大豆初生根凯氏带对铜离子的通透性

采用在根内生成有色铜沉淀的方法研究大豆(Glycine max)初生根凯氏带对铜离子的通透性。用真空泵抽取浓度为200 μmol·L^–1 的CuSO₄溶液进入根中, 然后在重力作用下从根基部灌注400 μmol·L^–1的K₄[Fe(CN)₆]溶液, 两种物质在根内相遇即可产生棕色的Cu₂[Fe(CN)₆]沉淀, 根据沉淀的位置来确定铜离子所经过的途径。结果表明: Cu²⁺可以穿过内皮层凯氏带, 在木质部导管壁以及凯氏带至木质部之间的细胞壁处产生棕色沉淀, 侧根发生的部位也产生了大量的沉淀; 当抽取K₄[Fe(CN)₆]溶液后再灌注CuSO₄溶液, 发现Cu²⁺仍然可以穿过凯氏带, 并在凯氏带外侧以及外皮层细胞的细胞壁处产生棕色沉淀。研究结果证明凯氏带并不是一个可以完全阻止离子进出的完美屏障。     

大豆初生根凯氏带对铜离子的通透性

采用在根内生成有色铜沉淀的方法研究大豆（Glycine max）初生根凯氏带对铜离子的通透性。用真空泵抽取浓度为200μmol·L^–1的CuSO4溶液进入根中,然后在重力作用下从根基部灌注400μmol·L^–1的K4[Fe（CN）6]溶液,两种物质在根内相遇即可产生棕色的Cu2[Fe（CN）6]沉淀,根据沉淀的位置来确定铜离子所经过的途径。结果表明：Cu^2＋可以穿过内皮层凯氏带,在木质部导管壁以及凯氏带至木质部之间的细胞壁处产生棕色沉淀,侧根发生的部位也产生了大量的沉淀;当抽取K4[Fe（CN）6]溶液后再灌注CuSO4溶液,发现Cu^2＋仍然可以穿过凯氏带,并在凯氏带外侧以及外皮层细胞的细胞壁处产生棕色沉淀。研究结果证明凯氏带并不是一个可以完全阻止离子进出的完美屏障。     

Permeability of Plant Young Root Endodermis to Cu Ions and Cu-Citrate Complexes in Corn and Soybean

The non-selective apoplastic passage of Cu and Cu-citrate complexes into the root stele of monocotyledonous corn and dicotyledonous soybean was investigated using an inorganic-salt-precipitation technique. Either Cu ions or Cu-citrate complexes were drawn into root through the apoplast from the root growth medium, and K₄[Fe(CN)₆] was subsequently perfused through xylem vessels or the entire root cross section. Based on microscopic identification of the reddish-brown precipitates of copper ferrocyanide in the cell walls of the xylem of corn and soybean roots, Cu²⁺ passed through the endodermal barrier into the xylem of both species. When the solution containing 200 μM CuSO₄ and 400 μM sodium citrate (containing 199.98 μM Cu-citrate, 0.02 μM Cu²⁺) was drawn via differential pressure gradients into the root xylem while being perfused with K₄[Fe(CN)₆] through the entire root cross-section, reddish-brown precipitates were observed in the walls of the stele of soybean, but not corn root. However, when a CuSO₄ solution containing 0.02 or 0.2 μM free Cu²⁺ was used, no reddish-brown precipitates were detected in the stele of either of the two plants. Results indicated that endodermis was permeable to Cu-citrate complexes in primary roots of soybean, but not corn. The permeability of the endodermal barrier to the Cu-citrate complex may vary between dicotyledonous and monocotyledonous plants, which has considerable implications for chelant-enhanced phytoextraction.     

Blockage of apoplastic bypass-flow of water in rice roots by insoluble salt precipitates analogous to a Pfeffer cell

Precipitates of insoluble inorganic salts were used to clog apoplastic pores in cell walls of the outer part of rice roots (OPR) in two rice cultivars (lowland cv. IR64 and upland cv. Azucena). Aerenchyma of two different root zones (20–50 and 50–100 mm from the apex) was perfused with 1 m m potassium ferrocyanide (K₄[Fe(CN)₆]) while the whole root segments were bathed in 0.5 m m copper sulphate (CuSO₄) medium. In another experiment, salts were applied on opposite sides of the OPR. The copper-ferrocyanide precipitation technique resembles the famous osmotic experiments of the German botanist Wilhelm Pfeffer, in which he used them with clay diaphragms. Precipitates were observed on the side where ferrocyanide was applied, suggesting that Cu²⁺ and SO₄^2– were passing the barrier including the Casparian bands of the exodermis much faster than ferrocyanide. There was a patchiness in the formation of precipitates, correlated with the maturation of the exodermis. The intensity of copper ferrocyanide staining decreased along developing rice roots. No precipitates were observed in mature parts beyond 70–80 mm from the root apex, except for sites around the emergence of secondary roots, which were fairly leaky to both water and ions. Blockage of the apoplastic pores with precipitates caused a three- to four-fold reduction of hydraulic conductivity of the OPR (Lp_OPR). The reflection coefficient of the OPR (σ_sOPR) increased in response to the blockage with precipitates. The osmotic versus diffusive water permeability ratios of the OPR (P_fOPR/P_dOPR) were around 600 for immature and 1200 for mature root segments. Treatment significantly affected the bulk rather than the diffusive water flow and caused a three- to five-fold reduction of the P_fOPR/P_dOPR ratios. Results indicated that despite the existence of an exodermis with Casparian bands, most of the water moved around cells rather than using the cell-to-cell passage.     

Structural Changes and Associated Reduction of Hydraulic Conductance in Roots of Sorghum bicolor L. following Exposure to Water Deficit

The effects of a severe water deficit on total root (L_t) and axial (L_x) hydraulic conductances and on the development of the hypodermis, endodermis, and xylem were studied in sorghum (Sorghum bicolor L.). Water deficit was imposed in the upper rooting zone while the lower zones were kept moist. L_t and L_x were based on water flow rates obtained by applying suction to proximal xylem ends of excised roots. The development of the hypodermis, endodermis, and other tissues were examined by staining with fluorescent berberine hemisulfate and phloroglucinol-HCl. The L_t value (x 10⁻⁸ meters per second per megapascal) for unstressed control roots was 22.0 and only 5.9 for stressed roots. The low L_t in stressed roots was attributed, in part, to accelerated deposition of lignin and suberin in the hypodermis and endodermis. Calcofluor, an apoplastic tracer that binds to cellulose, was blocked in stressed roots at the lignified and suberized outer tangential walls of the hypodermis but readily penetrated the cortical walls of similar root regions in controls where the casparian band was not developed. L_x per unit root length was about 100 times lower in stressed roots than in controls because of the persistence of late metaxylem cross-walls and the smaller diameter and lower number of conductive protoxylem and early metaxylem vessels.     

Net sodium fluxes change significantly at anatomically distinct root zones of rice (Oryza sativa L.) seedlings

Casparian bands of endodermis and exodermis play crucial roles in blocking apoplastic movement of ions and water into the stele of roots through the cortex. These apoplastic barriers differ considerably in structure and function along the developing root. The present study assessed net Na⁺ fluxes in anatomically distinct root zones of rice seedlings and analyzed parts of individual roots showing different Na⁺ uptake. The results indicated that anatomically distinct root zones contributed differently to the overall uptake of Na⁺. The average Na⁺ uptake in root zones in which Casparian bands of the endo- and exo-dermis were interrupted by initiating lateral root primordia (root zone III) was significantly greater than that at the root apex, where Casparian bands were not yet formed (root zone I), or in the region where endo- and exo-dermis with Casparian bands were well developed (root zone II). The measurement of net Na⁺ fluxes using a non-invasive scanning ion-selective electrode technique (SIET) demonstrated that net Na⁺ flux varied significantly in different positions along developing rice roots, and a net Na⁺ influx was obvious at the base of young lateral root primordia. Since sodium fluxes changed significantly along developing roots of rice seedlings, we suggest that the significantly distinct net Na⁺ flux profile may be attributed to different apoplastic permeability due to lateral root primordia development for non-selective apoplastic bypass of ions along the apoplast.     

Branch Roots of Zea. V. Structural Features that may Influence Water and Nutrient Transport

First-order branch roots of field-grown Zea mays L. were examined by optical and electron microscopy. They were small-scale versions of nodal roots except for the usual retention of a live epidermis throughout their length. The Casparian strips and suberized lamellae of hypodermis and endodermis developed closer to the root tip than reported for main roots (in the zone 0.5 to 5.5 cm from the tip for the hypodermis, and 0.5 to 4 cm for the endodermis), in branches retaining an apical meristem. The hydrophobic deposits were in place to the distal ends of determinate branches. All hydrophobic deposits were fully formed before the late metaxylem elements were mature. Gaps in the suberized lamellae of both hypodermis and endodermis may permit apoplastic diffusion of solutes through these layers. Pit frequency in the outer tangential walls of the hypodermis and endodermis was 0.3 per 100 μm², and 0.6 to 0.7 per 100 μm², respectively, in both branch and main roots. Numbers of plasmodesmata per pit in the branches were 60 and 30 in the hypodermis and endodermis, respectively. Water fluxes from published data were used to calculated the possible flux through plasmodesmata on a symplastic path. Values up to 0.2 pl h^?1 for the hypodermis and twice this for the endodermis were obtained.     

Analytical electron microscopical investigations on the apoplastic pathways of lanthanum transport in barley roots

 In transmission electron microscopy studies, lanthanum ions have been used as electron-opaque tracers to delineate the apoplastic pathways for ion transport in barley (Hordeum vulgare L.) roots. To localize La³⁺ on the subcellular level, e.g. in cell walls and on the surface of membranes, electron-energy-loss spectroscopy and electron-spectroscopic imaging were used. Seminal and nodal roots were exposed for 30 min to 1 mM LaCl₃ and 10 mM LaCl₃, respectively. In seminal roots, possessing no exodermis, La³⁺ diffusion through the apoplast was stopped by the Casparian bands of the endodermis. In nodal roots with an exodermis, however, La³⁺ diffusion through the cortical apoplast had already stopped at the tight junctions of the exodermal cell walls resembling the Casparian bands of the endodermis. Therefore, we conclude that in some specialized roots such as the nodal roots of barley, the physiological role of the endodermis is largely performed by the exodermis. Received: 28 July 1999 / Accepted: 24 February 2000     

OBSERVATIONS ON THE ROOT ANATOMY OF RICE (ORYZA SATIVA L.)

Rice plants were grown hydroponically and roots were prepared for light and electron microscopy using standard techniques. The roots are bounded by an epidermis, exodermis, and fibrous layer. The exodermis has a suberin lamella along its inner tangential wall. The fibrous layer is composed of thick-walled lignified cells with little pitting. The cortical parenchyma is compact when young, but expands and separates to form a zone of cell walls and air spaces in a spoked arrangement. Supporting columns of living parenchyma cells are occasionally present, particularly near lateral roots. The endodermis is typical for grasses with Casparian strips, suberin lamellae, and tertiary state walls with numerous pits. The pericycle and pith become sclerified. Protoxylem elements alternate with protophloem in the young root; later, early metaxylem, late metaxylem, and metaphloem proliferate. The exodermis, fibrous layer, lacunate cortex, and endodermis appear to present a formidable barrier to radial ion movement in the mature portions of the root.     

菰适应湿地环境的解剖和屏障结构特征研究

菰(Zizania latifolia)是一种多年生挺水植物,为了探讨该植物根、茎和叶的解剖结构、组织化学及其质外体屏障的通透性生理。该文利用光学显微镜和荧光显微镜,对菰的根、茎、叶进行了解剖学和组织化学研究。结果表明:(1)菰不定根解剖结构由外而内分别为表皮、外皮层、单层细胞的厚壁机械组织层、皮层、内皮层和维管柱;茎结构由外而内分别为角质层、表皮、周缘厚壁机械组织层、皮层、具维管束的厚壁组织层和髓腔。叶鞘具有表皮和具维管束皮层,叶片具有表皮,叶肉和维管束。(2)不定根具有位于内侧的内皮层及其邻近栓质化细胞和外侧的外皮层组成的屏障结构;茎具内侧厚壁机械组织层,外侧的角质层和周缘厚壁机械组织层组成的屏障结构,屏障结构的细胞壁具凯氏带、木栓质和木质素沉积的组织化学特点,叶表面具有角质层。(3)菰通气组织包括根中通气组织,茎、叶皮层的通气组织和髓腔。(4)菰的屏障结构和解剖结构是其适应湿地环境的重要特征,但其茎周缘厚壁层和厚壁组织层较薄。由此推测,菰适应湿地环境,但在旱生环境中分布有一定的局限性。     

On the evidence of a diffusion barrier in the outer cortex apoplast of cress-roots (Lepidium sativum), demonstrated by analytical electron microscopy

To mark the apoplastic pathway of ions in the root of the dicotyledonous plant Lepidium sativum we used the heavy element lanthanum, which can be identified by analytical electron microscopy (EELS and ESI). In the front root tip, the primary walls of all meristematic cells contained lanthanum. 10-15 mm behind the root apex, lanthanum was found in the cortex cell walls up to the endodermis, but not in the stele. 20-25 mm from the tip, lanthanum was accumulated in the radial cell walls of the hypodermis, which, however, is not a complete diffusion barrier for ions, so that traces of lanthanum also were found in the cortex cell walls up to the endodermis. This study provides evidence for the presence of two apolastic diffusion barriers in the region of highest water uptake in cress roots.     

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     

In vivo and in vitro effects of copper and cadmium on the plasma membrane H+-ATPase from cucumber (Cucumis sativus L.) and maize (Zea mays L.) roots

The plasmalemma vesicles isolated from cucumber and maize roots were used to study the effect of Cu²⁺ and Cd²⁺ on the hydrolytic and proton pumping activities of ATPase. In vivo application of metal ions to the plant growth solutions resulted in stimulation of the proton transport in maize. In cucumber roots the action of metals was not the same: cadmium stimulated the H⁺ transport through plasmalemma whereas Cu²⁺ almost completely inhibited it. Copper ions decreased the hydrolytic activity of H⁺-ATPase in cucumber, without any effect on this activity in membranes isolated from maize roots. The effect of cadmium on the hydrolytic activities was opposite: ATP-hydrolysis activity in plasmalemma was not altered in cucumber, whereas in maize its stimulation was observed. The amount of accumulated metals was not the main reason of different influence of metals on H⁺-ATPase activity in tested plants. In in vitro experiments Cu²⁺ inhibited H⁺ transport in the cucumber, to a higher degree than Cd²⁺ and both metals did not change this H⁺-ATPase activity of plasmalemma isolated from corn roots. Cu²⁺ added into the incubation medium reduced the hydrolytic activity of ATPase in the plasma membrane isolated from cucumber as well as from corn roots. Cd²⁺ diminished the hydrolytic activity of ATPase in cucumber, and no effect of Cd²⁺ in the plasmalemma isolated from corn roots was found. Our results indicated different in vitro and in vivo action of both metals on H⁺-ATPase and different response of this enzyme to Cu²⁺ and Cd²⁺ in maize and cucumber.     

Root Browning in Pinus banksiana Lamb. and Eucalyptus pilularis Sm. 1. Anatomy and Permeability of the White and Tannin Zones

Growing tree roots are characteristically brown with white tips. The browning process, which occurs as the white region matures, has often been attributed to the deposition of suberin in various tissues. However, in pouch-grown tree seedlings of jack pine (Pinus banksiana Lamb.) and eucalyptus (Eucalyptus pilularis Sm.), browning was not linked to suberization but was caused by the deposition of condensed tannins in the walls of all cells external to the stele. Therefore, we propose using the term “tannin zone” to refer to this region of the root. Vitality tests indicated that the cells of the epidermis and cortex were alive in white regions but were dead in brown regions. Following sequential treatment with berberine hemisulfate and potassium thiocyanate, the cortical walls external to the endodermal Casparian band were full of berberine thiocyanate crystals, indicating that they were permeable to berberine. These walls should also be permeable to water and ions, which have smaller molecular dimensions than the tracer dye. Based on the anatomy and permeability of the tannin zone, we predict that its capacity for ion uptake would be reduced compared to the white zone because of a reduced absorptive plasmalemma surface area. In jack pine, some uptake could be effected by the passage cells of the endodermis. The tannin zone should be even less absorptive in eucalyptus because the exodermis remains an apoplastic barrier and the endodermis lacks passage cells. It is difficult to predict the difference between the tannin and white zones with respect to water uptake. Death of the cells external to the endodermis would reduce the resistance of the root to water movement, but deposition of tannins would increase it. The deposition of suberin lamellae in increasing numbers of endodermal cells may also retard water flow. The anatomy and physiological properties of the tannin zone are unique from those of the distal, white zone and the proximal, cork-clad zone.     

Root Browning in Pinus banksiana Lamb. and Eucalyptus pilularis Sm. 2. Anatomy and Permeability of the Cork Zone

The periderm in roots of Pinus banksiana Lamb. and the polyderm in roots of Eucalyptus pilularis Sm. originate from the pericycle. This occurs after the roots have turned brown due to deposition of tannins in the walls of cells external to the endodermis. In both species, cork cells form a continuous sheath around the vascular tissues. The cork cell walls are modified by the presence of suberin, lignin and tannin and it is the latter which imparts a brown colour to the tissue. The first layer of cork cells in both species constitutes an apoplastic barrier which prevents the fluorescent dye, berberine, from entering the vascular tissues, despite the absence of an identifiable Casparian band in the cells. Because the roots are still covered with the cortex and epidermis during early stages of periderm and polyderm formation, it is not possible to tell from the external aspect of the root when it makes a transition from the tannin zone to the cork zone.     

The chemical composition of suberin in apoplastic barriers affects radial hydraulic conductivity differently in the roots of rice (Oryza sativa L. cv. IR64) and corn (Zea mays L. cv. Helix)

Apoplastic transport barriers in the roots of rice (Oryza sativa L. cv. IR64) and corn (Zea mays L. cv. Helix) were isolated enzymatically. Following chemical degradation (monomerization, derivatization), the amounts of aliphatic and aromatic suberin monomers were analysed quantitatively by gas chromatography and mass spectrometry. In corn, suberin was determined for isolated endodermal (ECW) and rhizo-hypodermal (RHCW) cell walls. In rice, the strong lignification of the central cylinder (CC), did not allow the isolation of endodermal cell walls. Similarly, exodermal walls could not be separated from the rhizodermal and sclerenchyma cell layers. Suberin analyses of ECW and RHCW of rice, thus, refer to either the entire CC or to the entire outer part of the root (OPR), the latter lacking the inner cortical cell layer. In both species, aromatic suberin was mainly composed of coumaric and ferulic acids. Aliphatic suberin monomers released from rice and corn belonged to five substance classes: primary fatty acids, primary alcohols, diacids, omega-hydroxy fatty acids, and 2-hydroxy fatty acids, with omega-hydroxy fatty acids being the most prominent substance class. Qualitative composition of aliphatic suberin of rice was different from that of corn; (i) it was much less diverse, and (ii) besides monomers with chain lengths of C(16), a second maximum of C(28) was evident. In corn, C(24) monomers represented the most prominent class of chain lengths. When suberin quantities were related to surface areas of the respective tissues of interest (hypodermis and/or exodermis and endodermis), exodermal cell walls of rice contained, on average, six-times more aliphatic suberin than those of corn. In endodermal cell walls, amounts were 34 times greater in rice than in corn. Significantly higher amounts of suberin detected in the apoplastic barriers of rice corresponded with a substantially lower root hydraulic conductivity (Lp(r)) compared with corn, when water flow was driven by hydrostatic pressure gradients across the apoplast. As the OPR of rice is highly porous and permeable to water, it is argued that this holds true only for the endodermis. The results imply that some caution is required when discussing the role of suberin in terms of an efficient transport barrier for water. The simple view that only the quantity of suberin present is important, may not hold. A more detailed consideration of both the chemical nature of suberins and of the microstructure of deposits is required, i.e. how suberins impregnate wall pores.     

Anatomy of seedling tap roots of loblolly pine (Pinus taeda L.)

The development of tap root anatomical features was investigated in seedlings of loblolly pine (Pinus taeda L.) under both pot and pouch growth regimes. The roots possessed the three anatomical zones previously observed in jack pine (Pinus banksiana Lamb) and Eucalyptus pilularis Sm. - white, condensed tannin (CT), and cork - suggesting that this developmental sequence is preserved over species and growth conditions. Xylem development was centripetal and similar to that found earlier in P. sylvestris. Tracheids with lignified, secondary walls were detected distal to the point of endodermal Casparian band deposition. However, tests for ability to conduct fluid indicated that the protoxylem was capable of transport only proximal to the Casparian bands. Detailed examination of suberin lamella deposition in the endodermis demonstrated that passage cells were present through the white and CT zones. Progressive, centripetal cortical death in the CT zone did not include the endodermis, which remained alive until the cork layer formed, at which point the endodermis was crushed. Therefore, passage cells remain as functional portals for nutrient and water uptake in the CT zone even though the central cortex is dead. Tracer tests indicated that the endodermis provides an apoplastic barrier to tracer diffusion into the stele and that this function was taken over by the young cork layers. Results of this study point to a strong role for the endodermis in the regulation of nutrient and water uptake until the maturation of the first cork layer.     

The role of root apoplastic transport barriers in salt tolerance of rice (<Emphasis Type="Italic">Oryza sativa</Emphasis> L.)

Increasing soil salinity reduces crop yields worldwide, with rice being particularly affected. We have examined the correlation between apoplastic barrier formation in roots, Na⁺ uptake into shoots and plant survival for three rice (Oryza sativa L.) cultivars of varying salt sensitivity: the salt-tolerant Pokkali, moderately tolerant Jaya and sensitive IR20. Rice plants grown hydroponically or in soil for 1 month were subjected to both severe and moderate salinity stress. Apoplastic barriers in roots were visualized using fluorescence microscopy and their chemical composition determined by gas chromatography and mass spectrometry. Na⁺ content was estimated by flame photometry. Suberization of apoplastic barriers in roots of Pokkali was the most extensive of the three cultivars, while Na⁺ accumulation in the shoots was the least. Saline stress induced the strengthening of these barriers in both sensitive and tolerant cultivars, with increase in mRNAs encoding suberin biosynthetic enzymes being detectable within 30 min of stress. Enhanced barriers were detected after several days of moderate stress. Overall, more extensive apoplastic barriers in roots correlated with reduced Na⁺ uptake and enhanced survival when challenged with high salinity. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effects of NO 3 deficiency and NaCl stress on suberin deposition in rhizo- and hypodermal (RHCW) and endodermal cell walls (ECW) of castor bean (Ricinus communis L.) roots

Castor bean (Ricinus communis L.) plants were hydroponically cultivated to achieve NO₃ deficiency (N starvation), salt stress (addition of 100 mM NaCl), or normal conditions. Endodermal (ECW) and rhizodermal and hypodermal cell walls (RHCW) were isolated enzymatically from roots, and suberin monomers were released by transesterification after solvent extraction. Aromatic and aliphatic suberin monomers were identified and quantified by gas chromatography and mass spectrometry. Between 90 and 95% of the released suberin monomers were linear, long-chain, aliphatic compounds (alcohols, acids, diacids, ω-hydroxy acids and 2-hydroxy acids) with an average chain length of 19 C-atoms. The remainder was an aromatic suberin fraction mainly composed of coumaric and ferulic acid. Suberin amounts were significantly increased in ECW and RHCW in the presence of NaCl. In contrast, N starvation led to significantly reduced levels of suberization in ECW and RHCW. It is concluded that R. communis plants reinforce their apoplastic transport barriers in roots in adaptation to NaCl stress in order to minimize NaCl uptake. Under conditions of N starvation the opposite occurs and plants reduce the suberization of their apoplastic transport barriers to facilitate nutrient uptake form the soil.     

RCN1/OsABCG5, an ATP‐binding cassette (ABC) transporter,is required for hypodermal suberization of roots in rice (Oryza sativa)

Suberin is a complex polymer composed of aliphatic and phenolic compounds. It is a constituent of apoplastic plant interfaces. In many plant species, including rice (Oryza sativa), the hypodermis in the outer part of roots forms a suberized cell wall (the Casparian strip and/or suberin lamellae), which inhibits the flow of water and ions and protects against pathogens. To date, there is no genetic evidence that suberin forms an apoplastic transport barrier in the hypodermis. We discovered that a rice reduced culm number1 (rcn1) mutant could not develop roots longer than 100 mm in waterlogged soil. The mutated gene encoded an ATP‐binding cassette (ABC) transporter named RCN1/OsABCG5. RCN1/OsABCG5 gene expression in the wild type was increased in most hypodermal and some endodermal roots cells under stagnant deoxygenated conditions. A GFP‐RCN1/OsABCG5 fusion protein localized at the plasma membrane of the wild type. Under stagnant deoxygenated conditions, well suberized hypodermis developed in wild types but not in rcn1 mutants. Under stagnant deoxygenated conditions, apoplastic tracers (periodic acid and berberine) were blocked at the hypodermis in the wild type but not in rcn1, indicating that the apoplastic barrier in the mutant was impaired. The amount of the major aliphatic suberin monomers originating from C₂₈ and C₃₀ fatty acids or ω‐OH fatty acids was much lower in rcn1 than in the wild type. These findings suggest that RCN1/OsABCG5 has a role in the suberization of the hypodermis of rice roots, which contributes to formation of the apoplastic barrier.