Development Of Apoplastic Barriers Research Articles

Salt tolerance in mungbean is associated with controlling Na and Cl transport across roots, regulating Na and Cl accumulation in chloroplasts and maintaining high K in root and leaf mesophyll cells.

Salinity tolerance requires coordinated responses encompassing salt exclusion in roots and tissue/cellular compartmentation of salt in leaves. We investigated the possible control points for salt ions transport in roots and tissue tolerance to Na+ and Cl- in leaves of two contrasting mungbean genotypes, salt-tolerant Jade AU and salt-sensitive BARI Mung-6, grown in nonsaline and saline (75 mM NaCl) soil. Cryo-SEM X-ray microanalysis was used to determine concentrations of Na, Cl, K, Ca, Mg, P, and S in various cell types in roots related to the development of apoplastic barriers, and in leaves related to photosynthetic performance. Jade AU exhibited superior salt exclusion by accumulating higher [Na] in the inner cortex, endodermis, and pericycle with reduced [Na] in xylem vessels and accumulating [Cl] in cortical cell vacuoles compared to BARI Mung-6. Jade AU maintained higher [K] in root cells than BARI Mung-6. In leaves, Jade AU maintained lower [Na] and [Cl] in chloroplasts and preferentially accumulated [K] in mesophyll cells than BARI Mung-6, resulting in higher photosynthetic efficiency. Salinity tolerance in Jade AU was associated with shoot Na and Cl exclusion, effective regulation of Na and Cl accumulation in chloroplasts, and maintenance of high K in root and leaf mesophyll cells.

The changes in the maize root cell walls after exogenous application of auxin in the presence of cadmium.

Cadmium (Cd) is a transition metal and hazardous pollutant that has many toxic effects on plants. This heavy metal poses a health risk for both humans and animals. The cell wall is the first structure of a plant cell that is in contact with Cd; therefore, it can change its composition and/or ratio of wall components accordingly. This paper investigates the changes in the anatomy and cell wall architecture of maize (Zea mays L.) roots grown for 10 days in the presence of auxin indole-3-butyric acid (IBA) and Cd. The application of IBA in the concentration 10-9 M delayed the development of apoplastic barriers, decreased the content of lignin in the cell wall, increased the content of Ca2+ and phenols, and influenced the composition of monosaccharides in polysaccharide fractions when compared to the Cd treatment. Application of IBA improved the Cd2+ fixation to the cell wall and increased the endogenous concentration of auxin depleted by Cd treatment. The proposed scheme from obtained results may explain the possible mechanisms of the exogenously applied IBA and its effects on the changes in the binding of Cd2+ within the cell wall, and on the stimulation of growth that resulted in the amelioration of Cd stress.

Role of roots in adaptation of soil-indifferent Proteaceae to calcareous soils in south-western Australia.

Very few of the >650 Proteaceae species in south-western Australia cope with the high calcium (Ca) levels in young, calcareous soils (soil indifferent); most are Ca sensitive and occur on nutrient-impoverished, acidic soils (calcifuge). We assessed possible control points for Ca transport across roots of two soil-indifferent (Hakea prostrata and Banksia prionotes) and two calcifuge (H. incrassata and B. menziesii) Proteaceae. Using quantitative X-ray microanalysis, we investigated cell-specific elemental Ca concentrations at two positions behind the apex in relation to development of apoplastic barriers in roots of plants grown in nutrient solution with low or high Ca supply. In H. prostrata, Ca accumulated in outer cortical cells at 20 mm behind the apex, but [Ca] was low in other cell types. In H. incrassata, [Ca] was low in all cells. Accumulation of Ca in roots of H. prostrata corresponded to development of apoplastic barriers in the endodermis. We found similar [Ca] profiles in roots and similar [Ca] in leaves of two contrasting Banksia species. Soil-indifferent Hakea and Banksia species show different strategies to inhabit calcareous soils: H. prostrata intercepts Ca in roots, reducing transport to shoots, whereas B. prionotes allocates Ca to specific leaf cells.

Low calcium-induced delay in development of root apoplastic barriers enhances Cd uptake and accumulation in Sedum alfredii

Mineral nutrients play an important role in heavy metal uptake and accumulation in plant. However, the effects of calcium (Ca) supply level on apoplastic transport in roots and consequences for uptake of cadmium (Cd) in hyperaccumulators are poorly understood. Here, we investigated how Ca regulated the development of apoplastic barriers in the roots of two ecotypes of Sedum alfredii and assessed its effects on Cd uptake. Results of correlation analysis indicated that Ca content was positively correlated with the development of Casparian strips (CSs) and suberin lamellae (SL) in the absence or presence of Cd. Simultaneously, low Ca supply was proven to delay the formation of endodermis CSs and suberin accumulation by decreasing the relative expressions of genes associated with CSs localization and lignin/suberin synthesis. Moreover, Cd in apoplastic fluid and cell walls (regarding the apoplastic transport) and symplastic fractions were elevated by low Ca supply. Contrary to high Ca supply, the expression levels of genes related to Cd influx and xylem loading were increased upon low Ca addition in roots of both ecotypes. All the results above suggested that low Ca supply promotes root Cd uptake via apoplastic pathway by delaying apoplastic barriers development and also regulating Cd transport to the xylem in S. alfredii.

Abscisic acid-mediated modifications of radial apoplastic transport pathway play a key role in cadmium uptake in hyperaccumulator Sedum alfredii.

Abscisic acid (ABA) is a key phytohormone underlying plant resistance to toxic metals. However, regulatory effects of ABA on apoplastic transport in roots and consequences for uptake of metal ions are poorly understood. Here, we demonstrate how ABA regulates development of apoplastic barriers in roots of two ecotypes of Sedum alfredii and assess effects on cadmium (Cd) uptake. Under Cd treatment, increased endogenous ABA level was detected in roots of nonhyperaccumulating ecotype (NHE) due to up-regulated expressions of ABA biosynthesis genes (SaABA2, SaNCED), but no change was observed in hyperaccumulating ecotype (HE). Simultaneously, endodermal Casparian strips (CSs) and suberin lamellae (SL) were deposited closer to root tips of NHE compared with HE. Interestingly, the vessel-to-CSs overlap was identified as an ABA-driven anatomical trait. Results of correlation analyses and exogenous applications of ABA/Abamine indicate that ABA regulates development of both types of apoplastic barriers through promoting activities of phenylalanine ammonialyase, peroxidase, and expressions of suberin-related genes (SaCYP86A1, SaGPAT5, and SaKCS20). Using scanning ion-selected electrode technique and PTS tracer confirmed that ABA-promoted deposition of CSs and SL significantly reduced Cd entrance into root stele. Therefore, maintenance of low ABA levels in HE minimized deposition of apoplastic barriers and allowed maximization of Cd uptake via apoplastic pathway.

Early development of apoplastic barriers and molecular mechanisms in juvenile maize roots in response to La2O3 nanoparticles

The increasing occurrence of engineered nanoparticles (NPs) in soils may decrease water uptake in crops, followed by lower crop yield and quality. As one of the most common rare earth oxide NPs, lanthanum oxide (La2O3) NPs may inhibit the relative expressions of aquaporin genes, thus reduce water uptake. In the present study, maize plants were exposed to different La2O3 NPs concentrations (0, 5, 50 mg L−1) for 72 h and 144 h right after the first leaf extended completely. Our results revealed that water uptake was reduced by La2O3 NPs through accelerating the development of apoplastic barriers in maize roots. The level of abscisic acid, determined by using ultra high performance liquid chromatography-tandem mass spectrometry, was increased upon La2O3 NPs exposure. Furthermore, ZmPAL, ZmCCR2 and ZmCAD6, the core genes specific for biosynthesis of lignin, were up-regulated by 3–13 fold in roots exposed to 50 mg L−1 La2O3 NPs. However, ZmF5H was suppressed, indicating that lignin with S units could be excluded for the formed lignin in apoplastic barriers upon La2O3 NPs exposure. The level of essential component of apoplastic barriers - lignin was increased by 1.5-fold. The early development of apoplastic barriers was observed, and stomatal conductance and transpiration rate of La2O3 NPs-treated plants were significantly decreased by 63%–83% and 42%–86%, respectively, as compared to the control. The lignin enriched apoplastic barriers in juvenile maize thus led to the reduction of water uptake, subsequently causing significant growth inhibition. This is the first study to show early development of root apoplastic barriers upon La2O3 NPs exposure. This study will help us better understand the function of apoplastic barriers in roots in response to NPs, further providing fundamental knowledge to develop safer and more efficient agricultural nanotechnology.

The role of root anatomy, organic acids and iron plaque on mercury accumulation in rice

Rice contaminated by mercury [Hg, especially methylmercury (MeHg)] has given rise to great concern in recent years. This study investigated variations in ecophysiological features (anatomy, organic acid secretions, Fe plaque formation) of rice roots and their effects on the uptake and accumulation of total mercury (THg) and MeHg by rice plants. The development of apoplastic barriers in roots of four rice cultivars was observed by a hydroponic experiment while the concentrations of five organic acids, Fe and THg in Fe plaque were determined using a rhizobag trial with different Hg treatments. Cultivars with low Hg accumulation tended to develop strong apoplastic barriers in endodermis, secrete less organic acids and form more Fe plaque on root surfaces and in rhizosphere. Fe concentrations were positively correlated with THg concentrations in rhizosphere’s Fe plaque (R2 = 0.60, P < 0.01), whereas the latter was negatively correlated with bioavailable Hg concentrations in rhizosphere (R2 = 0.40, P < 0.01). Organic acids and Fe plaque formation of rice roots play important roles in Hg uptake and accumulation. The development of apoplastic barriers in root restricts Hg uptake but the significance of suberin deposition on Hg uptake needs further investigations.

Effects of silicon on growth, root anatomy, radial oxygen loss (ROL) and Fe/Mn plaque of Aegiceras corniculatum (L.) Blanco seedlings exposed to cadmium

Biomass, root anatomy, the location of Cd, radial oxygen loss (ROL) and Fe/Mn plaque in Aegiceras corniculatum (L.) blanco were investigated under Si and Cd treatments. The results revealed that Si alleviated the inhibition of growth due to Cd stress. Furthermore, Si prompted the development of apoplastic barriers in roots under Cd stress. Promotion of the apoplastic barrier caused the reduction of ROL. The effect of Si on the formation of Fe/Mn plaque was contrary, but Si reduced the content of Fe plaque and increased the content of Mn plaque. However, the content of Cd on Fe/Mn plaque was significantly increased by Si, which could possibly block the absorption of Cd in the root from the growth media. The present study proposed new evidence of adaptive strategy on metal tolerance and the effect of Si on metal tolerance by A. corniculatum seedlings.

Apoplastic barrier development and water transport in Zea mays seedling roots under salt and osmotic stresses.

The development of apoplastic barriers was studied in Zea mays seedling roots grown in hydroculture solution supplemented with 0-200 mM NaCl or 20% polyethylene glycol (PEG). Casparian bands in the endodermis of both NaCl- and PEG-treated roots were observed closer to the root tip in comparison with those of control roots, but the cell wall modifications in the endodermis and exodermis induced by salt and osmotic stresses differed. High salinity induced the formation of a multiseriate exodermis, which ranged from several cell layers to the entire cortex tissue but did not noticeably influence cell wall suberization in the endodermis. In contrast, osmotic stress accelerated suberization in both the endodermis and exodermis, but the exodermis induced by osmotic stress was limited to several cell layers in the outer cortex adjacent to the epidermis. The hydrostatic hydraulic conductivity (Lp) had decreased significantly after 1 day of PEG treatment, whereas in NaCl-treated roots, Lp decreased to a similar level after 5 days of treatment. Peroxidase activity in the roots increased significantly in response to NaCl and PEG treatments. These data indicate that salt stress and osmotic stress have different effects on the development of apoplastic barriers and water transport in Z. mays seedling roots.

Selenium addition alters mercury uptake, bioavailability in the rhizosphere and root anatomy of rice (Oryza sativa).

Mercury (Hg) is an extremely toxic pollutant, especially in the form of methylmercury (MeHg), whereas selenium (Se) is an essential trace element in the human diet. This study aimed to ascertain whether addition of Se can produce rice with enriched Se and lowered Hg content when growing in Hg-contaminated paddy fields and, if so, to determine the possible mechanisms behind these effects. Two cultivars of rice (Oryza sativa, japonica and indica) were grown in either hydroponic solutions or soil rhizobags with different Se and Hg treatments. Concentrations of total Hg, MeHg and Se were determined in the roots, shoots and brown rice, together with Hg uptake kinetics and Hg bioavailability in the soil. Root anatonmy was also studied. The high Se treatment (5 μg g(-1)) significantly increased brown rice yield by 48 % and total Se content by 2·8-fold, and decreased total Hg and MeHg by 47 and 55 %, respectively, compared with the control treatments. The high Se treatment also markedly reduced 'water-soluble' Hg and MeHg concentrations in the rhizosphere soil, decreased the uptake capacity of Hg by roots and enhanced the development of apoplastic barriers in the root endodermis. Addition of Se to Hg-contaminated soil can help produce brown rice that is simultaneously enriched in Se and contains less total Hg and MeHg. The lowered accumulation of total Hg and MeHg appears to be the result of reduced bioavailability of Hg and production of MeHg in the rhizosphere, suppression of uptake of Hg into the root cells and an enhancement of the development of apoplastic barriers in the endodermis of the roots.

Root anatomy and element distribution vary between two Salix caprea isolates with different Cd accumulation capacities

The understanding of the influence of toxic elements on root anatomy and element distribution is still limited. This study describes anatomical responses, metal accumulation and element distribution of rooted cuttings of Salix caprea after exposure to Cd and/or Zn. Differences in the development of apoplastic barriers and tissue organization in roots between two distinct S. caprea isolates with divergent Cd uptake and accumulation capacities in leaves might reflect an adaptive predisposition based on different natural origins. Energy-dispersive X-ray spectroscopy (EDX) revealed that Cd and Zn interfered with the distribution of elements in a tissue- and isolate-specific manner. Zinc, Ca, Mg, Na and Si were enriched in the peripheral bark, K and S in the phloem and Cd in both vascular tissues. Si levels were lower in the superior Cd translocator. Since the cuttings originated from stocks isolated from polluted and unpolluted sites we probably uncovered different strategies against toxic elements.

Root responses to cadmium in the rhizosphere: a review

This article reviews the responses of plant roots to elevated rhizosphere cadmium (Cd) concentrations. Cadmium enters plants from the soil solution. It traverses the root through symplasmic or apoplasmic pathways before entering the xylem and being translocated to the shoot. Leaf Cd concentrations in excess of 5-10 μg g(-1) dry matter are toxic to most plants, and plants have evolved mechanisms to limit Cd translocation to the shoot. Cadmium movement through the root symplasm is thought to be restricted by the production of phytochelatins and the sequestration of Cd-chelates in vacuoles. Apoplasmic movement of Cd to the xylem can be restricted by the development of the exodermis, endodermis, and other extracellular barriers. Increasing rhizosphere Cd concentrations increase Cd accumulation in the plant, especially in the root. The presence of Cd in the rhizosphere inhibits root elongation and influences root anatomy. Cadmium concentrations are greater in the root apoplasm than in the root symplasm, and tissue Cd concentrations decrease from peripheral to inner root tissues. This article reviews current knowledge of the proteins involved in the transport of Cd across root cell membranes and its detoxification through sequestration in root vacuoles. It describes the development of apoplastic barriers to Cd movement to the xylem and highlights recent experiments indicating that their maturation is accelerated by high Cd concentrations in their immediate locality. It concludes that accelerated maturation of the endodermis in response to local Cd availability is of functional significance in protecting the shoot from excessive Cd loads.

The Role of Plasma Membrane Intrinsic Protein Aquaporins in Water Transport through Roots: Diurnal and Drought Stress Responses Reveal Different Strategies between Isohydric and Anisohydric Cultivars of Grapevine

We report physiological and anatomical characteristics of water transport across roots grown in soil of two cultivars of grapevine (Vitis vinifera) differing in response to water stress (Grenache, isohydric; Chardonnay, anisohydric). Both cultivars have similar root hydraulic conductances (Lo; normalized to root dry weight) that change diurnally. There is a positive correlation between Lo and transpiration. Under water stress, both cultivars have reduced minimum daily Lo (predawn) attributed to the development of apoplastic barriers. Water-stressed and well-watered Chardonnay had the same diurnal change in amplitude of Lo, while water-stressed Grenache showed a reduction in daily amplitude compared with well-watered plants. Hydraulic conductivity of root cortex cells (Lpcell) doubles in Chardonnay but remains unchanged in Grenache. Of the two most highly expressed plasma membrane intrinsic protein (PIP) aquaporins in roots (VvPIP1;1 and VvPIP2;2), only VvPIP2;2 functions as a water channel in Xenopus laevis oocytes. VvPIP1;1 interacts with VvPIP2;2 to induce 3-fold higher water permeability. These two aquaporins are colocated in the root from in situ hybridization and immunolocalization of VvPIP1 and VvPIP2 subfamily members. They occur in root tip, exodermis, root cortex (detected up to 30 mm), and stele. VvPIP2;2 mRNA does not change diurnally or with water stress, in contrast to VvPIP1;1, in which expression reflects the differences in Lo and Lpcell between cultivars in their responses to water stress and rewatering. VvPIP1;1 may regulate water transport across roots such that transpirational demand is matched by root water transport capacity. This occurs on a diurnal basis and in response to water stress that corresponds to the difference in drought tolerance between the cultivars.