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不同品质类型小麦HMW-GS积累及GMP粒度分布对灌浆期高温胁迫的响应

Response of HMW-GS Accumulation and GMP Particle Size Distribution from Different Wheat Genotypes to Heat Stress During Grain Filling

【作者】 孟范玉

【导师】 王振林;

【作者基本信息】 山东农业大学 , 作物栽培学与耕作学, 2010, 硕士

【摘要】 本研究于2007~2009年在山东农业大学试验农场进行,选用不同品质类型的小麦藁城8901(GC8901,强筋)、济南17(JN17,强筋)、山农1391(SN1391,弱筋)和豫麦50(YM50,弱筋)为供试品种,通过改变籽粒灌浆阶段温度,系统研究小麦籽粒氮素转移效率、蛋白质组分含量、谷蛋白大聚合体积累、粒度分布及高分子量谷蛋白亚基积累的变化,明确高温影响籽粒蛋白质形成的生理生化机理。主要研究结果如下:1.灌浆期高温胁迫对小麦产量的影响灌浆期高温胁迫降低了籽粒穗粒数、千粒重,从而造成产量下降。高温处理对亩穗数影响较小,对粒重影响比对穗粒数影响大,说明高温条件下产量的下降主要是由于粒重下降引起的。灌浆前期、中期、后期高温处理后产量降幅分别为14.25%、17.32%、8.37%(山农1391)和17.87%、17.40%、23.38%(豫麦50);22.67%、27.46%、19.05%(藁城8901)和19.31%、12.11%、25.72%(济南17),不同品种处理间差异显著。前期高温处理后不同品种产量下降顺序为藁城8901>济南17>豫麦50>山农1391;中期高温处理后产量下降顺序为藁城8901>山农1391>豫麦50>济南17,后期高温处理后产量下降顺序为济南17>豫麦50>藁城8901>山农1391,说明不同品种对不同时期高温胁迫的响应存在差异。中期高温处理对豫麦50和济南17产量影响较前期和后期大,后期高温处理对山农1391和藁城8901的影响较前期和中期大。2.灌浆期高温胁迫对氮素转移效率和蛋白质含量的影响花后不同时期高温处理均降低了氮素转移效率,减少了氮素从植株向籽粒的转移,因而植株需要从环境中吸收更多的氮素来满足籽粒生长发育的需要。灌浆前期、中期、后期高温处理后弱筋小麦氮素转移效率降幅为8.05%、3.23%、9.36%(山农1391)和9.89%、13.01%、、8.20%(豫麦50);强筋小麦氮素转移效率降幅为4.63%、2.49%、6.60%(藁城8901)和14.25%、17.73%、7.63%(济南17)。在开花后各个时期对小麦进行高温处理,籽粒蛋白质含量均呈现升高的趋势。灌浆前期、中期、后期高温处理后不同品种籽粒蛋白质含量上升幅度不一致。高温胁迫对弱筋小麦山农1391和强筋小麦藁城8901蛋白质含量影响顺序为:后期>前期>中期,对弱筋小麦豫麦50和强筋小麦济南17的影响顺序为中期>前期>后期。山农1391和藁城8901对后期高温胁迫响应比较敏感,而豫麦50和济南17对中期高温胁迫比较敏感。说明不同品种对不同时期温度处理的适应性不同。3.灌浆期高温胁迫对蛋白质组分的影响对小麦开花后各个时期进行高温处理,籽粒清蛋白、球蛋白、醇溶蛋白以及可溶性谷蛋白含量均提高,谷蛋白/醇溶蛋白比值下降。不同时期对不同品种的影响存在差异,花后24~27天高温胁迫对弱筋小麦山农1391和强筋小麦藁城8901蛋白质组分的影响比花后7~9天和14~17天的影响大,花后14~17天高温胁迫对弱筋小麦豫麦50和强筋小麦济南17蛋白质组分含量影响比花后7~9天和24~27天高温处理影响大。4.灌浆期高温胁迫对谷蛋白大聚合体含量的影响灌浆期谷蛋白大聚合体(GMP)含量呈现先下降后上升成熟期又略有下降的趋势。山农1391和藁城8901GMP含量在花后25天出现最低值,豫麦50和济南17GMP含量在花后20天左右出现最低值。灌浆前、中、后期高温处理均提高了4个品种GMP含量,其中后期高温胁迫对弱筋小麦山农1391和强筋小麦藁城8901GMP含量的影响比对豫麦50和济南17的影响大,而中期高温处理后弱筋小麦豫麦50和强筋小麦济南17GMP含量上升幅度比弱筋小麦山农1391和强筋小麦藁城8901大。不同时期高温处理间比较,前期高温对GMP含量影响较小。5.灌浆期高温胁迫对谷蛋白大聚合体颗粒粒度分布的影响谷蛋白大聚合体的体积和表面积分布呈双峰曲线,数目分布呈单峰曲线变化。GMP<10μm颗粒所占体积为21.80%~47.91%,10~100μm颗粒和>100μm颗粒对体积的贡献分别为33.85%~58.81%和9.49%~31.95%。体积加权平均粒径D(4.3)为33.64~78.99μm。<10μm颗粒所占表面积为81.30%~92.81%,10~100μm颗粒和>100μm颗粒对表面积的贡献分别为6.61%~15.30%和0.65%~10.91%,表面积加权平均粒径D(3.2)为5.97~ 11.25μm。<10μm颗粒数目所占比例为99.88%~99.96%,>10μm颗粒数目所占比例为0.06%~0.12%。灌浆期高温胁迫降低了<10μm颗粒所占体积、表面积和数目比例,显著提高了10~100μm颗粒和>100μm颗粒所占体积、表面积和数目比例,同时体积加权平均粒径D(4.3)和表面积加权平均粒径D(3.2)也增大。不同时期高温处理间比较,花后7~9天高温胁迫对GMP粒度分布影响较小;花后24~27天高温胁迫对弱筋小麦山农1391和强筋小麦藁城8901GMP粒度分布的影响比对豫麦50和济南17的影响大,而花后14~17天高温处理后弱筋小麦豫麦50和强筋小麦济南17GMP粒度分布变化幅度大,较另外两个品种差异显著。6.灌浆期高温胁迫对高分子量谷蛋白亚基含量的影响灌浆前期高温胁迫后4个品种高低分子量谷蛋白亚基含量均提高。灌浆中期高温胁迫降低了山农1391和藁城8901 HMW-GS含量但提高了LMW-GS含量,从而使H/L降低;豫麦50和济南17 HMW-GS含量和LMW-GS含量在灌浆中期高温处理后均升高,但LMW-GS含量上升幅度较大,因而H/L比值也降低。后期高温胁迫后山农1391和藁城8901 HMW-GS含量和LMW-GS含量均升高,豫麦50和济南17 HMW-GS含量降低,LMW-GS含量上升,4个品种H/L比值均降低。强筋小麦藁城8901和济南17位于D位点上的不同类型的亚基对中、后期高温胁迫的响应不同,x-亚基比y-亚基更敏感。弱筋小麦山农1391和豫麦50位于B位点亚基对中、后期高温胁迫的响应不同,y-亚基比x-亚基对高温胁迫的响应更敏感。

【Abstract】 The effects of heat stress during grain filling on nitrogen remobilization efficiency (NRE, %), protein content, glutenin macro polymer (GMP) particle size distribution and high molecular weight glutenin subunit (HMW-GS) accumulation were studied, using four winter wheat (Triticum aestivum L.) cultivars, Shannong1391 (SN1391), Yumai50 (YM50), Gaocheng8901 (GC8901) and Jinan17 (JN17) with different HMW-GS compositions, which were grown at Tai’an Experimental Station of Shandong Agriculture University, Tai’an, Shandong, P R China (36°09’N, 117°09’E) during the 2007~2009 growing seasons. Three heat stress treatments were conducted with transparent plastic increasing temperature sheds from 7 days after anthesis (DAA, T1), 14 DAA (T2) and 24 DAA (T3), respectively, and each treatment last for three days during grain filling. The main results were as follows:1. Effects of heat stress during grain filling on grain yield and yield components of wheat cultivarsThe results showed that there was significant effect of heat stress on grain yield and yield components of wheat. Heat stress during grain filling reduced grain yield, grain numbers per spike, and kernel weight, but not spike number. The damage degree of heat stress on kernel weight was higher than grain numbers per spike, the results suggested that the decrease of grain yield was result from the reduction of kernel weight under heat stress conditions. For the four cultivars, the reduction of grain yield after T1, T2 and T3 treatments was found with the following rank: 14.25%、17.32%、8.37% for SN1391, 17.87%、17.40%、23.38% for YM50, 22.67%、27.46%、19.05% for GC8901 and 19.31%、12.11%、25.72% for JN17.The conclusion also showed that there was a difference in reduction extent of grain yield and grain weight from the four different wheat cultivars under heat stress. The reduction of grain yield after T1 treatment was found with the following rank: GC8901> JN17> YM50> SN1391. And GC8901> SN1391> YM50> JN17 after T2 treatment, JN17> YM50> GC8901>SN1391 after T3 treatment. The damage degree of heat stress on grain yield of YM50 and JN17 at middle grain filling stages was higher than early and later filling stages. On the other hand, heat stress at later grain filling stages affects grain yield of SN1391 and GC8901 more severity.2. Effects of heat stress during grain filling on nitrogen remobilization efficiency NRE (%) and protein content in grainsThe NRE (%) varied with heat stress treatments for all the cultivars. In general, comparison of NRE (%) between non-stressed and heat stress conditions, showed a systematic difference. No significant differences of NRE (%) among cultivars were found in the normal condition while the differences were significant under heat stress condition. All heat stress treatments negatively affected the overall remobilization efficiency of nitrogen. For SN 1391 and GC 8901, the largest reduction of NRE (9.36% and 6.60%, respectively) was found in T3 treatment with the following rank: CK>T2>T1>T3. Interestingly, there is a similar trend for both YM 50 and JN 17 under the T2 condition.Heat stress during grain filling improved protein content, there was a difference in protein content from the four different wheat cultivars. All heat stress treatments positively affected the protein content. For SN 1391 and GC 8901, the highest r protein content was found in T3 treatment with the following rank: CK>T2>T1>T3. On the other hand, a rank of CK>T2>T1>T3 was fond in both YM 50 and JN 17. Since growing conditions are the same, this can be due to the HMW-GS composition differences.3. Effects of heat stress during grain filling on protein components in grainsThe results showed that albumin, globulin, gliadin and soluble glutenin were significantly increased by all heat stress treatments. But glutenin/gliadin ratio was reduced. The effects of heat stress on protein components of SN1391 and GC8901 at later grain filling stages was higher than early and middle filling stages. On the other hand, heat stress at middle grain filling stages affects grain yield of YM50 and JN17 more severity.4. Effects of heat stress during grain filling on glutenin macro polymer (GMP) contentThere was a trend of“V”trend in GMP content during grain filling. The lowest content of SN1391 and GC8901 was found at 25 DAA, and for YM50 and JN17, the lowest content was found at 20DAA. The results showed that heat stress during grain filling positively affected GMP content of the four cultivars. And there was a similar trend like protein content under heat stress conditions.5. Effects of heat stress during grain filling on GMP size distributionThe glutenin particle sizes from the four different wheat genotypes ranged from 0.4 and to ca.325μm. The distribution of GMP volume and surface area showed a pattern of two-peak curve, and the GMP number distributed in the pattern of single-peak curve. The number of GMP particle mainly composed by <10μm particle (accounting for 99.88%~99.96%), the volume distribution percentage of <10μm particle ranged from 21.80% to 47.91%. 33.85%~58.81% and 9.49%~31.95% for 10~100μm and >100μm particle. On the other hand, the range of surface area <10μm, 10~100μm and >100μm was 81.30%~92.81%, 6.61%~15.30% and 0.65%~10.91%, 8901, the highest r protein content was found in T3 treatment with the following rank: CK>T2>T1>T3. On the other hand, a rank of CK>T2>T1>T3 was fond in both YM 50 and JN 17. Since growing conditions are the same, this can be due to the HMW-GS composition differences.3. Effects of heat stress during grain filling on protein components in grainsThe results showed that albumin, globulin, gliadin and soluble glutenin were significantly increased by all heat stress treatments. But glutenin/gliadin ratio was reduced. The effects of heat stress on protein components of SN1391 and GC8901 at later grain filling stages was higher than early and middle filling stages. On the other hand, heat stress at middle grain filling stages affects grain yield of YM50 and JN17 more severity.4. Effects of heat stress during grain filling on glutenin macro polymer (GMP) contentThere was a trend of“V”trend in GMP content during grain filling. The lowest content of SN1391 and GC8901 was found at 25 DAA, and for YM50 and JN17, the lowest content was found at 20DAA. The results showed that heat stress during grain filling positively affected GMP content of the four cultivars. And there was a similar trend like protein content under heat stress conditions.5. Effects of heat stress during grain filling on GMP size distributionThe glutenin particle sizes from the four different wheat genotypes ranged from 0.4 and to ca.325μm. The distribution of GMP volume and surface area showed a pattern of two-peak curve, and the GMP number distributed in the pattern of single-peak curve. The number of GMP particle mainly composed by <10μm particle (accounting for 99.88%~99.96%), the volume distribution percentage of <10μm particle ranged from 21.80% to 47.91%. 33.85%~58.81% and 9.49%~31.95% for 10~100μm and >100μm particle. On the other hand, the range of surface area <10μm, 10~100μm and >100μm was 81.30%~92.81%, 6.61%~15.30% and 0.65%~10.91%, another meaningful trait also found in our experiment was that GS belongs to y-type from SN 1391 and YM 50 reacted to heat stress more sensitively than x-type, and heat stress affected x-type from GC 8901 and JN 17 rapidly than y-types. These results may indicate the essential diversification in HMW-GS accumulation after heat stress is that HMW-GS at different locus have a variety response to the ambient temperature changes.

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