Hemicellulose 1 Research Articles

Boron modulates cell wall structure that alleviate aluminum toxicity-induced root growth defects in citrus

Based on the soil acidification situation in citrus plantation of southern China and even worldwide, it's essential to explore improved methods for resistant aluminum (Al) toxicity in plant. Trifoliate (Poncirus trifoliate (L) Raf.) is the most common rootstock in China. The ambition of this research was to illustrate the influence of boron (B) on Al bound in cell wall (CW) and the mechanism in trifoliate seedlings. Healthy seedlings were exposed to 0 or 300 µM AlCl3·6H2O with 10 µM B (H3BO3, control) and 50 µM B, respectively. The results displayed that Al accumulated in roots and root tips during Al exposure. However, Al content in root rips was reduced under B sufficient that reflected by lower morin fluorescence intensity compared to B10-treated roots. More specifically, B addition abates the Al content in root CW, manifested in lower Al content in pectin and hemicellulose 1 (HC1), with HC1 being more prominent. Moreover, B addition inhibited the pectin methyl-esterase (PME) activity to reduce pectin content, while increasing the xyloglucan endotransglucosylase (XET) and xyloglucan endotransglucosylase-hydrolase (XEH) activity. Thereby HC1 content was decreased, ultimately suppressing the bound site of Al in pectin and HC1, and diminished Al content in CW with B supply. Collectively, our results illustrate that B supply was attributed to the mitigation effect of B on Al toxicity by induced devaluation of Al accumulation in CW, which was associated with the reduction of binding Al with pectin and HC1 by regulating compounds of CW through adjusting CW enzymes (PME, XET, and XEH).

Nitrate Increases Aluminum Toxicity and Accumulation in Root of Wheat

Aluminum (Al) toxicity inhibits root growth, while nitrogen is an essential nutrient for plant growth and development. To explore the effects of nitrate (N) on Al toxicity and accumulation in root of wheat, two wheat genotypes, Shengxuan 6 hao (SX6, Al-tolerant genotype) and Zhenmai 168 (ZM168, Al-sensitive genotype), were used in a hydroponic experiment with four treatments (control without N or Al, N, Al, and Al+N, respectively). The results showed that N increased the inhibition of root elongation and aluminum accumulation in root. The Al-sensitive genotype suffered more serious Al toxicity than the Al-tolerant genotype. Histochemical observation clearly showed that Al prefers binding on the root apex 7–10 mm zones, and the Al-sensitive genotype accumulated more Al in these zones. Compared with other treatments, the Al+N treatment had significantly higher O2−, superoxides dismutase (SOD), catalase (CAT), peroxidase (POD) activities, H2O2, Evans blue uptake, malondialdehyde (MDA), ascorbic acid (AsA), pectin, and hemicellulose 1 (HC1) contents in both genotypes. Under Al+N treatment, O2− activity, Evans blue uptake, MDA, and HC1 contents of SX6 were significantly lower than those of ZM168, but SOD, CAT, and POD activities and AsA content exhibited an opposite trend. Therefore, aluminum toxicity and accumulation in root of wheat seedlings were aggravated by nitrate.

Short-chain aldehydes increase aluminum retention and sensitivity by enhancing cell wall polysaccharide contents and pectin demethylation in wheat seedlings

Upon environmental stimuli, aldehydes are generated downstream of reactive oxygen species and thereby contribute to severe cell damage. In this study, using two wheat genotypes differing in aluminum (Al) tolerance, we investigated the effects of lipid peroxidation-derived aldehydes on cell wall composition and subsequent Al-binding capacities. The spatial accumulation of Al along wheat roots was found to the generation of reactive aldehydes, which are highly localized to the apical regions of roots. Elimination of aldehydes by carnosine significantly reduced Al contents in root tips, with a concomitant alleviation of root growth inhibition. In contrast, root growth and Al accumulation were exacerbated by application of the short-chain aldehyde (E)−2-hexenal. We further confirmed that cell wall binding capacity, rather than malate efflux or pH alteration strategies, is associated with the aldehyde-induced accumulation of Al. Scavenging of lipid-derived aldehydes reduced Al accumulation in the pectin and hemicellulose 1 (HC1) fractions of root cell walls, whereas exposure to (E)−2-hexenal promoted a further accumulation of Al, particularly in the cell wall HC1 fraction of the Al-sensitive genotype. Different strategies were introduced by pectin and HC1 to accumulate Al in response to aldehydes in wheat roots. Accumulation in pectin is based on a reduction of methylation levels in response to elevated pectin methylesterase activity and gene expression, whereas that in HC1 is associated with an increase in polysaccharide contents. These findings indicate that aldehydes exacerbate Al phytotoxicity by enhancing Al retention in cell wall polysaccharides.

The role and its transcriptome mechanisms of cell wall polysaccharides in vanadium detoxication of rice

Cell wall-polysaccharides play a crucial role in heavy metals binding, and hence, contribute to heavy metal detoxication in plants. However, there is no data regarding the molecular mechanisms of vanadium (V) binding to root cell walls in plants, especially in rice (Oryza sativa L.). Taking two rice cultivars with various V tolerance as the research material, the present study investigated the effect of various V concentrations on subcellular distribution of V and revealed the regulatory mechanism of cell wall polysaccharides to V exposure. The results showed that rice roots inhibited the upward movement of V, and root cell walls accumulated 69.85–82.71% of V in roots. Furthermore, hemicellulose-1 (HC-1) in cell walls shared up to 67.72% and 66.95% of the cell-wall-bound V in tolerant and sensitive cultivars, respectively. FTIR spectroscopy demonstrated that V stress induced the remolding of cell wall polysaccharides. Under V stress, V-tolerant rice generated up to 19.3% pectin, 40.9% HC-1, and 49.34% HC-2, which were higher than V-sensitive cultivar. The genes encoding UGDH, UGE, and AXS for polysaccharide biosynthesis were higher expressed in V-tolerant rice than V-sensitive rice when exposed to V. The results could provide novel insight for phytoremediation and food security guarantees.

The role of polysaccharides functional groups in cadmium binding in root cell wall of a cadmium-safe rice line

Exploring the mechanism of cadmium (Cd) accumulation in Cd-safe rice lines is beneficial for ensuring rice safety. D62B, a Cd-safe rice line, accumulates less than 0.2 mg Cd kg-1 in the brown rice due to strong capacity of Cd retention in the roots, and the root cell wall (RCW) polysaccharides play important roles. However, specific underlying mechanism of Cd binding on the polysaccharides is little known. In this study, the role of polysaccharides, especially pectin and hemicellulose 1 (HC1), in RCW of D62B was investigated by adsorption experiments and Fourier Transform Infrared Spectroscopy (FTIR) analysis compared with a common rice line (Luhui17). Cadmium was adsorbed on RCW of two rice lines by a multilayer and inhomogeneous chemisorption way with the force of ion transfer or exchange. Cadmium was adsorbed on RCW rapidly at first stage with the limit of internal and external diffusion, and gradually reached saturation. With the removal of pectin, the Cd adsorption rate, maximum Cd adsorption amount and the shift degree of carboxyl groups in the RCW of D62B sharply decreased, which showed advantages compared with Luhui17. Sequential removal of HC1 further decreased the maximum Cd adsorption amount and the shift degree of hydroxyl groups. The results showed that more available functional groups, especially carboxyl groups in pectin and hydroxyl groups in HC1, contributed to Cd immobilization within the RCW of Cd-safe rice line, thus limiting Cd translocation to the shoot and reducing Cd accumulation in the brown rice.

Root cell wall chemistry remodelling enhanced arsenic fixation of a cabbage cultivar

The low- and high-arsenic (As) transferring cultivars (LTC and HTC) of cabbage showed significant differences in As uptake and distribution. We hypothesise that chemistry of root cell wall matrix plays a critical role. LTC and HTC were treated with As and grown for 60 days. As concentration and distribution at subcellular and cell wall component (pectin, hemicellulose and lignin) levels were determined. Remodelling enzymes (PME and PAL) and functional groups of cell wall were analysed. Results showed that shoot biomass of LTC was not affected by As. Less As was accumulated in shoot of LTC than HTC. LTC allocated more As in root and majority of As was deposited in cell wall. LTC had more hemicellulose 1 (HC1) and lignin, PME and PAL activities. The uronic acid contents of pectin, HC1 or HC2 were all positively (P < 0.05) correlated with As concentrations in each component, respectively. Chemistry of LTC root cell wall was remodelled in terms of changes in porosity, HC and lignin contents, and functional groups, which potentially exerted coupling effects on As entering and deposition. The LTC can restrain As in roots through changing characteristics of root cell wall matrix.

Cell wall modification induced by an arbuscular mycorrhizal fungus enhanced cadmium fixation in rice root

The chemistry of root cell wall of rice could be changed by inoculation of arbuscular mycorrhizal fungi (AMF). Hydroponic experiments were conducted to investigate the roles of such changes on cadmium (Cd) uptake and distribution in rice. Results showed that inoculation of AM fungus Rhizophagus intraradices (RI) significantly enhanced (p < 0.05) shoot biomass, plant height and root length of rice, and decreased Cd concentration in shoot and root under Cd stress. Moreover, Cd in root was mainly found in pectin and hemicellulose 1 (HC1) components of root cell wall. Inoculation of RI increased the levels of pectin, HC1 and lignin content, accompanied by the increments of L-phenylalanine ammonia-lyase (PAL) and pectin methylesterase (PME) activities. Results of Fourier transform infrared spectroscopy further showed that more hydroxyl and carboxyl groups in root cell wall were observed in mycorrhizal treatment, compared with control. This study demonstrates that cell wall components could be the locations for Cd fixation, which reduced Cd transportation from root to shoot. Inoculation of AMF may remodel root cell wall biosynthesis and affect the characteristics of Cd fixation. The entering and fixing mechanisms should be further studied.

The regulatory role of root in cadmium accumulation in a high cadmium-accumulating rice line (Oryza sativa L.).

There are some key processes that regulate cadmium (Cd) accumulation in rice. Understanding the characteristics and mechanisms of Cd accumulation in high Cd-accumulating rice lines benefits for excavating relevant genes. Cd accumulation and distribution in roots of Lu527-8, a high Cd-accumulating rice line, were investigated by a hydroponic experiment, with a control of a normal rice line (Lu527-4). Lu527-8 showed significantly higher Cd concentrations in roots than Lu527-4. More than 81% of Cd in roots of two rice lines is distributed in soluble fraction and cell wall. In soluble fraction, there were more organic acids, amino acids, and phytochelatins in Lu527-8, benefiting Cd accumulation. Pectin and hemicellulose 1 (HC1), especially pectin, were main polysaccharides in cell wall. Lu527-8 showed more pectin and HC1 along with higher pectin methylesterase (PME) activity compared with Lu527-4, promoting Cd accumulation. Besides, Lu527-8 showed higher Cd translocation from root to shoot due to more amounts of ethanol-extractable Cd in roots than Lu527-4. In conclusion, specific characteristics of Cd chemical forms and subcellular distribution in roots of high Cd-accumulating rice line are important for Cd accumulation and translocation.

Influence of Cd toxicity on subcellular distribution, chemical forms, and physiological responses of cell wall components towards short-term Cd stress in Solanum nigrum.

Solanum nigrum is a well-documented cadmium (Cd) hyperaccumulator; however, its Cd-induced tolerance capability and detoxification mechanism remain elusive. Hence, a short-term hydroponic experiment was performed in a multiplane glasshouse to determine the influence of Cd toxicity on subcellular distribution, chemical forms, and the physiological responses of cell wall towards Cd stress in a 4-week-old plant. The experiment was conducted following completely randomized design (CRD) with five treatments (n = 4 replicates). The results showed that Cd stress showed dose-dependent response towards growth inhibition. The subcellular distribution of Cd in S. nigrum was in the order of cell wall > soluble fractions > organelles, and Cd was predominantly extracted by 1 M NaCl (29.87~43.66%). The Cd contents in different plant tissues and cell wall components including pectin, hemicellulose 1 (HC1), hemicellulose 2 (HC2), and cellulose were increased with the increase in Cd concentrations; however, the percentage of Cd concentration decreased in pectin and cellulose. Results of the polysaccharide components such as uronic acid, total sugar contents, and pectin methylesterase (PME) activity showed Cd-induced dose-dependent increase relative to exposure Cd stress. The pectin methylesterase (PME) activity was significantly (p < 0.05) enhanced by 125.78% at 75 μM Cd in root, 105.78% and 73.63% at 100 μM Cd in stem and leaf, respectively. In addition, the esterification, amidation, and pectinase treatment of cell wall and Fourier transform infrared spectroscopy (FTIR) assay exhibited many functional groups that were involved in cell wall retention Cd, especially on carboxyl and hydroxyl groups of cell wall components that indicated that the -OH and -COOH groups of S. nigrum cell wall play a crucial role in Cd fixation. In summary, results of the current study will add a novel insight to understand mobilization/immobilization as well as detoxification mechanism of cadmium in S. nigrum.

MsPG1 alleviated aluminum-induced inhibition of root growth by decreasing aluminum accumulation and increasing porosity and extensibility of cell walls in alfalfa (Medicago sativa)

Pectins are abundant in the primary cell walls and involve in regulating intercellular adhesion. MsPG1gene was cloned from alfalfa and its function of Al tolerance was tested in vivo and vitro. Results showed that the expression of MsPG1 was increased by Al, and the MsPG1 was localized in the plasma membrane of epidermal cells in root apexes. Overexpression of MsPG1 in hairy roots of alfalfa increased root growth in the present of Al, which was attributed to the decreased Al accumulations in pectins, hemicellulose 1 (HC1), cell walls and root apexes conferred by the decreased content of HC1 and ratios of negatively charged functional groups (−OH and −COOH) in cell wall. The decreased Al accumulations in pectins and cell walls were also observed in incubation MsPG1 with pectins and cell walls in vitro. TEM and FESEM assays showed that overexpression of MsPG1 protected the cell microstructure of root apexes, and increased the porosity and pore size of porous lattice-like structure of cell wall in the present of Al. These results indicated that MsPG1 could increase Al tolerance of alfalfa via decreasing Al accumulation in cell wall and increasing the porosity and extensibility of cell wall under excess Al stress.

The combined effects of Cd and Pb enhanced metal binding by root cell walls of the phytostabilizer Athyrium wardii (Hook.)

Cell wall acts as a major metal sink in plant roots, while a few studies focused on root cell wall binding in plants for the phytostabilization of multi-metal contaminated soils. A pot experiment was performed to characterize root cell wall properties of the mining ecotype (ME) and non-mining ecotype (NME) of Athyrium wardii (Hook.) in response to Cd and Pb. The cell wall was found to be the major sink for Cd (41.3–54.3%) and Pb (71.4–73.8%) accumulation in roots of the ME when exposed to Cd and/or Pb. The ME showed more Cd and Pb accumulation in root cell walls when exposed to Cd and Pb simultaneously, compared with those exposed to single Cd or Pb as well as the NME, suggesting some modifications for cell walls. The uronic acid contents of pectin and hemicellulose 1 (HC1) in root cell walls of the ME increased significantly when exposed to Cd and Pb simultaneously, suggesting enhanced cell wall binding capacity, thus resulting in more Cd and Pb bound to pectin and HC1. In particular, pectin was found to be the predominant binding site for Cd and Pb. Greater pectin methylesterase activity along with a lower degree of methylesterification were observed in the cell walls of the ME when exposed to Cd and Pb simultaneously. Furthermore, the ME present more O–H, N–H, C–OH, C–O–C, C–C and/or Ar–H in root cell walls when exposed to Cd and Pb simultaneously. These changes of root cell wall properties of the ME lead to enhanced cell wall binding ability in response to the co-contamination of Cd and Pb, thus could be considered a key process for enhanced Cd and Pb accumulation in roots of the ME when exposed to Cd and Pb simultaneously.

Lead-induced oxidative stress triggers root cell wall remodeling and increases lead absorption through esterification of cell wall polysaccharide

Tall fescue (Festuca arundinacea Schreb) shows remarkable tolerance to lead (Pb), but the mechanisms involved in metal tolerance are not yet well understood. Here, tall fescue were firstly cultivated hydroponically with Pb2+ (0, 50, 200 and 1000 mg/L) for 14 days. The results showed that remodeling of root architecture plays important roles in tolerance of tall fescue to Pb2+ stress. Increased cell wall (CW) components contribute to restrict high amount of Pb2+ in roots. Additionally, the uronic acid contents of pectin, hemicellulose 1 (HC1) and hemicellulose 2 (HC2) increased under Pb2+ stress. We further observed that tall fescue cultivated with H2O2 showed similar remodeling of root architecture as Pb2+ treatment. Furthermore, pectin, HC1 and HC2 fractions were sequentially extracted from 0 and 10 mM H2O2 treated roots, and Pb2+ adsorption capacity and contents of carboxyl groups of pectin and HC2 fractions were steadily increased under H2O2 treatment in vitro. Our results suggest that degrees of esterification of pectin and HC2 are regulated by H2O2. High amount of low-esterified pectin and HC2 offer more carboxyl groups, provide more Pb2+ binding sites, and restrict more Pb2+ in the CW, which may enhance tolerance of tall fescue to Pb2+ stress.

Characteristics of cadmium immobilization in the cell wall of root in a cadmium-safe rice line (Oryza sativa L.).

Cultivating cadmium (Cd)-safe rice lines, which show low Cd accumulation in brown rice, is generally beneficial to ensure food safety. The Cd retention in root of Cd-safe rice line D62B plays an important role in its low Cd translocation from root to shoot. To understand the mechanism of Cd retention in root, a hydroponic experiment was conducted to investigate the subcellular distribution of Cd and the contribution of polysaccharides to Cd binding to the root cell wall of a Cd-safe rice line D62B with a common rice line Luhui17 as a control material. D62B retained more Cd in the root by sequestrated a higher proportion of Cd in the cell wall, further it transferred less Cd to shoot. Close to half of the Cd in the root cell wall of D62B was accumulated in the hemicellulose 1 (HC1), and the proportions of HC1 in it were 1.2-1.7 times higher than these of Luhui17. The proportion of Cd in the pectin showed a dose-dependent increase in two rice lines. D62B contained significantly higher uronic acid concentrations of the pectin and greater pectin methyl esterase (PME) activities than Luhui17 in the root cell wall. These results indicated that a superior Cd binding capacity of the cell wall polysaccharides in D62B played an important role in its Cd retention in root.

Contributions of root cell wall polysaccharides to Cu sequestration in castor (Ricinus communis L.) exposed to different Cu stresses

Cell wall polysaccharides play a vital role in binding with toxic metals such as copper (Cu) ions. However, it is still unclear whether the major binding site of Cu in the cell wall varies with different degrees of Cu stresses. Moreover, the contribution of each cell wall polysaccharide fraction to Cu sequestration with different degrees of Cu stresses also remains to be verified. The distribution of Cu in cell wall polysaccharide fractions of castor (Ricinus communis L.) root was investigated with various Cu concentrations in the hydroponic experiment. The results showed that the hemicellulose1 (HC1) fraction fixed 44.9%–67.8% of the total cell wall Cu under Cu stress. In addition, the pectin fraction and hemicelluloses2 (HC2) fraction also contributed to the Cu binding in root cell wall, accounting for 11.0%–25.9% and 14.1%–26.6% of the total cell wall Cu under Cu treatments, respectively. When the Cu levels were ≤25 μmol/L, pectin and HC2 contributed equally to Cu storage in root cell wall. However, when the Cu level was higher than 25 μmol/L, the ability of the pectin to bind Cu was easy to reach saturation. Much more Cu ions were bound on HC1 and HC2 fractions, and the HC2 played a much more important role in Cu binding than pectin. Combining fourier transform infrared (FT-IR) and two-dimensional correlation analysis (2D-COS) techniques, the hemicellulose components were showed not only to accumulate most of Cu in cell wall, but also respond fastest to Cu stress.

Bacillus amyloliquefaciens SAY09 Increases Cadmium Resistance in Plants by Activation of Auxin-Mediated Signaling Pathways.

Without physical contact with plants, certain plant growth-promoting rhizobacteria (PGPR) can release volatile organic compounds (VOCs) to regulate nutrient acquisition and induce systemic immunity in plants. However, whether the PGPR-emitted VOCs can induce cadmium (Cd) tolerance of plants and the underlying mechanisms remain elusive. In this study, we probed the effects of Bacillus amyloliquefaciens (strain SAY09)-emitted VOCs on the growth of Arabidopsis plants under Cd stress. SAY09 exposure alleviates Cd toxicity in plants with increased auxin biosynthesis. RNA-Seq analyses revealed that SAY09 exposure provoked iron (Fe) uptake- and cell wall-associated pathways in the Cd-treated plants. However, SAY09 exposure failed to increase Cd resistance of plants after treatment with 1-naphthylphthalamic acid (NPA) or 2-(4-carboxyphenyl)-4,4,5,5-tetramethylimidazoline-1-oxyl-3-oxide (c-PTIO). Under Cd stress, SAY09 exposure markedly promoted Fe absorption in plants with the increased hemicellulose 1 (HC1) content and Cd deposition in root cell wall, whereas these effects were almost abrogated by treatment with NPA or c-PTIO. Moreover, exogenous NPA remarkably repressed the accumulation of nitric oxide (NO) in the SAY09-exposed roots under Cd stress. Taken together, the findings indicated that NO acted as downstream signals of SAY09-induced auxin to regulate Fe acquisition and augment Cd fixation in roots, thereby ameliorating Cd toxicity.

Pectin enhances rice (Oryza sativa) root phosphorus remobilization.

Plants growing in phosphorus (P)-deficient conditions can either increase their exploration of the environment (hence increasing P uptake) or can solubilize and reutilize P from established tissue sources. However, it is currently unclear if P stored in root cell wall can be reutilized. The present study shows that culture of the rice cultivars 'Nipponbare' (Nip) and 'Kasalath' (Kas) in P-deficient conditions results in progressive reductions in root soluble inorganic phosphate (Pi). However, Nip consistently maintains a higher level of soluble Pi and lower relative cell wall P content than does Kas, indicating that more cell wall P is released in Nip than in Kas. P-deficient Nip has a greater pectin and hemicellulose 1 (HC1) content than does P-deficient Kas, consistent with the significant positive relationship between pectin and root-soluble Pi levels amongst multiple rice cultivars. These observations suggest that increased soluble Pi might result from increased pectin content during P starvation. In vitro experiments showed that pectin releases Pi from insoluble FePO4. Furthermore, an Arabidopsis thaliana mutant with reduced pectin levels (qua1-2), has less root soluble Pi and is more sensitive to P deficiency than the wild type (WT) Col-0, whereas NaCl-treated WT plants exhibit both an increased root pectin content and an elevated soluble Pi content during P-starvation. These observations indicate that pectin can facilitate the remobilization of P deposited in the cell wall. This is a previously unknown mechanism for the reutilization of P in P-starved plants.

Endogenous nitric oxide mediates alleviation of cadmium toxicity induced by calcium in rice seedlings

The effect of calcium chloride (CaCl2) on rice seedling growth under cadmium chloride (CdCl2) stress, as well as the possible role of endogenous nitric oxide (NO) in this process, was studied. The growth of rice seedlings was seriously inhibited by CdCl2, and the inhibition was significantly mitigated by CaCl2. However, hemoglobin (Hb) and 2-(4-carboxyphenyl)-4, 4,5,5-tetramethylimidazoline-l-oxyl-3-oxide (cPTIO) weakened the promotion effect of CaCl2. The results of NO fluorescence localization suggest that growth accelerated by CaCl2 might be associated with elevated NO levels. The content of Cd, protein thiols (PBT), and nonprotein thiols (NPT) in cell walls, cell organelles, and soluble fractions, respectively, of rice seedlings decreased considerably in the presence of CaCl2, whereas the content of pectin, hemicellulose 1 (HC1), and hemicellulose 2 (HC) increased significantly. Elimination of endogenous NO in Cd+Ca treatment could promote the transportation of Cd2+ to cell organelles and soluble fractions and increase the content of NPT and PBT in leaves. In addition, transportation of Cd2+ to cell organelles and soluble fractions was retarded in roots, the content of NPT increased, and the content of PBT decreased. With elimination of endogenous NO in Cd+Ca treatment, the content of pectin, HC1, and HC2 decreased significantly. Thus, Ca may alleviate Cd toxicity via endogenous NO with variation in the levels of NPT, PBT, and matrix polysaccharides.