BACKGROUND: Climate change induces perturbation in the global water cycle, profoundly impacting water availability for agriculture and therefore global food security. Water stress encompasses both drought (i.e. water scarcity) that causes the drying of soil and subsequent plant desiccation, and flooding, which results in excess soil water and hypoxia for plant roots. Terrestrial plants have evolved diverse mechanisms to cope with soil water stress, with the root system serving as the first line of defense. The responses of roots to water stress can involve both structural and physiological changes, and their plasticity is a vital feature of these adaptations. Genetic methodologies have been extensively employed to identify numerous genetic loci linked to water stress-responsive root traits. This knowledge is immensely important for developing crops with optimal root systems that enhance yield and guarantee food security under water stress conditions.
RESULTS: This review focused on the latest insights into modifications in the root system architecture and anatomical features of legume roots in response to drought and flooding stresses. Special attention was given to recent breakthroughs in understanding the genetic underpinnings of legume root development under water stress. The review also described various root phenotyping techniques and examples of their applications in different legume species. Finally, the prevailing challenges and prospective research avenues in this dynamic field as well as the potential for using root system architecture as a breeding target are discussed.
CONCLUSIONS: This review integrated the latest knowledge of the genetic components governing the adaptability of legume roots to water stress, providing a reference for using root traits as the new crop breeding targets.

Transformation of plants using wild strains of agrobacteria is termed natural transformation and is not covered by GMO legislation in e.g. European Union and Japan. In the current study, offspring lines (A11 and B3) of Rhizobium rhizogenes naturally transformed oilseed rape (Brassica napus) were randomly selected to characterize the morphological traits, and the implications of such morphological changes on plant drought resilience. It was found that the introduction of Ri-genes altered the biomass partitioning to above- and under-ground parts of oilseed rape plants. Compared to the wild type (WT), the A11 and B3 lines possessed 1.2-4.0 folds lower leaf and stem dry weight, leaf area and plant height, but had 1.3-5.8 folds greater root dry weight, root length and root surface area, resulting in a significantly enhanced root: shoot dry mass ratio and root surface area: leaf area ratio. In addition, the introduction of Ri-genes conferred reduced stomatal pore aperture and increased stomatal density in the B3 line, and increased leaf thickness in A11 line, which could benefit plant drought resilience. Finally, the modulations in morphological traits as a consequence of transformation with Ri-genes are discussed concerning resilience in water-limited conditions. These findings reveal the potential of natural transformation with R. rhizogenes for drought-targeted breeding in crops.

Some citrus orchards in China often experience nitrogen (N) deficiency. For the first time, targeted metabolomics was used to examine N-deficient effects on hormones in sweet orange (Citrus sinensis (L.) Osbeck cv. Xuegan) leaves and roots. The purpose was to validate the hypothesis that hormones play a role in N deficiency tolerance by regulating root/shoot dry weight ratio (R/S), root system architecture (RSA), and leaf and root senescence. N deficiency-induced decreases in gibberellins and indole-3-acetic acid (IAA) levels and increases in cis(+)-12-oxophytodienoic acid (OPDA) levels, ethylene production, and salicylic acid (SA) biosynthesis might contribute to reduced growth and accelerated senescence in leaves. The increased ethylene formation in N-deficient leaves might be caused by increased 1-aminocyclopropanecarboxylic acid and OPDA and decreased abscisic acid (ABA). N deficiency increased R/S, altered RSA, and delayed root senescence by lowering cytokinins, jasmonic acid, OPDA, and ABA levels and ethylene and SA biosynthesis, increasing 5-deoxystrigol levels, and maintaining IAA and gibberellin homeostasis. The unchanged IAA concentration in N-deficient roots involved increased leaf-to-root IAA transport. The different responses of leaf and root hormones to N deficiency might be involved in the regulation of R/S, RSA, and leaf and root senescence, thus improving N use efficiency, N remobilization efficiency, and the ability to acquire N, and hence conferring N deficiency tolerance.

Plant phenotypic plasticity plays an important role in nitrogen (N) acquisition and use under nitrogen-limited conditions. However, this role has never been quantified as a function of N availability, leaving it unclear whether plastic responses should be considered as potential targets for selection. A combined modelling and experimentation approach was adopted to quantify the role of plasticity on N uptake and plant yield. Based on a greenhouse experiment we considered plasticity in two maize traits: root-to-leaf biomass allocation ratio and emergence rate of axial roots. In a simulation experiment we individually enabled or disabled both plastic responses for maize stands grown across six N levels. Both plastic responses contributed to maintaining a higher N uptake and plant productivity as N-availability declined, compared to stands in which plastic responses were disabled. We conclude that plastic responses quantified in this study may be a potential target trait in breeding programs for greater N uptake across N levels while it may only be important for the internal use of N under N-limited conditions in maize. Given the complexity of breeding for plastic responses, an a priori model analysis is useful to identify which plastic traits to target for enhanced plant performance.

Strengthening rhizosphere effects to enhance pollutant removal is a hotspot of constructed wetlands (CWs) research in recent years, and improving the root traits and metabolic capacity of macrophytes is crucial for strengthening rhizosphere effects. In the field experiment, two types of subsurface flow (SSF) CWs (CW10 and CW20, with substrate depths of 10 and 20 cm, respectively) under the vertical spatial stress of roots (VSSR) and two types of non-VSSR SSF CWs (CW40 and CW60) were adopted with Typha orientalis as cultivated plants to investigate the variability of root development, metabolism, and pollutant removal at different substrate depths. VSSR induced substantial redundant root development, which significantly increased root-shoot ratio, fine and lateral root biomass, root porosity, and root activity, with lateral and fine root biomass of CW20 reaching 409.17 and 237.42 g/m2, respectively, which were 3.18 and 5.28 times those of CW60. The radical oxygen loss (ROL) and dissolved organic carbon (DOC) levels of CW20 single plant were 1.36 and 4.57 times higher than those of CW60, respectively, and more types of root exudates were determined (e.g., aldehydes, ketones and amides). More aerobic heterotrophs (e.g., Massilia, Planomicrobium), nitrification bacteria (e.g., Ellin6067, Nitrospira), aerobic denitrification bacteria (e.g., Bacillu, Chryseobacterium, Pseudomonas) and denitrification phosphorus accumulating organisms (e.g., Flavobacterium) were enriched in the rhizosphere of CW20. This changed the main transformation pathways of pollutants and enhanced the removal of pollutants, with the COD, TN and TP average removal rates of CW20 increasing by 9.99%, 13.28% and 8.92%, respectively, compared with CW60. The ideotype root system architecture CW (RSACW; CW20) constructed in this study, which consists of a large number of fine and lateral roots, can stimulate more efficient rhizosphere effects stably and continuously.

Transcription factors in coordination with phytohormones form an intricate regulatory network modulating vital cellular mechanisms like development, growth and senescence in plants. In this study, we have functionally characterized the transcription factor OsNAC121 by developing gene silencing and overexpressing transgenic rice plants, followed by detailed analyses of the plant architecture. Transgenic lines exhibited remodelling in crown root development, lateral root structure and density, tiller height and number, panicle and grain morphologies, underpinning the imbalanced auxin: cytokinin ratio due to perturbed auxin transportation. Application of cytokinin, auxin and abscisic acid increased OsNAC121 gene expression nearly 17-, 6- and 91-folds, respectively. qRT-PCR results showed differential expressions of auxin and cytokinin pathway genes, implying their altered levels. A 47-fold higher expression level of OsNAC121 during milky stage in untransformed rice, compared to 14-day old shoot tissue, suggests its crucial role in grain filling; as evidenced by a large number of undeveloped grains produced by the gene silenced lines. Crippled gravitropic response by the transgenic plants indicates their impaired auxin transport. Bioinformatics revealed that OsNAC121 interacts with co-repressor (TOPLESS) proteins and forms a part of the inhibitor complex OsIAA10, an essential core component of auxin signalling pathway. Therefore, OsNAC121 emerges as an important regulator of various aspects of plant architecture through modulation of crosstalk between auxin and cytokinin, altering their concentration gradient in the meristematic zones, and consequently modifying different plant organogenesis processes.

CONCLUSIONS: This review article highlights a broader perspective of NPs and plant-root interaction by focusing on their beneficial and deleterious impacts on root system architecture (RSA). The root performs a vital function by securing itself in the soil, absorbing and transporting water and nutrients to facilitate plant growth and productivity. In dicots, the architecture of the root system (RSA) is markedly shaped by the development of the primary root and its branches, showcasing considerable adaptability in response to changes in the environment. For promoting agriculture and combating global food hunger, the use of nanoparticles (NPs) may be an exciting option, for which it is essential to understand the behaviour of plants under NPs exposure. The nature of NPs and their physicochemical characteristics play a significant role in the positive/negative response of roots and shoots. Root morphological features, such as root length, root mass and root development features, may regulated positively/negatively by different types of NPs. In addition, application of NPs may also enhance nutrient transport and soil fertility by the promotion of soil microorganisms including plant growth-promoting rhizobacteria (PGPRs) and also soil enzymes. Interestingly the interaction of nanomaterials (NMs) with rhizospheric bacteria can enhance plant development and soil health. However, some studies also suggested that the increased use of several types of engineered nanoparticles (ENPs) may disrupt the equilibrium of the soil-root interface and unsafe morphogenesis by causing the browning of roots and suppressing the growth of root and soil microbes. Thus, this review article has sought to compile a broader perspective of NPs and plant-root interaction by focusing on their beneficial or deleterious impacts on RSA.

OBJECTIVE: Root system architecture (RSA) plays a key role in plant adaptation to drought because deep rooting enables better water uptake than shallow rooting under terminal drought. Understanding RSA during early plant development is essential for improving crop yields, as early drought can affect subsequent shoot growth. Herein, we demonstrate that root distribution in the topsoil significantly impacts shoot growth during the early stages of rice (Oryza sativa) development under drought, as assessed through three-dimensional (3D) image analysis.
METHODS: We used 109 F12 recombinant inbred lines (RILs) obtained from a cross between shallow-rooting lowland rice and deep-rooting upland rice, representing a population with diverse RSA. We applied a moderate drought during the early development of rice grown in a plant pot (25 cm height) by stopping irrigation 14 days after sowing (DAS). Time-series RSA at 14, 21, and 28 DAS was visualized by X-ray computed tomography, and subsequently compared between drought and well-watered conditions. Following this analysis, we further investigated drought-avoidant RSA by testing 20 randomly selected RILs under drought conditions.
RESULTS: We inferred the root location that most influences shoot growth using a hierarchical Bayes approach: the root segment depth, which positively impacted shoot growth, ranged between 1.7-3.4 cm under drought conditions and between 0.0-1.7 cm under well-watered conditions. Drought-avoidant RILs had a higher root density in the lower layers of the topsoil compared to the others.
CONCLUSIONS: Fine classification of soil layers using 3D image analysis revealed that increasing root density in the lower layers of the topsoil, rather than in the subsoil, is advantageous for drought avoidance during the early growth stage of rice.

UNASSIGNED: Root system architecture (RSA) plays an important role in the plant\'s ability to sustain yield under abiotic stresses such as drought. Preceding crops (precrops) can affect the yield of the proceeding crop, partially by affecting the RSA. This experiment aims to explore the interactions between precrop identity, crop genotype and drought at early growth stages.
UNASSIGNED: Rhizotrons, sized 60 × 80 × 3.5 cm, were used to assess the early root growth of two winter wheat (Triticum aestivum L.) genotypes, using precrop-treated soil around the seedlings and differing water regimes. The rhizotrons were automatically imaged 3 times a week to track root development.
UNASSIGNED: Precrop-treated soil affected the RSA and changes caused by the reduced water treatment (RWT) were different depending on the precrop. Largest of these was the 36% reduction in root depth after wheat, but 44% after OSR. This indicates that effects caused by the precrop can be simulated, at least partially, by transferring precrop-treated soils to controlled environments. The genotypes had differential RSA and reacted differently to the RWT, with Julius maintaining an 8.8-13.1% deeper root system compared to Brons in the RWT. In addition, the combined environmental treatment affected the genotypes differently.
UNASSIGNED: Our results could help explain discrepancies found from using precrops to enhance yield as they indicate differences in the preceding crop effect when experiencing drought stress. Further, these differences are affected by genotypic interactions, which can be used to select and adapt crop genotypes for specific crop rotations, depending on the year. Additionally, we have shown a viable method of stimulating a partial precrop effect at the seedling stage in a controlled greenhouse setting using field soil around the germinated seed.

Plant growth and survival in their natural environment require versatile mitigation of diverse threats. The task is especially challenging due to the largely unpredictable interaction of countless abiotic and biotic factors. To resist an unfavorable environment, plants have evolved diverse sensing, signaling, and adaptive molecular mechanisms. Recent stress studies have identified molecular elements like secondary messengers (ROS, Ca2+, etc.), hormones (ABA, JA, etc.), and signaling proteins (SnRK, MAPK, etc.). However, major gaps remain in understanding the interaction between these pathways, and in particular under conditions of stress combinations. Here, we highlight the challenge of defining \"stress\" in such complex natural scenarios. Therefore, defining stress hallmarks for different combinations is crucial. We discuss three examples of robust and dynamic plant acclimation systems, outlining specific plant responses to complex stress overlaps. (a) The high plasticity of root system architecture is a decisive feature in sustainable crop development in times of global climate change. (b) Similarly, broad sensory abilities and apparent control of cellular metabolism under adverse conditions through retrograde signaling make chloroplasts an ideal hub. Functional specificity of the chloroplast-associated molecular patterns (ChAMPs) under combined stresses needs further focus. (c) The molecular integration of several hormonal signaling pathways, which bring together all cellular information to initiate the adaptive changes, needs resolving.