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Plant invasion has severely damaged ecosystem stability and species diversity worldwide. The cooperation between arbuscular mycorrhizal fungi (AMF) and plant roots is often affected by changes in the external environment. Exogenous phosphorus (P) addition can alter the root absorption of soil resources, thus regulating the root growth and development of exotic and native plants. However, it remains unclear how exogenous P addition regulates the root growth and development of exotic and native plants mediated by AMF, affecting the exotic plant invasion. In this experiment, the invasive plant Eupatorium adenophorum and native plant Eupatorium lindleyanum were selected and cultured under intraspecific (Intra-) competition and interspecific (Inter-) competition conditions, involving inoculation with (M+) and without AMF (M−) and three different levels of P addition including no addition (P0), addition with 15 mg P kg−1 soil (P15), and addition with 25 mg P kg−1 soil (P25) for the two species. Root traits of the two species were analyzed to study the response of the two species’ roots to AMF inoculation and P addition. The results showed that AMF significantly promoted the root biomass, length, surface area, volume, tips, branching points, and carbon (C), nitrogen (N), and P accumulation of the two species. Under M+ treatment, the Inter- competition decreased the root growth and nutrient accumulation of invasive E. adenophorum but increased the root growth and nutrient accumulation of native E. lindleyanum relative to the Intra- competition. Meanwhile, the exotic and native plants responded differently to P addition, exhibiting root growth and nutrient accumulation of invasive E. adenophorum increased with P addition, whereas native E. lindleyanum reduced with P addition. Further, the root growth and nutrition accumulation of native E. lindleyanum were higher than invasive E. adenophorum under Inter- competition. In conclusion, exogenous P addition promoted the invasive plant but reduced the native plant in root growth and nutrient accumulation regulated by AMF, although the native plant outcompeted the invasive plant when the two species competed. The findings provide a critical perspective that the anthropogenic P fertilizer addition might potentially contribute to the successful invasion of exotic plants.

Soybean [Glycine max (L.) Merr.], an important oilseed crop, is a low-cost source of protein and oil. In Southeast Asia and Africa, soybeans are widely cultivated for use as traditional food and feed and industrial purposes. Given the ongoing changes in global climate, developing crops that are resistant to climatic extremes and produce viable yields under predicted climatic conditions will be essential in the coming decades. To develop such crops, it will be necessary to gain a thorough understanding of the genetic basis of agronomic and plant root traits. As plant roots generally lie beneath the soil surface, detailed observations and phenotyping throughout plant development present several challenges, and thus the associated traits have tended to be ignored in genomics studies. In this study, we phenotyped 357 soybean landraces at the early vegetative (V2) growth stages and used a 180 K single-nucleotide polymorphism (SNP) soybean array in a genome-wide association study (GWAS) conducted to determine the phenotypic relationships among root traits, elucidate the genetic bases, and identify significant SNPs associated with root trait-controlling genomic regions/loci. A total of 112 significant SNP loci/regions were detected for seven root traits, and we identified 55 putative candidate genes considered to be the most promising. Our findings in this study indicate that a combined approach based on SNP array and GWAS analyses can be applied to unravel the genetic basis of complex root traits in soybean, and may provide an alternative high-resolution marker strategy to traditional bi-parental mapping. In addition, the identified SNPs, candidate genes, and diverse variations in the root traits of soybean landraces will serve as a valuable basis for further application in genetic studies and the breeding of climate-resilient soybeans characterized by improved root traits.

Mechanical hill wet-seeded rice machine is beneficial for establishing and growing uniform rows of seedlings. However, there is limited knowledge regarding the effects of the establishment of furrows on growth, lodging and yield, and their relationships with root traits. In this study, field experiments were conducted during 2012 and 2013 using two super rice varieties (hybrid rice Peizataifeng and inbred rice Yuxiangyouzhan) under three furrow establishment treatments (T1, both water and seed furrows were established by the machine; T2, only seed furrows were established by the machine; and T3, neither water nor seed furrows were established by the machine). Lodging index, lodging-related traits, grain yield, above-ground dry weight and root traits were measured. The results showed that the lodging index was significantly affected by the treatments with furrows (T1 and T2). The strongest lodging resistance was detected in the mechanical hill wet-seeded rice with furrow treatment (T1) in both 2012 and 2013. Lodging resistance was strongly related to the breaking resistance. No significant difference was found in grain yield or dry weight of the mechanical hill wet-seeded rice. Therefore, the mechanical hill wet-seeded rice with furrow treatment increased rice lodging resistance, which was related to root traits.

Non-structural carbohydrates (NSCs) facilitate plant adaptation to drought stress, characterize tree growth and survival ability, and buffer against external disturbances. Previous studies have focused on the distribution and dynamics of NSCs among different plant organs under drought conditions. However, discussion about the NSC levels of fine roots in different root branch orders is limited, especially the relationship between fine root trait variation and NSC content. The objective of the study was to shed light on the synergistic variation in fine root traits and NSC content in different root branch orders under different drought and soil substrate conditions. The 2-year-old Fraxinus mandshurica Rupr. potted seedlings were planted in three different soil substrates (humus, loam, and sandy–loam soil) and subjected to four drought intensities (CK, mild drought, moderate drought, and severe drought) for 2 months. With increasing drought intensity, the biomass of fine roots decreased significantly. Under the same drought intensity, seedlings in sandy–loam soil had higher root biomass, and the coefficient of variation of 5th-order roots (37.4, 44.5, and 53% in humus, loam, and sandy–loam soil, respectively) was higher than that of lower-order roots. All branch order roots of seedlings in humus soil had the largest specific root length (SRL) and specific root surface area (SRA), in addition to the lowest diameter. With increasing drought intensity, the SRL and average diameter (AD) of all root branch orders increased and decreased, respectively. The fine roots in humus soil had a higher soluble sugar (SS) content and lower starch (ST) content compared to the loam and sandy–loam soil. Additionally, the SS and ST contents of fine roots showed decreasing and increasing tendencies with increasing drought intensities, respectively. SS and ST explained the highest degree of the total variation in fine root traits, which were 32 and 32.1%, respectively. With increasing root order, the explanation of the variation in root traits by ST decreased (only 6.8% for 5th-order roots). The observed response in terms of morphological traits of different fine root branch orders of F. mandshurica seedlings to resource fluctuations ensures the maintenance of a low cost-benefit ratio in the root system development.

Assessing differences in plant functional traits (PFTs) along climatic gradients is potentially useful for understanding variation within and across populations, and for predicting their responses to climate change. This study investigates the intraspecific variability of several PFTs in Stipa tenacissima (Alpha grass) seedlings from different populations distributed across a climatic gradient. Seven populations from Tunisia to Spain within a 100–600 mm/year rainfall range were selected. Seedlings from each population were grown in a common garden. We expected the functional characteristics to differ among seedling populations according to their climatic gradient. The response patterns were helpful to predict acclimation and fitness under future climatic conditions in these populations. The seedling development analysis showed differences in PFTs among S. tenacissima populations. The biomass traits analysis revealed that higher above-ground biomass was related to higher below-ground development. The leaf traits proved that seedlings with longer leaf length would have less sclerophyllous leaves, a trade-off between productivity and drought resistance. The root traits analysis reflects seedling strategies to maximize resource uptake efficiency. PFTs showed several significant relationships with climatic conditions. The less rainfall, the higher plant allocation to root systems exploring soil. Higher mean temperatures were related to reduced root/plant development. The PFT analysis proves that species followed the ‘optimal partitioning theory’, in that plants preferentially allocate biomass to acquire the resource that most limits their development. However, both the environmental conditions and genetic diversity in S. tenacissima populations influenced seedling growth and behaviour to face ongoing climate change. AV’s work was partially funded by Projects CGL-2011-30531-CO2-02 and CGL2015-69773-C2-2-P MINECO/FEDER by the Spanish Government, and by the Prometeo Program (FEEDBACK project, 2009/006) by the Generalitat Valenciana. CEAM is supported by the Generalitat Valenciana.

Non-structural carbohydrates (NSCs) facilitate plant adaptation to drought stress, characterize tree growth and survival ability, and buffer against external disturbances. Previous studies have focused on the distribution and dynamics of NSCs among different plant organs under drought conditions. However, discussion about the NSC levels of fine roots in different root branch orders is limited, especially the relationship between fine root trait variation and NSC content. The objective of the study was to shed light on the synergistic variation in fine root traits and NSC content in different root branch orders under different drought and soil substrate conditions. The 2-year-old Fraxinus mandshurica Rupr. potted seedlings were planted in three different soil substrates (humus, loam, and sandy–loam soil) and subjected to four drought intensities (CK, mild drought, moderate drought, and severe drought) for 2 months. With increasing drought intensity, the biomass of fine roots decreased significantly. Under the same drought intensity, seedlings in sandy–loam soil had higher root biomass, and the coefficient of variation of 5th-order roots (37.4, 44.5, and 53% in humus, loam, and sandy–loam soil, respectively) was higher than that of lower-order roots. All branch order roots of seedlings in humus soil had the largest specific root length (SRL) and specific root surface area (SRA), in addition to the lowest diameter. With increasing drought intensity, the SRL and average diameter (AD) of all root branch orders increased and decreased, respectively. The fine roots in humus soil had a higher soluble sugar (SS) content and lower starch (ST) content compared to the loam and sandy–loam soil. Additionally, the SS and ST contents of fine roots showed decreasing and increasing tendencies with increasing drought intensities, respectively. SS and ST explained the highest degree of the total variation in fine root traits, which were 32 and 32.1%, respectively. With increasing root order, the explanation of the variation in root traits by ST decreased (only 6.8% for 5th-order roots). The observed response in terms of morphological traits of different fine root branch orders of F. mandshurica seedlings to resource fluctuations ensures the maintenance of a low cost-benefit ratio in the root system development.

Mangroves are among the world's most carbon-dense ecosystems, but have suffered extensive deforestation, prompting reforestation projects. The effects of mangrove reforestation on belowground carbon dynamics are poorly understood. In particular, we do not know how fine root production develops following mangrove reforestation, despite fine root production being a major carbon sink and an important control of mangrove soil accretion. Using minirhizotrons, we investigated fine root production and its depth variation along a chronosequence of mature Vietnamese mangroves. Our results showed that fine root production decreases strongly with stand age in the uppermost 32 cm of our soil profiles. In younger mangrove stands, fine root production declines with depth, possibly due to a vertical gradient in soil nutrient availability; while root production in the oldest stand is low at all depths and exhibits no clear vertical pattern. A major fraction of fine root production occurs deeper than 30 cm, depths that are commonly omitted from calculations of mangrove carbon budgets. Younger mangroves may accrue shallow soil organic matter faster than older mangroves. Therefore, root productivity and forest stand age should be accounted for when forecasting mangrove carbon budgets and resistance to sea-level rise.

International audience; Background and aims Identifying root types helps understand the diversity of the morphogenetic capacities of root axes. Unfortunately, root types are generally defined arbitrarily. We used an unsupervised clustering method to define root types and examine the ontogeny of the Pinus pinaster root system.Methods K-clustering was first used on four root traits, including three geometric architectural traits: basal cross-sectional area, root tropism, parent root tropism, and branching angle. Based on these groups, we assigned all the root axes to a root type.Results Clustering yielded the same five groups of lateral roots and explained the same percentage of variance (70%) whatever the tree age. This way, 11,004 root axes from 69 excavated root systems ranging from 3 to 50 years were assigned in 5 types. P. pinaster root axes showed large differences based on root tropism and parent root tropism intensity. The larger horizontal shallow roots represented most of the woody volume throughout the life cycle of the trees. The frame of the central part of the root system was almost completed in 4-year-old trees.Conclusions In addition to the taproot, five types of lateral roots were identified in P. pinaster based on root geometry and size, and no consistent age effect was reported. This method could improve the connexion between root trait data and root modelling.

Significance Genetically improving the root system architectures of plants is an effective strategy for developing climate-resilient crops. In this study, we revealed that a cloned rice quantitative trait locus associated with root growth angle, qSOR1, is a DRO1 homolog involved in root gravitropic responses. The loss-of-function allele qsor1 resulted in roots that developed on the soil surface and enabled plants to avoid the reducing stress found in saline paddy soils and, consequently, increased yields. We show that the DRO1 homologs could be useful for the controlled breeding of root system architectures that are adapted to the abiotic stress conditions caused by global climate change., The root system architecture (RSA) of crops can affect their production, particularly in abiotic stress conditions, such as with drought, waterlogging, and salinity. Salinity is a growing problem worldwide that negatively impacts on crop productivity, and it is believed that yields could be improved if RSAs that enabled plants to avoid saline conditions were identified. Here, we have demonstrated, through the cloning and characterization of qSOR1 (quantitative trait locus for SOIL SURFACE ROOTING 1), that a shallower root growth angle (RGA) could enhance rice yields in saline paddies. qSOR1 is negatively regulated by auxin, predominantly expressed in root columella cells, and involved in the gravitropic responses of roots. qSOR1 was found to be a homolog of DRO1 (DEEPER ROOTING 1), which is known to control RGA. CRISPR-Cas9 assays revealed that other DRO1 homologs were also involved in RGA. Introgression lines with combinations of gain-of-function and loss-of-function alleles in qSOR1 and DRO1 demonstrated four different RSAs (ultra-shallow, shallow, intermediate, and deep rooting), suggesting that natural alleles of the DRO1 homologs could be utilized to control RSA variations in rice. In saline paddies, near-isogenic lines carrying the qSOR1 loss-of-function allele had soil-surface roots (SOR) that enabled rice to avoid the reducing stresses of saline soils, resulting in increased yields compared to the parental cultivars without SOR. Our findings suggest that DRO1 homologs are valuable targets for RSA breeding and could lead to improved rice production in environments characterized by abiotic stress.

Root growth and development has become an important research topic for breeders and researchers based on a growing need to adapt plants to changing and more demanding environmental conditions worldwide. Over the last few years, genome-wide association studies (GWASs) became an important tool to identify the link between traits in the field and their genetic background. Here we give an overview of the current literature concerning GWASs performed on root system architecture (RSA) in plants. We summarize which root traits and approaches have been used for GWAS, mentioning their respective success rate towards a successful gene discovery. Furthermore, we zoom in on the current technical hurdles in root phenotyping and GWAS, and discuss future possibilities in this field of research.

Carbon (C) sequestration is receiving increasing scientific and political attention in a framework of greenhouse gasses mitigation. However, geotechnical soils have been neglected for their C sequestration potential, with the global attention focusing on agricultural and natural soils. In the present thesis project we aim to assess the potential of geotechnical embankments as C sink, and, through the study of plant species and soils showing contrasting features, shed light on SOC sequestration mechanisms and the role of the different actor involved. We aim not only to quantify the C gained and lost in soil, but even its origin (fresh new C input or old preexistent C) and how it is partitioned in different C pools characterized by different C stability (quality of stored C). First, we evaluated the C storage in different pools under soil sowed with 12 different herbaceous species in a 10 months experiment. Assessing different root traits allowed understanding the influence of root economic spectrum on C storage. We showed how traits linked to high labile C are linked to a higher C increase in the stable SILT+CLAY pool (

Carbon (C) sequestration is receiving increasing scientific and political attention in a framework of greenhouse gasses mitigation. However, geotechnical soils have been neglected for their C sequestration potential, with the global attention focusing on agricultural and natural soils. In the present thesis project we aim to assess the potential of geotechnical embankments as C sink, and, through the study of plant species and soils showing contrasting features, shed light on SOC sequestration mechanisms and the role of the different actor involved. We aim not only to quantify the C gained and lost in soil, but even its origin (fresh new C input or old preexistent C) and how it is partitioned in different C pools characterized by different C stability (quality of stored C). First, we evaluated the C storage in different pools under soil sowed with 12 different herbaceous species in a 10 months experiment. Assessing different root traits allowed understanding the influence of root economic spectrum on C storage. We showed how traits linked to high labile C are linked to a higher C increase in the stable SILT+CLAY pool (

Breeding new crop cultivars with efficient root systems carries great potential to enhance resource use efficiency and plant adaptation to unstable climates. Here, we evaluated the natural variation of root system architectural traits in a diverse spring barley association panel and conducted genome-wide association mapping to identify genomic regions associated with root traits. For six studied traits, root system depth, root spreading angle, seminal root number, total seminal root length, and average seminal root length 1.9- to 4.2-fold variations were recorded. Using a mixed linear model, 55 QTLs were identified cumulatively explaining between 12.1% of the phenotypic variance for seminal root number to 48.1% of the variance for root system depth. Three major QTLs controlling root system depth, root spreading angle and total seminal root length were found on Chr 2H (56.52 cM), Chr 3H (67.92 cM), and Chr 2H (76.20 cM) and explained 12.4%, 18.4%, and 22.2% of the phenotypic variation, respectively. Meta-analysis and allele combination analysis indicated that root system depth and root spreading angle are valuable candidate traits for improving grain yield by pyramiding of favorable alleles.

BACKGROUND AND AIMS: In several disciplines, identifying relevant root traits to characterize the root system architecture of species or genotypes is a crucial step. To address this question, we analysed the inter-specific variations of root architectural traits in two contrasting environments. METHODS: We sampled 60 species in natura, at two sites, each presenting homogeneous soil conditions. We estimated for each species and site a set of five traits used for the modelling of the root system architecture: extreme tip diameters (D(min) and D(max)), relative diameter range (Drange), mean inter-branch distance (IBD) and dominance slope between the diameters of parent and lateral roots (DlDm). KEY RESULTS: The five traits presented a highly significant species effect, explaining between 77 and 98 % of the total variation. D(min), D(max) and Drange were particularly determined by the species, while DlDm and IBD exhibited a higher percentage of environmental variations. These traits make it possible to confirm two main axes of variation: ‘fineness–density’ (defined by D(min) and IBD) and ‘dominance–heterorhizy’ (DlDm and Drange), that together accounted for 84 % of the variations observed. CONCLUSIONS: We confirmed the interest of these traits in the characterization of the root system architecture in ecology and genetics, and suggest using them to enrich the ‘root economic spectrum’.

Background and aims: In order to analyse root system architectures (RSAs) from captured images, a variety of manual (e.g. Data Analysis of Root Tracings, DART), semi-automated and fully automated software packages have been developed. These tools offer complementary approaches to study RSAs and the use of the Root System Markup Language (RSML) to store RSA data makes the comparison of measurements obtained with different (semi-) automated root imaging platforms easier. The throughput of the data analysis process using exported RSA data, however, should benefit greatly from batch analysis in a generic data analysis environment (R software). Methods: We developed an R package (archiDART) with five functions. It computes global RSA traits, root growth rates, root growth directions and trajectories, and lateral root distribution from DART-generated and/or RSML files. It also has specific plotting functions designed to visualise the dynamics of root system growth. Results: The results demonstrated the ability of the package’s functions to compute relevant traits for three contrasted RSAs (Brachypodium distachyon [L.] P. Beauv., Hevea brasiliensis Müll. Arg. and Solanum lycopersicum L.). Conclusions: This work extends the DART software package and other image analysis tools supporting the RSML format, enabling users to easily calculate a number of RSA traits in a generic data analysis environment.

Highlight Roots in narrow-leafed lupin (Lupinus angustifolius) exhibit large phenotypic and genetic diversity. An association between root traits and DArT markers demonstrates potential for a marker-assisted selection programme for this species., Narrow-leafed lupin (Lupinus angustifolius L.) is the predominant grain legume crop in southern Australia, contributing half of the total grain legume production of Australia. Its yield in Australia is hampered by a range of subsoil constraints. The adaptation of lupin genotypes to subsoil constraints may be improved by selecting for optimal root traits from new and exotic germplasm sources. We assessed root trait diversity and genetic diversity of a core collection of narrow-leafed lupin (111 accessions) using 191 Diversity Arrays Technology (DArT) markers. The genetic relationship among accessions was determined using the admixture model in STRUCTURE. Thirty-eight root-associated traits were characterized, with 21 having coefficient of variation values >0.5. Principal coordinate analysis and cluster analysis of the DArT markers revealed broad diversity among the accessions. An ad hoc statistics calculation resulted in 10 distinct populations with significant differences among and within them (P < 0.001). The mixed linear model test in TASSEL showed a significant association between all root traits and some DArT markers, with the numbers of markers associated with an individual trait ranging from 2 to 13. The percentage of phenotypic variation explained by any one marker ranged from 6.4 to 21.8%, with 15 associations explaining >10% of phenotypic variation. The genetic variation values ranged from 0 to 7994, with 23 associations having values >240. Root traits such as deeper roots and lateral root proliferation at depth would be useful for this species for improved adaptation to drier soil conditions. This study offers opportunities for discovering useful root traits that can be used to increase the yield of Australian cultivars across variable environmental conditions.

Background and aims : Our objectives were to evaluate changes in soil aggregate stability along a successional gradient, located in severely eroded Mediterranean gully bed ecosystems and to identify predictors of soil aggregate stability variations among several soil, root traits and plant community characteristics.Methods : We selected 75 plots in gully beds, representing five successional stages that differ in plant community composition, dominated by herbs, shrubs or trees according to successional stage. In each plot, we measured soil aggregate stability, basic soil characteristics, root traits and plant diversity indices.Results : Soil aggregate stability increased along the successional gradient, being thrice higher in tree-dominated communities as compared to grass-dominated communities. This increase was mainly driven by soil organic carbon (SOC) accumulation. In early successional stages showing low SOC (below 24 g.kg−1 or 12 g.kg−1 in some cases), fine sand content and the percentage of fine roots acted as co-drivers enhancing soil aggregate stability while silt content decreased it.Conclusion : Plant succession in severely eroded Mediterranean gully bed ecosystems is accompanied by a strong stabilization of soil aggregates, mainly driven by SOC accumulation and for early successional stages, by soil granulometry and root traits as co-drivers. Stimulating succession thus appears as a promising restoration strategy for severely eroded ecosystems

Summary Our objective was to calibrate a model of the root system architecture on several Poaceae species and to assess its value to simulate several ‘integrated’ traits measured at the root system level: specific root length (SRL), maximum root depth and root mass. We used the model ArchiSimple, made up of sub-models that represent and combine the basic developmental processes, and an experiment on 13 perennial grassland Poaceae species grown in 1.5-m-deep containers and sampled at two different dates after planting (80 and 120 d). Model parameters were estimated almost independently using small samples of the root systems taken at both dates. The relationships obtained for calibration validated the sub-models, and showed species effects on the parameter values. The simulations of integrated traits were relatively correct for SRL and were good for root depth and root mass at the two dates. We obtained some systematic discrepancies that were related to the slight decline of root growth in the last period of the experiment. Because the model allowed correct predictions on a large set of Poaceae species without global fitting, we consider that it is a suitable tool for linking root traits at different organisation levels.

13 pages, 3 figures, 4 tables, 81 references., 1. Plant species composition and community functional structure (i.e. trait composition at the community level) result from a hierarchy of environmental filters that constrain which species and traits tend to be dominant in a given habitat. 2. We quantified variation in community functional structure along natural gradients of soil resources using several above- and below-ground parameters and explored links among these attributes to determine whether plant resource economics can be applied at the community level in a Mediterranean rangeland of southern France. 3. Limitation by nitrogen, soil water and soil depth were the main ecological factors driving the functional response at the community level. Most of the community functional parameters considered in this study were more dependent on nitrogen limitation than on the other two factors, mostly related with the acquisition–conservation trade-off at both the leaf and the root level. 4. We found a strong coordination between above-ground and below-ground components, with a high level of concordance along the resource gradients explored. As an example, tissue dry matter content – both in leaves and roots – was positively related to nitrogen limitation. These findings indicate that the leaf economic spectrum paradigm (resource conservation in resource-poor habitats versus resource acquisition in resource-rich habitats) can be extrapolated to the below-ground component and extends to a plant community spectrum. 5. Changes in the functional structure of communities were promoted by two complementary components of variation: (i) the replacement of species with highly contrasting resource-use strategies and, to a lesser extent, (ii) the intraspecific variation in several above-ground traits. 6. Synthesis. This study showed that soil water and nutrient limitations are the main drivers controlling functional community structure in the Mediterranean rangelands studied and that shifts in this structure were mainly due to species turnover. In addition, we provided evidence for a plant community economics spectrum, based on a strong coordination between above- and below-ground components in these resource-limited communities., This study was supported by a postdoctoral MEC contract to I. M.P.R. and by the DIVHERBE project (French National Programme ECOGER). This is a paper from GDR 2574 (‘TRAITS’) of CNRS.