Response of nitrogen utilization to root interaction and plant density in barley-pea intercropping system
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摘要: 针对禾豆间作密植机理研究薄弱问题，以大麦间作豌豆为研究对象，设施氮[不施氮：0 mg（N）·kg-1（土）；施氮：100 mg（N）·kg-1（土）]、隔根（不隔根、隔根）和密度[低密度：15株（大麦）·盆-1；高密度：25株（大麦）·盆-1]3个参试因子，通过盆栽试验探讨了施氮和根系分隔对密植间作群体氮素竞争互补关系和利用效率的影响，以期为禾豆间作密植和氮素高效利用提供调控依据。结果表明：1）施氮、根间互作和增加大麦密度均可提高大麦||豌豆间作群体的吸氮量，其中施氮较不施氮处理提高33.8%，不隔根处理较隔根处理提高81.1%，高密度较低密度处理提高4.2%；根间互作在低氮条件下对间作吸氮量的贡献相对较高，不施氮和施氮条件下，根间互作提高间作吸氮量的比例分别为92.4%和11.0%；根间互作条件下增大大麦种植密度可显著提高间作群体吸氮量。2）大麦为氮素竞争优势种，密植使大麦氮素竞争比率显著提高，施氮能弱化大麦氮素竞争比率，抽穗期大麦相对于豌豆的氮素竞争优势达到最大值。3）根间互作使大麦、豌豆籽粒氮含量在施氮条件下分别提高126.7%、26.9%，不施氮时分别提高188.5%、46.5%，且施氮水平和根间作用方式对间作籽粒氮含量有显著的交互作用。4）高密度大麦和根间互作可显著提高间作群体的氮肥利用率，根间互作条件下增加大麦密度使间作群体氮肥利用率提高59.8%；大麦相对于豌豆的氮素竞争比率与间作群体氮肥利用率呈显著正相关关系。本研究表明，施氮、根间作用与大麦密度对大麦||豌豆间作氮素利用呈显著的交互作用，适宜的施氮量和充分的根间作用是支撑间作密植、优化种间对氮素的竞争关系，最终提高群体吸氮量和氮肥利用率的重要途径。Abstract: To investigate the mechanism of high planting density in cereal-legume intercropping system, a pot experiment of barley-pea intercropping system was carried out and the effects of nitrogen (N) application and root barrier on nitrogen and fertilizer use efficiency under high planting density of barley were determined. In the experiment, three factors, each with two treatment levels-N fertilizer application[no N application and N application with 100 mg(N)·kg-1], root barrier[no barrier with root interaction and root barrier without root interaction) and planting density (low density with 15 barley plants per pot and high density with 25 barley plants per pot) were set up. The results showed that:1) N application, plant root interaction and high barley planting density improved N uptake of barley-pea intercropping system. Compared with no N application treatment, N uptake increased by 33.8% in N application treatment. There was also 81.1% increase in N uptake under no root barrier treatment over root barrier treatment. N uptake under high planting density treatment increased by 4.2% compared with low planting density treatment. Plant root interaction improved N uptake by 92.4% and 11.0%, respectively, under no N application and N application treatments. Increasing planting density with root interaction significantly increased N uptake of the intercropping system. 2) Barely plant performed better for N competition, and its' N competition ratio significantly increased under high planting density. However, N application reduced barley N competition ratio. Compared with pea, barley was highest in competitive advantage at heading stage. 3) Root interaction improved grain N content of barley and pea, respectively, by 126.7% and 26.9% under N application treatment. Also barley and pea kernel N content increased, respectively, by 188.5% and 46.5% under no N treatment. There was a significant interaction between N application and root interaction for kernel N content. 4) High barley planting density significantly improved N use efficiency by 59.8% under root interaction treatment of the intercropping system. N competition between barley and pea was positively correlated with N use efficiency in the intercropping plant population. In conclusion, interactions of N application, root barrier and barley planting density enhanced crop productivity of barley-pea intercropping system. Proper N application and sufficient root interaction made feasible high planting density intercropping system, optimized competition between barley and pea, and improved N uptake and use efficiency.
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图 1 大麦||豌豆间作系统不同处理下大麦相对于豌豆的氮素竞争比率
N0和N1分别代表不施氮和施氮; D1和D2代表大麦低和高种植密度。N0: no nitrogen applied; N1: 100 mg(N)×kg-1applied; D1: low barley planting density; D2: high barley planting density.
Figure 1. Dynamics on N competitive ratios of barley to pea in the barley-pea intercropping system under different treatments
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图 2 大麦||豌豆间作氮肥利用率对不同密度和施氮水平的响应
N1代表施氮; P代表根系分隔; D1和D2代表大麦低和高种植密度。N1: 100 mg(N)×kg-1 applied; P: root barrier; D1: low barley planting density; D2: high barley planting density.
Figure 2. Response of N use efficiency of barly-pea intercropping to different treatments of barley planting densitiesy of barly and nitrogen application treatments
表 1 不同施氮、根间作用和密度处理下大麦||豌豆间作系统不同大麦生育时期植株吸氮量
Table 1 N uptake of the baeley-pea intercropping system at different barley growth stages under different treatments of N application, root barrier and barley planting density mg(N)·pot-1
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表 2 大麦||豌豆间作系统不同处理下大麦、豌豆籽粒氮素含量及氮素收获指数
Table 2 Grain N contents and N harvest indexes of barley and pea in the barley-pea intercropping system under different treatments
年份 Year 处理 Treatment 籽粒氮含量 Grain N content (mg×pot-1) 氮素收获指数 N harvest index (NHI) 大麦 Barley 豌豆 Pea 间作 Intercropping 大麦 Barley 豌豆 Pea 间作 Intercropping 2012 N0D1 300.2±19.7 b 407.5±6.0ab 707.7±14.1 b 0.775±0.04a 0.619±0.01 b 0.676±0.05a N0D2 318.9±15.7b 352.0±9.4bc 670.9±16.8 b 0.781±0.08a 0.555±0.02 b 0.642±0.09a N0PD1 117.1±17.9d 247.2±6.1d 364.3±23.3c 0.620±0.06ab 0.618±0.02b 0.620±0.04a N0PD2 98.0±7.5d 277.7±46.4cd 375.6±45.3c 0.542±0.07b 0.702±0.06 ab 0.653±0.04a N1D1 450.8±86.2a 475.1±87.5a 926.0±59.2 a 0.800±0.04a 0.690±0.14ab 0.738±0.13a N1D2 513.9±53.9 a 473.6±97.3a 987.5±43.6a 0.752±0.06ab 0.625±0.09b 0.685±0.08a N1PD1 219.4±8.0 c 385.1±38.4ab 604.4±45.2b 0.721±0.02ab 0.817±0.03a 0.775±0.02a N1PD2 227.5±28.3c 372.8±53.3abc 600.4±73.0b 0.722±0.05ab 0.784±0.14a 0.759±0.16a 2013 N0D1 260.5±23.2bc 411.0±39.4b 672.1±26.3b 0.734±0.04bc 0.595±0.05de 0.643±0.04e N0D2 294.2±3.1b 357.5±21.3c 651.7±18.5b 0.785±0.08abc 0.531±0.00e 0.621±0.08e N0PD1 113.6±10.2e 251.6±10.1d 365.2±7.3d 0.697±0.06c 0.743±0.06ab 0.728±0.11bcd N0PD2 85.9±5.1e 273.7±23.8d 359.5±18.7d 0.504±0.07d 0.803±0.07a 0.701±0.00d N1D1 511.3±46.2a 506.6±27.0a 1017.9±23.9a 0.888±0.04a 0.699±0.02bc 0.783±0.04ab N1D2 527.5±13.3a 466.1±24.8a 993.7±14.2a 0.806±0.06ab 0.627±0.05cd 0.711±0.04cd N1PD1 212.2±35.9d 369.3±45.8bc 581.5±11.3c 0.688±0.02c 0.807±0.03a 0.759±0.01abc N1PD2 224.9±11.4cd 388.1±23.3bc 612.9±29.6c 0.734±0.05bc 0.823±0.04a 0.788±0.09a 显著性检验 Significance due to: 年际 Year NS NS NS NS NS NS 施氮 N application (N) *** *** *** ** *** *** 密度 Planting density (D) NS NS NS NS NS NS 根系分隔 Root barrier (B) *** *** *** *** *** * 施氮×密度 N × D NS NS NS NS NS NS 施氮×根系分隔 N × B *** NS * NS NS NS 密度×根系分隔 D × B * * NS NS * NS 施氮×密度×根系分隔 N × D × B NS NS NS * NS NS N0和N1分别代表不施氮和施氮; P代表根系分隔; D1和D2代表大麦低种植密度和高种植密度。显著性检验是分作物和年份进行; 各处理每年数据3次重复的平均值进行比较。同列不同字母表示差异显著。NS表示不显著; ***表示P<0.001显著; **表示P<0.01显著; *表示P<0.05显著。N0: no N applied; N1: 100 mg(N)×kg-1 applied; P: root barrier; D1: low barley planting density; D2: high barley planting density. The test of significance was performed on the same plant and year; Values are means of three triplicates of one treatments in every year. Different letters within the same column indicate significant differences at P < 0.05 level. NS: not significant; ***: significant at P < 0.001 level; **: significant at P < 0.01 level; *: significant atP < 0.05 level.
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表 3 大麦||豌豆间作系统大麦不同生育期氮素竞争比率与间作籽粒氮含量的关联度排序
Table 3 Degrees of association between seasonal N competitive ratio and grain nitrogen content of the barly-pea intercropping system at different barley growth stages under different treatments
处理 Treatment 间作作物籽粒氮含量 Grain nitrogen content of barly-pea intercropping (mg×pot-1) 氮素竞争比率 N competitive ratio 苗期 Seeding stage 分蘖期 Tillering stage 抽穗期 Heading stage 成熟期 Maturity stage N0D1 689.90 -0.22 0.05 0.58 0.27 N0D2 661.29 -0.03 0.22 0.75 0.44 N1D1 971.93 0.00 0.21 0.62 0.32 N1D2 990.58 0.20 0.49 0.91 0.58 关联度 Degree of association 0.6112 0.9611 0.9874 0.9781 排序 Ranking 4 3 1 2 N0和N1分别代表不施氮和施氮; D1和D2代表大麦低和高种植密度。N0: no nitrogen applied; N1: 100 mg(N)×kg-1 applied; D1: low barley planting density; D2: high barley planting density.
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表 4 大麦豌豆共生期不同大麦生育时期氮素竞争比率与籽粒氮含量的相关系数
Table 4 Correlation coefficients between seasonal N competitive ratio and grain nitrogen content in barly-pea intercropping systems under different barley growth stageperieds
参数 Parameter 生育时期 Growth stage 籽粒氮含量 Grain nitrogen content 苗期 Seeding 分蘖期 Tillering 抽穗期 Heading 成熟期 Maturity 大麦 Barley 豌豆 Pea 间作 Intercropping 苗 期 Seeding 1 0.649** 0.531** 0.502 0.371 -0.059 0.224 分蘖期 Tillering 1 0.360 0.525** 0.499 -0.062 0.310 抽穗期 Heading 1 0.635 0.091 -0.052 成熟期 Maturity 1 0.250 -0.189 0.090
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