Belowground Strategies Research Articles

A trait-based root acquisition-defence-decomposition framework in angiosperm tree species

To adapt to the complex belowground environment, plants make trade-offs between root resource acquisition and defence ability. This includes forming partnerships with different types of root associating microorganisms, such as arbuscular mycorrhizal and ectomycorrhizal fungi. These trade-offs, by mediating root chemistry, exert legacy effects on nutrient release during decomposition, which may, in turn, affect the ability of new roots to re-acquire resources, thereby generating a feedback loop. However, the linkages at the basis of this potential feedback loop remain largely unquantified. Here, we propose a trait-based root ‘acquisition-defence-decomposition’ conceptual framework and test the strength of relevant linkages across 90 angiosperm tree species. We show that, at the plant species level, the root-fungal symbiosis gradient within the root economics space, root chemical defence (condensed tannins), and root decomposition rate are closely linked, providing support to this framework. Beyond the dichotomy between arbuscular mycorrhizal-dominated versus ectomycorrhizal-dominated systems, we suggest a continuous shift in feedback loops, from ‘high arbuscular mycorrhizal symbiosis-low defence-fast decomposition-inorganic nutrition’ by evolutionarily ancient taxa to ‘high ectomycorrhizal symbiosis-high defence-slow decomposition-organic nutrition’ by more modern taxa. This ‘acquisition-defence-decomposition’ framework provides a foundation for testable hypotheses on multidimensional linkages between species’ belowground strategies and ecosystem nutrient cycling in an evolutionary context.

Above‐ and belowground strategies of tropical montane tree species are coordinated and driven by small‐scale nitrogen availability

Abstract The question whether the strategies of above‐ and belowground plant organs are coordinated as predicted by the plant economics spectrum theory is still under debate. We aim to determine the leading dimensions of tree trait variation for above‐ and belowground functional traits, and test whether they represent spectra of adaptation along a soil fertility gradient in tropical Andean forests. We measured leaf, stem and fine root functional traits, and individual‐level soil nutrient availability for 433 trees from 52 species at three elevations between 1000 and 3000 m a.s.l. We found close coordination between above– and belowground functional traits related to the trade‐off between resource acquisition and conservation, whereas root diameter and specific root length formed an independent axis of covarying traits. The position of a tree species along the acquisition–conservation axis of the trait space was closely associated with local soil nitrogen, but not phosphorus, availability. Our results imply that above‐ and belowground plant functional traits determine at which edaphic microhabitats coexisting tree species can grow, which is potentially crucial for understanding community assembly in species‐rich tropical montane forests. Read the free Plain Language Summary for this article on the Journal blog.

The plant root economics space in relation to nutrient limitation in Eurasian herbaceous plant communities.

Plant species occupy distinct niches along a nitrogen-to-phosphorus (N:P) gradient, yet there is no general framework for belowground nutrient acquisition traits in relation to N or P limitation. We retrieved several belowground traits from databases, placed them in the "root economics space" framework, and linked these to a dataset of 991 plots in Eurasian herbaceous plant communities, containing plant species composition, aboveground community biomass and tissue N and P concentrations. Our results support that under increasing N:P ratio, belowground nutrient acquisition strategies shift from "fast" to "slow" and from "do-it-yourself" to "outsourcing", with alternative "do-it-yourself" to "outsourcing" strategies at both ends of the spectrum. Species' mycorrhizal capacity patterns conflicted with root economics space predictions based on root diameter, suggesting evolutionary development of alternative strategies under P limitation. Further insight into belowground strategies along nutrient stoichiometry is crucial for understanding the high abundance of threatened plant species under P limitation.

Trait divergence and opposite above- and below-ground strategies facilitate moso bamboo invasion into subtropical evergreen broadleaf forest.

Understanding the invasion of moso bamboo (Phyllostachys edulis) into adjacent evergreen broadleaf forest based on functional traits is crucial due to its significant influence on ecosystem processes. However, existing research has primarily focused on above- or below-ground traits in isolation, lacking a comprehensive integration of both. In this study, we conducted a trait-based analysis including 23 leaf traits and 11 root traits in three forest types - bamboo forest, mixed bamboo and broadleaf forest, and evergreen broadleaf forest - to investigate trait differences, phenotypic integration, and above- and below-ground resource strategies in bamboo and broadleaf species. Our findings demonstrated significant differences in leaf and root key traits between bamboo and broadleaf species, strongly supporting the "phenotypic divergence hypothesis". Bamboo exhibited stronger trait correlations compared to broadleaf species, indicating higher phenotypic integration. Above- and below-ground strategies were characterized by trade-offs rather than coordination, resulting in a multi-dimensional trait syndrome. Specifically, a unidimensional leaf economics spectrum revealed that bamboo with higher leaf N concentrations (LNC), P concentrations (LPC), and specific leaf area (SLA) adopted a "fast acquisitive" above-ground strategy, while broadleaf species with thicker leaves employed a "slow conservative" above-ground strategy. A two-dimensional root trait syndrome indicated a "conservation" gradient with bamboo adopting a "slow conservative" below-ground strategy associated with higher root tissue density (RTD), and broadleaf species exhibiting a "fast acquisitive" below-ground strategy linked to higher root N concentrations (RNC) and P concentrations (RPC), and a "collaboration" gradient probably ranging from broadleaf species with a "do-it-yourself" strategy characterized by high specific root length (SRL), to bamboo adopting an "outsourcing" strategy with thicker roots. In conclusion, key trait divergence from coexisting broadleaf species, higher phenotypic integration, and multi-dimensional opposite above- and below-ground resource strategies confer competitive advantages to moso bamboo, shedding light on the mechanistic understanding of its invasion into subtropical evergreen broadleaf forest and providing theoretical guidance for maintaining the stability of subtropical forest ecosystem.

To protect or to hide: Why not both? An investigation of fire-related strategies in Cerrado woody species

Two key strategies enable woody species persistence and survival in fire-prone ecosystems after fire: the aboveground protection of stems and buds by thick bark and the allocation of biomass belowground to specialized bud bearing storage organs – both strategies allowing plants to resprout new branches after the aboveground parts are damaged by fire. Here we investigate whether those two strategies can be combined with each other. We compared 24 woody species from the Cerrado (tropical savannas from Brazil) and analyzed their underground storage organs (USOs) in relation to their aboveground bark production and aerial bud protection – two key traits allowing species to first survive and then persist after fire – together with plant potential height. We then compared if bark growth rate, aerial bud protection and plant potential height are linked to the ecological function of the underground storage organs (on spot persistence vs clonal growth through lateral spreading). Species with woody rhizomes (capable of spreading laterally) better protected their aboveground stems with thicker bark when compared to species with xylopodium and root-crown organs (adapted for on spot basal resprouting). A clear division was found concerning how well species are protecting their aerial buds and the type of underground storage organ, with species spreading laterally displaying a greater aerial bud protection. The results suggest that the presence of a specialized organ belowground does not appear to be mutually exclusive with producing thick bark on aboveground stems. It does exist, however, a different expression of bark production and aerial bud protection between species displaying lateral spread and those persisting on spot, suggesting the existence of a trade-off between above- and belowground strategies in species displaying on spot persistence. This highlights that Cerrado species can combine different fire-survival strategies, further questioning which fire conditions promote each strategy and their combination and in which cases trade-offs occur.

Above‐ and belowground traits along a stress gradient: trade‐off or not?

The role of plant traits in shaping community assembly along environmental gradients is a topic of ongoing research. It is well accepted that plant traits of aboveground organs tend to be conservative in stressful conditions. However, there is limited understanding of how belowground traits respond. Plants may have similar strategies above and belowground, but an intriguing possibility is that there is a tradeoff between above and belowground traits of communities to both ensure efficient resource‐use and limit niche overlap along the gradient. To test this, we asked whether the response of above and belowground traits of communities is coordinated or not along a stress gradient in Mediterranean sand dune communities. We analyzed 80 vegetation plots in central Italy to test for coordinated vs independent patterns in above vs belowground plant traits using community weighted mean and standardized effect size of functional richness. Our results show that plant communities close to the sea, which experience higher stress, were characterized by higher convergence towards aboveground resource conservation and conservative water‐use strategies but belowground resource acquisition, consistent with a strong effect of habitat filtering and an above–belowground tradeoff favoring adaptation to harsh and dry conditions. At the opposite end of the gradient with lower stress, plants exhibited higher trait diversity for both above and belowground traits, but overall a dominance of aboveground fast resource acquisition and generally acquisitive water‐use strategies, combined with conservative belowground strategies. This suggests that fast growth rate aboveground was compensated by more conservative fine‐root strategies, but processes such as competition limited niche overlap overall. Our findings provide new insights into the relationship between functional traits and environmental gradients in plant communities, shedding light on the tradeoffs between the above and belowground dimensions.

Enhanced nodulation and phosphorus acquisition from sparingly-soluble iron phosphate upon treatment with arbuscular mycorrhizal fungi in chickpea.

The coordination/trade-off among below-ground strategies for phosphorus (P) acquisition, including root morphology, carboxylate exudation and colonisation by arbuscular mycorrhizal fungi (AMF), is not well understood. This is the first study investigating the relationships between root nodulation, morphology, carboxylates and colonisation by an indigenous community of AMF under varying P levels and source. Two chickpea genotypes with contrasting amounts of rhizosheath carboxylates were grown in pots at six P levels (from 0 to 160 μg g-1 ) as KH2 PO4 (KP, highly soluble) or FePO4 (FeP, sparingly soluble), with or without AMF (±AMF) treatment. Under both FeP and KP, the presence of AMF inhibited shoot growth and shoot branching, decreased total root length and specific root length, increased mean root diameter and root tissue density and reduced carboxylates. However, the role of AMF in acquiring P differed between the two P sources, with the enhanced P acquisition under FeP while not under KP. Co-inoculation of AMF and rhizobia enhanced nodulation under FeP, but not under KP. Our results suggest that the effects of AMF on shoot branching were mediated by cytokinins as the reduced shoot branching in FeP40 and KP40 under +AMF relative to -AMF coincided with a decreased concentration of cytokinins in xylem sap for both genotypes.

Complementary belowground strategies underlie species coexistence in an early successional forest.

Unravelling belowground strategies is critical for understanding species coexistence and successional dynamics; yet, our knowledge of nutrient acquisition strategies of forest species at different successional stages remains limited. We measured morphological (diameter, specific root length, and root tissue density), architectural (branching ratio), physiological (ammonium, nitrate, and glycine uptake rates) root traits, and mycorrhizal colonisation rates of eight coexisting woody species in an early successional plantation forest in subtropical China. By incorporating physiological uptake efficiency, we revealed a bi-dimensional root economics space comprising of an 'amount-efficiency' dimension represented by morphological and physiological traits, and a 'self-symbiosis' dimension dominated by architectural and mycorrhizal traits. The early pioneer species relied on root-fungal symbiosis, developing densely branched roots with high mycorrhizal colonisation rates for foraging mobile soil nitrate. The late pioneer species invested in roots themselves and allocated effort towards improving uptake efficiency of less-mobile ammonium. Within the root economics space, the covariation of axes with soil phosphorus availability also distinguished the strategy preference of the two successional groups. These results demonstrate the importance of incorporating physiological uptake efficiency into root economics space, and reveal a trade-off between expanding soil physical space exploration and improving physiological uptake efficiency for successional species coexistence in forests.

Differential aboveground-belowground adaptive strategies to alleviate N addition-induced P deficiency in two alpine coniferous forests

Increasing atmospheric nitrogen (N) deposition has resulted in phosphorus (P) limitation in multiple terrestrial ecosystems, yet how plants coordinate aboveground and belowground strategies to adapt to such P deficiency remains unclear. In this study, we conducted a field N fertilization experiment in two alpine coniferous plantations (Picea asperata Mast. and Pinus armandii Franch.) with different soil N availability on the eastern Tibetan Plateau of China, to examine N addition effects on plant nutrient limiting status and plant adaptive strategies corresponding to aboveground P conservation and belowground P acquisition. The results showed that N addition aggravated P deficiency in both plantations, as indicated by decreased needle P concentrations and increased N:P ratios, and that plant strategies for addressing such P deficiency differed in the two plantations with different initial soil N availabilities. In the P. asperata plantation with relatively high N availability, significantly enhanced needle phosphatase activity and shifts in P fraction allocation (downregulation of the structural P fraction and increased allocation to the residual P fraction) co-occurred with increased rhizosphere effects on phosphatase activity under N addition, indicating a synergistic strategy of aboveground P conservation and belowground P mining to alleviate P deficiency. In the P. armandii plantation with relatively low N availability, however, N addition only enhanced phosphatase activity and increased allocation to residual P fraction in the aboveground but had little effect on belowground P acquisition-associated traits, suggesting a decoupling relationship between aboveground P conservation and belowground P acquisition. This study highlights the vital significance of initial soil nutrient availability in regulating the coordination of aboveground and belowground strategic alternatives, emphasizing the need to integrate soil nutrient conditions for a holistic understanding of forest adaptation to anthropogenic N enrichment.

Transcriptome analysis reveals infection strategies employed by Fusarium graminearum as a root pathogen

The fungal pathogen Fusarium graminearum (Fg) infects both heads and roots of cereal crops causing several economically important diseases such as head blight, seedling blight, crown rot and root rot. Trichothecene mycotoxins such as deoxynivalenol (DON), a well-known virulence factor, produced by Fg during disease development is also an important health concern. Although how Fg infects above-ground tissues is relatively well studied, very little is known about molecular processes employed by the pathogen during below-ground infection. Also unknown is the role of DON during root infection. In the present study, we analyzed the transcriptome of Fg during root infection of the model cereal Brachypodium distachyon (Bd). We also compared our Fg transcriptome data obtained during Bd root infection with those reported during wheat head infection. These analyses suggested that both shared and unique infection strategies were employed by the pathogen during colonization of different host tissues. Several metabolite biosynthesis genes induced in Fg during root infection could be linked to phytohormone production, implying that the pathogen likely interferes with root specific defenses. In addition, to understand the role of DON in Fg root infection, we analyzed the transcriptome of the DON deficient Tri5 mutant. These analyses showed that the absence of DON had a significant effect on fungal transcriptional responses. Although DON was produced in infected roots, this mycotoxin did not act as a Fg virulence factor during root infection. Our results reveal new mechanistic insights into the below-ground strategies employed by Fg that may benefit the development of new genetic tools to combat this important cereal pathogen.

Linking soil nutrients and traits to seedling growth: A test of the plant economics spectrum

Investigating the link among plant growth rates, traits and local environmental heterogeneity is necessary for understanding forest dynamics and community assembly. Although recent results showed that aboveground traits are key for determining organism’s performance and main resource-use strategies, it is clear that organism performance is influenced by both above- and below-ground traits. However, we have limited understanding about how belowground strategies change in response to soil fertility and determine plant performance. We hypothesize that belowground traits have significant effects on plant performance and that shifts in soil nutrient variability are associated to shifts belowground traits. We tested these hypotheses by surveying plant communities including 3969 seedlings represented by 49 common species over a 10-year period, measured soil nutrients, including total nitrogen (TN), phosphorus (TP) and potassium (TK), and leaf and root traits to examine the links among soil nutrients, traits and growth. We first examined the coordination of leaf and root traits of community level, then used traits to predict individual and species mean height growth rate (RGRh). Finally, we tested for shifts in traits in response to gradients of soil nutrients. We found that community-level traits tended to be multidimensional. Species with acquisitive leaf traits, e.g., high leaf area ratio (LAR), phosphorus content (LP), specific leaf area (SLA) and low leaf dry matter content (LDMC), exhibited high growth rates. However, root traits were weak predictors of RGRh, all root traits were not significantly correlated to RGRh of species, only root tissue density (RTD), specific root area (SRA) and length (SRL) was significantly correlated with RGRh of individuals. Community-weighted mean traits only significantly changed along the gradients of limiting soil nutrients, especially for TK and TP. Species with high LN, SLA, root nitrogen content, SRA and SRL, and low tissue density associated with high TK and TP. Ultimately, multidimensional trait variations, and weak links between root traits and growth only partly support the plant economics spectrum (correlation among traits along one axis), but emphasizes that, beyond root traits, other resource uptake processes of roots should be linked to plant performances. Our findings provide further insights into the understanding of how ecological strategies regulate plant performances and shape potential responses of plant communities to environmental change.

Vertical fine-root distributions in five subalpine forest types shifts with soil properties across environmental gradients

Vertical fine-root distribution determines the potential for acquisition of resources throughout soil profiles; yet, variation among forest types and changes in vertical distribution with environments are poorly understood. We examined vertical root distributions of different forest communities to determine how belowground strategies shift across different forest types and along edaphic gradients. Specific root length and diameter of fine roots as well as fine-root biomass, length and area densities were measured in sequential soil layers at 10 cm depth increments across 118 forest plots representing five subalpine forest types. Evergreen forest types, including evergreen oaks, were more deeply rooted than birch forests. Differences in rooting depth were due to the dominant tree species identity, not to variations in shrub or herbaceous components. Within forest types, soil nutrients and physical properties contributed to shifts rooting depth but not root morphology. Vertical distributions of fine roots represent critical inputs of plant carbon into soils as well as different capacities for the acquisition of soil resources. Our findings identify consistent patterns of rooting distributions among forest types that may be predictable based on more easily measured root and soil properties and can improve efforts to model rooting depth profiles in forest communities.

Climate and soil nutrients differentially drive multidimensional fine root traits in ectomycorrhizal‐dominated alpine coniferous forests

Abstract Fine root traits vary greatly with environmental changes, but the understanding of root trait variation and its drivers is limited over broad geographical scales, especially for ectomycorrhizal (ECM)‐dominated conifers in alpine forests. Herein, the covariation patterns of and environmental controls for fine root traits among ECM‐dominated conifers were examined to test whether and how climate and soil nutrients differentially affect fine root trait variations. Eight traits of first‐ and second‐order roots were measured, that is, root diameter (RD), specific root length (SRL), branching intensity (BRI), root tissue density (RTD), mycorrhizal colonization rate (MCR) and concentrations of carbon (C), nitrogen (N) and phosphorus (P), across 76 alpine coniferous populations on the eastern Tibetan Plateau, China. Our results showed that variations of the fine root traits fell into two major dimensions: the first dimension (32.39% of the total variance) was mainly represented by RD and SRL, potentially conveying a trade‐off between root life span and efficiency of resource foraging; the second dimension (23.70% of the variance) represented coordinated variation for root nutrients (i.e. N and P) and RTD, which depicts the conservation‐acquisition trade‐off in resource uptake, that is, root economic spectrum. Variations in RD and SRL were mainly driven by climatic variables, characterized by a significant increase in RD and a decrease in SRL with increasing mean annual precipitation. In contrast, variations in fine root nutrients (i.e. N and P) and RTD were primarily driven by soil fertility, showing a significant increase in root N and P concentrations but a decrease in RTD with increasing soil resource levels. Synthesis. Our study clearly shows two distinct dimensions of the variation of fine root traits in ECM‐dominated alpine coniferous forests, providing further evidence of the inherent multidimensionality of root traits. Moreover, our findings highlight different roles of climatic and soil variables in driving the variation of fine root traits, potentially leading to the multidimensionality of root traits. This study provides new insights for understanding and predicting shifts in plant belowground strategies in climate‐sensitive alpine forests world‐wide.

Global plant-symbiont organization and emergence of biogeochemical cycles resolved by evolution-based trait modelling.

One of the most distinct but unresolved global patterns is the apparent variation in plant-symbiont nutrient strategies across biomes. This pattern is central to our understanding of plant-soil-nutrient feedbacks in the land biosphere, which, in turn, are essential for our ability to predict the future dynamics of the Earth system. Here, we present an evolution-based trait-modelling approach for resolving (1) the organization of plant-symbiont relationships across biomes worldwide and (2) the emergent consequences for plant community composition and land biogeochemical cycles. Using game theory, we allow plants to use different belowground strategies to acquire nutrients and compete within local plant-soil-nutrient cycles in boreal, temperate and tropical biomes. The evolutionarily stable strategies that emerge from this analysis allow us to predict the distribution of belowground symbioses worldwide, the sequence and timing of plant succession, the bistability of ecto- versus arbuscular mycorrhizae in temperate and tropical forests, and major differences in the land carbon and nutrient cycles across biomes. Our findings imply that belowground symbioses have been central to the evolutionary assembly of plant communities and plant-nutrient feedbacks at the scale of land biomes. We conclude that complex global patterns emerge from local between-organism interactions in the context of Darwinian natural selection and evolution, and that the underlying dynamics can be mechanistically probed by our low-dimensional modelling approach.

Role of way of life, latitude, elevation and climate on the richness and distribution of orchid species

We are experiencing climate warming that is likely to affect all regions worldwide, although in a different manner, when its consequences (e.g. increase of temperature, lower seasonality, lower environmental stochasticity) are considered. Thus, our prediction of how global change will affect distribution and survival of species can be estimated by using our knowledge, how species richness and distribution is related with latitude and elevation. We used 193 terrestrial orchid species and subspecies as an example and we classified them according to their root system. This trait represents the evolution of strategies for underground storage of resources and resource acquisition as well as a characteristics that can be described as a life history trait. Classification of orchid life history traits that focus on belowground strategies has never been examined in a macroecological study. We then explored the associations between species richness, mean niche breadth and mean distribution on one hand and selected predictors on the other hand, using regression techniques for all orchids, and then for their subsets with different root systems. The predictive power, as well as the significance of the predictors, was also tested using polynomial second order generalized linear models. Species richness for the three belowground strategies was significantly affected by the predictors, whereas their mean niche breadth and mean distribution were largely dependent on their evolutionary history. The correlations of mean niche breadth and mean distribution with maximum elevation, latitude and longitude were significant for all orchid taxa and their subsets. All the variables together accounted for almost 50% or more of the variance in each of the subsets. The maximum elevation was the most significant factor for rhizomatous and intermediate orchids, whereas minimum temperature in the coldest month was highly significant for the tuberous orchids. Spatial distribution of Greek orchids is associated with a combination of elevation, latitude and climate. The distributions of rhizomatous and intermediate orchids are mainly associated with the orographic configuration of Greece, whereas the tuberous orchids are widely distributed in the southern, central and north-western areas of Greece, where most of them are limited by the harsh winter climate.

Diverse belowground resource strategies underlie plant species coexistence and spatial distribution in three grasslands along a precipitation gradient.

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.

Functional groups of Sahelian trees in a semiarid agroforestry system of Senegal

Addressing plant responses to water stress is critical to understand the structure of plant communities in water-limited environments and to forecast their resilience to future changes in climate. In a semiarid agroforestry system in the Sahelian savannah of Leona (Senegal), we selected nine common tree species and explored their stress-resistance mechanisms. These species represent a variety of life forms and are of high regional socio-economic importance. We hypothesized that different species would show different suites of traits to cope with water stress and expected to identify functional groups differing in strategies to withstand water shortage. Along a dry and a wet season, we monitored four traits reflecting above- and below-ground strategies of resource acquisition such as predawn leaf water potential (ψₚd), specific leaf area (SLA), leaf thickness and leaf area index (LAI). We also measured two morphological traits: trunk diameter and tree height. LAI and ψₚd were measured six times during the dry and rainy seasons, and the other traits were measured once. We identified two functional classes subdivided into two functional groups of each class. The first class included deciduous and semi-deciduous species that generally had large SLA, low leaf thickness and small-to-intermediate inter-seasonal variations in ψₚd. The second class included evergreen species of two functional groups that differ in SLA, leaf thickness and the magnitude of inter-seasonal variations of ψₚd throughout the year. The four functional groups identified in this study represent plant strategies differing in their response to changing environmental conditions.

Seedling root responses to soil moisture and the identification of a belowground trait spectrum across three growth forms.

Root trait variation and plasticity could be key factors differentiating plant performance under drought. However, water manipulation and root measurements are rarely coupled empirically across growth forms to identify whether belowground strategies are generalizable across species. We measured seedling root traits across three moisture levels in 18 Mediterranean forbs, grasses, and woody species. Drought increased the root mass fraction (RMF) and decreased the relative proportion of thin roots (indicated by increased root diameters and decreased specific root length (SRL)), rates of root elongation and growth, plant nitrogen uptake, and plant growth. Although responses varied across species, plasticity was not associated with growth form. Woody species differed from forbs and grasses in many traits, but herbaceous groups were similar. Across water treatments, trait correlations suggested a single spectrum of belowground trade-offs related to resource acquisition and plant growth. While effects of SRL and RMF on plant growth shifted with drought, root elongation rate consistently represented this spectrum. We demonstrate that general patterns of root morphology and plasticity are identifiable across diverse species. Root trait measurements should enhance our understanding of belowground strategy and performance across growth forms, but it will be critical to incorporate plasticity and additional aspects of root function into these efforts.

Patterns in root traits of woody species hosting arbuscular and ectomycorrhizas: implications for the evolution of belowground strategies.

Root traits vary enormously among plant species but we have little understanding of how this variation affects their functioning. Of central interest is how root traits are related to plant resource acquisition strategies from soil. We examined root traits of 33 woody species from northeastern US forests that form two of the most common types of mutualisms with fungi, arbuscular mycorrhizas (AM) and ectomycorrhizas (EM). We examined root trait distribution with respect to plant phylogeny, quantifying the phylogenetic signal (K statistic) in fine root morphology and architecture, and used phylogenetically independent contrasts (PICs) to test whether taxa forming different mycorrhizal associations had different root traits. We found a pattern of species forming roots with thinner diameters as species diversified across time. Given moderate phylogenetic signals (K = 0.44–0.68), we used PICs to examine traits variation among taxa forming AM or EM, revealing that hosts of AM were associated with lower branching intensity (rPIC = −0.77) and thicker root diameter (rPIC = −0.41). Because EM evolved relatively more recently and intermittently across plant phylogenies, significant differences in root traits and colonization between plants forming AM and EM imply linkages between the evolution of these biotic interactions and root traits and suggest a history of selection pressures, with trade-offs for supporting different types of associations. Finally, across plant hosts of both EM and AM, species with thinner root diameters and longer specific root length (SRL) had less colonization (rPIC = 0.85, −0.87), suggesting constraints on colonization linked to the evolution of root morphology.