Water availability is undoubtedly one of the most important environmental factors affecting crop production. Drought causes a gradual deprivation of water in the soil from top to deep layers and can occur at diverse stages of plant development. Roots are the first organs that perceive water deficit in soil and their adaptive development contributes to drought adaptation. Domestication has contributed to a bottleneck in genetic diversity. Wild species or landraces represent a pool of genetic diversity that has not been exploited yet in breeding program. In this study, we used a collection of 230 two-row spring barley landraces to detect phenotypic variation in root system plasticity in response to drought and to identify new quantitative trait loci (QTL) involved in root system architecture under diverse growth conditions. For this purpose, young seedlings grown for 21 days in pouches under control and osmotic-stress conditions were phenotyped and genotyped using the barley 50k iSelect SNP array, and genome-wide association studies (GWAS) were conducted using three different GWAS methods (MLM GAPIT, FarmCPU, and BLINK) to detect genotype/phenotype associations. In total, 276 significant marker-trait associations (MTAs; p-value (FDR)< 0.05) were identified for root (14 and 12 traits under osmotic-stress and control conditions, respectively) and for three shoot traits under both conditions. In total, 52 QTL (multi-trait or identified by at least two different GWAS approaches) were investigated to identify genes representing promising candidates with a role in root development and adaptation to drought stress.

Rootstocks are fundamental for peach production, and their architectural root traits determine their performance. Root-system architecture (RSA) analysis is one of the key factors involved in rootstock selection. However, there are few RSA studies on Prunus spp., mostly due to the tedious and time-consuming labor of measuring below-ground roots. A root-phenotyping experiment was developed to analyze the RSA of seedlings from \'Okinawa\' and \'Guardian\'™ peach rootstocks. The seedlings were established in rhizoboxes and their root systems scanned and architecturally analyzed. The root-system depth:width ratio (D:W) throughout the experiment, as well as the root morphological parameters, the depth rooting parameters, and the root angular spread were estimated. The \'Okinawa\' exhibited greater root morphological traits, as well as the other parameters, confirming the relevance of the spatial disposition and growth pattern of the root system.

The root system architecture (RSA), being a key characteristic of the root economic spectrum, describes the spatial arrangement and positioning of roots that determines the plant\'s exploration of water and nutrients in the soil. Still, it remains poorly understood how the RSA of woody plants responds to the demand for water and nutrients in different soil environments and how the uptake of these resources is optimized. Here we selected single-species plantations of Cupressus funebris and determined their topological index (TI), revised topological index (q a and q b ), root link length (RLL), root branching rate (R b and R i :R i+1), and in situ soil physicochemical properties to assess which root foraging strategies adopt in different soil environments among Guang\'an City (GA), Suining City (SN), Mianyang City (MY), and Deyang City (DY) in China. We also tested the potential effects of different nutrients upon RSA according to its plastic phenotype. Principal component analysis (PCA) showed that levels of soil nutrients were the highest at DY, followed by MY and SN, and lower at GA. A dichotomous branching pattern was observed for GA, SN, and MY, but a herringbone branching pattern for DY. The RLL was ranked as GA, > SN, > MY > DY. The R b of GA, SN, and MY was significantly lower than that of DY (p < 0.05). Among the different city regions, values of R 1 /R 2 were the largest in different regions and those of R 4 /R 5 the smallest. The cross-sectional area of the root system did not differ between any two connected branch orders. The TI, q a , and RLL were significantly and negatively correlated with soil\'s water content, porosity, total nitrogen, total potassium, available nitrogen, and available phosphorus (p < 0.05), whereas they all had significant, positive relationships with soil temperature (p < 0.05). The R b was significantly and positively correlated with total potassium in soil (p < 0.05). Redundancy analysis showed that total potassium was the main factor driving variation in RSA. Our results emphasize that the RSA is capable of corresponding plastic alterations by changing its number of internal or external links and the root link length of fine roots vis-à-vis a heterogeneous environment, thereby optimizing the rates of water capture and space utilization.

Durum wheat (Triticum turgidum, 2n = 4x = AABB) includes several subspecies with differential characteristics in their root system architecture (RSA). Subspecies durum has longer and more vertical roots, while subspecies turgidum has smaller and shallower roots. The homeologous genes TtDro1A and TtDro1B of both subspecies have been identified and found to differ in their sizes, sequences and the proteins they encode. To determine whether there is a relationship between the level of expression of these two genes and the angle adopted by the roots of durum wheat seedlings, their expressions has been studied by RT-qPCR, both in the primary seminal root and in the other seminal roots. The results of the analyses showed that the TtDro1A gene is expressed 1.4 times more in the primary seminal root than in the other seminal roots. Furthermore, this gene is expressed 2.49 to 8.76 times more than TtDro1B depending on root type (primary or seminal) and subspecies. There are positive correlations between the expression ratio of both genes (TtDro1A/TtDro1B) and the mean of all root angles, the most vertical root angle and the most horizontal root angle of the seedlings. The higher the expression of TtDro1B gene, the lower the root growth angles.

Roots are important plant organs for the absorption of water and nutrients. To date, there have been few genome-wide association studies of maize root system architecture (RSA) in the field. The genetic basis of maize RSA is poorly understood, and the maize RSA-related genes that have been cloned are very limited. Here, 421 maize inbred lines of an association panel were planted to measure the root systems at the maturity stage, and a genome-wide association study was performed. There was a strong correlation among eight RSA traits, and the RSA traits were highly correlated with the aboveground plant architecture traits (e.g., plant height and ear leaf length, r = 0.13-0.25, p < 0.05). The RSA traits of the stiff stalk subgroup (SS) showed lower values than those of the non-stiff stalk subgroup (NSS) and tropical/subtropical subgroup (TST). Using the RSA traits, the genome-wide association study identified 63 SNPs and 189 candidate genes. Among them, nine candidate genes co-localized between RSA and aboveground architecture traits. A further co-expression analysis identified 88 candidate genes having high confidence levels. Furthermore, we identified four highly reliable RSA candidate genes, GRMZM2G099797, GRMZM2G354338, GRMZM2G085042, and GRMZM5G812926. This research provides theoretical support for the genetic improvement of maize root systems, and it identified candidate genes that may act as genetic resources for breeding.

The root tissues play important roles in water and nutrient acquisition, environmental adaptation, and plant development. In this study, a diversity panel of 388 wheat accessions was collected to investigate nine root system architecture (RSA) traits at the three-leaf stage under two growing environments: outdoor pot culture (OPC) and indoor pot culture (IPC). Phenotypic analysis revealed that root development was faster under OPC than that under IPC and a significant correlation was observed between the nine RSA traits. The 660K single-nucleotide polymorphism (SNP) chip was used for a genome-wide association study (GWAS). Significant SNPs with a threshold of -log10 (p-value) ≥ 4 were considered. Thus, 36 quantitative trait loci (QTLs), including 13 QTL clusters that were associated with more than one trait, were detected, and 31 QTLs were first identified. The QTL clusters on chromosomes 3D and 5B were associated with four and five RSA traits, respectively. Two candidate genes, TraesCS2A01G516200 and TraesCS7B01G036900, were found to be associated with more than one RSA trait using haplotype analysis, and preferentially expressed in the root tissues. These favourable alleles for RSA traits identified in this study may be useful to optimise the root system in wheat.

Water and nutrient acquisition is a critical function of plant root systems. Root system architecture (RSA) traits are often complex and controlled by many genes. This is the first genome-wide association study reporting genetic loci for RSA traits for field-grown soybean (Glycine max). A collection of 289 soybean genotypes was grown in three environments, root crowns were excavated, and 12 RSA traits assessed. The first two components of a principal component analysis of these 12 traits were used as additional aggregate traits for a total of 14 traits. Marker-trait association for RSA traits were identified using 31,807 single-nucleotide polymorphisms (SNPs) by a genome-wide association analysis. In total, 283 (non-unique) SNPs were significantly associated with one or more of the 14 root traits. Of these, 246 were unique SNPs and 215 SNPs were associated with a single root trait, while 26, four, and one SNPs were associated with two, three, and four root traits, respectively. The 246 SNPs marked 67 loci associated with at least one of the 14 root traits. Seventeen loci on 13 chromosomes were identified by SNPs associated with more than one root trait. Several genes with annotation related to processes that could affect root architecture were identified near these 67 loci. Additional follow-up studies will be needed to confirm the markers and candidate genes identified for RSA traits and to examine the importance of the different root characteristics for soybean productivity under a range of soil and environmental conditions.

Root hairs (RHs) are single-celled elongated epidermal cells and play a vital role in nutrient absorption, particularly for immobile minerals like phosphorus (P). As an adaptive response to P deficiency, an increase in RH length enhances root-soil contact and absorptive area for P absorption. Genetic variations have been reported for RH length and its response to P deficiency in plants. However, only a few association studies have been conducted to identify genes and genetic loci associated with RH length. Here, we screened desi chickpea accessions for RH length and its plasticity under P deficiency. Further, the genome-wide association study (GWAS) was conducted to identify the genetic loci associated with RH length in P deficient and sufficient conditions. Although high variability was observed in terms of RH length in diverse genotypes, majority of the accessions showed typical response of increase in RH length in low P. Genome-wide association mapping identified many SNPs with significant associations with RH length in P-sufficient and P-deficient conditions. A few candidate genes for RH length in P deficient (SIZ1-like and HAD superfamily protein) and sufficient (RSL2-like and SMAP1-like) conditions were identified which have known roles in RH development and P deficiency response or both. Highly associated loci and candidate genes identified in this study would be useful for genomic-assisted breeding to develop P-efficient chickpea.

Root systems can display variable architectures that contribute to nutrient foraging or to increase the tolerance of abiotic stress conditions. Root tip excision promotes the developmental progression of previously specified lateral root (LR) founder cells, which allows to easily measuring the branching capacity of a given root as regards its genotype and/or growth conditions. Here, we describe the natural variation among 120 Arabidopsis thaliana accessions in root system architecture (RSA) after root tip excision. Wound-induced changes in RSA were associated with 19 genomic loci using genome-wide association mapping. Three candidate loci associated with wound-induced LR formation were investigated. Sequence variation in the hypothetical protein encoded by the At4g01090 gene affected wound-induced LR development and its loss-of-function mutants displayed a reduced number of LRs after root tip excision. Changes in a histidine phosphotransfer protein putatively involved in cytokinin signaling were significantly associated with LR number variation after root tip excision. Our results provide a better understanding of some of the genetic components involved in LR capacity variation among accessions.

OBJECTIVE: Lateral root (LR) initiation (LRI) is a central process in root branching. Based on LR and/or LR primordium densities, it has been shown that nitric oxide (NO) promotes LRI. However, because NO inhibits primary root growth, we hypothesized that NO may have an opposite effect if the analysis is performed on a cellular basis. Using a previously proposed parameter, the LRI index (which measures how many LRI events take place along a root portion equivalent to the length of a single file of 100 cortical cells of average length), we addressed this hypothesis and illustrate here that the LRI index provides a researcher with a tool to uncover hidden but important information about root initiation. •
RESULTS: Arabidopsis thaliana roots were treated with an NO donor (sodium nitroprusside [SNP]) and/or an NO scavenger (2-(4-carboxyphenyl)-4,4,5,5-tetramethylimidazoline-1-oxyl-3-oxide [cPTIO]). LRI was analyzed separately in the root portions formed before and during the treatment. In the latter, SNP caused root growth inhibition and an increase in the LR density accompanied by a decrease in LRI index, indicating overall inhibitory outcome of the NO donor on branching. The inhibitory effect of SNP was reversed by cPTIO, showing the NO-specific action of SNP on LRI. •
CONCLUSIONS: Analysis of the LRI index permits the discovery of otherwise unknown modes of action of a substance on the root system formation. NO has a dual action on root branching, slightly promoting it in the root portion formed before the treatment and strongly inhibiting it in the root portion formed during the treatment.