Effects of potassium and silicon fertilization on lignin metabolism and lodging resistance of oil flax stem
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摘要: 通过田间试验, 探讨了钾硅养分耦合对胡麻茎秆木质素代谢及抗倒伏特性的调控效应, 以期为肥料运筹抗倒伏提供依据。选用三因素裂区试验设计, 以两个胡麻品种‘陇亚11号’(V1)、‘定亚23号’(V2)为主区因素; 钾肥为副区因素, 设0 kg(K2O)∙hm−2 (K0)、52.5 kg(K2O)∙hm−2 (K1)、105 kg(K2O)∙hm−2 (K2) 3个水平; 硅肥为副副区因素, 设0 kg(SiO2)∙hm−2 (Si0)、90 kg(SiO2)∙hm−2 (Si1)两个水平。结果表明, V1和V2的茎秆木质素含量及代谢酶活性具有显著差异。施钾显著提高了木质素含量及苯丙氨酸解氨酶(PAL)、4-香豆酸辅酶A连接酶(4CL)和肉桂醇脱氢酶(CAD)的活性, 且始终保持K1>K2>K0的趋势; 施硅对上述试验指标无显著的主效应, 但钾硅肥互作效应显著提升了现蕾前的木质素含量及整个生育期的PAL、4CL和CAD的活性, 且K1Si1处理的提升效果最佳。木质素含量与整个生育期的CAD活性极显著或显著正相关, CAD活性的提高是胡麻茎秆木质素含量增加的重要酶学基础。V1的抗折力和抗倒伏指数极显著高于V2; 与K0相比, K1显著提高了茎秆抗折力和抗倒伏指数, K2水平下变化趋势相反, 且始终保持K1>K0>K2的趋势; 施硅可提高茎秆的抗折力和抗倒伏指数, 且低钾配施硅肥可显著提升茎秆抗折力和抗倒伏指数。木质素含量与抗折力、抗倒伏指数显著正相关, 与实际倒伏率负相关。低钾处理及低钾配施硅肥均显著提高了籽粒产量。综上, ‘陇亚11号’胡麻的抗倒伏性能显著优于‘定亚23号’; 52.5 kg(K2O)∙hm−2+90 kg(SiO2)∙hm−2对木质素含量及其代谢酶活性、抗倒伏能力和籽粒产量的提升效果最佳, 且单施钾肥对木质素代谢及抗倒伏特性的提升效果优于单施硅肥。Abstract: The potassium and silicon nutritional status of crops is closely related to lodging resistance, which is often evaluated by using lignin content. Investigating the regulatory effects of potassium and silicon nutrient coupling on lignin metabolism and lodging resistance of oil flax stems can provide a foundation for lodging resistance through fertilizer management. A three-factor split plot design was used, with variety, potassium dosage, and silicon dosage as the main treatment, split treatment, and sub-split treatment, respectively. In the study, the two varieties were ‘Longya No. 11’ (V1) and ‘Dingya No. 23’ (V2); three potassium dosages were 0 (K0), 52.5 (K1), and 105 kg(K2O)·hm−2 (K2); and two silicon dosages were 0 (Si0) and 90 kg(SiO2)·hm−2 (Si1). The results showed that there were significant differences in the stem lignin content and related metabolic enzyme activities between V1 and V2. Potassium application significantly increased lignin content, as well as the activities of phenylalanine deaminase (PAL), 4-coumaric acid: CoA ligase (4CL), and cinnamyl alcohol dehydrogenase (CAD); the order of the influence effect was K1>K2>K0. Silicon application did not have a significant effect on these test indicators, but the interaction of potassium and silicon significantly increased the lignin content before the budding stage and improved the activities of PAL, 4CL, and CAD throughout the growth stage, with the best enhancing effect derived from K1Si1 treatment. The lignin content was significantly and positively correlated with CAD activity during the entire growth period, and the increase in CAD activity was an important enzymatic basis for the increase in lignin content. The breaking resistance and lodging resistance index of V1 were higher than those of V2. Compared with K0, K1 significantly increased breaking resistance and lodging resistance index, while K2 had the opposite effect, and the influence effect was K1>K0>K2. Silicon application improved stem breaking resistance and lodging resistance index, and lower potassium accompanied by silicon significantly improved the above-mentioned two indicators. Lignin content was significantly and positively correlated with breaking resistance and lodging resistance index, and negatively correlated with the actual lodging rate. Both lower potassium treatment and lower potassium accompanied by silicon fertilizer significantly increased seed yield. In conclusion, ‘Longya No. 11’ had better lodging resistance than ‘Dingya No. 23’; 52.5 kg(K2O)·hm−2 accompanied by 90 kg(SiO2)·hm−2 most improved the lignin content, as well as the metabolic enzyme activity, lodging resistance properties, and seed yield; applying potassium fertilizer alone had a better promotion effect on lignin metabolism and lodging resistance than fertilizing silicon individually.
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Key words:
- Potassium fertilizer /
- Silicon fertilizer /
- Oil flax /
- Lignin /
- Lignin metabolizing enzymes /
- Lodging resistance /
- Yield
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图 1 钾硅肥配施下不同胡麻品种各生育时期的茎秆木质素含量
V1和V2分别为胡麻品种‘陇亚11号’和‘定亚23号’。K0、K1和K2分别表示施钾量为0 kg(K2O)∙hm−2、52.5 kg(K2O)∙hm−2和105 kg(K2O)∙hm−2; Si0和Si1分别表示施硅量为0 kg(SiO2)∙hm−2和90 kg(SiO2)∙hm−2。同一时期不同小写字母表示处理间在P<0.05水平差异显著。V1 and V2 are flax varieties ‘Longya 11’ and ‘Dingya 23’, respectively. K0, K1 and K2 represent potassium application rates of 0, 52.5 and 105 kg(K2O)∙hm−2, respectively; Si0 and Si1 represent silicon application rates of 0 and 90 kg(SiO2)∙hm−2, respectively. Different lowercase letters in the same growth stage indicate significant differences among treatments at P<0.05 level.
Figure 1. Stem lignin contents of two flax varieties at different growth stages under combined application of potassium and silicon fertilizers
图 2 钾硅肥配施对胡麻不同生育时期茎秆木质素代谢酶活性的影响
V1和V2分别为胡麻品种‘陇亚11号’和‘定亚23号’。K0、K1和K2分别表示施钾量为0 kg(K2O)∙hm−2、52.5 kg(K2O)∙hm−2和105 kg(K2O)∙hm−2; Si0和Si1分别表示施硅量为0 kg(SiO2)∙hm−2和90 kg(SiO2)∙hm−2。同一时期不同小写字母表示处理间在P<0.05水平差异显著。V1 and V2 are flax varieties ‘Longya 11’ and ‘Dingya 23’, respectively. K0, K1 and K2 represent potassium application rates of 0, 52.5 and 105 kg(K2O)∙hm−2, respectively; Si0 and Si1 represent silicon application rates of 0 and 90 kg(SiO2)∙hm−2, respectively. Different lowercase letters in the same growth stage indicate significant differences among treatments at P<0.05 level.
Figure 2. Effects of combined application of potassium and silicon fertilizers on lignin metabolism enzymes activities in stems at different growth stages of different flax varities
图 3 钾硅肥配施下胡麻不同生育时期的茎秆抗折力
V1和V2分别为胡麻品种‘陇亚11号’和‘定亚23号’。K0、K1和K2分别表示施钾量为0 kg(K2O)∙hm−2、52.5 kg(K2O)∙hm−2和105 kg(K2O)∙hm−2; Si0和Si1分别表示施硅量为0 kg(SiO2)∙hm−2和90 kg(SiO2)∙hm−2。同一时期不同小写字母表示处理间在P<0.05水平差异显著。V1 and V2 are flax varieties ‘Longya 11’ and ‘Dingya 23’, respectively. K0, K1 and K2 represent potassium application rates of 0, 52.5 and 105 kg(K2O)∙hm−2, respectively; Si0 and Si1 represent silicon application rates of 0 and 90 kg(SiO2)∙hm−2, respectively. Different lowercase letters in the same growth stage indicate significant differences among treatments at P<0.05 level.
Figure 3. Stems breaking resistance under combined application of potassium and silicon fertilizers at different growth stages
图 4 钾硅肥配施下胡麻不同生育时期茎秆的抗倒伏指数
V1和V2分别为胡麻品种‘陇亚11号’和‘定亚23号’。K0、K1和K2分别表示施钾量为0 kg(K2O)∙hm−2、52.5 kg(K2O)∙hm−2和105 kg(K2O)∙hm−2; Si0和Si1分别表示施硅量为0 kg(SiO2)∙hm−2和90 kg(SiO2)∙hm−2。同一时期不同小写字母表示处理间在P<0.05水平差异显著。V1 and V2 are flax varieties ‘Longya 11’ and ‘Dingya 23’, respectively. K0, K1 and K2 represent potassium application rates of 0, 52.5 and 105 kg(K2O)∙hm−2, respectively; Si0 and Si1 represent silicon application rates of 0 and 90 kg(SiO2)∙hm−2, respectively. Different lowercase letters in the same growth stage indicate significant differences among treatments at P<0.05 level.
Figure 4. Lodging resistance index of flax stem under combined application of potassium and silicon fertilizers at different growth stages
图 5 钾硅肥配施对不同胡麻品种产量的影响
V1和V2分别为胡麻品种‘陇亚11号’和‘定亚23号’。K0、K1和K2分别表示施钾量为0 kg(K2O)∙hm−2、52.5 kg(K2O)∙hm−2和105 kg(K2O)∙hm−2; Si0和Si1分别表示施硅量为0 kg(SiO2)∙hm−2和90 kg(SiO2)∙hm−2。不同小写字母表示处理间在P<0.05水平差异显著。V1 and V2 are flax varieties ‘Longya 11’ and ‘Dingya 23’, respectively. K0, K1 and K2 represent potassium application rates of 0, 52.5 and 105 kg(K2O)∙hm−2, respectively; Si0 and Si1 represent silicon application rates of 0 and 90 kg(SiO2)∙hm−2, respectively. Different lowercase letters indicate significant differences among treatments at P<0.05 level.
Figure 5. Effect of combined application of potassium and silicon fertilizers on yield of different flax varieties
表 1 钾硅肥配施对胡麻不同生育时期茎秆木质素含量影响的方差分析(P值)
Table 1. P values in variance analysis of the effects of potassium and silicon fertilizers on lignin content of stems at different growth stages of different flax varieties
处理 Treatment 分茎期 Branching 现蕾期 Budding 盛花期 Anthesis 青果期 Kernel 成熟期 Maturity 品种 Variety (V) 0.115 0.086 0.032* 0.083 0.386 钾肥 Potassium fertilizer (K) 0.027* 0.049* 0.010** 0.000** 0.035* 硅肥 Silicon fertilizer (Si) 0.913 0.345 0.481 0.109 0.103 V×K 0.531 0.355 0.940 0.673 0.795 V×Si 0.856 0.408 0.700 0.974 0.755 K×Si 0.010** 0.011* 0.282 0.114 0.443 V×K×Si 0.032* 0.153 0.987 0.849 0.676 *: P<0.05; **: P<0.01. 表 2 钾硅肥配施对胡麻不同生育时期茎秆木质素合成相关酶活性影响的方差分析(P值)
Table 2. P values in variance analysis of the effects of potassium and silicon fertilizers on lignin metabolism-related enzymes activities of stems at different growth stages of different flax varieties
酶 Enzyme 处理 Treatment 分茎期 Branching 现蕾期 Budding 盛花期 Anthesis 青果期 Kernel 苯丙氨酸解氨酶
Phenylalnine ammonialyase (PAL)品种 Variety (V) 0.600 0.331 0.166 0.003** 钾肥 Potassium fertilizer (K) 0.000** 0.000** 0.000** 0.000** 硅肥 Silicon fertilizer (Si) 0.552 0.219 0.417 0.039* V×K 0.002** 0.282 0.000** 0.042* V×Si 0.584 0.210 0.523 0.003** K×Si 0.000** 0.000** 0.000** 0.000** V×K×Si 0.067 0.542 0.616 0.012* 4-香豆酸: CoA连接酶
4-coumaric acid: COA ligase (4CL)品种 Variety (V) 0.001** 0.020* 0.001** 0.011* 钾肥 Potassium fertilizer (K) 0.000** 0.000** 0.011* 0.004** 硅肥 Silicon fertilizer (Si) 0.010** 0.403 0.164 0.213 V×K 0.107 0.072 0.080 0.104 V×Si 0.788 0.409 0.104 0.662 K×Si 0.000** 0.002** 0.002** 0.010** V×K×Si 0.104 0.755 0.033* 0.646 肉桂醇脱氢酶
Cinnamyl alcohol dehydrogenase (CAD)品种 Variety (V) 0.800 0.036* 0.011* 0.171 钾肥 Potassium fertilizer (K) 0.000** 0.000** 0.000** 0.000** 硅肥 Silicon fertilizer (Si) 0.771 0.890 0.001** 0.118 V×K 0.000** 0.059 0.093 0.000** V×Si 0.113 0.054 0.610 0.000** K×Si 0.000** 0.000** 0.011* 0.000** V×K×Si 0.059 0.048* 0.237 0.001** *: P<0.05; **: P<0.01. 表 3 胡麻不同生育时期茎秆木质素含量和代谢相关酶活性的相关系数
Table 3. Correlation coefficients between lignin content and its metabolism-related enzymes activities in flax stem at different growth stages
酶
Enzyme木质素 Lignin 分茎期
Branching现蕾期
Budding盛花期
Anthesis青果期
Kernel苯丙氨酸解氨酶 Phenylalnine ammonialyase (PAL) 0.427** 0.535** 0.347* −0.228 4-香豆酸: CoA连接酶 4-coumaric acid:COA ligase (4CL) 0.257 0.421* −0.269 −0.015 肉桂醇脱氢酶 Cinnamyl alcohol dehydrogenase (CAD) 0.393* 0.451** 0.502** 0.402* *: P<0.05; **: P<0.01. 表 4 钾硅肥配施对胡麻不同生育时期茎秆抗折力影响的方差分析(P值)
Table 4. P value in variance analysis of the effects of potassium and silicon fertilizers on stem breaking resistance of different flax varieties
处理 Treatment 现蕾期 Budding 盛花期 Anthesis 青果期 Kernel 成熟期 Maturity 品种 Variety (V) 0.001** 0.001** 0.001** 0.005** 钾肥 Potassium fertilizer (K) 0.002** 0.000** 0.000** 0.019* 硅肥 Silicon fertilizer (Si) 0.066 0.000** 0.000** 0.067 V×K 0.022* 0.000** 0.010** 0.045* V×Si 0.075 0.660 0.001** 0.016* K×Si 0.000** 0.000** 0.000** 0.009** V×K×Si 0.001** 0.002** 0.037* 0.024* *: P<0.05; **: P<0.01. 表 5 钾硅肥处理对胡麻不同生育时期抗倒伏指数影响的方差分析(P值)
Table 5. P value in variance analysis of the effects of potassium and silicon fertilizers on lodging resistance index at different growth stages of different flax varieties
处理 Treatment 现蕾期 Budding 盛花期 Anthesis 青果期 Kernel 成熟期 Maturity 品种 Variety (V) 0.000** 0.000** 0.000** 0.000** 钾肥 Potassium fertilizer (K) 0.072 0.045* 0.000** 0.406 硅肥 Silicon fertilizer (Si) 0.104 0.002** 0.129 0.756 V×K 0.266 0.850 0.044* 0.424 V×Si 0.207 0.202 0.028* 0.146 K×Si 0.685 0.016* 0.046* 0.569 V×K×Si 0.148 0.016* 0.061 0.693 *: P<0.05; **: P<0.01. 表 6 胡麻不同生育时期茎秆木质素含量与抗倒伏能力相关分析
Table 6. Correlation analysis of lignin content and lodging resistance indexes in flax stem at different growth stages
抗倒伏指标
Lodging resistance properties木质素 Lignin 现蕾期 Budding 盛花期 Anthesis 青果期 Kernel 成熟期 Maturity 抗折力 Stem breaking resistance 0.495** 0.726** 0.490** 0.222 抗倒伏指数 Lodging resistance index 0.322 0.720** 0.555** 0.091 倒伏率 Lodging rate — — −0.556** −0.314 **: P<0.01. -
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