灌浆期高温与干旱胁迫对小麦籽粒淀粉合成关键酶活性及淀粉积累的影响

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胡阳阳, 卢红芳, 刘卫星, 康娟, 马耕, 李莎莎, 褚莹莹, 王晨阳. 灌浆期高温与干旱胁迫对小麦籽粒淀粉合成关键酶活性及淀粉积累的影响. 作物学报, 2018, 44(4): 591-600
HU Yang-Yang, LU Hong-Fang, LIU Wei-Xing, KANG Juan, MA Geng, LI Sha-Sha, CHU Ying-Ying, WANG Chen-Yang. Effects of High Temperature and Water Deficiency during Grain Filling on Activities of Key Starch Synthesis Enzymes and Starch Accumulation in Wheat. ACTA AGRONOMICA SINICA, 2018, 44(4): 591-600 复制到剪切板

DOI:10.3724/SP.J.1006.2018.00591
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作物学报编辑部

灌浆期高温与干旱胁迫对小麦籽粒淀粉合成关键酶活性及淀粉积累的影响

胡阳阳^1,^**, 卢红芳^1,^**, 刘卫星¹, 康娟¹, 马耕¹, 李莎莎¹, 褚莹莹¹, 王晨阳^1,^2,^*

¹河南农业大学, 河南郑州 450002

²国家小麦工程技术研究中心, 河南郑州 450002

^* 通信作者(Corresponding author): 王晨阳, E-mail: xmzxwang@163.com ^**同等贡献(Contributed equally to this work)

作者简介: E-mail: 18238802035@163.com

收稿日期:2017-07-02 接受日期:2018-01-08 网络出版日期:2018-01-23

基金:本研究由国家重点研发计划项目(2017YFD0300204), 国家科技支撑计划项目(2015BAD26B01)和河南省小麦产业技术体系建设专项(S2010-01-G07)资助

摘要

高温与干旱是小麦灌浆期的主要胁迫逆境, 影响小麦同化物的生成及其在籽粒中的积累。本文以郑麦366为试验材料, 采用盆栽和人工气候室模拟相结合的方式, 研究了灌浆期高温(HT)、干旱(DS)和高温干旱复合胁迫(HT+DS)对小麦籽粒淀粉合成关键酶活性、淀粉及其组分的影响。于花后10 d将长势均匀一致的盆栽小麦转移至人工气候室, 至小麦成熟。人工气候室设适温(昼25℃/夜15℃)和高温(昼32℃/夜22℃)两种温度模式, 每种温度模式下又设置正常水分(土壤相对含水量75%左右)和轻度干旱胁迫(土壤相对含水量50%左右)两个土壤水分处理, 以适温、正常水分处理为对照。与对照相比, HT、DS及HT+DS条件下籽粒可溶性淀粉合酶(SSS)和结合态淀粉合酶(GBSS)活性在胁迫初期显著升高, 之后迅速下降; 淀粉分支酶(SBE)、ADPG焦磷酸化酶(AGPase)和蔗糖合酶(SS)活性在小麦籽粒整个灌浆过程中均低于对照; 上述酶活性还受高温与干旱互作的显著影响。HT、DS及HT+DS逆境下, 小麦籽粒淀粉积累速率减小, 籽粒直、支链淀粉和总淀粉含量下降, 生育期缩短, 粒重和产量降低。其中, HT的影响大于DS, 复合胁迫的影响大于单一胁迫。相关分析表明, SSS和GBSS活性与籽粒直、支链淀粉和总淀粉含量在多数测定时期呈正相关( P<0.01), AGPase、SBE和SS活性在灌浆后期(花后22~26 d)与籽粒直、支链淀粉和总淀粉含量呈正相关( P<0.01)。表明高温干旱胁迫通过淀粉合成关键酶而影响籽粒淀粉的合成与积累, 最终影响小麦产量和品质。

关键词: 冬小麦; 复合胁迫; 高温; 干旱; 淀粉合成酶; 淀粉积累

Effects of High Temperature and Water Deficiency during Grain Filling on Activities of Key Starch Synthesis Enzymes and Starch Accumulation in Wheat

HU Yang-Yang^1,^**, LU Hong-Fang^1,^**, LIU Wei-Xing¹, KANG Juan¹, MA Geng¹, LI Sha-Sha¹, CHU Ying-Ying¹, WANG Chen-Yang^1,^2,^*

¹Henan Agricultural University, Zhengzhou 450002, Henan, China

²National Wheat Engineering Research Center, Zhengzhou 450002, Henan, China

Fund:This study was supported by the National Key Research and Development Program of China (2017YFD0300204), the National Key Technologies R&D Program of China (2015BAD26B01), and the earmarked fund for Modern Wheat Industrial Technology System of Henan Province (S2010-01-G07).

Abstract

High temperature and drought, the major abiotic stresses during grain filling of winter wheat, have great influence on photosynthate accumulation in grains. In this study, we evaluated the effects of high temperature, drought and the compound stress at grain-filling stage on the activities of key starch synthesis enzymes and the contents of starch and its components in grains of winter wheat cultivar ‘Zhengmai 366’. The pot-grown plants were moved from field to the climate chamber at 10 days after anthesis and exposed to the stresses until maturity. The climate chamber was desired with the normal growth condition (25°C/15°C, day/night) and high temperature (HT) stress (32°C/22°C, day/night) treatments. Under each temperature treatment, the soil moisture was adjusted to ~75% (normal) and ~50% (drought stress, DS), respectively. The normal temperature and soil moisture condition was taken as the control. Compared with the control, HT, DS, and HT+DS significantly increased activities of soluble starch synthase (SSS) and granule bound starch synthase (GBSS) at early period of stress and rapidly declined them subsequently. However, all stress treatments reduced the activities of starch branching enzymes (SBE), ADPG pyrophosphorylase (AGPase) and sucrose synthase (SS) during the whole grain-filling stage. The interaction between HT and DS was significant on all enzyme activities. Under HT, DS and HT+DS conditions, starch accumulation rate of grain, and the contents of amylose, amylopectin and total starch decreased and growth duration shortened, leading to reduced grain weight and yield at maturity. HT showed greater influence than DS, and the effect of compound stress was larger than that of each individual stress. Correlation analysis revealed that the activities of SSS and GBSS were positively correlated with the contents of amylose, amylopectin and total starch at most stages ( P < 0.01), whereas, those of SBE, AGPase and SS at late grain-filling stage (22-26 days after anthesis) ( P < 0.01). These results indicate that high temperature and drought stresses cause yield loss and degrade grain quality by affecting activities of key starch synthesis enzymes and starch accumulation in wheat grains.

Keyword: winter wheat; compound stress; high temperature; drought; starch accumulation; starch synthesis enzymes

Show Figures

淀粉作为小麦籽粒主要组成成分, 其含量及积累动态决定着小麦产量和品质状况。籽粒淀粉积累受到一系列淀粉合成关键酶的调节。有研究表明, 蔗糖合酶(SS)、ADPG焦磷酸化酶(AGPase)、可溶性淀粉合酶(SSS)、结合态淀粉合酶(GBSS)和淀粉分支酶(SBE)是控制淀粉合成代谢的关键酶^{[1, 2]}, 在小麦籽粒淀粉合成与积累中起重要作用。其中, 小麦籽粒直链、支链和总淀粉的积累速率与SBE、SSS、GBSS和AGPase活性均存在显著的相关性^{[3, 4]}。

温度和水分是影响小麦生长发育的主要环境因子, 尤其是在生育后期, 对产量形成和品质优劣有重要的影响。花后高温和干旱往往导致小麦穗粒数减少, 粒重和产量显著降低^{[5, 6]}。黄淮海麦区灌浆后期还经常遭遇高温与干旱复合胁迫的危害, 甚至形成干热风天气, 导致小麦减产10%~30% ^[7]。随着全球气候变暖, 后期高温、干旱及其复合胁迫对小麦生产的影响呈加重趋势。

针对花后高温、干旱单因子胁迫对小麦籽粒灌浆和品质的影响已有较多研究。花后不同时期的高温胁迫均会抑制GBSS、SSS、SBE、AGPase的活性, 导致淀粉积累发生改变; 胁迫发生的越早, 对淀粉积累影响越大^[8]。干旱胁迫降低籽粒SSS、SBE、AGPase的活性, 减少籽粒直、支链淀粉含量, 导致总淀粉含量显著下降^{[9, 10]}。当小麦花后遭遇极端高温(35℃以上)和干旱双重胁迫时, 会导致灌浆期明显缩短, 籽粒直链、支链淀粉和总淀粉含量降低, 影响淀粉的糊化特性, 并使籽粒蛋白质积累量和产量下降, 而且高温与干旱存在显著的互作效应^{[11, 12, 13, 14]}。然而, 小麦籽粒灌浆期较长, 黄淮麦区一般35~45 d, 花后高温发生时间存在较大的不确定性, 以往对短暂高温(2~4 d)的模拟与生产实际出入较大, 不仅夜间温度难以确定, 而且在返还大田后受到整个灌浆期气候条件的显著影响, 难以定量描述。本研究设置了灌浆期不同昼/夜温度模式和干旱胁迫处理, 以观测小麦籽粒淀粉合成关键酶活性及淀粉积累的变化, 探讨昼/夜高温模式与干旱复合胁迫下小麦籽粒产量形成机制, 为小麦抗逆保优栽培提供理论依据。

1 材料与方法

1.1 试验材料和试验设计

2015— 2016和2016— 2017年度在河南农业大学科教试验园区, 选用高产强筋小麦品种郑麦366进行盆栽试验, 该品种蛋白质含量16%, 湿面筋含量33%。试验用土为耕层土壤(pH 8.0), 含有机质18 g kg^-1、全氮1 g kg^-1、碱解氮58 mg kg^-1、速效磷68 mg kg^-1、速效钾205 mg kg^-1, 田间持水量为26%。试验用盆的高27 cm, 盆口直径24 cm, 每盆装土10 kg; 播种前将装土盆钵埋于大田, 使盆内土面与大田基本齐平。于2015年10月20日和2016年10月17日播种, 播种前每盆施纯氮1.15 g、P₂O₅ 1.35 g和K₂O 1.15 g; 三叶期定苗, 每盆12株; 拔节期结合浇水, 每盆追施纯氮1.15 g。

开花期选择同一天开花、大小均匀的麦穗挂牌标记, 于开花后第10天, 选长势均匀一致的盆栽小麦转移至人工气候室进行逆境处理, 小麦生长至成熟。设置人工气候室光照时间为8:00— 18:00, 自然光照强度为500 μ mol m^-2 s^-1, 自动控温控湿, 温度变幅为± 1℃, 相对湿度为50%± 10%。模拟适温(25℃/15℃)和高温(32℃/22℃)两种昼/夜温度模式, 在每种温度模式下设正常水分处理(土壤相对含水量75%左右)和轻度干旱处理(土壤相对含水量50%左右), 形成4个温度水分处理组合, 即对照(CK)、干旱(DS)、高温(HT)和高温干旱复合胁迫(HT+DS), 每处理组合15盆。对于干旱处理自高温处理前7 d开始控制浇水, 采用称重法与土壤水分测定仪TDR300相结合的方法测定土壤含水量, 确保高温处理时达到目标含水量。

逆境胁迫处理前一天取样一次, 其后每4 d取样一次, 直至小麦成熟。每次取15个单茎(穗), 一部分籽粒样品于105℃杀青20 min, 70℃烘至恒重, 用高速万能粉碎机(天津泰斯特仪器有限公司)磨粉, 用以测定直链和支链淀粉含量。另一部分样品经液氮速冻后保存于-80℃超低温冰箱, 用于测定淀粉合成关键酶活性。

1.2 淀粉合成相关酶活性测定方法

采用淀粉合成酶试剂盒(苏州科铭生物技术有限公司, 货号为SS-2-Y、GBSS-2A-Y、SSS-S-Y、AGP-2A-Y和SBE-2-Y)分别测定SS、GBSS、SSS、AGPase和SBE活性, 其中GBSS、SSS和AGPase活性用每克植物鲜重每分钟催化产生NADPH的量(nmol)来表示, SS活性用每克植物鲜重每分钟催化产生的蔗糖量(μ g)来表示, SBE活性以波长660 nm下的吸光度下降百分率表示, 每克植物鲜重在反应体系中每降低1%碘蓝值为一个酶活力单位。

1.3 籽粒淀粉含量及淀粉积累速率测定方法

采用双波长法^[15]测定直链、支链淀粉含量, 两者之和为总淀粉含量。

^{${{R}_{n}}=\frac{{{C}_{n+4}}-{{C}_{n-4}}}{8}$ [11]}

式中, n为开花后天数, R_n为第n天的淀粉积累速率, C_n₊₄和C_n_-4分别为第n+4天和第n-4天的淀粉积累量。

1.4 数据处理和做图

用SPSS 21.0软件进行方差分析和相关分析, 用Microsoft Excel 2003绘制折线图。

2 结果与分析

2.1 高温与干旱胁迫对小麦籽粒淀粉合成关键酶活性的影响

2.1.1 高温与干旱胁迫下GBSS活性变化灌浆期不同处理小麦籽粒GBSS活性变化均为单峰曲线, CK和DS处理的GBSS活性于花后26 d达到峰值, 而HT及HT+DS处理峰值分别提前至花后18 d和14 d (图1-A)。总的来看, 无论单因子或复合胁迫, 胁迫初期(花后14 d) GBSS活性均高于CK, 自花后18 d开始, HT和HT+DS处理GBSS活性迅速降低, 与CK差距很大, 而DS处理在花后30 d才显著低于CK (P< 0.05)。

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	图1 灌浆期不同胁迫处理小麦籽粒淀粉合成关键酶活性的变化 HT: 高温; DS: 干旱; HT+DS: 高温与干旱复合胁迫。Fig. 1 Dynamics of starch synthesis key enzyme activities during grain filling under different stress treatments DS: drought stress; HT: high temperature stress; HT+DS: compound stress of HT and DS.

2.1.2 高温与干旱胁迫下SSS活性变化不同处理小麦籽粒SSS活性的变化趋势均呈单峰曲线(图1-B)。CK、DS、HT及HT+DS处理SSS活性分别于花后26 d、22 d、18 d、14 d达最大值, 表明HT、DS及HT+DS均使SSS活性峰值提前。与CK相比, 逆境胁迫下在花后10~22 d提高了籽粒SSS活性, 但之后开始显著低于对照(P< 0.05)。

2.1.3 高温与干旱胁迫下SBE活性变化在胁迫4 d (花后14 d)时, HT处理使SBE活性略高于CK, 之后迅速降低, 平均比CK低23.3% (2015— 2016年度)和31.2% (2016— 2017年度); 而DS及HT+DS处理全程导致SBE活性降低, 尤其是从花后14 d开始, 2015— 2016年度平均分别下降28.8%和46.8%, 2016— 2017年度平均下降23.3%和41.2% (图1-C)。

2.1.4 高温与干旱胁迫下AGPase活性变化小麦籽粒AGPase活性变化趋势与GBSS活性一致, CK和DS处理AGPase活性于花后22 d达到最大值, 而HT及HT+DS处理的峰值分别提前到花后18 d和14 d。花后18 d, HT处理AGPase活性略高于对照, 之后各处理AGPase活性均迅速降低, HT+DS处理酶活性最低, 其次是HT处理, 再次是DS处理(图1-D)。说明AGPase活性受高温胁迫的影响大于干旱胁迫, 并且复合胁迫表现出显著的叠加效应。

2.1.5 高温与干旱胁迫下SS活性变化不同处理籽粒SS活性的变化均呈单峰曲线, 除DS处理SS活性在花后14 d达到最大值, 其他处理均于花后18 d达到峰值, 之后迅速下降(图1-E)。HT及HT+DS处理使小麦籽粒SS活性显著降低, 2015— 2016年度平均比CK低23.8%和32.1%, 2016— 2017年度平均低18.2%和29.5%; 而DS处理SS活性在花后10~18 d高于CK, 之后平均比CK低19.1% (2015— 2016年度)和25.6% (2016— 2017年度)。HT对SS活性的影响大于DS, HT+DS处理, SS活性降幅最大。

2.2 高温与干旱互作对小麦籽粒淀粉合成关键酶活性影响的方差分析

HT处理对花后14 d的SBE和AGPase活性影响不显著, 对其他各时期各种所测淀粉合成关键酶的活性影响均达显著水平; DS除对花后14 d的GBSS影响不显著外, 对所有关键酶活性的影响均达显著水平; 高温与干旱对淀粉合成关键酶活性的影响存在显著的互作效应(表1)。

表1 高温与干旱胁迫对小麦籽粒淀粉合成关键酶活性影响的互作效应方差分析(F值) Table 1 Variance analysis on interaction of HT and DS on activity of key starch synthesis enzymes (F-value)

2.3 高温与干旱胁迫对小麦籽粒淀粉积累的影响

2.3.1 高温与干旱胁迫对小麦籽粒淀粉含量的影响

小麦籽粒中直、支链淀粉及总淀粉含量变化趋势一致(图2), HT、DS及HT+DS均使籽粒淀粉含量显著下降。与CK相比, DS、HT及HT+DS处理的直链淀粉含量下降5.9%、10.2%、19.7% (2015— 2016年度)和5.5%、9.5%、19.2% (2016— 2017年度), 支链淀粉含量分别低7.2%、11.2%、18.2% (2015— 2016年度)和6.7%、11.2%、18.3% (2016— 2017年度), 总淀粉含量下降6.9%、10.99%、18.6% (2015— 2016年度)和6.4%、10.8%、18.5% (2016— 2017年度)。可见, HT对籽粒淀粉含量的影响大于DS, HT+DS大于单一胁迫。同时, 逆境胁迫缩短了小麦生育期, DS、HT及HT+DS较CK分别缩短4 d、8 d和12 d。

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	图2 不同处理籽粒直链淀粉(A)、支链淀粉(B)和总淀粉含量(C)的变化 HT: 高温; DS: 干旱; HT+DS: 高温与干旱复合胁迫。Fig. 2 Changes of amylose (A) amylopection (B), and total starch (C) contents under different treatments DS: drought stress; HT: high temperature stress; HT+DS: compound stress of HT and DS.

2.3.2 高温与干旱胁迫对小麦籽粒淀粉积累速率的影响不同处理下小麦籽粒淀粉积累速率均呈单峰曲线变化, HT和HT+DS在花后14 d达到峰值, CK和DS于花后18 d达到峰值(图3)。HT、DS及HT+DS均降低了小麦籽粒淀粉积累速率, 分别较CK下降6.0%、11.4%、21.8% (2015— 2016年度)和8.4%、11.9%、26.4% (2016— 2017年度)。

	Figure Option View Download New Window
	图3 不同处理小麦籽粒淀粉积累速率的变化 HT: 高温; DS: 干旱; HT+DS: 高温与干旱复合胁迫。Fig. 3 Changes of starch accumulating rate in wheat grains under different treatments DS: drought stress; HT: high temperature stress; HT+DS: compound stress of HT and DS.

2.4 高温与干旱胁迫对成熟期小麦籽粒淀粉及其组分含量的影响

HT、DS及其复合胁迫(HT+DS)均显著降低小麦籽粒直链、支链和总淀粉含量(表2)。与CK相比, HT、DS和HT+DS直链淀粉的含量分别下降10.6%、4.1%、14.5% (2015— 2016年度)和10.2%、4.2%、14.9% (2016— 2017年度), 支链淀粉含量分别下降13.8%、6.9%、19.7% (2015— 2016年度)和13.4%、6.0%、18.7% (2016— 2017年度), 总淀粉含量分别下降13.0%、6.2%、18.4% (2015— 2016年度)和12.7%、5.6%、17.8% (2016— 2017年度)。HT、DS及HT+DS使成熟期籽粒淀粉直/支比分别较CK提高3.7%、3.5%、6.4% (2015— 2016年度)和3.7%、1.9%、4.6% (2016— 2017年度)。表明高温胁迫对淀粉组成的影响大于干旱, 而复合胁迫具有叠加效应。

表2 不同处理成熟期小麦籽粒淀粉及其组分含量 Table 2 Starch content and its components in wheat grain under different treatments at maturity

2.5 高温与干旱胁迫对小麦籽粒产量及其构成因素的影响

HT和DS处理从花后10 d开始, 小麦穗数、穗粒数已基本确定, 因此各处理的穗数和穗粒数差异不显著, 但千粒重和籽粒产量都显著低于对照, 以复合胁迫造成的减产最明显(表3)。HT+DS处理的千粒重和盆产量两年度降幅为33.4%~36.6%和33.5%~38.1%; HT处理使千粒重和盆产量降低17.6%~22.5%和20.5%~21.5%; DS处理的减产幅度相对较小, 千粒重下降13.7%~14.5%, 盆产量下降18.1%~18.3%。

表3 不同处理对成熟期小麦籽粒产量及其构成因素的影响 Table 3 Effects of different treatments on grain yield and its components of wheat

2.6 小麦籽粒淀粉合成关键酶活性与淀粉积累量的相关性

在多数测定时期, 籽粒GBSS、SSS活性与支链淀粉、直链淀粉和总淀粉的含量均呈极显著正相关(花后22 d直链淀粉除外); 在灌浆后期(花后22~26 d), SBE、AGPase活性与籽粒支链淀粉、直链淀粉和总淀粉的含量亦呈极显著正相关; SS活性在灌浆初期(花后14 d)与籽粒支链淀粉、直链淀粉和总淀粉的含量均呈极显著负相关, 而后期(花后22~26 d)则呈显著正相关(表4)。由此可见, GBSS、SSS、SBE、AGPase和SS共同参与籽粒淀粉的合成与积累, 但其功能与作用时期有所不同。

表4 小麦籽粒淀粉合成关键酶活性与淀粉积累的相关系数 Table 4 Correlation coefficients of activities of key starch synthesis enzymes with contents of starch and its components

3 讨论

3.1 高温干旱胁迫下小麦籽粒淀粉合成关键酶活性与淀粉积累

小麦籽粒的充实过程主要是淀粉的合成与积累过程, 受一系列与淀粉合成有关酶的调控^[16]。AGPase、SSS、GBSS、SBE和SS被认为是调控淀粉合成的关键酶^{[17, 18]}。灌浆期高温、干旱导致小麦籽粒淀粉积累量下降, 一方面是由于籽粒蔗糖供应减少引起糖原不足, 另一方面则因灌浆中后期淀粉合成关键酶活性下降^{[8, 19, 20, 21, 22]}。逆境胁迫对籽粒淀粉合成关键酶活性的影响与籽粒发育阶段有关, 高温、干旱诱导了胁迫初期籽粒GBSS、SSS、SBE、AGPase和SS的活性, 而对胁迫后期的酶活性起抑制作用, 从而导致籽粒淀粉积累速率下降^{[23, 24, 25]}。孙立影等^[26]对春小麦花后不同时期施以短暂高温(35± 3, 3 d), 结果籽粒AGPase、SSS 和SBE活性较对照下降7%~13%。石慧清等^[19]在大田条件下于花后设置增温棚(增温2~5℃), 观测到高温处理使持绿型小麦品种籽粒SS、AGPase和SSS活性峰值分别下降7.7%、10.0%和21.5%, 非持绿型小麦品种分别下降14.1%、27.7%和27.9%, 表明小麦对高温胁迫的响应存在着基因型差异。本研究在昼夜高温模式下, 籽粒SS、AGPase、SSS和SBE活性的峰值较对照平均分别下降11.1%、15.6%、3.6%和3.5%, 并在多数测定时期低于对照。高温、干旱及其复合胁迫提高了灌浆前期籽粒GBSS、SSS活性, 而降低了灌浆后期的酶活性。从影响程度上看, 高温胁迫的影响大于干旱, 复合胁迫大于单一胁迫, 表现出明显的叠加效应。研究证明, 籽粒SSS、SBE、AGPase和SS活性与支链淀粉的合成具有显著的相关性, 而GBSS、AGPase和SS的活性与直链淀粉合成关系密切^{[2, 10, 27, 28]}。在本研究中, SSS和GBSS活性与籽粒支链、直链和总淀粉含量在多个测定时期均存在极显著正相关, 而SS、AGPase和SBE活性在灌浆后期(22~26 d)与籽粒支链、直链和总淀粉的含量呈极显著正相关, 表明高温、干旱及其复合胁迫对淀粉积累的影响是淀粉合成相关酶综合作用的结果。

3.2 高温干旱胁迫下小麦籽粒淀粉积累与产量

籽粒淀粉的合成与积累既受遗传因素的影响, 也易受环境条件的影响^{[29, 30, 31]}。高温和干旱胁迫对小麦内部结构和功能会造成一定的损伤^{[32, 33, 34]}, 灌浆期超过30℃的温度显著影响小麦籽粒支链淀粉、直链淀粉和总淀粉含量, 导致粒重和产量下降^{[22, 35, 36, 37]}。胡吉帮等^[38]研究表明, 灌浆前中期短暂高温(38℃, 2~4 d)可导致小麦的成熟期提前3~5 d。本研究在不同昼夜温度模式下, 高温、干旱及复合胁迫分别使小麦生育期缩短8、4和12 d。可见, 灌浆期持续的逆境胁迫(如昼夜温度模式)对小麦植株的影响大于短期极端高温。在本试验干旱胁迫下, 2015— 2016年度在灌浆前期籽粒直链、支链和总淀粉含量与对照差异不显著, 而2016— 2017年度则使其显著增加, 这种差异性可能与处理之前的气候条件差异或水分控制有关, 但随着胁迫时间的延长, 最终均导致籽粒直链、支链和总淀粉含量的显著下降, 两年结果一致。试验结果还表明, 高温胁迫初期会提高淀粉积累速率, 但随着胁迫时间的延长, 籽粒发育进程加快, 成熟期提前, 最终淀粉积累速率下降, 直链、支链和总淀粉含量低于对照, 粒重降低, 这与李春艳等^[39]研究结果一致。杨晓娟等^[33]研究认为干旱胁迫对小麦籽粒淀粉积累和粒重的影响比短暂高温胁迫更为明显, 而岳鹏莉等^[12]通过设置花后短暂高温(38℃)、干旱(55%)及其复合胁迫处理, 使小麦粒重分别下降34.4%、7.6%和43.2%, 表明高温胁迫的影响大于干旱处理。本研究中, 高温、干旱及其复合胁迫使小麦淀粉含量平均较对照分别下降10.9%、6.7%和18.6%, 粒重平均分别下降20.2%、14.1%和35.1%, 产量平均分别下降20.1%、18.2%和35.8%, 即高温的影响大于干旱胁迫, 复合胁迫具有明显的叠加效应。上述试验结果的差异与胁迫强度、持续时间(如短暂高温或持续昼夜温度模式)及胁迫处理时期有关, 同时还因基因型不同而有所差异。

4 结论

小麦灌浆期持续高温、干旱及其复合胁迫均导致小麦籽粒淀粉合成关键酶活性降低, 籽粒总淀粉及其组分含量减少, 进而粒重下降, 产量降低。本试验条件下, 高温处理对小麦籽粒淀粉合成关键酶活性、淀粉积累及产量的影响大于干旱胁迫, 且高温与干旱具显著的互作效应。高温与干旱胁迫通过影响籽粒中GBSS、SSS、SBE、AGPase和SS活性而改变淀粉积累特性, 最终影响小麦籽粒产量。

The authors have declared that no competing interests exist.

作者已声明无竞争性利益关系。

参考文献

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[1]	李永庚, 于振文, 姜东, 余松烈. 冬小麦旗叶蔗糖和籽粒淀粉合成动态及与其有关的酶活性的研究. 作物学报, 2001, 27: 658-664 Li Y G, Yu Z W, Jiang D, Yu S L. Studies on the dynamic changes of the synthesis of sucrose in the flag leaf and starch in the grain and related enzymes of high-yielding wheat. Acta Agron Sin, 2001, 27: 658-664 (in Chinese with English abstract) [本文引用:1]
[2]	闫素辉, 王振林, 戴忠民, 李文阳, 付国占, 贺明荣, 尹艳枰. 两个直链淀粉含量不同的小麦品种籽粒淀粉合成酶活性与淀粉积累特征的比较. 作物学报, 2007, 33: 84-89 Yan S H, Wang Z L, Dai Z M, Li W Y, Fu G Z, He M R, Yin Y P. Activity of enzymes involved in starch synthesis and accumulation in grains of two wheat cultivars with a different amylose content. Acta Agron Sin, 2007, 33: 84-89 (in Chinese with English abstract) [本文引用:2]
[3]	Wang Z B, Li W H, Qi J C, Shi P C, Yin Y A. Starch accumulation, activities of key enzyme and gene expression in starch synthesis of wheat endosperm with different starch contents. J Food Sci Technol, 2014, 51: 419-429 [本文引用:1]
[4]	李春燕, 封超年, 张影, 郭文善, 朱新开, 彭永欣. 氮肥基追比对弱筋小麦宁麦9号籽粒淀粉合成及相关酶活性的影响. 中国农业科学, 2005, 38: 1120-1125 Li C Y, Feng C N, Zhang Y, Guo W S, Zhu X K, Peng Y X. Effects of the ratio between basal N and top dressing N on grain starch formation in weak gluten wheat variety Ningmai 9 and its enzymes activities. Sci Agric Sin, 2005, 38: 1120-1125 (in Chinese with English abstract) [本文引用:1]
[5]	王晨阳, 郭天财, 阎耀礼, 朱云集, 夏国军, 王化岑, 周继泽. 花后短期高温胁迫对小麦叶片光合性能的影响. 作物学报, 2004, 30: 88-91 Wang C Y, Guo T C, Yan Y L, Zhu Y J, Xia G J, Wang H C, Zhou J Z. *Effects of short post-anthesis high temperature stress on leaf photosynthetic potential in winter wheat ( Triticum aestivum* L. )*. Acta Agron Sin*, 2004, 30: 88-91 (in Chinese with English abstract) [本文引用:1]
[6]	张洪华, 贺明荣, 刘永环, 李飞, 吴翠平, 张宾, 王晓英. 氮、硫肥与灌浆后期高温胁迫对小麦籽粒产量和品质的影响. 生态学杂志, 2008, 27: 162-166 Zhang H H, He M R, Liu Y H, Li F, Wu C P, Zhang B, Wang X Y. Effect of high temperature stress at late grain-filling stage and nitrogen and sulphur fertilization on wheat grain yield and quality. Chin J Ecol, 2008, 27: 162-166 (in Chinese with English abstract) [本文引用:1]
[7]	刘祖贵, 孙景生, 张寄阳, 王景雷, 孟兆江, 李晓东, 刘小飞. 不同时期干旱对强筋小麦产量与品质特性的影响. 麦类作物学报, 2008, 28: 877-882 Liu Z G, Sun J S, Zhang J Y, Wang J L, Meng Z J, Li X D, Liu X F. Effect of drought at different growing stages on yield and quality characteristics of strong-gluten wheat. J Trit Crops, 2008, 28: 877-882 (in Chinese with English abstract) [本文引用:1]
[8]	杨毅, 李昱, 康建宏, 刘萍. 花后高温胁迫对春小麦籽粒淀粉合成的影响. 麦类作物学报, 2015, 35: 1535-1541 Yang Y, Li Y, Kang J H, Liu P. Effect of heat stress after anthesis on starch synthesis in spring wheat. J Trit Crops, 2015, 35: 1535-1541 (in Chinese with English abstract) [本文引用:2]
[9]	李亚婷, 康建宏, 吴宏亮, 李昱, 姚珊. 花后水分胁迫对春小麦淀粉形成及相关酶活性的影响. 草业科学, 2016, 33: 917-925 Li Y T, Kang J H, Wu H L, Li Y, Yao S. Effect of water stress after anthesis on starch formation and the activity of associated enzymes in spring wheat. Pratacult Sci, 2016, 33: 917-925 (in Chinese with English abstract) [本文引用:1]
[10]	赵晶晶, 姚珊, 李昱, 康建宏, 吴宏亮, 许强. 花后干旱对春小麦籽粒淀粉含量及淀粉形成关键酶活性的影响. 干旱地区农业研究, 2015, 33: 104-110 Zhao J J, Yao S, Li Y, Kang J H, Wu H L, Xu Q. Effects of drought on the starch contents and activities of key enzymes after anthesis in spring wheat grains. Agric Res Arid Areas, 2015, 33: 104-110 (in Chinese with English abstract) [本文引用:2]
[11]	苗建利, 王晨阳, 郭天财, 马冬云, 胡吉帮, 冯辉. 高温与干旱互作对两种筋力小麦品种籽粒淀粉及其组分含量的影响. 麦类作物学报, 2008, 28: 254-259 Miao J L, Wang C Y, Guo T C, Ma D Y, Hu J B, Feng H. Effects of post-anthesis interactions of high temperature and drought stress on content and composition of grain starch in two wheat cultivars with different gluten strength. J Trit Crops, 2008, 28: 254-259 (in Chinese with English abstract) [本文引用:2]
[12]	岳鹏莉, 王晨阳, 卢红芳, 刘卫星, 马耕, 王强, 胡阳阳. 灌浆期高温干旱胁迫对小麦籽粒淀粉积累的影响. 麦类作物学报, 2016, 36: 1489-1496 Yue P L, Wang C Y, Lu H F, Liu W X, Ma G, Wang Q, Hu Y Y. Effect of heat and drought stress on starch accumulation during grain filling stage. J Trit Crops, 2016, 36: 1489-1496 (in Chinese with English abstract) [本文引用:2]
[13]	卢红芳, 王晨阳, 郭天财, 尹云星. 灌浆前期高温和干旱胁迫对小麦籽粒蛋白质含量和氮代谢关键酶活性的影响. 生态学报, 2014, 34: 3612-3619 Lu H F, Wang C Y, Guo T C, Yin Y X. Effects of high- temperature and drought stress on protein concentration and keys enzymes activities in relation to nitrogen metabolism in wheat grains during the early stage of grain filling. Acta Ecol Sin, 2014, 34: 3612-3619 (in Chinese with English abstract) [本文引用:1]
[14]	王晨阳, 苗建利, 张美微, 马冬云, 冯伟, 谢迎新, 郭天财. 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. 生态学报, 2014, 34: 4882-4890 Wang C Y, Miao J L, Zhang M W, Ma D Y, Feng W, Xie Y X, Guo T C. Effects of post-anthesis high temperature, drought stress and their interaction on the starch pasting properties of two wheat cultivars with different gluten strength. Acta Ecol Sin, 2014, 34: 4882-4890 (in Chinese with English abstract) [本文引用:1]
[15]	何照范. 粮油籽粒品质及其分析技术. 北京: 中国农业出版社, 1985. pp 144-150 He Z F. Analysis Technique for Grain Quality in Cereals and Oils. Beijing: China Agriculture Press, 1985. pp 144-150(in Chinese) [本文引用:1]
[16]	Rahman S, Regina A, Li Z Y, Mukai Y, Yamamoto M, Kosar- Hashemi B, Abrahams S, Morell M K. Comparison of starch branching enzyme genes reveals evolutionary relationships among isoform. Characterization of a gene for starch-branching enzymes IIa from the wheat D genome donor Aegilops tauschii. Plant Physiol, 2001, 125: 1314-1324 [本文引用:1]
[17]	Nakamura Y, Yuki K, Park S Y, Ohya T. Carbohydrate metabolism in the developing endosperm of rice grains. Plant Cell Physiol, 1989, 30: 833-839 [本文引用:1]
[18]	曹颖妮, 胡卫国, 王根平, 刘录祥, 王成社. 糯性和非糯性小麦灌浆期胚乳直/支链淀粉积累及其相关酶活性研究. 西北植物学报, 2010, 30: 1995-2001 Cao Y N, Hu W G, Wang G P, Liu L X, Wang C S. Dynamic changes of starch accumulation and enzymes relating to starch biosynthesis of kernel during grain filling in waxy and non-waxy winter wheat. Acta Bot Boreali-Occid Sin, 2010, 30: 1995-2001 (in Chinese with English abstract) [本文引用:1]
[19]	石慧清, 龚月桦, 张东武. 花后高温对持绿型小麦叶片衰老及籽粒淀粉合成相关酶的影响. 植物生态学报, 2011, 35: 769-778 Shi H Q, Gong Y H, Zhang D W. Effect of high temperature on leaf senescence and related enzymes of grain starch synthesis in stay-green wheat after anthesis. J Plant Ecol, 2011, 35: 769-778 (in Chinese with English abstract) [本文引用:2]
[20]	Dai Z M, Yin Y P, Wang Z L. Comparison of starch accumulation and enzyme activity in grains of wheat cultivars differing in kernel type. Plant Growth Regul, 2009, 57: 153-162 [本文引用:1]
[21]	Ahmadi A, Baker D A. The effect of water stress on the activities of key regulatory enzymes of the sucrose to starch pathway in wheat. Plant Growth Regul, 2001, 35: 81-91 [本文引用:1]
[22]	包刚, 覃志豪, 周义, 包玉海, 萨楚拉. 气候变化对中国农业生产影响的模拟评价进展. 中国农学通报, 2012, 28(2): 303-307 Bao G, Tan Z H, Zhou Y, Bao Y H, Sa C L. Advances of evaluation of climate changes impact on crop yield. Chin Agric Sci Bull, 2012, 28(2): 303-307 (in Chinese with English abstract) [本文引用:2]
[23]	赵辉, 戴廷波, 荆奇, 姜东, 曹卫星, 陆玮, 田孝威. 灌浆期高温对两种品质类型小麦品种籽粒淀粉合成关键酶活性的影响. 作物学报, 2006, 32: 423-429 Zhao H, Dai T B, Jing Q, Jiang D, Cao W X, Lu W, Tian X W. Effects of high temperature during grain filling on key enzymes involved in starch synthesis in two wheat cultivars with different quality types. Acta Agron Sin, 2006, 32: 423-429 (in Chinese with English abstract) [本文引用:1]
[24]	闫素辉, 尹燕枰, 李文阳, 梁太波, 李勇, 邬云海, 王平, 耿庆辉, 戴忠民, 王振林. 灌浆期高温对小麦籽粒淀粉的积累、粒度分布及相关酶活性的影响. 作物学报, 2008, 34: 1092-1096 Yan S H, Yin Y P, Li W Y, Liang T B, Li Y, Wu Y H, Wang P, Geng Q H, Dai Z M, Wang Z L. Effect of high temperature during grain filling on starch accumulation, starch granule distribution, and activities of related enzymes in wheat grains. Acta Agron Sin, 2008, 34: 1092-1096 (in Chinese with English abstract) [本文引用:1]
[25]	王征宏, 郭秀璞, 吕淑芳, 李友军, 邓西平. 土壤干旱对不同小麦品种籽粒淀粉积累和相关酶活性的影响. 干旱地区农业研究, 2011, 29: 67-74 Wang Z H, Guo X P, Lyu S F, Li Y J, Deng X P. Effects of soil drought on starch accumulation of grain and activity of corresponding starch synthesis enzymes in different winter wheat varieties. Agric Res Arid Areas, 2011, 29: 67-74 (in Chinese with English abstract) [本文引用:1]
[26]	孙立影, 苏玮, 吴宏亮, 李昱, 王雪, 康建宏. 花后不同时期高温对春小麦淀粉形成的影响. 农业科学研究, 2015, 36: 54-60 Sun L Y, Su W, Wu H L, Li Y, Wang X, Kang J H. Effects of the high temperature after the flowering period on starch formation in spring wheat. J Agric Sci, 2015, 36: 54-60 (in Chinese with English abstract) [本文引用:1]
[27]	王月福, 于振文, 李尚霞, 余松烈. 小麦籽粒灌浆过程中有关淀粉合成酶的活性及其效应. 作物学报, 2003, 29: 75-81 Wang Y F, Yu Z W, Li S X, Yu S L. Activity of enzymes related to starch synthesis and their effect during the filling of winter wheat. Acta Agron Sin, 2003, 29: 75-81 (in Chinese with English abstract) [本文引用:1]
[28]	李建敏, 王振林, 高荣岐, 李圣福, 蔡瑞国, 闫素辉, 于安玲, 尹艳枰. 强、弱筋小麦籽粒形成期蔗糖、淀粉合成相关酶活性及其与氮代谢的关系. 作物学报, 2008, 34: 1019-1026 Li J M, Wang Z L, Gao R Q, Li S F, Cai R G, Yan S H, Yu A L, Yin Y P. Activities of enzymes involved in sucrose and starch synthesis during grain filling and the relation to nitrogen metabolism in strong-and weak-gluten wheat cultivars. Acta Agron Sin, 2008, 34: 1019-1026 (in Chinese with English abstract) [本文引用:1]
[29]	Panozzo J F, Eagles H A. Cultivar and environmental effects on quality characters in wheat: I. Starch. Aust J Agric Res, 1998, 49: 757-766 [本文引用:1]
[30]	Morris C F, Shackley B J, King G E, Kidwell K K. Genotypicand environmental variation for flour swelling volume in wheat. Cereal Chem, 1997, 74: 16-21 [本文引用:1]
[31]	Jenner C F. Starch synthesis in the kernel of wheat under high temperature conditions. Aust J Plant Physiol, 1994, 21: 791-806 [本文引用:1]
[32]	许振柱, 于振文, 张永丽. 土壤水分对小麦籽粒淀粉合成和积累特性的影响. 作物学报, 2003, 29: 595-600 Xu Z Z, Yu Z W, Zhang Y L. The effects of soil moisture on grain starch synthesis and accumulation of winter wheat. Acta Agron Sin, 2003, 29: 595-600 (in Chinese with English abstract) [本文引用:1]
[33]	杨晓娟, 居辉, 王治世, 郭安廷, 杨佑明. 花后高温和干旱对冬小麦光合、抗氧化特性及粒重的影响. 麦类作物学报, 2015, 35: 958-963 Yang X J, Ju H, Wang Z S, Guo A T, Yang Y M. Effect of high temperature and drought after anthesis on photosynthesis, antioxidant properties and grain weight in winter wheat. J Trit Crops, 2015, 35: 958-963 (in Chinese with English abstract) [本文引用:2]
[34]	苏玮, 孙立影, 康建宏, 吴宏亮, 李昱. 高温干旱对春小麦光合特性及产量的影响. 农业科学研究, 2015, 36: 49-53 Su W, Sun L Y, Kang J H, Wu H L, Li Y. Effect of high temperature and drought treatment on photosynthetic characteristics and yield of different varieties of spring wheat. J Agric Sci, 2015, 36: 49-53 (in Chinese with English abstract) [本文引用:1]
[35]	Jenner C F. Effects of exposure of wheat ears to high temperature on dry matter accumulation and carbohydrate metabolism in the grain of two cultivars: II. Carry-over effects. Aust J Plant Physiol, 1991, 18: 179-190 [本文引用:1]
[36]	Rane J, Nagarajan S. High temperature index-for field evaluation of heat tolerance in wheat varieties. Agric Systems, 2004, 79: 243-255 [本文引用:1]
[37]	Shah N H, Paulsen G M. Interaction of drought and high temperature on photosynthesis and grain-filling of wheat. Plant Soil, 2003, 257: 219-226 [本文引用:1]
[38]	胡吉帮, 王晨阳, 郭天财, 苗建利, 朱云集, 贺德先. 灌浆期高温和干旱对小麦灌浆特性的影响. 河南农业大学学报, 2008, 42: 597-601 Hu J B, Wang C Y, Guo T C, Miao J L, Zhu Y J, He D X. Effects of high temperature and drought stress on grain filling characteristics in wheat during grain filling period. J Henan Agric Univ, 2008, 42: 597-601 (in Chinese with English abstract) [本文引用:1]
[39]	李春艳, 付凯勇, 张润琪, 徐芳芳, 史晓艳, 朱永琪, 覃安祥, 李诚. 灌浆期高温对冬小麦淀粉粒发育的影响. 麦类作物学报, 2015, 35: 1395-1402 Li C Y, Fu K Y, Zhang R Q, Xu F F, Shi X Y, Zhu Y Q, Qin A X, Li C. Effect of high temperature post anthesis on the development of starch granules in winter wheat. J Trit Crops, 2015, 35: 1395-1402 (in Chinese with English abstract) [本文引用:1]

2001

0.0

李永庚, 于振文, 姜东, 余松烈. 冬小麦旗叶蔗糖和籽粒淀粉合成动态及与其有关的酶活性的研究. 作物学报, 2001, 27: 658-664

Li Y

, Yu Z

, Jiang

, Yu S

. Studies on the dynamic changes of the synthesis of sucrose in the flag leaf and starch in the grain and related enzymes of high-yielding wheat. Acta Agron Sin, 2001, 27: 658-664 (in Chinese with English abstract)

鲁麦22和鲁麦14相比,旗叶的光合能力强,蔗糖含量高,催化蔗糖合成的磷酸蔗糖合成酶(SPS)活性高;籽粒中蔗糖供应充足,降解蔗糖的蔗糖合成酶(SS)活性高;与淀粉合成有关的酶:腺苷二磷酸葡萄糖焦磷酸化酶(ADPG-PPase)、游离态(或可溶性)淀粉合成酶(S-STS)和束缚态淀粉合成酶(B-STS)均有较高的活性,这是其粒重高的原因.通过对两个品

... 有研究表明, 蔗糖合酶(SS)、ADPG焦磷酸化酶(AGPase)、可溶性淀粉合酶(SSS)、结合态淀粉合酶(GBSS)和淀粉分支酶(SBE)是控制淀粉合成代谢的关键酶^[1,2], 在小麦籽粒淀粉合成与积累中起重要作用 ...

2007

0.0

闫素辉, 王振林, 戴忠民, 李文阳, 付国占, 贺明荣, 尹艳枰. 两个直链淀粉含量不同的小麦品种籽粒淀粉合成酶活性与淀粉积累特征的比较. 作物学报, 2007, 33: 84-89

Yan S

, Wang Z

, Dai Z

, Li W

, Fu G

, He M

, Yin Y

. Activity of enzymes involved in starch synthesis and accumulation in grains of two wheat cultivars with a different amylose content. Acta Agron Sin, 2007, 33: 84-89 (in Chinese with English abstract)

The amylose content of wheat grain is significantly correlated with amount of Waxy protein subunit. However, there is a great difference in amylose content among different wheat cultivars which all have three Waxy genes. In order to investigate the activity changes of enzymes involved in starch biosynthesis, the characteristics of starch accumulation in grains, and the relationship between them, two wheat ( Triticum aestivum L.) cultivars, JM20 (lower amylose content in grains) and LM21 (higher amylose content in grains) were used in the study, which both carry three Waxy protein subunits. The results indicated that the activities of sucrose synthase (SS), adenosine diphosphorate glucose pyrophrylase (AGPP), soluble starch synthase (SSS), granule-bound starch synthase (GBSS) changed in the pattern of a single-peak curve during grain filling. The activities of SS, AGPP, GBSS, SSS and SBE in LM21, which had higher starch content, were higher than those in JM20. The accumulation rate of amylopectin was significantly or highly significantly correlated with the activities of SS, AGPP, SSS and SBE, and the accumulation rate of amylose was highly significantly correlated with the activities of SS, AGPP and GBSS. The accumulation courses of both amylopectin and amylose were well fitted to the logistic equation by relating amylopectin and amylose contents against days post anthesis. The simulation parameters revealed that the higher contents of amylopectin and amylose resulted from earlier initiating accumulation time and greater accumulation rate, but accumulation duration probably played a less important role for them. The amylose content in grains was mainly determined by GBSS activity as well as the activities of SS and AGPP. The change of GBSS activity reflected expression level of waxy gene was closely associated with the accumulation rate of amylose, indicating that amylose content in grains was determined by GBSS activity, especially at late grain filling stage. The difference in expression of waxy gene might be a major reason for the large difference in amylose contents between JM20 and LM21.

以均含有3个Waxy蛋白亚基的普通小麦品种济麦20（低直链淀粉含量）和鲁麦21（高直链淀粉含量）为材料，对灌浆期籽粒淀粉合成相关酶活性的变化及淀粉积累特征进行了研究，并分析了两者之间的关系。结果表明，蔗糖合成酶(SS)、腺苷二磷酸葡萄糖焦磷酸化酶(AGPP)、束缚态淀粉合成酶(GBSS)、可溶性淀粉合成酶(SSS)和淀粉分支酶(SBE)活性均呈单峰曲线变化。鲁麦21的上述酶活性均高于济麦20。相关分析表明，支链淀粉积累速率与SS、AGPP、SSS和SBE呈显著或极显著正相关；直链淀粉积累速率与SS、AGPP和GBSS呈极显著正相关。Logistic方程拟合淀粉积累过程发现，支、直链淀粉最终积累量的高低取决于积累启动时间的早晚和积累速率的高低，而积累持续期的调节作用较小。直链淀粉的积累速率除受GBSS活性影响外，还受SS和AGPP活性的影响，其中，GBSS活性的变化与2品种籽粒直链淀粉积累量的变化情况基本吻合。籽粒灌浆后期的GBSS活性对直链淀粉最终积累量的调节作用大于灌浆前期，说明对同时具有3个Waxy蛋白亚基的不同品种，Waxy蛋白亚基表达量(GBSS活性)的差异可能是导致品种间籽粒直链淀粉含量较大差异的一个关键原因。

... 研究证明, 籽粒SSS、SBE、AGPase和SS活性与支链淀粉的合成具有显著的相关性, 而GBSS、AGPase和SS的活性与直链淀粉合成关系密切^[2,10,27,28] ...

2014

0.0

... 其中, 小麦籽粒直链、支链和总淀粉的积累速率与SBE、SSS、GBSS和AGPase活性均存在显著的相关性^[3,4] ...

2005

0.0

李春燕, 封超年, 张影, 郭文善, 朱新开, 彭永欣. 氮肥基追比对弱筋小麦宁麦9号籽粒淀粉合成及相关酶活性的影响. 中国农业科学, 2005, 38: 1120-1125

Li C

, Feng C

, Zhang

, Guo W

, Zhu X

, Peng Y

. Effects of the ratio between basal N and top dressing N on grain starch formation in weak gluten wheat variety Ningmai 9 and its enzymes activities. Sci Agric Sin, 2005, 38: 1120-1125 (in Chinese with English abstract)

... 其中, 小麦籽粒直链、支链和总淀粉的积累速率与SBE、SSS、GBSS和AGPase活性均存在显著的相关性^[3,4] ...

2004

0.0

王晨阳, 郭天财, 阎耀礼, 朱云集, 夏国军, 王化岑, 周继泽. 花后短期高温胁迫对小麦叶片光合性能的影响. 作物学报, 2004, 30: 88-91

Wang C

, Guo T

, Yan Y

, Zhu Y

, Xia G

, Wang H

, Zhou J

. Effects of short post-anthesis high temperature stress on leaf photosynthetic potential in winter wheat ( Triticum aestivum L. ). Acta Agron Sin, 2004, 30: 88-91 (in Chinese with English abstract)

温度是小麦生长发育的主要生态因子之一.在华北地区,小麦生育后期温度回升较快,常出现高温天气,致使小麦植株衰老加速,灌浆期缩短,粒重和产量显著下降 [1].据对河南省气象资料分析,小麦灌浆后期(5月22～28日)出现日均温26℃、日最高气温超过32℃天气的年份占35%～55%,其中西部地区高于北部和南部地区[2].

... 花后高温和干旱往往导致小麦穗粒数减少, 粒重和产量显著降低^[5,6] ...

2008

0.0

张洪华, 贺明荣, 刘永环, 李飞, 吴翠平, 张宾, 王晓英. 氮、硫肥与灌浆后期高温胁迫对小麦籽粒产量和品质的影响. 生态学杂志, 2008, 27: 162-166

Zhang H

, He M

, Liu Y

, Li

, Wu C

, Zhang

, Wang X

. Effect of high temperature stress at late grain-filling stage and nitrogen and sulphur fertilization on wheat grain yield and quality. Chin J Ecol, 2008, 27: 162-166 (in Chinese with English abstract)

灌浆期高温胁迫通常导致小麦籽粒产量降低、品质变差.为了探索有效缓解高温胁迫不利影响的有效措施,在大田条件下,以济麦20为试验材料,运用后期高温棚增温的方法,研究了氮、硫肥与灌浆后期高温对小麦籽粒产量和品质的影响.结果表明:高温处理后小麦千粒质量、籽粒产量显著降低(19.7%～28.3%),蛋白质含量升高,蛋白质组分中不溶性谷蛋白含量下降,可溶性谷蛋白含量上升,面团形成时间和稳定时间缩短;氮肥追施比例由50%增加到70%,非高温处理的籽粒产量无显著变化,不溶性谷蛋白质含量和谷蛋白聚合指数 (不溶性谷蛋白含量/可溶性谷蛋白含量)升高,面团形成时间和稳定时间延长;高温处理的千粒质量和籽粒产量因氮肥追施比例提高而提高 (2.07%～3.58%),面团形成时间和稳定时间缩短;不论高温处理还是非高温处理,与追施硫肥相比,硫肥基施提高籽粒产量、谷蛋白聚合指数,延长面团形成时间和稳定时间;为了有效缓解灌浆后期高温对小麦籽粒产量和品质的不利影响,硫肥基施、氮肥施用基追比50%:50%是较为适宜的施肥模式.

... 花后高温和干旱往往导致小麦穗粒数减少, 粒重和产量显著降低^[5,6] ...

2008

0.0

刘祖贵, 孙景生, 张寄阳, 王景雷, 孟兆江, 李晓东, 刘小飞. 不同时期干旱对强筋小麦产量与品质特性的影响. 麦类作物学报, 2008, 28: 877-882

Liu Z

, Sun J

, Zhang J

, Wang J

, Meng Z

, Li X

, Liu X

. Effect of drought at different growing stages on yield and quality characteristics of strong-gluten wheat. J Trit Crops, 2008, 28: 877-882 (in Chinese with English abstract)

... 黄淮海麦区灌浆后期还经常遭遇高温与干旱复合胁迫的危害, 甚至形成干热风天气, 导致小麦减产10%~30% ^[7] ...

2015

0.0

杨毅, 李昱, 康建宏, 刘萍. 花后高温胁迫对春小麦籽粒淀粉合成的影响. 麦类作物学报, 2015, 35: 1535-1541

Yang

, Li

, Kang J

, Liu

. Effect of heat stress after anthesis on starch synthesis in spring wheat. J Trit Crops, 2015, 35: 1535-1541 (in Chinese with English abstract)

To understand the effects of heat stress on starch synthesis in wheat,high temperature (35℃±3℃) was induced to Ningchun 4 wheat at different growth stages after anthesis (5 d,10 d,15 d,20 d). The expression levels of four starch synthase genes,namely,ADP-glucose pyrophosphorylase gene ( AGPase ),granule-bound starch synthase Ⅰ (GBSSⅠ)gene,starch branching enzyme (SBE) gene,Soluble starch synthase (SSS)gene were investigated by quantitative real-time PCR (qRT-PCR). Meanwhile the change of enzyme (AGPase,SBE,GBSS,SSS) activity in starch biosynthesis and the accumulation of starch were detected during the wheat grain filling stage. Correlation analysis between the gene expression level and the activity of starch synthesis enzyme has been conducted. The results showed that all the genes encoding starch biosynthesis enzymes had high expression level during grain filling stage under normal temperature. Under heat stress,however,only the GBSSⅠ remained its expression level. The AGPase,SBE ,and SSS gene expression levels and the activities of the four starch synthase,on the other hand,were all inhibited by high temperature during the grain filling stages. The gene expression level of GBSSⅠ did not correlate with the activity of its starch synthases,while the relative gene expression level of SBE correlated positively with the activity of its starch synthase ( P AGPase and SSS correlated positively with the activity of their starch synthase under normal temperature ( P SBE,AGPase,SSS expression and the activities of starch synthase enzyme decreased under high temperature. These results suggested that AGPase,SBE,GBSⅠS and SSS may control starch synthesis in wheat grain filling stage,and heat stress can inhibited the accumulation of starch by affecting starch synthase gene expression,thereby affecting the enzyme activities of starch synthase. The inhibitive effects were more significant if high temperature occurred earlier at grain filling stage.

为揭示高温胁迫对春小麦籽粒淀粉合成的影响机制，以宁春4号小麦为材料，分别于花后5、10、15和20 d进行高温（35 ℃±3 ℃）胁迫，利用实时荧光定量PCR法（qPCR）研究花后不同时期高温胁迫下小麦籽粒淀粉合成相关酶基因表达，并将基因表达量与对应籽粒淀粉合成酶活性进行相关性分析。结果表明，束缚态淀粉合成酶Ⅰ（GBSSⅠ）基因对高温不敏感，而腺苷二磷酸葡萄糖焦磷酸化酶（AGPase）基因、淀粉分支酶（SBE）基因、可溶性淀粉合成酶（SSS）基因的表达均被高温胁迫抑制；被测的4种淀粉合成相关酶的活性均被高温协迫抑制。自然生长温度下，除GBSSⅠ酶外，SBE、AGPase和SSS酶活性与其对应基因的表达量极显著或显著正相关；高温胁迫使4种被测的淀粉合成相关酶活性与其对应基因表达量的相关性降低。高温胁迫通过抑制小麦籽粒淀粉合成相关酶基因的表达而影响对应酶活性，使小麦籽粒淀粉的积累发生改变；高温胁迫发生时间越早，对小麦籽粒淀粉积累影响越大。

... 胁迫发生的越早, 对淀粉积累影响越大^[8] ...

... 灌浆期高温、干旱导致小麦籽粒淀粉积累量下降, 一方面是由于籽粒蔗糖供应减少引起糖原不足, 另一方面则因灌浆中后期淀粉合成关键酶活性下降^{[8,19,20,21,22]} ...

2016

0.0

李亚婷, 康建宏, 吴宏亮, 李昱, 姚珊. 花后水分胁迫对春小麦淀粉形成及相关酶活性的影响. 草业科学, 2016, 33: 917-925

Li Y

, Kang J

, Wu H

, Li

, Yao

. Effect of water stress after anthesis on starch formation and the activity of associated enzymes in spring wheat. Pratacult Sci, 2016, 33: 917-925 (in Chinese with English abstract)

Recently, frequent drought and waterlogging due to climate change caused severe yield loss in spring wheat. The research about the effects of water stress after anthesis on starch formation had great importance for cultivation of spring wheat with stable production and high quality. In the present study, two spring wheat varieties Ningchun 4 and Ningchun 47 which were domain varieties in Ningxia province were selected as material. Four different soil moisture treatments including heavy drought, light drought, normal and water logging were performed by manipulating the post-anthesis soil moisture in pots. The results showed that post-anthesis drought significantly increased ( P <0.05) the grain starch content during the earlier filling stage and significantly reduced ( P <0.05) the amylose and amylopectin content during the mid and late filing stage so that significantly reduced ( P <0.05) the total starch content during the late filing stage. The activities of four key enzymes associated with starch formation, AGPP(ADP-glucose pyrophosphorylase), UGPP( UDP-glucose pyrophosphorylase), SBE (starch branching Enzyme)and SSS(soluble starch synthase), varied with a single peak curve. Water stress depressed these four enzyme activities in spring wheat during the mid and late filling stage. The rate of amylose accumulation in the two spring wheat was significant positively correlated with the activity of SSS ( P <0.01).

近年来,气候变化导致农田干旱与渍水频繁发生。研究花后水分胁迫对春小麦( Triticum aestivum )淀粉形成的影响,对栽培稳产优质的春小麦品种具有重要意义。本研究以宁夏平原主栽春小麦宁春4号和宁春47号为试材,采用盆栽种植,人工控制土壤含水量,设置重度干旱、轻度干旱、正常处理和过度灌溉4个试验梯度。结果表明,花后干旱显著( P <0.05)提高了宁春4号灌浆前期籽粒中直链淀粉含量,显著降低了两种春小麦灌浆中后期直链、支链淀粉含量,最终使收获期籽粒中总淀粉含量显著降低。与籽粒中淀粉形成相关的4种关键酶,AGPP、UGPP、SBE及SSS活性在整个灌浆期均呈单峰曲线,两种春小麦灌浆中后期受花后水分胁迫显著降低了酶活性,两种春小麦的直链淀粉积累速率与SSS活性活性均呈极显著正相关( P <0.01)。

... 干旱胁迫降低籽粒SSS、SBE、AGPase的活性, 减少籽粒直、支链淀粉含量, 导致总淀粉含量显著下降^[9,10] ...

2015

0.0

赵晶晶, 姚珊, 李昱, 康建宏, 吴宏亮, 许强. 花后干旱对春小麦籽粒淀粉含量及淀粉形成关键酶活性的影响. 干旱地区农业研究, 2015, 33: 104-110

Zhao J

, Yao

, Li

, Kang J

, Wu H

, Xu

. Effects of drought on the starch contents and activities of key enzymes after anthesis in spring wheat grains. Agric Res Arid Areas, 2015, 33: 104-110 (in Chinese with English abstract)

... 干旱胁迫降低籽粒SSS、SBE、AGPase的活性, 减少籽粒直、支链淀粉含量, 导致总淀粉含量显著下降^[9,10] ...

... 研究证明, 籽粒SSS、SBE、AGPase和SS活性与支链淀粉的合成具有显著的相关性, 而GBSS、AGPase和SS的活性与直链淀粉合成关系密切^[2,10,27,28] ...

2008

0.0

苗建利, 王晨阳, 郭天财, 马冬云, 胡吉帮, 冯辉. 高温与干旱互作对两种筋力小麦品种籽粒淀粉及其组分含量的影响. 麦类作物学报, 2008, 28: 254-259

Miao J

, Wang C

, Guo T

, Ma D

, Hu J

, Feng

. Effects of post-anthesis interactions of high temperature and drought stress on content and composition of grain starch in two wheat cultivars with different gluten strength. J Trit Crops, 2008, 28: 254-259 (in Chinese with English abstract)

... 当小麦花后遭遇极端高温(35℃以上)和干旱双重胁迫时, 会导致灌浆期明显缩短, 籽粒直链、支链淀粉和总淀粉含量降低, 影响淀粉的糊化特性, 并使籽粒蛋白质积累量和产量下降, 而且高温与干旱存在显著的互作效应^{[11,12,13,14]} ...

... ^{${{R}_{n}}=\frac{{{C}_{n+4}}-{{C}_{n-4}}}{8}$[11]} ...

2016

0.0

岳鹏莉, 王晨阳, 卢红芳, 刘卫星, 马耕, 王强, 胡阳阳. 灌浆期高温干旱胁迫对小麦籽粒淀粉积累的影响. 麦类作物学报, 2016, 36: 1489-1496

Yue P

, Wang C

, Lu H

, Liu W

, Ma

, Wang

, Hu Y

. Effect of heat and drought stress on starch accumulation during grain filling stage. J Trit Crops, 2016, 36: 1489-1496 (in Chinese with English abstract)

摘　要：为探讨花后高温和干旱逆境胁迫对小麦淀粉组分的影响，采用盆栽和人工气候室相结合的方式研究了灌浆期短暂高温、干旱及其复合胁迫对两个不同品质类型小麦品种籽粒总淀粉、直链淀粉和支链淀粉含量的影响。结果表明，小麦籽粒淀粉的积累量符合Logistic方程，但逆境胁迫会缩短籽粒淀粉积累持续时间，降低其积累速率，使其最终淀粉积累量减少。与对照相比，高温、干旱及其复合胁迫显著降低两个小麦品种籽粒的支链淀粉和总淀粉含量，直链淀粉含量受影响较小，淀粉的直／支比增加。高温、干旱及其复合胁迫下，洛麦24的千粒重分别较对照下降42．03％、21．95％和50．01％，产量分别较对照下降57．88％、40．75％和61．08％；郑麦366的千粒重分别较对照下降34．39％、7．64％和43．16％，产量分别较对照下降41．58％、30．97％和48．49％。综上所述，高温、干旱胁迫抑制支链淀粉积累是造成产量下降的重要原因；复合胁迫对小麦籽粒淀粉积累量的影响大于干旱或高温单独胁迫；高温胁迫的影响大于干旱；洛麦24较郑麦366对高温和干旱胁迫更敏感。

... 杨晓娟等^[33]研究认为干旱胁迫对小麦籽粒淀粉积累和粒重的影响比短暂高温胁迫更为明显, 而岳鹏莉等^[12]通过设置花后短暂高温(38℃)、干旱(55%)及其复合胁迫处理, 使小麦粒重分别下降34 ...

2014

0.0

卢红芳, 王晨阳, 郭天财, 尹云星. 灌浆前期高温和干旱胁迫对小麦籽粒蛋白质含量和氮代谢关键酶活性的影响. 生态学报, 2014, 34: 3612-3619

Lu H

, Wang C

, Guo T

, Yin Y

. Effects of high- temperature and drought stress on protein concentration and keys enzymes activities in relation to nitrogen metabolism in wheat grains during the early stage of grain filling. Acta Ecol Sin, 2014, 34: 3612-3619 (in Chinese with English abstract)

To explore the physiological mechanisms of stresses on nitrogen metabolism and protein accumulation in wheat grains, a pot experiment was performed at the Scientific & Educational Park of Henan Agricultural University from 2010 to 2012. Zhengmai 366 and Yunong 949, wheat cultivars with different gluten-strengths, were grown in 24-cm pots with 10 kg loam soil for the experiment, which was performed using a completely randomized design in three replicates. Plants were initially grown in the field environment and were then transferred to a climate-controlled greenhouse for the high temperature (HT) treatment. HT began from the 8 th day after anthesis (DAA) and ended at the 11 th DAA, with a high temperature of 38℃for five hours from 11:00 to 16:00, after which the plants were returned to the field environment. The concentration of protein and its components, and the activities of glutamine synthetase (GS) and glutamic-pyruvic transaminase (GPT) in the grains were measured to evaluate the effects of the stresses. The results showed that HT, drought stress (DS) and their combination significantly increased protein and its components' concentrations, yet decreased the ratio of glutenin to gliadin. The protein accumulation was also reduced by stresses. For example, for Zhengmai 366, the protein yield significantly decreased by 20.7% under HT, by 16.2% under DS, and by 26.1% under a combination of HT and DS. In Yunong 949, the protein yield decreased by 12.4% under HT, by 11.9% under DS and by 15.8% under a combination of HT and DS. The grain weight significantly declined by 23.2% under HT, by 18.0% under DS and by 29.9% under a combination of HT and DS, in Zhengmai 366, and decreased by 24.0% under HT, by 16.0% under DS, and by 29.9% under a combination of HT and DS, in Yunong 949. However, the activities of GS and GPT were promoted under HT, DS and their combination. A correlation analysis showed that the activity of GS was positively correlated with the protein concentration in mature wheat grains at significant or extremely significant levels at 8, 17, 23, and 29 DAA. However, GS activity was negatively correlated with protein yield at extremely significant levels at 23 and 35 DAA, and it was negatively correlated with grain yield at significant or extremely significant levels at 8, 17, 23, 29 and 35 DAA. The activity of GPT was positively correlated with protein concentration at extremely significant levels at 11 and 17 DAA. However, GPT activity was negatively correlated with protein yield at extremely significant levels at 8, 17 and 23 DAA and negatively correlated with grain yield at significant or extremely significant levels at 8, 11, 17 and 23 DAA. The results also indicated that, under the present experimental conditions, the influence of the HT treatment was greater than that of the DS, and an overlap effect existed between HT and DS. In addition, Zhengmai 366, a high-gluten cultivar, showed more sensitivity to stresses than Yunong 949, a mid-gluten cultivar. These results may aid in understanding how HT and DS influence protein accumulation and may be useful in improving wheat quality.

为探讨花后逆境胁迫影响小麦籽粒氮代谢及蛋白质合成的生理机制，采用盆栽和人工气候室模拟花后高温的方式，研究了灌浆前期短暂高温和干旱胁迫对两个不同品质类型小麦品种籽粒蛋白质含量、组分及谷氨酰胺合成酶（GS）、谷丙转氨酶（GPT）活性的影响。结果表明，灌浆前期高温、干旱及其复合胁迫均显著提高两品种籽粒蛋白质及组分含量，但降低谷/醇比。逆境胁迫使蛋白质积累量和粒重显著下降，其中高温处理使两品种蛋白质产量分别下降20.7%和12.4%，粒重下降23.2%和24.0%；干旱胁迫使两品种蛋白质产量分别下降16.2%和11.9%，粒重下降18.0%和16.0%；复合胁迫使两品种蛋白质产量分别下降26.1%和15.8%，粒重下降29.9%和28.9%。高温、干旱及其复合胁迫下两品种籽粒氮代谢关键酶活性升高。花后8，17，23，29 d的GS活性和花后11，17 d的GPT活性与蛋白质含量呈显著或极显著正相关，花后23，35 d的GS和花后8，17，23 d的GPT活性与蛋白质产量呈显著或极显著负相关，花后8，17，23，29，35 d的GS和花后8，11，17，23 d的GPT活性与籽粒产量呈显著或极显著负相关。试验条件下，高温处理对籽粒蛋白质合成的影响大于干旱胁迫，二者具有叠加效应，强筋小麦品种郑麦366受逆境胁迫的影响较大。

2014

0.0

王晨阳, 苗建利, 张美微, 马冬云, 冯伟, 谢迎新, 郭天财. 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. 生态学报, 2014, 34: 4882-4890

Wang C

, Miao J

, Zhang M

, Ma D

, Feng

, Xie Y

, Guo T

. Effects of post-anthesis high temperature, drought stress and their interaction on the starch pasting properties of two wheat cultivars with different gluten strength. Acta Ecol Sin, 2014, 34: 4882-4890 (in Chinese with English abstract)

To evaluate the effects of post-anthesis high-temperature (HT) and drought stress (DS) and their interactions on starch pasting properties in wheat grains, two winter wheat cultivars differing in gluten strength, Yumai 34 (a strong-gluten cultivar) and Yumai 50 (a weak-gluten cultivar) were investigated in pot experiments at the experimental farm of Henan Agricultural University (E113°35', N34°51'). The experiment used a split-plot design of two factors, with three temperature regimes and two soil water treatments. HT treatments were performed in a climate-controlled greenhouse at 38℃ for 2 days (treatment 2, T 2 ) or 4 days (treatment 3, T 3 ) at 5 days after anthesis (5 DAA), 15 DAA or 25 DAA, while the control was treated at 28℃ (treatment 1, T 1 ). For the two soil water treatments, the soil relative water content was maintained at (55±5)% (W 2 ) for DS treatment and at (75±5)% for the control. The results indicated that both HT and DS and their interactions significantly influenced starch pasting properties in the grains of the two wheat cultivars. However, the responses of pasting parameters to HT and DS were very different between the two wheat cultivars. In the strong-gluten wheat cultivar Yumai 34, DS applied at any grain-filling stage increased most parameters such as peak viscosity, final viscosity and hold-through, among which the final viscosity was significantly improved at the early (5 DAA) and late grain-filling stages (25 DAA), and the peak viscosity was significantly increased at the late grain-filling stage. In Yumai 50, DS applied at any grain-filling stage (early, middle or late) decreased peak viscosity, setback and breakdown. Drought stress applied at the early and middle grain-filling stages (15 DAA) clearly improved hold-through and final viscosity, but significantly decreased both parameters when applied at the late grain-filling stage. Heat stress significantly increased peak viscosity, final viscosity (except when applied at 5 DAA), set-back and breakdown (except when applied at 15 DAA) in grains of the strong-gluten wheat cultivar Yumai 34, while the parameter of hold-through in grains of Yumai 34 had much different responses to HT applied at various stages: it was significantly decreased in both HT treatments (both T 1 and T 2 ) at 5 DAA, and was decreased in T 2 (2 days HT) but significantly enhanced in T 3 (4 days HT) at 15 DAA. In the weak-gluten wheat cultivar Yumai 50, HT significantly decreased hold-through and final viscosity in the grains, but had little influence on peak viscosity and setback. From the treatment stage results, we concluded that HT applied at the middle grain-filling stage (15 DAA) had the greatest impact on pasting properties in both wheat cultivars. The analysis also indicated that interaction between HT and DS had a significant effect on viscosity parameters in both cultivars, which was much greater in Yumai 50 than in Yumai 34. The effect of HT and DS interaction on pasting properties in Yumai 50 was greater when they were applied at the early grain-filling stage compared with the middle or late grain-filling stages. From our results, we conclude that different wheat genotypes show different responses in grain pasting properties to various environmental stresses, implying that such stresses have a complex effect on starch quality in the grains of winter wheat.

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。

1985

0.0

... 3 籽粒淀粉含量及淀粉积累速率测定方法采用双波长法^[15]测定直链、支链淀粉含量, 两者之和为总淀粉含量 ...

2001

0.0

... 1 高温干旱胁迫下小麦籽粒淀粉合成关键酶活性与淀粉积累小麦籽粒的充实过程主要是淀粉的合成与积累过程, 受一系列与淀粉合成有关酶的调控^[16] ...

1989

0.0

... AGPase、SSS、GBSS、SBE和SS被认为是调控淀粉合成的关键酶^[17,18] ...

2010

0.0

曹颖妮, 胡卫国, 王根平, 刘录祥, 王成社. 糯性和非糯性小麦灌浆期胚乳直/支链淀粉积累及其相关酶活性研究. 西北植物学报, 2010, 30: 1995-2001

Cao Y

, Hu W

, Wang G

, Liu L

, Wang C

. Dynamic changes of starch accumulation and enzymes relating to starch biosynthesis of kernel during grain filling in waxy and non-waxy winter wheat. Acta Bot Boreali-Occid Sin, 2010, 30: 1995-2001 (in Chinese with English abstract)

... AGPase、SSS、GBSS、SBE和SS被认为是调控淀粉合成的关键酶^[17,18] ...

2011

0.0

石慧清, 龚月桦, 张东武. 花后高温对持绿型小麦叶片衰老及籽粒淀粉合成相关酶的影响. 植物生态学报, 2011, 35: 769-778

Shi H

, Gong Y

, Zhang D

. Effect of high temperature on leaf senescence and related enzymes of grain starch synthesis in stay-green wheat after anthesis. J Plant Ecol, 2011, 35: 769-778 (in Chinese with English abstract)

... 石慧清等^[19]在大田条件下于花后设置增温棚(增温2~5℃), 观测到高温处理使持绿型小麦品种籽粒SS、AGPase和SSS活性峰值分别下降7 ...

2009

0.0

2001

0.0

2012

0.0

包刚, 覃志豪, 周义, 包玉海, 萨楚拉. 气候变化对中国农业生产影响的模拟评价进展. 中国农学通报, 2012, 28(2): 303-307

Bao

, Tan Z

, Zhou

, Bao Y

, Sa C

. Advances of evaluation of climate changes impact on crop yield. Chin Agric Sci Bull, 2012, 28(2): 303-307 (in Chinese with English abstract)

The assessment of the potential impacts of climate change on agricultural productivity has become a hot issue of many scientific researches, governments and public circles. In recent years, scientists carried out many quantitative studies of evaluation of climate change impact on agricultural system from different aspects and using different methods in this field. In this paper, we mainly addressed the recent advances in evaluation of climate change impacts on crop yields, crop growth and adaptation strategy using crop growth simulation model, and briefly discussed the research tendency and study work required to focus in the future.

气候变化对农业生产影响评价研究已成为众多学者和政府部门关注的热点问题。近几十年来，国内外学者从不同角度用不同方法开展了大量的气候变化对农业生产影响的评估研究。该文重点论述了利用模型模拟方法评价气候变化对中国农业生产、作物生长发育的影响，以及适应对策的最近研究进展，并简要地讨论了该领域研究存在的问题和今后有待进一步深入的研究内容。

... 高温和干旱胁迫对小麦内部结构和功能会造成一定的损伤^[32,33,34], 灌浆期超过30℃的温度显著影响小麦籽粒支链淀粉、直链淀粉和总淀粉含量, 导致粒重和产量下降^{[22,35,36,37]} ...

2006

0.0

赵辉, 戴廷波, 荆奇, 姜东, 曹卫星, 陆玮, 田孝威. 灌浆期高温对两种品质类型小麦品种籽粒淀粉合成关键酶活性的影响. 作物学报, 2006, 32: 423-429

Zhao

, Dai T

, Jing

, Jiang

, Cao W

, Lu

, Tian X

. Effects of high temperature during grain filling on key enzymes involved in starch synthesis in two wheat cultivars with different quality types. Acta Agron Sin, 2006, 32: 423-429 (in Chinese with English abstract)

... 逆境胁迫对籽粒淀粉合成关键酶活性的影响与籽粒发育阶段有关, 高温、干旱诱导了胁迫初期籽粒GBSS、SSS、SBE、AGPase和SS的活性, 而对胁迫后期的酶活性起抑制作用, 从而导致籽粒淀粉积累速率下降^[23,24,25] ...

2008

0.0

闫素辉, 尹燕枰, 李文阳, 梁太波, 李勇, 邬云海, 王平, 耿庆辉, 戴忠民, 王振林. 灌浆期高温对小麦籽粒淀粉的积累、粒度分布及相关酶活性的影响. 作物学报, 2008, 34: 1092-1096

Yan S

, Yin Y

, Li W

, Liang T

, Li

, Wu Y

, Wang

, Geng Q

, Dai Z

, Wang Z

. Effect of high temperature during grain filling on starch accumulation, starch granule distribution, and activities of related enzymes in wheat grains. Acta Agron Sin, 2008, 34: 1092-1096 (in Chinese with English abstract)

In order to investigate the effects of high temperature at early grain filling stages (5–9 d after anthesis) on grain starch formation of wheat ( Triticum aestivum L.), cultivars Jimai 20 (weak heat tolerance) and Lumai 21 (strong heat tolerance), were used with a heat stress treatment in the field by plastic sheds. Starch accumulation, starch granule distribution, and changes of enzymes activities involved in starch synthesis were examined. The results showed that heat stress after anthesis remarkably reduced starch accumulation at maturity. The total starch and amylopectin contents decreased significantly, but amylose content increased in heat treatments compared to control. Accordingly the ratio of amylose to amylopectin of heat treantment was significantly higher than that of control. The effect of high temperature on starch accumulation in Jimai 20 was greater than that in Lumai 21. The volume, number, and surface area percentage of A type starch granule were enhanced significantly, but those of B type starch granule reduced greatly under high temperature. After 5 d heating stress, the sucrose content, the activities of sucrose synthase (SS), adnosine diphosphate glucose pyrophosphorylase (AGPP), soluble starch synthase (SSS), and granule-bounded starch synthase (GBSS) increased in heat-treated grains of Lumai 21, but changed slightly in treated grains of Jimai 20 with no significant difference to control. These indexes of Jimai 20 and Lumai 21 became lower than those of control at 15 and 20 d post anthesis (6 and 11 d after heat stress removal), respectively. The GBSS activity was affected slightly by heat treatment. The results indicate that the changes in grain sucrose content and the activities of SS, AGPP, SSS, and GBSS are companied with the changes in grain starch accumulation, and high temperature restrains starch synthesis at middle and late grain filling stages, which is caused by inferior sucrose providing capacity and low activities of enzymes (SS, AGPP, SSS, and GBSS) involved in starch synthesis.

以不同耐热性品种济麦20和鲁麦21为材料, 于花后5~9 d进行高温处理, 研究了小麦灌浆期高温对籽粒淀粉的积累、粒度分布及合成相关酶活性的影响。结果表明, 灌浆期高温显著降低籽粒淀粉积累量, 显著降低籽粒淀粉、支链淀粉含量, 提高直链淀粉含量和直/支链淀粉比例。高温对济麦20籽粒淀粉积累的影响程度较鲁麦21大。灌浆期高温使小麦籽粒A型淀粉粒的体积、数量和表面积百分比显著增加, B型淀粉粒这3指标则显著降低。高温处理后, 济麦20籽粒蔗糖合酶(SS)、ADPG焦磷酸化酶(AGPP)、可溶性淀粉合酶(SSS)、束缚态淀粉合酶(GBSS)活性与对照无显著差异, 而鲁麦21上述酶活性则高于对照。济麦20、鲁麦21籽粒上述酶活性分别于花后15 d和20 d开始低于对照。与其他淀粉合成相关酶相比, 高温对籽粒GBSS活性的影响程度较小。两品种处理间籽粒蔗糖含量及SS、AGPP、SSS和GBSS活性的变化趋势, 与其籽粒淀粉积累量的变化趋势基本一致。灌浆期高温使籽粒淀粉积累量降低, 主要因高温抑制了籽粒灌浆中后期的淀粉合成, 这是由籽粒蔗糖供应不足和籽粒淀粉合成相关酶活性下降所造成的。

2011

0.0

王征宏, 郭秀璞, 吕淑芳, 李友军, 邓西平. 土壤干旱对不同小麦品种籽粒淀粉积累和相关酶活性的影响. 干旱地区农业研究, 2011, 29: 67-74

Wang Z

, Guo X

, Lyu S

, Li Y

, Deng X

. Effects of soil drought on starch accumulation of grain and activity of corresponding starch synthesis enzymes in different winter wheat varieties. Agric Res Arid Areas, 2011, 29: 67-74 (in Chinese with English abstract)

选用小麦旱地品种长武134和水地品种陕253,研究土壤干旱对小麦籽粒淀粉积累及相关酶活性的影响,探讨土壤干旱影响淀粉积累的酶学机制.结果表明,干旱处理提高了灌浆前期小麦籽粒淀粉积累速率、籽粒SS、AGPase、UGPase、SSS和GBSS酶活性.灌浆后期旱地品种籽粒淀粉积累速率及AGPase、UGPase降低幅度大于水地品种,而SS降低幅度则小于水地品种;旱地品种灌浆后期SSS活性与对照无明显差异,而水地品种SSS活性则大于对照;与其它淀粉合成酶活性相比,干旱对灌浆后期GBSS活性影响程度较小.分析结果表明,SS、AGPase是干旱影响小麦淀粉积累速率的主要调节酶,抗旱性强的小麦品种AGPase的调节作用较大,而抗旱性弱的小麦品种SS的调节作用较大.

2015

0.0

孙立影, 苏玮, 吴宏亮, 李昱, 王雪, 康建宏. 花后不同时期高温对春小麦淀粉形成的影响. 农业科学研究, 2015, 36: 54-60

Sun L

, Su

, Wu H

, Li

, Wang

, Kang J

. Effects of the high temperature after the flowering period on starch formation in spring wheat. J Agric Sci, 2015, 36: 54-60 (in Chinese with English abstract)

以宁夏灌区主栽品种宁春4号和宁春47号为材料,研究花后不同时期高温对春小麦淀粉形成的影响.结果表明:不同时期高温处理使籽粒的直链淀粉、支链淀粉、总淀粉含量呈逐渐上升趋势.处理间比较,高温处理越早支链淀粉含量在后期降低越快,而对直链淀粉含量影响不显著.不同品种间比较,宁春47号受高温胁迫的影响较大,不同品种间籽粒淀粉酶活性变化趋势与淀粉积累趋势基本一致,花后高温处理后籽粒中的ADPG焦磷酸化酶(AGP)、UDPG焦磷酸化酶(UGP)、可溶性淀粉合成酶(SSS)及淀粉合成关键酶(SBE)酶活性下降7%~13%,但AGP、UGP与SSS受高温影响相对较大.花后高温出现越早,对春小麦淀粉形成关键酶和淀粉的含量影响越大,导致灌浆不足,千粒质量降低,产量下降.采取相应栽培技术缓解花后高温危害,可以在一定程度上提高宁夏春小麦的产量.

... 孙立影等^[26]对春小麦花后不同时期施以短暂高温(35#cod#x000b1 ...

2003

0.0

王月福, 于振文, 李尚霞, 余松烈. 小麦籽粒灌浆过程中有关淀粉合成酶的活性及其效应. 作物学报, 2003, 29: 75-81

Wang Y

, Yu Z

, Li S

, Yu S

. Activity of enzymes related to starch synthesis and their effect during the filling of winter wheat. Acta Agron Sin, 2003, 29: 75-81 (in Chinese with English abstract)

在不同土壤肥力条件下, 对不同品质类型小麦品种的籽粒淀粉积累及淀粉合成过程中的关键酶-ADPG焦磷酸化酶、可溶性淀粉合成酶与淀粉粒结合态淀粉合成酶活性变化进行了研究. 结果表明, ADPG焦磷酸化酶、可溶性淀粉合成酶与总淀粉和支链淀粉积累速率呈极显著正相关; 淀粉粒结合态淀粉合成酶与直链淀粉积累速率呈极显著正相关,

... 研究证明, 籽粒SSS、SBE、AGPase和SS活性与支链淀粉的合成具有显著的相关性, 而GBSS、AGPase和SS的活性与直链淀粉合成关系密切^[2,10,27,28] ...

2008

0.0

李建敏, 王振林, 高荣岐, 李圣福, 蔡瑞国, 闫素辉, 于安玲, 尹艳枰. 强、弱筋小麦籽粒形成期蔗糖、淀粉合成相关酶活性及其与氮代谢的关系. 作物学报, 2008, 34: 1019-1026

Li J

, Wang Z

, Gao R

, Li S

, Cai R

, Yan S

, Yu A

, Yin Y

. Activities of enzymes involved in sucrose and starch synthesis during grain filling and the relation to nitrogen metabolism in strong-and weak-gluten wheat cultivars. Acta Agron Sin, 2008, 34: 1019-1026 (in Chinese with English abstract)

... 研究证明, 籽粒SSS、SBE、AGPase和SS活性与支链淀粉的合成具有显著的相关性, 而GBSS、AGPase和SS的活性与直链淀粉合成关系密切^[2,10,27,28] ...

1998

0.0

... 2 高温干旱胁迫下小麦籽粒淀粉积累与产量籽粒淀粉的合成与积累既受遗传因素的影响, 也易受环境条件的影响^[29,30,31] ...

1997

0.0

... 2 高温干旱胁迫下小麦籽粒淀粉积累与产量籽粒淀粉的合成与积累既受遗传因素的影响, 也易受环境条件的影响^[29,30,31] ...

1994

0.0

... 2 高温干旱胁迫下小麦籽粒淀粉积累与产量籽粒淀粉的合成与积累既受遗传因素的影响, 也易受环境条件的影响^[29,30,31] ...

2003

0.0

许振柱, 于振文, 张永丽. 土壤水分对小麦籽粒淀粉合成和积累特性的影响. 作物学报, 2003, 29: 595-600

Xu Z

, Yu Z

, Zhang Y

. The effects of soil moisture on grain starch synthesis and accumulation of winter wheat. Acta Agron Sin, 2003, 29: 595-600 (in Chinese with English abstract)

以防雨池栽的方式研究了限量灌溉对冬小麦籽粒淀粉合成和积累的影响.结果表明,冬小麦籽粒中可溶性淀粉合成酶(SSS)、淀粉粒结合态合成酶(GBSS)和Q酶 (一种分支酶)均是在灌浆中期(花后14 d)活性最高.中度、严重干旱显著地降低了它们的活性.适宜的灌水处理(处理C和处理D)使三种酶保持较高的活性,这是形成较高产量的生理基础.三

2015

0.0

杨晓娟, 居辉, 王治世, 郭安廷, 杨佑明. 花后高温和干旱对冬小麦光合、抗氧化特性及粒重的影响. 麦类作物学报, 2015, 35: 958-963

Yang X

, Ju

, Wang Z

, Guo A

, Yang Y

. Effect of high temperature and drought after anthesis on photosynthesis, antioxidant properties and grain weight in winter wheat. J Trit Crops, 2015, 35: 958-963 (in Chinese with English abstract)

High temperature and drought occur simultaneously during grain filling,which significantly reduce grain starch content and grain yield of wheat. This research was conducted to determine the effects of high temperature,dought and their combination on flag leaf photosynthetic potential,MDA content and activities of antioxidant enzymes in flag leaf and grain starch content and grain weight. Results showed that on the 7th day (DAA21) of stress,high temperature and drought and their combination significantly reduced flag leaf photosynthetic rate and SPAD value,the decline was significantly reduced when the stress was removed. High temperature stress reduced photosynthetic rate and SPAD value,but differences were not significant. The F v / F m of flag leaf was not sensitive to single high temperature or drought,but was sensitive to combined stress significantly. F v / F m values of flag leaf decreased because of the combination of high temperature and drought and represented a smaller degree of recovery after the stress was removed. The MDA content and POD activity of flag leaf continuously increased from DAA14 to DAA21,yet SOD and CAT enzyme activity increased and then decreased with the increase of stress days. From the 7th day (DAA21)，combination stress significantly declined the activity of SOD and CAT enzymes. The MDA content and POD activity of wheat flag leaf were still significantly higher than control under the combination stress. The change tendency of P n ,SPAD,starch content and grain weight were basically similar,which indicated that the decrease of starch content and grain weight associated with membrane lipid peroxidation,chlorophyll degradation and early senescence of flag leaf under high temperature and drought. The damage of photosynthetic performance has been intensified by the high temperature and drought.

为了解花后高温和干旱双重胁迫对小麦的效应，以石麦15为材料，于花后15 d到21 d进行高温（34~36 ℃）和干旱（土壤含水量为田间持水量的40%~45%）处理，研究了高温和干旱胁迫对小麦旗叶光合性能、旗叶丙二醛（MDA）含量和抗氧化酶活性、籽粒淀粉含量和千粒重的影响。结果表明，胁迫第7 d（花后21 d）时，干旱和高温-干旱双重胁迫显著降低了小麦旗叶净光合速率（ P n ）和叶绿素相对含量(SPAD值)，胁迫解除后 P n 降幅缩小，高温胁迫下 P n 和SPAD值与对照的差异不显著。高温-干旱双重胁迫显著降低了旗叶 F v / F m 值，且胁迫解除后恢复程度较小。胁迫期间，旗叶MDA含量和过氧化物酶（POD）活性持续升高，超氧化物歧化酶（SOD）和过氧化氢酶（CAT）活性均呈先升后降趋势，SOD和CAT活性对干旱、高温-干旱双重胁迫更敏感，胁迫第7天时高温-干旱双重胁迫下降幅度最大。旗叶 P n 和SPAD的变化趋势与籽粒淀粉积累量和千粒重变化趋势基本一致，说明灌浆期高温、干旱胁迫下小麦籽粒淀粉积累量和粒重的降低与质膜过氧化、叶绿素的降解和旗叶早衰导致的光合性能下降密切相关，在双重胁迫下光合性能受抑加剧。

2015

0.0

苏玮, 孙立影, 康建宏, 吴宏亮, 李昱. 高温干旱对春小麦光合特性及产量的影响. 农业科学研究, 2015, 36: 49-53

, Sun L

, Kang J

, Wu H

, Li

. Effect of high temperature and drought treatment on photosynthetic characteristics and yield of different varieties of spring wheat. J Agric Sci, 2015, 36: 49-53 (in Chinese with English abstract)

1991

0.0

2004

0.0

2003

0.0

2008

0.0

胡吉帮, 王晨阳, 郭天财, 苗建利, 朱云集, 贺德先. 灌浆期高温和干旱对小麦灌浆特性的影响. 河南农业大学学报, 2008, 42: 597-601

Hu J

, Wang C

, Guo T

, Miao J

, Zhu Y

, He D

. Effects of high temperature and drought stress on grain filling characteristics in wheat during grain filling period. J Henan Agric Univ, 2008, 42: 597-601 (in Chinese with English abstract)

在盆栽和人工气候箱模拟条件下,研究了高温与干旱胁迫对两个不同筋力小麦品种豫麦34号(强筋) 和豫麦50号(弱筋) 灌浆特性的影响.结果表明,高温胁迫抑制了子粒灌浆速率,使最大灌浆速率出现时间提前,灌浆持续期缩短,导致子粒干物质积累量下降.不同时段高温胁迫对粒重的影响有差异,豫麦34号和豫麦50号分别以前期和中期受高温胁迫影响最大,后期相对小.豫麦34号以中期干旱胁迫粒重降幅最大,而豫麦50号在前期受干旱胁迫影响最大,后期干旱使两品种的粒重略有增加.高温和干旱对小麦子粒灌浆特性及粒重的影响存在明显的水、温互作效应.研究还表明,渐增期平均灌浆速率的降低及快增期、缓增期持续时间的缩短是最终导致粒重降低的主要原因.

... 胡吉帮等^[38]研究表明, 灌浆前中期短暂高温(38℃, 2~4 d)可导致小麦的成熟期提前3~5 d ...

2015

0.0

李春艳, 付凯勇, 张润琪, 徐芳芳, 史晓艳, 朱永琪, 覃安祥, 李诚. 灌浆期高温对冬小麦淀粉粒发育的影响. 麦类作物学报, 2015, 35: 1395-1402

Li C

, Fu K

, Zhang R

, Xu F

, Shi X

, Zhu Y

, Qin A

, Li

. Effect of high temperature post anthesis on the development of starch granules in winter wheat. J Trit Crops, 2015, 35: 1395-1402 (in Chinese with English abstract)

... 试验结果还表明, 高温胁迫初期会提高淀粉积累速率, 但随着胁迫时间的延长, 籽粒发育进程加快, 成熟期提前, 最终淀粉积累速率下降, 直链、支链和总淀粉含量低于对照, 粒重降低, 这与李春艳等^[39]研究结果一致 ...