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.
Keywords：winter wheat;compound stress;high temperature;drought;starch accumulation;starch synthesis enzymes -->0 PDF (660KB)元数据多维度评价相关文章收藏文章 本文引用格式导出EndNoteRisBibtex收藏本文--> 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[J]. 作物学报, 2018, 44(04): 591-600 https://doi.org/10.3724/SP.J.1006.2018.00591 HUYang-Yang, LUHong-Fang, LIUWei-Xing, KANGJuan, MAGeng, LISha-Sha, CHUYing-Ying, WANGChen-Yang. Effects of High Temperature and Water Deficiency during Grain Filling on Activities of Key Starch Synthesis Enzymes and Starch Accumulation in Wheat[J]. Acta Agronomica Sinica, 2018, 44(04): 591-600 https://doi.org/10.3724/SP.J.1006.2018.00591 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。
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)。 Table 1 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 1Variance analysis on interaction of HT and DS on activity of key starch synthesis enzymes (F-value)
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**
*和**分别表示在0.05和0.01概率水平(双侧)上差异显著。DAA: 开花后天数; DS×HT: 干旱与高温互作。* Significant difference at the 0.05 probability level; ** significant difference at the 0.01 probability level. DAA: days after anthesis; DS: drought stress; HT: high temperature stress; DS×HT: interaction of HT and DS. 新窗口打开
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。 显示原图|下载原图ZIP|生成PPT 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. 2Changes 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)。 显示原图|下载原图ZIP|生成PPT 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. 3Changes 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。 Table 2 表2 表2不同处理成熟期小麦籽粒淀粉及其组分含量 Table 2Starch content and its components in wheat grain under different treatments at maturity
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
数据后不同字母表示处理间差异显著(P<0.05)。HT+DS: 高温与干旱复合胁迫。Values within a column followed by different letters are significantly different among treatments at P < 0.05. DS: drought stress; HT: high temperature stress; HT+DS: compound stress of HT and DS. 新窗口打开
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%。 Table 3 表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 3Effects of different treatments on grain yield and its components of wheat
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
数据后不同字母表示处理间差异显著(P<0.05)。SN: 穗数; GNS: 穗粒数; TGW: 千粒重; GY: 籽粒产量; HT+DS: 高温与干旱复合胁迫。Values within a column followed by different letters are significantly different among treatments at P < 0.05. SN: spike number; GNS: grain number per spike; TGW: thousand-grain weight; GY: grain yield; DS: drought stress; HT: high temperature stress; HT+DS: compound stress of HT and DS. 新窗口打开
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。 Table 4 表4 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 4Correlation coefficients of activities of key starch synthesis enzymes with contents of starch and its components
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**
*和**分别表示在0.05和0.01概率水平显著相关。* and ** indicate significant correlation at the 0.05 and 0.01 probability levels, respectively. 新窗口打开
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。
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