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The endodermis in roots acts as a selectivity filter for nutrient and water transport essential for growth and development. This selectivity is enabled by the formation of lignin-based Casparian strips. Casparian strip formation is initiated by the localization of the Casparian strip domain proteins (CASPs) in the plasma membrane, at the site where the Casparian strip will form. Localized CASPs recruit Peroxidase 64 (PER64), a Respiratory Burst Oxidase Homolog F, and Enhanced Suberin 1 (ESB1), a dirigent-like protein, to assemble the lignin polymerization machinery. However, the factors that control both expression of the genes encoding this biosynthetic machinery and its localization to the Casparian strip formation site remain unknown. Here, we identify the transcription factor, MYB36, essential for Casparian strip formation. MYB36 directly and positively regulates the expression of the Casparian strip genesCASP1, PER64,andESB1. Casparian strips are absent in plants lacking a functionalMYB36and are replaced by ectopic lignin-like material in the corners of endodermal cells. The barrier function of Casparian strips in these plants is also disrupted. Significantly, ectopic expression ofMYB36in the cortex is sufficient to reprogram these cells to start expressingCASP1–GFP, correctly localize the CASP1–GFP protein to form a Casparian strip domain, and deposit a Casparian strip-like structure in the cellwall at this location. These results demonstrate that MYB36 is controlling expression of the machinery required to locally polymerize lignin in a fine band in the cell wall for the formation of the Casparian strip.

Suberin is a hydrophobic biopolymer that can be deposited at the periphery of cells, forming protective barriers against biotic and abiotic stress. In roots, suberin forms lamellae at the periphery of endodermal cells where it plays crucial roles in the control of water and mineral transport. Suberin formation is highly regulated by developmental and environmental cues. However, the mechanisms controlling its spatiotemporal regulation are poorly understood. Here, we show that endodermal suberin is regulated independently by developmental and exogenous signals to fine-tune suberin deposition in roots. We found a set of four MYB transcription factors (MYB41, MYB53, MYB92, and MYB93), each of which is individually regulated by these two signals and is sufficient to promote endodermal suberin. Mutation of these four transcription factors simultaneously through genome editing leads to a dramatic reduction in suberin formation in response to both developmental and environmental signals. Most suberin mutants analyzed at physiological levels are also affected in another endodermal barrier made of lignin (Casparian strips) through a compensatory mechanism. Through the functional analysis of these four MYBs, we generated plants allowing unbiased investigation of endodermal suberin function, without accounting for confounding effects due to Casparian strip defects, and were able to unravel specific roles of suberin in nutrient homeostasis.

Deposition of silica in plant cell walls improves their mechanical properties and helps plants to withstand various stress conditions. Its mechanism is still not understood and silica-cell wall interactions are elusive. The objective of this study was to investigate the effect of silica deposition on the development and structure of sorghum root endodermis and to identify the cell wall components involved in silicification. Sorghum bicolor seedlings were grown hydroponically with (Si+) or without (Si-) silicon supplementation. Primary roots were used to investigate the transcription of silicon transporters by quantitative RT-PCR. Silica aggregation was induced also under in vitro conditions in detached root segments. The development and architecture of endodermal cell walls were analysed by histochemistry, microscopy and Raman spectroscopy. Water retention capability was compared between silicified and non-silicified roots. Raman spectroscopy analyses of isolated silica aggregates were also carried out. Active uptake of silicic acid is provided at the root apex, where silicon transporters Lsi1 and Lsi2 are expressed. The locations of silica aggregation are established during the development of tertiary endodermal cell walls, even in the absence of silicon. Silica aggregation takes place in non-lignified spots in the endodermal cell walls, which progressively accumulate silicic acid, and its condensation initiates at arabinoxylan-ferulic acid complexes. Silicification does not support root water retention capability; however, it decreases root growth inhibition imposed by desiccation. A model is proposed in which the formation of silica aggregates in sorghum roots is predetermined by a modified cell wall architecture and takes place as governed by endodermal development. The interaction with silica is provided by arabinoxylan-ferulic acid complexes and interferes with further deposition of lignin. Due to contrasting hydrophobicity, silicification and lignification do not represent functionally equivalent modifications of plant cell walls.

Paddy rice [Oryza sativa) is able to accumulate high concentrations of Mn without showing toxicity; however, the molecular mechanisms underlying Mn uptake are unknown. Here, we report that a member of the Nramp (for the Natural Resistance-Associated Macrophage Protein) family, Nramp5, is involved in Mn uptake and subsequently the accumulation of high concentrations of Mn in rice. Nramp5 was constitutively expressed in the roots and encodes a plasma membrane-localized protein. Nramp5 was polarly localized at the distal side of both exodermis and endodermis cells. Knockout of Nramp5 resulted in a significant reduction in growth and grain yield, especially when grown at low Mn concentrations. This growth reduction could be partially rescued by supplying high concentrations of Mn but not by the addition of Fe. Mineral analysis showed that the concentration of Mn and Cd in both the roots and shoots was lower in the knockout line than in wild-type rice. A short-term uptake experiment revealed that the knockout line lost the ability to take up Mn and Cd. Taken together, Nramp5 is a major transporter of Mn and Cd and is responsible for the transport of Mn and Cd from the external solution to root cells.

This review focuses on the regulation of root water uptake in plants which are exposed to salt stress. Root water uptake is not considered in isolation but is viewed in the context of other potential tolerance mechanisms of plants—tolerance mechanisms which relate to water relations and gas exchange. Plants spend between one third and half of their lives in the dark, and salt stress does not stop with sunset, nor does it start with sunrise. Surprisingly, how plants deal with salt stress during the dark has received hardly any attention, yet any growth response to salt stress over days, weeks, months and years is the integrative result of how plants perform during numerous, consecutive day/night cycles. As we will show, dealing with salt stress during the night is a prerequisite to coping with salt stress during the day. We hope to highlight with this review not so much what we know, but what we do not know; and this relates often to some rather basic questions.

Most plant-parasitic nematodes are obligate biotrophs feeding on the roots of their hosts. Whereas ectoparasites remain on the root surface and feed on the outer cell layers, endoparasitic nematodes enter the host to parasitize cells around or within the central cylinder. Nematode invasion and feeding causes tissue damage which may, in turn, lead to the activation of host basal defence responses. Hitherto, research interests in plant–nematode interaction have emphasized effector-triggered immunity rather than basal plant defence responses. However, some recent investigations suggest that basal defence pathways are not only activated but also play an important role in determining interaction outcomes. In this review we discuss the major findings and point out future directions to dissect the molecular mechanisms underlying plant basal defence to nematodes further.

Salt tolerance and the possible functions of suberization on salt exclusion and secretion were examined in a dominant mangrove plant, Avicennia marina. The results showed that low salinities (10‰ and 20‰) almost has no negative effect on A. marina, however significant growth inhibitions were observed in the seedlings grown in higher salinities (30‰ and 40‰). With the increases of salinity, increased tissue Na+ content and enhanced salt secretion by glands were observed. Obvious suberization thickening were detected both in the exodermis and endodermis of the roots after salt pretreatment when compared to the roots without salt treatment. More importantly, the present data further confirmed that these root apoplastic barriers would directly decrease Na+ loading into xylem. Higher salt tolerance was observed in the seedlings pre-cultivated by salty tide when compared to fresh water cultivated A. marina. In summary, this study suggests a barrier property of suberization in dealing with salt exclusion in mangroves, a moderate salt pre-treatment may benefit plant withstanding high salinity.

• Background and Aims Adventitious roots (ARs) are part of the root system in numerous plants, and are required for successful micropropagation. In the Arabidopsis thaliana primary root (PR) and lateral roots (LRs), the quiescent centre (QC) in the stem cell niche of the meristem controls apical growth with the involvement of auxin and cy tokinin. In arabidopsis, ARs emerge in planta from the hypocotyl pericycle, and from different tissues in in vitro cultured expiants, e.g. from the stem endodermis in thin cell layer (TCL) expiants. The aim of this study was to investigate the establishment and maintenance of the QC in arabidopsis ARs, in planta and in TCL expiants, because information about this process is still lacking, and it has potential use for biotechnological applications. • Methods Expression of PR/LR QC markers and auxin influx (LAX3)/efftux (PINI) genes was investigated in the presence/absence of exogenous auxin and cytokinin. Auxin was monitored by the DR5∷GUS system and cytokinin by immunolocalization. The expression of the auxin-biosynthetic YUCCA6 gene was also investigated by in situ hybridization in planta and in AR-forming TCLs from the indole acetic acid (IAA)-overproducing superroot2-1 mutant and its wild type. • Key Results The accumulation of auxin and the expression of the QC marker WOX5 characterized the early derivatives of the AR founder cells, in planta and in in vitro cultured TCLs. By determination of PINI auxin efflux carrier and LAX3 auxin influx carrier activities, an auxin maximum was determined to occur at the AR tip, to which WOX5 expression was restricted, establishing the positioning of the QC. Cytokinin caused a restriction of LAX3 and PINI expression domains, and concomitantly the auxin biosynthesis YUCCA6 gene was expressed in the apex. • Conclusions In ARs formed in planta and TCLs, the QC is established in a similar way, and auxin transport and biosynthesis are involved through cytokinin tuning.

Background Homoeologs are defined as homologous genes resulting from allopolyploidy. Bread wheat, Triticum aestivum , is an allohexaploid species with many homoeologs. Homoeolog expression bias, referring to the relative contribution of homoeologs to the transcriptome, is critical for determining the traits that influence wheat growth and development. Asymmetric transcription of homoeologs has been so far investigated in a tissue or organ-specific manner, which could be misleading due to a mixture of cell types. Results Here, we perform single nuclei RNA sequencing and ATAC sequencing of wheat root to study the asymmetric gene transcription, reconstruct cell differentiation trajectories and cell-type-specific gene regulatory networks. We identify 22 cell types. We then reconstruct cell differentiation trajectories that suggest different origins between epidermis/cortex and endodermis, distinguishing bread wheat from Arabidopsis . We show that the ratio of asymmetrically transcribed triads varies greatly when analyzing at the single-cell level. Hub transcription factors determining cell type identity are also identified. In particular, we demonstrate that TaSPL14 participates in vasculature development by regulating the expression of BAM1 . Combining single-cell transcription and chromatin accessibility data, we construct the pseudo-time regulatory network driving root hair differentiation. We find MYB3R4, REF6, HDG1, and GATAs as key regulators in this process. Conclusions Our findings reveal the transcriptional landscape of root organization and asymmetric gene transcription at single-cell resolution in polyploid wheat.

The formation of Casparian strips (CS) and the deposition of suberin at the endodermis of plant roots are thought to limit the apoplastic transport of water and ions. We investigated the specific role of each of these apoplastic barriers in the control of hydro-mineral transport by roots and the consequences on shoot growth.A collection of Arabidopsis thaliana mutants defective in suberin deposition and/or CS development was characterized under standard conditions using a hydroponic system and the Phenopsis platform.Mutants altered in suberin deposition had enhanced root hydraulic conductivity, indicating a restrictive role for this compound in water transport. In contrast, defective CS directly increased solute leakage and indirectly reduced root hydraulic conductivity. Defective CS also led to a reduction in rosette growth, which was partly dependent on the hydro-mineral status of the plant. Ectopic suberin was shown to partially compensate for defective CS phenotypes. Altogether, our work shows that the functionality of the root apoplastic diffusion barriers greatly influences the plant physiology, and that their integrity is tightly surveyed.