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作物学报 ›› 2018, Vol. 44 ›› Issue (04): 591-600.doi: 10.3724/SP.J.1006.2018.00591

• 耕作栽培·生理生化 • 上一篇    下一篇

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. 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
  • 收稿日期:2017-07-02 接受日期:2018-01-08 出版日期:2018-01-23 网络出版日期:2018-01-23
  • 通讯作者: 胡阳阳,卢红芳,王晨阳
  • 作者简介:

    18238802035@163.com

  • 基金资助:
    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

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

Yang-Yang HU1,**(), Hong-Fang LU1,**, Wei-Xing LIU1, Juan KANG1, Geng MA1, Sha-Sha LI1, Ying-Ying CHU1, Chen-Yang WANG1,2,*()   

  1. 1 Henan Agricultural University, Zhengzhou 450002, Henan, China
    2 National Wheat Engineering Research Center, Zhengzhou 450002, Henan, China
  • Received:2017-07-02 Accepted:2018-01-08 Published:2018-01-23 Published online:2018-01-23
  • Contact: Yang-Yang HU,Hong-Fang LU,Chen-Yang WANG
  • Supported by:
    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).

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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

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.

Key words: winter wheat, compound stress, high temperature, drought, starch accumulation, starch synthesis enzymes

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。"

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)"

酶
Enzyme		 来源
Source		 2015-2016 2016-2017
14DAA 18DAA 22DAA 26DAA 14DAA 18DAA 22DAA 26DAA
GBSS 高温HT 22.8** 12.7** 1290.8** 4956.8** 12.6** 19.8** 728.5** 3297.7**
干旱DS 1.5 142.6** 139.3** 5.0 0.3 46.6** 78.8** 40.5**
DS×HT 8.3* 301.4** 199.3** 134.1** 11.4** 63.1** 158.1** 147.6**
SSS 高温HT 319.1** 213.3** 216.0** 344.1** 113.7** 50.0** 277.7** 902.8**
干旱DS 9.1* 91.6** 42.9** 49.2** 10.0* 72.5** 0.3 130.2**
DS×HT 0.0 168.7** 50.4** 10.8* 2.2 148.7** 42.7** 11.6**
SBE 高温HT 0.6 141.4** 531.3** 116.6** 0.3 190.5** 174.8** 621.4**
干旱DS 46.5** 114.3** 560.8** 84.4** 17.9** 127.6** 186.2** 368.0**
DS×HT 20.9** 13.2** 99.8** 0.7 7.0** 7.6* 11.5** 9.2*
AGPase 高温HT 1.5 10.0* 433.4** 358.7** 0.3 6.3* 1682.4** 1390.1**
干旱DS 6.0* 98.0** 216.2** 223.9** 2.2 107.6** 194.6** 726.5**
DS×HT 15.0** 20.7** 3.3 26.2** 7.1* 20.7** 0.1 27.4**
SS 高温HT 174.0** 40.1** 449.2** 378.9** 154.8** 52.9** 66.0** 297.3**
干旱DS 13.3** 3.8 25.2** 51.0** 23.6** 9.6* 34.4** 146.4**
DS×HT 43.5** 5.6* 0.2 20.4** 99.2** 9.0* 1.1 11.5**

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。"

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。"

表2

不同处理成熟期小麦籽粒淀粉及其组分含量"

处理
Treatment		 2015-2016 2016-2017
直链淀粉Amylose
(%)		 支链淀粉Amylopectin (%) 总淀粉
Total starch
(%)		 直/支比
Amylose/ amylopectin		 直链淀粉Amylose
(%)		 支链淀粉Amylopectin (%) 总淀粉
Total starch
(%)		 直/支比
Amylose/
amylopectin
对照CK 15.9 a 50.8 a 66.7 a 0.31 c 15.6 a 50.0 a 65.6 a 0.31 c
干旱DS 15.3 b 47.1 b 62.4 b 0.32 b 15.0 b 46.9 b 61.9 b 0.32 bc
高温HT 14.2 c 43.8 c 58.1 c 0.33 b 14.0 c 43.2 c 57.3 c 0.32 ab
HT+DS 13.6 d 40.8 d 54.4 d 0.33 a 13.3 d 40.6 d 53.9 d 0.33 a

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"

处理
Treatment		 2015-2016 2016-2017
SN (pot-1) GNP TGW (g) GY (g pot-1) SN (pot-1) GNP TGW (g) GY (g pot-1)
对照CK 35.7 a 45.5 a 48.1 a 72.6 a 43.0 a 46.0 a 42.9 a 67.7 a
干旱DS 37.0 a 43.3 a 41.5 b 59.3 b 42.3 a 46.6 a 36.7 b 55.4 b
高温HT 38.0 a 45.8 a 37.3 c 56.9 b 39.3 a 43.5 a 35.4 c 53.8 b
HT+DS 38.0 a 46.5 a 30.5 d 48.2 c 40.7 a 46.1 a 28.6 d 41.9 c

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"

淀粉
Amylose		 淀粉合成关键酶
Key starch synthesis
enzymes		 开花后天数 Days after anthesis (DAA)
花后14 d
14 DAA		 花后18 d
18 DAA		 花后22 d
22 DAA		 花后26 d
26 DAA
2015-2016
支链淀粉含量
Amylopectin content		 GBSS 0.792** 0.420 0.767** 0.830**
SSS 0.962** 0.668* 0.322 0.938**
SBE 0.031 -0.274 0.824** 0.941**
AGP -0.316 0.165 0.903** 0.943**
SS -0.792** -0.500 0.818** 0.939**
直链淀粉含量
Amylose content		 GBSS 0.881** 0.516 0.552 0.825**
SSS 0.853** 0.852** 0.157 0.884**
SBE 0.133 -0.477 0.457 0.867**
AGP -0.314 0.174 0.547 0.883**
SS -0.685* -0.320 0.310 0.905**
总淀粉含量
Total starch content		 GBSS 0.820** 0.530 0.812** 0.836**
SSS 0.953** 0.881** 0.291 0.934**
SBE 0.051 -0.417 0.818** 0.933**
AGP -0.320 0.189 0.916** 0.943**
SS -0.781** -0.546 0.744** 0.940**
2016-2017
支链淀粉含量
Amylopectin content		 GBSS 0.711** 0.031 0.830** 0.909**
SSS 0.880** 0.637* 0.669* 0.953**
SBE 0.184 -0.305 0.914** 0.968**
AGP -0.099 0.274 0.884** 0.968**
SS -0.673* -0.531 0.925** 0.962**
直链淀粉含量
Amylose content		 GBSS 0.829** 0.829** 0.550 0.791**
SSS 0.756** 0.756** 0.355 0.759**
SBE 0.309 0.309 0.456 0.836**
AGP -0.225 -0.225 0.380 0.809**
SS -0.463 -0.463 0.526 0.834**
总淀粉含量
Total starch content		 GBSS 0.739** 0.042 0.821** 0.902**
SSS 0.864** 0.716** 0.643* 0.926**
SBE 0.209 -0.376 0.870** 0.959**
AGP -0.123 0.244 0.828** 0.951**
SS -0.640* -0.553 0.895** 0.953**
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