Casparian Strip Research Articles

In vivo cytological and chemical analysis of Casparian strips using stimulated Raman scattering microscopy

The Casparian strip, a barrier to the apoplastic movement of solutes from the cortex to the stele, is essential for the exclusion of salts, selective nutrient uptake, and many other processes. To date, extensive studies have focused on the physiological functions of endodermal Casparian strips. However, the chemical deposition nature of Casparian strips, as well as its relevance with respect to diffusion barrier functions, remains to be further elucidated. Here, we revealed three developmental stages of Casparian strips in maize primary roots using a traditional fluorescent staining method. Apoplastic permeability tests demonstrated that the barrier function of Casparian strips is largely related to their developmental stage and the pattern of lignin and suberin deposits. Fourier transform infrared (FTIR) analysis showed that the Casparian strips from the roots exhibited significant absorption bands characteristic of lignin and suberin, implying that the Casparian strips in maize primary roots consist largely of lignin and suberin. Furthermore, we developed a new method for label-free, in vivo structural, and biochemical analysis of Casparian strips based on stimulated Raman scattering (SRS) microscopy. Using SRS microscopy, we found that lignin and suberin accumulate simultaneously during the Casparian strip formation process. Based on these results, we propose a potential application of SRS for the chemical composition analysis of plant Casparian strips in situ.

De novo assembly of Phlomis purpurea after challenging with Phytophthora cinnamomi

BackgroundPhlomis plants are a source of biological active substances with potential applications in the control of phytopathogens. Phlomis purpurea (Lamiaceae) is autochthonous of southern Iberian Peninsula and Morocco and was found to be resistant to Phytophthora cinnamomi. Phlomis purpurea has revealed antagonistic effect in the rhizosphere of Quercus suber and Q. ilex against P. cinnamomi. Phlomis purpurea roots produce bioactive compounds exhibiting antitumor and anti-Phytophthora activities with potential to protect susceptible plants. Although these important capacities of P. purpurea have been demonstrated, there is no transcriptomic or genomic information available in public databases that could bring insights on the genes underlying this anti-oomycete activity.ResultsUsing Illumina technology we obtained a de novo assembly of P. purpurea transcriptome and differential transcript abundance to identify putative defence related genes in challenged versus non-challenged plants. A total of 1,272,600,000 reads from 18 cDNA libraries were merged and assembled into 215,739 transcript contigs. BLASTX alignment to Nr NCBI database identified 124,386 unique annotated transcripts (57.7%) with significant hits. Functional annotation identified 83,550 out of 124,386 unique transcripts, which were mapped to 141 pathways.39% of unigenes were assigned GO terms. Their functions cover biological processes, cellular component and molecular functions. Genes associated with response to stimuli, cellular and primary metabolic processes, catalytic and transporter functions were among those identified.Differential transcript abundance analysis using DESeq revealed significant differences among libraries depending on post-challenge times.Comparative cyto-histological studies of P. purpurea roots challenged with P. cinnamomi zoospores and controls revealed specific morphological features (exodermal strips and epi-cuticular layer), that may provide a constitutive efficient barrier against pathogen penetration. Genes involved in cutin biosynthesis and in exodermal Casparian strips formation were up-regulated.ConclusionsThe de novo assembly of transcriptome using short reads for a non-model plant, P. purpurea, revealed many unique transcripts useful for further gene expression, biological function, genomics and functional genomics studies.The data presented suggest a combination of a constitutive resistance and an increased transcriptional response from P. purpurea when challenged with the pathogen. This knowledge opens new perspectives for the understanding of defence responses underlying pathogenic oomycete/plant interaction upon challenge with P. cinnamomi.

Evidence for the Root-Uptake of Arsenite at Lateral Root Junctions and Root Apices in Rice (Oryza sativa L.)

The uptake of arsenite (As(III)i) at the Casparian band via Lsi1 and Lsi2 Si transporters is responsible for ~75% of shoot As(III)i uptake in rice and, therefore, ~25% of shoot As(III)i is taken up by other transport pathways. We hypothesized that areas devoid of Casparian bands—lateral root junctions and root apices—can transport As(III)i into roots. We analyzed the elemental distribution and As concentration, speciation, and localization in rice roots from soil-grown and solution-grown plants. With solution-grown plants dosed with As(III)i, we sectioned roots as a function of distance from the root apex and analyzed the cross-sections using confocal microscopy coupled to synchrotron X-ray fluorescence imaging and spectroscopy. We observed elevated As(III)i associated with lateral root junctions and root apices in rice. As(III)i entered the stele at lateral root junctions and radially permeated the root interior in cross-sections 130–140 µm from the root apex that are devoid of Casparian bands. Our findings suggest that lateral root junctions and rice root apices are hot-spots for As(III)i transport into rice roots, but the contribution to shoot As requires further research.

A Comprehensive Biophysical Model of Ion and Water Transport in Plant Roots. I. Clarifying the Roles of Endodermal Barriers in the Salt Stress Response.

In this paper, we present a detailed and comprehensive mathematical model of active and passive ion and water transport in plant roots. Two key features are the explicit consideration of the separate, but interconnected, apoplastic, and symplastic transport pathways for ions and water, and the inclusion of both active and passive ion transport mechanisms. The model is used to investigate the respective roles of the endodermal Casparian strip and suberin lamellae in the salt stress response of plant roots. While it is thought that these barriers influence different transport pathways, it has proven difficult to distinguish their separate functions experimentally. In particular, the specific role of the suberin lamellae has been unclear. A key finding based on our simulations was that the Casparian strip is essential in preventing excessive uptake of Na+ into the plant via apoplastic bypass, with a barrier efficiency that is reflected by a sharp gradient in the steady-state radial distribution of apoplastic Na+ across the barrier. Even more significantly, this function cannot be replaced by the action of membrane transporters. The simulations also demonstrated that the positive effect of the Casparian strip of controlling Na+ uptake, was somewhat offset by its contribution to the osmotic stress component: a more effective barrier increased the detrimental osmotic stress effect. In contrast, the suberin lamellae were found to play a relatively minor, even non-essential, role in the overall response to salt stress, with the presence of the suberin lamellae resulting in only a slight reduction in Na+ uptake. However, perhaps more significantly, the simulations identified a possible role of suberin lamellae in reducing plant energy requirements by acting as a physical barrier to preventing the passive leakage of Na+ into endodermal cells. The model results suggest that more and particular experimental attention should be paid to the properties of the Casparian strip when assessing the salt tolerance of different plant varieties and species. Indeed, the Casparian strip appears to be a more promising target for plant breeding and plant genetic engineering efforts than the suberin lamellae for the goal of improving salt tolerance.

Casparian bands and suberin lamellae in exodermis of lateral roots: an important trait of roots system response to abiotic stress factors.

Root absorptive characteristics rely on the presence of apoplastic barriers. However, little is known about the establishment of these barriers within a complex root system, particularly in a major portion of them - the lateral roots. In Zea mays L., the exodermis differentiates under the influence of growth conditions. Therefore, the species presents a suitable model to elucidate the cross-talk among environmental conditions, branching pattern and the maturation of barriers within a complex root system involved in the definition of the plant-soil interface. The study describes the extent to which lateral roots differentiate apoplastic barriers in response to changeable environmental conditions. The branching, permeability of the outer cell layers and differentiation of the endo- and exodermis were studied in primary roots and various laterals under different types of stress of agronomic importance (salinity, heavy metal toxicity, hypoxia, etc.). Histochemical methods, image analysis and apoplastic tracer assays were utilized. The results show that the impact of growth conditions on the differentiation of both the endodermis and exodermis is modulated according to the type/diameter of the root. Fine laterals clearly represent that portion of a complex root system with a less advanced state of barrier differentiation, but with substantial ability to modify exodermis differentiation in response to environmental conditions. In addition, some degree of autonomy in exodermal establishment of Casparian bands (CBs) vs. suberin lamellae (SLs) was observed, as the absence of lignified exodermal CBs did not always fit with the lack of SLs. This study highlights the importance of lateral roots, and provides a first look into the developmental variations of apoplastic barriers within a complex root system. It emphasizes that branching and differentiation of barriers in fine laterals may substantially modulate the root system-rhizosphere interaction.

Identification of genes related to skin development in potato.

Newly identified genes that are preferentially expressed in potato skin include genes that are associated with the secondary cell wall and stress-related activities and contribute to the skin's protective function. Microarrays were used to compare the skin and tuber-flesh transcriptomes of potato, to identify genes that contribute to the unique characteristics of the skin as a protective tissue. Functional gene analysis indicated that genes involved in developmental processes such as cell division, cell differentiation, morphogenesis and secondary cell wall formation (lignification and suberization), and stress-related activities, are more highly expressed in the skin than in the tuber flesh. Several genes that were differentially expressed in the skin (as verified by qPCR) and had not been previously identified in potato were selected for further analysis. These included the StKCS20-like, StFAR3, StCYP86A22 and StPOD72-like genes, whose sequences suggest that they may be closely related to known suberin-related genes; the StHAP3 transcription factor that directs meristem-specific expression; and the StCASP1B2-like and StCASP1-like genes, which are two orthologs of a protein family that mediates the formation of Casparian strips in the suberized endodermis of Arabidopsis roots. An examination of microtubers induced from transgenic plants carrying GUS reporter constructs of these genes indicated that these genes were expressed in the skin, with little to no expression in the tuber flesh. Some of the reporter constructs were preferentially expressed in the inner layers of the skin, the root endodermis, the vascular cambium and the epidermis of the stem. Cis-regulatory elements within the respective promoter sequences support this gene-expression pattern.

An ABC Transporter Is Involved in the Silicon-Induced Formation of Casparian Bands in the Exodermis of Rice.

Silicon (Si) promotes the formation of Casparian bands (CB) in rice and reduces radial oxygen loss (ROL). Further transcriptomic approaches revealed several candidate genes involved in the Si-induced formation of CB such as ATP binding cassette (ABC) transporter, Class III peroxidases, ligases and transferases. Investigation of these genes by means of overexpression (OE) and knockout (KO) mutants revealed the contribution of the ABC transporter (OsABCG25) to CB formation in the exodermis, which was also reflected in the expression of other OsABCG25 in the Si-promoted formation of CB genes related to the phenylpropanoid pathway, such as phenylalanine-ammonia-lyase, diacylglycerol O-acyltransferase and 4-coumarate-CoA ligase. Differential CB development in mutants and Si supply also affected the barrier function of the exodermis. OE of the ABC transporter and Si supply reduced the ROL from roots and Fe uptake. No effect on ROL and Fe uptake could be observed for the KO mutant. The presented research confirms the impact of the OsABCG25 in the Si-promoted formation of CB and its barrier functions.

Transient cell-specific EXO70A1 activity in the CASP domain and Casparian strip localization.

In a striking case of evolutionary convergence, polarized cell layers with ring-like diffusion barriers have evolved in both plant and animal lineages independently. In plants, ring-like Casparian strips become localized by the CASPARIAN STRIP MEMBRANE DOMAIN PROTEINS (CASPs). The mechanism of this striking localization, however, has remained enigmatic. Here we present a genetic screen aimed at isolating determinants of CASP localization. One of the mutants, lord of the rings 2 (lotr2)/exo70a1, displays dramatic de-localization of CASPs into randomly localized microdomains. EXO70A1 is a subunit of the exocyst complex, a central component of secretion in eukaryotes. Irradiation of EXO70 subunit genes in plants has suggested specialization of this conserved complex. Intriguingly, lotr2/exo70a1 does neither affect secretion of the CASPs, nor that of other membrane proteins in the endodermis, thus separating exocyst activity in localization from a general defect in secretion. Our results establish EXO70A1 as a central player in Casparian strip formation, generating a transient positional information that will be translated into a precisely localized cell wall modification.

Genome-wide expression profiling in leaves and roots of date palm (Phoenix dactylifera L.) exposed to salinity

BackgroundDate palm, as one of the most important fruit crops in North African and West Asian countries including Oman, is facing serious growth problems due to salinity, arising from persistent use of saline water for irrigation. Although date palm is a relatively salt-tolerant plant species, its adaptive mechanisms to salt stress are largely unknown.ResultsIn order to get an insight into molecular mechanisms of salt tolerance, RNA was profiled in leaves and roots of date palm seedlings subjected to NaCl for 10 days. Under salt stress, photosynthetic parameters were differentially affected; all gas exchange parameters were decreased but the quantum yield of PSII was unaffected while non-photochemical quenching was increased. Analyses of gene expression profiles revealed 2630 and 4687 genes were differentially expressed in leaves and roots, respectively, under salt stress. Of these, 194 genes were identified as commonly responding in both the tissue sources. Gene ontology (GO) analysis in leaves revealed enrichment of transcripts involved in metabolic pathways including photosynthesis, sucrose and starch metabolism, and oxidative phosphorylation, while in roots genes involved in membrane transport, phenylpropanoid biosynthesis, purine, thiamine, and tryptophan metabolism, and casparian strip development were enriched. Differentially expressed genes (DEGs) common to both tissues included the auxin responsive gene, GH3, a putative potassium transporter 8 and vacuolar membrane proton pump.ConclusionsLeaf and root tissues respond differentially to salinity stress and this study has revealed genes and pathways that are associated with responses to elevated NaCl levels and thus may play important roles in salt tolerance providing a foundation for functional characterization of salt stress-responsive genes in the date palm.

Plant Biology: Building Barriers… in Roots

The Casparian strip is an important barrier regulating water and nutrient uptake into root tissues. New research reveals two peptide signals and their co-receptors play critical roles patterning and maintaining barrier integrity.

Role of LOTR1 in Nutrient Transport through Organization of Spatial Distribution of Root Endodermal Barriers

The formation of Casparian strips and suberin lamellae at the endodermis limits the free diffusion of nutrients and harmful substances via the apoplastic space between the soil solution and the stele in roots [1-3]. Casparian strips are ring-like lignin polymers deposited in the middle of anticlinal cell walls between endodermal cells and fill the gap between them [4-6]. Suberin lamellae are glycerolipid polymers covering the endodermal cells and likely function as a barrier to limit transmembrane movement of apoplastic solutes into the endodermal cells [7, 8]. However, the current knowledge on the formation of these two distinct endodermal barriers and their regulatory role in nutrient transport is still limited. Here, we identify an uncharacterized gene, LOTR1, essential for Casparian strip formation in Arabidopsis thaliana. The lotr1 mutants display altered localization of CASP1, an essential protein for Casparian strip formation [9], disrupted Casparian strips, ectopic suberization of endodermal cells, and low accumulation of shoot calcium (Ca). Degradation by expression of a suberin-degrading enzyme in the mutants revealed that the ectopic suberization at the endodermal cells limits Ca transport through the transmembrane pathway, thereby causing reduced Ca delivery to the shoot. Moreover, analysis of the mutants showed that suberin lamellae function as an apoplastic diffusion barrier to the steleat sites of lateral root emergence where Casparian strips are disrupted. Our findings suggest that thetransmembrane pathway through unsuberized endodermal cells, rather than the sites of lateral root emergence, mediates the transport of apoplastic substances such as Ca into the xylem.

Structural features of Phalaris arundinacea in the Jianghan Floodplain of the Yangtze River, China

We examined reed canary grass, Phalaris arundinacea, under shallowly flooded conditions along the Jianghan Floodplain of the Yangtze River to determine structural traits with an emphasis on histochemistry of cell walls in endodermal, exodermal, and epidermal layers and to determine if these traits are consistent with those of other wetland plants of the Floodplain. Roots have an endodermis with Casparian bands, suberin lamellae, and secondarily lignified walls. Outside schizo-lysigenous aerenchyma of the cortex, a biseriate exodermis has Casparian bands and suberin lamellae and lignified walls, and often an internal suberized layer is present. Rhizomes have a thick cuticle, lignified peripheral mechanical ring with an exodermis, large cortical cavities, an endodermis delimiting a central cylinder and sclerenchyma ring, and pith cavities. Culms with leaf sheaths have a thick cuticle and an exodermis in a narrower peripheral mechanical ring, small cortical cavities, a central cylinder with a sclerenchyma ring and pith cavities, but no endodermis. Most vascular bundles are located in the central cylinder and its sclerenchyma ring. These structural features of reed canary grass are characteristic of plants adapted to the seasonal flooding of the Floodplain below the Three Gorges Reservoir Region of the Yangtze River in China.

A peptide hormone required for Casparian strip diffusion barrier formation in Arabidopsis roots.

Plants achieve mineral ion homeostasis by means of a hydrophobic barrier on endodermal cells called the Casparian strip, which restricts lateral diffusion of ions between the root vascular bundles and the soil. We identified a family of sulfated peptides required for contiguous Casparian strip formation in Arabidopsis roots. These peptide hormones, which we named Casparian strip integrity factor 1 (CIF1) and CIF2, are expressed in the root stele and specifically bind the endodermis-expressed leucine-rich repeat receptor kinase GASSHO1 (GSO1)/SCHENGEN3 and its homolog, GSO2. A mutant devoid of CIF peptides is defective in ion homeostasis in the xylem. CIF genes are environmentally responsive. Casparian strip regulation is not merely a passive process driven by root developmental cues; it also serves as an active strategy to cope with adverse soil conditions.

Root diffusion barrier control by a vasculature-derived peptide binding to the SGN3 receptor.

The root endodermis forms its extracellular diffusion barrier by developing ringlike impregnations called Casparian strips. A factor responsible for their establishment is the SCHENGEN3/GASSHO1 (SGN3/GSO1) receptor-like kinase. Its loss of function causes discontinuous Casparian strips. SGN3 also mediates endodermal overlignification of other Casparian strip mutants. Yet, without ligand, SGN3 function remained elusive. Here we report that schengen2 (sgn2) is defective in an enzyme sulfating peptide ligands. On the basis of this observation, we identified two stele-expressed peptides (CASPARIAN STRIP INTEGRITY FACTORS, CIF1/2) that complement sgn2 at nanomolar concentrations and induce Casparian strip mislocalization as well as overlignification-all of which depend on SGN3. Direct peptide binding to recombinant SGN3 identifies these peptides as SGN3 ligands. We speculate that CIF1/2-SGN3 is part of a barrier surveillance system, evolved to guarantee effective sealing of the supracellular Casparian strip network.

Nanoscale measurement of trace element distributions in Spartina alterniflora root tissue during dormancy

This paper reports a nanometer-scale investigation of trace element (As, Ca, Cr, Cu, Fe, Mn, Ni, S and Zn) distributions in the root system Spartina alterniflora during dormancy. The sample was collected on a salt marsh island in Jamaica Bay, New York, in April 2015 and the root was cross-sectioned with 10 μm resolution. Synchrotron X-ray nanofluorescence was applied to map the trace element distributions in selected areas of the root epidermis and endodermis. The sampling resolution was 60 nm to increase the measurement accuracy and reduce the uncertainty. The results indicate that the elemental concentrations in the epidermis, outer endodermis and inner endodermis are significantly (p < 0.01) different. The root endodermis has relatively higher concentrations of these elements than the root epidermis. Furthermore, this high resolution measurement indicates that the elemental concentrations in the outer endodermis are significantly (p < 0.01) higher than those in the inner endodermis. These results suggest that the Casparian strip may play a role in governing the aplastic transport of these elements. Pearson correlation analysis on the average concentrations of each element in the selected areas shows that most of the elements are significantly (p < 0.05) correlated, which suggests that these elements may share the same transport pathways.

The composite water and solute transport of barley (Hordeum vulgare) roots: effect of suberized barriers.

Background and Aims Roots have complex anatomical structures, and certain localized cell layers develop suberized apoplastic barriers. The size and tightness of these barriers depend on the growth conditions and on the age of the root. Such complex anatomical structures result in a composite water and solute transport in roots. Methods Development of apoplastic barriers along barley seminal roots was detected using various staining methods, and the suberin amounts in the apical and basal zones were analysed using gas chromatography–mass spectometry (GC-MS). The hydraulic conductivity of roots (Lpr) and of cortical cells (Lpc) was measured using root and cell pressure probes. Key Results When grown in hydroponics, barley roots did not form an exodermis, even at their basal zones. However, they developed an endodermis. Endodermal Casparian bands first appeared as ‘dots’ as early as at 20 mm from the apex, whereas a patchy suberin lamellae appeared at 60 mm. The endodermal suberin accounted for the total suberin of the roots. The absolute amount in the basal zone was significantly higher than in the apical zone, which was inversely proportional to the Lpr. Comparison of Lpr and Lpc suggested that cell to cell pathways dominate for water transport in roots. However, the calculation of Lpr from Lpc showed that at least 26 % of water transport occurs through the apoplast. Roots had different solute permeabilities (Psr) and reflection coefficients (σsr) for the solutes used. The σsr was below unity for the solutes, which have virtually zero permeability for semi-permeable membranes. Conclusions Suberized endodermis significantly reduces Lpr of seminal roots. The water and solute transport across barley roots is composite in nature and they do not behave like ideal osmometers. The composite transport model should be extended by adding components arranged in series (cortex, endodermis) in addition to the currently included components arranged in parallel (apoplastic, cell to cell pathways).

Impact of TiO2 nanoparticles on lead uptake and bioaccumulation in rice (Oryza sativa L.)

Titanium dioxide nanoparticles (TiO2 NPs) are among the most widely used metal oxide nanoparticles, and their release into the environment is inevitable. The impact of this NPs release on the fate and toxicity of coexisting heavy metals is largely unknown. In this work, a series of hydroponic experiments were conducted to investigate impact of four types of TiO2 NPs, including one anatase (NAnT), one pristine rutile (NRuT) and two rutiles with hydrophilic and hydrophobic surfaces (NLRuT, NBRuT), as well as the bulk particles (BT), on the bioaccumulation of lead (Pb) by rice (Oryza sativa) seedlings. The results indicate that TiO2 NPs exposure at the levels of 10 and 1000mg/L had no significant adverse effects on the growth and development of rice seedlings. Due to its small crystallite size (252.2Å), only one type of TiO2 NPs (NAnT) entered rice seedling roots through an apoplastic route. However, NAnT particles were not translocated from roots to shoots, likely due to the obstruction effects of casparian strip in the root tissues. Both nano and bulk TiO2 reduced the bioaccumulation of Pb in rice at high TiO2 exposure levels (1000mg/L) but the extent of the reduction was TiO2-type dependent. NAnT, NRuT and NLRuT reduced the Pb concentration in rice roots by >80% and by 77–97% in shoots, likely due to the particles high sorption potential for Pb in the nutrient solution. NBRuT and BT reduced Pb levels by 45–61% in roots and 11–38% in shoots. Although NAnT was able to effectively reduce the bioaccumulation of Pb in rice tissues, this particle did accumulate in rice roots, which poses a potential risk to food safety. Accordingly, NAnT is not recommended for Pb pollution control. NRuT and NLRuT are more appropriate agents to reduce Pb contamination threats to rice plants, which in turn may mitigate health risk to humans through the food chain.

Control of Arabidopsis lateral root primordium boundaries by MYB36.

Root branching in plants relies on the de novo formation of lateral roots. These are initiated from founder cells, triggering new formative divisions that generate lateral root primordia (LRP). The LRP size and shape depends on the balance between positive and negative signals that control cell proliferation. The mechanisms controlling proliferation potential of LRP cells remains poorly understood. We found that Arabidopsis thaliana MYB36, which have been previously shown to regulate genes required for Casparian strip formation and the transition from proliferation to differentiation in the primary root, plays a new role in controlling LRP development at later stages. We found that MYB36 is a novel component of LR development at later stages. MYB36 was expressed in the cells surrounding LRP where it controls a set of peroxidase genes, which maintain reactive oxygen species (ROS) balance. This was required to define the transition between proliferating and arrested cells inside the LRP, coinciding with the change from flat to dome-shaped primordia. Reducing the levels of hydrogen peroxide (H2 O2 ) in myb36-5 significantly rescues the mutant phenotype. Our results uncover a role for MYB36 outside the endodermis during LRP development through a mechanism analogous to regulating the proliferation/differentiation transition in the root meristem.

A morel improved growth and suppressed Fusarium infection in sweet corn.

A post-fire morel collected from Populus simonii stands in Mt. Qingling was identified as Morchella crassipes Mes-20 by using nuclear ribosomal DNA internal transcribed spacer phylogeny. It was inoculated into sweet corn to observe colonized roots in purified culture and in greenhouse experiments. The elongation and maturation zones of sweet corn were remarkably colonized at the cortex intercellular and intracellular cells, vessel cells, and around the Casparian strip, forming ectendomycorrhiza-like structures. Colonization was also observed in the zone of cell division proximal to the root cap. Greenhouse assays with sweet corn showed that this morel stimulated the development of the root system and significantly increased the dry root biomass. M. crassipes also significantly reduced the incidence of Fusarium verticillioides in the kernels of mature ears when inoculated into young ears before Fusarium inoculation and prevented Fusarium infection in corn ears compared with that of the control in the greenhouse. When grown under axenic conditions, M. crassipes produced the phytohormones abscisic acid, indole-3-acetic acid, and salicylic acid. The benefits to plants elicited by M. crassipes may result from these phytohormones which may improve the drought resistance, biomass growth and resistance to Fusarium.

The endodermis as a checkpoint for nutrients.

Contents 1604 I. 1604 II. 1604 III. 1605 IV. 1608 V. 1609 1609 References 1609 SUMMARY: Plant roots forage the soil for nutrients and transport them upwards to the aerial parts. Nutrients entering the plant are transported through the concentric layers of epidermis, cortex and endodermis before reaching the central vasculature. The endodermis is the innermost cortical cell layer that surrounds the vasculature. The endodermis forms barriers, the Casparian strips and suberin lamellae, which have been assumed to play a major role in controlling nutrient acquisition. However, the molecular network controlling its differentiation has started to be investigated only recently, giving an unprecedented opportunity to address the role of these barriers in plant nutrition. This insight aims to present recent advances regarding endodermis differentiation, its function as a barrier for nutrients and its developmental plasticity, all pointing to a pivotal role of the endodermis as a checkpoint for nutrients.