Effects of Gibberellin and 2, 4-epbrassinolide on Grain-filling and Yield of Tartary Buckwheat
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摘要:
目的 明确赤霉素(GA3)与2,4-表油菜素内酯(EBR)对苦荞灌浆及产量形成的影响。 方法 设置GA3和EBR不同配比6个处理,研究其对苦荞强弱籽粒之间灌浆特性和淀粉合成酶活性之间的差异,灌浆阶段叶片中抗氧化酶活性和光合特性的变化,以及对农艺性状及产量的影响。 结果 单施GA3处理与清水对照相比,显著增加了苦荞的百粒重以及籽粒中腺苷二磷酸葡萄糖焦磷酸化酶和可溶性淀粉合成酶的活性、叶片的抗氧化酶活性、净光合速率、气孔导度和蒸腾速率,显著提高了单株及最终的产量。80 mg·L−1 GA3配施不同质量浓度的EBR处理时,随EBR施用量的增加,苦荞强势粒的淀粉合成酶活性、叶片抗氧化酶活性、光合特性、主茎分枝数、主茎直径、单株粒数、单株粒重和产量均表现为先增加后降低;苦荞的主茎节数和株高呈持续增加的变化趋势;弱势粒的淀粉合成酶活性则呈持续下降的变化趋势。 结论 赤霉素配施适宜的2,4-表油菜素内酯处理(80 mg·L−1 GA3+0.5 mg·L−1 EBR)能促进苦荞强势粒的灌浆,延缓叶片衰老,提高粒重与最终的产量。 -
关键词:
- 苦荞 /
- 2,4-表油菜素内酯 /
- 赤霉素 /
- 产量
Abstract:Objective Effects of gibberellic acid (GA3) and 2, 4-ebrassinolide (EBR) on grain-filling and yield of Tartary buckwheat were studied. Method Grain-filling and starch synthase activity of dominant and weak Tartary buckwheat grains under 6 treatments of GA3 and/or EBR were compared. Antioxidant enzyme activity and photosynthetic properties of the leaves during grain-filling stage and agronomic traits associated with grain yield of the plants were monitored. Result Compared to control, GA3 alone significantly increased the 100-grain weight and activities of adenosine diphosphate glucose coke phosphorylase and soluble starch synthase in the grains as well as the antioxidant enzyme activities, net photosynthetic rate, stomatal conductance, and transpiration rate in the leaves. It also significantly increased the yield per plant and final grain yield. Increasing EBR application, on the other hand, increased the activities of starch synthase, leaf antioxidant enzymes, leaf photosynthetic characteristics, main stem branch number and diameter, grains per plant, grain weight per plant, and final grain yield initially but declined after reaching a peak. Meanwhile, the number of main stem nodes and plant height rose and the starch synthase activity of the weak grains lowered continuously. Conclusion By combining GA3 at the rate of 80 mg·L−1 and EBR at 0.5 mg·L−1 for treating Tartary buckwheat, the grain-filling of dominant grains was enhanced, the leaf senescence delayed, and the grain weight and final yield increased. -
Key words:
- Tartary buckwheat /
- 2,4-epbrassinolide /
- gibberellin /
- yield
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表 1 赤霉素与2,4-表油菜素内酯对苦荞籽粒增重过程的影响
Table 1. Effect of GA3 and EBR treatments on weight gain of buckwheat grains
粒位
Grain position处理
Treatment籽粒百粒干重 Dry weight of 100 grains/g 7 d 14 d 21 d 28 d 35 d 强势粒 Superior grain CK 0.47e 1.81e 2.07c 2.35b 2.26d T1 0.54c 1.87d 2.11b 2.36b 2.35c T2 0.54c 2.08b 2.12b 2.43a 2.45b T3 0.64a 2.18a 2.23a 2.44a 2.49a T4 0.58b 2.09b 2.11b 2.43a 2.46b T5 0.50d 2.01c 2.04d 2.36b 2.33c 弱势粒 Inferior grain CK 0.29b 0.41a 0.66b 1.31b 1.82b T1 0.32a 0.42a 0.69a 1.39a 1.87a T2 0.29b 0.40b 0.64c 1.27c 1.78c T3 0.28b 0.40b 0.63c 1.28c 1.73d T4 0.26c 0.39b 0.63c 1.28c 1.68e T5 0.22d 0.37c 0.59d 1.18d 1.53f 注:同列数字后不同字母表示显著差异(P<0.05),下表同。
Note: The different letters after the numbers in the same column indicate significant difference (P<0.05),the same table below.表 2 赤霉素与2,4-表油菜素内酯对苦荞籽粒灌浆参数的影响
Table 2. Effect of GA3 and EBR treatments on buckwheat grain-filling parameters
粒位
Grain location处理
TreatmentA B K N R2 R0 Tmax.G/d Vmax/(g· hd−1) 强势粒 Superior grain CK 2.264 d 1.203 c 0.324 d 0.088 b 0.989 6 b 3.68 e 8.06 a 0.22 d T1 2.323 c 1.174 d 0.320 e 0.101 a 0.990 7 a 3.17 f 7.68 c 0.23 d T2 2.359 b 1.220 b 0.391 b 0.063 c 0.976 4 c 6.18 c 7.58 d 0.26 c T3 2.402 a 1.220 b 0.422 a 0.058 d 0.985 5 b 7.24 b 7.20 e 0.28 b T4 2.356 b 1.272 a 0.378 c 0.066 c 0.973 4 5.77 d 7.84 b 0.23 d T5 2.260 d 1.164 e 0.427 a 0.055 d 0.976 6 c 7.79 a 7.16 f 0.30 a 弱势粒 Inferior grain CK 16.749 a 0.626 d 0.079 d 0.138 e 0.988 7 a 0.57 d 19.21 a 0.20 a T1 6.852 d 0.580 f 0.125 a 0.165 b 0.979 0 c 0.76 a 10.05 f 0.12 d T2 14.798 b 0.605 e 0.085 c 0.142 d 0.987 0 a 0.60 c 17.07 c 0.18 b T3 7.528 c 0.721 a 0.080 c 0.158 c 0.983 3 b 0.51 e 18.93 b 0.14 c T4 5.469 e 0.671 c 0.103 b 0.165 b 0.982 4 b 0.63 b 13.60 d 0.12 d T5 4.046 f 0.693 b 0.110 b 0.169 a 0.983 5 b 0.65 b 12.83 e 0.10 e 注:A:终极生长量;B:初值参数;K:生长速率参数;N:形状参数; R2:判断系数;表示其配合度;R0:灌浆起始势;Tmax.G:达到最大灌浆速率的时间;Vmax:最大灌浆速率。
Note: A: Final weight of 1000 grain; B: The initial parameter; K: The growth-rate parameter; N: The shape parameter; R2: The compatibility; R0: The initial grouting potential; Tmax.G: The time to reach the maximum grouting rate; Vmax: The maximum grouting rate.表 3 赤霉素与2,4-表油菜素内酯对苦荞籽粒灌浆速率的影响
Table 3. Effect of GA3 and EBR treatments on buckwheat grain-filling rate
粒位
Grain location处理
Treatment前期
Early filling stage中期
Middle filling stage后期
Later filling stage平均速率
average rate/
(g·hd−1)贡献率
contribution/
%平均速率
average rate/
(g·hd−1)贡献率
contribution/
%平均速率
average rate/
(g·hd−1)贡献率
contribution/
%强势粒 Superior grain CK 0.10 bc 20.90 c 0.24 d 64.69 a 0.03 b 14.19 b T1 0.09 c 18.41 d 0.24 d 65.38 a 0.03 b 15.94 a T2 0.11 b 24.04 b 0.29 b 63.05 c 0.03 b 12.73 c T3 0.12 ab 24.54 b 0.32 a 62.70 c 0.04 a 12.57 c T4 0.13 a 28.27 a 0.27 c 61.12 d 0.03 b 10.49 d T5 0.09 bc 20.03 c 0.31 a 64.15 b 0.04 a 15.56 a 弱势粒 Inferior grain CK 0.04 b 1.58 d 0.11 a 17.12 d 0.21 a 57.17 a T1 0.08 a 2.03 c 0.06 d 14.05 f 0.12 c 50.54 f T2 0.04 b 1.52 d 0.09 b 14.81 e 0.19 b 54.77 e T3 0.04 b 3.25 a 0.09 b 31.91 a 0.09 d 55.15 d T4 0.04 b 2.83 b 0.07 c 25.52 c 0.09 d 56.72 c T5 0.04 b 3.23 a 0.06 d 28.81 b 0.07 e 55.79 b 表 4 赤霉素与2,4-表油菜素内酯对苦荞籽粒淀粉合成酶活性的影响
Table 4. Effect of GA3 and EBR treatments on starch synthase activity in buckwheat grains
指标
Item粒位
Grain location处理
Treatment7 d 14 d 21 d 28 d 35 d 腺苷二磷酸葡萄糖焦磷酸
ADP-glucose pyrophosphorylase /( U·g−1·min−1)强势粒 Superior grain CK 0.69 e 1.26 e 1.65 f 1.30 c 1.09 c T1 0.96 c 1.68 d 2.26 e 1.31 c 1.15 b T2 0.95 c 1.76 c 2.35 d 1.34 b 1.16 b T3 1.29 a 2.39 a 2.85 a 1.38 a 1.21 a T4 1.00 b 1.84 b 2.71 b 1.17 d 1.08 c T5 0.91 d 1.67 d 2.40 c 1.09 e 1.05 d 弱势粒 Inferior grain CK 0.38 a 0.71 a 1.42 a 1.76 a 1.30 a T1 0.38 a 0.67 b 1.18 b 1.70 b 1.24 b T2 0.37 ab 0.58 c 1.05 c 1.65 c 1.05 c T3 0.36 b 0.52 d 1.02 d 1.57 d 0.93 d T4 0.35 bc 0.44 e 0.95 e 1.50 e 0.92 d T5 0.34 c 0.41 f 0.81 f 1.42 f 0.88 f 可溶性淀粉合成酶
Soluble starch synthase/ (U·mg−1·min−1)强势粒 Superior grain CK 3.27 e 5.27 d 11.51 d 9.99 d 2.60 e T1 4.24 d 6.79 c 13.15 c 10.87 c 3.57 d T2 4.66 c 7.42 b 13.44 b 11.38 b 3.80 b T3 5.40 a 8.49 a 15.24 a 11.67 a 3.84 a T4 4.71 b 7.59 b 11.40 d 9.96 d 3.72 c T5 4.24 d 6.85 c 10.75 e 9.62 e 2.19 f 弱势粒 Inferior grain CK 3.13 a 7.12 a 9.18 a 7.29 a 3.31 a T1 2.90 b 6.82 b 8.87 b 6.89 b 3.02 b T2 2.64 c 6.34 c 8.18 c 6.81 b 2.89 c T3 2.43 d 5.86 d 7.92 d 6.62 c 2.85 c T4 2.38 d 5.83 d 7.77 e 6.49 d 2.79 d T5 1.92 e 3.61 e 7.24 f 6.26 e 2.63 e 表 5 赤霉素与2,4-表油菜素内酯对苦荞叶片抗氧化酶活性的影响
Table 5. Effect of GA3 and EBR treatments on antioxidant enzyme activities in buckwheat leaves
指标
Item处理
Treatment7 d 14 d 21 d 28 d 35 d 超氧化物歧化酶
Superoxide dismutase /(U·g−1·h−1)CK 280.62 e 409.97 b 379.15 c 284.85 f 256.76 e T1 320.10 c 436.36 a 394.64 a 307.04 e 283.54 d T2 321.10 c 406.97 c 386.25 b 309.62 d 284.07 d T3 323.39 b 402.14 d 367.53 d 368.39 a 313.66 a T4 344.36 a 383.56 e 349.46 e 354.76 b 309.36 b T5 292.37 d 313.16 f 324.93 f 334.28 c 307.66 c 过氧化物酶
Peroxidase/ (U·g−1·h−1)CK 1.32 c 1.61 b 1.81 d 4.82 e 1.91 f T1 1.35 c 1.66 a 3.10 a 8.33 b 2.62 d T2 1.34 c 1.59 b 2.85 b 8.39 b 2.56 e T3 1.53 b 1.55 c 1.88 c 8.79 a 2.79 c T4 2.08 a 1.45 d 1.84 c 7.51 c 4.64 a T5 1.53 b 1.40 e 1.77 e 6.71 d 2.96 b 表 6 赤霉素与2,4-表油菜素内酯对苦荞叶片光合特性的影响
Table 6. Effect of GA3 and EBR treatments on photosynthetic properties of buckwheat leaves
指标
Item处理
Treatment7 d 14 d 21 d 28 d 35 d 净光合速率
Pn/(μmol · m−2·s−1)CK 10.678 b 14.010 b 12.028 d 3.017 e 1.650 d T1 11.833 a 14.823 a 12.149 c 3.988 d 2.007 c T2 11.723 a 13.269 c 12.248 b 4.212 c 2.071 c T3 10.597 b 13.192 c 12.541 a 6.668 a 2.364 a T4 8.972 c 12.877 d 11.439 e 5.715 b 2.100 b T5 8.666 c 12.894 d 9.459 f 4.154 c 2.353 a 气孔导度
Gs/(mol·m−2·s−1)CK 0.266 d 0.415 e 0.108 e 0.082 f 0.053 e T1 0.273 d 0.451 c 0.398 b 0.180 c 0.081 d T2 0.333 c 0.514 b 0.429 a 0.111 e 0.107 c T3 0.406 a 0.518 a 0.434 a 0.265 a 0.147 a T4 0.392 a 0.515 b 0.329 c 0.196 b 0.114 b T5 0.374 b 0.430 d 0.310 d 0.155 d 0.098 c 蒸腾速率
Tr/(mmol·m−2·s−1)CK 1.216 f 3.686 f 3.084 d 0.739 e 0.501 c T1 1.242 e 3.985 e 3.247 c 1.602 b 0.578 b T2 1.363 d 4.506 c 3.361 c 1.088 d 0.606 b T3 1.838 a 5.257 a 4.884 a 2.156 a 0.870 a T4 1.537 b 5.026 b 4.255 b 1.360 c 0.425 d T5 1.425 c 4.368 d 4.230 b 1.156 d 0.403 e 表 7 赤霉素与2,4-表油菜素内酯对苦荞农艺性状及产量的影响
Table 7. Effect of GA3 and EBR treatments on agronomic traits and yield of buckwheat
处理
Treatment主茎节数
Number of
nodes of
main stem主茎分枝数
number of
main stem
branching主茎直径
Diameter of the
main stem/mm株高
stem
height/cm单株粒数
number of
Grains per
plant单株粒重
Number of
weight per
plant/g千粒重
1000-grain
weight/g产量
Yield/
(kg·hm−2)CK 14.3d 7.7c 4.58e 109.66e 350.4d 6.64d 17.94d 1515.78d T1 15.3c 8.3b 4.83d 125.91d 375.7c 6.87c 18.24c 1598.92c T2 17.7b 8.3b 4.90d 137.40c 385.1b 7.30b 19.12b 1698.78b T3 18.0ab 9.0a 5.93a 147.64b 406.5a 7.41a 19.63a 1792.51a T4 18.0ab 7.3d 5.54b 154.63a 342.2e 6.18e 17.19e 1458.10e T5 18.7a 7.0e 5.16c 157.71a 283.3f 5.45f 16.25f 1387.73f -
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