Silicon ameliorates manganese toxicity by regulating manganese transport and antioxidant reactions in rice (Oryza sativa L.)

Background and aims

This study aimed to investigate the roles of silicon (Si) in ameliorating manganese (Mn) toxicity in two rice (Oryza sativa L.) cultivars: i.e. cv. Xinxiangyou 640 (XXY), a Mn-sensitive cultivar and cv. Zhuliangyou 99 (ZLY), a Mn-tolerant cultivar.

Methods

Plants were cultured in nutrient solution containing normal Mn (6.7 μM) or high Mn (2.0 mM), both with or without Si supply at 1.5 mM Si.

Results

Plant growth was severely inhibited by high Mn in cv. XXY, but was enhanced by Si supply. In cv. XXY, Si-enhanced tolerance resulted from a restriction of Mn transport, whereas in cv. ZLY Mn uptake was depressed. In cv. XXY, high Mn significantly increased superoxide dismutase (SOD), catalase and ascorbate peroxidase activities but decreased non-protein thiols and glutathione concentrations, leading to accumulation of H₂O₂ and malondialdehyde. The addition of Si significantly counteracted high Mn-elevated malondialdehyde and H₂O₂ concentrations and enhanced plant growth. In cv. ZLY, high Mn considerably raised SOD activities and glutathione concentrations, thus leading to relatively low oxidative damage.

Conclusions

Si-enhanced Mn tolerance was attributed mainly to restricted Mn transport in cv. XXY but to depressed Mn uptake in cv. ZLY. Silicon mainly influenced non-enzymatic antioxidants in these two rice cultivars under high Mn stress.     

Arsenic toxicity to rice (Oryza sativa L.) in Bangladesh

Natural contamination of groundwater with arsenic (As) occurs around the world but is most widespread in the river basin deltas of South and Southeast Asia. Shallow groundwater is extensively used in the Bengal basin for irrigation of rice in the dry winter season, leading to the possibility of As accumulation in soils, toxicity to rice and increased levels of As in rice grain and straw. The impact of As contaminated irrigation water on soil-As content and rice productivity was studied over two winter-season rice crops in the command area of a single tubewell in Faridpur district, Bangladesh. After 16–17 years of use of the tubewell, a spatially variable build up of As and other chemical constituents of the water (Fe, Mn and P) was observed over the command area, with soil-As levels ranging from about 10 to 70 mg kg^?1. A simple mass balance calculation using the current water As level of 0.13 mg As L^?1 suggested that 96% of the added arsenic was retained in the soil. When BRRI dhan 29 rice was grown in two successive years across this soil-As gradient, yield declined progressively from 7–9 to 2–3 t ha^?1 with increasing soil-As concentration. The average yield loss over the 8 ha command area was estimated to be 16%. Rice-straw As content increased with increasing soil-As concentration; however, the toxicity of As to rice resulted in reduced grain-As concentrations in one of the 2 years. The likelihood of As-induced yield reductions and As accumulation in straw and grain has implications to agricultural sustainability, food quality and food security in As-affected regions throughout South and Southeast Asia.     

Straighthead disease of rice (Oryza sativa L.) induced by arsenic toxicity

Straighthead disease is a physiological disorder of rice (Oryza sativa L.) characterized by sterility of the florates/spikelets leading to reduced grain yield. Though the exact cause of straighthead is unknown, a glass house experiment was conducted to investigate the effect of inorganic arsenic on straighthead disease in rice (Oryza sativa L.). BRRI dhan 29, a popular Bangladeshi rice strain, was grown in soils spiked with arsenic (prepared from sodium arsenate, Na₂HAsO₄·7H₂O) at the rate of 10, 20, 30, 40, 50, 60, 70, 80 and 90 mg of As kg^?1 and one control treatment was also run to compare the results. Although there may be some other soil physico-chemical factors involved, arsenic concentration was found to be closely associated with straighthead of rice. With the increase of soil arsenic concentration, the severity of straighthead increased significantly. Up to the 50 mg of As kg^?1 soil treatments, the severity of straighthead incidences were not prevalent. Straighthead resulted in sterile florets with distorted lemma and palea, reduced plant height, tillering, panicle length and grain yield. Straighthead caused approximately 17–100% sterile florates/spikelets formation and about 16–100% loss of grain yield. Straighthead also causes the reduction of panicle formation and panicle length significantly (p < 0.01). In the present study, panicle formation was found to be reduced by 21–95% by straighthead.     

Zinc ameliorates copper-induced oxidative stress in developing rice (Oryza sativa L.) seedlings

Rice (Oryza sativa L.) seedlings were treated with different concentrations of copper (Cu) either in presence or absence of zinc (Zn), and different events were investigated to evaluate the ameliorative effect of Zn on Cu stress. In presence of high Cu concentration, growth of both root and shoots were considerably reduced. Decline in elongation and fresh mass was observed in root and shoot. Zn alone did not show any considerable difference as compared to control, but when supplemented along with high concentration Cu, it prompted the growth of both root and shoot. After 7 days, root growth was 9.36 and 9.59 cm, respectively, at 200 and 500 μM of Cu alone as compared to 10.59 and 12.26 cm at similar Cu concentrations, respectively, in presence of Zn. Cu accumulation was considerably high after 7 days of treatment. In absence of Zn, significant accumulation of Cu was observed. Zn supplementation ameliorated the toxic impact of Cu and minimized its accumulation. Cu treatment for 1 and 7 days resulted in a dose-dependent increase in hydrogen peroxide (H₂O₂). When Cu was added in presence of Zn, the H₂O₂ production in root and shoot was reduced significantly. The increase in H₂O₂ production under Cu stress was accompanied by augmentation of lipid peroxidation. In absence of Zn, Cu alone enhanced the malondialdehyde (MDA) production in both root and shoot after 1 and 7 days of treatment. The MDA content drastically reduced in root and shoot as when Zn was added during Cu treatment. The activities of antioxidant enzymes like superoxide dismutase (SOD), catalase (CAT), and guaiacol peroxidase (GPX) were elevated under Cu stress both in root and shoot. Addition of Zn further stimulated the activities of these enzymes. Both ascorbate (AsA) and glutathione (GSH) contents were high under Cu stress either in presence or absence of Zn. The results suggests that Zn supplementation improves plant survival capacity under high Cu stress by modulating oxidative stress through stimulation of antioxidant mechanisms and restricts the accumulation of toxic concentrations of Cu.     

Silicon-mediated amelioration of zinc toxicity in rice (Oryza sativa L.) seedlings

Background and aims

Silicon (Si) was suggested to enhance plant resistance to toxic elements, and its beneficial role was mainly based on external and internal plant mechanisms. This work aimed at investigating the internal effect of Si on zinc (Zn) detoxification to rice (Oryza sativa L., cv. Tian You 116) seedlings.

Methods

In a hydroponic experiment, we examined the uptake, xylem loading and localization of Zn in rice seedlings under the condition of 200?μM Zn contamination with the additional silicate supply at three levels ( 0, 0.5 and 1.8?mM).

Results

The silicate addition significantly increased the seedling biomass, and decreased Zn concentration in both root and shoot of seedlings and in xylem sap flow. Zinpyr-1 fluorescence test and Energy-dispersive X-ray spectroscopy analysis showed the concentration of biologically active Zn²⁺ decreased, and Zn and Si co-localized in the cell wall of metabolically less active tissues, especially in sclerenchyma of root. The fractionation analysis further supported silicate supply increased about 10% the cell wall bound fraction of Zn.

Conclusions

This study suggests the Si-assisted Zn tolerance of rice is mainly due to the reduction of uptake and translocation of excess Zn, and a stronger binding of Zn in the cell wall of less bioactive tissues might also contribute to some degree.     

Shoot tolerance mechanisms to iron toxicity in rice (Oryza sativa L.)

Iron toxicity frequently affects lowland rice and leads to oxidative stress via the Fenton reaction. Tolerance mechanisms were investigated in contrasting genotypes: the intolerant IR29 and the tolerant recombinant inbred line FL483. Seedlings were exposed to 1000 mg L^‐1 ferrous iron, and the regulation of genes involved in three hypothetical tolerance mechanisms was investigated (I) Iron uptake, partitioning and storage. The iron concentration and speciation in different plant tissues did not differ significantly between genotypes. Sub‐cellular iron partitioning genes such as vacuolar iron transporters or ferritin showed no genotypic differences. (II) Antioxidant biosynthesis. Only one gene involved in carotenoid biosynthesis showed genotypic differences, but carotenoids are unlikely to scavenge the reactive oxygen species (ROS) involved in Fe toxicity, i.e. H₂O₂ and hydroxyl radicals. (III) Enzymatic activities for ROS scavenging and antioxidants turnover. In shoots, glutathione‐S‐transferase and ascorbate oxidase genes showed genotypic differences, and consistently, the tolerant FL483 had lower dehydroascorbate reductase and higher ascorbate oxidase activity, suggesting that high rates ascorbate reduction confer sensitivity. This hypothesis was confirmed by application of exogenous reduced ascorbate or L‐galactono‐1,4‐lactone, which increased lipid peroxidation under iron toxic conditions. Our results demonstrate in planta pro‐oxidant activity of reduced ascorbate in the presence of iron.     

Indel polymorphism in sugar translocation and transport genes associated with grain filling of rice (Oryza sativa L.)

Molecular Breeding - Depending on the position on the panicle, grain filling differs for spikelets on primary branches and secondary branches of upper and lower portions in rice (Oryza sativa L.)....     

Identification of salt-stress responsive genes in rice (Oryza sativa L.) by cDNA array

To identify salt stress-responsive genes, we constructed a cDNA library with the salt-tolerant rice cultivar, Lansheng. About 15000 plasmids were extracted and dotted on filters with Biomeck 2000 HDRT system or by hand. Thirty genes were identified to display altered expression levels responding to 150 mmol/L NaCl. Among them eighteen genes were up-regulated and the remainders down-regulated. Twenty-seven genes have their homologous genes in Gen-Bank Databases. The expression of twelve genes was studied by Northern analysis. Based on the functions, these genes can be classified into five categories, including photosynthesis-related gene, transport-related gene, metabolism-related gene, stress- or resistance-related gene and the others with various functions. The results showed that salt stress influenced many aspects of rice growth. Some of these genes may play important roles in plant salt tolerance.     

Auxin signaling is closely associated with Zn-efficiency in rice (Oryza sativa L.)

This study elucidates the involvement of auxin with Zn-efficiency (ZE) in Zn-efficient rice var. Pokkali. Pokkali showed no significant decrease in morpho-physiological features, electrolyte leakage and total soluble proteins due to Zn deficiency as compared with Zn-sufficient seedlings. However, auxin inhibitor under Zn deficiency severely affected these characteristics, suggesting that ZE is associated with auxin signaling in rice. Results further revealed significant decreases in the expression of Zn transporter genes (OsIRT1, OsZIP4 and OsZIP1), OsDMAS1 (deoxymugeneic acid synthase) and phytochelatin in roots due to auxin inhibitor. It implies that auxin signaling may trigger Zn uptake, transport and chelation in rice seedlings to withstand Zn-deficiency. Further, significant reduction of major S-metabolites (cysteine, methionine, glutathione) and antioxidant enzymes (superoxide dismutase and glutathione reductase) along with increased H₂O₂ content, due to auxin inhibitor under Zn deficiency compared with controls. Taken together, these findings reveal that mechanisms associated with ZE in Pokkali are dependent on auxin signaling.     

Morphological and physiological characteristics of a root-hairless mutant in rice (Oryza sativa L.)

This paper reports morphological and physiological characteristics of a first root-hairless mutant (RH2) of rice (Oryza sativa L.), which can be useful in advancing knowledge on the role of root hairs in water and nutrient uptake, and genetics of root hairs. The mutant was selected among NaN₃ mutagenized progeny of the rice cultivar Oochikara. Microscopic observations showed absence of root hairs in RH2. At the seedling stage, RH2 showed shorter seedling height and shorter roots compared to the wild type variety Oochikara. Because of the differences in seedling growth, all comparisons between Oochikara and RH2 in uptake-related characters were made on the basis of values adjusted by the dry weight of either the shoot or the root. When grown at low water potential in soil, Oochikara and RH2 were similar in shoot water content and transpiration per unit shoot dry weight, and similarly, at low water potential in solution culture, there was no significant difference between Oochikara and RH2 in transpiration per unit shoot dry weight. These results suggest that at the seedling stage, root hairs do not significantly contribute to uptake of water. In solution culture, Oochikara and RH2 did not significantly differ in phosphate uptake per unit root dry weight. This result supports the previous work that root hairs do not contribute to phosphate uptake in solution culture. Regarding to response to plant hormones, RH2 showed a higher level of resistance to two synthetic auxins, 2,4-dichlorophenoxyacetic acid (2,4-D) and 1-naphthaleneacetic acid (NAA) than Oochikara. NAA treatment induced very short root hairs in RH2, suggesting that the absence of root hairs in RH2 may be due to a shortage of endogenous auxin. Genetic analysis showed that the root hairless character in RH2 is inherited as a single recessive gene.     

Heterosis breeding in rice (Oryza sativa L.)

Summary Studies conducted at the International Rice Research Institute (IRRI) during 1980 and 1981 have shown up to 73% heterosis, 59% heterobeltiosis and 34% standard heterosis for yield in rice. The latter was estimated in comparison to commercial varieties: IR36 and IR42 (yield 4–5 t/ha in wet season trials and 7–8 t/ha in dry season trials). Generally speaking, absolute yield was lower and extent of standard heterosis was higher in wet season than in dry season with some exception. Yields up to 5.9 t/ha (22% standard heterosis) in the wet season and 10.4 t/ha (34% standard heterosis) in the dry season were obtained. Most of the hybrids performed better in some season while some performed better in both seasons. Hybrids showed better lodging resistance although they were 5–10 cm taller. F₁ hybrids had significant positive correlations with the parental traits viz., yield (r = 0.446), tillering (r = 0.746), height (r = 0.810) and flowering (r = 0.843). Selection of parents among elite breeding lines on the basis of their per se yield performance, diverse origin and resistance to insects and diseases should give heterotic combination. Yield advantage of hybrids was due primarily to increase in number of spikelets per unit area even though tiller number was reduced. Grain weight was either the same or slightly higher. High yielding hybrids also showed significant heterosis and heterobeltiosis for total dry matter and harvest index. For commercial utilization of heterosis in rice, effective male sterility and fertility restoration systems are available and up to 45% natural outcrossing on male sterile lines has been observed. Consequently, F₁ rice hybrid have been successfully developed and used in China. Prospects of developing hybrid rice varieties elsewhere appear bright especially in countries that have organized seed production, certification and distribution programs and where hybrid seed can be produced at a reasonable cost.     

Variability and genetics of tolerance for aluminum toxicity in rice (Oryza sativa L.)

A study was undertaken to investigate the variability among lowland rice cultivars and the mode of gene action of aluminum (Al) toxicity tolerance in rice. Pregerminated seeds were grown in a nutrient solution containing 30 ppm Al and in normal nutrient solution, and relative root length (RRL) was determined at the 14-day-old stage to characterize genotypes for tolerance. Sixty-two traditional rice cultivars grown on lowland acid sulfate soil areas of Asia and West Africa were tested. Tolerant varieties Azucena, IRAT104, and Moroberekan, moderately sensitive IR29 and IR43, and sensitive IR45 and IR1552 were used to investigate the genetics of tolerance by diallel analysis. Of the 62 cultivars tested, only 3 were found to be sensitive to A l toxicity. Among the tolerant cultivars identified, 11 (Siyam Kuning, Gudabang Putih, Siyam, Lemo, Khao Daeng, Siyamhalus, Bjm-12, Ketan, Seribu Gantang, Bayer Raden Rati, and Padi Kanji) were found to possess higher levels of tolerance than the improved tolerant upland cultivar IRAT104. Diallel analysis revealed that high RRL is governed by both additive and dominance effects with a preponderance of additive effects. The trait exhibited partial dominance, and one group of genes was detected. Heritability was high, and environmenal effects were low. Findings suggest that when breeding for A1 toxicity tolerance, selection can be made in early generations. The pedigree method of breeding would be suitable. Combining ability analysis revealed the importance of both general combining ability (GCA) and specific combining ability (SCA) in the genetics of A1 toxicity tolerance in rice. GCA was more prevalent than SCA. Tolerant parens Azucena, IRAT104, and Moroberekan were the best general combiners. The presence of reciprocal effects among crosses suggested the proper choice of parents in hybridization programs. Results indicated that Azucena, IRAT 104, and Moroberekan should be used as the female in crosses for A1 toxicity tolerance.     

Molecular mapping of genes conferring aluminum tolerance in rice (Oryza sativa L.)

Crop productivity on acid soil is restricted by multiple abiotic stress factors. Aluminum (Al) tolerance seems to be a key to productivity on soil with a pH below 5.0, but other factors such as Mn toxicity and the deficiency of P, Ca and Mg also play a role. The development of Al-tolerant genotypes of rice is an urgent necessity for improving crop productivity in developing countries. Inhibition of root growth is a primary and early symptom of Al toxicity. The present study was conducted to identify genetic factors controlling the aluminum tolerance of rice. Several parameters related to Al tolerance, most importantly the relative root growth under Al stress versus non-stress conditions, were scored in 188 F₃ selfed families from a cross between an Al-tolerant Vietnamese local variety, Chiembau, and an Al-susceptible improved variety, Omon269–65. The two varieties are both Oryza sativa ssp. indica, but showed a relatively high level of DNA polymorphism, permitting the assembly of an RFLP map consisting of 164 loci spanning 1,715.8 cM, and covering most of the rice genome. A total of nine different genomic regions on eight chromosomes have been implicated in the genetic control of root and shoot growth under aluminum stress. By far the greatest effects on aluminum tolerance were associated with the region near WG110 on chromosome 1. This region does not seem to correspond to most of the genes that have been mapped for aluminum tolerance in other species, nor do they correspond closely to one another. Most results, both from physiological studies and from molecular mapping studies, tend to suggest that aluminum tolerance is a complex multi-genic trait. The identification of DNA markers (such as WG110) that are diagnostic for aluminum tolerance in particular gene pools provides an important starting point for transferring and pyramiding genes that may contribute to the sustainable improvement of crop productivity in aluminum-rich soils. The isolation of genes responsible for aluminum tolerance is likely to be necessary to gain a comprehensive understanding of this complex trait. Received: 29 March 2000 / Accepted: 16 August 2000     

Candidate genes for drought tolerance and improved productivity in rice (Oryza sativa L.)

Candidate genes are sequenced genes of known biological action involved in the development or physiology of a trait. Twenty-one putative candidate genes were designed after an exhaustive search in the public databases along with an elaborate literature survey for candidate gene products and/or regulatory sequences associated with enhanced drought resistance. The downloaded sequences were then used to design primers considering the flanking sequences as well. Polymerase chain reaction (PCR) performed on 10 diverse cultivars that involvedJaponica, Indica and local accessions, revealed 12 polymorphic candidate genes. Seven polymorphic candidate genes were then utilized to genotype 148 individuals of CT9993 × IR62266 doubled haploid (DH) mapping population. The segregation data were tested for deviation from the expected Mendelian ratio (1:1) using a Chi-square test (<1%). Based on this, four candidate genes were assessed to be significant and the remaining three, as non-significant. All the significant candidate genes were biased towards CT9993, the female parent in the DH mapping population. Single-marker analysis strongly associated (<1%) them to different traits under both well-watered and low-moisture stress conditions. Two candidate genes,EXP15 andEXP13, were found to be associated with root number and silicon content in the stem respectively, under both well-watered and low-moisture stress conditions