Plants interact with root microbes via chemical signaling, which modulates competence or symbiosis. Although several volatile organic compounds (VOCs) from fungi may affect plant growth and ...development, the signal transduction pathways mediating VOC sensing are not fully understood. 6‐pentyl‐2H‐pyran‐2‐one (6‐PP) is a major VOC biosynthesized by Trichoderma spp. which is probably involved in plant–fungus cross‐kingdom signaling. Using microscopy and confocal imaging, the effects of 6‐PP on root morphogenesis were found to be correlated with DR5:GFP, DR5:VENUS, H2B::GFP, PIN1::PIN1::GFP, PIN2::PIN2::GFP, PIN3::PIN3::GFP and PIN7::PIN7::GFP gene expression. A genetic screen for primary root growth resistance to 6‐PP in wild‐type seedlings and auxin‐ and ethylene‐related mutants allowed identification of genes controlling root architectural responses to this metabolite. Trichoderma atroviride produced 6‐PP, which promoted plant growth and regulated root architecture, inhibiting primary root growth and inducing lateral root formation. 6‐PP modulated expression of PIN auxin‐transport proteins in a specific and dose‐dependent manner in primary roots. TIR1, AFB2 and AFB3 auxin receptors and ARF7 and ARF19 transcription factors influenced the lateral root response to 6‐PP, whereas EIN2 modulated 6‐PP sensing in primary roots. These results indicate that root responses to 6‐PP involve components of auxin transport and signaling and the ethylene‐response modulator EIN2.
Summary
Phosphate (Pi) is a critical macronutrient for the biochemical and molecular functions of cells. Under phosphate limitation, plants manifest adaptative strategies to increase phosphate ...scavenging. However, how low phosphate sensing links to the transcriptional machinery remains unknown.
The role of the MEDIATOR (MED) transcriptional co‐activator, through its MED16 subunit in Arabidopsis root system architecture remodeling in response to phosphate limitation was assessed. Its critical function acting over the SENSITIVE TO PROTON RHIZOTOXICITY1 (STOP1)‐ALUMINUM‐ACTIVATED MALATE TRANSPORT1 (ALMT1) signaling module was tested through a combination of genetic, biochemical, and genome‐wide transcriptomic approaches.
Root system configuration in response to phosphate scarcity involved MED16 functioning, which modulates the expression of a large set of low‐phosphate‐induced genes that respond to local and systemic signals in the Arabidopsis root tip, including those directly activated by STOP1. Biomolecular fluorescence complementation analysis suggests that MED16 is required for the transcriptional activation of STOP1 targets, including the membrane permease ALMT1, to increase malate exudation in response to low phosphate.
Our results unveil the function of a critical transcriptional component, MED16, in the root adaptive responses to a scarce plant macronutrient, which helps understanding how plant cells orchestrate root morphogenesis to gene expression with the STOP1‐ALMT1 module.
Plants adapt to soil injury and biotic stress via cell regeneration. In Arabidopsis, root tip damage by genotoxic agents, antibiotics, UV light and cutting induces a program that recovers the missing ...tissues through activation of stem cells and involves ethylene response factor 115 (ERF115), which triggers cell replenishment. Here, we show that mutation of the gene encoding an MED18 subunit of the transcriptional MEDIATOR complex and chromate Cr(VI), an environmental pollutant, synergistically trigger a developmental program that enables the splitting of the meristem in vivo to produce twin roots. Expression of the quiescent centre gene marker WOX5, auxin‐inducible DR5:GFP reporter and the ERF115 factor traced the changes in cell identity during the conversion of single primary root meristems into twin roots and were induced in an MED18 and chromate‐dependent manner during the root twinning events, which also required auxin redistribution and signalling mediated by IAA14/SOLITARY ROOT (SLR1). Splitting of the root meristem allowed dichotomous root branching in Arabidopsis, a poorly understood process in which stem cells may act to enable whole organ regeneration.
This manuscript describes the twinning of the primary root meristem, also termed root fasciation, in Arabidopsis med18 mutants treated with chromate. Root regeneration takes place through cell re‐specification programs, which correlates with quiescent centre, auxin re‐distribution and cell renewal.
Plants host a diverse microbiome and differentially react to the fungal species living as endophytes or around their roots through emission of volatiles. Here, using divided Petri plates for ...Arabidopsis‐T. atroviride co‐cultivation, we show that fungal volatiles increase endogenous sugar levels in shoots, roots and root exudates, which improve Arabidopsis root growth and branching and strengthen the symbiosis. Tissue‐specific expression of three sucrose phosphate synthase‐encoding genes (AtSPS1F, AtSPS2F and AtSPS3F), and AtSUC2 and SWEET transporters revealed that the gene expression signatures differ from those of the fungal pathogens Fusarium oxysporum and Alternaria alternata and that AtSUC2 is largely repressed either by increasing carbon availability or by perception of the fungal volatile 6‐pentyl‐2H‐pyran‐2‐one. Our data point to Trichoderma volatiles as chemical signatures for sugar biosynthesis and exudation and unveil specific modulation of a critical, long‐distance sucrose transporter in the plant.
T. atroviride volatiles trigger Arabidopsis biomass production and root branching by modulating genes encoding proteins for sugar biosynthesis and transport. A single volatile, namely 6‐pentyl‐2H‐pyran‐2‐one directly influences the major sucrose transporter AtSUC2, which may be critical for the development of the symbiosis.
The DNA damage response avoids mutations into dividing cells. Here, we analysed the role of photoreceptors on the restriction of root growth imposed by genotoxic agents and its relationship with cell ...viability and performance of meristems. Comparison of root growth of Arabidopsis WT, phyA-211, phyB-9, and phyA-211phyB-9 double mutants unveiled a critical role for phytochrome A (PhyA) in protecting roots from genotoxic stress, regeneration and cell replenishment in the meristematic zone. PhyA was located on primary root tips, where it influences genes related to the repair of DNA, including ERF115 and RAD51. Interestingly, phyA-211 mutants treated with zeocin failed to induce the expression of the repressor of cell cycle MYB3R3, which correlated with expression of the mitotic cyclin CycB1, suggesting that PhyA is required for safeguarding the DNA integrity during cell division. Moreover, the growth of the primary roots of PhyA downstream component HY5 and root growth analyses in darkness suggest that cell viability and DNA damage responses within root meristems may act independently from light and photomorphogenesis. These data support novel roles for PhyA as a key player for stem cell niche maintenance and DNA damage responses, which are critical for proper root growth.
Serotonin (5‐hydroxytryptamine) is a bioactive indoleamine with neurotransmitter function in vertebrates, which represents an emerging signaling molecule in plants, playing key roles in the ...development and defense. In this study, the role of reactive oxygen species (ROS) and jasmonic acid (JA)–ethylene (Et) signaling in root developmental alterations induced by serotonin was investigated. An Arabidopsis thaliana mutant defective at the RADICAL‐INDUCED CELL DEATH1 (RCD1) locus was resistant to paraquat‐induced ROS accumulation in primary roots and showed decreased inhibition or root growth in response to serotonin. A suite of JA‐ and Et‐related mutants including coronatine insensitive1, jasmonic acid resistant1 (jar1), etr1, ein2 and ein3 showed tolerance to serotonin in the inhibition of primary root growth and ROS redistribution within the root tip when compared with wild‐type (WT) seedlings. Competence assays between serotonin and AgNO3, a well‐known blocker of Et action, showed that primary root growth in medium supplemented with serotonin was normalized by AgNO3, whereas roots of eto3, an Et overproducer mutant, were oversensitive to serotonin. Comparison of ROS levels in WT, etr1, jar1 and rcd1 primary root tips using the ROS‐specific probe 2′,7′‐dichlorofluorescein diacetate and confocal imaging showed that serotonin inhibition of primary root growth likely occurs independently of its conversion into melatonin. Our results provide compelling evidence that serotonin affects ROS distribution in roots, involving RCD1 and components of the JA–Et signaling pathways.
Aims The aims of this work were to investigate the aluminum (Al) and phosphate (P) interactions in the regulation of root system architecture of Arabidopsis thaliana seedlings and the contribution of ...auxin signaling in primary and lateral root growth in response to Al toxicity. Methods Detailed analyses of root system architecture and cell division were performed in Arabidopsis WT seedlings and in low phosphorus insensitive mutants lpi1-3 and lpr1-1 lpr2-1 in response to Al. Expression studies of P-deficiency regulated phosphate transporter AtPT2 were also conducted. The role of auxin as a mediator of root morphogenetic changes by Al was evaluated by using the auxin-signaling mutants tir1, tir1 afb2 qrb3, and arf7 arf19. Results Al inhibited primary root growth by affecting cell cycle progression and causing differentiation of cells in the root meristem. These effects were reduced in low phosphorus insensitive lpi1-3 and low phosphate resistant lpr1-1 Ipr2-1 Arabidopsis mutants. A1 also activated the expression of the low phosphate-induced P transporter AtPT2 in roots. Lateral root formation by Al decreased in tir1 afb 2 afb 3 while arf7 arf19 mutants were highly resistant to Al in both primary root inhibition and lateral root induction. Conclusions Our results suggest that lateral root formation in response to Al toxicity and P deficiency may involve common signaling mechanisms, while a pathway involving ARF7 and ARF 19 is important for primary root growth inhibition by Al.
Root hairs are epidermal cell extensions that increase the root surface for water and nutrient acquisition. Thus, both the initiation and elongation of root hairs are critical for soil exploration ...and plant adaptation to ever changing growth conditions. Here, we describe the critical roles of two subunits of the Mediator complex, MED12 and MED13, in root hair growth in response to sucrose and abscisic acid, which are tightly linked to abiotic stress resistance. When compared to the WT, med12 and med13 mutants showed increased sensitivity to sucrose and ABA treatments on root meristem and elongation zones that were accompanied with alterations in root hair length and morphology, leading to the isodiametric growth of these structures. The swollen root hair phenotype appeared to be specific, since med8 or med16 mutants did not develop rounded hairs when supplied with 4.8% sucrose. Under standard growth medium, MED12 and MED13 were mainly expressed in root vascular tissues and cotyledons, and their expression was repressed by sucrose or ABA. Interestingly, med12 and med13 mutants manifested exacerbated levels of nitric oxide under normal growth conditions, and upon sucrose supplementation in trichoblast cells, which coincided with root hair deformation. Our results indicate that MED12 and MED13 play non-redundant functions for maintenance of root hair integrity in response to sucrose and ABA and involve nitric oxide as a cellular messenger in Arabidopsis thaliana.
Pyocyanin acts as a virulence factor in Pseudomonas aeruginosa, a plant and animal pathogen. In this study, we evaluated the effect of pyocyanin on growth and development of Arabidopsis seedlings. ...Root inoculation with P. aeruginosa PAO1 strain inhibited primary root growth in wild-type (WT) Arabidopsis seedlings. In contrast, single lasI- and double rhlI-/lasI- mutants of P. aeruginosa defective in pyocyanin production showed decreased root growth inhibition concomitant with an increased phytostimulation. Treatment with pyocyanin modulates root system architecture, inhibiting primary root growth and promoting lateral root and root hair formation without affecting meristem viability or causing cell death. These effects correlated with altered proportions of hydrogen peroxide and superoxide in root tips and with an inhibition of cell division and elongation. Mutant analyses showed that pyocyanin modulation of root growth was likely independent of auxin, cytokinin, and abscisic acid but required ethylene signaling because the Arabidopsis etr1-1, ein2-1, and ein3-1 ethylene-related mutants were less sensitive to pyocyanin-induced root stoppage and reactive oxygen species (ROS) distribution. Our findings suggest that pyocyanin is an important factor modulating the interplay between ROS production and root system architecture by an ethylene-dependent signaling.
Root system architecture is a major determinant of water and nutrient acquisition as well as stress tolerance in plants. The Mediator complex is a conserved multiprotein complex that acts as a ...universal adaptor between transcription factors and the RNA polymerase II. In this article, we characterize possible roles of the MEDIATOR8 (MED8) and MED25 subunits of the plant Mediator complex in the regulation of root system architecture in Arabidopsis (Arabidopsis thaliana). We found that loss-of-function mutations in PHYTOCHROME AND FLOWERING TIME1 (PFT1)/MED25 increase primary and lateral root growth as well as lateral and adventitious root formation. In contrast, PFT1/MED25 overexpression reduces these responses, suggesting that PFT1/MED25 is an important element of meristematic cell proliferation and cell size control in both lateral and primary roots. PFT1/MED25 negatively regulates auxin transport and response gene expression in most parts of the plant, as evidenced by increased and decreased expression of the auxin-related reporters PIN-FORMED1 (PIN1)::PIN1::GFP (for green fluorescent protein), DR5:GFP, DR5:uidA, and BA3:uidA in pft1-2 mutants and in 35S:PFT1 seedlings, respectively. No alterations in endogenous auxin levels could be found in pft1-2 mutants or in 35S:PFT1 -overexpressing seedlings. However, detailed analyses of DR5:GFP and DR5:uidA activity in wildtype, pft1-2, and 35S:PFT1 seedlings in response to indole-3-acetic acid, naphthaleneacetic acid, and the polar auxin transport inhibitor 1-N-naphthylphthalamic acid indicated that PFT1/MED25 principally regulates auxin transport and response. These results provide compelling evidence for a new role for PFT1/MED25 as an important transcriptional regulator of root system architecture through auxin-related mechanisms in Arabidopsis.
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