小麦抗旱机理研究进展

干旱是影响小麦产量和品质的重要环境因素,而小麦在干旱胁迫下的抗旱机理非常复杂,不同程度以及不同发育时期的干旱胁迫对不同品种的影响各异。对近年来有关干旱胁迫下小麦的形态结构与渗透调节物质的变化,特别是基因和蛋白表达量的改变进行了综述,对小麦在干旱胁迫下如何提高品质及产量等问题提出建议。     

干旱低磷胁迫对不同品种小麦根系导水率的影响

控制磷素水平,采用控制灌水量(正常供水、中度及重度干旱胁迫)的盆栽试验法,选择抗旱性小麦品种陕合6号(W1)和水分敏感型品种郑引1号(W2)为供试材料。用压力室法测定了三叶期的两品种小麦根系导水率(LPr)的变化规律。结果表明：陕合6号,在有磷正常供水处理( PH)下具有较高的导水率,干旱胁迫时LPr降低较少,且复水后有较强的恢复能力。郑引1号, PH的LPr值相对较小,干旱导致的根系导水率下降非常突出,复水后的恢复能力也较弱。另外,干旱胁迫对小麦苗期根系导水率的影响大于磷胁迫对其导水率的影响,且两品种小麦无磷止常供水处理(-PH)的LPr分别为 PH的31．9%和53．6％,即磷对前者LPr的影响大于后者。     

低磷和干旱胁迫对大豆植株干物质积累及磷效率的影响

土壤缺磷和季节性干旱已经成为南方酸性红壤地区大豆生产的主要限制因素之一。选取2个大豆品种巴西10号(磷高效)和本地2号(磷低效),研究其在不同磷素(0,15, 30 mg/kg P)和水分处理(分别在开花期和结荚期进行干旱胁迫)下的反应,从植株生物量、叶绿素含量、磷效率指标等方面研究不同基因型大豆对水磷耦合胁迫的适应机制。研究结果表明,随着土壤磷素水平的增加,两个品种的生物量和叶片叶绿素含量显著增加,根冠比则显著下降。在同一磷素水平处理下,干旱胁迫则导致较高的根冠比,对叶片叶绿素含量影响不大,两个品种表现一致。两个基因型大豆受到干旱胁迫后,其产量均显著低于正常水分处理。中等施磷能显著提高两个大豆品种的产量,但高磷处理对产量的增加幅度有限,甚至高磷处理还造成本地2号减产。巴西10号的产量随土壤中磷素的增加而增加,而本地2号的产量则为中磷>高磷>低磷,不管是磷处理还是水分处理,巴西10号的产量均高于本地2号。无论是花期干旱还是结荚期干旱,巴西10号和本地2号的根磷效率比、磷吸收效率及磷转移效率均随土壤磷浓度的增加而增加,磷利用效率则降低。总体上来讲,巴西10号的磷吸收效率和利用效率高于本地2号,而根磷效率比、磷转移效率则小于本地2号。     

干旱胁迫对小麦幼苗根系生长和叶片光合作用的影响

采用水培试验方法,以2个耐旱性不同的小麦品种(敏感型望水白和耐旱型洛旱7号)为材料,研究了干旱胁迫对小麦幼苗根系形态、生理特性以及叶片光合作用的影响,以期揭示小麦幼苗对干旱胁迫的适应机制.结果表明： 干旱胁迫下,2个小麦品种幼苗的根系活力显著增大,而根数和根系表面积受到抑制;干旱胁迫降低了望水白的叶片相对含水量,提高了束缚水/自由水,而对洛旱7号无显著影响;干旱胁迫降低了2个小麦品种叶片的叶绿素含量、净光合速率、蒸腾速率、气孔导度和胞间CO2浓度,但随胁迫时间的延长,洛旱7号的叶绿素含量和净光合速率与对照差异不显著;干旱胁迫降低了2个小麦品种幼苗的单株叶面积,以及望水白的根系、地上部和植株生物量,而对洛旱7号无显著影响.水分胁迫下,耐旱型品种可以通过提高根系活力、保持较高的根系生长量来补偿根系吸收面积的下降,保持较高的根系吸水能力,进而维持较高的光合面积和光合速率,缓解干旱对生长的抑制.     

干旱,半干旱地区作物育种的困惑与出路

粮食问题主要取决于一年生谷类作物产量。作物产量低而不稳的原因主要是病虫害及各种胁迫生境,其中干旱缺水为最大的产量限制因素,提高作物生产力的途径有二：其一是改善作物的生长环境,其二是通过育种手段选育在各肿胁迫环境中具有优良表现的基因型（品种）。矮秆化育种手段使水肥充裕区小麦产量有显著的提高,是通过提高收获指数获得的。干旱、半干旱地区育种却未能获得显著效果,要提高干旱、半干旱地区小麦育种的成效,对干旱     

干旱胁迫对小麦幼苗根系生长和叶片光合作用的影响

采用水培试验方法,以2个耐旱性不同的小麦品种(敏感型望水白和耐旱型洛旱7号)为材料,研究了干旱胁迫对小麦幼苗根系形态、生理特性以及叶片光合作用的影响,以期揭示小麦幼苗对干旱胁迫的适应机制.结果表明： 干旱胁迫下,2个小麦品种幼苗的根系活力显著增大,而根数和根系表面积受到抑制;干旱胁迫降低了望水白的叶片相对含水量,提高了束缚水/自由水,而对洛旱7号无显著影响;干旱胁迫降低了2个小麦品种叶片的叶绿素含量、净光合速率、蒸腾速率、气孔导度和胞间CO2浓度,但随胁迫时间的延长,洛旱7号的叶绿素含量和净光合速率与对照差异不显著;干旱胁迫降低了2个小麦品种幼苗的单株叶面积,以及望水白的根系、地上部和植株生物量,而对洛旱7号无显著影响.水分胁迫下,耐旱型品种可以通过提高根系活力、保持较高的根系生长量来补偿根系吸收面积的下降,保持较高的根系吸水能力,进而维持较高的光合面积和光合速率,缓解干旱对生长的抑制.     

稀土浸种对水旱地小麦根苗生长及产量的影响

以山西农业大学培育的冬小麦031165为供试材料,对水、旱地小麦采用浓度均为1 000 mg/kg的稀土溶液[La(NO3)3,Ce(NO3)3]进行浸种,研究其对小麦根苗生长、抗氧化酶系活性及产量的影响。结果表明:干旱胁迫抑制了小麦根苗生长,降低了小麦的实际产量。无论干旱胁迫与否,稀土浸种均能显著提高不同生育期小麦株高、地上干重、叶面积及根系生物量;稀土浸种可提高部分生育期旗叶中SOD、POD活性,降低胞内MDA含量,显著提高小麦产量和经济系数,且在干旱胁迫下作用较明显。从整体上看,Ce浸种对小麦生长的促进作用比La浸种明显。     

干旱胁迫对小麦幼苗抗氰呼吸和活性氧代谢的影响

研究了干旱胁迫对抗旱性强弱不同的两种小麦幼苗的抗氰呼吸和活性氧代谢的影响。干旱胁迫导致了两种小麦抗氰呼吸活性及基因转录水平的下降,但抗旱品种在轻度干旱胁迫下表现出一定的适应能力,其抗氰呼吸活性及基因转录水平均高于不抗旱品种。干旱胁迫下,对干旱敏感的小麦幼苗叶片中活性氧含量高于抗旱小麦;3种抗氧化酶的活性低于抗旱小麦的3种抗氧化酶的活性。据此认为,严重的干旱胁迫引起活性氧含量的增加扰动了活性氧与抗氰呼吸之间的应答平衡,但抗氰呼吸可能通过清除活性氧等机制而起了抗旱的作用。     

高温、干旱及其互作对两个筋力小麦品种淀粉糊化特性的影响

以强筋和弱筋的两个代表性小麦品种为材料,采用盆栽与人工气候箱模拟的方法,研究了花后不同时期高温、干旱及其互作对籽粒淀粉糊化特性的影响。结果表明,高温和干旱均显著影响淀粉糊化特性,但两品种表现有所不同:高温胁迫使强筋小麦品种豫麦34的峰值黏度、最终黏度(除花后5 d外)、稀懈值(除花后15 d外)和反弹值显著增大;而弱筋小麦品种豫麦50低谷黏度和最终黏度显著下降,其峰值黏度、反弹值变化不明显,从不同时期看,花后15 d影响较大。干旱胁迫使豫麦34多数黏度参数增大;而使豫麦50峰值黏度、反弹值和稀懈值下降,其低谷黏度和最终黏度在灌浆前期和中期干旱胁迫下增大,后期干旱胁迫则明显下降。研究结果还表明,花后高温与干旱胁迫对小麦黏度参数的影响存在显著的互作效应。从F值大小看,互作对弱筋小麦品种豫麦50多数黏度参数影响较大,而对强筋小麦品种豫麦34淀粉黏度参数影响较小,反映了高温、干旱及其互作对小麦淀粉特性的影响存在着显著的基因型差异。     

渗透胁迫和缺磷对小麦幼苗生长的影响

水培条件下缺磷小麦幼苗的叶片含水量和叶绿素含量显著降低,丙二醛含量增加,小麦幼苗生长受抑,地上部分生长受到的影响大于根系,上述指标的变化幅度还与缺磷程度呈正相关,且不耐低磷品种中国春大于耐低磷品种烟中144。在相同条件下,渗透胁迫和缺磷两者表现出胁迫加剧的现象。     

Salinity and drought interaction in wheat (Triticum aestivum L.) is affected by the genotype and plant growth stage

Salinity and drought are important agro-environmental problems occurring separately as well as together with the combined occurrence increasing with time due to climate change. Screening of bread wheat genotypes against salinity or drought alone is common; however, little information is available on the response of wheat genotypes to a combination of these stresses. This study investigates the response of a salt-resistant (SARC-1) and a salt-sensitive (7-Cerros) wheat genotype to drought at different growth stages under non-saline (ECe 2.1 dS m^?1) and saline soil (ECe 15 dS m^?1) conditions. Drought was applied by withholding water for 21 days at a particular growth stage viz. tillering, booting, and grain filling stages. At booting stage measurements regarding water relations, leaf ionic composition and photosynthetic attributes were made. At maturity grain yield and different yield, components were recorded. Salinity and drought significantly decreased grain yield and different yield components with a higher decrease in the case of combined stress of salinity × drought. The complete drought treatment (drought at tillering + booting + grain filling stages) was most harmful for wheat followed by drought at booting stage and grain filling–tillering stages, respectively. The salt-resistant wheat genotype SARC-1 performed better than the salt-sensitive genotype 7-Cerros in different stress treatments. A decrease in the water and turgor potentials, photosynthetic and transpiration rates, stomatal conductance, leaf K⁺, and increased leaf Na⁺ were the apparent causes of growth and yield reduction of bread wheat due to salinity, drought, and salinity × drought.     

不同生育时期增铵营养对小麦生长及氮素利用的影响

通过两年的田间试验,研究了不同生育时期增铵营养(EAN)对小麦生长和氮素利用的影响．结果表明,田间增铵营养促进了小麦植株的生长和氮素吸收．其中基肥、分蘖期、拔节期EAN提高了小麦的干物质积累量、地上部氮积累量、有效穗数、叶面积指数、叶片叶绿素含量以及小麦的籽粒产量;孕穗期EAN效果不明显;全生育期EAN在促进生长方面的效果并无明显优势,但可有效降低土壤NO3^--N的淋溶损失．与对照相比,EAN提高了氮流效率和吸收效率,但以拔节前处理最为明显．拔节期EAN主要在于改善后期的叶片光合性能,并促进同化物向籽粒的再分配,而基肥和分蘖期EAN主要在于提高有效分蘖数．     

不同生育时期增铵营养对小麦生长及氮素利用的影响

通过两年的田间试验,研究了不同生育时期增铵营养(EAN)对小麦生长和氮素利用的影响.结果表明,田间增铵营养促进了小麦植株的生长和氮素吸收.其中基肥、分蘖期、拔节期EAN提高了小麦的干物质积累量、地上部氮积累量、有效穗数、叶面积指数、叶片叶绿素含量以及小麦的籽粒产量;孕穗期EAN效果不明显;全生育期EAN在促进生长方面的效果并无明显优势,但可有效降低土壤N₃^--N的淋溶损失.与对照相比,EAN提高了氮流效率和吸收效率,但以拔节前处理最为明显.拔节期EAN主要在于改善后期的叶片光合性能,并促进同化物向籽粒的再分配,而基肥和分蘖期EAN主要在于提高有效分蘖数.     

稻-麦轮作系统土壤水解酶及氧化还原酶活性对开放式空气CO2浓度增高的响应

研究了FACE条件下(CO2浓度增加200μmol·mol^-1)水稻、小麦不同生育期0～10cm土层土壤脲酶、磷酸酶、芳基硫酸酯酶、脱氢酶活性的变化．结果表明,FACE条件下,土壤脲酶活性在冬小麦生育前期低于对照,在孕穗期高于对照;在水稻生育前期高于对照,在成熟期低于对照．磷酸单酯酶活性在冬小麦生育期高于对照;在水稻分蘖期高于对照,在生育后期(拔节期、抽穗期和成熟期)低于对照．芳基硫酸酯酶活性在小麦越冬期和孕穗期低于对照,在分蘖期和成熟期高于对照;在水稻生育期间均高于对照．脱氢酶活性在小麦和水稻的生育前期低于对照,在后期高于对照．     

Drought stress amelioration in wheat through inoculation with Burkholderia phytofirmans strain PsJN

Plant growth promoting endophytic bacteria Burkholderia phytofirmans PsJN was used to investigate the potential to ameliorate the effects of drought stress on growth, physiology and yield of wheat (Triticum aestivum L.) under natural field conditions. Inoculated and uninoculated (control) seeds of wheat cultivar Sahar 2006 was sown in the field. The plants were exposed to drought stress at different stages of growth (tillering stage and flowering stage) by skipping the respective irrigation. The results showed that drought stress adversely affected the physiological, biochemical and growth parameters of wheat seedlings. It decreased the CO₂ assimilation, stomatal conductance, relative water content, transpiration rate and chlorophyll contents in wheat. Inoculation of wheat with PsJN significantly diluted the adverse effects of drought on relative water contents and CO₂ assimilation rate thus improving the photosynthetic rate, water use efficiency and chlorophyll content over the uninoculated control. Grain yield was also decreased when plants were exposed to drought stress at the tillering and flowering stage, but inoculation resulted in better grain yield (up to 21 and 18 % higher, respectively) than the respective uninoculated control. Similarly, inoculation improved the ionic balance, antioxidant levels, and also increased the nitrogen, phosphorus, potassium and protein concentration in the grains of wheat. The results suggested that B. phytofirmans strain PsJN could be effectively used to improve the growth, physiology and quality of wheat under drought conditions.     

干旱胁迫下耐旱性小麦在不同生育期脱水素的表达分析

以2种耐旱性不同的盆栽小麦陕合6号(干旱耐受型)和郑引1号(干旱敏感型)为材料,分别在其苗期、分蘖期、拔节期、开花期对土壤实施不同程度的自然干旱胁迫和复水处理,采用SDS-PAGE和Western blotting技术研究其叶片脱水素的表达规律,探究小麦整个生长期脱水素的表达与干旱胁迫的关系.结果表明:2种小麦的脱水素均仅在干旱胁迫时表达,其中45 kD和37 kD的脱水素在2种小麦的4个发育期的叶片中均有表达,28 kD的脱水素仅在特定发育时期表达.在干旱耐受型小麦(陕合6号)中,脱水素在胁迫初期少量表达,随着胁迫程度加剧表达量急剧增加,在重度干旱胁迫下达到峰值,复水后小麦叶片中脱水素含量迅速下降;在干旱敏感型小麦(郑引1号)中,脱水素在胁迫初期大量表达,中度胁迫表达量小幅度回落,到复水1 d达到峰值,此后随着复水时间增加小麦叶片中脱水素的量逐渐下降.研究表明,小麦叶片脱水素表达与干旱胁迫程度和生育期迫密切相关,不同耐旱型小麦材料中叶片脱水素表达的差异与品种之间的干旱耐受能力密切相关.     

Effects of silicon on defense of wheat against oxidative stress under drought at different developmental stages

The effects of silicon application before sowing on the drought-induced oxidative stress and antioxidant defense in wheat (Triticum aestivum L.) were investigated. Drought stress was applied by withholding watering till sampling at booting or filling stage. Application of Si increased the water potential of drought-stressed plants at filling stage, whereas it did not at booting stage. The superoxide dismutase (SOD) activity was inhibited and peroxidase (POD) activity was enhanced by drought at booting stage, and no differences were observed due to the Si treatment. At filling stage, however, application of Si increased the SOD activity and decreased the POD activity of drought-stressed plants. The catalase (CAT) activity was slightly increased by drought only in the absence of Si and at booting stage. The activity of glutathione reductase (GR) was not greatly influenced. Application of Si did not change the contents of H₂O₂, total soluble protein and protein carbonyl of drought-stressed plants at booting stage, whereas at filling stage, it decreased the content of H₂O₂ and protein carbonyl and increased the content of total soluble protein. The content of thiobarbituric acid reactive substances (TBARS) and the activities of acid phospholipase (AP) and lipoxygenase (LOX) in drought-stressed plants were also decreased by application of Si at both stages.     

Glycine betaine application modifies biochemical attributes of osmotic adjustment in drought stressed wheat

Nineteen wheat genotypes were used to examine the effects of foliar applied glycine betaine (GB, 100 mM) on concentration of various osmolytes (such as proline, choline, GB and sucrose) under drought stress conditions. Drought stress caused a significant increase in proline content and GB content of wheat genotypes, both at maximum tillering and anthesis stages. Choline and sucrose were accumulated significantly at higher levels under stress conditions at both the stages. GB application increased the proline content and endogenous levels of GB in comparison to their stressed counterparts both at maximum tillering and anthesis stages but this increase was observed to be genotype specific. Furthermore, significant decrease in choline levels and sucrose contents of GB treated plants at anthesis stage and enhanced levels of proline questioned about involvement of GB in production of other osmolytes as well as stage specific response of wheat genotypes to GB spray. But these changes in osmolyte accumulation (OA) were not correlated with relative water content and stress tolerance index observed, under both GB sprayed and non-sprayed drought stressed conditions. So OA could not be considered as a selection criteria for drought tolerance in wheat.     

秸秆还田下氮肥运筹对水稻各生育期土壤微生物群落结构的影响

运用磷脂脂肪酸(phospholipid fatty acid,PLFA)和Biolog方法,研究了秸秆不还田不施肥(CK)、秸秆还田+尿素1(N分配:麦收后∶水稻移栽前∶分蘖期∶孕穗期=0∶6∶2∶2,T₁)、秸秆还田+尿素2(N分配:麦收后∶水稻移栽前∶分蘖期∶孕穗期=3∶3∶2∶2,T₂)、秸秆还田+沼液+尿素(N分配:麦收后∶水稻移栽前∶分蘖期∶孕穗期=3(沼液)∶3(2沼液+1尿素)∶2(尿素)∶2(尿素),T₃) 4种氮肥运筹方式对水稻各生育期(分蘖期、孕穗期、成熟期)土壤微生物群落结构的影响。结果表明: 1)T₃处理显著提高了各生育期土壤中的有效氮含量,其中成熟期有效氮含量显著高于分蘖期和孕穗期;T₁～T₃处理的有效磷和速效钾含量在各生育期均高于CK,且分蘖期的含量高于孕穗期和成熟期;稻田各生育期与各处理的交互作用对土壤有效氮、有效磷、速效钾含量均有显著影响;2)T₃能提高稻田土壤中微生物碳源代谢强度,碳水化合物、氨基酸、聚合物、羧酸是稻田土壤微生物利用的主要碳源,稻田各生育期与各处理的交互作用对微生物利用碳水化合物和羧酸的能力有显著影响;3)T₂、T₃能显著提高土壤微生物生物量;T₂处理真菌/细菌比较高,以真菌为主导,更有利于稻田土壤生态系统的稳定。表明秸秆还田同步施用氮肥(尿素或沼液)能提高土壤微生物活性,改善土壤环境。     

Changes of dehydrin profiles induced by drought in winter wheat at different developmental stages

Two cultivars of winter wheat (Triticum aestivum L.) differing in their drought tolerance (KTC86211 and ND7532) were subjected to a progressive soil water stress and recovery at four developmental stages. Dehydrins with molecular masses of 45 and 37 kDa were constitutively accumulated during all stages in both genotypes. The 28 kDa dehydrin accumulated exclusively at seedling stage in both genotypes. The 49 and 40 kDa dehydrins accumulated at both tillering and jointing stages but showed a genotype-specific pattern. The content of most dehydrins increased with decreased soil moisture and then decreased during recovery. These results suggest that accumulation pattern of dehydrins during water stress was related to the genotype and developmental stage.