Silicon isotope fractionation in rice plants, an experimental study on rice growth under hydroponic conditions

Silicon (Si) isotope composition and Si distribution among different rice plant organs and different parts of rice leaf at maturity were studied, which may provide new insights into the mechanism of Si accumulation in plants and biogeochemical Si cycle. An isotope ratio mass spectrometer (IRMS) was used to examine Si isotope fractionation by rice plant grown in a hydroponic system. The observed ³⁰Si-depletion (about 0.3‰) of whole plant relative to external nutrient solutions suggested biologically mediated Si isotope fractionation occurred during uptake. However, it was not possible to judge the Si uptake mechanism with the data. For δ³⁰Si variation within plant, there was a consistent increasing trend from lower to upper tissues (stem < leaf < husk < grain; leaf sheath < leaf blade base <leaf blade middle < leaf blade top). The phenomenon, reflecting kinetic isotope effects, could be explained that isotope fractionation during Si deposition in rice plant was a Rayleigh-like behavior. The range (−2.7‰ to 2.3‰) of δ³⁰Si variation among rice plant tissues in present experiment exceeded that (−1.7‰ to 2.5‰) of phytoliths observed previously in continents, which would enhance understanding the role of phytoliths on globe Si isotope balance.     

硅素对水稻幼苗镉积累及抗胁迫应答的调节效应

施硅(Si)可以显著缓解镉(Cd)胁迫对水稻生长发育的毒害效应。本研究通过水培分根试验,研究了Si对水稻幼苗Cd积累及胁迫应答的调节效应。结果表明: Cd胁迫下水稻幼苗的生物量显著降低,加Si可以显著缓解Cd对水稻幼苗生长的抑制效应。水稻幼苗对Cd的吸收、转运和积累明显受到Si的影响,单侧根系Cd胁迫下加Si(Si-Cd+Si,Si-Cd)使根系对Cd的滞留系数达83.3%～83.6%,限制了Cd从根向地上部转移。单侧根系Cd胁迫下非胁迫侧加Si(Si-Cd)处理的植株对Cd的吸收和累积明显增加,尤其是根中Cd的积累量较单侧根系Cd胁迫下无Si(CK-Cd)处理增加了48.2%;而单侧根系Cd胁迫下双侧加Si(Si-Cd+Si)处理则显著降低了根和地上部对Cd的吸收,较CK-Cd处理分别降低了36.7%和54.9%。双侧Cd胁迫下单侧加Si(Cd-Cd+Si)则使根和地上部对Cd的吸收量显著减少,较双侧根Cd胁迫(Cd-Cd)处理分别降低了57.8%和46.5%。Cd胁迫下水稻幼苗根中含较高浓度的Si,加Si则使Cd胁迫下根和地上部积累更多的Si。加Si也影响了水稻幼苗对其他金属元素如钙(Ca)、镁(Mg)、锰(Mn)的吸收,Cd-Cd+Si处理显著增加了根系和地上部的Ca、Mg浓度,但Mn浓度的变化则因Cd胁迫程度而表现不同。加Si对Cd胁迫下根系超氧化物歧化酶(SOD)和过氧化物酶(POD)活性有一定的影响,尤其是Si-Cd处理的胁迫侧POD和非胁迫侧SOD活性显著上升,有利于清除Cd胁迫产生的氧自由基。总之,Si对Cd胁迫下水稻幼苗生长、Cd和Si等的吸收及根系的抗氧化反应有一定的调节效应,植株体内较高的Si浓度有利于增强植株对Cd的耐受性。     

Silicon deficiency and the adaptation of tropical rice ecotypes

Although silicon (Si) is found at much higher concentrations in healthy rice crops than N, P or K, it has received far less study, particularly for upland rice. There are few reports on the existence, causes, and effects of varying Si supplies in different environments. Chemical analyses of soil, water and plant tissue samples from experiments grown on a typical weathered, acidic upland soil in Colombia found concentrations of Si which were 80-90% lower than those in a typical lowland environment. These results corroborate published findings from West Africa and Hawaii, and lend support to a conclusion that acid-soil upland rice environments in the tropics tend to be deficient in Si, increasing disease damage, among other effects. Critical values for diagnosis of Si deficiency in soils, water and rice husk tissue are suggested. These are reasonably consistent with, but extend the application of previously published values derived from lowland rice studies to upland environments, and use simpler sampling and analysis methods. A strong correlation was found (r = -0.91) between high husk Si concentration and low husk discoloration disease damage, among diverse rice genotypes grown in the uplands. These genotypic differences were mainly explained by their ecotypic affinities: those belonging to the tropical japonica ecotype exhibited 93% higher husk Si concentrations than indica ecotypes (ecotypic means of 23 vs. 12 mg kg^-1 ). This is consistent with a hypothesis that the tropical japonicas may have adapted to Si deficiency in their native upland environment by evolving mechanisms to attain relatively higher tissue Si concentrations than indicas, which are believed to have evolved in the lowlands, where the Si supply is generally ample. Increased understanding of Si-mediated disease resistance in different rice environments and ecotypes could help breeders combine the high yield potential of indica types with the more durable disease resistance of the japonicas, and could contribute to the development of integrated disease management strategies.     

硅介导的水稻对二化螟幼虫钻蛀行为的影响

采用对二化螟敏感(汕优63)和中抗(盐丰47)的水稻品种,设置硅酸钙处理,观察二化螟蚁螟和三龄幼虫在不同处理稻茎上的钻蛀率、蛀入率和蛀入耗时,同时测定不同处理植株的硅细胞数量及植株与土壤的二氧化硅含量,旨在明确水稻施用硅肥对二化螟幼虫钻蛀行为的影响以及这种影响在不同龄期幼虫和不同抗虫性水稻品种间是否存在差异。蚁螟和三龄幼虫钻蛀率随硅肥施用量增加而降低(幅度为5%—28%)。蚁螟蛀入率在硅肥处理之间和水稻品种之间均没有差异;三龄幼虫蛀入率随硅肥施用量增加而显著下降10%—40%,盐丰47上的蛀入率显著低于汕优63(差异10%—30%)。蚁螟蛀入耗时随硅肥施用量增加而显著延长,三龄幼虫蛀入耗时与品种抗性有显著关系。稻茎硅含量随硅肥施用量增加而增大,并且与三龄幼虫蛀入率呈负相关、与三龄幼虫蛀入耗时呈正相关关系。因此,施用硅肥可直接抑制二化螟幼虫钻蛀,蛀入耗时的延长可间接地延长幼虫暴力于其它防治措施的时间;施用硅肥对三龄幼虫成功蛀入的影响大于对蚁螟的影响;相对于抗虫品种,施用硅肥能在更大程度上增强感虫品种对二化螟幼虫钻蛀行为的抑制作用。     

A rice mutant defective in Si uptake

Rice (Oryza sativa) accumulates silicon (Si) in the tops to levels up to 10.0% of shoot dry weight, but the mechanism responsible for high Si uptake by rice roots is not understood. We isolated a rice mutant (GR1) that is defective in active Si uptake by screening M(2) seeds (64,000) of rice cv Oochikara that were treated with 10(-3) M sodium azide for 6 h at 25 degrees C. There were no phenotypic differences between wild type (WT) and GR1 except that the leaf blade of GR1 remained droopy when Si was supplied. Uptake experiments showed that Si uptake by GR1 was significantly lower than that by WT at both low and high Si concentrations. However, there was no difference in the uptake of other nutrients such as phosphorus and potassium. Si concentration in the xylem sap of WT was 33-fold that of the external solution, but that of GR1 was 3-fold higher than the external solution at 0.15 mM Si. Si uptake by WT was inhibited by metabolic inhibitors including NaCN and 2,4-dinitrophenol and by low temperature, whereas Si uptake by GR1 was not inhibited by these agents. These results suggest that an active transport system for Si uptake is disrupted in GR1. Analysis of F(2) populations between GR1 and WT showed that roots with high Si uptake and roots with low Si uptake segregated at a 3:1 ratio, suggesting that GR1 is a recessive mutant of Si uptake.     

水稻施用硅肥对稻纵卷叶螟幼虫取食和成虫产卵选择性的影响

采用感虫水稻品种TN1,设置3种施硅水平,即高硅(0.32 g Si/kg土壤)、低硅(0.16 g Si/kg土壤)和不施硅对照(0 g Si/kg土壤),研究了施用硅肥对稻纵卷叶螟产卵和取食选择性的影响。结果表明:稻纵卷叶螟幼虫对硅处理水稻叶片的取食选择性和成虫在硅处理水稻上的着卵量、着卵率均显著低于对照水稻。高硅处理水稻叶片中的硅含量、可溶性糖含量和碳氮比高于对照,而氮含量低于对照;低硅处理水稻叶片的碳氮比高于对照、氮含量低于对照。同时,硅处理显著降低水稻的卷叶株率和卷叶率。这些结果表明,施硅能增强稻纵卷叶螟对水稻的不选择性,从而增强水稻对稻纵卷叶螟的抗性。     

Physiological and cytological mechanisms of silicon-induced resistance in rice against blast disease

Rice (Oryza sativa L.) blast disease caused by Magnaporthe grisea is one of the most destructive diseases in the rice-growing areas of the world. Silicon is an important nutritional element especially for rice. Two near-isogenic lines of rice with different resistance to blast disease, i.e. CO39 (susceptible) and C101LAC (Pi-1) (resistant), were selected to determine the effects of Si amendment on the severity and incidence of rice blast disease. The physiological and cytological mechanisms involved in the induced disease resistance by silicon were investigated. Exogenous Si application at a concentration of 2 mM reduced the disease index by 45% for CO39 and 56% for C101LAC (Pi-1). Si application alone did not change lignin content and the activities of defense-related enzymes including peroxidase (POD), polyphenol oxidase (PPO) and phenylalanine ammonia-lyase (PAL) in rice leaves of both isogenic lines. However, after inoculation with M. grisea, Si-treated rice plants significantly increased the activities of POD, PPO and PAL in leaves of both isogenic lines. Si and lignin content were also significantly increased in Si-treated inoculated seedlings. Environmental scanning electron microscope observations revealed that Si amendment resulted in higher Si deposit on dumbbell bodies in the rice leaves and silicon papilla accumulation on the guard cell of stoma. These results suggest that silicon-induced defense response and cell silicification of rice leaves altogether contribute to the silicon-induced rice resistance to blast disease.     

水稻纹枯病菌的矿质营养生态位

运用生态位原理与方法,研究了水稻纹枯病菌的矿质元素营养生态位.结果表明,在分蘖盛期、孕穗期、抽穗期和蜡熟期,水稻纹枯病菌矿质营养生态位宽度指数分别为0.2710、0.3865、0.4252和0.4817,即随着水稻的生长而逐渐增大,但各生育期的生态位宽度总体上仍偏小.表明在水稻生长的各时期水稻纹枯病菌只占有了矿质营养类型中的较少部分.水稻纹枯病菌总是优先占有低Mg、低Zn、低Si的营养位,说明Mg、Zn、Si的含量与水稻对纹枯病的抗性紧密相关.     

Electron-probe microanalysis of silicon in the epidermis of rice (Oryza sativa L.) internodes

Summary Electron-probe X-ray microanalysis showed that significant amounts of silicon are accumulated in the entire epidermal system of the rice internode except in the stomatal apparatuses. Thus, there is a lack of specific sites for Si deposition from levels just above the base to the tip of the rice internode. In the intercalary meristem region, 1 cm above the base of the internode, point-count data indicate more Si accumulation in the dumb-bell shaped silica cells than in the long epidermal cells. Above this region, Si is accumulated essentially in a uniform pattern in all epidermal cells. Such a pattern for Si accumulation in rice internodes markedly contrasts with that for Avena internodes and may explain, in part, why rice plants have a higher percentage Si (dry weight basis) in their shoots. The adaptive significance of this silicification pattern in rice is discussed.     

Effects of silicon on aluminum toxicity in upland rice plants

Aims

This study aimed to determine the capacity of Si to mitigate Al toxicity in upland rice plants (Oryza sativa L.) by evaluating plant growth and the Si and Al uptake kinetics.

Methods

Plants were grown for 40 days, after which the Si and Al uptake kinetics (Cmin, Km and Imax) were analyzed. Then, the shoots and roots were separated, and the dry matter, root morphology and Si and Al concentration and accumulation in the plant were evaluated.

Results

Aluminum decreased plant growth and the Si uptake capacity by decreasing the root growth and Si transport system efficiency in the upland rice roots (> Km and > Cmin). Silicon mitigated Al toxicity in the upland rice plants by decreasing Al transport to the plant shoots, although it did not reduce the Al uptake rate (Imax). Si treatment increased the growth of upland rice plant shoots grown in the presence of Al without influencing the root growth. The alleviation of Al toxicity by Si is more evident in the susceptible upland rice cultivar Maravilha.

Conclusions

Silicon mitigated Al toxicity in the upland rice plants by decreasing Al transport to the plant shoots but did not reduce the Al uptake rate by roots.

     

Silicon induces phytochelatin and ROS scavengers facilitating cadmium detoxification in rice

• Cadmium (Cd) is detrimental to crops and the environment. This work examines the natural mechanisms underlying silicon‐ (Si‐)directed Cd detoxification in rice plants.
• The addition of Si to plants under Cd stress caused significant improvements in morphological parameters, chlorophyll score, F_v/F_m and total soluble protein concentration compared to controls, confirming that Si is able to ameliorate Cd‐induced damage in rice plants. This morpho‐physiological evidence was correlated with decreased cell death and electrolyte leakage after Si application.
• The results showed no critical changes in root Cd concentration, while shoot Cd decreased significantly after Si supplementation in comparison with Cd‐stressed rice. Additionally, expression of Cd transporters (OsNRAMP5 and OsHMA2) was significantly down‐regulated while the concentration of phytochelatin, cysteine and glutathione, together with expression of OsPCS1 (phytochelatin synthase) in roots of Cd‐stressed rice was significantly induced when subjected to Si treatment. This confirms that the alleviation of Cd stress is not only limited to the down‐regulation of Cd transporters but also closely related to the phytochelatin‐driven vacuolar storage of Cd in rice roots.
• The enzymatic analysis further revealed the role of SOD and GR enzymes in protecting rice plants from Cd‐induced oxidative harm. These findings suggest a mechanistic basis in rice plants for Si‐mediated mitigation of Cd stress.

     

取食施硅水稻对褐飞虱成虫体内保护酶和解毒酶活性的影响

【目的】本研究旨在明确褐飞虱Nilaparvata lugens成虫取食施硅水稻对其体内保护酶及解毒酶活性的影响,为硅介导的水稻抗虫性的应用提供证据。【方法】以TN1(感虫品种)为供试水稻品种,采用营养液添加Na₂SiO₃·9H₂O的方法设置112 mg Si/L(Si+)和0 mg Si/L(Si-)2种施硅水平,通过酶活性分析测定取食Si+或Si-水稻植株24, 48, 72和96 h后褐飞虱成虫体内保护酶[过氧化氢酶(catalase, CAT)、过氧化物酶(peroxidase, POD)和超氧化物歧化酶(superoxide, SOD)]以及解毒酶[谷胱甘肽S-转移酶(glutathione-S-transferase, GST)、羧酸酯酶(carboxylesterase, CarE)和多功能氧化酶(mixed-functional oxidase, MFO)]活性的动态变化。【结果】与取食Si-水稻植株的褐飞虱成虫相比,取食Si+水稻植株24和48 h时,褐飞虱成虫POD活性分别显著增加101.2%和55....     

硅素分期施用对镉污染水稻光合特性及物质积累的影响

硅(Si)可有效提高水稻对镉(Cd)的抗性,但关于优化硅肥管理对水稻耐Cd性、光合及物质积累等响应机制尚不明晰。采用受污染的农田土壤,通过盆栽试验研究移栽期施Si(T)、拔节期施Si(J)、移栽和拔节期等量分期施Si(TJ)对Cd污染水稻光合作用及物质积累的影响,以不施Si(CK)为对照。结果表明：施Si可明显提高Cd污染下水稻净光合速率,延长叶片光合功能,促进叶片Si沉积,增加Cd在叶中的固定,减少其向籽粒转移。与CK相比,TJ处理在全生育期具有较高的光合速率,产量显著高于其他处理。T、TJ、J处理叶片细胞壁Cd的固持量分别增加11.45%、24.16%和30.15%,且叶片Cd更多以惰性形态(包括果胶和蛋白质结合态Cd、不溶解性磷酸Cd和残渣态Cd)存在,导致叶片Cd转移系数降低,T、TJ、J较CK分别降低33.91%、56.67%和52.16%。此外,主成分分析结果表明,3种Si处理对水稻耐Cd性和光合特性的综合影响大小为TJ>J>T。综合考虑Si调控光合作用、产量、叶片和籽粒Cd浓度的效应,推荐Si素于移栽期与拔节期分期施用。     

Fly ash effect on improving soil properties and rice productivity in Korean paddy soils

Paddy soils in Korea generally require the addition of Si to enhance rice productivity. Coal combustion fly ash, which has a high available Si content and alkaline pH, was selected as a potential source of Si in this study. Two field experiments were carried out to evaluate rice (Oryza sativa) productivity in silt loam and loamy sand soils to which 0, 40, 80, and 120 Mg ha(-1) of fly ash were added with 2 Mg ha(-1) Si as a control. Fly ash increased the soil pH and available Si and P contents of both soils. The amount of available B increased to a maximum of 2.57 mg kg(-1), and the B content of the rice plants increased to a maximum of 52-53 mg kg(-1) following the addition of 120 Mg ha(-1) fly ash. The rice plants did not show toxicity effects. The highest rice yields were achieved following the addition of around 90 Mg ha(-1) fly ash. The application of fly ash increased Si, P and K uptake by the rice plants, but did not result in an excessive uptake of heavy metals in the submerged paddy soil. In conclusion, fly ash could be a good supplement to other inorganic soil amendments to improve the nutrient balance in paddy soils.     

Reexamination of silicon effects on rice growth and production under field conditions using a low silicon mutant

Silicon (Si) is a beneficial element for healthy growth and high and sustainable production of rice, but the mode of action of the beneficial effects has not been well understood. We carried out field trials for four years at two different locations to re-examine the effects of Si on the growth and production of rice using a low silicon rice (lsi1) mutant. The mutant accumulated much lower Si at each growth stage compared with the wild-type rice (Oryza sativa L. cv Oochikara), but there was no difference in the accumulation of other nutrients including N, P, and K. Measurements at different growth stages showed that low Si in the mutant hardly affected the tiller number, chlorophyll content (SPAD value), and root growth. The plant height and shoot dry weight of the wild-type rice were slightly higher than those of the mutant at a later growth stage, but the difference was not significant between the two lines. However, grain yield was reduced by 79–98%, depending on year, due to a low Si accumulation in the mutant, which showed the largest effect of Si on rice production among all studies reported so far. Among the yield components, the percentage of filled spikelets was mostly affected, being only 13.9% of the wild-type rice in the mutant. The grain color of the mutant became brown because of excessive transpiration and infection of pathogens. These results indicate that Si increases rice yield mainly by enhancing the fertility of spikelets.     

Silicon application promotes rice growth and negatively affects development of Spodoptera frugiperda (J. E. Smith)

Silicon (Si) has been reported to enhance plant resistance against biotic and abiotic stressors and also benefit plant growth. These effects are more pronounced in grass species, especially with soil‐applied Si. This study investigated the effects of Si application on rice resistance to Spodoptera frugiperda development and plant vegetative growth. Effects of Si on rice were assessed via soil and foliar applications and compared with untreated plants (control). Si was soil‐ and foliar‐applied as 1% silicic acid solution at a dosage equivalent to 1.4 t Si per ha. After application, leaf material was collected from Si‐treated and untreated plants and placed in Petri dishes with individual S. frugiperda neonate larvae, where development was followed to adult emergence and biological parameters recorded. Vegetative growth parameters recorded in rice plants were the height, chlorophyll content, fresh and dry weights of shoots, and shoot Si content. No effects of Si application were observed on the durations of larval and pupal stages, larval and pupal survival, and sex ratio of S. frugiperda. Insects fed leaves from Si‐treated plants exhibited lower leaf consumption, larval and pupal weights, longevity of males and females, number of eggs, and egg viability. The negative effects were correlated with higher rice Si content. Si application to rice increased plant height, chlorophyll content and dry weight. Our study demonstrates that foliar‐applied Si is as efficient as soil‐applied Si in negatively affecting S. frugiperda development and providing beneficial effects on rice plant growth.     

Effect of silicon on the growth and phosphorus uptake of rice

A pot experiment was conducted to measure the effect of silicon on phosphorus uptake and on the growth of rice at different P levels. Rice (Oryza sativa L. cv. Akebono) was cultured in Kimura B nutrient solution without and with silicon (1.66 mM Si) and with three phosphorus levels (0.014 mM P, low; 0.21 mM, medium; and 0.70 mM, high).Shoot dry weight with Si (+Si) in solution increased with increasing P level, while shoot weight without Si (–Si) was maximum at 0.21 mM P, suggesting that +Si raised the optimum P level for rice. +Si increased shoot weight more when P was low or high than when P was medium.The concentration and amount of inorganic P in shoots increased with increasing P level. +Si did not significantly decrease P uptake by rice at 0.014 mM P, however, uptake at 0.21 and 0.70 mM P was 27 and 30 percent less than uptake with –Si, respectively. In –Si with 0.21 and 0.70 mM P, inorganic P in shoots was more than double the concentration in shoots grown in +Si solutions.The Si concentration in shoots decreased slightly with increasing P level, although Si uptake was not significantly affected by P. +Si decreased the uptake of Fe and Mn by an average of 20 and 50 percent, respectively, thus P/Mn and P/Fe ratios increased in the shoot when P was low.From the results above, the beneficial effect of Si on the growth of rice was clearly shown when P was low or high. This effect may have resulted from decreased Mn and Fe uptake, and thus increased P availability within P deficient plants, or from reduced P uptake when P was high.     

水稻-雍菜间作系统中种间关系和水稻的硅、氮营养状况

间作是一种优良的生态农业模式,但以水稻为主的水生作物间作研究很少.本试验以水稻和雍菜为研究对象,开展了连续两年(2014—2015年)四季的大田试验,研究水稻和雍菜间作条件下产量、种间关系以及水稻的硅、氮营养状况.试验共设置5个处理,包括水稻单作、雍菜单作、水稻-雍菜2∶2间作、3∶2间作、4∶2间作.结果表明: 水稻和雍菜间作显著增加水稻的产量,其中2∶2间作、3∶2间作和4∶2间作在单位面积上的增产幅度分别为77.5%～120.6%、64.9%～80.9%、37.7%～56.0%,但间作显著降低了雍菜的产量.从土地当量比(LER)来看,水稻和雍菜间作可增加系统的综合产量,3种间作的LER全部大于1,其中3∶2间作模式最佳.从竞争指数来看,在间作系统中,水稻比雍菜有更强的竞争能力,特别是在早季.与水稻单作相比,水稻-雍菜间作显著增加水稻叶片对硅和氮的吸收量以及成熟期水稻叶片的硅含量,但是并没有增加水稻叶片的氮含量,甚至在成熟期略低于单作;而对土壤有效硅、铵态氮和硝态氮含量没有显著影响.在整个试验期间,与水稻单作和间作相比,雍菜单作始终有最高的土壤有效硅、铵态氮和硝态氮含量.本研究表明,水稻和雍菜间作可以促进水稻对硅和氮的吸收,增加水稻的竞争能力.     

水稻土施硅对土壤-水稻系统中镉的降低效果

水稻中镉的积累造成人类健康的风险,增加水稻硅素能减轻镉中毒症状,降低稻米镉积累,但是硅对重金属的作用机理尚不清楚。主要研究了在中度和高度镉污染的土壤中,通过施用固态和液态的富硅物质对土壤-水稻系统中镉的吸收和转运的影响,探明决定镉和硅在根与芽的质外体和共质体中的作用机理。试验结果表明:(1)在中度和高度污染的土壤中,镉在土壤-作物系统中的转移和积累情况是不同的,可以通过富硅物质中的单硅酸与镉离子的相互作用,增加镉在硅物质表面的吸附来减少镉在土壤中的流动;(2)富硅物质可以降低水稻根和芽中镉的积累,在高度镉污染的情况下,施用硅可以使镉大量积累在水稻根及其共质体中,并降低根及其共质体中镉的转换和积累;(3)新鲜土壤中水萃取态的单硅酸含量与镉在土壤-作物系统中的流动性、转运以及积累等主要参数密切相关。