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Summary: Plant root system is significantly influenced by high soil levels of ammonium nitrogen, leading to reduced root elongation and enhanced lateral root branching. In Arabidopsis, these processes have been reported to be mediated by phytohormones and their downstream signaling pathways, while the controlling mechanisms remain elusive in crops.Through a transcriptome analysis of roots subjected to high/low ammonium treatments, we identified a cytokinin oxidase/dehydrogenase encoding gene, CKX3, whose expression is induced by high ammonium. Knocking out CKX3 and its homologue CKX8 results in shorter seminal roots, fewer lateral roots, and reduced sensitivity to high ammonium.Endogenous cytokinin levels are elevated by high ammonium or in ckx3 mutants. Cytokinin application results in shorter seminal roots and fewer lateral roots in wild‐type, mimicking the root responses of ckx3 mutants to high ammonium. Furthermore, CKX3 is transcriptionally activated by type‐B RR25 and RR26, and ckx3 mutants have reduced auxin content and signaling in roots under low ammonium.This study identified RR25/26‐CKX3‐cytokinin as a signal module that mediates root responses to external ammonium by modulating of auxin signaling in the root meristem and lateral root primordium. This highlights the critical role of cytokinin metabolism in regulating rice root development in response to ammonium. [ABSTRACT FROM AUTHOR]

Targeted cellular auxin distribution is required for morphogenesis and adaptive responses of plant organs. In Arabidopsis thaliana (Arabidopsis), this involves the prototypical auxin influx facilitator AUX1 and its LIKE-AUX1 (LAX) homologs, which act partially redundantly in various developmental processes. Interestingly, AUX1 and its homologs are not strictly essential for the Arabidopsis life cycle. Indeed, aux1 lax1 lax2 lax3 quadruple knock-outs are mostly viable and fertile, and strong phenotypes are only observed at low penetrance. Here we investigated the Brachypodium distachyon (Brachypodium) AUX1 homolog BdAUX1 by genetic, cell biological and physiological analyses. We report that BdAUX1 is essential for Brachypodium development. Bdaux1 loss-of-function mutants are dwarfs with aberrant flower development, and consequently infertile. Moreover, they display a counter-intuitive root phenotype. Although Bdaux1 roots are agravitropic as expected, in contrast to Arabidopsis aux1 mutants they are dramatically longer than wild type roots because of exaggerated cell elongation. Interestingly, this correlates with higher free auxin content in Bdaux1 roots. Consistently, their cell wall characteristics and transcriptome signature largely phenocopy other Brachypodium mutants with increased root auxin content. Our results imply fundamentally different wiring of auxin transport in Brachypodium roots and reveal an essential role of BdAUX1 in a broad spectrum of developmental processes, suggesting a central role for AUX1 in pooideae.

Background: Root system architecture (RSA) is an essential characteristic for efficient water and nutrient absorption in terrestrial plants; its plasticity enables plants to respond to different soil environments. Better understanding of root plasticity is important in developing stress-tolerant crops. Non-invasive techniques that can measure roots in soils nondestructively, such as X-ray computed tomography (CT), are useful to evaluate RSA plasticity. However, although RSA plasticity can be measured by tracking individual root growth, only a few methods are available for tracking individual roots from time-series three-dimensional (3D) images. Results: We developed a semi-automatic workflow that tracks individual root growth by vectorizing RSA from time-series 3D images via two major steps. The first step involves 3D alignment of the time-series RSA images by iterative closest point registration with point clouds generated by high-intensity particles in potted soils. This alignment ensures that the time-series RSA images overlap. The second step consists of backward prediction of vectorization, which is based on the phenomenon that the root length of the RSA vector at the earlier time point is shorter than that at the last time point. In other words, when CT scanning is performed at time point A and again at time point B for the same pot, the CT data and RSA vectors at time points A and B will almost overlap, but not where the roots have grown. We assumed that given a manually created RSA vector at the last time point of the time series, all RSA vectors except those at the last time point could be automatically predicted by referring to the corresponding RSA images. Using 21 time-series CT volumes of a potted plant of upland rice (Oryza sativa), this workflow revealed that the root elongation speed increased with age. Compared with a workflow that does not use backward prediction, the workflow with backward prediction reduced the manual labor time by 95%. Conclusions: We developed a workflow to efficiently generate time-series RSA vectors from time-series X-ray CT volumes. We named this workflow 'RSAtrace4D' and are confident that it can be applied to the time-series analysis of RSA development and plasticity. [ABSTRACT FROM AUTHOR]

The phytohormones ethylene and jasmonate play important roles in the adaptation of rice plants to salt stress. However, the molecular interactions between ethylene and jasmonate on rice seminal root growth under salt stress are unknown. In this study, the effects of NaCl on the homeostasis of ethylene and jasmonate, and on rice seminal root growth were investigated. Our results indicate that NaCl treatment promotes ethylene biosynthesis by up-regulating the expression of ethylene biosynthesis genes, whereas NaCl-induced ethylene does not inhibit rice seminal root growth directly, but rather indirectly, by promoting jasmonate biosynthesis. NaCl treatment also promotes jasmonate biosynthesis through an ethylene-independent pathway. Moreover, NaCl-induced jasmonate reduces meristem cell number and cell division activity via down-regulated expression of Oryza sativa PLETHORA (OsPLT) and cell division-related genes, respectively. Additionally, NaCl-induced jasmonate inhibits seminal root cell elongation by down-regulating the expression of cell elongation-related genes. Overall, salt stress promotes jasmonate biosynthesis through ethylene-dependent and -independent pathways in rice seminal roots, and jasmonate inhibits rice seminal root growth by inhibiting root meristem cell proliferation and root cell elongation. [ABSTRACT FROM AUTHOR]

The root of wheat consists of seminal and nodal roots. Comparatively speaking, fewer studies have been carried out on the nodal root system because of its disappearance at the early seedling stage under indoor environments. In this study, 196 accessions from the Huanghuai Wheat Region (HWR) were used to identify the characteristics of seminal and nodal root traits under different growth environments, including indoor hydroponic culture (IHC), outdoor hydroponic culture (OHC), and outdoor pot culture (OPC), for three growing seasons. The results indicated that the variation range of root traits in pot environment was larger than that in hydroponic environment, and canonical coefficients were the greatest between OHC and OPC (0.86) than those in other two groups, namely, IHC vs. OPC (0.48) and IHC vs. OHC (0.46). Most root traits were negatively correlated with spikes per area (SPA), grains per spike (GPS), and grain yield (GY), while all the seminal root traits were positively correlated with thousand-kernel weight (TKW). Genome-wide association study (GWAS) was carried out on root traits by using a wheat 660K SNP array. A total of 35 quantitative trait loci (QTLs)/chromosomal segments associated with root traits were identified under OPC and OHC. In detail, 11 and 24 QTLs were significantly associated with seminal root and nodal root traits, respectively. Moreover, 13 QTLs for number of nodal roots per plant (NRP) containing 14 stable SNPs, were distributed on chromosomes 1B, 2B, 3A, 4B, 5D, 6D, 7A, 7B, and Un. Based on LD and bioinformatics analysis, these QTLs may contain 17 genes closely related to NRP. Among them, TraesCS2B02G552500 and TraesCS7A02G428300 were highly expressed in root tissues. Moreover, the frequencies of favorable alleles of these 14 SNPs were confirmed to be less than 70% in the natural population, suggesting that the utilization of these superior genes in wheat root is still improving.

The root of wheat consists of seminal and nodal roots. Comparatively speaking, fewer studies have been carried out on the nodal root system because of its disappearance at the early seedling stage under indoor environments. In this study, 196 accessions from the Huanghuai Wheat Region (HWR) were used to identify the characteristics of seminal and nodal root traits under different growth environments, including indoor hydroponic culture (IHC), outdoor hydroponic culture (OHC), and outdoor pot culture (OPC), for three growing seasons. The results indicated that the variation range of root traits in pot environment was larger than that in hydroponic environment, and canonical coefficients were the greatest between OHC and OPC (0.86) than those in other two groups, namely, IHC vs. OPC (0.48) and IHC vs. OHC (0.46). Most root traits were negatively correlated with spikes per area (SPA), grains per spike (GPS), and grain yield (GY), while all the seminal root traits were positively correlated with thousand-kernel weight (TKW). Genome-wide association study (GWAS) was carried out on root traits by using a wheat 660K SNP array. A total of 35 quantitative trait loci (QTLs)/chromosomal segments associated with root traits were identified under OPC and OHC. In detail, 11 and 24 QTLs were significantly associated with seminal root and nodal root traits, respectively. Moreover, 13 QTLs for number of nodal roots per plant (NRP) containing 14 stable SNPs, were distributed on chromosomes 1B, 2B, 3A, 4B, 5D, 6D, 7A, 7B, and Un. Based on LD and bioinformatics analysis, these QTLs may contain 17 genes closely related to NRP. Among them, TraesCS2B02G552500 and TraesCS7A02G428300 were highly expressed in root tissues. Moreover, the frequencies of favorable alleles of these 14 SNPs were confirmed to be less than 70% in the natural population, suggesting that the utilization of these superior genes in wheat root is still improving. [ABSTRACT FROM AUTHOR]

Nitrogen (N) is an important nutrient influencing the growth and yield in rice. Root plasticity is a key trait in the higher uptake of nutrient from the soil, especially in conditions where some major nutrients such as N is under optimal. In this study, a total of 168 BC1F4 mapping population from a cross between Malay 2 and US-2 and their parents were grown in hydroponics with deficient (5 µM) and sufficient (500 µM) N for 8 days to identify putative quantitative trait loci (QTLs) associated with root elongation. Eighty-three (83) markers were used in QTL detection using composite interval mapping. Results showed that US-2 had significantly greater shoot length and number of nodal roots than Malay 2 under N-deficient conditions. Around 19.6 and 26.8% of the population in N-deficient and N-sufficient conditions, respectively, had either longer or similar seminal root length compared with US-2. For the allele distribution, 43.1% of the population had homozygous allele for Malay 2, 20.9% had homozygous allele for US-2, and 8.4% had heterozygous allele for both parents. A total of eight putative QTLs associated with seminal root length (qSRL11.1 and qSRL2.1); number of nodal roots (qNNR6.1, qNNR11.1, and qNNR11.2); root dry weight (qRDW11.1 and qRDW11.2); and shoot length (qSL2.1) in chromosome 2, 6, and 11 regions were detected under both N treatments. [ABSTRACT FROM AUTHOR]

Background: Roots are essential for drought adaptation because of their involvement in water and nutrient uptake. As the study of the root system architecture (RSA) is costly and time-consuming, it is not generally considered in breeding programs. Thus, the identification of molecular markers linked to RSA traits is of special interest to the breeding community. The reported correlation between the RSA of seedlings and adult plants simplifies its assessment. Methods: In this study, a panel of 170 bread wheat landraces from 24 Mediterranean countries was used to identify molecular markers associated with the seminal RSA and related traits: seminal root angle, total root number, root dry weight, seed weight and shoot length, and grain yield (GY). Results: A genome-wide association study identified 135 marker-trait associations explaining 6% to 15% of the phenotypic variances for root related traits and 112 for GY. Fifteen QTL hotspots were identified as the most important for controlling root trait variation and were shown to include 31 candidate genes related to RSA traits, seed size, root development, and abiotic stress tolerance (mainly drought). Co-location for root related traits and GY was found in 17 genome regions. In addition, only four out of the fifteen QTL hotspots were reported previously. Conclusions: The variability found in the Mediterranean wheat landraces is a valuable source of root traits to introgress into adapted phenotypes through marker-assisted breeding. The study reveals new loci affecting root development in wheat.

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Maize forms a complex root system with structurally and functionally diverse root types that are formed at different developmental stages to extract water and mineral nutrients from soil. In recent years proteomics has been intensively applied to identify proteins involved in shaping the three-dimensional architecture and regulating the function of the maize root system. With the help of developmental mutants, proteomic changes during the initiation and emergence of shoot-borne, lateral and seminal roots have been examined. Furthermore, root hairs were surveyed to understand the proteomic changes during the elongation of these single cell type structures. In addition, primary roots have been used to study developmental changes of the proteome but also to investigate the proteomes of distinct tissues such as the meristematic zone, the elongation zone as well as stele and cortex of the differentiation zone. Moreover, subcellular fractions of the primary root including cell walls, plasma membranes and secreted mucilage have been analyzed. Finally, the superior vigor of hybrid seedling roots compared to their parental inbred lines was studied on the proteome level. In summary, these studies provide novel insights into the complex proteomic interactions of the elaborate maize root system during development. [ABSTRACT FROM AUTHOR]

The relationships between allelochemicals and environmental factors are a key factor for the growth of plants under rotation. We investigated the allelopathic potential of Amaranthus cruentus L, grown under different temperature conditions in in vitro bioassays. An inhibitory effect on germination and growth of lettuce (Lactuca sativa L.), tomato (Solanum lycopersicum L.), pepper (Capsicum annuum L.) and cucumber (Cucumis sativus L.) was observed when seeds were subjected to the leaf litter of Amaranthus cruentus. Analysis from our study indicated that germination percentage was significantly affected by growth temperatures (T) of the amaranth (P ˂ 0.0001), litter concentration (C) (P ˂ 0.0001), vegetable type (V) (P ˂ 0.0001), the T × V interaction (P = 0.0041) and V × C interaction (P ˂ 0.0001). Pepper was the most sensitive with a decline in germination percentage at increasing concentrations (0, 1 and 5 mg ml-1) of litter. Hypocotyl and seminal root lengths were adversely influenced by the plant litter for all the temperature treatments, although effects were most severe when exposed to the leaf litter of the hot temperature treatment. The inhibition caused by the litter was dependent on growth temperature and concentration, while each vegetable species showed different levels of sensitivity.

Breeding for favourable root traits in food and energy crops should be considered in the context of interactions with soil biota, notably those that can increase the nutrient use efficiency of crops. Arbuscular mycorrhizal (AM) fungi can provide services to plants but a better understanding of the interactions between root traits and AM fungi is required to maximise these benefits. One source of intraspecific variation in root architecture is the allocation of resources to different root types. We hypothesized that different root types would have different traits and be colonised differently by AM fungi, either in the amount of colonisation or identity of colonisers. We studied communities colonising the seminal, coleoptile nodal and leaf nodal roots of seven Brachypodium distachyon accessions grown in three crop or pasture soils. Leaf nodal roots had low specific root length compared to coleoptile nodal and seminal roots, yet all three root types harboured similar AM fungal communities and levels of colonisation. Most of the variation in the AM fungal communities was explained by soil; significant proportions were explained by plant accession and root type but these effects were weak. Differential allocation of resources between root types is not a trait that could be selected to maximise beneficial interactions with AM fungi and we found no link between a root phenotypic trait (specific root length) and AM fungal colonisation. Accessions did vary in extent of colonisation by arbuscules, meaning B. distachyon may be a useful model to study mechanisms underlying the symbiotic interface and mycorrhizal growth response of cereals. [ABSTRACT FROM AUTHOR]

Morphological adaptation of roots is critical for plants to survive under waterlogging. In this study, we evaluated the capacity of wheat to form aerenchyma in seminal roots in combination with the growth angle of the roots. We used five Japanese cultivars from the waterlogging-prone Kanto-Kyushu region in Japan, and a non-Japanese cultivar, Bobwhite for comparison. Seedlings in pot culture were waterlogged at a 3-cm depth for 7 days. The first adverse effect of waterlogging on plant growth was a significant reduction of root dry mass. The reduction rate varied with the cultivar, and it was 19.2% in cv. Shiroganekomugi and 40.0% in cv. Norin 61. Root aerenchyma was initially observed on the 2nd day of waterlogging and developed until the 7th day, in all 6 cultivars. Quantitative analysis of the aerenchyma development revealed no significant difference in radial distribution among the cultivars, whereas a slight difference was found in the axial distribution. As a consequence, the heavier root weight of Shiroganekomugi was not related to either the radial or axial developing capacity of aerenchyma but might be due to the effect of its shallow root angle in the soil. These results suggest that the capacity to form aerenchyma in the seminal root is not sufficient for expression of waterlogging tolerance in the Japanese wheat cultivars.

Seminal roots of maize are pivotal for early seedling establishment. The maize mutant rootless concerning crown and seminal roots (rtcs) is defective in seminal root initiation during embryogenesis. In this study, the transcriptomes of wild-type and rtcs embryos were analyzed by RNA-Seq based on histological results at three stages of seminal root primordia formation. Hierarchical clustering highlighted that samples of each genotype grouped together along development. Determination of their gene activity status revealed hundreds of genes specifically transcribed in wildtype or rtcs embryos, while K-mean clustering revealed changes in gene expression dynamics between wild-type and rtcs during embryo development. Pairwise comparisons of rtcs and wild-type embryo transcriptomes identified 131 transcription factors among 3526 differentially expressed genes [false discovery rate (FDR) <5% and llog2Fcl≥1]. Among those, functional annotation highlighted genes involved in cell cycle control and phytohormone action, particularly auxin signaling. Moreover, in silico promoter analyses identified putative RTCS target genes associated with transcription factor action and hormone metabolism and signaling. Significantly, non-syntenic genes that emerged after the separation of maize and sorghum were over-represented among genes displaying RTCS-dependent expression during seminal root primordia formation. This might suggest that these non-syntenic genes came under the transcriptional control of the syntenic gene rtcs during seminal root evolution. Taken together, this study provides first insights into the molecular framework underlying seminal root initiation in maize and provides a starting point for further investigations of the molecular networks underlying RTCS-dependent seminal root initiation. [ABSTRACT FROM AUTHOR]

Polar auxin transport was inhibited in rice seedlings when they were treated with N- 1-naphthylphthalamic acid (NPA). The treatment reduced total root length and the number of lateral roots, and negatively affected gravitropism. The auxin content at the base of the seminal root significantly increased in the NPA treatment seedlings compared to the control. Lateral roots elongated along the seminal root axis after NPA treatment, but growth remained within the cortex. Lignin content in the basal region also increased at the same time and accumulated in the epidermis. These results suggest that NPA treatment prevents lateral roots from penetrating the hypodermis due to the hardening of hypodermis cell walls through the enhanced lignification, and the disturbed gravitropism caused by NPA treatment affected auxin flow. [ABSTRACT FROM AUTHOR]

Deep rooting of wheat has been suggested that it influences the tolerance to various environmental stresses. In this study, the nodes from which the deepest penetrated roots had emerged were examined in winter wheat. The wheat was grown in long tubes with or without mechanical stress and in large root boxes. The length and growth angle of each axile root were examined to analyze the difference in the vertical distribution of the roots between the two wheat cultivars, one with a deep and one with a shallow root system. In Shiroganekomugi, a Japanese winter wheat cultivar with a shallow root system, the rooting depths of the seminal and nodal roots decreased as the rooting nodes advanced acropetally. Six out of nine deepest roots were seminal root in the non-mechanical stress conditions. In Mutsubenkei, a Japanese winter wheat cultivar with a deep root system, grown in root boxes, not only the seminal roots but also the coleoptilar and the first nodal roots penetrated to a depth of more than 1.3 m in the root box, and became the deepest roots. In both cultivars, the seminal roots became the deepest roots under the mechanical stress conditions. There were no clear tendencies in the root growth angles among the rooting nodes in the wheat root system. This indicates that the length of the axile roots can explain the differences in the rooting depths among axile roots in a wheat root system. On the other hand, the axile roots of Mutsubenkei elongated significantly more vertically than those of Shiroganekomugi. This suggests that not only seminal but also nodal roots exhibit strong positive gravitropism and penetrate deeply in a cultivar with a deep root system. In wheat cultivars, it is likely that the extent of its Root Depth Index results partly from the gravitropic responses of both seminal and nodal roots.

Maize develops a complex root system composed of embryonic and post-embryonic roots. Spatio-temporal differences in the formation of these root types imply specific functions during maize development. A comparative transcriptomic study of embryonic primary and seminal, and post-embryonic crown roots of the maize inbred line B73 by RNA sequencing along with anatomical studies were conducted early in development. Seminal roots displayed unique anatomical features, whereas the organization of primary and crown roots was similar. For instance, seminal roots displayed fewer cortical cell files and their stele contained more meta-xylem vessels. Global expression profiling revealed diverse patterns of gene activity across all root types and highlighted the unique transcriptome of seminal roots. While functions in cell remodeling and cell wall formation were prominent in primary and crown roots, stress-related genes and transcriptional regulators were over-represented in seminal roots, suggesting functional specialization of the different root types. Dynamic expression of lignin biosynthesis genes and histochemical staining suggested diversification of cell wall lignification among the three root types. Our findings highlight a cost-efficient anatomical structure and a unique expression profile of seminal roots of the maize inbred line B73 different from primary and crown roots. [ABSTRACT FROM AUTHOR]

Optimization of root system architecture (RSA) traits is an important objective for modern wheat breeding. Linkage and association mapping for RSA in two recombinant inbred line populations and one association mapping panel of 183 elite durum wheat (Triticum turgidum L. var. durum Desf.) accessions evaluated as seedlings grown on filter paper/polycarbonate screening plates revealed 20 clusters of quantitative trait loci (QTLs) for root length and number, as well as 30 QTLs for root growth angle (RGA). Divergent RGA phenotypes observed by seminal root screening were validated by root phenotyping of field-grown adult plants. QTLs were mapped on a high-density tetraploid consensus map based on transcriptassociated Illumina 90K single nucleotide polymorphisms (SNPs) developed for bread and durum wheat, thus allowing for an accurate cross-referencing of RSA QTLs between durum and bread wheat. Among the main QTL clusters for root length and number highlighted in this study, 15 overlapped with QTLs for multiple RSA traits reported in bread wheat, while out of 30 QTLs for RGA, only six showed co-location with previously reported QTLs in wheat. Based on their relative additive effects/significance, allelic distribution in the association mapping panel, and co-location with QTLs for grain weight and grain yield, the RSA QTLs have been prioritized in terms of breeding value. Three major QTL clusters for root length and number (RSA_QTL_cluster_5#, RSA_QTL_cluster_6#, and RSA_QTL_cluster_12#) and nine RGA QTL clusters (QRGA.ubo- 2A.1, QRGA.ubo-2A.3, QRGA.ubo-2B.2/2B.3, QRGA.ubo-4B.4, QRGA.ubo-6A.1, QRGA.ubo-6A.2, QRGA.ubo-7A.1, QRGA. ubo-7A.2, and QRGA.ubo-7B) appear particularly valuable for further characterization towards a possible implementation of breeding applications in marker-assisted selection and/or cloning of the causal genes underlying the QTLs. [ABSTRACT FROM AUTHOR]

Developmental plasticity in lateral roots may be one of the key traits for the growth of rice plants under soil moisture fluctuations. We aimed to examine responses in seminal root system development to changing soil moisture for diverse rice cultivars. Special attention was paid to the two different types of lateral roots ; the generally long, thick L type capable of branching into higher orders, and the non-branching S type. Plants were grown in half-split polyvinyl chloride tubes fixed with transparent acrylic plate for root observation under glasshouse conditions. When plants were grown first under drought conditions, then rewatered, the seminal root system development in terms of dry weight and total length was promoted as compared with plants grown under continuously well-watered conditions in IR AT 109 and Dular, drought tolerant cultivars. Promoted production of L type lateral roots mainly contributed to the development of the longer seminal root system. Plants exposed to soil submergence before they were grown under drought conditions did not show such promoted responses in these two cultivars. However, in KDML 105, a drought tolerant cultivar, the production of especially L type laterals was substantially promoted under drought and rewatered conditions. Honenwase was characterized by the shallow root system and great reduction in root system length when soil moisture becomes limited. These facts show that genotypic variations exist in the plastic response of rice seminal root system and that the L type lateral root plays a key role in manifestation of this plasticity.

The growth and panicle characters of bread wheat (Triticum aestivum L.), in which only a single primary seminal root was allowed to grow by suppressing the growth of nodal roots, were analyzed in pots. The growth of the aboveground parts was not inferior to that of the control until the panicle formation stage due to vigorous growth of the primary seminal root. At the booting stage, however, the T-R ratio in weight was doubled as compared with the control, although the ratio of leaf area to root length was hardly changed. The grain weight per plant of the treated plants was about 80% of that in the controls, and the grains in the treated plants were more vitreous than the control. These results indicate that the primary seminal root remained functional until the ripening period in the plants without development of any other roots.