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作物学报 ›› 2018, Vol. 44 ›› Issue (04): 554-568.doi: 10.3724/SP.J.1006.2018.00554

• 耕作栽培·生理生化 • 上一篇    下一篇

干湿交替灌溉对水稻花后同化物转运和籽粒灌浆的影响

徐云姬1,2(), 许阳东1, 李银银1, 钱希旸1, 王志琴1, 杨建昌1,*()   

  1. 1 扬州大学江苏省作物遗传生理重点实验室 / 粮食作物现代产业技术协同创新中心, 江苏扬州 225009
    2 扬州大学教育部农业与农产品安全国际合作联合实验室, 江苏扬州 225009
  • 收稿日期:2018-09-04 接受日期:2018-01-08 出版日期:2018-01-26 网络出版日期:2018-01-26
  • 通讯作者: 杨建昌
  • 作者简介:

    xuyunji19881004@163.com

  • 基金资助:
    本研究由国家自然科学基金项目(31461143015, 31471438), 国家科技支撑计划项目(2014AA10A605), 国家重点研发计划(2016YFD0300206-4), 江苏高校优势学科建设工程项目(PAPD)和扬州大学高端人才支持计划项目(2015-1)资助

Effect of Alternate Wetting and Drying Irrigation on Post-anthesis Remobilization of Assimilates and Grain Filling of Rice

Yun-Ji XU1,2(), Yang-Dong XU1, Yin-Yin LI1, Xi-Yang QIAN1, Zhi-Qin WANG1, Jian-Chang YANG1,*()   

  1. 1 Key Laboratory of Crop Genetics and Physiology of Jiangsu Province / Yangzhou University, Yangzhou 225009, Jiangsu, China
    2 Joint International Research Laboratory of Agriculture and Agri-product Safety, the Ministry of Education of China, Yangzhou University, Yangzhou 225009, Jiangsu, China
  • Received:2018-09-04 Accepted:2018-01-08 Published:2018-01-26 Published online:2018-01-26
  • Contact: Jian-Chang YANG
  • Supported by:
    This study was supported by the grants from the National Natural Science Foundation of China (31461143015, 31471438), the National Key Technology Support Program of China (2014AA10A605), the National Key Research and Development Support Program of China (2016YFD0300206-4), the Priority Academic Program Development of Jiangsu Higher Education Institutions (PAPD), and the Top Talent Supporting Program of Yangzhou University (2015-01).

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摘要：

为阐明干湿交替灌溉对水稻花后同化物转运和籽粒灌浆的影响及其生理原因, 大田种植扬两优6号、武运粳24和旱优8号3个水稻品种, 自移栽后10 d起, 设置常规灌溉(CI)、轻干-湿交替灌溉(WMD)和重干-湿交替灌溉(WSD) 3种灌溉模式, 观察产量及构成因素、强弱势粒灌浆动态、籽粒中酶活性变化、剑叶光合性能、茎鞘非结构性碳水化合物(NSC)运转及其淀粉水解酶活性变化, 并利用13C同位素示踪茎鞘物质运转动态。结果表明, 与CI相比, WMD显著增加了3个水稻品种的每穗粒数、千粒重和结实率, 进而提高籽粒产量。WMD显著提高千粒重和结实率的主要原因是促进了弱势粒的灌浆。WMD和WSD显著增强了茎鞘α-淀粉酶和β-淀粉酶活性, 促进同化物质再运转与分配, 提高茎鞘储藏物质对粒重的贡献率。其中WMD提高了剑叶光合性能以及增强了水稻弱势粒中蔗糖-淀粉代谢途径关键酶活性, WSD的作用相反。表明较好的叶片性能、花后茎中较多的同化物向籽粒转运、较高的茎鞘淀粉水解酶活性以及弱势粒中较高的糖代谢酶活性是WMD促进弱势粒灌浆的重要生理原因。

关键词: 水稻, 干湿交替灌溉, 籽粒灌浆, 13C同位素示踪, 强势粒, 弱势粒, 茎鞘物质运转

Abstract:

Alternate wetting and drying irrigation (AWD) has been widely adopted in rice production for saving water and increasing water use efficiency. However, there is limited information about how AWD affects post-anthesis remobilization of assimilates and grain filling. To elucidate this issue, we planted three rice cultivars, including Yangliangyou 6 (indica hybrid), Wuyunjing 24 (japonica) and Hanyou 8 (japonica) in the field. With treatments of conventional irrigation (CI), alternate wetting and moderate soil drying irrigation (WMD) and alternate wetting and severe soil drying irrigation (WSD) from 10 days after transplanting to maturity. Grain yield and its components, grain filling of superior and inferior spikelets, changes in activities of the key enzymes involved in the conversion from sucrose to starch in grains, photosynthetic traits of flag leaf, the remobilization of non-structural carbohydrates in stems (culms and sheaths) and changes in starch hydrolytic enzymes in stems were investigated and isotope 13C was applied to trace redistribution of stem reserves. WMD significantly increased number of spikelets per panicle, 1000-grain weight, percentage of filled grains and grain yield of all the tested cultivars as compared with CI. Increases in 1000-grain weight and percentage of filled grains under WMD were mainly due to the enhancement of grain filling in inferior spikelets. Both WMD and WSD significantly enhanced the activities of α-amylase and β-amylase in stems and promoted translocation and redistribution of stem reserves, and increased the contribution of reserved carbohydrates in stems to grain yield. Moreover, WMD strengthened photosynthetic efficiency of flag leaf and enhanced the activities of the key enzymes involved in the conversion from sucrose to starch in inferior spikelets, whereas WSD exhibited the opposite effects. The results suggest that better leaf performance and higher activities of starch hydrolytic enzymes in stems, more remobilization of assimilates from stems to grains, and stronger activities of the key enzymes involved in sugar metabolism in inferior spikelets under the WMD are important physiological reasons for the enhancement of grain-filling in inferior spikelets of rice.

Key words: rice, alternate wetting and drying irrigation, grain filling, tracer of isotope 13C, superior spikelets, inferior spikelets, remobilization of reserves in stems

图1

干湿交替灌溉处理下水稻灌浆期土壤水势的变化 WMD: 轻干-湿交替灌溉; WSD: 重干-湿交替灌溉; D1和D2表示土壤落干期; 箭头指示复水期。"

图2

干湿交替灌溉对灌浆期剑叶水势的影响 CI: 常规灌溉; WMD: 轻干-湿交替灌溉; WSD: 重干-湿交替灌溉。D1和D2表示土壤落干期; W1和W2表示复水期。"

表1

干湿交替灌溉对供试水稻产量及其构成因素的影响"

品种
Cultivar		 灌溉
方式
Irrigation regime		 产量
Grain yield
(t hm-2)		 穗数
Panicles
(×104 hm-2)		 每穗粒数
Number of spikelets per panicle		 千粒重
1000-grain weight
(g)		 结实率
Filled grains
(%)		 整个生长期
灌溉水量
Irrigation water during the whole growing season (mm)		 抽穗后
灌溉水量
Irrigation water after heading (mm)		 灌溉水
利用效率
Water use
efficiency for
irrigation
(kg grain m-3)
扬两优6号 CI 9.83 b 218.88 a 189.50 b 28.40 b 83.42 b 922 a 286 a 1.07 c
Yangliangyou 6 WMD 10.88 a 205.83 b 206.04 a 29.96 a 85.63 a 704 b 140 b 1.55 a
WSD 7.22 c 186.69 c 177.40 c 26.88 c 81.06 c 516 c 108 c 1.40 b
武运粳24 CI 8.76 b 230.25 a 175.13 b 26.06 b 83.35 b 914 a 275 a 0.96 c
Wuyunjing 24 WMD 9.72 a 210.17 b 188.68 a 27.96 a 87.68 a 722 b 124 b 1.35 a
WSD 5.99 c 189.18 c 160.30 c 24.36 c 81.12 c 536 c 104 c 1.12 b
旱优8号 CI 9.03 b 236.01 a 172.84 b 25.20 b 87.84 b 908 a 279 a 0.99 c
Hanyou 8 WMD 9.73 a 220.47 b 183.71 a 26.85 a 89.45 a 687 b 133 b 1.42 a
WSD 6.97 c 208.85 c 162.42 c 24.16 c 85.02 c 528 c 94 c 1.32 b

图3

干湿交替灌溉对供试水稻品种强、弱势粒增重动态和籽粒灌浆速率的影响 CI: 常规灌溉; WMD: 轻干-湿交替灌溉; WSD: 重干-湿交替灌溉; S: 强势粒; I: 弱势粒。"

表2

干湿交替灌溉对水稻强、弱势粒灌浆特征参数的影响"

品种
Cultivar		 灌溉方式
Irrigation regime		 粒位
Grain position		 粒重
Grain weight (A)
(mg grain-1)		 最大灌浆速率
The maximum grain-filling rate (Gmax) (mg grain-1 d-1)		 平均灌浆速率
The mean grain-filling rate (Gmean) (mg grain-1 d-1)
扬两优6号 CI S 27.34 a 1.87 a 1.19 a
Yangliangyou 6 I 19.06 d 1.27 d 0.84 d
WMD S 27.64 a 1.89 a 1.21 a
I 21.60 c 1.32 c 0.95 c
WSD S 24.82 b 1.61 b 1.03 b
I 15.00 e 1.01 e 0.67 e
武运粳24 CI S 26.84 a 1.89 a 1.22 a
Wuyunjing 24 I 16.91 d 1.16 d 0.77 d
WMD S 26.99 a 1.84 a 1.18 a
I 18.80 c 1.32 c 0.88 c
WSD S 24.33 b 1.69 b 1.09 b
I 15.00 e 1.01 e 0.67 e
旱优8号 CI S 26.85 a 1.88 a 1.13 a
Hanyou 8 I 16.78 d 1.21 d 0.81 d
WMD S 26.92 a 1.86 a 1.16 a
I 18.56 c 1.33 c 0.92 c
WSD S 24.67 b 1.63 b 1.08 b
I 15.19 e 1.00 e 0.70 e

图4

干湿交替灌溉对水稻籽粒中蔗糖合酶(SuSase, a~c)、腺苷二磷酸葡萄糖焦磷酸化酶(AGPase, d~f)、淀粉合酶(StSase, g~i)和淀粉分支酶(SBE, j~l)活性的影响 CI: 常规灌溉; WMD: 轻干-湿交替灌溉; WSD: 重干-湿交替灌溉。S: 强势粒; I: 弱势粒。D1和D2表示土壤落干期。W1和W2表示复水期。"

表3

干湿交替灌溉对水稻剑叶光合速率的影响"

品种
Cultivar		 灌溉方式
Irrigation regime		 光合速率 Photosynthetic rate (μmol m-2 s-1)
D1 W1 D2 W2
扬两优6号 CI 20.59 a 21.13 b 15.89 a 15.96 b
Yangliangyou 6 WMD 21.00 a 24.08 a 16.20 a 17.81 a
WSD 18.42 b 19.43 c 13.37 b 14.16 c
武运粳24 CI 20.61 a 20.30 b 16.23 a 16.56 b
Wuyunjing 24 WMD 20.75 a 23.05 a 16.56 a 18.03 a
WSD 19.02 b 18.72 c 14.03 b 14.81 c
旱优8号 CI 21.38 a 20.43 b 15.92 a 15.38 b
Hanyou 8 WMD 20.83 a 22.94 a 16.11 a 17.09 a
WSD 18.72 b 18.22 c 14.63 b 12.27 c

图5

干湿交替灌溉对供试水稻品种剑叶超氧化物歧化酶(SOD)活性和丙二醛(MDA)含量的影响 CI: 常规灌溉; WMD: 轻干-湿交替灌溉; WSD: 重干-湿交替灌溉。D1和D2表示土壤落干期, W1和W2表示复水期。"

表4

干湿交替灌溉对供试水稻品种茎鞘NSC积累和运转的影响"

品种
Cultivar		 灌溉方式
Irrigation regime		 茎鞘NSC积累 NSC accumulation in stems (g m-2) NSC运转率
NSC remobilization (%)		 NSC对籽粒产量
的贡献率
Contribution of NSC to grain yield (%)
抽穗期
Heading		 成熟期
Maturity		 抽穗-成熟
Heading-maturity
扬两优6号 CI 282.89 a 214.05 a 68.84 c 24.33 b 7.00 c
Yangliangyou 6 WMD 275.06 b 170.45 b 104.61 a 38.03 a 9.61 b
WSD 238.15 c 147.12 c 91.03 b 38.22 a 12.60 a
武运粳24 CI 248.12 a 192.58 a 55.54 b 22.38 b 6.34 c
Wuyunjing 24 WMD 236.84 b 152.54 b 84.30 a 35.59 a 8.67 b
WSD 224.85 c 144.26 c 80.59 a 35.84 a 13.45 a
旱优8号 CI 256.00 a 198.13 a 57.87 c 22.61 b 6.41 c
Hanyou 8 WMD 241.88 b 156.02 b 85.86 a 35.50 a 8.83 b
WSD 220.59 c 140.58 c 80.01 b 36.27 a 11.48 a

图6

水稻花后茎鞘中淀粉含量变化 CI: 常规灌溉; WMD: 轻干-湿交替灌溉; WSD: 重干-湿交替灌溉。"

图7

水稻花后茎鞘中淀粉水解酶活性变化 CI: 常规灌溉; WMD: 轻干-湿交替灌溉; WSD: 重干-湿交替灌溉。D1和D2表示土壤落干期, W1和W2表示复水期。"

图8

水稻灌浆期茎鞘和籽粒中13C分配的变化 CI: 常规灌溉; WMD: 轻干-湿交替灌溉; WSD: 重干-湿交替灌溉。"

表5

水稻灌浆期叶片中13C的分配比例"

品种
Cultivar		 灌溉方式
Irrigation regime		 叶片中13C分配比例 13C-distribution in rice leaves (%)
抽穗期 Heading 花后10 d 10 DPA 花后30 d 30 DPA
扬两优6号 CI 15.67 a 11.88 a 4.62 a
Yangliangyou 6 WMD 16.20 a 9.74 b 3.04 b
WSD 16.89 a 7.00 c 1.97 c
武运粳24 CI 18.25 a 12.39 a 5.12 a
Wuyunjing 24 WMD 17.88 a 11.86 a 2.38 b
WSD 18.43 a 6.87 b 2.00 b
旱优8号 CI 19.03 a 14.21 a 6.05 a
Hanyou 8 WMD 18.41 a 13.69 ab 4.97 b
WSD 19.22 a 12.43 b 4.33 b

表6

籽粒中蔗糖-淀粉代谢途径关键酶活性和剑叶光合性能与籽粒灌浆的相关系数"

项目
Item		 粒重
Grain weight		 最大灌浆速率
The maximum grain-filling rate (Gmax)		 平均灌浆速率
The mean grain-filling rate (Gmean)
蔗糖合酶活性 SuSase activity 0.56** 0.55** 0.59**
腺苷二磷酸葡萄糖焦磷酸化酶活性 AGPase activity 0.59** 0.62** 0.65**
淀粉合酶活性 StSase activity 0.55** 0.58** 0.57**
淀粉分支酶活性 SBE activity 0.35 0.40 0.43
剑叶光合速率 Photosynthetic rate of flag leaf 0.82** 0.91** 0.89**
剑叶超氧化物歧化酶活性 SOD activity of flag leaf 0.87** 0.94** 0.93**
剑叶丙二醛含量 MDA content of flag leaf -0.82** -0.88** -0.90**

表7

茎鞘中淀粉水解酶活性与物质运转的相关系数"

与相关
Correlations with		 茎鞘中α-淀粉酶活性
α-amylase activity in stems		 茎鞘中β-淀粉酶活性
β-amylase activity in stems
茎鞘中NSC运转 NSC remobilization 0.90** 0.67*
茎鞘中NSC对产量的贡献率 Contribution of NSC to grain yield 0.93** 0.80**
茎鞘中淀粉运转 Starch remobilization in stems 0.94** 0.79**
茎鞘中13C 13C in stems -0.92** -0.77**
茎鞘中13C运转 13C remobilization in grains 0.92** 0.79**
籽粒中13C 13C in grains 0.91** 0.84**
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