双基因型玉米间作控制气传病害的生态生理基础
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摘要
双基因型玉米间作能够有效控制玉米的气传病害,但对其机制缺乏系统研究。本文以郑单958(株型紧凑,感南方锈病)、鲁单981(株型半紧凑,高抗南方锈病)、周单041(株型半紧凑,抗南方锈病)、豫玉19(株型半紧凑,感南方锈病)、浚单20(株型紧凑,中感小斑病)和滑丰9(株型半紧凑,中抗小斑病)为材料,采用大田试验和室内试验相结合的方法,研究了双基因型玉米间作的群体质量、对气传病害(玉米小斑病和南方锈病)的防控及抗病代谢反应,提取并分析了不同基因型玉米叶片挥发性物质的组分,同时探讨了玉米叶片挥发性物质对南方锈病和小斑病病原物的抑制作用。目的是从作物栽培学、植物病理学和化学生态学相结合的角度,揭示双基因型玉米间作控制气传病害的生态生理基础,为开发可持续控制玉米气传病害的保健栽培技术积累理论依据。主要研究结果如下：
(1)双基因型玉米间作有利于改善群体结构、减少病原物侵染机率、降低病情指数。在ZD958‖LD98和HF9‖XD20两种间作模式中,4个品种间作植株的平均叶龄均有不同程度的增加；吐丝前,群体叶面积指数(leaf area index,LAI)单间作无明显差异,吐丝后,HF9和LD981的LAI分别大于和显著大于单作群体,而ZD958和XD20则分别小于和显著小于单作群体；吐丝后10d,4个品种棒三叶叶色值(soil and plant analyzer development readings,SPADRs)均有所增加,并且除ZD958外,其余3个品种棒三叶净光合速率均有所增加,其中LD981增加显著。间作对吐丝以前的群体干物质积累量影响不大,吐丝后,半紧凑型品种(HF9和LD981)的干物质积累量增加,其中LD981增加显著,而紧凑型品种(XD20和ZD958)的干物质积累量减少,其中ZD958显著减少。在ZD958‖LD98和YY19‖ZD041两种间作模式中,在吐丝后10d,4个玉米品种叶片中的SOD和POD活性均有所提高或显著提高,ZD041和ZD958叶片中的CAT活性提高或显著提高,YY19和LD981变化不显著；4个玉米品种叶片中的MDA含量降低。在吐丝后40d,SOD和POD活性及MDA含量的变化与吐丝后10d基本一致,CAT活性均有所提高,其中以YY19较为显著。紧凑型和半紧凑型品种间作增加了群体透光率。间作降低了间作群体引起继代感染的病原菌孢子数量,ZD958和XD20病情指数分别下降59.75%和50.38%,HF9和XD20小斑病病情也有所下降。间作还增加了复合群体产量,3种间作模式的土地当量比(land equivalent ratio,LER)均大于1,籽粒品质也有所改善。
(2)双基因型玉米间作增强了群体的抗病代谢。人工接种试验表明,抗南方锈病基因型LD981基因型与感南方锈病基因型ZD958间作,病原菌侵染后,能够及时启动间作群体的抗病代谢,LOX、PAL和PPO活性迅速提高,并维持较长时间的高活性水平；但抗、感病品种的表现有所差异,在感病品种ZD958抗南方锈病的代谢中,LOX、PAL和PPO均具有重要作用,而在抗病品种LD981抗南方锈病的代谢中,PPO所起的作用更大一些。
(3)双基因型玉米间作能够增加群体遗传多样性、丰富群体叶片挥发性物组分、增强对南方锈病和小斑病的抵抗能力。采用蒸馏法提取了玉米叶片挥发性物质,并对其组分进行了GC-MS分析。结果表明,玉米叶片挥发性组分及组分比在品种间和提取方法间存在较大差异,间作也能改变其组分及组分比。染毒试验结果表明,叶片挥发性物质能够抑制南方锈病和小斑病病原菌孢子萌发,并能抑制小斑病菌丝的生长和分生孢子的形成,对南方锈病病原菌孢子萌发的抑制作用尤为显著。5种挥发组分对南方锈病病原菌夏孢子萌发的抑制率从大到小依次为顺-3-己烯醇>反-2-己烯醛>苯甲醇>香草醛>α-蒎烯,对小斑病病原菌孢子萌发的抑制率从大到小依次为苯甲醇>香草醛>顺-3-己烯醇>反-2-己烯醛>α-蒎烯。香草醛、反-2-己烯醛和顺-3-己烯醇对小斑病病原菌菌丝生长的抑制作用最强,但低浓度和高浓度的顺-3-己烯醇处理对孢子的形成与萌发具有促进作用；反-2-己烯醛显著抑制孢子的形成,但高浓度处理对菌丝所产孢子的萌发无影响；香草醛显著抑制孢子的形成,但所产孢子的萌发率相对较高；苯甲醇对菌丝生长的影响较小,但菌丝颜色变异较大,气生菌丝发达呈灰白色至白色,难以产生分生孢子；α-蒎烯对菌丝生长的抑制作用最弱,菌丝性状基本正常,但产孢量较少,孢子的萌发率也较低。挥发组分组合作用的效果要优于单剂处理,这符合植物抗病的自然状况。
总之,双基因型玉米间作抗病具有深厚的生态生理基础,是群体生态(包括化学生态)和群体抗病生理共同作用的结果。双基因型玉米间作,不仅优化群体结构、改善群体通风透光条件、营造了一个有利于植株生长而不利于病原物滋生的微生态环境,而且增加了群体抗病基因的遗传多样性,群体次生代谢变得丰富多彩,群体中挥发性物质及其所建立起来的化学通讯网络更加复杂,提高了群体抵抗病原菌侵染的能力,从而延缓了叶片衰老,减少了病原物侵染的机率及玉米植株为抵御病害所付出的能量损失,提高了群体的光合速率和物质生产能力,促进了籽粒产量的提高和品质的改善。
Maize airborne diseases can be controlled efficiently via double-genotypes intercropping, but the mechanisms are little known. We presuppose that the eco-physiological factors such as compound population quality, disease resistant metabolism and maize leaf volatile organic compounds (LVOCs) may be of importance for controlling maize airborne diseases in this intercropped system. In this study, we conducted a field experiment on the farm of Henan Agricultural University, Zhengzhou, China, using six maize hybrids, Zhengdan 958(ZD958, compact plant type, susceptible to maize leaf south rust), Ludan 981(LD981, semi-compact plant type, highly resistant to maize leaf south rust), Zhoudan 041(ZD041, semi-compact plant type, resistant to maize leaf south rust disease), Yuyu 19(YY19, semi-compact plant type, susceptible to maize leaf south rust), Xundan 20(XD20, compact plant type, medium susceptible to maize leaf spot), Huafeng 9(HF9, semi-compact plant type, medium resistant to maize leaf spot). The experimental plots of 6 m×6 m were separated by 1 m. The experiments were planted by hand with rows oriented north-south and spacing of 0.70 and 0.50 m. The ratio of two maize varieties intercropped was 1:1 per row. The planting density was 67 500 plant·hm-2. There were nine treatments: each of the varieties alone, YY19 intercropped with ZD041 (YY19‖ZD041), ZD958 intercropped with LD981 (ZD958‖LD981), and HF9 intercropped with XD20 (HF9‖XD20). There were four replicate plots for each treatment, arranged in a randomized complete block design. We investigated the population quality, resistant physiology and relationship between airborne diseases resistance and maize leaf volatile organic compounds. Our objective was to reveal the eco-physiological basic for controlling maize airborne diseases in intercropping with double-genotypes. The results are as follows:
1) Maize intercropping with double-genotypes is favorable for improving plant population structure, enhancing population quality and reducing disease index. In the intercropping treatments both ZD958 II LD981 and HF9 II XD20, the average leaf life of four varieties increased. The leaf area indexes (LAI) of four varieties were no significant difference between sole-cropping and Intercropping before silking. But, the LAI of HF9 and LD981 increased while that of ZD958 and XD20 decreased in the intercropping treatments after silking. The soil and plant analyzer development readings (SPADRs) of ear-leaves of four varieties increased in the treatments 10 days after silking, and the photosynthesis rate of ear-leaves in LD981, HF9 and XD20 increased or increased significantly. There were no differences in the dry matter accumulation (DMA) of four varieties between the treatments of sole-cropping and intercropping before silking. The DMA of semi-compact plant types both HF9 and LD981 increased or increased markedly, while that of compact plant types both XD20 and ZD958 decreased or decreased significantly after silking. In the intercropping treatments both ZD958‖LD98 and YY19‖ZD041, the activities of SOD and POD of leaves increased in the intercropping treatments 10 days after silking, which resulted in decreased MDA contents of leaves. The activities of CAT of leaves in ZD041 and ZD958 increased or increased markedly, while YY19 and LD981 decreased slightly. The changes of activities of SOD and POD and contents of MDA in leaves remained similar 40 days after silking. Meanwhile, the activities of CAT of leaves increased, among them YY19 increased significantly. Transmittance enhanced in the treatment of compact plant type intercropped with semi-compact plant type, which resulted in reduced pathogen spore concentrations in the intercropped colonies, which led to reduced disease index of the intercropped colonies. For example, the disease index of southern rust in the intercropped colonies of ZD958 and HF9 reduced by 59.75% and 50.38%, respectively. Also, yields, land equivalent ratio (LER) and grain quality were enhanced in the intercropped colonies. Results suggested that suitable maize intercropping with different genotypes would increase the activities of SOD, POD and CAT of leaf, prolong leaf life, enhance photosynthesis rate, increase disease index, increase yield and improve quality.
2) Maize intercropping with double-genotypes is favorable for enhancing resistant metabolism. The artificial inoculation test showed that, in the intercropping treatment of ZD958‖LD98, the activities LOX, PAL and PPO of leaves increased rapidly and maintaining at the high activity level for a long period after inoculation. There was a significant difference for resistance action against southern rust between susceptible variety (ZD958) and resistant variety (LD981). LOX, PAL and PPO would play a key role in ZD958 against southern rust, while PPO would be very important in the same action for LD981. Results suggested that maize intercropping with different resistance genotypes would be favorable for improving defense ability of the intercropped colonies.
3) Maize intercropping with double-genotypes increases the genetic diversity of colony, which results in enriching leaf volatile organic compounds (VOCs) and the ability to resistant maize leaf south rust and maize leaf spot maize leaf spot. Maize leaf VOCs were extracted by steam distillation and analyzed by GC-MS. Results showed that there were great differences of the leaf VOCs among varieties, and the VOCs were changed in the intercropping treatments. The results of exposure to VOCs test indicate that the urediospore germinations of Puccinia polysora Underw were inhibited by leaf VOCs of LD981, HF9 andXD20, while conidia germinations of Helminthos porium maydis Nisik & Miy were marked inhibited by leaf VOCs of HF9 andXD20. In order to prove antimicrobial activity of maize leaf VOCs, a further test was conducted by five maize leaf volatile organic components such as (Z)-3-hexenol, (E)-2-hexenal, benzyl alcohol, vanillin and a-pinene. Results revealed that the urediospore germinations of Puccinia polysora Underw were inhibited significantly by the five components, the inhibitory rate from high to low were (Z)-3-hexenol, (E)-2-hexenal, benzyl alcohol, vanillin and a-pinene. Also, the conidia germinations of Helminthos porium maydis Nisik & Miy were inhibited markedly by the five components, and the inhibitory rates of the conidia germination were benzyl alcohol>vanillin>(Z)-3-hexenol> (E)-2-hexenal>α-pinene. Meanwhile, the mycelium growth of Helminthos porium maydis Nisik & Miy were significant inhibited markedly by the five components. Among them, the inhibitory rates of mycelium growth of vanillin, (E)-2-hexenal and (Z)-3-hexenol were more significant. The sporulation of the mycelium treated with high and low concentration of (Z)-3-hexenol was promoted, and the germination of spores generated from the mycelium was also promoted. In the treatments both (E)-2-hexenal and vanillin, the sporulations of mycelium were inhibited significantly, while the germinations of spores inhibited slightly. In the treatment of benzyl alcohol, the growth and sporulation of mycelium were mightily inhibited, and the mycelium was significant variation, the color of aerial mycelium was white with hardly sporulated. The treatment of a-pinene had a slight inhibition effect on the mycelium growth, while the sporulation quantity and the spore germination rate were lower. Results also indicated that the inhibitory effects of the treatments of different combination with the five volatile components on Puccinia polysora Underw and Helminthos porium maydis Nisik & Miy were more significant than the treatment of a single volatile organic component, which accord with natural condition of plant resistance.
In conclusion, the higher resistance against Puccinia polysora Underw and Helminthos porium maydis Nisik & Miy in maize intercropped system is based on the intercropped colonies ecology (including chemistry ecology) and physiological resistance. Suitable maize intercropping with different genotypes not only optimizes the population structure, improves the conditions of ventilation and transmission in the intercropped colonies, creates a micro-environment which is benefit for plant growth and makes against the pathogen growth. It also increases the genetic diversity of disease-resistant gene in the compound population, which results in increasing secondary metabolisms and leaf VOCs, and the chemical communication net of the compound population becomes more sophisticated and consequently improves the population resistance. Thus, leaf life would be prolonged, which be favor of improving grain yield and quality.
(1)双基因型玉米间作有利于改善群体结构、减少病原物侵染机率、降低病情指数。在ZD958‖LD98和HF9‖XD20两种间作模式中,4个品种间作植株的平均叶龄均有不同程度的增加；吐丝前,群体叶面积指数(leaf area index,LAI)单间作无明显差异,吐丝后,HF9和LD981的LAI分别大于和显著大于单作群体,而ZD958和XD20则分别小于和显著小于单作群体；吐丝后10d,4个品种棒三叶叶色值(soil and plant analyzer development readings,SPADRs)均有所增加,并且除ZD958外,其余3个品种棒三叶净光合速率均有所增加,其中LD981增加显著。间作对吐丝以前的群体干物质积累量影响不大,吐丝后,半紧凑型品种(HF9和LD981)的干物质积累量增加,其中LD981增加显著,而紧凑型品种(XD20和ZD958)的干物质积累量减少,其中ZD958显著减少。在ZD958‖LD98和YY19‖ZD041两种间作模式中,在吐丝后10d,4个玉米品种叶片中的SOD和POD活性均有所提高或显著提高,ZD041和ZD958叶片中的CAT活性提高或显著提高,YY19和LD981变化不显著；4个玉米品种叶片中的MDA含量降低。在吐丝后40d,SOD和POD活性及MDA含量的变化与吐丝后10d基本一致,CAT活性均有所提高,其中以YY19较为显著。紧凑型和半紧凑型品种间作增加了群体透光率。间作降低了间作群体引起继代感染的病原菌孢子数量,ZD958和XD20病情指数分别下降59.75%和50.38%,HF9和XD20小斑病病情也有所下降。间作还增加了复合群体产量,3种间作模式的土地当量比(land equivalent ratio,LER)均大于1,籽粒品质也有所改善。
(2)双基因型玉米间作增强了群体的抗病代谢。人工接种试验表明,抗南方锈病基因型LD981基因型与感南方锈病基因型ZD958间作,病原菌侵染后,能够及时启动间作群体的抗病代谢,LOX、PAL和PPO活性迅速提高,并维持较长时间的高活性水平；但抗、感病品种的表现有所差异,在感病品种ZD958抗南方锈病的代谢中,LOX、PAL和PPO均具有重要作用,而在抗病品种LD981抗南方锈病的代谢中,PPO所起的作用更大一些。
(3)双基因型玉米间作能够增加群体遗传多样性、丰富群体叶片挥发性物组分、增强对南方锈病和小斑病的抵抗能力。采用蒸馏法提取了玉米叶片挥发性物质,并对其组分进行了GC-MS分析。结果表明,玉米叶片挥发性组分及组分比在品种间和提取方法间存在较大差异,间作也能改变其组分及组分比。染毒试验结果表明,叶片挥发性物质能够抑制南方锈病和小斑病病原菌孢子萌发,并能抑制小斑病菌丝的生长和分生孢子的形成,对南方锈病病原菌孢子萌发的抑制作用尤为显著。5种挥发组分对南方锈病病原菌夏孢子萌发的抑制率从大到小依次为顺-3-己烯醇>反-2-己烯醛>苯甲醇>香草醛>α-蒎烯,对小斑病病原菌孢子萌发的抑制率从大到小依次为苯甲醇>香草醛>顺-3-己烯醇>反-2-己烯醛>α-蒎烯。香草醛、反-2-己烯醛和顺-3-己烯醇对小斑病病原菌菌丝生长的抑制作用最强,但低浓度和高浓度的顺-3-己烯醇处理对孢子的形成与萌发具有促进作用；反-2-己烯醛显著抑制孢子的形成,但高浓度处理对菌丝所产孢子的萌发无影响；香草醛显著抑制孢子的形成,但所产孢子的萌发率相对较高；苯甲醇对菌丝生长的影响较小,但菌丝颜色变异较大,气生菌丝发达呈灰白色至白色,难以产生分生孢子；α-蒎烯对菌丝生长的抑制作用最弱,菌丝性状基本正常,但产孢量较少,孢子的萌发率也较低。挥发组分组合作用的效果要优于单剂处理,这符合植物抗病的自然状况。
总之,双基因型玉米间作抗病具有深厚的生态生理基础,是群体生态(包括化学生态)和群体抗病生理共同作用的结果。双基因型玉米间作,不仅优化群体结构、改善群体通风透光条件、营造了一个有利于植株生长而不利于病原物滋生的微生态环境,而且增加了群体抗病基因的遗传多样性,群体次生代谢变得丰富多彩,群体中挥发性物质及其所建立起来的化学通讯网络更加复杂,提高了群体抵抗病原菌侵染的能力,从而延缓了叶片衰老,减少了病原物侵染的机率及玉米植株为抵御病害所付出的能量损失,提高了群体的光合速率和物质生产能力,促进了籽粒产量的提高和品质的改善。
Maize airborne diseases can be controlled efficiently via double-genotypes intercropping, but the mechanisms are little known. We presuppose that the eco-physiological factors such as compound population quality, disease resistant metabolism and maize leaf volatile organic compounds (LVOCs) may be of importance for controlling maize airborne diseases in this intercropped system. In this study, we conducted a field experiment on the farm of Henan Agricultural University, Zhengzhou, China, using six maize hybrids, Zhengdan 958(ZD958, compact plant type, susceptible to maize leaf south rust), Ludan 981(LD981, semi-compact plant type, highly resistant to maize leaf south rust), Zhoudan 041(ZD041, semi-compact plant type, resistant to maize leaf south rust disease), Yuyu 19(YY19, semi-compact plant type, susceptible to maize leaf south rust), Xundan 20(XD20, compact plant type, medium susceptible to maize leaf spot), Huafeng 9(HF9, semi-compact plant type, medium resistant to maize leaf spot). The experimental plots of 6 m×6 m were separated by 1 m. The experiments were planted by hand with rows oriented north-south and spacing of 0.70 and 0.50 m. The ratio of two maize varieties intercropped was 1:1 per row. The planting density was 67 500 plant·hm-2. There were nine treatments: each of the varieties alone, YY19 intercropped with ZD041 (YY19‖ZD041), ZD958 intercropped with LD981 (ZD958‖LD981), and HF9 intercropped with XD20 (HF9‖XD20). There were four replicate plots for each treatment, arranged in a randomized complete block design. We investigated the population quality, resistant physiology and relationship between airborne diseases resistance and maize leaf volatile organic compounds. Our objective was to reveal the eco-physiological basic for controlling maize airborne diseases in intercropping with double-genotypes. The results are as follows:
1) Maize intercropping with double-genotypes is favorable for improving plant population structure, enhancing population quality and reducing disease index. In the intercropping treatments both ZD958 II LD981 and HF9 II XD20, the average leaf life of four varieties increased. The leaf area indexes (LAI) of four varieties were no significant difference between sole-cropping and Intercropping before silking. But, the LAI of HF9 and LD981 increased while that of ZD958 and XD20 decreased in the intercropping treatments after silking. The soil and plant analyzer development readings (SPADRs) of ear-leaves of four varieties increased in the treatments 10 days after silking, and the photosynthesis rate of ear-leaves in LD981, HF9 and XD20 increased or increased significantly. There were no differences in the dry matter accumulation (DMA) of four varieties between the treatments of sole-cropping and intercropping before silking. The DMA of semi-compact plant types both HF9 and LD981 increased or increased markedly, while that of compact plant types both XD20 and ZD958 decreased or decreased significantly after silking. In the intercropping treatments both ZD958‖LD98 and YY19‖ZD041, the activities of SOD and POD of leaves increased in the intercropping treatments 10 days after silking, which resulted in decreased MDA contents of leaves. The activities of CAT of leaves in ZD041 and ZD958 increased or increased markedly, while YY19 and LD981 decreased slightly. The changes of activities of SOD and POD and contents of MDA in leaves remained similar 40 days after silking. Meanwhile, the activities of CAT of leaves increased, among them YY19 increased significantly. Transmittance enhanced in the treatment of compact plant type intercropped with semi-compact plant type, which resulted in reduced pathogen spore concentrations in the intercropped colonies, which led to reduced disease index of the intercropped colonies. For example, the disease index of southern rust in the intercropped colonies of ZD958 and HF9 reduced by 59.75% and 50.38%, respectively. Also, yields, land equivalent ratio (LER) and grain quality were enhanced in the intercropped colonies. Results suggested that suitable maize intercropping with different genotypes would increase the activities of SOD, POD and CAT of leaf, prolong leaf life, enhance photosynthesis rate, increase disease index, increase yield and improve quality.
2) Maize intercropping with double-genotypes is favorable for enhancing resistant metabolism. The artificial inoculation test showed that, in the intercropping treatment of ZD958‖LD98, the activities LOX, PAL and PPO of leaves increased rapidly and maintaining at the high activity level for a long period after inoculation. There was a significant difference for resistance action against southern rust between susceptible variety (ZD958) and resistant variety (LD981). LOX, PAL and PPO would play a key role in ZD958 against southern rust, while PPO would be very important in the same action for LD981. Results suggested that maize intercropping with different resistance genotypes would be favorable for improving defense ability of the intercropped colonies.
3) Maize intercropping with double-genotypes increases the genetic diversity of colony, which results in enriching leaf volatile organic compounds (VOCs) and the ability to resistant maize leaf south rust and maize leaf spot maize leaf spot. Maize leaf VOCs were extracted by steam distillation and analyzed by GC-MS. Results showed that there were great differences of the leaf VOCs among varieties, and the VOCs were changed in the intercropping treatments. The results of exposure to VOCs test indicate that the urediospore germinations of Puccinia polysora Underw were inhibited by leaf VOCs of LD981, HF9 andXD20, while conidia germinations of Helminthos porium maydis Nisik & Miy were marked inhibited by leaf VOCs of HF9 andXD20. In order to prove antimicrobial activity of maize leaf VOCs, a further test was conducted by five maize leaf volatile organic components such as (Z)-3-hexenol, (E)-2-hexenal, benzyl alcohol, vanillin and a-pinene. Results revealed that the urediospore germinations of Puccinia polysora Underw were inhibited significantly by the five components, the inhibitory rate from high to low were (Z)-3-hexenol, (E)-2-hexenal, benzyl alcohol, vanillin and a-pinene. Also, the conidia germinations of Helminthos porium maydis Nisik & Miy were inhibited markedly by the five components, and the inhibitory rates of the conidia germination were benzyl alcohol>vanillin>(Z)-3-hexenol> (E)-2-hexenal>α-pinene. Meanwhile, the mycelium growth of Helminthos porium maydis Nisik & Miy were significant inhibited markedly by the five components. Among them, the inhibitory rates of mycelium growth of vanillin, (E)-2-hexenal and (Z)-3-hexenol were more significant. The sporulation of the mycelium treated with high and low concentration of (Z)-3-hexenol was promoted, and the germination of spores generated from the mycelium was also promoted. In the treatments both (E)-2-hexenal and vanillin, the sporulations of mycelium were inhibited significantly, while the germinations of spores inhibited slightly. In the treatment of benzyl alcohol, the growth and sporulation of mycelium were mightily inhibited, and the mycelium was significant variation, the color of aerial mycelium was white with hardly sporulated. The treatment of a-pinene had a slight inhibition effect on the mycelium growth, while the sporulation quantity and the spore germination rate were lower. Results also indicated that the inhibitory effects of the treatments of different combination with the five volatile components on Puccinia polysora Underw and Helminthos porium maydis Nisik & Miy were more significant than the treatment of a single volatile organic component, which accord with natural condition of plant resistance.
In conclusion, the higher resistance against Puccinia polysora Underw and Helminthos porium maydis Nisik & Miy in maize intercropped system is based on the intercropped colonies ecology (including chemistry ecology) and physiological resistance. Suitable maize intercropping with different genotypes not only optimizes the population structure, improves the conditions of ventilation and transmission in the intercropped colonies, creates a micro-environment which is benefit for plant growth and makes against the pathogen growth. It also increases the genetic diversity of disease-resistant gene in the compound population, which results in increasing secondary metabolisms and leaf VOCs, and the chemical communication net of the compound population becomes more sophisticated and consequently improves the population resistance. Thus, leaf life would be prolonged, which be favor of improving grain yield and quality.
引文
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