetic capacity, and high L(L) and M(A), had higher T(drought), consistent with the general stress tolerance strategy, whereas variation in T(shade) did not concur with the postulated stress tolerance strategy. As drought and shade often interact in natural communities, reverse effects of foliar traits on these key environmental stress tolerances demonstrate that species niche differentiation is inherently constrained in temperate woody species. Different combinations of traits among key plant functional types further explain the contrasting bivariate correlations often observed in studies seeking functional explanation of variation in species environmental tolerances.

The dynamics of fine (<2.0 mm) roots were measured in two sugar maple (Acersaccharum Marsh.) dominated ecosystems (northern and southern sites) during 1989 and 1990 using a combination of minirhizotrons and destructive harvests of fine root biomass and N content. Greater than 50% of annual length production occurred before midsummer in both ecosystems, while the period of greatest mortality was from late summer through winter. About one third of annual fine root production and mortality occur simultaneously, with little observable change in total root length pools. Using fine root length dynamics to derive biomass production and mortality, we calculated annual biomass production values of approximately 8000 and 7300 kg•ha−1•year−1, respectively, at the southern and northern sites. Corresponding biomass mortality (i.e., turnover) values were 6700 and 4800 kg•ha−1•year−1, and total nitrogen returns to the soil from fine root mortality were 72 kg•ha−1•year−1 at the southern site and 54 kg•ha−1•year−1 at the northern site. Fine roots dominated total biomass and N litter inputs to the soil in both ecosystems, accounting for over 55% of total biomass and nearly 50% of total N returns. In both ecosystems, roots <0.5 mm comprised the bulk of fine root biomass and N pools, and the contribution of these roots to northern hardwood ecosystem carbon and nitrogen budgets may have been underestimated in the past.

Understanding and predicting ecosystem functioning (e.g., carbon and water fluxes) and the role of soils in carbon storage requires an accurate assessment of plant rooting distributions. Here, in a comprehensive literature synthesis, we analyze rooting patterns for terrestrial biomes and compare distributions for various plant functional groups. We compiled a database of 250 root studies, subdividing suitable results into 11 biomes, and fitted the depth coefficient β to the data for each biome (Gale and Grigal 1987). β is a simple numerical index of rooting distribution based on the asymptotic equation Y=1-β, where d = depth and Y = the proportion of roots from the surface to depth d. High values of β correspond to a greater proportion of roots with depth. Tundra, boreal forest, and temperate grasslands showed the shallowest rooting profiles (β=0.913, 0.943, and 0.943, respectively), with 80-90% of roots in the top 30 cm of soil; deserts and temperate coniferous forests showed the deepest profiles (β=0.975 and 0.976, respectively) and had only 50% of their roots in the upper 30 cm. Standing root biomass varied by over an order of magnitude across biomes, from approximately 0.2 to 5 kg m. Tropical evergreen forests had the highest root biomass (5 kg m), but other forest biomes and sclerophyllous shrublands were of similar magnitude. Root biomass for croplands, deserts, tundra and grasslands was below 1.5 kg m. Root/shoot (R/S) ratios were highest for tundra, grasslands, and cold deserts (ranging from 4 to 7); forest ecosystems and croplands had the lowest R/S ratios (approximately 0.1 to 0.5). Comparing data across biomes for plant functional groups, grasses had 44% of their roots in the top 10 cm of soil. (β=0.952), while shrubs had only 21% in the same depth increment (β=0.978). The rooting distribution of all temperate and tropical trees was β=0.970 with 26% of roots in the top 10 cm and 60% in the top 30 cm. Overall, the globally averaged root distribution for all ecosystems was β=0.966 (r =0.89) with approximately 30%, 50%, and 75% of roots in the top 10 cm, 20 cm, and 40 cm, respectively. We discuss the merits and possible shortcomings of our analysis in the context of root biomass and root functioning.

One of the most dramatic effects of infection by vesicular-arbuscular mycorrhizal fungi on the physiology of the host plant is an increase in phosphorus absorption. When phosphorus is limiting, the maximum extent to which mycorrhizal infection can improve plant performance is thus predicted to be a function of the phosphorus deficit of the plant, the difference between phosphorus demand and phosphorus supply. Phosphorus demand is defined as the rate of phosphorus absorption that would result in optimum performance of the plant as measured by growth rate, reproduction or fitness. The phosphorus supply is defined as the actual rate of phosphorus absorption under the prevailing conditions. Variation among plant taxa in morphological, physiological or phenological traits which affect either phosphorus demand or phosphorus supply (and thus phosphorus deficit) is predicted to lead to variation in potential response to mycorrhizal infection. The actual response to mycorrhizal infection is predicted to be a function of the increase in phosphorus uptake due to mycorrhizal infection and the phosphorus utilization efficiency of the plant. Demonstrated variability in responsiveness to mycorrhizal infection among plant taxa suggests that mycorrhizal fungi may play an important role in determining the structure of plant communities. Mycorrhizal infection may alter the phosphorus deficit or phosphorus utilization efficiency independently from its direct effect on phosphorus uptake, making the prediction of response to mycorrhizal infection based on the traits of non-mycorrhizal plants quite difficult. For example, infection may at times increase the rate of phosphorus accumulation beyond that which can be currently utilized in growth, reducing the current phosphorus utilization efficiency. Such momentary 'luxury consumption' of phosphorus may, however, serve a storage function and be utilized subsequently, allowing mycorrhizal plants ultimately to outperform non-mycorrhizal plants. CONTENTS Summary 365 I. Introduction 366 II. The concepts of phosphorus supply, phosphorus demand and phosphorus deficit 367 III. Factors which affect phosphorus supply 369 IV. Plant traits which affect phosphorus demand 370 V. Variation in response to mycorrhizal infection 372 VI. The effect of infection on inherent traits which influence phosphorus supply, phosphorus demand or phosphorus utilization efficiency 378 VII. Mycorrhizal effects not related to phosphorus 380 VIII. Conclusions 380 Acknowledgements 381 References 381.

Absorptive root traits show remarkable cross-species variation, but major root trait dimensions across species have not been defined. We sampled first-order roots and measured 14 root traits for 96 angiosperm woody species from subtropical China, including root diameter, specific root length, stele diameter, cortex thickness, root vessel size and density, mycorrhizal colonization rate, root branching intensity, tissue density, and concentrations of carbon and nitrogen ([N]). Root traits differed in the degree of variation and phylogenetic conservatism, but showed predictable patterns of cross-trait coordination. Root diameter, cortex thickness and stele diameter displayed high variation across species (coefficient of variation (CV)=0.51-0.69), whereas the stele:root diameter ratio and [N] showed low variation (CV<0.32). Root diameter, cortex thickness and stele diameter showed a strong phylogenetic signal across species, whereas root branching traits did not, and these two sets of traits were segregated onto two nearly orthogonal (independent) principal component analysis (PCA) axes. Two major dimensions of root trait variation were found: a diameter-related dimension potentially integrating root construction, maintenance, and persistence with mycorrhizal colonization, and a branching architecture dimension expressing root plastic responses to the environment. These two dimensions may offer a promising path for better understanding root trait economics and root ecological strategies world-wide.© 2014 The Authors. New Phytologist © 2014 New Phytologist Trust.

. Plant roots typically vary along a dominant ecological axis, the root economics spectrum, depicting a tradeoff between resource acquisition and conservation. For absorptive roots, which are mainly responsible for resource acquisition, we hypothesized that root economic strategies differ with increasing root diameter. To test this hypothesis, we used seven plant species (a fern, a conifer, and five angiosperms from south China) for which we separated absorptive roots into two categories: thin roots (thickness of root cortex plus epidermis < 247 µm) and thick roots. For each category, we analyzed a range of root traits related to resource acquisition and conservation, including root tissue density, different carbon (C), and nitrogen (N) fractions (i.e., extractive, acid-soluble, and acid-insoluble fractions) as well as root anatomical traits. The results showed significant relationships among root traits indicating an acquisition-conservation tradeoff for thin absorptive roots while no such trait relationships were found for thick absorptive roots. Similar results were found when reanalyzing data of a previous study including 96 plant species. The contrasting economic strategies between thin and thick absorptive roots, as revealed here, may provide a new perspective on our understanding of the root economics spectrum.\n

The root economics spectrum (RES), a common hypothesis postulating a tradeoff between resource acquisition and conservation traits, is being challenged by conflicting relationships between root diameter, tissue density (RTD) and root nitrogen concentration (RN). Here, we analyze a global trait dataset of absorptive roots for over 800 plant species. For woody species (but not for non-woody species), we find nonlinear relationships between root diameter and RTD and RN, which stem from the allometric relationship between stele and cortical tissues. These nonlinear relationships explain how sampling bias from different ends of the nonlinear curves can result in conflicting trait relationships. Further, the shape of the relationships varies depending on evolutionary context and mycorrhizal affiliation. Importantly, the observed nonlinear trait relationships do not support the RES predictions. Allometry-based nonlinearity of root trait relationships improves our understanding of the ecology, physiology and evolution of absorptive roots.

Contents 1597 I. 1597 II. 1597 III. 1598 IV. 1598 V. 1600 VI. 1601 VII. 1601 VIII. 1601 1602 References 1602 SUMMARY: Trait-based approaches have led to significant advances in plant ecology, but are currently biased toward above-ground traits. It is becoming clear that a stronger emphasis on below-ground traits is needed to better predict future changes in plant biodiversity and their consequences for ecosystem functioning. Here I propose six 'below-ground frontiers' in trait-based plant ecology, with an emphasis on traits governing soil nutrient acquisition: redefining fine roots; quantifying root trait dimensionality; integrating mycorrhizas; broadening the suite of root traits; determining linkages between root traits and abiotic and biotic factors; and understanding ecosystem-level consequences of root traits. Focusing research efforts along these frontiers should help to fulfil the promise of trait-based ecology: enhanced predictive capacity across ecological scales.© 2016 The Authors. New Phytologist © 2016 New Phytologist Trust.

Increasing attention has recently been focused on the linkages between plant functional traits and ecosystem functioning. A comprehensive understanding of these linkages can facilitate to address the ecological consequences of plant species loss induced by human activities and climate change, and provide theoretical support for ecological restoration and ecosystem management. In recent twenty years, the evidence of strong correlations between plant functional traits and changes in ecosystem processes is growing. More importantly, ecosystem functioning can be predicted more precisely, using plant functional trait diversity (i.e., functional diversity) than species diversity. In this paper, we first defined plant functional traits and their important roles in determining ecosystem processes. Then, we review recent advances in the relationships between ecosystem functions and plant functional traits and their diversity. Finally, we propose several important future research directions, including (1) exploration of the relationships between aboveground and belowground plant traits and their roles in determining ecosystem functioning, (2) incorporation of the impacts of consumer and global environmental change into the correlation between plant functional traits and ecosystem functioning, (3) effects of functional diversity on ecosystem multifunctionality, and (4) examination of the functional diversity-ecosystem functioning relationship at different temporal and spatial scales.

植物功能性状与生态系统功能是生态学研究的一个重要领域和热点问题。开展植物功能性状与生态系统功能的研究不仅有助于人类更好地应对全球变化情景下生物多样性丧失的生态学后果,而且能为生态恢复实践提供理论基础。近二十年来,该领域的研究迅速发展,并取得了一系列的重要研究成果,增强了人们对植物功能性状-生态系统功能关系的认识和理解。本文首先明确了植物功能性状的概念, 评述了近年来植物功能性状-生态系统功能关系领域的重要研究结果, 尤其是植物功能性状多样性-生态系统功能关系研究现状; 提出了未来植物功能性状与生态系统功能关系研究中应加强植物地上和地下性状之间关系及其与生态系统功能、植物功能性状与生态系统多功能性、不同时空尺度上植物功能性状与生态系统功能, 以及全球变化和消费者的影响等方面。

Functional traits and their variation mediate plant species coexistence and spatial distribution. Yet, how patterns of variation in belowground traits influence resource acquisition across species and plant communities remains obscure. To characterize diverse belowground strategies in relation to species coexistence and abundance, we assessed four key belowground traits - root diameter, root branching intensity, first-order root length and mycorrhizal colonization - in 27 coexisting species from three grassland communities along a precipitation gradient. Species with thinner roots had higher root branching intensity, but shorter first-order root length and consistently low mycorrhizal colonization, whereas species with thicker roots enhanced their capacity for resource acquisition by producing longer first-order roots and maintaining high mycorrhizal colonization. Plant species observed across multiple sites consistently decreased root branching and/or mycorrhizal colonization, but increased lateral root length with decreasing precipitation. Additionally, the degree of intraspecific trait variation was positively correlated with species abundance across the gradient, indicating that high intraspecific trait variation belowground may facilitate greater fitness and chances of survival across multiple habitats. These results suggest that a small set of critical belowground traits can effectively define diverse resource acquisition strategies in different environments and may forecast species survival and range shifts under climate change.© 2017 The Authors. New Phytologist © 2017 New Phytologist Trust.

In most cases, both roots and mycorrhizal fungi are needed for plant nutrient foraging. Frequently, the colonization of roots by arbuscular mycorrhizal (AM) fungi seems to be greater in species with thick and sparsely branched roots than in species with thin and densely branched roots. Yet, whether a complementarity exists between roots and mycorrhizal fungi across these two types of root system remains unclear. We measured traits related to nutrient foraging (root morphology, architecture and proliferation, AM colonization and extramatrical hyphal length) across 14 coexisting AM subtropical tree species following root pruning and nutrient addition treatments. After root pruning, species with thinner roots showed more root growth, but lower mycorrhizal colonization, than species with thicker roots. Under multi-nutrient (NPK) addition, root growth increased, but mycorrhizal colonization decreased significantly, whereas no significant changes were found under nitrogen or phosphate additions. Moreover, root length proliferation was mainly achieved by altering root architecture, but not root morphology. Thin-root species seem to forage nutrients mainly via roots, whereas thick-root species rely more on mycorrhizal fungi. In addition, the reliance on mycorrhizal fungi was reduced by nutrient additions across all species. These findings highlight complementary strategies for nutrient foraging across coexisting species with contrasting root traits. © 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.

Plant roots are a highly heterogeneous and hierarchical system. Although the root-order method is superior to the root diameter method for revealing differences in the morphology and physiology of fine roots, its complex partitioning limits its application. Whether root order can be determined by partitioning the main root based on its diameter remains uncertain. Four methods were employed for studying the morphological characteristics of seedling roots of two Pinus species in a natural and nitrogen-enriched environment. The intrinsic relationships among categories of roots by root order and diameter were systematically compared to explore the possibility of using the latter to describe root morphology. The normal transformation method proved superior to the other three in that the diameter intervals corresponded most closely (at least 68.3%) to the morphological characteristics. The applied methods clearly distinguished the results from the natural and nitrogen-rich environments. Considering both root diameter and order simplified the classification of fine roots, and improved the estimation of root lifespan and the data integrity of field collection, but failed to partition all roots into uniform diameter intervals.

We measured the influences of soil fertility and plant community composition on Glomeromycota, and tested the prediction of the functional equilibrium hypothesis that increased availability of soil resources will reduce the abundance of arbuscular mycorrhizal (AM) fungi. Communities of plants and AM fungi were measured in mixed roots and in Elymus nutans roots across an experimental fertilization gradient in an alpine meadow on the Tibetan Plateau. As predicted, fertilization reduced the abundance of Glomeromycota as well as the species richness of plants and AM fungi. The response of the glomeromycotan community was strongly linked to the plant community shift towards dominance by Elymus nutans. A reduction in the extraradical hyphae of AM fungi was associated with both the changes in soil factors and shifts in the plant community composition that were caused by fertilization. Our findings highlight the importance of soil fertility in regulating both plant and glomeromycotan communities, and emphasize that high fertilizer inputs can reduce the biodiversity of plants and AM fungi, and influence the sustainability of ecosystems.© 2012 The Authors. New Phytologist © 2012 New Phytologist Trust.

A hypothetical ideotype is presented to optimize water and N acquisition by maize root systems. The overall premise is that soil resource acquisition is optimized by the coincidence of root foraging and resource availability in time and space. Since water and nitrate enter deeper soil strata over time and are initially depleted in surface soil strata, root systems with rapid exploitation of deep soil would optimize water and N capture in most maize production environments. • THE IDEOTYPE: Specific phenes that may contribute to rooting depth in maize include (a) a large diameter primary root with few but long laterals and tolerance of cold soil temperatures, (b) many seminal roots with shallow growth angles, small diameter, many laterals, and long root hairs, or as an alternative, an intermediate number of seminal roots with steep growth angles, large diameter, and few laterals coupled with abundant lateral branching of the initial crown roots, (c) an intermediate number of crown roots with steep growth angles, and few but long laterals, (d) one whorl of brace roots of high occupancy, having a growth angle that is slightly shallower than the growth angle for crown roots, with few but long laterals, (e) low cortical respiratory burden created by abundant cortical aerenchyma, large cortical cell size, an optimal number of cells per cortical file, and accelerated cortical senescence, (f) unresponsiveness of lateral branching to localized resource availability, and (g) low K(m) and high Vmax for nitrate uptake. Some elements of this ideotype have experimental support, others are hypothetical. Despite differences in N distribution between low-input and commercial maize production, this ideotype is applicable to low-input systems because of the importance of deep rooting for water acquisition. Many features of this ideotype are relevant to other cereal root systems and more generally to root systems of dicotyledonous crops.

Fine root respiration is a significant component of carbon cycling in forest ecosystems. Although fine roots differ functionally from coarse roots, these root types have been distinguished based on arbitrary diameter cut-offs (e.g., 2 or 5 mm). Fine root morphology is directly related to physiological function, but few attempts have been made to understand the relationships between morphology and respiration of fine roots. To examine relationships between respiration rates and morphological traits of fine roots (0.15-1.4 mm in diameter) of mature Quercus serrata Murr., we measured respiration of small fine root segments in the field with a portable closed static chamber system. We found a significant power relationship between mean root diameter and respiration rate. Respiration rates of roots<0.4 mm in mean diameter were high and variable, ranging from 3.8 to 11.3 nmol CO2 g(-1) s(-1), compared with those of larger diameter roots (0.4-1.4 mm), which ranged from 1.8 to 3.0 nmol CO2 g(-1) s(-1). Fine root respiration rate was positively correlated with specific root length (SRL) as well as with root nitrogen (N) concentration. For roots<0.4 mm in diameter, SRL had a wider range (11.3-80.4 m g(-1)) and was more strongly correlated with respiration rate than diameter. Our results indicate that a more detailed classification of fine roots<2.0 mm is needed to represent the heterogeneity of root respiration and to evaluate root biomass and root morphological traits.

Soil N fertility has an effect on belowground C allocation, but the physiological and morphological responses of individual fine root segments to variations in N availability under field conditions are still unclear. In this study, the direction and magnitude of the physiological and morphological function of fine roots in response to variable in situ soil N fertility in a forest site were determined. We measured the specific root respiration (Rr) rate, N concentration and morphology of fine root segments with 1-3 branching orders in a 100-year-old coniferous forest of Chamaecyparis obtusa. Higher soil N fertility induced higher Rr rates, root N concentration, and specific root length (SRL), and lower root tissue density (RTD). In all fertility levels, the Rr rates were significantly correlated positively with root N and SRL and negatively with RTD. The regression slopes of respiration with root N and RTD were significantly higher along the soil N fertility gradient. Although no differences in the slopes of Rr and SRL relationship were found across the levels, there were significant shifts in the intercept along the common slope. These results suggest that a contrasting pattern in intraspecific relationships between specific Rr and N, RTD, and SRL exists among soils with different N fertility. Consequently, substantial increases in soil N fertility would exert positive effects on organ-scale root performance by covarying the Rr, root N, and morphology for their potential nutrient and water uptake.

Estimates of forest net primary production (NPP) demand accurate estimates of root production and turnover. We assessed root turnover with the use of an isotope tracer in two forest free-air carbon dioxide enrichment experiments. Growth at elevated carbon dioxide did not accelerate root turnover in either the pine or the hardwood forest. Turnover of fine root carbon varied from 1.2 to 9 years, depending on root diameter and dominant tree species. These long turnover times suggest that root production and turnover in forests have been overestimated and that sequestration of anthropogenic atmospheric carbon in forest soils may be lower than currently estimated.

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 (R² = 0.62) or in combination with root N : C ratio (R² = 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.© 2012 The Authors. New Phytologist © 2012 New Phytologist Trust.

Fine roots acquire essential soil resources and mediate biogeochemical cycling in terrestrial ecosystems. Estimates of carbon and nutrient allocation to build and maintain these structures remain uncertain because of the challenges of consistently measuring and interpreting fine-root systems. Traditionally, fine roots have been defined as all roots ≤ 2 mm in diameter, yet it is now recognized that this approach fails to capture the diversity of form and function observed among fine-root orders. Here, we demonstrate how order-based and functional classification frameworks improve our understanding of dynamic root processes in ecosystems dominated by perennial plants. In these frameworks, fine roots are either separated into individual root orders or functionally defined into a shorter-lived absorptive pool and a longer-lived transport fine-root pool. Using these frameworks, we estimate that fine-root production and turnover represent 22% of terrestrial net primary production globally - a c. 30% reduction from previous estimates assuming a single fine-root pool. Future work developing tools to rapidly differentiate functional fine-root classes, explicit incorporation of mycorrhizal fungi into fine-root studies, and wider adoption of a two-pool approach to model fine roots provide opportunities to better understand below-ground processes in the terrestrial biosphere.© 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.

Trait-based approaches provide a useful framework to investigate plant strategies for resource acquisition, growth, and competition, as well as plant impacts on ecosystem processes. Despite significant progress capturing trait variation within and among stems and leaves, identification of trait syndromes within fine-root systems and between fine roots and other plant organs is limited. Here we discuss three underappreciated areas where focused measurements of fine-root traits can make significant contributions to ecosystem science. These include assessment of spatiotemporal variation in fine-root traits, integration of mycorrhizal fungi into fine-root-trait frameworks, and the need for improved scaling of traits measured on individual roots to ecosystem-level processes. Progress in each of these areas is providing opportunities to revisit how below-ground processes are represented in terrestrial biosphere models. Targeted measurements of fine-root traits with clear linkages to ecosystem processes and plant responses to environmental change are strongly needed to reduce empirical and model uncertainties. Further identifying how and when suites of root and whole-plant traits are coordinated or decoupled will ultimately provide a powerful tool for modeling plant form and function at local and global scales.© 2017 The Authors. New Phytologist © 2017 New Phytologist Trust.

Since their emergence onto land, terrestrial plants have developed diverse strategies to acquire soil resources. However, we lack a framework that adequately captures how these strategies vary among species. Observations from around the world now allow us to quantify the variation observed in commonly-measured fine-root traits but it is unclear how root traits are interrelated and whether they fall along an "economic" spectrum of acquisitive to conservative strategies. We assessed root trait variation and mycorrhizal colonization rates by leveraging the largest global database of fine-root traits (the Fine-Root Ecology Database; FRED). We also developed a heuristic model to explore the role of mycorrhizal fungi in defining belowground exploration efficiency across a gradient of thin- to thick-diameter roots. In support of the expectations of the "root economic spectrum," we found that root diameter was negatively related to specific root length (Pearson's =-0.76). However, we found an unexpected negative relationship between root diameter and root tissue density (Pearson's = -0.40), and we further observed that root nitrogen content was largely unrelated to other economic traits. Mycorrhizal colonization was most closely associated with root diameter (Pearson's = 0.62) and was unrelated to root tissue density and root nitrogen. The heuristic model demonstrated that while thinner roots have inherently greater capacity to encounter soil resources based on higher surface area per unit mass, the potential for increased associations with mycorrhizal fungi in thicker roots, combined with greater hyphal growth, can result in equally acquisitive strategies for both thin- and thick roots. Taken together, our assessments of root trait variation, trade-offs with mycorrhizal fungi, and broader connections to root longevity allowed us to propose a series of fundamental constraints on belowground resource acquisition strategies. Physical tradeoffs based on root construction (i.e., economic traits) and functional limitations related to the capacity of a root to encounter and acquire soil resources combine to limit the two-dimensional belowground trait space. Within this trait space there remains a diversity of additional variation in root traits that facilitates a wide range of belowground resource acquisition strategies.Copyright © 2019 McCormack and Iversen.

植物性状反映了植物对生长环境的响应和适应，将环境、植物个体和生态系统结构、 过程与功能联系起来（所谓的&ldquo;植物功能性状&rdquo;）。该文介绍了植物功能性状的分类体系，综述了国内外植物功能性状与气候（包括气温、降水、光照）、地理空间变异（包括地形地 貌、生态梯度、海拔）、营养、干扰（包括火灾、放牧、生物入侵、土地利用）等环境因素，以及与生态系统功能之间关系的研究进展，探讨了全球变化（气候变化和CO₂浓度升高 ） 对个体和群落植物功能性状的影响。植物功能性状的研究已经取得很多成果，并应用于全球变化、古植被恢复和古气候定量重建、环境监测与评价、生态保护和恢复等研究中，但大尺度、多生境因子下的植物功能性状研究仍有待于加强，同时需要改进性状的测量手段；我国 的植物功能性状研究还需要更加明朗化和系统化。

Absorption roots usually refer to several terminal branch orders in a root branch that are comprised of primary tissues and are responsible for resource uptake. Functional traits of absorption roots have been widely used to assess and predict a range of functions and processes from individual to ecosystem scale. Mycorrhizal fungi colonization is one of the key traits exerting significant influence on root morphology, structure, and the inter-relationships among root traits. In this paper, we first review the relationships of mycorrhizal fungi with two traits closely related to resource uptake: root hair and root diameter; a hypothesis is proposed to describe relationships among mycorrhizal fungi, root hair and chemical defense. Then, we review how functional traits in different absorption roots are altered by mycorrhizal fungi adapting to environments differing in precipitation, temperature, soil fertility, and energy consumption. Finally, we identify several topics and research outlooks for guiding future studies to facilitate studies on the relationship between mycorrhizal fungi and root functional traits.

吸收根(absorption root)一般是指根枝系统末端少数几级具有初生结构、负责物质吸收的根。吸收根功能性状被广泛用于评价和预测植物个体到生态系统水平上的一系列功能和过程。菌根真菌侵染是吸收根的一个关键性状, 它可以深刻影响吸收根的形态、结构, 以及功能性状之间的关系。该文针对与吸收功能密切相关的菌根真菌与根毛和根直径之间的关系进行了研究综述, 提出了真菌侵染、根毛和化学防御之间关系的一个假说; 探讨了温带和热带不同类型的吸收根如何通过菌根真菌影响根的功能性状, 从而适应不同的水热条件、养分状况和能量消耗; 提出一些需要关注的议题和研究方向, 以期为菌根真菌与吸收根功能性状之间关系的研究提供借鉴。

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.

Changes in the production and turnover of roots in forests and grasslands in response to rising atmospheric CO2 \nconcentrations, elevated temperatures, altered precipitation, or nitrogen deposition could be a key link between \nplant responses and longer‐term changes in soil organic matter and ecosystem carbon balance. Here we summarize \nthe experimental observations, ideas, and new hypotheses developed in this area in the rest of this volume. Three \ncentral 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.

Contents Summary 670 I. Introduction 671 II. Principle 1 - Plant respiration performs three distinct functions 673 III. Principle 2 - Metabolic pathway flexibility underlies plant respiratory performance 676 IV. Principle 3 - Supply and demand interact over time to set plant respiration rate 677 V. Principle 4 - Plant respiratory acclimation involves adjustments in enzyme capacities 679 VI. Principle 5 - Respiration is a complex trait that helps to define, and is impacted by, plant lifestyle strategies 680 VII. Future directions 680 Acknowledgements 682 References 682 SUMMARY: Respiration is a core biological process that has important implications for the biochemistry, physiology, and ecology of plants. The study of plant respiration is thus conducted from several different perspectives by a range of scientific disciplines with dissimilar objectives, such as metabolic engineering, crop breeding, and climate-change modelling. One aspect in common among the different objectives is a need to understand and quantify the variation in respiration across scales of biological organization. The central tenet of this review is that different perspectives on respiration can complement each other when connected. To better accommodate interdisciplinary thinking, we identify distinct mechanisms which encompass the variation in respiratory rates and functions across biological scales. The relevance of these mechanisms towards variation in plant respiration are explained in the context of five core principles: (1) respiration performs three distinct functions; (2) metabolic pathway flexibility underlies respiratory performance; (3) supply and demand interact over time to set respiration rates; (4) acclimation involves adjustments in enzyme capacities; and (5) respiration is a complex trait that helps to define, and is impacted by, plant lifestyle strategies. We argue that each perspective on respiration rests on these principles to varying degrees and that broader appreciation of how respiratory variation occurs can unite research across scales.© 2018 The Authors. New Phytologist © 2018 New Phytologist Trust.

Although fine roots are important components of the global carbon cycle, there is limited understanding of root structure-function relationships among species. We determined whether root respiration rate and decomposability, two key processes driving carbon cycling but always studied separately, varied with root morphological and chemical traits, in a coordinated way that would demonstrate the existence of a root economics spectrum (RES). Twelve traits were measured on fine roots (diameter ≤ 2 mm) of 74 species (31 graminoids and 43 herbaceous and dwarf shrub eudicots) collected in three biomes. The findings of this study support the existence of a RES representing an axis of trait variation in which root respiration was positively correlated to nitrogen concentration and specific root length and negatively correlated to the root dry matter content, lignin : nitrogen ratio and the remaining mass after decomposition. This pattern of traits was highly consistent within graminoids but less consistent within eudicots, as a result of an uncoupling between decomposability and morphology, and of heterogeneity of individual roots of eudicots within the fine-root pool. The positive relationship found between root respiration and decomposability is essential for a better understanding of vegetation-soil feedbacks and for improving terrestrial biosphere models predicting the consequences of plant community changes for carbon cycling.© 2016 CNRS. New Phytologist © 2016 New Phytologist Trust.

As sessile organisms, root plasticity enables plants to forage for and acquire nutrients in a fluctuating underground environment. Here, we use genetic and genomic approaches in a "split-root" framework--in which physically isolated root systems of the same plant are challenged with different nitrogen (N) environments--to investigate how systemic signaling affects genome-wide reprogramming and root development. The integration of transcriptome and root phenotypes enables us to identify distinct mechanisms underlying "N economy" (i.e., N supply and demand) of plants as a system. Under nitrate-limited conditions, plant roots adopt an "active-foraging strategy", characterized by lateral root outgrowth and a shared pattern of transcriptome reprogramming, in response to either local or distal nitrate deprivation. By contrast, in nitrate-replete conditions, plant roots adopt a "dormant strategy", characterized by a repression of lateral root outgrowth and a shared pattern of transcriptome reprogramming, in response to either local or distal nitrate supply. Sentinel genes responding to systemic N signaling identified by genome-wide comparisons of heterogeneous vs. homogeneous split-root N treatments were used to probe systemic N responses in Arabidopsis mutants impaired in nitrate reduction and hormone synthesis and also in decapitated plants. This combined analysis identified genetically distinct systemic signaling underlying plant N economy: (i) N supply, corresponding to a long-distance systemic signaling triggered by nitrate sensing; and (ii) N demand, experimental support for the transitive closure of a previously inferred nitrate-cytokinin shoot-root relay system that reports the nitrate demand of the whole plant, promoting a compensatory root growth in nitrate-rich patches of heterogeneous soil.

The promise of "trait-based" plant ecology is one of generalized prediction across organizational and spatial scales, independent of taxonomy. This promise is a major reason for the increased popularity of this approach. Here, we argue that some important foundational assumptions of trait-based ecology have not received sufficient empirical evaluation. We identify three such assumptions and, where possible, suggest methods of improvement: (i) traits are functional to the degree that they determine individual fitness, (ii) intraspecific variation in functional traits can be largely ignored, and (iii) functional traits show general predictive relationships to measurable environmental gradients.

Processes in the soil remain among the least well-characterized components of the carbon cycle. Arbuscular mycorrhizal (AM) fungi are ubiquitous root symbionts in many terrestrial ecosystems and account for a large fraction of photosynthate in a wide range of ecosystems; they therefore play a key role in the terrestrial carbon cycle. A large part of the fungal mycelium is outside the root (the extraradical mycelium, ERM) and, because of the dispersed growth pattern and the small diameter of the hyphae (<5 micrometers), exceptionally difficult to study quantitatively. Critically, the longevity of these fine hyphae has never been measured, although it is assumed to be short. To quantify carbon turnover in these hyphae, we exposed mycorrhizal plants to fossil ("carbon-14-dead") carbon dioxide and collected samples of ERM hyphae (up to 116 micrograms) over the following 29 days. Analyses of their carbon-14 content by accelerator mass spectrometry (AMS) showed that most ERM hyphae of AM fungi live, on average, 5 to 6 days. This high turnover rate reveals a large and rapid mycorrhizal pathway of carbon in the soil carbon cycle.

The residence time of fine-root carbon in soil is one of the least understood aspects of the global carbon cycle, and fine-root dynamics are one of the least understood aspects of plant function. Most recent studies of these belowground dynamics have used one of two methodological strategies. In one approach, based on analysis of carbon isotopes, the persistence of carbon is inferred; in the other, based on direct observations of roots with cameras, the longevity of individual roots is measured. We show that the contribution of fine roots to the global carbon cycle has been overstated because observations of root lifetimes systematically overestimate the turnover of fine-root biomass. On the other hand, isotopic techniques systematically underestimate the turnover of individual roots. These differences, by virtue of the separate processes or pools measured, are irreconcilable.

•  Since mycorrhizal fungi constitute an important component of the soil-plant interface, their responses to changes in nutrient availability may mediate shifts in ecosystem function. We tested the hypothesis that initial soil nutrient availability may determine effects of nitrogen (N) and phosphorus (P) additions on the growth and community of arbuscular mycorrhizal (AM) fungi. •  Extraradical hyphal lengths and degree of root colonization of AM fungi were measured in control and fertilized plots along a soil fertility gradient in Hawaii. Responses of individual AM genera were assessed through immunofluorescent labeling. •  The AM biomass was increased by N and P additions in the N- and P-limited sites, respectively, and reduced by P fertilization in the fertile site only. The abundance of Scutellospora was lower under N than under P fertilization, whereas the incidence of Glomus was higher in the fertile site than the N-limited site. Gigaspora and Acaulospora did not vary among sites or treatments. •  Our results indicate that a decrease in AM abundance following nutrient additions cannot be assumed to occur and the effects may differ among AM genera and ecosystems with varying soil nutrients. Limitation of N and P may be one possible explanation.

Understanding root processes at the whole-plant or ecosystem scales requires an accounting of the range of functions within a root system. Studying root traits based on their branching order can be a powerful approach to understanding this complex system. The current study examined the highly branched root system of the ericoid plant, Vaccinium corymbosum L. (highbush blueberry) by classifying its root orders with a modified version of the morphometric approach similar to that used in hydrology for stream classification. Root anatomy provided valuable insight into variation in root function across orders. The more permanent portion of the root system occurred in 4th- and higher-order roots. Roots in these orders had radial growth; the lowest specific root length, N:C ratios, and mycorrhizal colonization; the highest tissue density and vessel number; and the coarsest root diameter. The ephemeral portion of the root system was mainly in the first three root orders. First- and 2nd-order roots were nearly anatomically identical, with similar mycorrhizal colonization and diameter, and also, despite being extremely fine, median lifespans were not very short (115-120 d; estimated with minirhizotrons). Our research underscores the value of examining root traits by root order and its implications to understanding belowground processes.

Fine-root traits play key roles in ecosystem processes, but the drivers of fine-root trait diversity remain poorly understood. The plant economic spectrum (PES) hypothesis predicts that leaf and root traits evolved in coordination. Mycorrhizal association type, plant growth form and climate may also affect root traits. However, the extent to which these controls are confounded with phylogenetic structuring remains unclear. Here we compiled information about root and leaf traits for > 600 species. Using phylogenetic relatedness, climatic ranges, growth form and mycorrhizal associations, we quantified the importance of these factors in the global distribution of fine-root traits. Phylogenetic structuring accounts for most of the variation for all traits excepting root tissue density, with root diameter and nitrogen concentration showing the strongest phylogenetic signal and specific root length showing intermediate values. Climate was the second most important factor, whereas mycorrhizal type had little effect. Substantial trait coordination occurred between leaves and roots, but the strength varied between growth forms and clades. Our analyses provide evidence that the integration of roots and leaves in the PES requires better accounting of the variation in traits across phylogenetic clades. Inclusion of phylogenetic information provides a powerful framework for predictions of belowground functional traits at global scales.© 2017 The Authors. New Phytologist © 2017 New Phytologist Trust.

Contents 1159 I. 1159 II. 1161 III. 1164 IV. 1166 1167 References 1167 SUMMARY: The search for a root economics spectrum (RES) has been sparked by recent interest in trait-based plant ecology. By analogy with the one-dimensional leaf economics spectrum (LES), fine-root traits are hypothesised to match leaf traits which are coordinated along one axis from resource acquisitive to conservative traits. However, our literature review and meta-level analysis reveal no consistent evidence of an RES mirroring an LES. Instead the RES appears to be multidimensional. We discuss three fundamental differences contributing to the discrepancy between these spectra. First, root traits are simultaneously constrained by various environmental drivers not necessarily related to resource uptake. Second, above- and belowground traits cannot be considered analogues, because they function differently and might not be related to resource uptake in a similar manner. Third, mycorrhizal interactions may offset selection for an RES. Understanding and explaining the belowground mechanisms and trade-offs that drive variation in root traits, resource acquisition and plant performance across species, thus requires a fundamentally different approach than applied aboveground. We therefore call for studies that can functionally incorporate the root traits involved in resource uptake, the complex soil environment and the various soil resource uptake mechanisms - particularly the mycorrhizal pathway - in a multidimensional root trait framework.© 2016 Wageningen University. New Phytologist © 2016 New Phytologist Trust.

Plant trait variation drives plant function, community composition, and ecosystem processes. However, our current understanding of trait variation disproportionately relies on aboveground observations. Here we integrate root traits into the global framework of plant form and function. We developed and tested an overarching conceptual framework that integrates two recently identified root trait gradients with a well-established aboveground plant trait framework. We confronted our novel framework with published relationships between above- and belowground trait analogues and with multivariate analyses of aboveground and belowground traits of 2510 species. Our traits represent the leaf- and root conservation gradients (specific leaf area, leaf and root nitrogen concentration and root tissue density), the root collaboration gradient (root diameter and specific root length), and the plant size gradient (plant height and rooting depth). We found that an integrated, whole-plant trait space required as much as four axes. The two main axes represented the fast-slow 'conservation' gradient on which leaf and fine-root traits were well aligned, and the 'collaboration' gradient in roots. The two additional axes were separate, orthogonal plant size axes for height and rooting depth. This perspective on the multi-dimensional nature of plant trait variation better encompasses plant function and influence on the surrounding environment.This article is protected by copyright. All rights reserved.

When ecologically important plant traits are correlated they may be said to constitute an ecological 'strategy' dimension. Through identifying these dimensions and understanding their inter-relationships we gain insight into why particular trait combinations are favoured over others and into the implications of trait differences among species. Here we investigated relationships among several traits, and thus the strategy dimensions they represented, across 2134 woody species from seven Neotropical forests.Six traits were studied: specific leaf area (SLA), the average size of leaves, seed and fruit, typical maximum plant height, and wood density (WD). Trait relationships were quantified across species at each individual forest as well as across the dataset as a whole. 'Phylogenetic' analyses were used to test for correlations among evolutionary trait-divergences and to ascertain whether interspecific relationships were biased by strong taxonomic patterning in the traits.The interspecific and phylogenetic analyses yielded congruent results. Seed and fruit size were expected, and confirmed, to be tightly related. As expected, plant height was correlated with each of seed and fruit size, albeit weakly. Weak support was found for an expected positive relationship between leaf and fruit size. The prediction that SLA and WD would be negatively correlated was not supported. Otherwise the traits were predicted to be largely unrelated, being representatives of putatively independent strategy dimensions. This was indeed the case, although WD was consistently, negatively related to leaf size.The dimensions represented by SLA, seed/fruit size and leaf size were essentially independent and thus conveyed largely independent information about plant strategies. To a lesser extent the same was true for plant height and WD. Our tentative explanation for negative WD-leaf size relationships, now also known from other habitats, is that the traits are indirectly linked via plant hydraulics.

Leaf size varies by over a 100,000-fold among species worldwide. Although 19th-century plant geographers noted that the wet tropics harbor plants with exceptionally large leaves, the latitudinal gradient of leaf size has not been well quantified nor the key climatic drivers convincingly identified. Here, we characterize worldwide patterns in leaf size. Large-leaved species predominate in wet, hot, sunny environments; small-leaved species typify hot, sunny environments only in arid conditions; small leaves are also found in high latitudes and elevations. By modeling the balance of leaf energy inputs and outputs, we show that daytime and nighttime leaf-to-air temperature differences are key to geographic gradients in leaf size. This knowledge can enrich "next-generation" vegetation models in which leaf temperature and water use during photosynthesis play key roles.Copyright © 2017, American Association for the Advancement of Science.

Ecosystems are vulnerable to large areas of rocky desertification, which results in patchy soils and stone-inlaid soils. Karst landforms are typically characterized by heterogeneous phosphorus (P) distributions in soils at high calcium (Ca), but root foraging behavior has not been fully documented in agronomical plants. In this study, Bidens pilosa L. and Plantago asiatica L. were raised in pots in a simulated soil environment with sands at high Ca (2 g kg−1) and low Ca (0.63 g kg−1) levels. Inner spaces were divided into four sections to receive P in homogeneous (Homo.) (four quarters: 2 mg P kg−1) or heterogenous (Hete.) (one quarter: 8 mg P kg−1; three quarters: no-P input) patterns. Both species had longer roots in high P sections compared to no P sections. Foraging scale (highest length or surface-area(SA)) was higher in P. asiatica plants subjected to the Hete. pattern than to the Homo. pattern in low Ca pots. Foraging precision (length or SA differences between P patches as a proportion of the total) was also higher for P. asiatica subjected to the Hete. pattern but did not change in response to Ca level or P placement pattern. Overall, P. asiatica has a higher foraging ability than B. pilosa because of higher levels of foraging scale and precision from high-P (8 mg kg−1) patches in soils subjected to low Ca (0.63 g kg−1).

Root anatomical traits play crucial roles in understanding root functions and root form-function linkages. However, the root anatomy and form-function linkages of monocotyledonous and dicotyledonous herbs remain largely unknown. We measured order-based anatomical traits and mycorrhizal colonization rates of 32 perennial herbs of monocotyledons and dicotyledons in a temperate steppe. For monocots, relative constant proportion of cortex and mycorrhizal colonization rates, but increased cell-wall thickening of the endodermis and proportion of stele were observed across root orders, indicating a slight reduction in absorption capacity and improvement in transportation capacity across orders. For dicots, the cortex and mycorrhizal colonization disappeared in the fourth-order and/or fifth-order roots, whereas the secondary vascular tissue increased markedly, suggesting significant transition of root functions from absorption to transportation across root orders. The allometric relationships between stele and cortex differed across root orders and plant groups, suggesting different strategies to coordinate the absorption and transportation functions among plant groups. In summary, our results revealed different functional transition patterns across root orders and distinct strategies for coordinating the absorption and transportation of root system between monocots and dicots. These findings will contribute to our understanding of the root form and functions in herbaceous species.© 2022 The Authors. New Phytologist © 2022 New Phytologist Foundation.