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Efficient acquisition and use of available phosphorus from the soil is crucial for plant growth, development, and yield. With an ever‐increasing acreage of croplands with suboptimal available soil phosphorus, genetic improvement of sorghum germplasm for enhanced phosphorus acquisition from soil is crucial to increasing agricultural output and reducing inputs, while confronted with a growing world population and uncertain climate. Sorghum bicolor is a globally important commodity for food, fodder, and forage. Known for robust tolerance to heat, drought, and other abiotic stresses, its capacity for optimal phosphorus use efficiency (PUE) is still being investigated for optimized root system architectures (RSA). Whilst a few RSA‐influencing genes have been identified in sorghum and other grasses, the epigenetic impact on expression and tissue‐specific activation of candidate PUE genes remains elusive. Here, we present transcriptomic, epigenetic, and regulatory network profiling of RSA modulation in the BTx623 sorghum background in response to limiting phosphorus (LP) conditions. We show that during LP, sorghum RSA is remodeled to increase root length and surface area, likely enhancing its ability to acquire P. Global DNA 5‐methylcytosine and H3K4 and H3K27 trimethylation levels decrease in response to LP, while H3K4me3 peaks and DNA hypomethylated regions contain recognition motifs of numerous developmental and nutrient responsive transcription factors that display disparate expression patterns between different root tissues (primary root apex, elongation zone, and lateral root apex).

Aims The objective of this research was to develop a three-dimensional (3D) rhizosphere modeling capability for plant-soil interactions by integrating plant biophysics, water and ion uptake and release from individual roots, variably saturated flow, and multicomponent reactive transport in soil. Methods We combined open source software for simulating plant and soil interactions with parallel computing technology to address highly-resolved root system architecture (RSA) and coupled hydrobiogeochemical processes in soil. The new simulation capability was demonstrated on a model grass, Brachypodium distachyon . Results In our simulation, the availability of water and nutrients for root uptake is controlled by the interplay between 1) transpiration-driven cycles of water uptake, root zone saturation and desaturation; 2) hydraulic redistribution; 3) multicomponent competitive ion exchange; 4) buildup of ions not taken up during kinetic nutrient uptake; and 5) advection, dispersion, and diffusion of ions in the soil. The uptake of water and ions by individual roots leads to dynamic, local gradients in ion concentrations. Conclusion By integrating the processes that control the fluxes of water and nutrients in the rhizosphere, the modeling capability we developed will enable exploration of alternative RSAs and function to advance the understanding of the coupled hydro-biogeochemical processes within the rhizosphere.

Root system architecture (RSA) - the spatial configuration of a root system - is an important developmental and agronomic trait, with implications for overall plant architecture, growth rate and yield, abiotic stress resistance, nutrient uptake, and developmental plasticity in response to environmental changes. Root architecture is modulated by intrinsic, hormone-mediated pathways, intersecting with pathways that perceive and respond to external, environmental signals. The recent development of several non-invasive 2D and 3D root imaging systems has enhanced our ability to accurately observe and quantify architectural traits on complex whole-root systems. Coupled with the powerful marker-based genotyping and sequencing platforms currently available, these root phenotyping technologies lend themselves to large-scale genome-wide association studies, and can speed the identification and characterization of the genes and pathways involved in root system development. This capability provides the foundation for examining the contribution of root architectural traits to the performance of crop varieties in diverse environments. This review focuses on our current understanding of the genes and pathways involved in determining RSA in response to both intrinsic and extrinsic (environmental) response pathways, and provides a brief overview of the latest root system phenotyping technologies and their potential impact on elucidating the genetic control of root development in plants.

Flooding causes oxygen deprivation in soils. Plants adapt to low soil oxygen availability by changes in root morphology, anatomy, and architecture to maintain root system functioning. Essential traits include aerenchyma formation, a barrier to radial oxygen loss, and outgrowth of adventitious roots into the soil or the floodwater. We highlight recent findings of mechanisms of constitutive aerenchyma formation and of changes in root architecture. Moreover, we use modelling of internal aeration to demonstrate the beneficial effect of increasing cortex-to-stele ratio on sustaining root growth in waterlogged soils. We know the genes for some of the beneficial traits, and the next step is to manipulate these genes in breeding in order to enhance the flood tolerance of our crops.

Carbon partitioning is important for understanding root development but little is known about its regulation. Existing models suggest that partitioning is controlled by the potential sink strength. They cannot, however, simulate hierarchical uptake other than by using absolute priorities. Moreover, they cannot explain that the changes in photoassimilate partitioning result from changes in photosynthesis. In this paper we present a model of phloem sieve circulation, based on the model of Minchin et al. (Journal of Experimental Botany44: 947–955, 1993). The root system was represented by a network of segments to which meristems were connected. The properties of the segments were determined by the differentiation stage. Photoassimilate import from each organ was assumed to be limited by a metabolic process and driven by Michaelis–Menten kinetics. The axial growth was proportional to meristem respiration, which drives the flux of new cells required for root elongation. We used the model to look at trophic apical dominance, determinate and indeterminate root growth, the effect of the activity of a root on competition with its neighbours, and the effect of photoassimilate availability on changes in partitioning. The simulated phloem mass flow yielded results of the same order of magnitude as those generally reported in the literature. For the main well vascularized axis, the model predicted that one single apical meristem larger than its neighbouring laterals, was enough to generate a taprooted system. Conversely, when the meristem of laterals close to the collar had a volume similar to that of the taproot, the predicted network became fibrous. The model predicted a hierarchical priority for organ photoassimilate uptake, similar to that described in the literature, during the decline in photosynthetic activity. Our model suggests that determinate growth of the first laterals resulted from a local shortage of photoassimilate at their meristem, as a result of the limited transport properties of the developed roots.

Ammonium is a major inorganic nitrogen source for plants. At low external supplies, ammonium promotes plant growth, while at high external supplies it causes toxicity. Ammonium triggers rapid changes in cytosolic pH, in gene expression, and in post-translational modifications of proteins, leading to apoplastic acidification, co-ordinated ammonium uptake, enhanced ammonium assimilation, altered oxidative and phytohormonal status, and reshaped root system architecture. Some of these responses are dependent on AMT-type ammonium transporters and are not linked to a nutritional effect, indicating that ammonium is perceived as a signaling molecule by plant cells. This review summarizes current knowledge of ammonium-triggered physiological and morphological responses and highlights existing and putative mechanisms mediating ammonium signaling and sensing events in plants. We put forward the hypothesis that sensing of ammonium takes place at multiple steps along its transport, storage, and assimilation pathways.

The architecture of the branched root system of plants is a major determinant of vigor. Water availability is known to impact root physiology and growth; however, the spatial scale at which this stimulus influences root architecture is poorly understood. Here we reveal that differences in the availability of water across the circumferential axis of the root create spatial cues that determine the position of lateral root branches. We show that roots of several plant species can distinguish between a wet surface and air environments and that this also impacts the patterning of root hairs, anthocyanins, and aerenchyma in a phenomenon we describe as hydropatterning. This environmental response is distinct from a touch response and requires available water to induce lateral roots along a contacted surface. X-ray microscale computed tomography and 3D reconstruction of soil-grown root systems demonstrate that such responses also occur under physiologically relevant conditions. Using early-stage lateral root markers, we show that hydropatterning acts before the initiation stage and likely determines the circumferential position at which lateral root founder cells are specified. Hydropatterning is independent of endogenous abscisic acid signaling, distinguishing it from a classic water-stress response. Higher water availability induces the biosynthesis and transport of the lateral root-inductive signal auxin through local regulation of TRYPTOPHAN AMINOTRANSFERASE OF ARABIDOPSIS 1 and PIN-FORMED 3, both of which are necessary for normal hydropatterning. Our work suggests that water availability is sensed and interpreted at the suborgan level and locally patterns a wide variety of developmental processes in the root.

Plants are exposed to increasingly severe drought events and roots play vital roles in maintaining plant survival, growth, and reproduction. A large body of literature has investigated the adaptive responses of root traits in various plants to water stress and these studies have been reviewed in certain groups of plant species at a certain scale. Nevertheless, these responses have not been synthesized at multiple levels. This paper screened over 2000 literatures for studies of typical root traits including root growth angle, root depth, root length, root diameter, root dry weight, root-to-shoot ratio, root hair length and density and integrates their drought responses at genetic and morphological scales. The genes, quantitative trait loci (QTLs) and hormones that are involved in the regulation of drought response of the root traits were summarized. We then statistically analyzed the drought responses of root traits and discussed the underlying mechanisms. Moreover, we highlighted the drought response of 1-D and 2-D root length density (RLD) distribution in the soil profile. This paper will provide a framework for an integrated understanding of root adaptive responses to water deficit at multiple scales and such insights may provide a basis for selection and breeding of drought tolerant crop lines.

OsPIN2 is identified as the casual gene responsible for the phenotype of lta1, a rice mutant that displays large root angles and a shallow root system architecture, affecting the polar transport of auxin in the root tip. Abstract Root system architecture is very important for plant growth and crop yield. It is essential for nutrient and water uptake, anchoring, and mechanical support. Root growth angle (RGA) is a vital constituent of root system architecture and is used as a parameter for variety evaluation in plant breeding. However, little is known about the underlying molecular mechanisms that determine root growth angle in rice (Oryza sativa). In this study, a rice mutant large root angle1 (lra1) was isolated and shown to exhibit a large RGA and reduced sensitivity to gravity. Genome resequencing and complementation assays identified OsPIN2 as the gene responsible for the mutant phenotypes. OsPIN2 was mainly expressed in roots and the base of shoots, and showed polar localization in the plasma membrane of root epidermal and cortex cells. OsPIN2 was shown to play an important role in mediating root gravitropic responses in rice and was essential for plants to produce normal RGAs. Taken together, our findings suggest that OsPIN2 plays an important role in root gravitropic responses and determining the root system architecture in rice by affecting polar auxin transport in the root tip.

To face future challenges in crop production dictated by global climate changes, breeders and plant researchers collaborate to develop productive crops that are able to withstand a wide range of biotic and abiotic stresses. However, crop selection is often focused on shoot performance alone, as observation of root properties is more complex and asks for artificial and extensive phenotyping platforms. In addition, most root research focuses on development, while a direct link to the functionality of plasticity in root development for tolerance is often lacking. In this paper we review the currently known root system architecture (RSA) responses in Arabidopsis and a number of crop species to a range of abiotic stresses, including nutrient limitation, drought, salinity, flooding, and extreme temperatures. For each of these stresses, the key molecular and cellular mechanisms underlying the RSA response are highlighted. To explore the relevance for crop selection, we especially review and discuss studies linking root architectural responses to stress tolerance. This will provide a first step toward understanding the relevance of adaptive root development for a plant's response to its environment. We suggest that functional evidence on the role of root plasticity will support breeders in their efforts to include root properties in their current selection pipeline for abiotic stress tolerance, aimed to improve the robustness of crops.