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Acta Agronomica Sinica ›› 2018, Vol. 44 ›› Issue (04): 554-568.doi: 10.3724/SP.J.1006.2018.00554

• TILLAGE & CULTIVATION·PHYSIOLOGY & BIOCHEMISTRY • Previous Articles     Next Articles

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 Online:2018-01-26 Published:2018-01-26
  • Contact: Jian-Chang YANG E-mail:xuyunji19881004@163.com;jcyang@yzu.edu.cn
  • 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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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

Fig. 1

Changes in soil water potential under alternate wetting and drying irrigation (AWD) during grain filling of riceWMD: alternate wetting and moderate soil drying; WSD: alternate wetting and severe soil drying; D1 and D2 represent soil water potential at -15 kPa and -30 kPa, respectively, under WMD and WSD. Arrows indicate the re-watering period."

Fig. 2

Effect of alternate wetting and drying irrigation on water potential of the flag leaf during grain filling of rice CI: conventional irrigation; WMD: alternate wetting and moderate soil drying; WSD: alternate wetting and severe soil drying. D1 and D2 represent soil water potential at -15 kPa and -30 kPa, respectively, under WMD and WSD. W1 and W2 indicate the re-watering period."

Table 1

Effect of alternate wetting and drying irrigation on grain yield and its components of rice"

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

Fig. 3

Effect of alternate wetting and drying irrigation on grain weight and grain filling rate for superior and inferior spikelets of riceCI: conventional irrigation; WMD: alternate wetting and moderate soil drying; WSD: alternate wetting and severe soil drying; S: superior spikelets; I: inferior spikelets."

Table 2

Effect of alternate wetting and drying irrigation on grain filling parameters of superior and inferior spikelets of rice"

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

Fig. 4

Effect of alternate wetting and drying irrigation on activities of sucrose synthase (SuSase), adenosine diphosphate glucose pyrophosphorylase (AGPase), starch synthase (StSase), and starch branching enzyme (SBE) in rice grainsCI: conventional irrigation; WMD: alternate wetting and moderate soil drying; WSD: alternate wetting and severe soil drying. S: superior spikelets; I: inferior spikelets. D1 and D2 represent soil water potential at -15 kPa and -30 kPa, respectively, under WMD and WSD. W1 and W2 indicate the re-watering period."

Table 3

Effect of alternate wetting and drying irrigation on photosynthetic rate of flag leaf of rice"

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

Fig. 5

Effect of alternate wetting and drying irrigation on SOD activity and MDA content of flag leaf of riceCI: conventional irrigation; WMD: alternate wetting and moderate soil drying; WSD: alternate wetting and severe soil drying. D1 and D2 represent soil water potential at -15 kPa and -30 kPa, respectively, under WMD and WSD. W1 and W2 indicate the re-watering period."

Table 4

Effect of alternate wetting and drying irrigation on NSC accumulation and remobilization of rice"

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

Fig. 6

Changes in starch content in rice stemsCI: conventional irrigation; WMD: alternate wetting and moderate soil drying; WSD: alternate wetting and severe soil drying."

Fig. 7

Changes in hydrolytic enzymes in rice stemsCI: conventional irrigation; WMD: alternate wetting and moderate soil drying; WSD: alternate wetting and severe soil drying. D1 and D2 represent soil water potential at -15 kPa and -30 kPa, respectively, under WMD and WSD. W1 and W2 indicate the re-watering period."

Fig. 8

Changes in 13C-distribution in rice stems and grains during grain fillingCI: conventional irrigation; WMD: alternate wetting and moderate soil drying; WSD: alternate wetting and severe soil drying."

Table 5

Proportion of 13C-distribution in rice leaves during grain filling"

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

Table 6

Correlation coefficients of key enzyme activities in sucrose-starch pathway and photosynthetic characteristics of flag leaf with grain weight and grain filling rate"

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**

Table 7

Correlation coefficients of starch hydrolytic enzymes activities in stems with carbon remobilization"

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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