Enhanced formation of aerenchyma and induction of a barrier to radial oxygen loss in adventitious roots of Zea nicaraguensis contribute to its waterlogging tolerance as compared with maize (Zea mays ssp. mays)

Enhancement of oxygen transport from shoot to root tip by the formation of aerenchyma and also a barrier to radial oxygen loss (ROL) in roots is common in waterlogging‐tolerant plants. Zea nicaraguensis (teosinte), a wild relative of maize (Zea mays ssp. mays), grows in waterlogged soils. We investigated the formation of aerenchyma and ROL barrier induction in roots of Z. nicaraguensis, in comparison with roots of maize (inbred line Mi29), in a pot soil system and in hydroponics. Furthermore, depositions of suberin in the exodermis/hypodermis and lignin in the epidermis of adventitious roots of Z. nicaraguensis and maize grown in aerated or stagnant deoxygenated nutrient solution were studied. Growth of maize was more adversely affected by low oxygen in the root zone (waterlogged soil or stagnant deoxygenated nutrient solution) compared with Z. nicaraguensis. In stagnant deoxygenated solution, Z. nicaraguensis was superior to maize in transporting oxygen from shoot base to root tip due to formation of larger aerenchyma and a stronger barrier to ROL in adventitious roots. The relationships between the ROL barrier formation and suberin and lignin depositions in roots are discussed. The ROL barrier, in addition to aerenchyma, would contribute to the waterlogging tolerance of Z. nicaraguensis.     

Contrasting dynamics of radial O2-loss barrier induction and aerenchyma formation in rice roots of two lengths

Background and Aims

Many wetland species form aerenchyma and a barrier to radial O₂ loss (ROL) in roots. These features enhance internal O₂ diffusion to the root apex. Barrier formation in rice is induced by growth in stagnant solution, but knowledge of the dynamics of barrier induction and early anatomical changes was lacking.

Methods

ROL barrier induction in short and long roots of rice (Oryza sativa L. ‘Nipponbare’) was assessed using cylindrical root-sleeving O₂ electrodes and methylene blue indicator dye for O₂ leakage. Aerenchyma formation was also monitored in root cross-sections. Microstructure of hypodermal/exodermal layers was observed by transmission electron microscopy (TEM).

Key Results

In stagnant medium, barrier to ROL formation commenced in long adventitious roots within a few hours and the barrier was well formed within 24 h. By contrast, barrier formation took longer than 48 h in short roots. The timing of enhancement of aerenchyma formation was the same in short and long roots. Comparison of ROL data and subsequent methylene blue staining determined the apparent ROL threshold for the dye method, and the dye method confirmed that barrier induction was faster for long roots than for short roots. Barrier formation might be related to deposition of new electron-dense materials in the cell walls at the peripheral side of the exodermis. Histochemical staining indicated suberin depositions were enhanced prior to increases in lignin.

Conclusions

As root length affected formation of the barrier to ROL, but not aerenchyma, these two acclimations are differentially regulated in roots of rice. Moreover, ROL barrier induction occurred before histochemically detectable changes in putative suberin and lignin deposits could be seen, whereas TEM showed deposition of new electron-dense materials in exodermal cell walls, so structural changes required for barrier functioning appear to be more subtle than previously described.     

Construction of intraspecific linkage maps, detection of a chromosome inversion, and mapping of QTL for constitutive root aerenchyma formation in the teosinte Zea nicaraguensis

The teosinte Zea nicaraguensis, a wild relative of maize, possesses a flooding tolerance-related trait: the formation of constitutive root aerenchyma under drained (non-flooded) soil conditions. A previous study suggested that the degree of constitutive aerenchyma formation varies within Z. nicaraguensis. The objectives of this study were to construct linkage maps, to determine the marker order in a region of chromosome 4 in which recombination between maize and Z. nicaraguensis is suppressed, and to identify quantitative trait loci (QTL) controlling constitutive root aerenchyma formation in two segregating populations of Z. nicaraguensis. A total of 236 simple sequence repeat (SSR) markers were screened for polymorphism in an S1 population of Z. nicaraguensis. Seventy-one polymorphic SSR markers were assigned to 10 chromosomes, and a linkage map was constructed covering 793.5 cM. In the S1 map, a paracentric inversion was detected on the long arm of chromosome 4; this rearrangement was confirmed in an S1 linkage map of a different Z. nicaraguensis accession. Composite interval mapping analysis in 96 S1 plants revealed QTL for aerenchyma formation on chromosomes 1 (bins 1.06–1.07) and 7 (bin 7.01), explaining 17 and 12% of the total phenotypic variance, respectively. The QTL on chromosome 1 was verified by using 156 S2 plants. Near-isogenic lines exhibiting the presence or absence of the aerenchyma QTL have been developed that should be useful for genetic and physiological analyses of root aerenchyma formation.     

Suberin lamella development in maize seedling roots grown in aerated and stagnant conditions

Hypoxia can stimulate the development of a suberized exodermis in aquatic plants; however, its influence on this aspect of terrestrial root development is sparsely documented. To determine the effects of hypoxia on maize (Zea mays cv. Seneca Horizon) roots, seedlings were grown in vermiculite (VERM), aerated hydroponics (AER), stagnant hydroponics with agar (STAG), or aerated hydroponics with agar (AERAG). The endo- and exodermis were examined for wall modifications. Lateral root emergence and aerenchyma formation were documented qualitatively. The endodermal Casparian band formation was unaffected by treatment. Endodermal and exodermal suberin lamella formation was earliest and most extensive in VERM. Suberization, especially in the exodermis of aerated treatments, was depressed in all hydroponic media. In comparison with AER, STAG exodermal lamellae were increased, but endodermal lamellae were decreased. Since the suberized exodermis forms a barrier to radial oxygen loss from roots to the medium, its stimulation in STAG roots (which also developed extensive aerenchyma) would help retain oxygen in the root. The reduction of endodermal lamellae should facilitate oxygen diffusion into the stele. Clearly, the response to environmental conditions is variable within individual cortical cell layers. Additionally, the observed patterns of lamellae, aerenchyma and lateral root development indicate a tight radial co-ordination of root development.     

Impact of the exodermis on infection of roots byFusarium culmorum

Patterns of infection withFusarium culmorum (W G Smith) Saccardo were observed in seedling roots of barley (Hordeum vulgare L.), wheat (Triticum aestivum L.), maize (Zea mays L.) and asparagus (Asparagus officinalis L). Apical regions of the main roots were not infected. Since penetration into the root occurred several days after inoculation and the roots were growing during the experiment, these regions had apparently not been in existence long enough to be infected. In older regions of barley, wheat and asparagus, hyphae entered through the tips of lateral roots. In barley and wheat, which had not developed any suberin lamellae in their subepidermal layer, infection occurred randomly over the remainder of the root. In maize, the fungus penetrated the epidermis at many sites but did not breach the exodermis in which all cells possessed both Casparian bands and suberin lamellae. Maize roots, therefore, sustained only minimal infections. In asparagus, the fungus grew through the short (passage) cells but never the long cells of the exodermis. In doing so, it penetrated cells possessing Casparian bands but lacking suberin lamellae. The results support the hypothesis that suberin lamellae provide effective barriers to the growth ofF. culmorum hyphae.     

Chemical composition of apoplastic transport barriers in relation to radial hydraulic conductivity of corn roots (Zea mays L.)

The hydraulic conductivity of roots (Lp_r) of 6- to 8-d-old maize seedlings has been related to the chemical composition of apoplastic transport barriers in the endodermis and hypodermis (exodermis), and to the hydraulic conductivity of root cortical cells. Roots were cultivated in two different ways. When grown in aeroponic culture, they developed an exodermis (Casparian band in the hypodermal layer), which was missing in roots from hydroponics. The development of Casparian bands and suberin lamellae was observed by staining with berberin-aniline-blue and Sudan-III. The compositions of suberin and lignin were analyzed quantitatively and qualitatively after depolymerization (BF₃/methanol-transesterification, thioacidolysis) using gas chromatography/mass spectrometry. Root Lp_r was measured using the root pressure probe, and the hydraulic conductivity of cortical cells (Lp) using the cell pressure probe. Roots from the two cultivation methods differed significantly in (i) the Lp_r evaluated from hydrostatic relaxations (factor of 1.5), and (ii) the amounts of lignin and aliphatic suberin in the hypodermal layer of the apical root zone. Aliphatic suberin is thought to be the major reason for the hydrophobic properties of apoplastic barriers and for their relatively low permeability to water. No differences were found in the amounts of suberin in the hypodermal layers of basal root zones and in the endodermal layer. In order to verify that changes in root Lp_r were not caused by changes in hydraulic conductivity at the membrane level, cell Lp was measured as well. No differences were found in the Lp values of cells from roots cultivated by the two different methods. It was concluded that changes in the hydraulic conductivity of the apoplastic rather than of the cell-to-cell path were causing the observed changes in root Lp_r. Received: 17 March 1999 / Accepted: 22 June 1999     

Apoplastic barriers to radial oxygen loss and solute penetration: a chemical and functional comparison of the exodermis of two wetland species, Phragmites australis and Glyceria maxima

Few studies have examined exodermal development in relation to the formation of barriers to both radial oxygen loss (ROL) and solute penetration along growing roots. Here, we report on the structural development, chemical composition and functional properties of the exodermis in two diverse wetland grasses, Glyceria maxima and Phragmites australis. Anatomical features, development, the biochemical composition of exodermal suberin and the penetration of apoplastic tracers and oxygen were examined. Striking interspecific differences in exodermal structure, suberin composition and quantity per unit surface area, and developmental changes along the roots were recorded. Towards the root base, ROL and periodic acid (H(5)IO(6)) penetration were virtually stopped in P. australis; in G. maxima, a tight ROL barrier restricted but did not stop H(5)IO(6) penetration and the exodermis failed to stain with lipidic dyes. Cultivation in stagnant deep hypoxia conditions or oxygenated circulating solution affected the longitudinal pattern of ROL profiles in G. maxima but statistically significant changes in exodermal suberin composition or content were not detected. Interspecific differences in barrier performance were found to be related to hypodermal structure and probably to qualitative as well as quantitative variations in suberin composition and distribution within exodermal cell walls. Implications for root system function are discussed, and it is emphasized that sufficient spatial resolution to identify the effects of developmental changes along roots is crucial for realistic evaluation of exodermal barrier properties.     

Chemical analysis and immunolocalisation of lignin and suberin in endodermal and hypodermal/rhizodermal cell walls of developing maize (Zea mays L.) primary roots

The composition of suberin and lignin in endodermal cell walls (ECWs) and in rhizodermal/hypodermal cell walls (RHCWs) of developing primary maize (Zea mays L.) roots was analysed after depolymerisation of enzymatically isolated cell wall material. Absolute suberin amounts related to root length significantly increased from primary ECWs (Casparian strips) to secondary ECWs (suberin lamella). During further maturation of the endodermis, reaching the final tertiary developmental state characterised by the deposition of lignified secondary cell walls (u-shaped cell wall deposits), suberin amounts remained constant. Absolute amounts of lignin related to root length constantly increased throughout the change from primary to tertiary ECWs. The suberin of Casparian strips contained high amounts of carboxylic and 2-hydroxy acids, and differed substantially from the suberin of secondary and tertiary ECWs, which was dominated by high contents of ω-hydroxycarboxylic and 1,ω-dicarboxylic acids. Furthermore, the chain-length distribution of suberin monomers in primary ECWs ranged from C₁₆ to C₂₄, whereas in secondary and tertiary ECWs a shift towards higher chain lengths (C₁₆ to C₂₈) was observed. The lignin composition of Casparian strips (primary ECWs) showed a high syringyl content and was similar to lignin in secondary cell walls of the tertiary ECWs, whereas lignin in secondary ECWs contained higher amounts of p-hydroxyphenyl units. The suberin and lignin compositions of RHCWs rarely changed with increasing root age. However, compared to the suberin in ECWs, where C₁₆ and C₁₈ were the most prominent chain lengths, the suberin of RHCWs was dominated by the higher chain lengths (C₂₄ and C₂₆). The composition of RHCW lignin was similar to that of secondary-ECW lignin. Using lignin-specific antibodies, lignin epitopes were indeed found to be located in the Casparian strip. Surprisingly, the mature suberin layers of tertiary ECWs contained comparable amounts of lignin-like epitopes. Received: 19 August 1998 / Accepted: 3 February 1999     

A linkage map of maize × teosinte <Emphasis Type="Italic">Zea luxurians</Emphasis> and identification of QTLs controlling root aerenchyma formation

The objectives of this study were to construct a linkage map and identify quantitative trait loci (QTLs) controlling root aerenchyma formation in drained soil conditions using 195 F₂ individuals derived from a cross between maize inbred line B73 × teosinte Zea luxurians. A 107 SSR marker based map covering 1,331 cM across all ten chromosomes was developed. One significant difference between the parents utilized in the study was that under non-flooding conditions, B73 exhibits a minor capacity to develop root aerenchyma, whereas Z. luxurians exhibits a high tendency to form aerenchyma. Linkage analysis indicated segregation distortion regions on chromosomes 2, 4 and 8, and severe recombination suppression on the long arm of chromosome 4. Multiple interval mapping analysis suggests that five QTLs for aerenchyma formation in non-flooding conditions are located on chromosomes 2, 3, 5, 9 and 10, and these explained 36.3% of the total phenotyphic variance. The Z. luxurians alleles in all five QTLs increased the capacity to form aerenchyma and the locations of these QTLs did not overlap those previously identified in the teosinte Z. nicaraguensis. By transferring aerenchyma-forming QTLs from both Z. luxurians and Z. nicaraguensis, it may be possible to pyramid these genes and develop a maize line with exceptional aerenchyma formation and a high level of tolerance to flooding conditions.     

Lead,zinc and iron (Fe2+) tolerances in wetland plants and relation to root anatomy and spatial pattern of ROL

Metal (Pb, Zn and Fe²⁺) tolerances, root anatomy and profile of radial oxygen loss (ROL) along the root (i.e., spatial pattern of ROL) were studied in 10 emergent wetland plants. The species studied could be classified into three groups. Group I included Alternanthera philoxeroides, Beckmannia syzigachne, Oenanthe javanica and Polypogon fugax, with high ROL along the whole length of root (‘partial barrier’ to ROL). Group II included Cyperus flabelliformis, Cyperus malaccensis, Juncus effusus, Leersia hexandra and Panicum paludosum, ROL of which was remarkably high just behind the root apex, but decreased significantly at relatively basal regions (‘tight barrier’ to ROL). Group III consisted of only Neyraudia reynaudiana, with extremely low ROL along the length of root. The results indicated that metal tolerance in wetland plants was related to root anatomy and spatial pattern of ROL. Co-evolution of metal (Fe and Zn) tolerance and flood tolerance possibly developed in wetland plants since species showing a ‘tight barrier’ to ROL (a common trait of flood-tolerant species) in basal root zones had higher Fe and Zn tolerances than those showing a ‘partial barrier’. Root anatomy such as lignin and suberin deposition contributed to a ‘tight barrier’ in root and conferred to exclusion ability in tolerant species.     

Early root growth plasticity in seedlings of three Mediterranean woody species

Using a 141 F₂ population generated from maize inbred B64 × teosinte Zea nicaraguensis cross, quantitative trait loci (QTLs) controlling aerenchyma formation in roots under non-flooding drained soil conditions were identified. Seedlings of Z. nicaraguensis formed clear aerenchyma in the cortex of adventitious roots in non-flooding conditions, whereas the maize inbred line B64 did not. In the F₂ population, the capacity to develop aerenchyma exhibited wide and continuous variation, suggesting the trait was controlled by multiple genes. A linkage map was developed using 85 SSR markers, covering 1,224 cM across all ten chromosomes. Composite interval mapping analysis revealed that four QTLs for aerenchyma formation under non-flooding conditions were located to two regions of chromosome 1 (identified as Qaer1.02-3 and Qaer1.07), chromosome 5 (Qaer5.09) and chromosome 8 (Qaer8.06-7), and these explained 46.5% of the total phenotypic variance. The multiple interval mapping approach identified additional QTLs on chromosomes 1 (Qaer1.01) and 5 (Qaer5.01). Using these results, it may be possible to use SSR markers linked to aerenchyma formation in a marker assisted selection approach to introduce aerenchyma formation in drained soil conditions into maize for the eventual development of flooding tolerant maize hybrids.     

Variation for Root Aerenchyma Formation in Flooded and Non-Flooded Maize and Teosinte Seedlings

Morphological and anatomical factors such as aerenchyma formation in roots and the development of adventitious roots are considered to be amongst the most important developmental characteristics affecting flooding tolerance. In this study we investigated the lengths of adventitious roots and their capacity to form aerenchyma in three- and four-week-old seedlings of two maize (Zea mays ssp. mays, Linn.) inbred accessions, B64 and Na4, and one teosinte, Z. nicaraguensis Iltis & Benz (Poaceae), with and without a flooding treatment. Three weeks after sowing and following a seven day flooding treatment, both maize and teosinte seedlings formed aerenchyma in the cortex of the adventitious roots of the first three nodes. The degree of aerenchyma formation in the three genotypes increased with a second week of flooding treatment. In drained soil, the two maize accessions failed to form aerenchyma. In Z. nicaraguensis, aerenchyma developed in roots located at the first two nodes three weeks after sowing. In the fourth week, aerenchyma developed in roots of the third node, with a subsequent increase in aerenchyma in the second node roots. In a second experiment, we investigated the capacity of aerenchyma to develop in drained soil. An additional three teosinte species and 15 maize inbred lines, among them a set of flooding-tolerant maize lines, were evaluated. Evaluations indicate that accessions of Z. luxurians (Durieu & Asch. Bird) and two maize inbreds, B55 and Mo20W, form aerenchyma when not flooded. These materials may be useful genetic resources for the development of flooding-tolerant maize accessions.     

Suberin lamellae in the hypodermis of maize (Zea mays) roots; development and factors affecting the permeability of hypodermal layers

Abstract The development of suberin lamellae in the hypodermis of Zea mays cv. LG 11 was observed by electron microscopy and the presence of suberin inferred from autoliuorescence and by Sudan black B staining in nodal (adventitious) and primary (seminal) root axes. Suberin lamellae were evident at a distance of 30–50 mm from the tip of roots growing at 20°C and became more prominent with distance from the tip. Both oxygen deficiency and growth at 13°C produced shorter roots in which the hypodermis was suberized closer to the root tip. There were no suberin lamellae in epidermal cells or cortical collenchyma adjacent to the hypodermis. Plasmodesmata were not occluded by the suberin lamellae: there were twice as many of them in the inner tangential hypodermal wall (1,14 μn^?2) as in the junction between the epidermis and hypodermis (0.54 μm^?2). Water uptake by seminal axes (measured by micropotometry) was greater at distances more than 100 mm from the root lip than in the apical zone where the hypodermis was unsuberized. In the more mature zones of roots grown at 13°C rates of water uptake were greater than in roots grown at 20°C even though hypodermal suberization was more marked. Sleeves of epidermal/hypodermal cells (plus some accessory collenchyma) were isolated from the basal 60 mm of nodal axes by enzymatic digestion (drisclase). The roots were either kept totally immersed in culture solution or had the basal 50 mm exposed to moist air above the solution surface. In both treatments the permeabilities to tritiated water and ₈₆Rb were low (circa 10^?5mms^?1) in sleeves isolated from the extreme base. In roots grown totally immersed, however, the permeability of sleeves increased 10 to 50-fold over a distance of 40 mm. In roots exposed to moist air the permeability remained at a low level until the point where the root entered the culture solution and then increased rapidly (> 50-fold in a distance of 8 mm). Growth of roots in oxygen depleted (5% O₂) solutions promoted the development of extensive cortical aerenchymas. These developments were not associated with any reduction in permeability of sleeves isolated from the basal 40 mm of the axis. It was concluded that the presence of suberin lamellae in hypodermal walls does not necessarily indicate low permeability of cells or tissues to water or solutes. The properties of the walls (lamellae?) can be greatly changed by exposure to moist air, perhaps due to increased oxygen availability.     

Development and Structure of the Root Cortex in Caltha palustris L. and Nymphaea odorata Ait.

Structural features of the mature root cortex and its apoplasticpermeability to dyes have been determined for two dicotyledonouswetland plants of differing habitats: Nymphaea odorata, growingrooted in water and mud, and Caltha palustris, growing in temporalwetlands among cattails. In mature roots, movement of the apoplasticdyes, berberine and safranin, into the roots was blocked atthe hypodermis, indicating the presence of an exodermis. A hypodermiswith an exodermis, i.e. Casparian bands in the outermost uniseriatelayer plus suberin lamellae, is present in both species. InN. odorata, hypodermal walls are further modified with cellulosicsecondary walls. Roots of N. odorata and C. palustris have anendodermis with Casparian bands only. A honeycomb aerenchymais produced by differential expansion in N. odorata and includesastrosclereids and diaphragms, while roots of C. palustris haveno aerenchyma, but some irregular lacunae are found in old roots.These aspects of cortex structure are related to an open meristemorganization, with unusual patterns of cell divisions in certainground meristem cells (called semi-regular hexagon cells) ofN. odorata. The correlation between aerenchyma pattern and hypodermalstructure appears to be related to habitat differences.Copyright2000 Annals of Botany Company Caltha palustris, Nymphaea odorata, root development, cortex, endodermis, aerenchyma, exodermis, hypodermis, permeability, wetland plants     

Stagnant deoxygenated growth enhances root suberization and lignifications, but differentially affects water and NaCl permeabilities in rice (Oryza sativa L.) roots

It has been shown that rice roots grown in a stagnant medium develop a tight barrier to radial oxygen loss (ROL), whereas aerated roots do not. This study investigated whether the induction of a barrier to ROL affects water and solute permeabilities. Growth in stagnant medium markedly reduced the root growth rate relative to aerated conditions. Histochemical studies revealed an early deposition of Casparian bands (CBs) and suberin lamellae (SL) in both the endodermis (EN) and exodermis, and accelerated lignification of stagnant roots. The absolute amounts of suberin, lignin and esterified aromatics (coumaric and ferulic acid) in these barriers were significantly higher in stagnant roots. However, correlative permeability studies revealed that early deposition of barriers in stagnant roots failed to reduce hydraulic conductivity (Lp(r) ) below those of aerated roots. In contrast to Lp(r) , the NaCl permeability (P(sr) ) of stagnant roots was markedly lower than that of aerated roots, as indicated by an increased reflection coefficient (σ(sr) ). In stagnant roots, P(sr) decreased by 60%, while σ(sr) increased by 55%. The stagnant medium differentially affected the Lp(r) and P(sr) of roots, which can be explained in terms of the physical properties of the molecules used and the size of the pores in the apoplast.     

Location of the major barriers to water and ion movement in young roots of Zea mays L.

The main barriers to the movement of water and ions in young roots of Zea mays were located by observing the effects of wounding various cell layers of the cortex on the roots' hydraulic conductivities and root pressures. These parameters were measured with a root pressure probe. Injury to the epidermis and cortex caused no significant change in hydraulic conductivity and either no change or a slight decline in root pressure. Injury to a small area of the endodermis did not change the hydraulic conductivity but caused an immediate and substantial drop in root pressure. When large areas of epidermis and cortex were removed (15–38% of total root mass), the endodermis was always injured and root pressure fell. The hydraulic conductance of the root increased but only by a factor of 1.2–2.7. The results indicate that the endodermis is the main barrier to the radial movement of ions but not water. The major barrier to water is the membranes and apoplast of all the living tissue. These conclusions were drawn from experiments in which hydrostatic-pressure differences were used to induce water flows across young maize roots which had an immature exodermis and an endodermis with Casparian bands but no suberin lamellae or secondary walls. The different reactions of water and ions to the endodermis can be explained by the huge difference in the permeability of membranes to these substances. A hydrophobic wall barrier such as the Casparian band should have little effect on the movement of water, which permeates membranes and, perhaps, also the Casparian bands easily. However, hydrophobic wall depositions largely prevent the movement of ions. Several hours after wounding the endodermis, root pressure recovered to some extent in most of the experiments, indicating that the wound in the endodermis had been partially healed.Abbreviations Lp_r hydraulic conductivity of root; T_1/2 = half-time of water exchange between root xylem and external medium This research was supported by a grant from EUROSILVA (project no. 39473C) to E.S., and by a Bilateral Exchange Grant jointly funded by the Deutsche Forschungsgemeinschaft and the Natural Sciences and Engineering Research Council of Canada to C.A.P. We thank Mr. Burkhard Stumpf for his excellent technicial assistance.     

Role of root hydrophobic barriers in salt exclusion of a mangrove plant Avicennia officinalis

Salt exclusion at the roots and salt secretion in the leaves were examined in a mangrove, Avicennia officinalis. The non‐secretor mangrove Bruguiera cylindrica was used for comparative study of hydrophobic barrier formation in the roots. Bypass flow was reduced when seedlings were previously treated with high salt concentration. A biseriate exodermis was detected in the salt‐treated roots, along with an enhanced deposition of hydrophobic barriers in the endodermis. These barriers reduced Na⁺ loading into the xylem, accounting for a 90–95% salt exclusion in A. officinalis. Prominent barriers were found in the roots of B. cylindrica even in the absence of salt treatment. A cytochrome P450 gene that may regulate suberin biosynthesis was up‐regulated within hours of salt treatment in A. officinalis roots and leaves, corresponding with increased suberin deposition. X‐ray microanalysis showed preferential deposition of Na⁺ and Cl^? in the root cortex compared with the stele, suggesting that the endodermis is the primary site of salt exclusion. Enhanced salt secretion and increased suberin deposition surrounding the salt glands were seen in the leaves with salt treatment. Overall, these data show that the deposition of apoplastic barriers increases resistance to bypass flow leading to efficient salt exclusion at the roots in mangroves.     

Chemical Composition of the Epidermal, Hypodermal, Endodermal and Intervening Cortical Cell Walls of Various Plant Roots

A survey of chemical modifications in the cells of the epidermis,hypodermis, cortical parenchyma and endodermis in roots of 27plant species was performed. Cortical parenchyma walls weregenerally free of modifying substances whereas the walls ofthe epidermis, hypodermis and endodermis were usually modifiedby the presence of lipids, phenols, suberin or lignin. In mostcases, wall-modifying components could be detected within 5mm of the root apex. lipids, phenols, suberin, lignin, ferulic acid, root, epidermis, hypodermis, cortex, endodermis, cell wall