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Imazethapyr, a widely used herbicide, exhibits a long persistence in soils and can cause injury to rotational crops. Here, we discovered that imazethapyr inhibits primary root elongation in Arabidopsis by inhibiting cell division and expansion rather than damaging the organization of root meristem. Integration of transcriptomic and metabolomic analysis revealed that imazethapyr downregulated multiple genes related to cell wall loosening and modification, leading to increased cell wall thickness and inhibited cellular expansion in Arabidopsis roots. Furthermore, imazethapyr upregulated auxin biosynthesis and transport, resulting in enhanced auxin accumulation at root tips. Elevated auxin concentrations triggered apoplast alkalization and the inactivation of wall-loosening enzymes, further suppressing root growth. This research provides new insights into the molecular mechanism underlying imazethapyr phytotoxicity and offers potential strategies for developing crops that are better adapted to soils contaminated with imidazolinone herbicides.

Plant growth depends on growth regulators, nutrient availability, and amino acid levels, all of which influence cell wall formation and cell expansion. Cell wall integrity and structures are surveyed and modified by a complex array of cell wall integrity sensors, including leucine-rich repeat (LRR)-extensins (LRXs) that bind RALF (rapid alkalinization factor) peptides with high affinity and help to compact cell walls. Expressing the Arabidopsis root hair-specific LRX1 without the extensin domain, which anchors the protein to the cell wall (LRX1ΔE14), has a negative effect on root hair development. The mechanism of this negative effect was investigated by a suppressor screen, which led to the identification of a sune (suppressor of dominant-negative LRX1ΔE14) mutant collection. The sune82 mutant was identified as an allele of HISN2, which encodes an enzyme essential for histidine biosynthesis. This mutation leads to reduced accumulation of histidine and an increase in several amino acids, which appears to have an effect on the TOR (target of rapamycin) network, a major controller of eukaryotic cell growth. It also represents an excellent tool to study the effects of reduced histidine levels on plant development, as it is a rare example of a viable partial loss-of-function allele in an essential biosynthetic pathway., (© The Author(s) 2024. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

Lung seven transmembrane receptor family is a small part of Arabidopsis gene family. So far, the function of some members of the this family is unknown. Plant elicitor peptide1 (Pep1) is one of damage-associated molecular patterns (DAMPs), which could trigger root growth inhibition and plant immunity responses. Here, we identified members of the Lung seven transmembrane family, and proved they are important for Pep1-induced root growth and development. We found that the expression levels of 7TM2 and 7TM6 were elevated in wild-type treated with Pep1. Phenotypic analysis showed that the growth phenotypes of 7tm2 and 7tm6 were similar to the wild-type under Pep1 treatment, but the 7tm2 7tm6 had a Pep1 hypersensitivity phenotype compared to wild-type. Furthermore, the complementation lines were able to restore the Pep1 phenotype of 7tm2 7tm6 to that similar to wild-type. These results suggest that 7TM2 and 7TM6 are involved in the regulation of root growth by Pep1. This study revealed the new functions of the Lung seven transmembrane receptor family and provided new ideas for further revealing the molecular mechanism of Pep1-regulated root growth and development., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2025 Elsevier Inc. All rights reserved.)

Folates are vital one carbon donors and acceptors for a whole range of key biochemical reactions, including the biosynthesis of DNA building blocks. Plants use one carbon metabolism as a jack of all trades in their growth and development. Depletion of folates impedes root growth in Arabidopsis thaliana, but the mechanistic basis behind this function is still obscure. A global transcriptomic study hinted that folate depletion may cause misregulation of cell cycle progression. However, investigations on a direct connection thereof are scarce. We confirmed the effect of methotrexate (MTX), a folate biosynthesis inhibitor, on the expression of cell cycle genes. Subsequently, we determined the effect of MTX on root morphology and cell cycle progression through phase-specific cell cycle reporter analyses. Our study reveals that folate depletion affects the expression of cell cycle regulatory genes in roots, thereby suppressing cell cycle progression. We confirmed, through DNA labelling by EdU, that MTX treatment leads to arrest in the S phase of meristematic cells, likely due to the lack of DNA precursors. Further, we noted an accumulation of the A-type CYCA3;1 cyclin at the root tip, suggesting a possible link with the observed loss of apical dominance. Overall, our study shows that the restricted cell division and cell cycle progression is one of the reasons behind the loss of primary root growth upon folate depletion., Competing Interests: Declarations. Conflict of interest: The authors have no conflicts of interest to declare., (© 2025. The Author(s).)

The use of herbicides is increasing because they are essential to ensure food safety, especially in the increasing need for food production. However, their misapplication leads to significant environmental problems. This study aimed to determine the cytotoxic effect of picloram on the bioindicator L. culinaris. Nine doses of picloram (0.1, 0.25, 0.5, 0.75, 1, 1.5, 1.5, 2.0, 2.5, and 5 mg/L) were evaluated for their impact on root growth, mitotic index, and frequency of chromosomal abnormalities. Treatment with 5 mg/L picloram significantly inhibited root growth (0.24 ± 0.1 cm, 0.4 ± 0.1 cm, and 0.5 ± 0.15 cm at 24, 48, and 72 h, respectively) and reduced the mitotic index (5.2 ± 10.8), evidencing an interference in cell division. The frequency of chromosomal abnormalities increased with higher concentrations of picloram, suggesting a genotoxic effect even at low concentrations. These findings highlight the need to thoroughly evaluate the effects of herbicides on various aspects of plant development and cellular integrity to gain a more complete understanding of their toxicity and the environmental risks associated with them., Competing Interests: Declarations. Ethical approval: In this study, the Ethical Approval section is marked as “not applicable” since the research does not entail the involvement of human participants, their data, or biological samples. Consent to participate: Individual participants or their guardians provided written informed consent. Consent to publish: The consent to publish section is marked as “not applicable” for this study since the manuscript does not incorporate any data derived from individuals. Competing interests: The authors declare no competing interests., (© 2025. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

This work presents an imaging testing system (software and hardware) that can generate repeatable images through a stabilized port in the soil for processes known to change with time. The system includes (i) a stabilized port in the ground made of standard PVC pipe, with sections lined with a borosilicate glass tube, and (ii) a digital imaging instrument to survey the optically transparent portion of the stabilized port. The instrument uses a probe containing a digital camera and two light sources, one using white lights and one using ultraviolet (UV) lights (365 nm). The main instrument controls the probe using a cable within the stabilized port to take overlapping pictures of the soil under the different light sources. Two examples are provided, one to document the distribution of soil and groundwater contaminants known as non-aqueous phase liquids (NAPL, which include petroleum) at variable water saturation levels and a second one to monitor the growth of a plant over a 2-week interval. In both examples, the system successfully identified critical changes in soil processes and showed a resolution of approximately 15 µm (in the order of the thickness of a human hair), demonstrating the potential for repeated imaging of soil processes known to experience temporal changes. Both examples are illustrative, as additional applications might be possible. The novelty of this system lies in its ability to generate repeated measurements at larger depths than the current shallow systems installed by hand.

Besides playing a crucial role in plant immunity via the nonexpressor of pathogenesis-related (NPR) proteins, increasing evidence shows that salicylic acid (SA) can also regulate plant root growth. However, the transcriptional regulatory network controlling this SA response in plant roots is still unclear. Here, we found that NPR1 and WRKY45, the central regulators of SA response in rice leaves, control only a reduced sector of the root SA signaling network. We demonstrated that SA attenuates root growth via a novel NPR1/WRKY45-independent pathway. Furthermore, using regulatory network analysis and mutant characterization, we identified a set of new NPR1/WRKY45-independent regulators that conservedly modulate the root development and root-associated microbiota composition in both Oryza sativa (monocot) and Arabidopsis thaliana (dicot) in response to SA. Our results established the SA signaling as a central element regulating plant root functions under ecologically relevant conditions. These results provide new insights to understand how regulatory networks control plant responses to abiotic and biotic stresses., (© 2024 Institute of Botany, Chinese Academy of Sciences.)

This study evaluates the impact of yellow mealworm frass in combination with an inorganic nitrogen fertilizer on growth, yield, and nutrient concentration in annual sowthistle plants. It was found that the combined application of yellow mealworm frass (YM-frass) as basal fertilizer and inorganic nitrogen fertilizer (INF) as top dressing increased the shoot biomass of annual sowthistle up to 67.2 % compared to the INF treatment (100 kg N/ha). The application of both YM-frass and INF increased P, Mg and Fe root concentration. Moreover, the lowest P and K concentration in shoot tissues were found in the unfertilized control treatment, while the reverse was the case for Ca and Fe. In conclusion, yellow mealworm frass can be used instead of inorganic basal fertilizer, whereas its combination with an inorganic nitrogen fertilizer applied as top-dressing during the vegetative growth can increase the yield and quality of leafy vegetables., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024. Published by Elsevier Ltd.)

Plants emit green leaf volatiles (GLVs) in response to biotic and abiotic stress. Receiver plants perceive GLVs as alarm cues resulting in activation of defensive or protective mechanisms. While this is well documented, it is not known how GLVs are perceived by receiver cells and what the structural determinants are for GLV activity. We tested whether the carbon chain length in (Z)-3-fatty alcohols with four to nine carbons and the double bonds in six-carbon alcohols contribute to bioactivity. In Solanum peruvianum suspension-cultured cells we found that (Z)-3-fatty alcohols, except (Z)-3-butenol, induce medium alkalinization and MAP kinase phosphorylation, two signaling responses often tied to the perception of molecular patterns that function in plant immunity and resistance to herbivores. In tomato (S. lycopersicum) seedlings, we found that (Z)-3-fatty alcohols induce inhibition of root growth. In both signaling and physiological responses, (Z)-3-octenol and (Z)-3-nonenol had a higher bioactivity than (Z)-3-heptenol and (Z)-3-hexenol, with (Z)-3-butenol only being active in root growth assays. Bioactivity correlated not only with chain length but also with lipophilicity of the fatty alcohols. The natural GLVs (E)-2-hexenol and the saturated 1-hexanol exhibited a higher bioactivity in pH assays than (Z)-3-hexenol, indicating that the presence and position of a double bond also contributes to bioactivity. Our results indicate that perceiving mechanisms for (Z)-3-fatty alcohols show a preference for longer chain fatty alcohols or that longer chain fatty alcohols are more accessible to receptors., Competing Interests: Declarations. Competing Interests: The authors declare no competing interests., (© 2025. The Author(s).)

Plants frequently encounter relatively low and fluctuating potassium (K + ) concentrations in soil, with roots serving as primary responders to this stress. Histone modifications, such as de-/acetylation, can function as epigenetic markers of stress-inducible genes. However, the signaling network between histone modifications and low-K + (LK) response pathways remains unclear. This study investigated the regulatory role of Histone Deacetylase Complex 1 (HDC1) in primary root growth of Arabidopsis thaliana under K + deficiency stress. Using a hdc1-2 mutant line, we observed that HDC1 positively regulated root growth under LK conditions. Compared to wild-type (WT) plants, the hdc1-2 mutant exhibited significantly inhibited primary root growth under LK conditions, whereas HDC1-overexpression lines displayed opposite phenotypes. No significant differences were observed under HK conditions. Further analysis revealed that the inhibition of hdc1-2 on root growth was due to reduced apical meristem cell proliferation rather than cell elongation. Notably, the root growth of hdc1-2 showed reduced sensitivity compared to WT after auxin treatment under LK conditions. HDC1 may regulate root growth by affecting auxin polar transport and subsequent auxin signaling, as evidenced by the altered expression of auxin transport genes. Moreover, the organ-specific RT-qPCR analyses unraveled that HDC1 negatively regulates the expression of CBL-CIPK-K + channel-related genes such as CBL1, CBL2 , CBL3 , AKT1, and TPK1 , thereby establishing a molecular link between histone deacetylation, auxin signaling, and CBLs-CIPKs pathway in response to K + deficiency.

Maintaining a stable basal level of salicylic acid (SA) is crucial for plant growth, development, and stress response, although basal levels of SA vary significantly among plant species. However, the molecular mechanisms by which basal SA regulates plant growth and stress response remain to be clarified. In this study, we performed a genetic screen to identify suppressors of the root growth defect in Osaim1, a rice mutant deficient in basal SA biosynthesis. We found that mutation of the E3 ligase OsPRT6, a key component of the Arg/N-degron pathway, can rescue the root growth defect of Osaim1. Further analysis revealed that OsWRKY62 and OsWRKY76 act as substrates of the OsPRT6 N-degron pathway to modulate root growth. We demonstrated that reducing the basal SA level activates the PRT6 N-degron pathway and that basal SA modulates the stress response in part through the PRT6 N-degron pathway. Importantly, the effects of basal SA levels on the PRT6 N-degron pathway are conserved across plant species. Taken together, these findings reveal a novel regulatory mechanism by which basal SA represses the PRT6 N-degron pathway to modulate root growth and abiotic stress response in rice., (Copyright © 2025 The Authors. Published by Elsevier Inc. All rights reserved.)

Roots absorb water and nutrients from the soil, support the plant's aboveground organs, and detect environmental changes, making them crucial targets for improving crop productivity. Particularly sensitive to soil salinity, a major abiotic stress, roots face significant challenges that threaten global agriculture. In response to salt stress, plants suppress root meristem size, thereby reducing root growth. However, the mechanisms underlying this growth restriction remain unclear. Here, we investigate the role of reactive oxygen species (ROS) in this process and reveal that LATERAL ORGAN BOUNDARIES DOMAIN 11 (LBD11) plays a central role in ROS-mediated regulation of meristem size and the salt stress-induced inhibition of root growth. Under normal conditions, LBD11 controls the expression of key ROS metabolic genes, maintaining ROS homeostasis within root developmental zones to control meristem size and overall root growth. Upon sensing salt stress, LBD11 undergoes rapid proteasome-mediated degradation, leading to decreased distribution of O 2 , which in turn curtails meristem size and limits root length. Our findings highlight an unexplored plant adaptation strategy, where the growth-promoting LBD11/ROS pathway is downregulated to finely regulate root growth under challenging conditions. We propose a strategy for developing crops with heightened resilience and increased yields in salt-affected environments.⋅ - , which in turn curtails meristem size and limits root length. Our findings highlight an unexplored plant adaptation strategy, where the growth-promoting LBD11/ROS pathway is downregulated to finely regulate root growth under challenging conditions. We propose a strategy for developing crops with heightened resilience and increased yields in salt-affected environments., (Copyright © 2025 The Authors. Published by Elsevier Inc. All rights reserved.)

Rhizospheric pH, an important environmental cue, severely impacts plant growth and fitness, therefore, has emerged as a major determinant of crop productivity. Despite numerous attempts, the key questions related to plants response against rhizospheric pH remains largely elusive. The present study provides a mechanistic framework for rhizospheric pH-mediated root growth inhibition (RGI). Utilizing various genetic resources combined with pharmacological agents and high-resolution confocal microscopy, the study provides direct evidence for the involvement of jasmonates and auxin in rhizospheric pH-mediated RGI. We show that auxin maxima at root tip is tightly regulated by the rhizospheric pH. In contrast, jasmonates (JAs) abundance inversely correlates with rhizospheric pH. Furthermore, JA-mediated regulation of auxin maxima through GRETCHEN HAGEN 3 (GH3) family genes explains the pattern of RGI observed over the range of rhizospheric pH. Our findings revealed auxin as the key regulator of RGI during severe pH conditions, while JAs antagonistically regulate auxin response against rhizospheric pH., (© 2024 John Wiley & Sons Ltd.)

Calcium signaling plays an essential role in integrating plant responses to diverse stimuli and regulating growth and development. While some signaling components and their roles are well-established, such as the ubiquitous calmodulin (CaM) sensor, plants possess a broader repertoire of calcium sensors. Notably, CaM-like proteins (CMLs) represent a poorly characterized class for which interacting partners and biological functions remain largely elusive. Our work investigates the role of Arabidopsis thaliana CML8 that exhibits a unique expression profile in seedlings. A reverse genetic approach revealed a function of CML8 in regulating root growth and hypocotyl elongation. RNA-seq analyses highlighted CML8 association with the regulation of numerous genes involved in growth and brassinosteroid (BR) signaling. Using co-immunoprecipitation experiments, we demonstrated that CML8 interacts with the BR receptor, BRI1, in planta in a ligand-dependent manner. This finding suggests the existence of a novel regulatory step in the BR pathway, involving calcium signaling., (© 2024 The Author(s). The Plant Journal published by Society for Experimental Biology and John Wiley & Sons Ltd.)

Posttranslational tyrosine sulfation of peptides and proteins is catalysed by tyrosylprotein sulfotransferases (TPSTs). In Arabidopsis, tyrosine sulfation is essential for the activities of peptide hormones, such as phytosulfokine (PSK) and root meristem growth factor (RGF). Here, we identified a TPST-encoding gene, MtTPST, from model legume Medicago truncatula. MtTPST expression was detected in all organs, with the highest level in root nodules. A promoter:GUS assay revealed that MtTPST was highly expressed in the root apical meristem, nodule primordium and nodule apical meristem. The loss-of-function mutant mttpst exhibited a stunted phenotype with short roots and reduced nodule number and size. Application of both of the sulfated peptides PSK and RGF3 partially restored the defective root length of mttpst. The reduction in symbiotic nodulation in mttpst was partially recovered by treatment with sulfated PSK peptide. MtTPST-PSK module functions downstream of the Nod factor signalling to promote nodule initiation via regulating accumulation and/or signalling of cytokinin and auxin. Additionally, the small-nodule phenotype of mttpst, which resulted from decreased apical meristematic activity, was partially complemented by sulfated RGF3 treatment. Together, these results demonstrate that MtTPST, through its substrates PSK, RGF3 and other sulfated peptide(s), positively regulates nodule development and root growth., (© 2024 John Wiley & Sons Ltd.)

Extracellular ATP (eATP) orchestrates vital processes in plants, akin to its role in animals. P2K1 is a crucial receptor mediating eATP effects. Immunoprecipitation tandem mass spectrometry data highlighted FERONIA's significant interaction with P2K1, driving us to explore its role in eATP signaling. Here, we investigated putative P2K1-interactor, FERONIA, which is a versatile receptor kinase pivotal in growth and stress responses. We employed a FERONIA loss-of-function mutant, fer-4 , to dissect its effects on eATP signaling. Interestingly, fer-4 showed distinct calcium responses compared to wild type, while eATP-responsive genes were constitutively upregulated in fer-4 . Additionally, fer-4 displayed insensitivity to eATP-regulated root growth and reduced cell wall accumulation. Together, these results uncover a role for FERONIA in regulating eATP signaling. Overall, our study deepens our understanding of eATP signaling, revealing the intricate interplay between P2K1 and FERONIA impacting the interface between growth and defense.

Biochar application emerges as a valuable soil management strategy for enhancing crop yield; however, the mechanisms underlying the relationships between soil and plants remain unclear after biochar application. In this study, soil pore characteristics and maize yield were assessed in a five-year biochar-application experiment on the Loess Plateau of China, including four treatments: Control (no biochar), low-dose biochar application (LB, 3 t ha -1 ), moderate-dose biochar application (MB, 6 t ha -1 ), and high-dose biochar application (HB, 9 t ha -1 ). Root growth traits were evaluated by cultivating maize in intact soil cores collected from field conditions using X-ray computed tomography. Our findings indicate that, compared to the Control, the HB treatment enhanced macroporosity (> 0.1 mm in diameter), porosity of 0.1-0.5 mm pores, and saturated water content, while reducing macropore connectivity and penetration resistance. However, biochar application treatments did not alter the water retention characteristics from field capacity to permanent wilting point or the plant-available water content (PAWC). Furthermore, the mean angle of primary and seminal roots as well as the length and surface area of entire roots increased in the HB treatment, showing a positive correlation with the porosity of 0.1-0.5 mm pores. The mean diameter of primary and seminal roots, leaf fresh and dry weights, and maize yield also increased in the HB treatment compared to the Control. Partial least squares path modeling analysis indicated that biochar application rates positively impacted on root growth and plant productivity through an indirect influence of soil pore size distribution, with 0.1-0.5 mm pores being particularly crucial for facilitating deeper root penetration and root elongation. These findings demonstrate that biochar application primarily augmented 0.1-0.5 mm pores, rather than affecting smaller pores capable of retaining plant-available water or larger macropores, enhancing deeper rooting and root elongation, thus improving plant productivity and crop yield., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier B.V. All rights reserved.)

Biostimulation and precision irrigation are strategies that increase the sustainability of agriculture, and both have been widely studied in table grapes, but their interaction is a new approach for viticulture. The objective of this field trial was to assess the physiological effects of water deficit on table grapes pretreated for two consecutive years with five different biostimulation programs. Therefore, during the first year, vines were preconditioned with biostimulants composed of microorganisms, seaweed, and plant extracts and compared to an untreated control. During the second year, the same biostimulation treatments were evaluated under two different irrigation schedules: (i) farmer irrigation (FI), according to a farmer's criteria; and (ii) a deficit irrigation program, precision irrigation (PI), in which irrigation water was reduced from the post-veraison period to harvest, setting a threshold for allowable soil water depletion of 10% with respect to field capacity in order to minimize water leaching. The water inputs in the treatments under PI were reduced by 30% with respect to the FI treatment. While the deficit irrigation treatment clearly affected the plant water status indicators, biostimulation enhanced the root colonization by mycorrhizae and showed a trend of increased new root density. The combined effect of biostimulation and PI was shown to be an efficient strategy for optimizing the available resources, promoting the yield precocity.

Plant roots are considered highly efficient soil explorers. As opposed to the push-driven penetration strategy commonly used by many digging organisms, roots penetrate by growing, adding new cells at the tip, and elongating over a well-defined growth zone. However, a comprehensive understanding of the mechanical aspects associated with root penetration is currently lacking. We perform penetration experiments following Arabidopsis thaliana roots growing into an agar gel environment, and a needle of similar dimensions pushed into the same agar. We measure and compare the environmental deformations in both cases by following the displacement of fluorescent beads embedded within the gel, combining confocal microscopy and Digital Volume Correlation (DVC) analysis. We find that deformations are generally smaller for growing roots. To better understand the mechanical differences between the two penetration strategies, we develop a computational model informed by experiments. Simulations show that, compared to push-driven penetration, grow-driven penetration reduces frictional forces and mechanical work, with lower propagation of displacements in the surrounding medium. These findings shed light on the complex interaction of plant roots with their environment, providing a quantitative understanding based on a comparative approach., (© 2024 The Author(s). Plant, Cell & Environment published by John Wiley & Sons Ltd.)

Root-type ferredoxin:NADP(H) oxidoreductase (RFNR) is believed to reduce ferredoxin using NADPH in nonphotosynthetic tissues, facilitating ferredoxin-dependent biological processes. However, the physiological functions of RFNR remain unclear due to the difficulty in obtaining mutants lacking redundant RFNR isoproteins. The present study successfully generated Arabidopsis homozygous rnfr1;2 double mutants by traditional crossing and selection. However, they displayed severely stunted roots, challenging subsequent growth and abundant seed recovery. Notably, grafted plants combining mutant scions with wild-type rootstocks exhibited normal growth and produced abundant mutant seeds. Growth analysis employing reciprocal grafts with the wild-type and mutant plants showed that primary root growth was inhibited only when the rootstock was derived from the mutants. Meanwhile, the absence of RFNR1 and 2 in the scion had no apparent impact on shoot and root growth. Root transcriptome analysis revealed that RFNR1 and 2 deficiency upregulated genes encoding ferredoxin-dependent enzymes and root-type ferredoxin, leading to genome-wide reprogramming associated with cell walls, lipids, photosynthesis, secondary metabolism, and biotic/abiotic stress responses. Thus, Arabidopsis RFNR1 and 2 are crucial for root growth and various ferredoxin-dependent biological processes., Competing Interests: Declaration of competing interest The authors declare no conflicts of interest., (Copyright © 2025 Elsevier Inc. All rights reserved.)

The plant UDP-glycosyltransferases (UGTs) regulate several metabolic processes during root growth and development by conjugating sugar moieties to various small molecules. RsUGT71B5 is a novel UDP-glycosyltransferase in Raphanus sativus L., but its biological function is not well established. In this study, we generated RsUGT71B5-overexpressing transgenic Arabidopsis lines to determine the mechanisms by which RsUGT71B5 regulated root growth and development. Ectopic overexpression of RsUGT71B5 significantly enhanced root growth and seedling development. In culture medium supplemented with 1-3% exogenous sucrose, RsUGT71B5 overexpression increased the root length and surface area in the transgenic Arabidopsis lines compared with the wild type. Furthermore, transgenic RsUGT71B5 overexpression partially suppressed the inhibitory effects of 12% sucrose on root growth and development. RNA sequencing data analysis identified 102 differential expressed genes (DEGs), including 56 upregulated and 46 downregulated genes, in the transgenic RsUGT71B5 overexpression lines (OE). QRT-PCR analyses confirmed significant upregulation of glutathione S-transferases such as AT1G02930 (GSTF6) and AT1G02920 (GSTF7) in the transgenic RsUGT71B5 overexpression lines. KEGG pathway analyses of the DEGs showed that RsUGT71B5 overexpression regulated glutathione and sugar metabolism. In summary, this study demonstrated that RsUGT71B5 regulated root growth and development by modulating glutathione and sugar metabolism., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2025 Elsevier Masson SAS. All rights reserved.)

Brassinosteroids (BRs) are essential phytohormones that play a crucial role in plant growth and development. However, our understanding of BR receptors and their functions in tree species is currently limited. In this study, we looked for potential BR receptor genes in the burflower-tree (Neolamarckia cadamba) genome. We identified five candidate gene from sequence analysis and phylogenetic reconstruction. Among these genes, Neolamarckia cadamba BRASSINOSTEROID-INSENSITIVE 1 (NcBRI1) is ubiquitously expressed in all tested tissues and encodes a functional BR receptor localized to the plasma membrane. Ectopic expression of NcBRI1 in the Arabidopsis (Arabidopsis thaliana) loss-of-function BRI1 mutant bri1-5 not only rescued its growth retardation phenotype but also facilitated vascular development by reactivating BR signal transduction. Furthermore, overexpression of NcBRI1 promoted vascular formation and cell elongation in transgenic hairy roots of Neolamarckia cadamba. By contrast, microRNA-mediated knockdown of NcBRI1 resulted in delayed vascular development and smaller cells. Importantly, we found that manipulation of NcBRI1 in Neolamarckia cadamba can enhance the biomass of hairy roots. These findings highlight the critical role of NcBRI1 in BR signaling and its significant influence on vascular development and rapid growth in Neolamarckia cadamba., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier B.V. All rights reserved.)

The plant peptide-containing sulfated tyrosine (PSY) family plays critical roles in plant cell proliferation and stress responses. However, the functional characterization of the PSY peptide family in wheat remains unclear. This study systematically identified a total of 29 TaPSY genes at the genome-wide level, classifying them into six subgroups based on PSY-like motifs. These peptides contain a highly conserved active peptide domain, closely resembling the Arabidopsis AtPSY1 motif. All TaPSY homologs are predicted to have a sulfated tyrosine catalyzed by plant tyrosylprotein sulfotransferase (TPST). The TaPSY genes displayed distinct expression patterns across various tissues, with most genes showing higher expression levels in roots and stems. Synthetic sulfated TaPSY peptides enhanced root growth in both wild-type Arabidopsis and the tpst-1 mutant plants. In wheat, exogenous application of TaPSY peptides also promoted root growth, with the synthetic TaPSY5 peptide affecting reactive oxygen species levels in wheat taproots to stimulate primary root growth. Furthermore, transgenic Arabidopsis plants overexpressing TaPSY10 exhibited longer primary roots and increased lateral root numbers. These findings provide insights into the physiological roles of TaPSY peptides in regulating wheat root growth., Competing Interests: The authors declare no conflicts of interest.

Heat stress constitutes a serious threat to sesame ( Sesamum indicum L.). Root development during seed germination plays an essential role in plant growth and development. Nevertheless, the regulatory mechanisms underlying heat stress remain poorly understood. In this study, two sesame varieties differing in leaf heat tolerance (Zheng Taizhi 3 (heat-tolerant) and SP19 (heat-sensitive)) have been employed to investigate the impact of heat stress on root growth during germination. The results showed that heat stress significantly reduced the radicle length by 35.71% and 67.02% in Zheng Taizhi 3 and SP19, respectively, while germination rates remained unchanged. In addition, heat stress induced oxidative stress, as evidenced by increased reactive oxygen species (ROS) production, malondialdehyde (MDA) content, and reduced indole-3-acetic acid (IAA) content, accompanied by enhanced antioxidant enzyme activities, including those of superoxide dismutase (SOD), peroxidase (POD), and catalase (CAT), and the abscisic acid (ABA) content significantly increased in both varieties. However, the oxidation resistance in the roots of Zheng Taizhi 3 was enhanced compared to that of SP19 under heat stress, while IAA production was maintained and ABA content was reduced. A comparative transcriptome analysis identified 6164 and 6933 differentially expressed genes (DEGs) in Zheng Taizhi 3 and SP19, respectively, with 4346 overlapping DEGs. These DEGs included those related to stress tolerance, such as heat-shock proteins (HSPs), the antioxidant defense system, hormone signal transduction, and the biosynthetic pathway of phenylpropanoid. These findings provide insights into the physiological and molecular mechanisms underlying the adaptation of sesame to heat stress, which could inform breeding strategies for developing heat-tolerant sesame varieties.

Climate change triggers abiotic stress, such as drought and high salinity, that can cause osmotic stress. Water availability can limit plant growth, and the root tip tissues initially sense it. Most experiments destined to understand root growth adaptation to osmotic stress apply homogeneous high osmotic potentials (osmotic shock) to shoots and roots. However, this treatment does not represent natural field conditions where a root may encounter increasing osmotic potentials while exploring the soil. Osmotic shock severely reduces root growth rate, decreasing cell division in the proximal meristem and reducing mature cell length. In this work, we developed an in vitro osmotic gradient experimental system with increasing osmotic potentials. The system generates a controlled osmotic gradient in the root growth zone while exposing the aerial tissues to control conditions. The osmotic gradient system allowed Arabidopsis seedlings of Col-0 and ttl1 mutant (affected in the gene TETRATRICOPEPTIDE THIOREDOXIN-LIKE 1 ( TTL1 )) to sustain proper root growth for 25 days, reaching osmotic potentials of -1.2 MPa. We demonstrated that roots of seedlings grown in the osmotic gradient sustain a higher root growth rate than those that were grown under a homogeneous high osmotic potential. Furthermore, we found out that the expression of some genes is modified in the roots grown in the osmotic gradient compared to those grown in osmotic shock. Our data indicate that using an osmotic gradient can improve our understanding of how plants respond to osmotic stress and help find new genes to improve plant field performance., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest. The author(s) declared that they were an editorial board member of Frontiers, at the time of submission. This had no impact on the peer review process and the final decision., (Copyright © 2024 Píriz-Pezzutto, Martínez-Moré, Sainz, Borsani and Sotelo-Silveira.)

Main Conclusion: In the angiosperm root apical meristem, the holoploid DNA content is not directly related to cell cycle time. Instead, ploidy, chromosome number, and taxa emerge as key factors that influence this interaction. It is commonly considered that cell cycle time in the angiosperm root apical meristem is directly related to the holoploid DNA content, and this is one of the manifestations of the nucleotypic effect. In this paper, we significantly expanded the previously reported data on cell cycle time using the thymidine method and the rate-of-cell-production method and investigated the nucleotypic effect in different taxonomic groups, in diploids with varying numbers of chromosomes, and in species of different ploidy levels. The nucleotypic effect was most pronounced in diploids with a chromosome number of less than 17, especially in the orders Asparagales and Liliales. In diploids with 17 and a greater number of chromosomes, and in polyploids, the nucleotypic effect was only evident in these two orders and was practically absent in the rest of angiosperms, which have generally lower the holoploid DNA content and average DNA content per chromosome. Overall, our work demonstrates that in angiosperms the relationship between cell cycle time in the root apical meristem and the holoploid DNA content is complex and not homogeneous and it is not directly related to cell cycle time. Instead, ploidy, chromosome number, and taxa emerge as key factors that influence this interaction., (© 2024. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Chromium (Cr) contamination in agricultural soils, largely due to industrial activities, poses a significant threat to plant growth and productivity. This study examines the effects of Cr stress at concentrations of 100 and 200 mg of K 2 Cr 2 O 7 per kg soil on rapeseed (Brassica napus) roots and evaluates the mitigating potential of biochar. Biochar, produced through both slow and fast pyrolysis and applied at 30 g per kg soil, was investigated for its ability to neutralize Cr toxicity. Our findings indicate that Cr stress significantly decreased the growth and physiological functions of rapeseed roots. However, biochar application improved soil pH, cation exchange capacity, and the uptake of essential nutrients such as nitrogen, phosphorus, potassium, calcium, and magnesium. Additionally, biochar enhanced the production of osmotic regulators like glycine betaine and soluble proteins, as well as indole acetic acid, promoting better root growth and water uptake under Cr stress. Notably, biochar reduced Cr availability and absorption in rapeseed roots, leading to lower levels of stress-related hormones such as abscisic acid, salicylic acid, and jasmonic acid. Among the biochars tested, slow pyrolysis biochar was more effective than fast pyrolysis biochar in mitigating Cr toxicity. These results highlight the potential of slow pyrolysis biochar as a sustainable strategy to alleviate Cr pollution and enhance plant resilience in contaminated soils., Competing Interests: Declaration of competing interest The authors declare no conflict of interest., (Copyright © 2024 Elsevier Masson SAS. All rights reserved.)

Introduction: Peanut production is being increasingly threatened by water stress with the context of global climate change. Film mulching have been reported to alleviate the adverse impact of drought on peanut. Lower phosphorus use efficiency is another key factor limiting peanut yield. Application of iron-modified and phosphorus-loaded biochar (B IP ) has been validated to enhance phosphorus utilization efficiency in crops. However, whether combined effect of film mulching and B IP could increase water use efficiency and enhance peanut production through regulating soil properties and root morphologies needs further investigation., Methods: A two-year (2021-2022) pot experiment using a split-plot design was conducted to investigate the effects of phosphorus fertilizer substitution using B IP on soil properties, root morphology, pod yield, and water use of peanut under film mulching. The main plots were two mulching methods, including no mulching (M0) and film mulching (M1). The subplots were four combined applications of phosphorus fertilizer with B IP , including conventional phosphorus fertilizer rates (PCR) without B IP , P1C0; 3/4 PCR with 7.5 t ha -1 B IP , P2C1; 3/4 PCR with 15 t ha -1 B IP , P2C2; 2/3 PCR with 7.5 t ha -1 B IP , P3C1; 2/3 PCR with 15 t ha -1 B IP , P3C2., Results and Discussion: The results indicated that regardless of biochar amendments, compared with M0, M1 increased soil organic matter and root morphology of peanut at different growth stages in both years. In addition, M1 increased peanut yield and water use efficiency (WUE) by 18.8% and 51.6%, respectively, but decreased water consumption by 25.0%, compared to M0 (two-year average). Irrespective of film mulching, P2C1 increased length, surface area, and volume of peanut root at seedling by 16.7%, 17.7%, and 18.6%, at flowering by 6.6%, 19.9%, and 29.5%, at pod setting by 22.9%, 33.8%, and 37.3%, and at pod filling by 48.3%, 9.5%, and 38.2%, respectively (two-year average), increased soil pH and organic matter content during peanut growing season, and increased soil CEC at harvest. In general, the M1P2C1 treatment obtained the optimal root morphology, soil chemical properties, WUE, and peanut yield, which increased peanut yield by 33.2% compared to M0P1C0. In conclusion, the combination of film mulching with 7.5 t ha -1 B IP (M1P2C1) effectively improved soil chemical properties, enhanced root morphology of peanut, and ultimately increased peanut yield and WUE., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2024 Luo, Wang, Liu, Qiu, Zheng, Xia, Elbeltagi and Chi.)

Key Message: SiDVLs inhibit auxin signaling to regulate root growth by enhancing the expression of Aux/IAAs and reducing the protein accumulation of PINs. The DEVIL/ ROTUNDIFOLIA (DVL/RTFL), a small polypeptide family, is conserved in seed plants and important in regulating plant growth and development. However, the molecular mechanisms remain largely unknown. Here, 27 SiDVLs were identified in foxtail millet genome. Overexpression of three SiDVLs in Arabidopsis (Arabidopsis thaliana) strongly repressed the plant growth, especially the root growth. We demonstrate that overexpression of SiDVLs enhances Auxin/Indole-3-Acetic Acids (Aux/IAAs) transcription, thereby weakening auxin signaling in the roots. Furthermore, SiDVLs reduced the protein levels of the auxin transporters PIN-formed 1 (PIN1), PIN2, and PIN7 in the roots. The impaired auxin signaling reduces the cell division and elongation. In conclusion, SiDVLs suppress cell division and elongation in root by inhibiting auxin signaling and transport, which lead to the reduced root growth., (© 2024. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Root-root communication effects on several physiological and metabolic aspects among Solanaceae relatives were studied. We examined cherry (C) and field (F) tomato (Solanum lycopersicum) and bell pepper (B) (Capsicum annuum), comprising three degrees of relatedness (DOR): high (H-DOR; CC, FF and BB), medium (M-DOR; CF) and low (L-DOR; CB and FB). Plants were grown in pairs of similar or different plants on a paper-based and non-destructive root growth system, namely, rhizoslides. Root growth, including the proliferation of fine roots, and respiration increased as the DOR decreased and were highest in paired L-DOR plants, as was shown for root respiration that increased by 63, 110 and 88 % for C, F, and B when grown with B, B and F, respectively. On the other hand, root exudates of L-DOR plants had significantly lower levels of total organic carbon and protein than those of H-DOR plants, indicating different root-root communication between individuals with different DOR. Our findings indicate, for the first time, that carbon allocation to root growth, exudation and respiration depends on the degree of genetic relatedness, and that the degree of relatedness between individual plants plays a key role in the root-root communication within Solanaceae., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier B.V. All rights reserved.)

Sugars derived from photosynthesis, specifically sucrose, are the primary source of plant energy. Sucrose is produced in leaves and transported to the roots through the phloem, serving as a vital energy source. Environmental conditions can result in higher or lower photosynthesis, promoting anabolism or catabolism, respectively, thereby influencing the sucrose budget available for roots. Plants can adjust their root system to optimize the search for soil resources and to ensure the plant's adaptability to diverse environmental conditions. Recently, emerging research indicates that SNF1-RELATED PROTEIN KINASE 1 (SnRK1), trehalose 6-phosphate (T6P), and TARGET OF RAPAMYCIN (TOR) collectively serve as fundamental regulators of root development, together forming a signaling module to interpret the nutritional status of the plant and translate this to growth adjustments in the below ground parts., Competing Interests: Declaration of interests No interests are declared., (Crown Copyright © 2024. Published by Elsevier Ltd. All rights reserved.)

Autopolyploidization, which refers to a polyploidization via genome duplication without hybridization, promotes growth in autotetraploids, but suppresses growth in high polyploids (autohexaploids or auto-octoploids). The mechanism underlying this growth suppression (i.e. 'high-ploidy syndrome') has not been comprehensively characterized. In this study, we conducted a kinematic analysis of the root apical meristem cells in Arabidopsis thaliana autopolyploids (diploid, tetraploid, hexaploid, and octoploid) to determine the effects of the progression of genome duplication on root growth. The results of the root growth analysis showed that tetraploidization increases the cell volume, but decreases cell proliferation. However, cell proliferation and volume growth are suppressed in high polyploids. Whole-mount fluorescence in situ hybridization analysis revealed extensive chromosome polytenization in the region where cell proliferation does not usually occur in the roots of high polyploids, which is likely to be at least partly correlated with the suppression of endoreduplication. The study findings indicate that chromosome polytenization is important for the suppressed growth of high polyploids., (© The Author(s) 2024. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For commercial re-use, please contact reprints@oup.com for reprints and translation rights for reprints. All other permissions can be obtained through our RightsLink service via the Permissions link on the article page on our site—for further information please contact journals.permissions@oup.com.)

The triple response phenotype is characteristic for seedlings treated with the phytohormone ethylene or its direct precursor 1-aminocyclopropane-carboxylic acid, and is often employed to find novel chemical tools to probe ethylene responses. We identified a benzoxazole-urea derivative (B2) partially mimicking ethylene effects in a triple response bioassay. A phenotypic analysis demonstrated that B2 and its closest analogue arinole (ARI) induced phenotypic responses reminiscent of seedlings with elevated levels of auxin, including impaired hook development and inhibition of seedling growth. Specifically, ARI reduced longitudinal cell elongation in roots, while promoting cell division. In contrast to other natural or synthetic auxins, ARI mostly acts as an inducer of adventitious root development, with only limited effects on lateral root development. Quantification of free auxins and auxin biosynthetic precursors as well as auxin-related gene expression demonstrated that ARI boosts global auxin levels. In addition, analyses of auxin reporter lines and mutants, together with pharmacological assays with auxin-related inhibitors, confirmed that ARI effects are facilitated by TRYPTOPHAN AMINOTRANSFERASE1 (TAA1)-mediated auxin synthesis. ARI treatment in an array of species, including Arabidopsis, pea, tomato, poplar, and lavender, resulted in adventitious root formation, which is a desirable trait in both agriculture and horticulture., (© The Author(s) 2024. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For commercial re-use, please contact reprints@oup.com for reprints and translation rights for reprints. All other permissions can be obtained through our RightsLink service via the Permissions link on the article page on our site—for further information please contact journals.permissions@oup.com.)

In sweet potato, rational nitrogen (N) assimilation and distribution are conducive to inhibiting vine overgrowth. Nitrate (NO 3 - ) is the main N form absorbed by roots, and cultivar is an important factor affecting N utilization. Herein, a hydroponic experiment was conducted that included four NO 3 - concentrations of 0 (N0), 4 (N1), 8 (N2) and 16 (N3) mmol L -1 with two cultivars of Jishu26 (J26, N-sensitive) and Xushu32 (X32, N-tolerant). For J26, with increasing NO 3 - concentrations, the root length and root surface area significantly decreased. However, no significant differences were observed in these parameters for X32. Higher NO 3 supplies on root growth and NO - concentrations upregulated the expression levels of the genes that encode nitrate reductase (NR2), nitrite reductase (NiR2) and nitrate transporter (NRT1.1) in roots for both cultivars. The trends in the activities of NR and NiR were subject to regulation of NR2 and NiR2 transcription, respectively. For both cultivars, N2 increased the N accumulated in leaves, growth points and roots. For J26, N3 further increased the N accumulation in these organs. Under higher NO 3 - nutrition, compared with X32, J26 exhibited higher expression levels of the NiR2, NR2 and NRT1.1 genes, a higher influx NO 3 - rate in roots, and higher activities of NR and NiR in leaves and roots. Conclusively, the regulated effects of NO 3 - supplies on root growth and NO 3 - utilization were more significant for J26. Under high NO 3 - conditions, J26 exhibited higher capacities of NO 3 - absorption and distributed more N in leaves and in growth points, which may contribute to higher growth potential in shoots and more easily cause vine overgrowth., (© 2024. The Author(s).)

Background: Root-zone hole fertilization has a positive impact on enhancing crop production and fertilization efficiency. However, a suitable spacing for hole fertilization in rapeseed cultivation is unclear. To explore an adaptive hole spacing for improving rapeseed yield and fertilization efficiency, field experiments were conducted. Four spacings of hole fertilization were designed: 10 (FD10), 20 (FD20), 30 (FD30) and 40 cm (FD40), using no fertilization (F0) and deep-banded placement of fertilizer (DBP) as controls. The burial depth was 10 cm for FD and DBP treatments., Results: Compared to DBP, hole fertilization impacted soil microenvironment, crop growth and yield components, resulting in a significant increase of 28.4% in seed yield and 25.6% in oil yield. Seed yield in FD20 (4345.43 kg ha -1 ) increased by 4.3%, 9.4% and 15.1% compared to FD10, FD30 and FD40, respectively. Fertilizer partial factor productivity under FD20 was 4.2%, 8.6% and 13.9% greater than FD10, FD30 and FD40, respectively; whereas the increase for agronomic efficiency was 6.0%, 12.7% and 21.0%, and the increase for N recovery efficiency was 39.5%, 52.5% and 62.9%, respectively., Conclusion: Fertilization with a hole spacing of 17 cm is a promising practice to maintain high production and fertilization efficiency when cultivating rapeseed. These results provide a theoretical foundation and scientific basis for improving rapeseed productivity and fertilizer utilization. © 2024 Society of Chemical Industry., (© 2024 Society of Chemical Industry.)

Introduction: The depth of fertilizer application significantly influences soil nitrate concentration (SNC), sunflower root length density (RLD), sunflower nitrogen uptake (SNU), and yield. However, current studies cannot precisely capture subtle nutrient variations between soil layers and their complex relationships with root growth. They also struggle to assess the impact of different fertilizer application depths on sunflower root development and distribution as well as their response to the spatial and temporal distribution of nutrients., Methods: The Agricultural Production Systems sIMulator (APSIM) model was employed to explore the spatial and temporal patterns of nitrogen distribution in the soil at three controlled-release fertilizer (CRF) placement depths: 5, 15, and 25 cm. This study investigated the characteristics of the root system regarding nitrogen absorption and utilization and analyzed their correlation with sunflower yield formation. Furthermore, this study introduced the modified Jaccard index (considering the compatibility between soil nitrate and root length density) to analyze soil-root interactions, providing a deeper insight into how changes in CRF placement depth affect crop growth and nitrogen uptake efficiency., Results: The results indicated that a fertilization depth of 15 cm improved the modified Jaccard index by 6.60% and 7.34% compared to 5 cm and 25 cm depths, respectively, maximizing sunflower yield (an increase of 9.44%) and nitrogen absorption rate (an increase of 5.40%). This depth promoted a greater Root Length Density (RLD), with an increases of 11.95% and 16.42% compared those at 5 cm and 25 cm, respectively, enhancing deeper root growth and improving nitrogen uptake. In contrast, shallow fertilization led to higher nitrate concentrations in the topsoil, whereas deeper fertilization increased the nitrate concentrations in the deeper soil layers., Discussion: These results provide valuable insights for precision agriculture and sustainable soil management, highlighting the importance of optimizing root nitrogen absorption through tailored fertilization strategies to enhance crop production efficiency and minimize environmental impact., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2024 Ren, Li, Liu, Chen, Xin, Liu, Qi and Li.)

Plant secreted peptides RAPID ALKALINISATION FACTORs (RALFs), which act through the receptor FERONIA (FER), play important roles in plant growth. However, it remains unclear whether and how RALF-FER contributes to the trade-off of plant growth-defense. Here, we used a variety of techniques such as CRISPR/Cas9, protein-protein interaction and transcriptional regulation methods to investigate the role of RALF2 and its receptor FER in regulating lignin deposition, root growth, and defense against Fusarium oxysporum f. sp. lycopersici (Fol) in tomato (Solanum lycopersicum). The ralf2 and fer mutants show reduced primary root length, elevated lignin accumulation, and enhanced resistance against Fol than the wild-type. FER interacts with and phosphorylates MYB63 to promote its degradation. MYB63 serves as an activator of lignin deposition by regulating the transcription of dirigent protein gene DIR19. Mutation of DIR19 suppresses lignin accumulation, and reverses the short root phenotype and Fol resistance in ralf2 or fer mutant. Collectively, our results demonstrate that the RALF2-FER-MYB63 module fine-tunes root growth and resistance against Fol through regulating the deposition of lignin in tomato roots. The study sheds new light on how plants maintain the growth-defense balance via RALF-FER., (© 2024 The Authors. New Phytologist © 2024 New Phytologist Foundation.)

Selenium (Se) has been widely reported to affect plant growth, nutrient cycling, and the rhizobiome. However, how Se shapes the rhizobiome and interacts with plants remains largely elusive. Pot and hydroponic experiments were employed to elucidate the regulatory mechanism of Se in the citrus rhizobiome. Compared to the control, soil Se application significantly increased the root biomass (34.7%) and markedly reduced rhizosphere HCl-P, H 2 O-P, NaHCO 3 -IP, and residual-P of citrus, which were related to the variation of citrus rhizobiome. Se primarily enriched Proteobacteria and Actinobacteria as well as the phosphorus (P) functional genes phod and pqqc . Further study revealed that Se altered the metabolite profile of root exudate, particularly enhancing the abundance of l-cyclopentylglycine, cycloleucine, l-proline, l-pipecolic acid, and inositol, which played a key role in reshaping the citrus rhizobiome. These metabolites could serve as both nutrient sources and signaling molecules, thus supporting the growth or chemotaxis of the functional microbes. These bacterial taxa have the potential to solubilize P or stimulate plant growth. These findings provide a novel mechanistic understanding of the intriguing interactions between Se, root exudate, and rhizosphere microbiomes, and demonstrate the potential for utilizing Se to regulate rhizobiome function and enhance soil P utilization in citrus cultivation.

Freezing stress can seriously affect plant growth and development, but the mechanisms of these effects and plant responses to freezing stress require further exploration. Here, we identified a NAM, ATAF1/2, and CUC2 (NAC)-family transcription factor (TF), NAC056, that can promote freezing tolerance in Arabidopsis. NAC056 mRNA levels are strongly induced by freezing stress in roots, and the nac056 mutant exhibits compromised freezing tolerance. NAC056 acts positively in response to freezing by directly promoting key C-repeat-binding factor (CBF) pathway genes. Interestingly, we found that CBF1 regulates nitrate assimilation by regulating the nitrate reductase gene NIA1 in plants; therefore, NAC056-CBF1-NIA1 form a regulatory module for the assimilation of nitrate and the growth of roots under freezing stress. In addition, 35S::NAC056 transgenic plants show enhanced freezing tolerance, which is partially reversed in the cbfs triple mutant. Thus, NAC056 confers freezing tolerance through the CBF pathway, mediating plant responses to balance growth and freezing stress tolerance., (Copyright © 2024 The Author(s). Published by Elsevier Inc. All rights reserved.)

Proline metabolism plays a crucial role in both environmental stress responses and plant growth. However, the specific mechanism by which proline contributes to abiotic stress processes remains to be elucidated. In this study, we utilized atrzf1 (Arabidopsis thaliana ring zinc finger 1) as a parental line for T-DNA tagging mutagenesis and identified a suppressor mutant of atrzf1, designated proline content alterative 31 (pca31). The pca31 mutant suppressed the insensitivity of atrzf1 to dehydration stress during early seedling growth. Using Thermal Asymmetric Interlaced-PCR, we found that the T-DNA of pca31 was inserted into the promoter region of the At2g22620 gene, which encodes the cell wall enzyme rhamnogalacturonan lyase 1 (RGL1). Enzymatic assays indicated that RGL1 exhibited rhamnogalacturonan lyase activity, influencing cell wall pectin composition. The decrease in RGL1 gene expression suppressed the transcriptomic perturbation of the atrzf1 mutant. Silencing of the RGL1 gene in atrzf1 resulted in a sensitive phenotype similar to pca31 under osmotic stress conditions. Treatment with mannitol, salt, hydrogen peroxide, and abscisic acid induced RGL1 expression. Furthermore, we uncovered that RGL1 plays a role in modulating root growth and vascular tissue development. Molecular, physiological, and genetic experiments revealed that the positive modulation of RGL1 during abiotic stress was linked to the AtRZF1 pathway. Taken together, these findings establish that pca31 acts as a suppressor of atrzf1 in abiotic stress responses through proline and cell wall metabolisms., (© 2024 The Authors. The Plant Journal published by Society for Experimental Biology and John Wiley & Sons Ltd.)

Cultivating high-yield wheat under limited water resources is crucial for sustainable agriculture in semiarid regions. Amid water scarcity, plants activate drought response signaling, yet the delicate balance between drought tolerance and development remains unclear. Through genome-wide association studies and transcriptome profiling, we identified a wheat atypical basic helix-loop-helix (bHLH) transcription factor (TF), TabHLH27-A1, as a promising quantitative trait locus candidate for both relative root dry weight and spikelet number per spike in wheat. TabHLH27-A1/B1/D1 knock-out reduced wheat drought tolerance, yield, and water use efficiency (WUE). TabHLH27-A1 exhibited rapid induction with polyethylene glycol (PEG) treatment, gradually declining over days. It activated stress response genes such as TaCBL8-B1 and TaCPI2-A1 while inhibiting root growth genes like TaSH15-B1 and TaWRKY70-B1 under short-term PEG stimulus. The distinct transcriptional regulation of TabHLH27-A1 involved diverse interacting factors such as TaABI3-D1 and TabZIP62-D1. Natural variations of TabHLH27-A1 influence its transcriptional responses to drought stress, with TabHLH27-A1 Hap-II associated with stronger drought tolerance, larger root system, more spikelets, and higher WUE in wheat. Significantly, the excellent TabHLH27-A1 Hap-II was selected during the breeding process in China, and introgression of TabHLH27-A1 Hap-II allele improved drought tolerance and grain yield, especially under water-limited conditions. Our study highlights TabHLH27-A1's role in balancing root growth and drought tolerance, providing a genetic manipulation locus for enhancing WUE in wheat., (© 2024 Institute of Botany, Chinese Academy of Sciences.)

Enhancing root development is pivotal for boosting crop yield and augmenting stress resilience. In this study, we explored the regulatory effects of xylooligosaccharides (XOSs) on lettuce root growth, comparing their impact with that of indole-3-butyric acid potassium salt (IBAP). Treatment with XOS led to a substantial increase in root dry weight (30.77%), total root length (29.40%), volume (21.58%), and surface area (25.44%) compared to the water-treated control. These enhancements were on par with those induced by IBAP. Comprehensive phytohormone profiling disclosed marked increases in indole-3-acetic acid (IAA), zeatin riboside (ZR), methyl jasmonate (JA-ME), and brassinosteroids (BRs) following XOS application. Through RNA sequencing, we identified 3807 differentially expressed genes (DEGs) in the roots of XOS-treated plants, which were significantly enriched in pathways associated with manganese ion homeostasis, microtubule motor activity, and carbohydrate metabolism. Intriguingly, approximately 62.7% of the DEGs responsive to XOS also responded to IBAP, underscoring common regulatory mechanisms. However, XOS uniquely influenced genes related to cutin, suberine, and wax biosynthesis, as well as plant hormone signal transduction, hinting at novel mechanisms of stress tolerance. Prominent up-regulation of genes encoding beta-glucosidase and beta-fructofuranosidase highlights enhanced carbohydrate metabolism as a key driver of XOS-induced root enhancement. Collectively, these results position XOS as a promising, sustainable option for agricultural biostimulation.

The pH of the solution in the rhizosphere is an important factor that determines the availability and mobility of nutrients for plant uptake. Solution pH may also affect the root distribution and architecture of apple rootstocks. In this study, we evaluated the effect of solution pH on root system development of apple rootstocks using an aeroponics system designed and developed at Cornell AgriTech Geneva, USA. Four Geneva ® apple rootstocks (G.210, G.214, G.41, and G.890) were grown in an aeroponic system under nutrient solution misting featuring continuously adjusted pH levels to three pH treatments (5.5, 6.5, and 8.0). Root development was monitored for 30 days and evaluated regularly for distribution and root mass. Images of the developed roots grown in the aeroponic system were collected at the end of the experiment using a high-resolution camera and analyzed using GiA Roots ® software, which generates root architecture parameter values in a semi-automated fashion. The resulting root architecture analysis showed that the Geneva ® rootstocks were significantly different for two architecture parameters. The length-to-width ratio analysis represented by two GiA Roots parameters (minor-to-major ellipse ratio and network width-to-depth ratio) showed that G.210 was flatter than G.890, which had a greater tendency to grow downward. Rootstocks G.214 and G.41 displayed similar growth values. The solution pH affected most root architecture parameter measurements where overall root growth was higher at pH 8 than at pH 5.5 and 6.5, which showed similar growth. In general, the average root width tended to decrease at higher pH values. While there were no significant differences in the leaf nutrient concentrations of P, K, Ca, Mg, S, B, Zn, Cu, and Fe within the four rootstocks, the pH level of the solution had a significant effect on P, Ca, and Mn. This study is the first of its kind to investigate the effect of pH on root architecture in a soil-free (aeroponic) environment and may have implications for apple root behavior under field conditions where pH levels are different., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2024 Farqani, Cheng, Robinson and Fazio.)

In crops like wheat, terminal drought is one of the principal stress factors limiting productivity in rain-fed systems. However, little is known about root development after heading, when water uptake can be critical to wheat crops. The impact of water-stress on root growth was investigated in two wheat cultivars, Scout and Mace, under well-watered and post-anthesis water stress in three experiments. Plants were grown outside in 1.5-m long pots at a density similar to local recommended farming practice. Differences in root development were observed between genotypes, especially for water stress conditions under which Scout developed and maintained a larger root system than Mace. While under well-watered conditions both genotypes had shallow roots that appeared to senesce after heading, a moderate water stress stimulated shallow-root growth in Scout but accelerated senescence in Mace. For deep roots, post-heading biomass growth was observed for both genotypes in well-watered conditions, while under moderate water stress, only Scout maintained net growth as Mace deep roots senesced. Water stress of severe intensity affected both genotypes similarly, with root senescence at all depths. Senescence was also observed above ground. Under well-watered conditions, Scout retained leaf greenness (i.e. stay-green phenotype) for slightly longer than Mace. The difference between genotypes accentuated under moderate water stress, with rapid post-anthesis leaf senescence in Mace while Scout leaf greenness was affected little if at all by the stress. As an overall result, grain biomass per plant ('yield') was similar in the two genotypes under well-watered conditions, but more affected by a moderate stress in Mace than Scout. The findings from this study will assist improvement in modelling root systems of crop models, development of relevant phenotyping methods and selection of cultivars with better adaptation to drought., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2024 Shazadi, Christopher and Chenu.)

MicroRNAs are known to play a crucial role in plant development and physiology and become a target for investigating the regulatory mechanism underlying plant low phosphate tolerance. ZmmiR528 has been shown to display significantly different expression levels between wild-type and low Pi-tolerant maize mutants. However, its functional role in maize low Pi tolerance remains unknown. In the present study, we studied the role and underlying molecular mechanism of miR528 in maize with low Pi tolerance. Overexpression of ZmmiR528 in maize resulted in impaired root growth, reduced Pi uptake capacity and compromised resistance to Pi deficiency. By contrast, transgenic maize plants suppressing ZmmiR528 expression showed enhanced low Pi tolerance. Furthermore, ZmLac3 and ZmLac5 which encode laccase were identified and verified as targets of ZmmiR528. ZmLac3 transgenic plants were subsequently generated and were also found to play key roles in regulating maize root growth, Pi uptake and low Pi tolerance. Furthermore, auxin transport was found to be potentially involved in ZmLac3-mediated root growth. Moreover, we conducted genetic complementary analysis through the hybridization of ZmmiR528 and ZmLac3 transgenic plants and found a favorable combination with breeding potential, namely anti-miR528:ZmLac3OE hybrid maize, which exhibited significantly increased low Pi tolerance and markedly alleviated yield loss caused by low Pi stress. Our study has thus identified a ZmmiR528-ZmLac3 module regulating auxin transport and hence root growth, thereby determining Pi uptake and ultimately low Pi tolerance, providing an effective approach for low Pi tolerance improvement through manipulating the expression of miRNA and its target in maize., (© 2024 Society for Experimental Biology and John Wiley & Sons Ltd.)

Potassium (K + ), being an essential macronutrient in plants, plays a central role in many aspects. Root growth is highly plastic and is affected by many different abiotic stresses including nutrient deficiency. The Shaker-type K + channel Arabidopsis (Arabidopsis thaliana) K + Transporter 1 (AKT1) is responsible for K + uptake under both low and high external K + conditions. However, the upstream transcription factor of AKT1 is not clear. Here, we demonstrated that the WRKY6 transcription factor modulates root growth to low potassium (LK) stress in Arabidopsis. WRKY6 showed a quick response to LK stress and also to many other abiotic stress treatments. The two wrky6 T-DNA insertion mutants were highly sensitive to LK treatment, whose primary root lengths were much shorter, less biomass and lower K + content in roots than those of wild-type plants, while WRKY6-overexpression lines showed opposite phenotypes. A further investigation showed that WRKY6 regulated the expression of the AKT1 gene via directly binding to the W-box elements in its promoter through EMSA and ChIP-qPCR assays. A dual luciferase reporter analysis further demonstrated that WRKY6 enhanced the transcription of AKT1. Genetic analysis further revealed that the overexpression of AKT1 greatly rescued the short root phenotype of the wrky6 mutant under LK stress, suggesting AKT1 is epistatic to WRKY6 in the control of LK response. Further transcriptome profiling suggested that WRKY6 modulates LK response through a complex regulatory network. Thus, this study unveils a transcription factor that modulates root growth under potassium deficiency conditions by affecting the potassium channel gene AKT1 expression., (© 2024 Society for Experimental Biology and John Wiley & Sons Ltd.)

Various plant species can be selected for environmental testing, including pearl millet (Pennisetum glaucum (L.) R. Br), a globally significant cereal crop. This study aims to assess millet's suitability as a species for ecotoxicological tests, examining (1) germination and initial development dynamics, (2) the minimum seed quantity for reliable sampling, (3) optimal experimental design with replication numbers, (4) suitability of positive control, and (5) the effectiveness of the protocol in evaluating toxic effects of environmental pollutants. Millet exhibited rapid and uniform germination as well as consistent initial seedling development. To establish the minimum number of seeds required for reliable experimentation, germination, and seedling growth were compared across plots containing 10, 25, and 50 seeds. Consequently, 10 seeds per plot were chosen for subsequent experiments to reduce labor and costs while maintaining reliability. To validate the selected experimental design, and to establish a positive control for assays, aluminum was used as a toxic element at concentrations ranging from 10 -2 to 10 -6 M. While aluminum did not affect the final percentage of germinated seeds, it did exhibit an impact on the Germination Speed Index (GSI). Significant differences in root and aerial growth, and with fresh weight, were observed. The 10 -3 M concentration was chosen as the positive control as the 10 -2 concentration showed extreme toxicity. To assess the applicability of the established protocol in determining the toxic effects of environmental pollutants, millet roots were exposed to the toxic agents atrazine, cadmium, methyl methane sulfonate (MMS), and Spent pot liner (SPL). Millet demonstrated sensitivity and efficiency in response to these tests. In conclusion, millet proves to be an effective species for the toxicological risk assessment of environmental pollutants., (© 2024. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Currently, the control of rhizosphere selection on farms has been applied to achieve enhancements in phenotype, extending from improvements in single root characteristics to the dynamic nature of entire crop systems. Several specific signals, regulatory elements, and mechanisms that regulate the initiation, morphogenesis, and growth of new lateral or adventitious root species have been identified, but much more work remains. Today, phenotyping technology drives the development of root traits. Available models for simulation can support all phenotyping decisions (root trait improvement). The detection and use of markers for quantitative trait loci (QTLs) are effective for enhancing selection efficiency and increasing reproductive genetic gains. Furthermore, QTLs may help wheat breeders select the appropriate roots for efficient nutrient acquisition. Single-nucleotide polymorphisms (SNPs) or alignment of sequences can only be helpful when they are associated with phenotypic variation for root development and elongation. Here, we focus on major root development processes and detail important new insights recently generated regarding the wheat genome. The first part of this review paper discusses the root morphology, apical meristem, transcriptional control, auxin distribution, phenotyping of the root system, and simulation models. In the second part, the molecular genetics of the wheat root system, SNPs, TFs, and QTLs related to root development as well as genome editing (GE) techniques for the improvement of root traits in wheat are discussed. Finally, we address the effect of omics strategies on root biomass production and summarize existing knowledge of the main molecular mechanisms involved in wheat root development and elongation., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2024 Alrajhi, Alharbi, Beecham and Alotaibi.)

The rhizotoxicity of protons (H+) in acidic soils is a fundamental constraint that results in serious yield losses. However, the mechanisms underlying H+-mediated inhibition of root growth are poorly understood. In this study, we revealed that H+-induced root growth inhibition in Arabidopsis depends considerably on excessive iron deposition in the root apoplast. Reducing such aberrant iron deposition by decreasing the iron supply or disrupting the ferroxidases LOW PHOSPHATE ROOT 1 (LPR) and LPR2 attenuates the inhibitory effect of H+ on primary root growth efficiently. Further analysis showed that excessive iron deposition triggers a burst of highly reactive oxygen species, consequently impairing normal root development. Our study uncovered a valuable strategy for improving the ability of plants to tolerate H+ toxicity by manipulating iron availability., (© The Author(s) 2024. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For commercial re-use, please contact reprints@oup.com for reprints and translation rights for reprints. All other permissions can be obtained through our RightsLink service via the Permissions link on the article page on our site—for further information please contact journals.permissions@oup.com.)

Auxins play crucial regulatory roles in plants coping with cadmium (Cd) stress. However, the regulatory mechanism by which auxins alleviate Cd toxicity in tomato seedlings remains unclear. Here, we demonstrate that exposure to Cd stress leads to dynamic changes in the auxin response in tomato roots, characterized by an initial increase followed by a subsequent weakening. Under Cd stress, tomato seedlings show primary root- and hypocotyl-growth inhibition, accompanied by the accumulation of Cd and reactive oxygen species (ROS) in the roots. The exogenous application of 1-naphthylacetic acid (NAA) does not mitigate the inhibitory effect of Cd toxicity on primary root growth, but it does significantly enhance lateral root development under Cd stress. Auxin transport inhibitors, such as 1-N-naphthylphthalamic acid (NPA) and 2,3,5-triiodobenoic acid (TIBA), aggravate the growth inhibition of primary roots caused by Cd stress. Additionally, lateral root development was inhibited by NPA. However, applying auxin synthesis inhibitors L-kynurenine (kyn) and yucasin alleviated the tomato root growth inhibition caused by Cd stress; between them, the effect of yucasin was more pronounced. Yucasin mitigates Cd toxicity in tomato seedlings by reducing Cd 2+ absorption and auxin accumulation, strengthening ROS scavenging, and reducing cell death in roots. These observations suggest that yucasin potentially mitigates Cd toxicity and improves the tolerance of tomato seedlings to Cd stress.