旱作水稻/西瓜间作抑制西瓜枯萎病的生理机制
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摘要
间作是我国传统农业的组成部分,具有增大光合面积,提高土地利用率及避免根系在同一层次的养分竞争等间作优势。此外,间作种植还可以抑制植物的病虫害。西瓜枯萎病是西瓜生产中易发的病害,而且危害严重。前期的盆栽试验表明,旱作水稻/西瓜间作能够抑制西瓜枯萎病的发生。对其作用机制进一步的研究,证实了水稻根分泌物能够抑制西瓜枯萎病病原菌尖孢镰刀菌的生长和繁殖,这是水稻根分泌物从改善西瓜生长环境方面抑制西瓜枯萎病发生的机制,但是,水稻根分泌物能否到达西瓜根际、到达根际后能否提高西瓜自身的抗病能力,需要我们做进一步的研究。香豆酸是水稻根系分泌的酚酸,能显著抑制西瓜专化型尖孢镰刀菌的生长和繁殖。阿魏酸是西瓜根系分泌的酚酸,它促进西瓜专化型尖孢镰刀菌的生长和繁殖。水杨酸是水稻和西瓜根系都分泌的酚酸,同时也是一种激素,对西瓜专化型尖孢镰刀菌的生长和繁殖没有显著的影响。香豆酸、阿魏酸和水杨酸很可能对西瓜植株的系统获得性抗性有不同的影响,从而影响西瓜自身的抗枯萎病的能力。本研究采用盆栽、分隔盆栽和14C标记等方法,从微生物环境到生理水平和转录水平,研究了14C的放射活性、西瓜根际病原菌数量、西瓜根际微生物区系、水稻和西瓜对磷素的吸收、西瓜植株几丁质酶基因和苯丙氨酸解氨酶(PAL)基因的表达、西瓜植株几丁质酶活性和β-1,3-葡聚糖酶活性,具体采用的方法和主要结果如下：
1.采用根箱和14C标记的方法,研究了水稻同化的碳在水稻和西瓜之间的传递,以及丛枝菌根真菌对旱作水稻同化的碳素向西瓜根际传递的影响。将根箱分为标记侧和采样侧,在箱子中间插入30μm尼龙网。结果表明,与旱作水稻单作相比,旱作水稻和西瓜间作时,有更多的C分配到根系并分泌到水稻根际土壤中。无论旱作水稻单作还是旱作水稻/西瓜间作,当接种丛枝菌根真菌时,水稻同化的C会有更多的量传递到邻近作物根际(单作时是水稻,间作时为西瓜),并被邻近植物吸收、转运到地上部分。
2.采用中间不同分隔的大盆钵试验,研究在西瓜／旱作水稻间作系统中西瓜和旱作水稻根系之间不同的分隔方式对西瓜枯萎病的影响,并从西瓜根际病原菌及微生物区系方面探讨其作用机制。结果表明,与隔膜相比,当西瓜和旱作水稻根系之间没有分隔时,西瓜枯萎病的发病时间晚6-7天,发病程度最轻；当西瓜和旱作水稻根系之间用尼龙网隔开时,西瓜和旱作水稻的根系之间没有根系接触,而根分泌物可以自由穿过尼龙网,与隔膜的处理相比,西瓜枯萎病的发病时间推迟3-4天,发病程度也减轻,根际尖孢镰刀菌数量降低,真菌数量降低,细菌数量升高。当西瓜和旱作水稻之间用塑料膜隔开时,西瓜枯萎病的发病时间最早,发病程度最深,与不分隔的处理相比,西瓜根际的尖孢镰刀菌数量升高,真菌和细菌数量升高,放线菌数量降低。因此,在西瓜/旱作水稻间作系统对西瓜枯萎病的抑制中,旱作水稻根分泌物对西瓜枯萎病病原菌尖孢镰刀菌的抑制,和旱作水稻根分泌物改善西瓜根际微生物区系都起着重要的作用。
3.采用盆栽的方法向西瓜外源施加不同浓度的香豆酸,研究在水稻分泌的酚酸作用下能否提高西瓜自身抗枯萎病的能力。结果表明,西瓜根际接种尖孢镰刀菌下调了西瓜叶片几丁质酶基因和PAL基因的表达,叶片β-1,3-葡聚糖酶活性降低了75.80%,降低了西瓜的抗病能力；在西瓜外源施加香豆酸提高了西瓜根系几丁质酶基因和叶片PAL基因的表达,提高了根系中的几丁质酶和β-1,3-葡聚糖酶的活性,提高了根系分泌肉桂酸的量,从而提高了西瓜抗枯萎病的能力。当西瓜根际接种尖孢镰刀菌同时施加香豆酸时,随着香豆酸施加量的增加,叶片中的几丁质酶基因的表达逐渐升高,PAL基因表达量逐渐提高。当香豆酸的施加量为100μmol L-1时根系β-1,3-葡聚糖酶活性达到最高,比对照提高了53.7%。所以,香豆酸的施加提高了西瓜抗枯萎病的能力,而且在施加量在100μmol L-1时效果最好。
4.采用盆栽的方法,向西瓜植株外源施加阿魏酸,以模拟西瓜根分泌物对西瓜自身生理代谢以及西瓜抗病能力的影响。结果表明,西瓜植株外源施加阿魏酸下调西瓜叶片几丁质酶基因和根系PAL基因的表达,当施加阿魏酸浓度为100μmol L-1时,对几丁质酶基因抑制最强,使西瓜叶片几丁质酶活性降低65.2%；施加30μmol L-1阿魏酸时,根系PAL基因下调最显著；单独接种尖孢镰刀菌,西瓜叶片β-1,3-葡聚糖酶活性降低75.8%,当接种尖孢镰刀菌同时施加30μmol L-1-300μmol L-1阿魏酸时,西瓜叶片β-1,3-葡聚糖酶活性比对照处理降低了32%-37%。总之,外源施加阿魏酸抑制西瓜叶片和根系几丁质酶活性,抑制西瓜叶片β-1,3-葡聚糖酶活性,从而降低西瓜植株的抗病能力,诱导西瓜枯萎病的发生。
5.采用盆栽的方法,向西瓜外源施加水杨酸,研究了西瓜叶片和根中几丁质酶基因和PAL基因的表达,以及几丁质酶活性和β-1,3-葡聚糖酶的活性,探讨外源激素对西瓜植株叶片和根系中与抗性相关基因的表达及其功能蛋白几丁质酶和β-1,3-葡聚糖酶活性的影响,揭示水杨酸的施用对西瓜自身抗枯萎病能力的影响。结果表明,西瓜外源施加300μmol L-1水杨酸9天后,刺激了西瓜根系的生长；上调了西瓜叶片和根系中的PAL基因的表达,也上调了根系几丁质酶基因的表达,而对西瓜根系中的几丁质酶基因却有下调的趋势。与对照相比,施加30μmol L-1和100μmol L-1水杨酸时,西瓜叶片几丁质酶的活性分别提高了2.25倍和2.45倍,施加300μmol L-1水杨酸时,西瓜苗根系几丁质酶活性提高了52.2%。施加30μmol L-1和100μmol L-1水杨酸时,西瓜叶片中β-1,3-葡聚糖酶的活性分别提高了56.7%和50.7%,当施加浓度达到300μmol L-1时,西瓜叶片β-1,3-葡聚糖酶活性反而比对照降低了76.6%；西瓜外源施加水杨酸提高了根系分泌肉桂酸的量,当施加浓度为100μmol L-1时,其分泌量最高,与对照处理相比,提高了9.63倍。因此,外源施加100μmol L-1水杨酸在上调西瓜植株几丁质酶和PAL基因的表达,提高几丁质酶和β-1,3-葡聚糖酶的活性,提高根系对肉桂酸的分泌等方面效果最好,以提高西瓜的抗病能力。
综上所述,本文阐述了碳素在土壤中的传递,AM菌根真菌对植物碳素传递的影响,旱作水稻和西瓜根系之间的分隔对西瓜根际微生物和西瓜枯萎病的影响,水稻根分泌的香豆酸对西瓜苗抗病能力的影响,西瓜根分泌物的阿魏酸对自身抗病能力的影响以及外源激素对西瓜苗抗病能力的影响。旱作水稻和西瓜间作时,旱作水稻的根分泌物一方面直接抑制尖孢镰刀菌的生长和繁殖,改善了西瓜生长的外界环境；另一方面,通过分泌的化感物质刺激西瓜根系,改善西瓜自身的生理代谢,产生系统获得抗性,提高抗病能力。
Intercropping mode is a traditional agricultural management in China, with the advantages such as enhancing photosynthesis areas, increasing land use efficiency and avoiding nutrient competition by roots. In addition, intercropping can also control plant disease. Watermelon is susceptive to wilt disease, which usually results in reducing its yield and quality. Previous pot experiment showed that intercropping with aerobic rice could suppress wilt disease occurrence in watermelon, which is related to that rice root exudates suppressed F. oxysporum. Intercropping with aerobic rice or arbuscular mycorrhizal fungi (AMF) colonization alleviated watermelon wilt disease, which is likely attributed to rice root exudates or AMF depressing watermelon wilt pathogen. Coumaric acid is the specific substance in rice root exudates, with the ability to inhibit spore germination and spore procreation of F. oxysporum. However, further research is needed to investigate whether rice root exudates can transfer to watermelon rhizosphere soil, whether rice root exudates can induce systemic resistance against wilt disease in watermelon and whether AMF can affect the transfer of rice root exudates to watermelon rhizosphere soil. Ferulic acid is the specific substance in watermelon root exudates, with the ability to improve spore germination and spore procreation of F. oxysporum. Salicylic acid is exudated by both rice and watermelon roots. It has no effect on spore germination and spore procreation of F. oxysporum. Salicylic acid is an essential hormone in plant immunity. A pot experiment with14C labeling method was performed to investigate the transfer of rice root exudates to watermelon rhizosphere. A pot experiment with separation between rice and watermelon roots was conducted to detect the role of rice root exudates in alleviating watermelon wilt disease. Exogenous coumaric acid, ferulic acid and salicylic acid were applied to watermelon to investigate the effect of these phenolic acids on watermelon disease resistance. Chitinase gene and PAL gene expression as well as enzymatic acitvities were determined to show watermelon disease resistance.The methods and results were listed as below:
1.A rhizobox experiment,with aerobic rice under14C02,was conducted to investigate the effect of AMF colonization on carbon(C)transfer from rice to watermelon and phosphorus(P)uptake by both watermelon and rice.The rhizobox was separated into labelling side(L side)and sampling side(S side)by inserting nylon mesh in the middle of the box.The L side was planted with aerobic rice,and the S side was aerobic rice (monocropping)or watermelon(intercropping).When14CO2was added to rice canopy at the L side,14C activities of rice roots and rhizosphere soils in the L side were increased by intercropping with watermelon or AMF colonization.The14C was detected in roots and rhizosphere soils of rice and watermelon in the S side,but no differences were found among different treatments.14C activities in leaves were improved by AMF inoculation in the S side,regardless of rice or watermelon.Mycorrhizal colonization stimulated P absorption and translocation to rice in intercropping system.These findings suggest that AMF colonization could increase C transfer from rice to watermelon while intercropping with watermelon could promote AMF colonization and P uptake by rice.
2. Pot experiment with different separation methods between aerobic rice and watermelon was performed to investigate the effect of separation methods on watermelon wilt disease and its mechanism.The results showed that the wilt disease occurrence was delayed6-7days and was alleviated in the treatment with no separation between aerobic rice and watermelon roots,compared with film separation treatment.The wilt disease occurrence was delayed3-4days in the treatment with nylon mesh separation between aerobic rice and watermelon roots.Compared to film separation,the F. oxysporum number, bacterial number and fungal number decreased while the actinomycetes number increased in watermelon rhizosphere soil in no separation treatment.The similar results were obtained in nylon mesh separation treatment.
3.Pot experiment was carried out to investigate the effect of exogenous coumaric acid supply to watermelon rhizosphere on chitinase and phenylalanine ammonia lyase (PAL) genes expression and related enzymatic activities.The results showed that chitinase and PAL genes in watermelon leaf were down regulated.The β-1,3-glucanase activityin leaf was decreased by75.8%with F. oxysproum inoculation.Chitinase gene in root and PAL gene in leaf were up-regulated with exogenous supply of coumaric acid to watermelon.The activities of chitinase and β-1,3-glucanase in roots were increased,and more cinnamic acid was excreted from watermelon root with supplying exogenous coumaric acid. Under supplying both F. oxysporum and coumaric acid, chitinase gene and PAL gene expression in leaf increased with increasing the supply. Compared with control, the activity of β-1,3-glucanase in leaf was increased by53.7%when100μmol L-1coumaric acid was applied to watermelon. Therefore, exogenous coumaric acid improved watermelon disease resistance and100μmol L-1is the best dose for exogenous supply.
4. Pot experiment was carried out to investigate the effect of exogenous ferulic acid supply on watermelon resistance. The results showed that chitinase gene in leaf and PAL gene in root were down-regulated with ferulic acid supply. Compared with control, chitinase activity in leaf was decreased by65.2%with100μmol L-1ferulic acid supply. The β-1,3-glucanase activity in watermelon leaf was reduced by75.8%with F. oxysporum inoculation, and decreased by32%-37%under supplying both F. oxysporum and30-300μmol L-1ferulic acid, compared with the control. Therefore, supplying exogenous ferulic acid decreased chitinase activity in leaves and roots and β-1,3-glucanase activity in leaves of watermelon, and thus decreased disease resistance throuth down regulating chitinase gene and PAL gene, and induced wilt disease occurrence in watermelon.
5. Exogenous supply of salicylic acid to watermelon rhizosphere was performed to study the effect of hormone secreted by rice and watermelon roots on watermelon disease resistance. The results showed that watermelon root growth was improved by300βmol L-1salicylic acid supply for9days. In addition, the PAL gene expression in watermelon leaf and root was up regulated, as well as the chitinase gene in root with300μmol L-1salicylic acid supply for9days. Compared with control, the chitinase activity in leaf was promoted by2.25and2.45times with30μmol L-1and100μmol L-1salicylic acid application, respectively. The chitinase activity in root was increased by52.2%with300μmol L-1exogenous salicylic acid. Compared with control, the β-1,3-glucanase activity in leaf was enhanced by56.7%and50.7%with adding exogenous30μmol L-1and100μumol L-1salicylic acid, respectively. However, the β-1,3-glucanase activity decreased by76.6%with300μmol L-1salicylic acid. Commanic acid concentration in watermelon root exudates was improved by salicylic acid, and with the highest at100μmol L-1. The commanic acid concentration was increased by9.63times with100μmol L-1salicylic acid supply, compared with the control. Therefore,100μmol L-1salicylic acid aupply was better in up-regulating chitinase gene and PAL gene expression, in improving chitinase and β-1,3-glucanase activities and in secreting more cinnamic acid to improve watermelon wilt disease resistance.
The present study demonstrated the mechanism of suppression watermelon wilt disease by intercropped with aerobic rice. Rice root exudates transferred to watermelon rhizosphere to inhibit F. oxysporum growth. On the other hand, rice root exudates induced systemic acquired resistance of watermelon to decrease wilt disease incidence.
1.采用根箱和14C标记的方法,研究了水稻同化的碳在水稻和西瓜之间的传递,以及丛枝菌根真菌对旱作水稻同化的碳素向西瓜根际传递的影响。将根箱分为标记侧和采样侧,在箱子中间插入30μm尼龙网。结果表明,与旱作水稻单作相比,旱作水稻和西瓜间作时,有更多的C分配到根系并分泌到水稻根际土壤中。无论旱作水稻单作还是旱作水稻/西瓜间作,当接种丛枝菌根真菌时,水稻同化的C会有更多的量传递到邻近作物根际(单作时是水稻,间作时为西瓜),并被邻近植物吸收、转运到地上部分。
2.采用中间不同分隔的大盆钵试验,研究在西瓜／旱作水稻间作系统中西瓜和旱作水稻根系之间不同的分隔方式对西瓜枯萎病的影响,并从西瓜根际病原菌及微生物区系方面探讨其作用机制。结果表明,与隔膜相比,当西瓜和旱作水稻根系之间没有分隔时,西瓜枯萎病的发病时间晚6-7天,发病程度最轻；当西瓜和旱作水稻根系之间用尼龙网隔开时,西瓜和旱作水稻的根系之间没有根系接触,而根分泌物可以自由穿过尼龙网,与隔膜的处理相比,西瓜枯萎病的发病时间推迟3-4天,发病程度也减轻,根际尖孢镰刀菌数量降低,真菌数量降低,细菌数量升高。当西瓜和旱作水稻之间用塑料膜隔开时,西瓜枯萎病的发病时间最早,发病程度最深,与不分隔的处理相比,西瓜根际的尖孢镰刀菌数量升高,真菌和细菌数量升高,放线菌数量降低。因此,在西瓜/旱作水稻间作系统对西瓜枯萎病的抑制中,旱作水稻根分泌物对西瓜枯萎病病原菌尖孢镰刀菌的抑制,和旱作水稻根分泌物改善西瓜根际微生物区系都起着重要的作用。
3.采用盆栽的方法向西瓜外源施加不同浓度的香豆酸,研究在水稻分泌的酚酸作用下能否提高西瓜自身抗枯萎病的能力。结果表明,西瓜根际接种尖孢镰刀菌下调了西瓜叶片几丁质酶基因和PAL基因的表达,叶片β-1,3-葡聚糖酶活性降低了75.80%,降低了西瓜的抗病能力；在西瓜外源施加香豆酸提高了西瓜根系几丁质酶基因和叶片PAL基因的表达,提高了根系中的几丁质酶和β-1,3-葡聚糖酶的活性,提高了根系分泌肉桂酸的量,从而提高了西瓜抗枯萎病的能力。当西瓜根际接种尖孢镰刀菌同时施加香豆酸时,随着香豆酸施加量的增加,叶片中的几丁质酶基因的表达逐渐升高,PAL基因表达量逐渐提高。当香豆酸的施加量为100μmol L-1时根系β-1,3-葡聚糖酶活性达到最高,比对照提高了53.7%。所以,香豆酸的施加提高了西瓜抗枯萎病的能力,而且在施加量在100μmol L-1时效果最好。
4.采用盆栽的方法,向西瓜植株外源施加阿魏酸,以模拟西瓜根分泌物对西瓜自身生理代谢以及西瓜抗病能力的影响。结果表明,西瓜植株外源施加阿魏酸下调西瓜叶片几丁质酶基因和根系PAL基因的表达,当施加阿魏酸浓度为100μmol L-1时,对几丁质酶基因抑制最强,使西瓜叶片几丁质酶活性降低65.2%；施加30μmol L-1阿魏酸时,根系PAL基因下调最显著；单独接种尖孢镰刀菌,西瓜叶片β-1,3-葡聚糖酶活性降低75.8%,当接种尖孢镰刀菌同时施加30μmol L-1-300μmol L-1阿魏酸时,西瓜叶片β-1,3-葡聚糖酶活性比对照处理降低了32%-37%。总之,外源施加阿魏酸抑制西瓜叶片和根系几丁质酶活性,抑制西瓜叶片β-1,3-葡聚糖酶活性,从而降低西瓜植株的抗病能力,诱导西瓜枯萎病的发生。
5.采用盆栽的方法,向西瓜外源施加水杨酸,研究了西瓜叶片和根中几丁质酶基因和PAL基因的表达,以及几丁质酶活性和β-1,3-葡聚糖酶的活性,探讨外源激素对西瓜植株叶片和根系中与抗性相关基因的表达及其功能蛋白几丁质酶和β-1,3-葡聚糖酶活性的影响,揭示水杨酸的施用对西瓜自身抗枯萎病能力的影响。结果表明,西瓜外源施加300μmol L-1水杨酸9天后,刺激了西瓜根系的生长；上调了西瓜叶片和根系中的PAL基因的表达,也上调了根系几丁质酶基因的表达,而对西瓜根系中的几丁质酶基因却有下调的趋势。与对照相比,施加30μmol L-1和100μmol L-1水杨酸时,西瓜叶片几丁质酶的活性分别提高了2.25倍和2.45倍,施加300μmol L-1水杨酸时,西瓜苗根系几丁质酶活性提高了52.2%。施加30μmol L-1和100μmol L-1水杨酸时,西瓜叶片中β-1,3-葡聚糖酶的活性分别提高了56.7%和50.7%,当施加浓度达到300μmol L-1时,西瓜叶片β-1,3-葡聚糖酶活性反而比对照降低了76.6%；西瓜外源施加水杨酸提高了根系分泌肉桂酸的量,当施加浓度为100μmol L-1时,其分泌量最高,与对照处理相比,提高了9.63倍。因此,外源施加100μmol L-1水杨酸在上调西瓜植株几丁质酶和PAL基因的表达,提高几丁质酶和β-1,3-葡聚糖酶的活性,提高根系对肉桂酸的分泌等方面效果最好,以提高西瓜的抗病能力。
综上所述,本文阐述了碳素在土壤中的传递,AM菌根真菌对植物碳素传递的影响,旱作水稻和西瓜根系之间的分隔对西瓜根际微生物和西瓜枯萎病的影响,水稻根分泌的香豆酸对西瓜苗抗病能力的影响,西瓜根分泌物的阿魏酸对自身抗病能力的影响以及外源激素对西瓜苗抗病能力的影响。旱作水稻和西瓜间作时,旱作水稻的根分泌物一方面直接抑制尖孢镰刀菌的生长和繁殖,改善了西瓜生长的外界环境；另一方面,通过分泌的化感物质刺激西瓜根系,改善西瓜自身的生理代谢,产生系统获得抗性,提高抗病能力。
Intercropping mode is a traditional agricultural management in China, with the advantages such as enhancing photosynthesis areas, increasing land use efficiency and avoiding nutrient competition by roots. In addition, intercropping can also control plant disease. Watermelon is susceptive to wilt disease, which usually results in reducing its yield and quality. Previous pot experiment showed that intercropping with aerobic rice could suppress wilt disease occurrence in watermelon, which is related to that rice root exudates suppressed F. oxysporum. Intercropping with aerobic rice or arbuscular mycorrhizal fungi (AMF) colonization alleviated watermelon wilt disease, which is likely attributed to rice root exudates or AMF depressing watermelon wilt pathogen. Coumaric acid is the specific substance in rice root exudates, with the ability to inhibit spore germination and spore procreation of F. oxysporum. However, further research is needed to investigate whether rice root exudates can transfer to watermelon rhizosphere soil, whether rice root exudates can induce systemic resistance against wilt disease in watermelon and whether AMF can affect the transfer of rice root exudates to watermelon rhizosphere soil. Ferulic acid is the specific substance in watermelon root exudates, with the ability to improve spore germination and spore procreation of F. oxysporum. Salicylic acid is exudated by both rice and watermelon roots. It has no effect on spore germination and spore procreation of F. oxysporum. Salicylic acid is an essential hormone in plant immunity. A pot experiment with14C labeling method was performed to investigate the transfer of rice root exudates to watermelon rhizosphere. A pot experiment with separation between rice and watermelon roots was conducted to detect the role of rice root exudates in alleviating watermelon wilt disease. Exogenous coumaric acid, ferulic acid and salicylic acid were applied to watermelon to investigate the effect of these phenolic acids on watermelon disease resistance. Chitinase gene and PAL gene expression as well as enzymatic acitvities were determined to show watermelon disease resistance.The methods and results were listed as below:
1.A rhizobox experiment,with aerobic rice under14C02,was conducted to investigate the effect of AMF colonization on carbon(C)transfer from rice to watermelon and phosphorus(P)uptake by both watermelon and rice.The rhizobox was separated into labelling side(L side)and sampling side(S side)by inserting nylon mesh in the middle of the box.The L side was planted with aerobic rice,and the S side was aerobic rice (monocropping)or watermelon(intercropping).When14CO2was added to rice canopy at the L side,14C activities of rice roots and rhizosphere soils in the L side were increased by intercropping with watermelon or AMF colonization.The14C was detected in roots and rhizosphere soils of rice and watermelon in the S side,but no differences were found among different treatments.14C activities in leaves were improved by AMF inoculation in the S side,regardless of rice or watermelon.Mycorrhizal colonization stimulated P absorption and translocation to rice in intercropping system.These findings suggest that AMF colonization could increase C transfer from rice to watermelon while intercropping with watermelon could promote AMF colonization and P uptake by rice.
2. Pot experiment with different separation methods between aerobic rice and watermelon was performed to investigate the effect of separation methods on watermelon wilt disease and its mechanism.The results showed that the wilt disease occurrence was delayed6-7days and was alleviated in the treatment with no separation between aerobic rice and watermelon roots,compared with film separation treatment.The wilt disease occurrence was delayed3-4days in the treatment with nylon mesh separation between aerobic rice and watermelon roots.Compared to film separation,the F. oxysporum number, bacterial number and fungal number decreased while the actinomycetes number increased in watermelon rhizosphere soil in no separation treatment.The similar results were obtained in nylon mesh separation treatment.
3.Pot experiment was carried out to investigate the effect of exogenous coumaric acid supply to watermelon rhizosphere on chitinase and phenylalanine ammonia lyase (PAL) genes expression and related enzymatic activities.The results showed that chitinase and PAL genes in watermelon leaf were down regulated.The β-1,3-glucanase activityin leaf was decreased by75.8%with F. oxysproum inoculation.Chitinase gene in root and PAL gene in leaf were up-regulated with exogenous supply of coumaric acid to watermelon.The activities of chitinase and β-1,3-glucanase in roots were increased,and more cinnamic acid was excreted from watermelon root with supplying exogenous coumaric acid. Under supplying both F. oxysporum and coumaric acid, chitinase gene and PAL gene expression in leaf increased with increasing the supply. Compared with control, the activity of β-1,3-glucanase in leaf was increased by53.7%when100μmol L-1coumaric acid was applied to watermelon. Therefore, exogenous coumaric acid improved watermelon disease resistance and100μmol L-1is the best dose for exogenous supply.
4. Pot experiment was carried out to investigate the effect of exogenous ferulic acid supply on watermelon resistance. The results showed that chitinase gene in leaf and PAL gene in root were down-regulated with ferulic acid supply. Compared with control, chitinase activity in leaf was decreased by65.2%with100μmol L-1ferulic acid supply. The β-1,3-glucanase activity in watermelon leaf was reduced by75.8%with F. oxysporum inoculation, and decreased by32%-37%under supplying both F. oxysporum and30-300μmol L-1ferulic acid, compared with the control. Therefore, supplying exogenous ferulic acid decreased chitinase activity in leaves and roots and β-1,3-glucanase activity in leaves of watermelon, and thus decreased disease resistance throuth down regulating chitinase gene and PAL gene, and induced wilt disease occurrence in watermelon.
5. Exogenous supply of salicylic acid to watermelon rhizosphere was performed to study the effect of hormone secreted by rice and watermelon roots on watermelon disease resistance. The results showed that watermelon root growth was improved by300βmol L-1salicylic acid supply for9days. In addition, the PAL gene expression in watermelon leaf and root was up regulated, as well as the chitinase gene in root with300μmol L-1salicylic acid supply for9days. Compared with control, the chitinase activity in leaf was promoted by2.25and2.45times with30μmol L-1and100μmol L-1salicylic acid application, respectively. The chitinase activity in root was increased by52.2%with300μmol L-1exogenous salicylic acid. Compared with control, the β-1,3-glucanase activity in leaf was enhanced by56.7%and50.7%with adding exogenous30μmol L-1and100μumol L-1salicylic acid, respectively. However, the β-1,3-glucanase activity decreased by76.6%with300μmol L-1salicylic acid. Commanic acid concentration in watermelon root exudates was improved by salicylic acid, and with the highest at100μmol L-1. The commanic acid concentration was increased by9.63times with100μmol L-1salicylic acid supply, compared with the control. Therefore,100μmol L-1salicylic acid aupply was better in up-regulating chitinase gene and PAL gene expression, in improving chitinase and β-1,3-glucanase activities and in secreting more cinnamic acid to improve watermelon wilt disease resistance.
The present study demonstrated the mechanism of suppression watermelon wilt disease by intercropped with aerobic rice. Rice root exudates transferred to watermelon rhizosphere to inhibit F. oxysporum growth. On the other hand, rice root exudates induced systemic acquired resistance of watermelon to decrease wilt disease incidence.
引文
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