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Improving chilling tolerance is a major target of rice breeding. The OsMAPK3-OsbHLH002-OsTPP1 signalling pathway enhances chilling tolerance in rice: the kinase is activated by cold stress, and subsequently the transcription factor is phosphorylated by the activated kinase, triggering the expression of cold response genes. However, it is largely unknown how this pathway is suppressed in time to avoid it being in a continuously activated state. We found that a novel type 2C protein phosphatase, OsPP2C27, functions as a negative regulator of the OsMAPK3-OsbHLH002-OsTPP1 pathway. A dynamic change in OsMAPK3 activity was found during cold treatment. We show that OsPP2C27 interacts physically with and dephosphorylates OsMAPK3 in vitro and in vivo. Interestingly, OsPP2C27 can also directly dephosphorylate OsbHLH002, the target of OsMAPK3. After cold treatment, survival rates were higher in OsPP2C27-RNAi lines and a T-DNA insertion mutant, and lower in OsPP2C27-overexpression lines, compared to wild type. Moreover, expression of the OsTPP1 and OsDREBs were increased in OsPP2C27-RNAi lines and decreased in OsPP2C27-overexpression lines. These results indicate that cold-induced OsPP2C27 negatively regulates the OsMAPK3-OsbHLH002-OsTPP1 signalling pathway by directly dephosphorylating both phospho-OsMAPK3 and phospho-OsbHLH002, preventing the sustained activation of a positive pathway for cold stress and maintaining normal growth under chilling conditions., (© 2020 John Wiley & Sons Ltd.)

Natural plants must actively allocate their limited resources for survival and reproduction. Although vegetative growth, sexual reproduction, asexual reproduction and defense are all basic processes in the life cycle of plants, the strategies used to allocate resources between these processes are poorly understood. These processes are conspicuous in naturally grown Aconitum kusnezoffii Reichb., which makes it a suitable study subject. Here, the morphology, dry matter, total organic carbon, total nitrogen and aconitum alkaloid levels of shoot, principal root (PR) and lateral roots were measured throughout the growing season. Then, transcriptome and metabolite content analyses were performed. We found that vegetative growth began first. After vegetative growth ceased, sexual development began. Flower organ development was accompanied by increased photosynthesis and the PR consumed temporarily stored resources after flower formation. Asexual propagule development initiated earlier than sexual reproduction and kept accumulating resources after that. Development was slow before flower formation, mainly manifesting as increasing length; then, after flower formation it accelerated via enhanced material transport and accumulation. Defense compounds were maintained at low levels before flowering. In particular, the turnover of defense compounds was enhanced before and after flower bud emergence, providing resources for other processes. After flower formation, defense compounds were accumulated. The pattern found herein provides a vivid example for further studies on resource allocation strategies. The exciting finding that the PR, as a more direct storage site for photosynthate, is a buffer unit for resources, and that defense compounds can be reused for other processes, suggests a need to explore potential mechanisms., (© 2020 Society for Experimental Biology and John Wiley & Sons Ltd.)

Environmental pH is a critical parameter for innumerable chemical reactions, myriad biological processes and all forms of life. The mechanisms that underlie the perception of external pH (pH e ) have been elucidated in detail for bacteria, fungi and mammalian cells; however, little information is available on whether and, if so, how pH e is perceived by plants. This is particularly surprising since hydrogen ion activity of the substrate is of paramount significance for plants, governing the availability of mineral nutrients, the structure of the soil microbiome and the composition of natural plant communities. Rapid changes in soil pH require constant readjustment of nutrient acquisition strategies, which is associated with dynamic alterations in gene expression. Referring to observations made in diverse experimental set-ups that unambiguously show that pH e per se affects gene expression, we hypothesize that sensing of pH e in plants is mandatory to prioritize responses to various simultaneously received environmental cues.

Although interactions between plants and microbes at the plant-soil interface are known to be important for plant nutrient acquisition, relatively little is known about how root exudates contribute to nutrient exchange over the course of plant development. In this study, root exudates from slow- and fast-growing stages of Arabidopsis thaliana plants were collected, chemically analysed and then applied to a sandy nutrient-depleted soil. We then tracked the impacts of these exudates on soil bacterial communities, soil nutrients (ammonium, nitrate, available phosphorus and potassium) and plant growth. Both pools of exudates shifted bacterial community structure. GeoChip analyses revealed increases in the functional gene potential of both exudate-treated soils, with similar responses observed for slow-growing and fast-growing plant exudate treatments. The fast-growing stage root exudates induced higher nutrient mineralization and enhanced plant growth as compared to treatments with slow-growing stage exudates and the control. These results suggest that plants may adjust their exudation patterns over the course of their different growth phases to help tailor microbial recruitment to meet increased nutrient demands during periods demanding faster growth., (© 2020 John Wiley & Sons Ltd.)

Globally, flooding is a major threat causing substantial yield decline of cereal crops, and is expected to be even more serious in many parts of the world due to climatic anomaly in the future. Understanding the mechanisms of plants coping with unanticipated flooding will be crucial for developing new flooding-tolerance crop varieties. Here we describe survival strategies of plants adaptation to flooding stress at the morphological, physiological and anatomical scale systemically, such as the formation of adventitious roots (ARs), aerenchyma and radial O 2 loss (ROL) barriers. Then molecular mechanisms underlying the adaptive strategies are summarized, and more than thirty identified functional genes or proteins associated with flooding-tolerance are searched out and expounded. Moreover, we elaborated the regulatory roles of phytohormones in plant against flooding stress, especially ethylene and its relevant transcription factors from the group VII Ethylene Response Factor (ERF-VII) family. ERF-VIIs of main crops and several reported ERF-VIIs involving plant tolerance to flooding stress were collected and analyzed according to sequence similarity, which can provide references for screening flooding-tolerant genes more precisely. Finally, the potential research directions in the future were summarized and discussed. Through this review, we aim to provide references for the studies of plant acclimation to flooding stress and breeding new flooding-resistant crops in the future.

Background: Strawberry (Fragaria × ananassa Duch.) is an important fruit crop worldwide. It was particularly sensitive to drought stress because of their fibrous and shallow root systems. Mutant rty of Arabidopsis thaliana ROOTY (RTY) results in increased endogenous auxin levels, more roots, and shoot growth. It is still unclear whether the rty gene improves stress tolerance in strawberry., Results: rty gene was isolated from Arabidopsis thaliana and placed under the control of the cauliflower mosaic virus (CaMV) 35S promoter in the pBI121-rty binary vector carrying the selectable marker of neomycin phosphotransferase II (NPT II). Seven transgenic lines were confirmed by PCR and western blot analysis. Accumulations of IAA and ABA were significantly increased in the transgenic plants. The endogenous IAA contents were 46.5 ng g - 1 and 66.0 ng g - 1 in control and transgenic plants respectively. The endogenous ABA contents in the control plant were 236.3 ng g - 1 and in transgenic plants were 543.8 ng g - 1 . The production of adventitious roots and trichomes were enhanced in the transgenic plants. Furthermore, transcript levels of the genes including IAA and ABA biosynthetic, and stress-responsive genes, were higher in the transgenic plants than in the control plants under drought conditions. Water use efficiency and a reduced water loss rate were enhanced in the transgenic strawberry plants. Additionally, peroxidase and catalase activities were significantly higher in the transgenic plants than in the control plants. The experiment results revealed a novel function for rty related to ABA and drought responses., Conclusions: The rty gene improved hormone-mediated drought tolerance in transgenic strawberry. The heterologous expression of rty in strawberry improved drought tolerance by promoting auxin and ABA accumulation. These phytohormones together brought about various physiological changes that improved drought tolerance via increased root production, trichome density, and stomatal closure. Our results suggested that a transgenic approach can be used to overcome the inherent trade-off between plant growth and drought tolerance by enhancing water use efficiency and reducing water loss rate under water shortage conditions.

Main Conclusion: Differential accumulation of root carbohydrates at defoliation have a higher impact than regrowth environmental conditions on the phenolic content and feed quality of the perennial forage legume Lotus corniculatus. The unpredictable nature of proanthocyanidin (condensed tannin) accumulation in regrowth vegetation of the perennial forage legume Lotus corniculatus represents a dilemma to the wider use of this species in agriculture, and a potential problem in the nutritional ecology of some terrestrial herbivores, as variable condensed tannin levels can result in either beneficial or detrimental effects on animal nutrition. However, the source of this variation has not been extensively explored. High levels of carbon allocation to roots during low-temperature preconditioning of clonal plants were found to significantly increase condensed tannin and flavonol levels in regrowth foliage, while low levels of carbon allocation to roots during periods of high-temperature preconditioning significantly decreased condensed tannin and flavonol levels. Phenolic accumulation and tissue digestibility were also differentially affected by regrowth of these defoliated plants at high CO 2 concentrations and by drought. Lower rates of digestion generally paralleled increases in tannin levels in regrowth leaves under the different environmental conditions, with rates of digestion falling in high tannin plants, despite correspondingly higher levels of leaf carbohydrates. Differential accumulation of root carbohydrates between seasons and years may therefore explain some of the variability found in the nutritional quality of the forage of this species.

Background: Despite significant limitations of growth medium reuse, a large amount of organic substrate is reused in soilless cultivation of horticultural crops in China. Arbuscular mycorrhizal fungi (AMF) can promote nutrient absorption and improve plant tolerance to biotic and abiotic stresses. However, the mechanisms governing the effects of AMF on crop growth in organic continuous cropping substrates have not been elucidated., Results: In this study, we showed that the inoculation of AMF in continuous cropping substrates promoted growth and root development, and increased the root and NADP-malic enzyme (NADP-ME) activity of tomato seedlings. Root transcriptome analysis demonstrated that the plant hormone signal transduction pathway was highly enriched, and 109 genes that positively correlated with the AMF-inoculated plant phenotype were obtained by gene set enrichment analysis (GSEA), which identified 9 genes related to indole acetic acid (IAA). Importantly, the levels of endogenous IAA in tomato seedlings significantly increased after AMF inoculation. Furthermore, the application of AMF significantly increased the expression levels of NADP-ME1 and NADP-ME2, as well as the activity of NADP-ME, and enhanced the root activity of tomato seedlings in comparison to that observed without inoculation of AMF. However, these effects were blocked in plants treated with 2,3,5-triiodobenzoic acid (TIBA), a polar transport inhibitor of IAA., Conclusions: These results suggest that IAA mediates the AMF-promoted tomato growth and expression of NADP-MEs in continuous cropping substrates. The study provides convincing evidence for the reuse of continuous cropping substrates by adding AMF as an amendment.

The yield and quality of the medicinal plant Achyranthes bidentata can be increased when it is replanted into a field cultivated previously with the same crop, however, fundamental aspects of its biology (so-called "replanting benefit") still remain to be elucidated. miRNAs are sRNA molecules involved in the post-transcriptional regulation of gene expression in plant biological processes. Here, 267 conserved and 36 novel miRNAs were identified in A. bidentata roots. We compared the miRNA content of the roots (R1) from first-year planting with that of the roots (R2) of second-year replanting, and screened 21 differentially expressed (DE) miRNAs. Based on in silico functional analysis, integrated miRNA-mRNA datasets allowed the identification of 10 miRNA-target family modules, which might participate in the benefit. The expression profiles of the miRNA-target modules were potentially correlated with the presence of the replanting benefit. The indication was that the miRNA-responsive continuous monoculture could reprogram miRNA-mRNA expression patterns, which possibly promote the root growth and development, enhance its transport activity and strengthen its tolerance to various stresses, thereby improving A. bidentata productivity as observed in the replanting benefit. Our study provides basic data for further research on the molecular mechanisms of the benefit in A. bidentata.

Roots have important roles for plants to withstand adverse environmental conditions, including salt stress. Biostimulant application was shown to enhance plant resilience towards abiotic stresses. Here, we studied the effect of a tannin-based biostimulant on tomato (Solanum lycopersicum L.) grown under salt stress conditions. We investigated the related changes at both root architecture (via imaging and biometric analysis) and gene expression (RNA-Seq/qPCR) levels. Moreover, in order to identify the main compounds potentially involved in the observed effects, the chemical composition of the biostimulant was evaluated by UV/Vis and HPLC-ESI-Orbitrap analysis. Sixteen compounds, known to be involved in root development and having a potential antioxidant properties were identified. Significant increase of root weight (+ 24%) and length (+ 23%) was observed when the plants were grown under salt stress and treated with the biostimulant. Moreover, transcriptome analysis revealed that the application of the biostimulant upregulated 285 genes, most of which correlated to root development and salt stress tolerance. The 171 downregulated genes were mainly involved in nutrient uptake. These data demonstrated that the biostimulant is able not only to restore root growth in salty soils, but also to provide the adequate plant nourishment by regulating the expression of essential transcription factors and stress responsive genes.

Plants encounter a variety of mechanical stimuli during their growth and development. It is currently believed that mechanosensitive ion channels play an essential role in the initial perception of mechanical force in plants. Over the past decade, the study of Piezo, a mechanosensitive ion channel in animals, has made significant progress. It has been proved that the perception of mechanical force in various physiological processes of animals is indispensable. However, little is still known about the function of its homologs in plants. In this study, by investigating the function of the AtPiezo gene in the model plant Arabidopsis thaliana , we found that AtPiezo plays a role in the perception of mechanical force in plant root cap and the flow of Ca 2+ is involved in this process. These findings allow us to understand the function of AtPiezo from the perspective of plants and provide new insights into the mechanism of plant root cap in response to mechanical stimuli.

At the genus and species level, variation in root anatomy and architecture may interact to affect strategies of drought avoidance. To investigate this idea, root anatomy and architecture of the drought-sensitive common bean (Phaseolus vulgaris) and drought-adapted tepary bean (Phaseolus acutifolius) were analyzed in relation to water use under terminal drought. Intraspecific variation for metaxylem anatomy and axial conductance was found in the roots of both species. Genotypes with high-conductance root metaxylem phenotypes acquired and transpired more water per unit leaf area, shoot mass, and root mass than genotypes with low-conductance metaxylem phenotypes. Interspecific variation in root architecture and root depth was observed where P. acutifolius has a deeper distribution of root length than P. vulgaris. In the deeper-rooted P. acutifolius, genotypes with high root conductance were better able to exploit deep soil water than genotypes with low root axial conductance. Contrastingly, in the shallower-rooted P. vulgaris, genotypes with low root axial conductance had improved water status through conservation of soil moisture for sustained water capture later in the season. These results indicate that metaxylem morphology interacts with root system depth to determine a strategy of drought avoidance and illustrate synergism among architectural and anatomical phenotypes for root function., (© 2020 John Wiley & Sons Ltd.)

This chapter describes a method used to analyze the behavior of histone modifications in S phase in Arabidopsis using a whole-mount immunostaining technique. Previous studies have demonstrated that dramatic changes in local chromatin structure are required for the initiation and progression of DNA replication, and that histone modifications play an essential role in the determination of chromatin structure in S phase. Since euchromatic and heterochromatic regions are replicated in distinct S-phase stages, it is important to identify histone modifications at each stage. Here, we introduce a protocol for whole-mount immunostaining combined with 5-ethynyl-2'-deoxyuridine (EdU) staining, which enables the visualization of spatial patterns in histone modifications in the early and late S-phase nuclei of Arabidopsis roots.

Silicon (Si) can alleviate salt stress by decreasing Na + bypass flow in rice (Oryza sativa L.), however, the mechanisms underpinning remain veiled. In this study, we investigated the roles of OsLsi1 and OsLsi2 in Si-induced reduction of bypass flow and its resultant alleviation of salt stress by using lsi1 and lsi2 mutants (defective in OsLsi1 and OsLsi2, respectively) and their wild types (WTs). Under salt stress, Si promoted plant growth and decreased root-to-shoot Na + translocation in WTs, but not in mutants. Simultaneously, quantitative estimation and fluorescent visualization of trisodium-8-hydroxy-1,3,6-pyrenetrisulphonic (PTS, an apoplastic tracer) showed Si reduced bypass flow in WTs, but not in mutants. Energy-dispersive X-ray microanalysis (EDX) showed Si was deposited at root endodermis in WTs, but not in mutants. Moreover, results obtained from root split experiment using lsi1 WT showed down-regulated expression of Si transport genes (OsLsi1 and OsLsi2) in root accelerated Si deposition at root endodermis. In summary, our results reveal that Si deposition at root endodermis and its resultant reduction of Na + bypass flow is modulated by OsLsi1 and OsLsi2 and regulated by the expression of OsLsi1 and OsLsi2, implying that root Si deposition could be an active and physiologically-regulated process in rice., (Copyright © 2020 Elsevier Masson SAS. All rights reserved.)

Oligogalacturonides (OGs) are fragments of pectin released from the plant cell wall during insect or pathogen attack. They can be perceived by the plant as damage signals, triggering local and systemic defence responses. Here, we analyse the dynamics of local and systemic responses to OG perception in tomato roots or shoots, exploring their impact across the plant and their relevance in pathogen resistance. Targeted and untargeted metabolomics and gene expression analysis in plants treated with purified OGs revealed that local responses were transient, while distal responses were stronger and more sustained. Remarkably, changes were more conspicuous in roots, even upon foliar application of the OGs. The treatments differentially activated the synthesis of defence-related hormones and secondary metabolites including flavonoids, alkaloids and lignans, some of them exclusively synthetized in roots. Finally, the biological relevance of the systemic defence responses activated upon OG perception was confirmed, as the treatment induced systemic resistance to Botrytis cinerea. Overall, this study shows the differential regulation of tomato defences upon OGs perception in roots and shoots and reveals the key role of roots in the coordination of the plant responses to damage sensing., (© 2020 John Wiley & Sons Ltd.)

Arabidopsis thaliana TRY is a negative regulator of trichome differentiation that promotes root hair differentiation. Here, we established that LbTRY, from the recretohalophyte Limonium bicolor, is a typical MYB transcription factor that exhibits transcriptional activation activity and locates in nucleus. By in situ hybridization in L. bicolor, LbTRY may be specifically positioned in salt gland of the expanded leaves. LbTRY expression was the highest in mature leaves and lowest under NaCl treatment. For functional assessment, we heterologously expressed LbTRY in wild-type and try29760 mutant Arabidopsis plants. Epidermal differentiation was remarkably affected in the transgenic wild-type line, as was increased root hair development. Complementation of try29760 with LbTRY under both 35S and LbTRY specific promoter restored the wild-type phenotype. qRT-PCR analysis suggested that AtGL3 and AtZFP5 promote root hair cell fate in lines heterologously producing LbTRY. In addition, four genes (AtRHD6, AtRSL1, AtLRL2, and AtLRL3) involved in root hair initiation and elongation were upregulated in the transgenic lines. Furthermore, LbTRY specifically increased the salt sensitivity of the transgenic lines. The transgenic and complementation lines showed poor germination rates and reduced root lengths, whereas the mutant unexpectedly fared the best under a range of NaCl treatments. Under salt stress, the transgenic seedlings accumulated more MDA and Na + and less proline and soluble sugar than try29760. Thus, when heterologously expressed in Arabidopsis, LbTRY participates in hair development, similar to other MYB proteins, and specifically reduces salt tolerance by increasing ion accumulation and reducing osmolytes. The expression of salt-tolerance marker genes (SOS1, SOS2, SOS3 and P5CS1) was significant reduced in the transgenic lines. More will be carried by downregulating expression of TRY homologs in crops to improve salt tolerance., (Copyright © 2020 The Authors. Published by Elsevier B.V. All rights reserved.)

Superoxide dismutases (SODs) are enzymes detoxifying superoxide to hydrogen peroxide while temporal developmental expression and subcellular localisation are linked to their functions. Therefore, we aimed here to reveal in vivo developmental expression, subcellular, tissue- and organ-specific localisation of iron superoxide dismutase 1 (FSD1) in Arabidopsis using light-sheet and Airyscan confocal microscopy. FSD1-GFP temporarily accumulated at the site of endosperm rupture during seed germination. In emerged roots, it showed the highest abundance in cells of the lateral root cap, columella, and endodermis/cortex initials. The largest subcellular pool of FSD1-GFP was localised in the plastid stroma, while it was also located in the nuclei and cytosol. The majority of the nuclear FSD1-GFP is immobile as revealed by fluorescence recovery after photobleaching. We found that fsd1 knockout mutants exhibit reduced lateral root number and this phenotype was reverted by genetic complementation. Mutant analysis also revealed a requirement for FSD1 in seed germination during salt stress. Salt stress tolerance was coupled with the accumulation of FSD1-GFP in Hechtian strands and superoxide removal. It is likely that the plastidic pool is required for acquiring oxidative stress tolerance in Arabidopsis. This study suggests new developmental and osmoprotective functions of SODs in plants., (© 2020 John Wiley & Sons Ltd.)

Like any other biological tissue, plant tissue also exhibits optical properties like refraction, transmission, absorption, coloration, scattering and so on. Several studies have been conducted using different parts of plants such as leaves, seedlings, roots, stems and so on, and their optical properties have been analyzed to study plant physiology, influence of environmental cues on plant metabolism, light propagation through plant parts and the like. Thus, it is essential to study in detail the optical properties of several plant parts to determine their structural relationship. In this backdrop, an experimental study was conducted to observe and analyze the optical properties of node and inter-nodal tissue cross-sections of the plant Alternanthera philoxeroides under a polarizing microscope constructed and standardized in the laboratory. The observed optical properties of the microscopic tissue sections have been then studied to determine a significant structural relationship between nodal and inter-nodal tissue arrangement patterns as a whole. Tissue sections that have undergone a sort of biological perturbation like loss of water (dried in air for 15 min) have also been studied to study the change in the pattern of tissue optical property when compared with that of normal plant-tissue cross-sections under a polarizing microscope. This type of biological perturbation was chosen for the study because water plays an important role in maintenance of the normal physiological processes in plants and most other forms of life.

As increasing application of nanoparticles, nanoparticles have been becoming a new emerging environmental pollution that attracts a lot of attention from the scientific community and also regulatory agents. In the past decade, studying the toxicity and environmental impacts of nanoparticles is becoming a hot research field and more and more researches have been published using both plant and animal system. In this chapter, using oxidized metal nanoparticles as an example, we introduce a detailed protocol for performing research on biochemical and physiological toxicity of nanoparticles in plant. We employ a hydroponics system to study phytotoxicity of nanoparticles, which makes it easier to study the impact of nanoparticles. In this chapter, we majorly focus on plant respiration and photosynthesis, root vigor as well as oxidative stress. Oxidative stress is one major physiological response to different environmental pollution, in which we present a detailed method for detecting free radical oxygen species as well as the major molecules and enzymes associating with oxidative stress, including SOD and POD. Although we introduce the methods using cotton as an example, the protocols presented in this chapter can be used almost any plant species to test the biochemical and physiological toxicity of an environmental pollution.

Root-knot disease, caused by Meloidogyne spp., alters histology as well as physiology of the roots thus influencing metabolism of vegetative and reproductive parts leading to huge losses in crop productivity. The experimental plant, Vigna unguiculata L. (cowpea of Fabaceae family) var. Gomti is an economically important pulse crop plant. An experiment was conducted to evaluate the effects of different concentrations (0, 25, 50 or 100 ppm) and various modes of applications (root dip, soil drench or foliar spray) of MgO nanoparticles on cowpea infected with M. incognita. The MgO nanoparticles were synthesized chemically and characterized by transmission and scanning electron microscopy (TEM, SEM), UV-Vis spectroscopy, X-ray diffraction (XRD) and Fourier transform infrared spectroscopy (FTIR). The scanning electron microscopy images of second stage juveniles of M. incognita treated with MgO nanoparticles (50 and 100 ppm) exhibited indentations, roughness and distortions in the cuticular surface, in comparison to the control untreated juveniles. MgO nanoparticles, in varying concentrations (50, 100 and 200 ppm), were dispensed into the plants by root dip, soil drench and foliar spray methods and their efficacy was assessed in terms of morphological characteristics, yield parameters and biochemical attributes of M. incognita infected plants. In planta trials revealed that 100 ppm dose of MgO nanoparticles, as root dip application, demonstrated reduced nematode fecundity, decreased number and smaller size of galls; enhanced plant growth, increased chlorophyll, carotenoid, seed protein, and root and shoot nitrogen contents. From these findings it could be inferred that MgO nanoparticles played twin roles, first as a nematicidal agent and the other as growth promotion inducer., (Copyright © 2020 Elsevier Inc. All rights reserved.)

Silicon (Si), a major contributing constituent for plant resistance against abiotic stresses. In spite of this, the detailed mechanisms underlying the potential of Si in mitigating salt toxicity in maize (Zea mays L.) are still poorly understood. The present study deals with the response of Si application on growth, gaseous exchange, ion homeostasis and antioxidant enzyme activities in two maize cultivars (P1574 and Hycorn 11) grown under saline conditions. Salt stress remarkably reduced the plant tissue (roots and shoots) biomass, relative water contents (RWC), membrane stability index (MSI), gaseous exchange characteristics, and antioxidant enzymatic activities i.e., superoxide dismutase (SOD), peroxidase (POD), ascorbate peroxidase (APX) and catalase (CAT). However, salt-induced phytotoxicity increased the plant tissue concentration of malondialdehyde (MDA), hydrogen peroxide (H 2 O 2 ), Na + /K + ionic ratio, Na + translocation (root to shoot), and its uptake. The detrimental effects were more prominent in Hycorn 11 cultivar than the P1574 cultivar at higher salinity level (S2; 160 mM NaCl). The addition of Si alleviated salt toxicity, which was more obvious in P1574 relative to Hycorn 11 as demonstrated by an increasing trend in RWC, MSI, and activities of SOD, POD, APX and CAT. Besides, Si-induced mitigation of salt stress was due to the depreciation in Na + /K + ratio, Na + ion uptake at the surface of maize roots, translocation in plant tissues and thereby significantly reduced Na + ion accumulation. The findings showed a new dimension regarding the beneficial role of Si in maize plants grown under salt toxicity., (Copyright © 2020 Elsevier Masson SAS. All rights reserved.)

Phenotyping root systems provide essential information for plant breeding, particularly aiming for better abiotic stress resistance. Rhizobox systems provide a field-near growth environment for in situ imaging of root systems in soil. A protocol for RGB and hyperspectral imaging of rhizobox-grown plants is presented that enables gathering of root structural (morphology, architecture) as well as functional (water content, decomposition) information. The protocol exemplifies the setup of a root phenotyping platform combining low-cost RGB with advanced short-wave infrared hyperspectral imaging. For both types of imaging approach, the essential steps of an image analysis pipeline are provided to retrieve biological information on breeding-relevant traits from the imaging datasets.

The screening of 862 T-DNA lines was carried out to approach the genetic dissection of indirect adventitious organogenesis in tomato. Several mutants defective in different phases of adventitious organogenesis, namely callus growth (tdc-1), bud differentiation (tdb-1, -2, -3) and shoot-bud development (tds-1) were identified and characterized. The alteration of the TDC-1 gene blocked callus proliferation depending on the composition of growth regulators in the culture medium. Calli from tds-1 explants differentiated buds but did not develop normal shoots. Histological analysis showed that their abnormal development is due to failure in the organization of normal adventitious shoot meristems. Interestingly, tdc-1 and tds-1 mutant plants were indistinguishable from WT ones, indicating that the respective altered genes play specific roles in cell proliferation from explant cut zones (TDC-1 gene) or in the organization of adventitious shoot meristems (TDS-1 gene). Unlike the previous, plants of the three mutants defective in the differentiation of adventitious shoot-buds (tdb-1, -2, -3) showed multiple changes in vegetative and reproductive traits. Cosegregation analyses revealed the existence of an association between the phenotype of the tdb-3 mutant and a T-DNA insert, which led to the discovery that the SlMAPKKK17 gene is involved in the shoot-bud differentiation process., (Copyright © 2020 The Authors. Published by Elsevier B.V. All rights reserved.)

In the context of plant regeneration, in vitro systems to produce embryos are frequently used. In many of these protocols, nonzygotic embryos are initiated that will produce shoot-like structures but may lack a primary root. By increasing the auxin-to-cytokinin ratio in the growth medium, roots are then regenerated in a second step. Therefore, in vitro systems might not or only partially execute a similar developmental program as employed during zygotic embryogenesis. There are, however, in vitro systems that can remarkably mimic zygotic embryogenesis such as Brassica microspore-derived embryos. In this case, the patterning process of these haploid embryos closely follows zygotic embryogenesis and all fundamental tissue types are generated in a rather similar manner. In this review, we discuss the most fundamental molecular events during early zygotic embryogenesis and hope that this brief summary can serve as a reference for studying and developing in vitro embryogenesis systems in the context of doubled haploid production., (© 2021. The Author(s), under exclusive license to Springer Science+Business Media, LLC, part of Springer Nature.)

Symbiosis between soybean (Glycine max) and rhizobia is essential for efficient nitrogen fixation. Rhizobial effectors secreted through the type-III secretion system are key for mediating the interactions between plants and rhizobia, but the molecular mechanism remains largely unknown. Here, our genome-wide association study for nodule number identified G. max Nodule Number Locus 1 (GmNNL1), which encodes a new R protein. GmNNL1 directly interacts with the nodulation outer protein P (NopP) effector from Bradyrhizobium USDA110 to trigger immunity and inhibit nodulation through root hair infection. The insertion of a 179 bp short interspersed nuclear element (SINE)-like transposon into GmNNL1 leads to the loss of function of GmNNL1, enabling bradyrhizobia to successfully nodulate soybeans through the root hair infection route and enhancing nitrogen fixation. Our findings provide important insights into the coevolution of soybean-bradyrhizobia compatibility and offer a way to design new legume-rhizobia interactions for efficient symbiotic nitrogen fixation.

The interaction of plants with complex microbial communities is the result of co-evolution over millions of years and contributed to plant transition and adaptation to land. The ability of plants to be an essential part of complex and highly dynamic ecosystems is dependent on their interaction with diverse microbial communities. Plant microbiota can support, and even enable, the diverse functions of plants and are crucial in sustaining plant fitness under often rapidly changing environments. The composition and diversity of microbiota differs between plant and soil compartments. It indicates that microbial communities in these compartments are not static but are adjusted by the environment as well as inter-microbial and plant-microbe communication. Hormones take a crucial role in contributing to the assembly of plant microbiomes, and plants and microbes often employ the same hormones with completely different intentions. Here, the function of hormones as go-betweens between plants and microbes to influence the shape of plant microbial communities is discussed. The versatility of plant and microbe-derived hormones essentially contributes to the creation of habitats that are the origin of diversity and, thus, multifunctionality of plants, their microbiota and ultimately ecosystems., (© 2020 The Authors. The Plant Journal published by Society for Experimental Biology and John Wiley & Sons Ltd.)

Grassroots have received more attention than the traditional method as soil reinforcement materials, especially the use of vetiver and other vegetation protection methods to treat expansive soil slope, have been tried and applied. To study the influence of grassroots on the strength properties of expansive soil, the laws of vetiver root growth over time and its vertical distribution of root content(δ) were firstly investigated by the experiment of planting vetiver. Then different δ and depth of planted soil were obtained. Simultaneously different δ and water content(ω) of grafted soil were made. With the direct shear test, the shear strength parameters of root-soil with different δ were analyzed. The shear test on root-soil composites with different δ was carried out to compare the strength characteristics of planted and grafted soil. The results showed that the δ of vetiver decreased with the increase of depth, and the δ of each layer increased with the growth period. The δ of 180d was 70.5% higher than that of 90d. The cohesion(c) of root-soil can be increased by more than 97%, and internal friction angle(φ) can be increased by more than 15.4% after 180 days. The c of 90 d vetiver root system can be increased by more than 18%, and the φ can be increased by more than 1.5%. At each depth, the c and φ of composite soil increases with the increase of δ, and the increment of cohesion (Δc) and the increment of internal friction angle (Δφ) increase with the increment of δ. But the increase in the ω will weaken the shear strength parameters of root-soil. Under the condition of the planted root system and grafted root system, the influence degree of δ on strength parameter of root-soil is different, and the law of strength parameters versus δ of grafted soil of 365d is similar to that of planted soil of 90d. And the root reinforcement of grafted soil is weaker than planted soil. Hence the grafted soil can´t accurately reflect the root-soil interaction of the existing root system., Competing Interests: The authors have read the journal’s policy and have the following potential competing interests: RL is a paid employee of China Railway Siyuan Engineering Croup Co., Ltd. JC is a paid employee of CCCC First Highway Fifth Engineering Co., Ltd. JF is a paid employee of China CEC Engineering Corporation. This does not alter our adherence to PLOS ONE policies on sharing data and materials. There are no patents, products in development or marketed products associated with this research to declare.

Plants benefit from associations with a diverse community of root-colonizing microbes. Deciphering the mechanisms underpinning these beneficial services are of interest for improving plant productivity. We report a plant-beneficial interaction between Arabidopsis thaliana and the root microbiota under iron deprivation that is dependent on the secretion of plant-derived coumarins. Disrupting this pathway alters the microbiota and impairs plant growth in iron-limiting soil. Furthermore, the microbiota improves iron-limiting plant performance via a mechanism dependent on plant iron import and secretion of the coumarin fraxetin. This beneficial trait is strain specific yet functionally redundant across phylogenetic lineages of the microbiota. Transcriptomic and elemental analyses revealed that this interaction between commensals and coumarins promotes growth by relieving iron starvation. These results show that coumarins improve plant performance by eliciting microbe-assisted iron nutrition. We propose that the bacterial root microbiota, stimulated by secreted coumarins, is an integral mediator of plant adaptation to iron-limiting soils., Competing Interests: Declaration of Interests The authors declare no competing interests., (Copyright © 2020 The Authors. Published by Elsevier Inc. All rights reserved.)

Shortening the juvenile stage in citrus and inducing early flowering has been the focus of several citrus genetic improvement programs. FLOWERING LOCUS T (FT) is a small phloem-translocated protein that regulates precocious flowering. In this study, two populations of transgenic Carrizo citrange rootstocks expressing either Citrus clementina FT1 or FT3 genes under the control of the Arabidopsis thaliana phloem specific SUCROSE SYNTHASE 2 (AtSUC2) promoter were developed. The transgenic plants were morphologically similar to the non-transgenic controls (non-transgenic Carrizo citrange), however, only AtSUC2-CcFT3 was capable of inducing precocious flowers. The transgenic lines produced flowers 16 months after transformation and flower buds appeared 30-40 days on juvenile immature scions grafted onto transgenic rootstock. Gene expression analysis revealed that the expression of SUPPRESSOR OF OVEREXPRESSION OF CONSTANS 1 (SOC1) and APETALA1 (AP1) were enhanced in the transgenics. Transcriptome profiling of a selected transgenic line showed the induction of genes in different groups including: genes from the flowering induction pathway, APETALA2/ETHYLENE RESPONSE FACTOR (AP2/ERF) family genes, and jasmonic acid (JA) pathway genes. Altogether, our results suggested that ectopic expression of CcFT3 in phloem tissues of Carrizo citrange triggered the expression of several genes to mediate early flowering.

Plant roots determine carbon uptake, survivorship, and agricultural yield and represent a large proportion of the world's vegetation carbon pool. Study of belowground competition, unlike aboveground shoot competition, is hampered by our inability to observe roots. We developed a consumer-resource model based in game theory that predicts the root density spatial distribution of individual plants and tested the model predictions in a greenhouse experiment. Plants in the experiment reacted to neighbors as predicted by the model's evolutionary stable equilibrium, by both overinvesting in nearby roots and reducing their root foraging range. We thereby provide a theoretical foundation for belowground allocation of carbon by vegetation that reconciles seemingly contradictory experimental results such as root segregation and the tragedy of the commons in plant roots., (Copyright © 2020 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works.)

To provide an appropriate tillage fertilization model for improving N utilization efficiency and increasing production, the field experiments were conducted to study the effects on root distributions and N utilization efficiency of summer maize involving different straw mulching modes combined with N fertilization. No (N0), low (N1), medium (N2), and high (N3) levels of N fertilization were incorporated into soil combined with the surface coverage straw (Treatment B) and the deeply buried straw (Treatment S). The traditional cultivation was used as control treatment. The results shown that treatments S had significantly promoted deep root growth, and the root length density (RLD) increased with increases in N application rate. SN2 and SN3 treatments' average RLD were significantly increased by 67.5% and 68.1% in the greater than 40 cm soil layers. While the Treatment B had significantly increased the RLD in 0 -30 cm soil layers only. With increases in N application rate, the effect on summer maize yields increase under Treatment B were not significantly, and only BN3 increased by 0.4%, while under Treatments S were found to first increase, and then decrease. The apparent recovery efficiency of applied N, N uptake and summer maize yield of SN2 had increased by 66.8%, 20.4%, and 9.3%. Therefore the rational tillage fertilization model was deeply buried straw combined with medium N fertilizer in Hetao Irrigation District.

Lateral roots (LRs) are indispensable for plant growth, adaptability and productivity. We previously reported a rice mutant, exhibiting a high density of thick and long LRs (L-type LRs) with long parental roots and herein referred to as promoted lateral root1 (plr1). In this study, we describe that the mutant exhibited decreased basal shoot starch accumulation, suggesting that carbohydrates might regulate the mutant root phenotype. Further analysis revealed that plr1 mutation gene regulated reduced starch accumulation resulting in increased root sugars for the regulation of promoted LR development. This was supported by the exogenous glucose application that promoted L-type LRs. Moreover, nitrogen (N) application was found to reduce basal shoot starch accumulation in both plr1 mutant and wild-type seedlings, which was due to the repressed expression of starch biosynthesis genes. However, unlike the wild-type that responded to N treatment only at seedling stage, the plr1 mutant regulated LR development under low to increasing N levels, both at seedling and higher growth stages. These results suggest that plr1 mutation gene is involved in reduced basal shoot starch accumulation and increased root sugar level for the promotion of L-type LR development, and thus would be very useful in improving rice root architecture., (Copyright © 2020 Elsevier B.V. All rights reserved.)

Salinity is an environmental stress that causes decline in crop yield. Avicennia officinalis and other mangroves have adaptations such as ultrafiltration at the roots aided by apoplastic cell wall barriers to thrive in saline conditions. We studied a cytochrome P450 gene from A. officinalis , AoCYP94B1 , and its putative ortholog in Arabidopsis ( Arabidopsis thaliana ), AtCYP94B1 , which are involved in apoplastic barrier formation. Both genes were induced by 30 min of salt treatment in the roots. Heterologous expression of AoCYP94B1 in the atcyp94b1 Arabidopsis mutant and wild-type rice ( Oryza sativa ) conferred increased NaCl tolerance to seedlings by enhancing root suberin deposition. Histochemical staining and gas chromatography-tandem mass spectrometry quantification of suberin precursors confirmed the role of CYP94B1 in suberin biosynthesis. Using chromatin immunoprecipitation and yeast one-hybrid and luciferase assays, we identified AtWRKY33 as the upstream regulator of AtCYP94B1 in Arabidopsis. In addition, atwrky33 mutants exhibited reduced suberin and salt-sensitive phenotypes, which were rescued by expressing 35S::AtCYP94B1 in the atwrky33 background. This further confirmed that AtWRKY33-mediated regulation of AtCYP94B1 is part of the salt tolerance mechanism. Our findings may help efforts aimed at generating salt-tolerant crops., (© 2020 American Society of Plant Biologists. All Rights Reserved.)

The LATERAL ORGAN BOUNDARIES DOMAIN (LBD)-containing genes are plant-specific genes that play important roles in lateral organ development. In this study, we identified LBD40 (Solyc02g085910), which belongs to subfamily II of the LBD family of genes in tomato. LBD40 was highly expressed in roots and fruit. LBD40 expression was significantly induced by PEG and salt. Moreover, SlLBD40 expression was induced by methyl jasmonate treatment, while SlLBD40 expression could not be induced in the jasmonic acid-insensitive1 (jai1) mutant or MYC2-silenced plants, in which jasmonic acid (JA) signaling was disrupted. These findings demonstrate that SlLBD40 expression was dependent on JA signaling and that it might be downstream of SlMYC2, which is the master transcription factor in the JA signal transduction pathway. Overexpressing and CRISPR/Cas9 mediated knockout transgenic tomato plants were generated to explore SlLBD40 function. The drought tolerance test showed that two SlLBD40 knockout lines wilted slightly, while SlLBD40 overexpressing plants suffered severe wilting. The statistical water loss rate and midday leaf water potential also confirmed that knockout of SlLBD40 improved the water-holding ability of tomato under drought conditions. Taken together, our study demonstrates that SlLBD40, involved in JA signaling, was a negative regulator of drought tolerance and that knockout of SlLBD40 enhanced drought tolerance in tomato. This study also provides a novel function of SlLBD40, which belongs to subfamily II of LBD genes., (Copyright © 2020 Elsevier B.V. All rights reserved.)

As sessile organisms, plants must adjust their growth to withstand several environmental conditions. The root is a crucial organ for plant survival as it is responsible for water and nutrient acquisition from the soil and has high phenotypic plasticity in response to a lack or excess of them. How plants sense and transduce their external conditions to achieve development, is still a matter of investigation and hormones play fundamental roles. Hormones are small molecules essential for plant growth and their function is modulated in response to stress environmental conditions and internal cues to adjust plant development. This review was motivated by the need to explore how Arabidopsis thaliana primary root differentially sense and transduce external conditions to modify its development and how hormone-mediated pathways contribute to achieve it. To accomplish this, we discuss available data of primary root growth phenotype under several hormone loss or gain of function mutants or exogenous application of compounds that affect hormone concentration in several abiotic stress conditions. This review shows how different hormones could promote or inhibit primary root development in A. thaliana depending on their growth in several environmental conditions. Interestingly, the only hormone that always acts as a promoter of primary root development is gibberellins.

This study is the first report on the biosynthesized iron oxide nanoparticles (IONPs) which mediate in-vitro callus induction and shoot regeneration in economically important recalcitrant chickpea crop (Cicer arietinum L.). Here, we used leaf extract of Cymbopogon jwarancusa for the synthesis of IONPs in order to achieve a better biocompatibility. The bioactive compounds in C. jwarancusa leaf extract served as both reducing and capping agents in the fabrication process of IONPs. Field emission scanning electron microscopy (FE-SEM) revealed rods like surface morphology of IONPs with an average diameter of 50±0.2 nm. Energy-dispersive X-ray spectroscopy (EDS) depicted formation of pure IONPs with 69.84% Fe and 30.16% O2. X-ray diffractometry (XRD) and attenuated total reflectance-fourier transform infrared (ATR-FTIR) validate the crystalline structure, chemical analysis detect the presence of various biomolecular fingerprints in the as synthesized IONPs. UV-visible absorption spectroscopy depicts activity of IONPs under visible light. Thermo-gravimetric analysis (TGA) displayed thermal loss of organic capping around 500°C and confirmed their stabilization. The biosynthesized IONPs revealed promising results in callus induction, shoot regeneration and root induction of chickpea plants. Both chickpea varieties Punjab-Noor 09 and Bittle-98 explants, Embryo axes (EA) and Embryo axes plus adjacent part of cotyledon (EXC) demonstrated dose-dependent response. Among all explants, EXC of Punjab-Noor variety showed the highest callogenesis (96%) and shoot regeneration frequency (88%), while root induction frequency was also increased to 83%. Iron content was quantified in regenerated chickpea varieties through inductively coupled plasma-optical emission spectrometry. The quantity of iron is significantly increased in Punjab-Noor regenerated plants (4.88 mg/g) as compare to control treated plants (2.42 mg/g). We found that IONPs enhance chickpea growth pattern and keep regenerated plantlets infection free by providing an optimum environment for rapid growth and development. Thus, IONPs synthesized through green process can be utilized in tissue culture studies in other important recalcitrant legumes crops., Competing Interests: The authors have declared that no competing interests exist.

Rice (Oryza sativa L.) ssp. indica is the most cultivated species in the South of Brazil. However, these plants face low temperature stress from September to November, which is the period of early sowing, affecting plant development during the initial stages of growth, and reducing rice productivity. This study aimed to characterize the root response to low temperature stress during the early vegetative stage of two rice genotypes contrasting in their cold tolerance (CT, cold-tolerant; and CS, cold-sensitive). Root dry weight and length, as well as the number of root hairs, were higher in CT than CS when exposed to cold treatment. Histochemical analyses indicated that roots of CS genotype present higher levels of lipid peroxidation and H 2 O 2 accumulation, along with lower levels of plasma membrane integrity than CT under low temperature stress. RNAseq analyses revealed that the contrasting genotypes present completely different molecular responses to cold stress. The number of over-represented functional categories was lower in CT than CS under cold condition, suggesting that CS genotype is more impacted by low temperature stress than CT. Several genes might contribute to rice cold tolerance, including the ones related with cell wall remodeling, cytoskeleton and growth, signaling, antioxidant system, lipid metabolism, and stress response. On the other hand, high expression of the genes SRC2 (defense), root architecture associated 1 (growth), ACC oxidase, ethylene-responsive transcription factor, and cytokinin-O-glucosyltransferase 2 (hormone-related) seems to be related with cold sensibility. Since these two genotypes have a similar genetic background (sister lines), the differentially expressed genes found here can be considered candidate genes for cold tolerance and could be used in future biotechnological approaches aiming to increase rice tolerance to low temperature., (Copyright © 2020 Elsevier GmbH. All rights reserved.)

In Arabidopsis thaliana , lateral roots initiate in a process preceded by periodic gene expression known as the root clock. We identified the vesicle-trafficking regulator GNOM and its suppressor, ADENOSINE PHOSPHATE RIBOSYLATION FACTOR GTPase ACTIVATION PROTEIN DOMAIN3, as root clock regulators. GNOM is required for the proper distribution of pectin, a mediator of intercellular adhesion, whereas the pectin esterification state is essential for a functional root clock. In sites of lateral root primordia emergence, both esterified and de-esterified pectin variants are differentially distributed. Using a reverse-genetics approach, we show that genes controlling pectin esterification regulate the root clock and lateral root initiation. These results indicate that the balance between esterified and de-esterified pectin states is essential for proper root clock function and the subsequent initiation of lateral root primordia., (Copyright © 2020 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works.)

The phytohormone auxin and microRNA-mediated regulation of gene expressions are key regulators of plant growth and development at both optimal and under low-temperature stress conditions. However, the mechanistic link between microRNA and auxin in regulating plant cold stress response remains elusive. To better understand the role of microRNA (miR) in the crosstalk between auxin and cold stress responses, we took advantage of the mutants of Arabidopsis thaliana with altered response to auxin transport and signal. Screening of the mutants for root growth recovery after cold stress at 4 °C revealed that the auxin signaling mutant, solitary root 1 ( slr1 ; mutation in Aux/IAA14 ), shows a hypersensitive response to cold stress. Genome-wide expression analysis of miRs in the wild-type and slr1 mutant roots using next-generation sequencing revealed 180 known and 71 novel cold-responsive microRNAs. Cold stress also increased the abundance of 26-31 nt small RNA population in slr1 compared with wild type. Comparative analysis of microRNA expression shows significant differential expression of 13 known and 7 novel miRs in slr1 at 4 °C compared with wild type. Target gene expression analysis of the members from one potential candidate miR, miR169, revealed the possible involvement of miR169/ NF-YA module in the Aux/IAA14-mediated cold stress response. Taken together, these results indicate that SLR/IAA14, a transcriptional repressor of auxin signaling, plays a crucial role in integrating miRs in auxin and cold responses.

Background: Chitosan (CTS), a natural polysaccharide, exhibits multiple functions of stress adaptation regulation in plants. However, effects and mechanism of CTS on alleviating salt stress damage are still not fully understood. Objectives of this study were to investigate the function of CTS on improving salt tolerance associated with metabolic balance, polyamine (PAs) accumulation, and Na + transport in creeping bentgrass (Agrostis stolonifera)., Results: CTS pretreatment significantly alleviated declines in relative water content, photosynthesis, photochemical efficiency, and water use efficiency in leaves under salt stress. Exogenous CTS increased endogenous PAs accumulation, antioxidant enzyme (SOD, POD, and CAT) activities, and sucrose accumulation and metabolism through the activation of sucrose synthase and pyruvate kinase activities, and inhibition of invertase activity. The CTS also improved total amino acids, glutamic acid, and γ-aminobutyric acid (GABA) accumulation. In addition, CTS-pretreated plants exhibited significantly higher Na + content in roots and lower Na + accumulation in leaves then untreated plants in response to salt stress. However, CTS had no significant effects on K + /Na + ratio. Importantly, CTS enhanced salt overly sensitive (SOS) pathways and also up-regulated the expression of AsHKT1 and genes (AsNHX4, AsNHX5, and AsNHX6) encoding Na + /H + exchangers under salt stress., Conclusions: The application of CTS increased antioxidant enzyme activities, thereby reducing oxidative damage to roots and leaves. CTS-induced increases in sucrose and GABA accumulation and metabolism played important roles in osmotic adjustment and energy metabolism during salt stress. The CTS also enhanced SOS pathway associated with Na + excretion from cytosol into rhizosphere, increased AsHKT1 expression inhibiting Na + transport to the photosynthetic tissues, and also up-regulated the expression of AsNHX4, AsNHX5, and AsNHX6 promoting the capacity of Na + compartmentalization in roots and leaves under salt stress. In addition, CTS-induced PAs accumulation could be an important regulatory mechanism contributing to enhanced salt tolerance. These findings reveal new functions of CTS on regulating Na + transport, enhancing sugars and amino acids metabolism for osmotic adjustment and energy supply, and increasing PAs accumulation when creeping bentgrass responds to salt stress.

In several cultivation areas, grapevine can suffer from Fe chlorosis due to the calcareous and alkaline nature of soils. This plant species has been described to cope with Fe deficiency by activating Strategy I mechanisms, hence increasing root H + extrusion and ferric-chelate reductase activity. The degree of tolerance exhibited by the rootstocks has been reported to depend on both reactions, but to date, little emphasis has been given to the role played by root exudate extrusion. We studied the behaviour of two hydroponically-grown, tolerant grapevine rootstocks (Ramsey and 140R) in response to Fe deficiency. Under these experimental conditions, the two varieties displayed differences in their ability to modulate morpho-physiological parameters, root acidification and ferric chelate reductase activity. The metabolic profiling of root exudates revealed common strategies for Fe acquisition, including ones targeted at reducing microbial competition for this micronutrient by limiting the exudation of amino acids and sugars and increasing instead that of Fe(III)-reducing compounds. Other modifications in exudate composition hint that the two rootstocks cope with Fe shortage via specific adjustments of their exudation patterns. Furthermore, the presence of 3-hydroxymugenic acid in these compounds suggests that the responses of grapevine to Fe availability are rather diverse and much more complex than those usually described for Strategy I plants.

The mitotic activity of root apical meristem (RAM) is critical to primary root growth and development. Previous studies have identified the roles of ROOT GROWTH FACTOR 1 (RGF1), a peptide ligand, and its receptors, RGF1 INSENSITIVEs (RGIs), a clade of five leucine-rich-repeat receptor-like kinases, in promoting cell division in the RAM, which determines the primary root length. However, the downstream signaling components remain elusive. In this study, we identify a complete mitogen-activated protein kinase (MAPK or MPK) cascade, composed of YDA, MKK4/MKK5, and MPK3/MPK6, that functions downstream of the RGF1-RGI ligand-receptor pair. Similar to the rgi1/2/3/4/5 quintuple mutant, loss-of-function mutants of MPK3 and MPK6, MKK4 and MKK5, or YDA show a short-root phenotype, which is associated with reduced mitotic activity and lower expression of PLETHORA 1 (PLT1)/PLT2 in the RAM. Furthermore, MPK3/MPK6 activation in response to exogenous RGF1 treatment is impaired in the rgi1/2/3/4/5 quintuple, yda single, and mkk4 mkk5 double mutants. Epistatic analyses demonstrated that the expression of constitutively active MKK4, MKK5, or YDA driven by the RGI2 promoter can rescue the short-root phenotype of the rgi1/2/3/4/5 mutant. Taken together, these results suggest that the YDA-MKK4/MKK5-MPK3/MPK6 cascade functions downstream of the RGF1-RGI ligand-receptor pair and upstream of PLT1/PLT2 to modulate the stem cell population and primary root growth in Arabidopsis., (Copyright © 2020 The Authors. Published by Elsevier Inc. All rights reserved.)

Low-molecular-weight organic acid not only serves as a nutrient of plants or as a source of energy metabolism but also a bioactive molecule, which can be linked with hormones, nitrogen (N), and other signals to participate in regulating gene expression, stress response, growth, and development of plants via mechanisms that remain poorly understood. We investigated how the supply of gluconate and sulfate affects rice plant growth and water absorption at the same N supply under osmotic stress. Using a greenhouse hydroponic experimental approach, we assessed the physiology of rice seedlings supplied with C 6 H 11 O 7 NH 4 (AG) and (NH 4 ) 2 SO 4 (AS) over multiple weeks. The root xylem sap rate in rice roots treated with AG increased significantly compared with AS treatment and the control under osmotic stress. The length of roots between 0.5 and 1.5 mm in diameter was obtained after treatment with polyethylene glycol (PEG) solutions containing AG, which was lower than those treated with PEG alone and PEG solutions containing AS. Compared with PEG alone and PEG solutions containing AS, AG induced a significant increase in root lignin under PEG-induced osmotic stress. However, relative to AS supply characteristics, the markedly reduced aerenchyma and porosity of roots, as well as higher root activity, increased fine root tips and length, and higher aquaporins and glutamate synthase (GS) activity in AG supply resulted in increased water uptake under osmotic stress. In addition, AG supply markedly increased leaf area and chlorophyll content. These results suggested that gluconate can enhance the water absorption capacity of the root system by promoting the growth and development of the root system, increasing the activity of aquaporin and GS and reducing the aeration tissue and porosity of the root system under osmotic stress., (Copyright © 2020 Elsevier Masson SAS. All rights reserved.)

Plants counteract Cd toxicity by activating cellular stress responses. The simultaneous exogenous application of methyl jasmonate (MeJA) and methyl-β-cyclodextrin (CD) before Cd exposure improved the response of Arachis hypogaea hairy root culture to the unfavourable effects of Cd toxicity. At 24 h after elicitation, genes that encode key enzymes in the phenylpropanoid biosynthesis pathway (i.e., PAL and RS3) were up-regulated to 3.2- and 5.4-fold changes respectively, thereby inducing stilbene production. The up-regulation of genes that encode transcription factors (i.e., ERF1 and ERF6) significantly increased the expression of several genes (PR4A, PR5, PR10, and chitinase) that encode the pathogenesis-related (PR) proteins to 25.8-, 45-, 5- and 12.6-fold changes, respectively. The more dramatic up-regulation of PR protein-encoding genes demonstrated the significant role of defence proteins in plant protective mechanisms. The prolonged (i.e., 72-h) treatment with MeJA + CD_Cd triggered adaptive responses by substantially increasing the levels of antioxidants, stilbenes, and other phenolic substances. These findings suggest that the interaction between signalling elicitors (MeJA and CD) and Cd modulates a complex signalling network for plant defence system., (Copyright © 2020 Elsevier GmbH. All rights reserved.)

Assessing cell proliferation dynamics is crucial to understand the spatiotemporal control of organogenesis. Here we have generated a versatile fluorescent sensor, PlaCCI (plant cell cycle indicator) on the basis of the expression of CDT1a-CFP, H3.1-mCherry and CYCB1;1-YFP, that identifies cell cycle phases in Arabidopsis thaliana. This tool works in a variety of organs, and all markers and the antibiotic resistance are expressed from a single cassette, facilitating the selection in mutant backgrounds. We also show the robustness of PlaCCI line in live-imaging experiments to follow and quantify cell cycle phase progression.

Recently, a novel type of abiotic stress caused by a prolongation of the light period-coined photoperiod stress-has been described in Arabidopsis. During the night after the prolongation of the light period, stress and cell death marker genes are induced. The next day, strongly stressed plants display a reduced photosynthetic efficiency and leaf cells eventually enter programmed cell death. The phytohormone cytokinin (CK) acts as a negative regulator of this photoperiod stress syndrome. In this study, we show that Arabidopsis wild-type plants increase the CK concentration in response to photoperiod stress. Analysis of cytokinin synthesis and transport mutants revealed that root-derived trans-zeatin (tZ)-type CKs protect against photoperiod stress. The CK signalling proteins ARABIDOPSIS HISTIDINE PHOSPHOTRANSFER PROTEIN 2 (AHP2), AHP3 and AHP5 and transcription factors ARABIDOPSIS RESPONSE REGULATOR 2 (ARR2), ARR10 and ARR12 are required for the protective activity of CK. Analysis of higher order B-type arr mutants suggested that a complex regulatory circuit exists in which the loss of ARR10 or ARR12 can rescue the arr2 phenotype. Together the results revealed the role of root-derived CK acting in the shoot through the two-component signalling system to protect from the negative consequences of strong photoperiod stress., (© 2020 The Authors. Plant, Cell & Environment published by John Wiley & Sons Ltd.)

High levels of ammonium nutrition reduce plant growth and different plant species have developed distinct strategies to maximize ammonium acquisition while alleviating ammonium toxicity through modulating root growth. To date, the mechanisms underlying plant tolerance or sensitivity towards ammonium remain unclear. Rice (Oryza sativa) uses ammonium as its main N source. Here we show that ammonium supply restricts rice root elongation and induces a helical growth pattern, which is attributed to root acidification resulting from ammonium uptake. Ammonium-induced low pH triggers the asymmetric distribution of auxin in rice root tips through changes in auxin signaling, thereby inducing a helical growth response. Blocking auxin signaling completely inhibited this root response. In contrast, this root response is not activated in ammonium-treated Arabidopsis. Acidification of Arabidopsis roots leads to the protonation of indole-3-acetic acid and dampening of the intracellular auxin signaling levels that are required for maintaining root growth. Our study suggests a different mode of action by ammonium on the root pattern and auxin response machinery in rice versus Arabidopsis, and the rice-specific helical root response towards ammonium is an expression of the ability of rice to moderate auxin signaling and root growth to utilize ammonium while confronting acidic stress., (© 2020 Society for Experimental Biology and John Wiley & Sons Ltd.)

Plants are not just passive living beings that exist in nature. They are complex and highly adaptable species that react sensitively to environmental forces/stimuli with movement, morphological changes and through the communication via volatile molecules. In a way, plants mimic some traits of animal and human behaviour; they compete for limited resources by gaining more area for more sunlight and spread their roots underground. Furthermore, in order to survive and thrive, they evolve and 'learn' to control various environmental stress factors in order to increase the yield of flowering, fertilization and germination processes. The concept of associating complex behaviour, such as intelligence, with plants is still a highly debatable topic among researchers worldwide. Recent studies have shown that plants are able to discriminate between positive and negative experiences and 'learn' from them. Some botanists have interpreted these behavioural data as a form of primitive cognitive processes. Others have evaluated these responses as biological automatisms of plants determined by adaptation to the environment and absence of intelligence. This review aims to explore adaptive behavioural aspects of various plant species distributed in different ecosystems by emphasizing their biological complexity and survival instincts., (© 2020 German Society for Plant Sciences and The Royal Botanical Society of the Netherlands.)

Iron (Fe) toxicity is a major nutritional disorder that affects growth and yield in plants. Understanding the responses or damages due to Fe-toxicity may provide useful knowledge to improve tomato varieties. This study investigates the physiological and molecular responses in Fe-toxic tomato plants. The tomato plants were grown in separate hydroponic containers with two concentrations of Fe-EDTA (25 μM and 5 mM) in addition to the other nutrient elements. Fe-toxicity showed a severe reduction in growth parameters, which was accompanied by the increased electrolyte leakage and cell death in tomato. However, the SPAD score, quantum efficiency of PSII, and photosynthesis performance index did not show any changes in leaves, suggesting that damages due to Fe-toxicity are not related to the photosynthetic disturbance in tomato. The FCR (ferric chelate reductase) activity in root along with the Fe concentration in root and shoot significantly increased, being consistent with the upregulation of Fe-related genes (SlNramp1 and SlFRO1) in roots. It suggests that inefficiency to cope with elevated Fe is closely linked to Fe mobilization and uptake in roots of tomato. Consequently, this sensitive genotype was more prone to oxidative damages because of the inefficient antioxidant defense linked to antioxidant enzymes and metabolites. In conclusion, the growth retardation in Fe-toxic tomato is not related to photosynthetic inefficiency but highly associated with oxidative injuries in cells. These findings could be targeted in breeding or transgenic program to improve tomato plants sensitive to Fe toxicity., (Copyright © 2020 Elsevier Masson SAS. All rights reserved.)

Tea plants are important economic perennial crops that can be negatively impacted by drought stress (DS). However, their survival strategies in long-term DS conditions and the accumulation and influence of metabolites and mineral elements (MEs) in their organs, when facing hydraulic vulnerability segmentation, require further investigation. The MEs and metabolites in the leaf, stem, and root after long-term DS (20 d) were examined here, using inductively coupled plasma optical emission spectrometry (ICP-OES) and liquid chromatograph-mass spectrometry (LC-MS). The accumulation patterns of 116 differentially accumulated metabolites (DAMs) and nine MEs were considerably affected in all organs. The concentration of all MEs varied significantly in at least one organ, while the K and Ca levels were markedly altered in all three. Most DAM levels increased in the stem but decreased in the root and leaf, implying that vulnerability segmentation may occur with long-term DS. The typical nitrogen- and carbon-compound levels similarly increased in the stem and decreased in the leaf and root, as the plant might respond to long-term DS by stabilizing respiration, promoting nitrogen recycling, and free radical scavenging. Correlation analysis showed several possible DAM-ME interactions and an association between Mn and flavonoids. Thus, survival strategies under long-term DS included sacrificing distal/vulnerable organs and accumulating function-specialized metabolites and MEs to mitigate drought-induced oxidative damage. This is the first study that reports substance fluctuations after long-term DS in different organs of plants, and highlights the need to use whole plants to fully comprehend stress response strategies., (Copyright © 2020 Elsevier Masson SAS. All rights reserved.)