外源硅对植物抗盐性影响的研究进展

盐胁迫是世界范围内影响作物产量和品质的主要非生物胁迫之一,如何提高作物的抗盐性已经引起全世界的关注。硅 (Si) 是地壳中含量仅次于氧的第二大丰富元素。在pH值低于9的介质中,硅通常以单硅酸[Si(OH)₄]的形式被高等植物吸收。尽管目前硅仍然未被认为是植物生长的必需元素,但是作为植物生长的“有益元素”,硅可以缓解各种生物胁迫和非生物胁迫对植物生长发育的抑制。大量的研究表明硅可参与调控植物抗盐的生理生化代谢过程,并与一些信号物质,如乙烯、水杨酸和多胺等存在互作。主要进展如下:1) 植物对硅的吸收存在主动、被动和拒绝吸收三种,硅转运蛋白在硅的吸收和转运中起到非常重要的作用,但是关于该蛋白的编码基因在更多物种中的克隆和功能研究有待于进一步开展。2) 硅可以调节盐胁迫下植物体内的离子平衡,降低植物根系对盐离子的吸收和向地上部的转运,并使盐离子更均匀的分布在根系中;改善盐胁迫下根系对钙、钾、氮等营养元素的吸收,缓解盐胁迫造成的营养失调。近期一些研究表明多胺可能参与硅对根系盐离子吸收的调控。3) 硅可以通过调节水通道蛋白的表达和渗透调节物质的积累提高根系对水分的吸收和向地上部的转运,改善植株的水分状况。4) 硅可通过调节抗氧化酶活性,降低活性氧的产生和积累,同时可以缓解盐胁迫对光合器官和光合色素造成的损伤,保证盐胁迫下植物光合作用的正常进行。5) 植物耐盐的分子机制非常复杂,涉及大量基因的表达和调控以及信号转导过程,包括蛋白质组学和转录组学在内的组学研究策略为从分子水平揭示硅缓解胁迫的机理提供了有力的技术手段。转录组和蛋白质组学的研究表明硅可以通过调控转录因子、激素等相关基因的表达及蛋白的翻译和修饰来调控植物对盐胁迫的快速响应,提高植物的抗盐能力。6) 硅吸收突变体的应用有助于我们更好的了解硅在调控植物生理生化代谢中所发挥的作用。     

白洋淀芦苇湿地生态系统中植硅体的产生和积累研究

植硅体(phytoliths),又称植物蛋白石,存在于大部分植物组织细胞中,主要是依靠植物的根系吸取土壤溶液中的可溶性二氧化硅,在植物细胞或细胞内沉淀硅化而形成的一种固体的非晶质含水二氧化硅颗粒物[1].植硅体主要组成部分是二氧化硅(67％～95％)、水(1％～12％)、碳(0.1％～6％)及少量的无机元素Na、K、 Ca、 Fe、 AL、 Ti等[2],由于其具有较强的抗分解、抗腐蚀和耐高温等特性,可以长时间较稳定地保存在一些岩石和土壤中[3-5],在硅的生物地球化学循环中有着重要的作用,是全球硅循环的重要参与者[6].虽然植硅体作为生物硅的重要组成部分,在全球硅的生物地球化学循环中占据着重要地位,但在湿地生态系统中植硅体产生和积累研究则鲜见报道[7].     

植物硅素营养与土壤硅素肥力研究现状和展望

硅作为重要的植物营养元素,近年来越来越受到人们的关注。本文从植物对硅的吸收及硅在植物体内的输送和分布、硅素在提高植物抗病虫害能力和增强植物抗逆性上的作用、影响土壤硅素肥力的因素以及硅素肥料的施用技术等几个方面,对土壤硅素肥力及植物硅素营养的研究进展进行了综述,并对今后一段时间该领域的研究趋势做了展望。     

土壤植硅体碳积累潜力影响因素分析

土壤植硅体封存有机碳(phytolith-occluded organic carbon,PhytOC)是植物在地质历史时期固碳的重要形式之一,不同植物、不同器官、不同组织、不同生长时期的植硅体形态、含量、大小、分布、组合有所不同。土壤中植硅体碳积累潜力主要受气候变化、植物生产力、植硅体固碳效率、植硅体碳稳定性、土地利用方式、农艺措施、国家宏观政策等因素的影响。本文对各因素进行了较深入的分析。同时指出,固碳机理、植硅体固碳高效品种选育、人为干扰下农林生态系统植硅体碳循环过程为今后植硅体碳汇研究的重点。     

硅调节植物抗病性的机理:进展与展望

【目的】硅素营养增强作物对病虫害的防御能力已得到充分证实,但其作用机理至今仍然没有明确。本文对国内外有关硅素营养与作物病害发展的相互关系及相关机理的最新研究进展进行了归纳总结,为通过植物营养调节技术来提高作物病害防御能力的研究提供理论支撑。【内容】土壤有效硅包括土壤溶液中的单硅酸和易转化为单硅酸的盐类,土壤中有效硅含量一般在50～260 mg/kg。硅虽然不是植物生长发育的必需矿质营养元素,但是硅在减轻植物多种生物和非生物胁迫以及提高植物对病菌的抵抗能力等方面起着重要作用。施硅可以显著地抑制水稻稻瘟病、 纹枯病、 白叶枯病、 胡麻叶斑病,小麦、黄瓜、番茄等植物白粉病等多种病害的发生。关于硅调节植物抗病性的机理,首先提出了机械或物理屏障假设,认为施硅促进了细胞硅化作用的增强,细胞壁角质-硅双层以及表皮细胞乳突的增强,对病菌的入侵起到了物理防御作用。但随着研究的深入,发现物理屏障并非唯一机制,而后提出硅积极参与了生物化学防御过程,发现硅可以诱导感病植物产生酚醛类、黄酮类等抗毒素物质,以及施硅可以提高植物中几丁质酶、过氧化物酶、多酚氧化酶的活性、苯丙氨酸解氨酶等感病植物中病程相关蛋白酶的活性,从而通过化学防御过程提高植物对病害的抵抗能力。随着现代分子技术的发展,从基因组、转录组水平对其防御机制进行了阐明。研究认为硅通过主动的上调感病植物防卫基因及病程相关蛋白基因的表达,以应对病菌侵染。硅诱导植物产生乙烯、茉莉酸、活性氧等系列信号,使植物处于预激活化状态,从而减轻生物胁迫,但是硅在调节植物胁迫信号转导方面的机制还需要深入的研究。【结论】在缺硅土壤中施用硅肥,可以增强作物对病害的抵抗能力,从而大量降低杀菌剂的使用。关于硅调节植物抗病性机理,不能单一归因于某一方面,物理屏障防御机制与生物化学防御过程兼在。硅可能与关键的植物胁迫信号系统相互作用,而最终诱导产生对病原菌的抵抗, 但是这方面的确切机制还不是很清楚,是今后的研究重点。     

不同吸硅型植物各器官硅素及氮、磷、钾素分布特征

通过采集浙江省不同生态地区玉米、黄瓜、冬瓜和番茄植株,研究不同吸硅型植物各器官硅含量分布特征及其与大量营养元素氮、磷、钾的关系。结果表明,喜硅植物玉米和不喜硅植物黄瓜、冬瓜不同器官硅含量均分别满足"末端分布规律",即从根到茎叶呈现逐渐增加的趋势;喜硅植物根、茎、叶中硅含量显著大于不喜硅植物,且各器官之间差异显著。喜硅植物叶片硅含量最高,种子中硅含量最低,表明大量硅可能以沉淀硅形态积累在茎叶中,导致其向生殖器官的转移受到抑制。不同吸硅型植物氮、磷含量在果实、茎叶中的累积较根中显著提高。喜硅植物各器官氮、钾含量与其硅含量呈一定的相关性,其含量变化呈现叶>茎>根的趋势。且喜硅植物不同器官硅含量受生长环境、土壤有效硅含量影响差异显著。     

逆境条件下硅肥调控效应研究进展

硅(Si)元素被认为是N、P、K之后的第四大元素。硅肥有利于促进作物的生长和土壤环境的改善。为了给硅肥利用的研究提供一定参考,通过文献综述的方法,总结了逆境条件下硅肥对低温胁迫的调控作用、对植株病害的防效、对水分胁迫的调控作用以及对重金属胁迫的缓解效应。综合现有研究结果,认为增施硅肥具有提高植株的耐盐胁迫、耐重金属胁迫、抗低温、抗病害等能力。此外,不同硅肥的调控效果不同,而且配施其他肥料或者农艺措施等效果更佳。最后,对今后硅肥利用研究方面提出一些建议:(1)加强硅肥的相关机理研究,如硅是如何改善土壤微环境的,纳米硅材料在植物体内的运输、积累及其对植物抗逆性能的作用机理研究;(2)开展硅肥与非常规水资源安全利用的耦合研究,如何利用硅肥的特性来解决微咸水灌溉和再生水灌溉及其二者耦合灌溉的问题;(3)加强硅肥的提质效应研究。     

非生物胁迫下硅素营养对植物的作用及其机理

硅是对植物生长有益的一种营养元素,能提高作物的产量,改善作物品质。目前有文献报道,硅还可以提高植物对非生物胁迫的抵抗能力。综述了国内外有关硅素营养在减轻铝、锰、盐分、重金属对植物毒害中的作用及机理方面的研究。     

镉胁迫下硅对高粱生长的影响及其作用机制

尽管硅提高植物的抗镉毒害已经被广泛的报道,但是其作用机理还不明确。该试验研究在镉胁迫的条件下,硅对高粱生长的影响及作用机制。结果发现:外源施加硅能显著地缓解镉对高粱无论地上部分还是地下部分鲜重及干重的抑制作用。抗氧化酶结果表明:在正常条件下,硅对抗氧化酶的活性没有显著的影响。在镉毒害的条件下,硅显著增强CAT和POD的活性,SOD的活性不受影响。与此相对应:硅显著地降低MDA的含量。对有机酸的分析发现:在镉胁迫的条件下,叶片中有3种有机酸能够检测到:草酸,丙二酸,丁二酸,并且其含量都在硅的调节下增加。而在根系中所检测的4种有机酸中,草酸和丙二酸的含量受到硅的调节而增加,柠檬酸的含量不受影响,而苹果酸的含量却有下降的趋势。上述结果表明:硅增强高粱幼苗的抗镉胁迫与其增强高粱抗氧化酶活性及增加高粱体内有机酸的含量相关。     

土壤硅磷元素交互作用研究进展

介绍并讨论了国内外有关土壤硅、磷两种元素交互作用的研究动态与进展。硅、磷元素在植物体内的交互作用并非由于硅在生理上代替了植物对磷的需要,而是由于硅的存在降低了过量铁、锰、铝等金属离子的毒害,从而改善了植物的磷素营养。硅、磷元素可以相互促进肥效,提高各自的生物利用率。二者在土壤中存在着竞争性吸附关系,其中土壤对磷的吸附结合能更大。磷的施用不仅降低了土壤对硅的吸附,而且增加了硅的解吸。     

Effects of Silicon on Photosynthesis of Young Cucumber Seedlings Under Osmotic Stress

Effects of silicon (Si) application on photosynthesis of solution-cultured cucumber seedlings were investigated under osmotic stress and unstressed conditions. In unstressed conditions, silicon application had no effect on growth and photosynthetic parameters. The responses of the photosynthetic parameters to abruptly imposed osmotic stress did not differ between silicon treatments. After 1 week exposure to osmotic stress, growth reduction was observed, but it was less severe in seedlings grown with silicon than in those without silicon. Although there were no differences between silicon treatments in stomatal conductance, transpiration rate, cuticular transpiration, or xylem sap exudation rate under osmotic stress, leaf intercellular carbon dioxide (CO₂) concentration was significantly lower and photosynthetic rate tended to be higher in seedlings supplied with silicon. These results suggested that the silicon-induced alleviation of growth reduction under osmotic stress in cucumber was due to amelioration of stress-induced damage of leaf tissues rather than to improvement of leaf water status.     

Effects of silicon on the growth of maize seedlings under normal,aluminum, and salinity stress conditions

The effects of silicon on seed germination and growth parameters of maize seedlings under normal, aluminum (Al), and sodium chloride (NaCl) stress conditions were investigated. The results indicated a dose-dependent relationship between silicon and growth parameters, except for germination attributes.Application of silicon (2 mM) accelerated amylase activity, decreased abscisic acid content, and increased indoleacetic acid and gibberellic acid contents of seedling, The results suggest that the beneficial effects of silicon, at least in part, are mediated by the balance of phytohormones. Exposure to Al (10 mM) significantly decreased all growth parameters of maize seedlings. Application of 2 mM silicon, however, alleviated Al stress and restored almost all growth attributes. Decrease of Al absorption, increase of fructan content, and improvement of amylase activity were considered as the mechanisms of ameliorative function of silicon in Al-stressed maize seedlings. Exposure to silicon, however, did not show beneficial effects on growth parameters of maize seedlings under salt stress conditions.     

硅对盐胁迫下黄瓜幼苗生长和矿质元素吸收的影响

采用幼苗水培实验,研究外源硅对盐胁迫下黄瓜幼苗生长过程中吸收某些矿质元素的影响。对植株地上部和地下部各元素的分析表明,盐胁迫条件下,外源硅有效调节了黄瓜根系对Na+、Ca2+、K+的吸收以及向地上部的转运。适量的硅降低了黄瓜根系从介质中吸收Na+量,并减少其向地上部的运输,而增加了植株体对K+、Ca2+的吸收和转运量,有效缓解了Na+对黄瓜植株体造成的盐胁迫伤害,保证了黄瓜幼苗的正常生长。在相同的处理条件下,盐分敏感品种的Na+在根系和地上部累积量都要比耐盐品种高,而K+的积累量却正好相反,这可能是耐盐品种减轻盐害的主要方式。对其它营养元素的研究表明,施硅抑制了对N素的吸收,对P素的吸收则影响不大。     

The Positive Effects of Silicon on Rice Seedlings Under Saline-Alkali Mixed Stress

Saline-alkali mixed stress is a widespread environmental problem in agriculture. To examine the effects of silicon on rice under saline-alkali stress, the effects of silicon on germination, morphogenesis, chlorophyll content, soluble protein content, malondialdehyde (MDA) content, and antioxidases including peroxidase (POD), superoxide dismutase (SOD), and catalase (CAT) activities were investigated in rice seedlings. The results showed that germination percentage, germination rate, and germination index were increased significantly by the silicon compared with the nonsilicon treatment. Meanwhile, silicon application improved the growth of rice seedlings remarkably under saline-alkali stress, which was consistent with the elevation of chlorophyll contents. The MDA contents were decreased markedly with different silicon treatments. In contrast, the activities of SOD and POD were enhanced significantly with appropriate silicon treatment, which were in accordance with the increasing of soluble protein contents. All the analysis showed that 5 and 10 mmol/L silicon concentrations were more effective in alleviating the injury of rice seedlings under saline-alkali stress.     

Foliar application of silicon at different growth stages alters growth and yield of selected wheat cultivars

To evaluate the response of some selected wheat cultivars to silicon application at different growth stages under drought stress, an experiment was carried out in the greenhouse of College of Agriculture, Shiraz University, Iran, during 2012 using a completely randomized factorial design with four replications. Experimental treatments included drought stress (100% F.C. as control and 40% F.C. as drought) and foliar application of 6 mM sodium silicate (control, application at mid tillering stage, at anthesis stage, and application at tillering + anthesis stages) and wheat cultivars (Sirvan and Chamran, relatively drought-tolerant, and Shiraz and Marvdasht, drought-sensitive cultivars). Drought stress significantly reduced chlorophyll content, leaf area, relative water content, grains per spike, 1000-grain weight, grain yield and biomass of all wheat cultivars. Furthermore, drought stress increased electrolyte leakage of the flag leaves of all cultivars. In contrast, foliar-applied silicon significantly increased these parameters and reduced electrolyte leakage. Furthermore, highest positive influence of silicon application was observed at combined use of silicon both at the tillering + anthesis stages in wheat plants under both stress and non-stress conditions. Significant differences were found in physiological responses among wheat cultivars. The drought tolerant cultivars (Sirvan and Chamran) had significantly higher growth and yield than those of drought sensitive cvs. Shiraz and Marvdasht under drought stress. In conclusion, foliar application of silicon especially at the tillering + anthesis stages was very effective in promoting resistance in wheat plants to drought conditions by maintaining cellular membrane integrity and relative water content, and increasing chlorophyll content.     

Silicon Application by Sorghum Through the Alleviation of Stress-Induced Increase in Hydraulic Resistance

In this study, it was verified whether silicon (Si) affected plant hydraulic resistance, which was one of the significant factors affecting water uptake. Sorghum bicolor (L.) Moench. was grown hydroponically under varying silicon levels and exposed to osmotic stresses. Under osmotic stress, reduction in growth, photosynthesis, and transpiration were alleviated as supplied silicon levels increased. These alleviative effects were ascribed to enhancement of water uptake. Although shoot/root ratio was not affected by silicon, estimated apparent hydraulic resistance was lower in silicon-supplied sorghum than silicon-deficient one under osmotic stress. Simultaneous measurement of transpiration and water uptake rates indicated that under osmotic stress silicon-deficient sorghums showed unbalanced water relation that transpiration rate exceeded water uptake rate, while they were balanced in silicon-supplied sorghums. The results indicated that silicon improved hydraulic resistance, allowing sorghum to avoid from decrease in water uptake rate that happens to silicon-deficient sorghum under water stress.     

钙、硅对铝胁迫下荞麦光合生理的影响

采用营养液培养法,研究了钙、硅对荞麦根长、光合作用和叶绿素荧光的影响。结果表明,15 d时0.5 mmol/L和5 mmol/L Al3 处理降低了荞麦根长、叶绿素含量、净光合速率(Pn)、气孔导度(Ci),对叶绿素荧光Fv/Fm、Fv/Fo的影响不大,30 d后以上伤害均加重。配施钙或硅的处理,能促进根的伸长和提高叶绿素含量,并使叶片的Pn、Ci保持在较高的水平,施硅能明显提高荞麦叶片的叶绿素荧光Fv/Fm、Fv/F,而钙的影响不大。硅对铝胁迫的缓解效果好于钙。     

Growth and Nutrient Use in Four Grasses Under Drought Stress as Mediated by Silicon Fertilizers

Field water stress is a common problem in crop production, especially in arid and semi-arid zones and it is widely hypothesized that silicon (Si) could reduce water stress in plants. We set up a greenhouse study to evaluate some silicon sources—potassium silicate (K₂SiO₃), calcium silicate (CaSiO₃) and silica gel for growth and nutrient uptake by four grass species under adequate and deficit irrigation. The four species studied were Rhodes grass (Chloris gayana), Timothy grass (Phleum pratense), Sudan grass (Sorghum sudanense) and Tall fescue (Festuca arundinacea). For all species, the biomass yield response to applied silicon under deficit irrigation was significantly better than under adequate irrigation. The yield response of Rhodes grass across silicon sources was 205% under deficit irrigation compared with only 59% under adequate irrigation; for Sudan grass it was 49% compared with 26% and for Timothy, it was 48% compared with a mere 1%. The higher responses under deficit irrigation suggest that the plants relied more on silicon to endure drought stress. Biomass yield of individual plants also differed according to soil water levels with Timothy grass being the most sensitive to water stress as it exhibited the highest yield response (209%) to adequate irrigation. This was followed by tall fescue (122%) and Rhodes grass (97%). Sudan grass was the least affected by deficit irrigation, possibly on account of improved root mass and its natural drought tolerance. Strong associations were noted between the uptake of silicon and those of nitrogen (N) and phosphorus (P) irrespective of soil water condition, but the uptake of potassium (K) was more strongly correlated with that of Si under deficit than adequate irrigation. Improvements in plant growth following Si application could therefore be linked to enhanced uptake of major essential nutrients.