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SUMMARY: The mechanistic basis by which boron (B) deprivation inhibits root growth via the mediation of root apical auxin transport and distribution remains elusive. This study showed that B deprivation repressed root growth of wild‐type Arabidopsis seedlings, which was related to higher auxin accumulation (observed with DII‐VENUS and DR5‐GFP lines) in B‐deprived roots. Boron deprivation elevated the auxin content in the root apex, coinciding with upregulation of the expression levels of auxin biosynthesis‐related genes (TAA1, YUC3, YUC9, and NIT1) in shoots, but not in root apices. Phenotyping experiments using auxin transport‐related mutants revealed that the PIN2/3/4 carriers are involved in root growth inhibition caused by B deprivation. B deprivation not only upregulated the transcriptional levels of PIN2/3/4, but also restrained the endocytosis of PIN2/3/4 carriers (observed with PIN‐Dendra2 lines), resulting in elevated protein levels of PIN2/3/4 in the plasma membrane. Overall, these results suggest that B deprivation not only enhances auxin biosynthesis in shoots by elevating the expression levels of auxin biosynthesis‐related genes but also promotes the polar auxin transport from shoots to roots by upregulating the gene expression levels of PIN2/3/4, as well as restraining the endocytosis of PIN2/3/4 carriers, ultimately resulting in auxin accumulation in root apices and root growth inhibition. Significance Statement: While boron (B) deprivation‐induced root growth inhibition is causally associated with auxin accumulation and transport, the underlying mechanisms remain unknown. This work showed that enhanced auxin biosynthesis resulting from upregulation of the expression levels of auxin biosynthesis‐related genes in shoots and increased protein levels of PIN2/3/4 carriers in the plasma membrane mediated by increased transcript levels of PIN2/3/4 genes in roots combined with restrained endocytosis of PIN2/3/4 carriers result in auxin overaccumulation in root apices and consequently root growth inhibition. [ABSTRACT FROM AUTHOR]

SUMMARY: The supply of boron (B) alleviates the toxic effects of aluminum (Al) on root growth; however, the mechanistic basis of this process remains elusive. This study filled this knowledge gap, demonstrating that boron modifies auxin distribution and transport in Al‐exposed Arabidopsis roots. In B‐deprived roots, treatment with Al induced an increase in auxin content in the root apical meristem zone (MZ) and transition zone (TZ), whereas in the elongation zone (EZ) the auxin content was decreased beyond the level required for adequate growth. These distribution patterns are explained by the fact that basipetal auxin transport from the TZ to the EZ was disrupted by Al‐inhibited PIN‐FORMED 2 (PIN2) endocytosis. Experiments involving the modulation of protein biosynthesis by cycloheximide (CHX) and transcriptional regulation by cordycepin (COR) demonstrated that the Al‐induced increase of PIN2 membrane proteins was dependent upon the inhibition of PIN2 endocytosis, rather than on the transcriptional regulation of the PIN2 gene. Experiments reporting on the profiling of Al3+ and PIN2 proteins revealed that the inhibition of endocytosis of PIN2 proteins was the result of Al‐induced limitation of the fluidity of the plasma membrane. The supply of B mediated the turnover of PIN2 endosomes conjugated with indole‐3‐acetic acid (IAA), and thus restored the Al‐induced inhibition of IAA transport through the TZ to the EZ. Overall, the reported results demonstrate that boron supply mediates PIN2 endosome‐based auxin transport to alleviate Al toxicity in plant roots. Significance Statement: Aluminum toxicity and boron deficiency are limiting factors for agricultural production in acid soils, which make up almost half of the world's arable land. Boron fertilization alleviates aluminum toxicity but the mechanism remains unclear. The study shows that boron supply releases aluminum‐induced root growth arrest by facilitating endosome‐based polar auxin transport via PIN2 polar localization in the upper half of the plasma membrane (shootward). We provide solid support for boron fertilizer in remediating aluminum toxicity in acid soils. [ABSTRACT FROM AUTHOR]

Hydrotropism is the directed growth of roots toward the water found in the soil. However, mechanisms governing interactions between hydrotropism and gravitropism remain largely unclear. In this study, we found that an air system and an agar–sorbitol system induced only oblique water-potential gradients; an agar–glycerol system induced only vertical water-potential gradients; and a sand system established both oblique and vertical water-potential gradients. We employed obliquely oriented and vertically oriented experimental systems to study hydrotropism in Arabidopsis and tomato plants. Comparative analyses using different hydrotropic systems showed that gravity hindered the ability of roots to search for obliquely oriented water, whilst facilitating roots' search for vertically oriented water. We found that the gravitropism-deficient mutant aux1 showed enhanced hydrotropism in the oblique orientation but impaired root elongation towards water in the vertical orientation. The miz1 mutant exhibited deficient hydrotropism in the oblique orientation but normal root elongation towards water in the vertical orientation. Importantly, in contrast to miz1 , the miz1 / aux1 double mutant exhibited hydrotropic bending in the oblique orientation and attenuated root elongation towards water in the vertical orientation. Our results suggest that gravitropism is required for MIZ1-regulated root hydrotropism in both the oblique orientation and the vertical orientation, providing further insight into the role of gravity in root hydrotropism. [ABSTRACT FROM AUTHOR]

In plants, growth of roots and root hairs is regulated by the fine cellular control of pH and reactive oxygen species (ROS). MES, 2-(N-morpholino)ethanesulfonic acid as one of the Good's buffers has broadly been used for buffering medium, and it is thought to suit for plant growth with the concentration at 0.1 % (w/v) because the buffer capacity of MES ranging pH 5.5-7.0 (for Arabidopsis, pH 5.8). However, many reports have shown that, in nature, roots require different pH values on the surface of specific root apex zones, namely meristem, transition zone, and elongation zone. Despite the fact that roots always grow on a media containing buffer molecule, little is known about impact of MES on root growth. Here, we have checked the effects of different concentrations of MES buffer using growing roots of Arabidopsis thaliana. Our results show that 1 % of MES significantly inhibited root growth, the number of root hairs and length of meristem, whereas 0.1% promoted root growth and root apex area (region spanning from the root tip up to the transition zone). Furthermore, superoxide generation in root apex disappeared at 1% of MES. These results suggest that MES disturbs normal root morphogenesis by changing the ROS homeostasis in root apex. [ABSTRACT FROM AUTHOR]

Plant 14-3-3 proteins are phosphoserine-binding proteins that regulate a wide array of targets via direct protein-protein interactions. In this study, the role of a 14-3-3 protein, GRF9, in plant response to water stress was investigated. Arabidopsis wild-type, GRF9-deficient mutant (grf9), and GRF9-overexpressing (OE) plants were treated with polyethylene glycol (PEG) to induce mild water stress. OE plant showed better whole-plant growth and root growth than the wild type under normal or water stress conditions while the grf9 mutant showed worse growth. In OE plants, GRF9 favours the allocation of shoot carbon to roots. In addition, GRF9 enhanced proton extrusion, mainly in the root elongation zone and root hair zone, and maintained root growth under mild water stress. Grafting among the wild type, OE, and grf9 plants showed that when OE plants were used as the scion and GRF9 was overexpressed in the shoot, it enhanced sucrose transport into the root, and when OE plants were used as rootstock and GRF9 was overexpressed in the root, it caused more release of protons into the root surface under water stress. Taken together, the results suggest that under PEG-induced water stress, GRF9 is involved in allocating more carbon from the shoot to the root and enhancing proton secretion in the root growing zone, and this process is important for root response to mild water stress. [ABSTRACT FROM AUTHOR]

Root hairs are fast-growing tubular protrusions on root epidermal cells that play important roles in water and nutrient uptake in plants. The tip-focused polarized growth of root hairs is accomplished by the secretion of newly synthesized materials to the tip via the polarized membrane trafficking mechanism. Here, we report the function of two different types of plasma membrane (PM) Qa-SNAREs (soluble N-ethylmaleimide-sensitive factor attachment protein receptors), SYP123 and SYP132, in the growth of root hair in Arabidopsis. We found that SYP123, but not SYP132, localizes in the tip region of root hairs by recycling between the brefeldin A (BFA)-sensitive endosomes and the PM of the expanding tip in an F-actin-dependent manner. The vesicle-associated membrane proteins VAMP721/722/724 also exhibited tip-focused localization in root hairs and formed ternary SNARE complexes with both SYP123 and SYP132. These results demonstrate that SYP123 and SYP132 act in a coordinated fashion to mediate tip-focused membrane trafficking for root hair tip growth. [ABSTRACT FROM AUTHOR]

Gravitropism plays a critical role in plant growth and development, plant stability and acclimation to changes in water and nutrient availability. Ammonium (NH4+) is well known to have profound effects on root growth, but its impacts on gravitropism are poorly understood., To determine which genes are essential for the maintenance of root gravitropism under NH4+ stress, we isolated and identified an NH4+-sensitive mutant, gsa-1 ( gravitropism sensitive to ammonium- 1), in Arabidopsis thaliana, using an agar plate root reorientation assay., We found that, under NH4+ stress, gsa-1 displayed increased loss of root gravitropism. Gene cloning and sequencing revealed that gsa-1 contains a G to C transversion mutation at the highly conserved 5′- GT splice position of intron 10 of ARG1 ( ALTERED RESPONSE TO GRAVITY 1), known to participate in the transduction of the root gravity signal. Genetic complement tests established the locus of GSA-1/ ARG1 and its role in resistance to NH4+ inhibition on root gravitropism. GSA-1/ ARG1 is required for normal AUX1 expression and basipetal auxin transport in root apices. In addition, PIN-FORMED2 ( PIN2) is proposed as a target in the reduction of root gravitropism under NH4+ stress, a response which can be antagonized by the GSA-1/ ARG1-dependent pathway., These results suggest that GSA-1/ ARG1 protects root gravitropism in Arabidopsis thaliana under ammonium stress. [ABSTRACT FROM AUTHOR]

Structural sterols are abundant in the plasma membrane of root apex cells in Arabidopsis thaliana. They specifically accumulate in trichoblasts during the prebulging and bulge stages and show a polar accumulation in the tip during root hair elongation but are distributed evenly in mature root hairs. Thus, structural sterols may serve as a marker for root hair initiation and growth. In addition, they may predict branching events in mutants with branching root hairs. Structural sterols were detected using the sterol complexing fluorochrome filipin. Application of filipin caused a rapid, concentration-dependent decrease in tip growth. Filipin-complexed sterols accumulated in globular structures that fused to larger FM4-64–positive aggregates in the tip, so-called filipin-induced apical compartments, which were closely associated with the plasma membrane. The plasma membrane appeared malformed and the cytoarchitecture of the tip zone was affected. Trans-Golgi network/early endosomal compartments containing molecular markers, such as small Rab GTPase RabA1d and SNARE Wave line 13 (VTI12), locally accumulated in these filipin-induced apical compartments, while late endosomes, endoplasmic reticulum, mitochondria, plastids, and cytosol were excluded from them. These data suggest that the local distribution and apical accumulation of structural sterols may regulate vesicular trafficking and plasma membrane properties during both initiation and tip growth of root hairs in Arabidopsis. [ABSTRACT FROM AUTHOR]

Root hairs are tip-growing tubes that emerge from trichoblasts (hair-forming epidermal cells) along the length of the root. Signalling events involved in the formation of root hairs are largely unknown. However, two recent studies have revealed that signalling enzymes such as NADPH oxidase and phospholipase D are crucial for root hair growth and development. Reactive oxygen species (ROS) produced by NADPH oxidase activate calcium ion channels in the apical plasma membrane leading to the tip-focused calcium gradient, an inherent feature of growing root hairs. [Copyright &y& Elsevier]

Proton (H+) fluxes in plant roots play critical roles in maintaining root growth and facilitating plant responses to multiple soil stresses, including fluctuations in nutrient supply, salt infiltration, and water stress. Soil mining for nutrients and water, rates of nutrient uptake, and the modulation of cell expansion all depend on the regulation of root H+ fluxes, particularly at the root apex, mediated primarily by the activity of plasma membrane (PM) H+-ATPases. Here, we summarize recent findings on the regulatory mechanisms of H+ fluxes at the root apex under three abiotic stress conditions – phosphate deficiency, salinity stress, and water deficiency – and present an integrated physiomolecular view of the functions of H+ fluxes in maintaining root growth in the acclimation to soil stress. Modulation of proton fluxes at the root apex plays roles in nutrient and water acquisition to overcome several types of abiotic stress conditions. Enhanced plasma membrane (PM) H+-ATPase activity in different root apex zones helps to maintain cell elongation, root hair formation, lateral root development, and the secretion of organic acids under phosphate-deficient conditions. Auxin and blue-light signaling pathways are involved in the low-phosphate responses in roots that are tightly linked with regulation of the proton fluxes at the root apex. PKS5- and PIN2-regulated PM H+-ATPase activity at the root apex is essential for salt tolerance and the salt avoidance response. Several signaling components that attribute to drought resistance and hydrotropic response control the proton fluxes at the root apex via ABA- and brassinosteroid-mediated pathways. [ABSTRACT FROM AUTHOR]

Manganese (Mn) is an essential microelement for plant growth but can be toxic when in excessive. Mn toxicity-inhibited root growth possible involves in auxin biosynthesis and transport in plants, but the mechanism remains elusive. Our results have suggested that Mn-inhibited root growth was possible associated with a reduced auxin levels in roots, as shown in experiments with (1) transgenic DII-VENUS and DR5rev::GFP ; (2) the application of L-kynurenine (an inhibitor of TAA1 activity), 1-naphthylacetic acid (NAA), 1-naphthylphthalamic acid (NPA) to manipulate auxin levels; (3) using taa1 (defection in auxin biosynthesis) mutants; and (4) determining IAA by enzyme-linked immunosorbent assay. Those results were explained via down-regulated transcriptional levels of auxin biosynthesis-related genes in plants, and reduced abundance of PM-localized auxin transport carriers in roots. Combination of auxin transport-related transgenic pPINs-GFP or pAUX1::AUX1-YFP lines and a real-time in vitro observation of PM-localized PIN2/3/4 carriers using transgenic Dendra2 lines have revealed that excessive Mn induced degradation of these proteins, which might involve in Mn toxicity-elicited up-regulation of partially ubiquitin-, clathrin-meidiated endocytosis (CME)-, and endosomal sorting complex required for transport (ESCRT)-related genes. Overall, our results suggest that Mn toxicity reduces auxin level in the root apex possible via down-regulation of auxin biosynthesis-related genes and post-translational stimulation of the degradation of auxin transport proteins, leading to root growth inhibition caused by Mn toxicity. [ABSTRACT FROM AUTHOR]