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灌浆期干旱对小麦籽粒品质影响研究进展

来源:用户上传      作者:宋维富 杨雪峰 赵丽娟 刘东军 宋庆杰 张春利 辛文利 肖志敏

  摘要  小麦品质决定小麦的最终用途,也是小麦品种品质分类的主要依据。为了探讨灌浆期干旱对小麦品质影响的内在原因,综述了灌浆期干旱胁迫对小麦籽粒品质影响的研究进展,并进一步探讨了抗旱育种策略,以期为培育品质稳定型小麦品种提供有用信息。
  关键词  小麦;品质;灌浆期;干旱;抗旱育种
  中图分类号  S  512文献标识码  A
  文章编号  0517-6611(2020)04-0018-02
  doi:10.3969/j.issn.0517-6611.2020.04.005
  开放科学(资源服务)标识码(OSID):
  Review on Effects of Drought at Grain Filling Stage on Grain Quality of Wheat
  SONG Wei-fu,YANG Xue-feng,ZHAO Li-juan et al  (Crop Resources Institute of Heilongjiang Academy of Agricultural Sciences,Harbin,Heilongjiang 150086)
  Abstract  Wheat grain quality determines the end-use of wheat and is also the main basis for quality classification of wheat varieties.In order to investigate the major cause for effects of post-anthesis drought at grain filling stage on wheat quality,the impact of  drought at grain-filling stage on wheat quality was reviewed in this paper.Futhermore,drought-resistant breeding strategies for drought at grain filling satge were discussed.The aims of this review were to provide useful information for wheat quality improvement in breeding programs.
  Key words  Wheat; Quality;Grain filling stage;Drought;Drought-resistant breeding
  在优质强筋小麦育种中,小麦品质性状可划分为营养品质性状和加工品质性状。小麦籽粒的营养品质由土壤条件决定,而加工品质主要受气候条件影响[1]。小麦花后灌浆期的环境条件胁迫对加工品质的影响最大[2-5]。灌浆期干旱能够显著影响小麦的新陈代谢进程,使灌浆期缩短,细胞死亡进程加速,导致籽粒中蛋白质组分和淀粉颗粒大小改变,是影响小麦产量和品质的2个主要气候因素[6-8]。在小麦生产中干旱可单独发生,亦可与各种环境胁迫同步发生,不仅高温和强光照可引起干旱,而且贫瘠的土壤条件限制植物根的渗透性也可引起干旱,所以干旱通常与各种环境因素互作影响小麦品质[9-11]。高温条件下干旱对小麦各种生理代谢参数指标的影响程度要高于低温条件下,干旱与高温同时发生影响显著[12-14]。
  小麦加工品质与其最终用途密切相关,是划分小麦品种品质类型的重要依据。小麦育种和生产实践表明,小麦品质受遗传因素、环境条件及遗传与环境互作的影响[4,9]。其中灌浆期高温和干旱胁迫是影响小麦品质的主要因素[5,14-15]。为了探讨灌浆期干旱对小麦品质影响的内在原因,拟综述灌浆期干旱胁迫对小麦籽粒品质影响的研究进展,并进一步探讨抗旱育种策略,以期为培育品质稳定型小麦品种提供有用信息。
  1  灌浆期干旱胁迫对小麦籽粒品质影响研究
  在世界范围内,小麦灌浆期遇到高温和干旱同步发生条件非常普遍[16-19]。Irmak等[20]和Don等[4]大量研究发现,当灌浆期遇到大于30 ℃条件形成高温胁迫时,醇溶蛋白和不溶性谷蛋白聚合体相对含量均增加,但增加幅度不同,导致2种蛋白比例变化,使面团强度大幅度下降。在高温胁迫条件下高分子麦谷蛋白亚基5+10较2+12亚基耐热性好,品质较为稳定[2,20]。研究表明,高温胁迫使小麦灌浆期缩短,籽粒干瘪,降低了籽粒中淀粉和蛋白质的总体产量,但对淀粉降低幅度大于蛋白质,致使蛋白质相对含量提高[21-23]。高温胁迫对小麦品质的影响不仅取决于品种间的抗(耐)热能力差异,很大程度上还取决于灌浆期不同阶段所遇到的温度条件[5,24-26]。
  研究认为,灌浆期干旱是影响小麦产量和品质的主要气候因素[27-29]。干旱不仅可由高溫和强光照引起,而且贫瘠的土壤条件限制植物根的渗透性也可引起干旱[30]。小麦生产中干旱可单独发生,亦可与各种环境胁迫同步发生,所以干旱通常与各种环境因素互作影响小麦品质。王月福等[31]研究表明,适宜的土壤水分是小麦高产和优质的保证,干旱和渍水都会影响小麦产量提高和品质改善。灌浆期不同阶段的干旱胁迫对籽粒蛋白含量、沉降值、湿面筋含量的提高幅度不同,灌浆早期影响较大[31-32]。Guttieri等[33-35]和王立秋等[36]研究认为,在严格控水条件下灌浆期干旱可显著影响小麦品质。干旱条件下小麦产量下降,籽粒蛋白质相对含量提高,但蛋白质总量下降。Gooding等[37]研究认为灌浆期干旱可使籽粒蛋白含量增加,特别是低分子量蛋白增加显著。范雪梅等[38]利用2个不同品质类型小麦品种研究了花后干旱对其品质形成的影响,结果发现干旱处理对小麦籽粒麦谷蛋白积累的影响大于醇溶蛋白,导致麦谷蛋白/醇溶蛋白的比值明显下降,但是灌浆期干旱提高了籽粒干、湿面筋含量、沉降值和降落值。Guttieri等[35]研究表明灌浆期干旱增加了面团揉混峰高和出峰时间,但是降低了耐揉性,并使籽粒的出粉率降低。Ozturk等[32]研究发现干旱分别使籽粒蛋白含量、SDS沉降值、湿面筋含量提高了8.3%、8.7%和10.8%。许振柱等[39]研究指出,严重水分亏缺降低籽粒容重、面筋含量、沉降值、面团形成时间、稳定时间和评价值,但一定程度土壤水分亏缺反而促其升高。Frank等[40]研究认为,灌浆期干旱对小麦品质的影响程度不仅取决于品种的抗旱性,在很大程度上还取决于灌浆期不同阶段所遇到的干旱胁迫程度以及干旱持续的时间。   2  抗旱性育种
  培育抗旱性品種是减少干旱胁迫对小麦品质影响最经济和有效的方法。提高品种抗旱能力对于充分发挥小麦品种,特别是强筋类小麦品种品质遗传潜力和保证其品质稳定性非常关键。目前研究认为,植物防御干旱胁迫的机制主要有3种,即避旱、御旱和耐旱。其中,御旱性和耐旱性又被统称为抗旱性[41]。Myers等[42]、Mambani等[43]认为小麦根是提高抗旱性的重要部位,增加种子根的长度有利于小麦吸收土壤深层的水分并提高利用率,保持体内水分平衡,抵御干旱。Morgan[44]研究表明具有高的渗透调节能力的小麦品种对干旱的适应性更强。东北春麦区小麦育种者针对“十年九春旱”特定生态条件,以小麦苗期和灌浆期抗旱性存在着高度正相关为依据,通过选择苗期发育较慢、抗旱性较强的光敏型小麦品种,增加根冠比,将躲旱和抗旱机制有机结合,在一定程度上实现了小麦品种苗期和灌浆期抗旱能力的同步选择。根据多年育种实践发现,利用肥水耦合和以肥调水原理,实施氮素后移等措施在一定程度上可提高小麦品种灌浆期的抗旱能力。近年来,很多学者开发出的多个与抗旱性状相关的数量性状基因座(QTL),现逐步被用到提高小麦灌浆期抗旱能力育种之中。
  3  展望
  综上所述,可以认为小麦灌浆期高温和干旱均可显著影响小麦品质。其中,小麦灌浆期遇到干旱可使籽粒贮藏蛋白含量相对增加,蛋白质组分比例发生改变,面团耐揉性降低,导致各品质参数不均衡,进而影响面粉的最终用途。同时干旱往往与高温条件胁迫互作,进一步增加了小麦品种品质遗传潜力表达的影响程度。选育抗旱性较好,特别是灌浆期抗旱性较强的品种,对于充分发挥优质强筋小麦新品种品质潜力非常重要。
  上述研究结果对于小麦品质育种,特别是对于优质强筋小麦育种具有重要参考价值。至于土壤肥力和气候条件变化对小麦营养品质和加工品质的影响程度,以及小麦灌浆期高温和干旱何者为影响优质强筋小麦品种品质潜力表达程度的主导因子等问题,还有待深入研究。
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