草莓连作根腐病发生机制与微生物及化学修复研究
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摘要
连作障碍是限制世界草莓产业持续发展的瓶颈。本研究从根际微生态角度出发，通过连作草莓病健株根区土壤化学性质及微生物区系组成的比较研究，揭示了草莓连作根腐病害发生的土壤微生态机制，探索利用植物病健株根区土壤微生物区系比较法快速筛选高效广谱且有实际防病促生作用生防微生物的可行性。同时采用盆栽与日光温室试验，探索了利用微生物菌剂及硅肥对草莓连作障碍的修复效应，并从抑病促生、促进养分吸收及土壤微生态修复等角度探讨了其作用机理，以期为草莓连作障碍的综合修复提供理论依据。主要研究结果如下：
1.草莓连作根腐病害发生的根域土壤微生态机制研究。
本研究表明，草莓连作根腐病害的发生与根区土壤化学性质及微生物区系组成及其多样性变化密切相关。
（1）连作草莓病株根区土壤水溶性盐分含量较健株提高了50.6%～156.5%，有机质，速效磷及速效钾含量较健株分别下降22.2%～27.5%，15.3%～17.6%及10.9%～37.9%。
（2）病株根表土壤细菌与真菌数量比值B/F降低了2.9%～50.0%，微生物群落结构趋向“真菌型”发展，细菌、真菌多样性指数H及丰富度指数S呈降低趋势。病健株根区、根表优势真菌种类、数量及比例差异明显，其中健株以具较强拮抗性的2株青霉（Penicillium oxalicum，P. griseofulvum）及1株土曲霉（Aspergillus terreus）为主，而病株则以枝孢霉（Cladosporium sp.）和大双孢柱孢（Cylindrocarpon macrodidyma）等有害病原菌为主。大双孢柱孢C. macrodidyma CF9对草莓根系具有强烈侵染致病作用。
2.草莓根腐病原真菌分离及其拮抗真菌筛选
本研究采用病健株根区及根表土壤微生物区系比较法获得对草莓根系具有强烈侵染致病作用的根腐病原真菌大双孢柱孢C. macrodidyma CF9，同时筛选到2株广谱拮抗真菌灰黄青霉（P. griseofulvum CF3）和土曲霉（A. terreus CF7）。证明此法是快速锁定植物根腐病病原有害微生物及有益拮抗微生物可靠与可行的方法。
3.硅酸钾对病原真菌抑菌作用及其机理研究
硅酸钾对病原真菌菌丝生长具有较强抑制作用，其抑菌机理是通过硅酸钾提高培养基pH实现的，该结果否定了现有研究提出的硅对病原真菌具有类似抗生素作用的观点。
4.微生物及化学防治措施对连作草莓的修复效应研究。
（1）微生物活菌制剂对连作草莓具有明显的防病促生作用。2株拮抗真菌CF3和CF7可使盆栽草莓根腐发病率分别降低52.4%和48.7%，根鲜重分别增加15.2%和19.6%，果实产量分别提高26.7%和31.3%；放线菌Act12活菌制剂可使日光温室草莓根鲜重增加135.5%，开花提前6d，果实产量提高47.5%，同时改善了果实品质，盛果期可溶性固形物和糖酸比分别增加了11.6%和11.3%。
（2）硅酸钾和粉煤灰对连作草莓也有明显的防病促生作用。施用3.33mmol·kg-1硅酸钾对第3年盆栽草莓根腐病的防效为36.5%，开花提前5d，根鲜重和果实产量增加了31.3%和47.4%。日光温室每株施用60mg硅酸钾草莓开花提前6d，根鲜重和果实产量增加了118.0%和102.3%。盆栽施用100g·kg-1粉煤灰对第3年草莓根腐病的防效为36.2%，开花提前6d，根鲜重和果实产量增加了95.5%和50.5%。。
（3）硅酸钾及粉煤灰与菌剂配施除具有三者单独施用的效果外，在部分处理中，硅肥与放线菌剂配施具有增效作用。硅酸钾与菌剂配施可使日光温室草莓植株死亡率降低至0%，开花提前10d，植株总鲜重，根鲜重及单株产量分别提高73.7%，71.3%及126.2%，同时盛果期果实可溶性固形物含量，糖酸比及维生素C含量分别提高21.7%，36.2%及27.0%。盆栽试验表明，3.33mmol·kg-1硅酸钾与菌剂配施对第3年草莓根腐病的防效为58.6%，开花提前8d，根鲜重及单株产量分别提高73.6%和76.6%；100g·kg-1粉煤灰与菌剂配施对第3年草莓根腐病的防效为56.2%，开花提前6d，根鲜重及单株产量分别提高105.0%和91.0%。
5.微生物及化学修复措施对草莓连作障碍的修复机理研究。
（1）抗生及溶菌作用。2株拮抗真菌及放线菌Act12均可产生抗菌活性产物，通过抗菌活性代谢产物抑制草莓连作土传病原真菌菌丝生长，其中CF3对大双孢柱孢CF9菌丝具有重寄生和溶菌作用。
（2）放线菌制剂能刺激草莓根系生长，促进植株对土壤养分的吸收利用。Act12菌剂单施或与2种硅肥配施均可促进连作草莓对氮、磷、钾元素吸收。
（3）提高根系硅元素含量，增强根系抗病性。施用硅肥增加了土壤有效硅含量，促进草莓植株根系硅素的吸收积累，增加了根系含硅量，增强了草莓对根腐病的抗病性。
（4）改善根系生理功能，增强植物的诱导抗性。Act12菌剂、硅酸钾及粉煤灰均可影响草莓根系生长发育，增强在根腐病原真菌胁迫下草莓的根系活力；诱导草莓产生抗病性，提高草莓叶片超氧化物歧化酶活性（SOD）、过氧化物酶活性（POD）、多酚氧化酶活性（PPO）及苯丙氨酸解氨酶（PAL）主要防御酶活性。
（5）恢复草莓根区、根表土壤微生物区系平衡。Act12菌剂、硅酸钾及粉煤灰均可增加土壤细菌、放线菌数量，减少土壤真菌数量，同时提高B/F和A/F值，提高土壤细菌、真菌及放线菌的多样性；硅肥与菌剂配施可有效提高Act12菌剂在草莓根区土壤的定殖数量。即本研究采用微生物与化学修复措施可促进草莓根区土壤微生态环境向健康协调性方向发展。
Replant disease is the key limiting factor in perennial strawberry (Fragaria x ananassaDuch.) production worldwide. In order to elucidate reasons for the occurrence of strawberryreplant root rot disease, changes of rhizosphere soil chemical properties and microbialcommunity composition between healthy and diseased strawberry plants were determined byroutine soil agro-chemistry analysis and culture-depended method, respectively. Meanwhile, arapid screening program for biological control agents based on the rhizosphere microfloradifferentiae between healthy and diseased plants were established and evaluated. Experimentin greenhouse or under pot conditions were conducted respectively to investigate thealleviating effect of antagonistic microbial agents and silicon fertilizer on the strawberryreplant disease, and the possible alleviating mechanisms were investigated from view point ofdisease suppression, plant growth and nutrition absorption promotion as well as soilmicrobial-ecology restoration. Related results and conclusions are listed as follows：
1. The strawberry root rot disease in monoculture system was associated with thechanges of rhizosphere soil chemical properties and microbial community composition.Comparing to the healthy plant, rhizosphere soil salt content in diseased plants increased witha range of50.6%-56.5%, content of organic matter, available phosphorus and availablepotassium decreased with range of2.2%-27.5%,15.3%-17.6%and10.9%-37.9%,respectively. The ratio of bacteria to fungi (B/F) in rhizoplane soil decreased with a range of2.9%-50.0%, the microbial community tended to a fungi dominant group system. Diversityindex (H) and richness index (S) of bacteria and fungi decreased. The fungi community andstructures in diseased rhizoshpere and rhizoplane soil was significant difference from thehealthy plant soil. Antagonistic fungi Penicillium spp. and Aspergillus sp. dominated inhealthy plant, while soil-borne fungi including Cladosporium sp. and Cylindrocarponmacrodidyma dominated in the root rot strawberry.
2. The strawberry root rot causal agent C. macrodidyma CF9was identificated and twopromising biological control fungi P. griseofulvum CF3and A. terreus CF7were obtainedfrom rhizsphere through a rapid screening program based on the root-zone mycobiotadifferentiae between healthy and diseased strawberry plants. This method is a simple and reliable way to seek for biological control agents against root rot disease.
3. In vitro test showed that potassium silicate inhibited the mycelial growth of fourstrains of soil-borne pathogenic fungi. However, when the pH of potassium silicate amendedPDA was reduced so that it was equivalent to unamended control PDA, there was nodifference in fungal growth between the two treatments. This indicated that the inhibition offungal growth on potassium silicate amended PDA was due to a pH effect.
4. The alleveling effect of biological control agent and silicon fertilizer on the replantdisease of monoculture strawberry.
(1) The pot experiment results showed that two antagonistic fungi isolate P.griseofulvum CF3and A. terreus CF7were effective for the control of strawberry root rotdisease and plant growth promotion and yield enhancement. Root disease incidence ininoculation treatment decreased by52.4%and48.7%, respectively. The root fresh weight andyield increased15.2%,19.6%and26.7%,31.3%respectively. A biological control agentStreptomyces pactum Act12showed significant enhancement on strawberry plant growth andfruit yield in greenhouse. The blooming time of inoculation plant was antedated for6dayscompared with the control. The root fresh weight and fruit yiled also increased by135.5%and47.5%, respectively. The fruit total soluble solids and sugar-to-acid ratio at full fruit stageincreased by11.6%and11.3%.
(2) Individual application with potassium silicate or coal fly ash used as two differentsilicon sources showed better growth and higher yield for replanting strawberry. Applicationwith3.33mmol·kg-1potassium silicate into soil decreased the root rot by36.5%in the thirdyear, antedated blooming for5days, enhanced root fresh weight and fruit yield by31.3%and47.4%. Application with60mg potassium silicate per plant promoted strawberry plant growthin greenhouse. The blooming time was antedated for6days, root fresh weight and fruit yieldincreased with118.0%and102.3%. Application with100g·kg-1fly ash into soil decreased theroot rot by36.2%the third year, antedated blooming for6days, root fresh weight and fruityield was increased by95.5and50.5%.
(3) Combined appilication of silicon fertilizer with Act12agent exhibited synergiesactions both under pot and greenhouse mononculture conditions. Potassium silicate combinedwith Act12agent decreased plant death rate to0%, and antedated blooming for10days ingreenhouse. The plant fresh weight, root fresh weight and fruit yield also increased by73.7%,71.3%and126.2%. Compared with the control, fruit contents of total soluble solid,sugar-to-acid ratio and vitamin C at full fruit stage increased by21.7%,36.2%and27.0%,respectively. Combined appilication of3.33mmol·kg-1potassium silicate with Act12agentinto soil decreased the root rot by58.6%in the third year, antedated blooming for8days, enhanced root fresh weight and fruit yield by73.6%and76.6%. Appilication of100g·kg-1flyash combined with Act12agent into soil decreased the root rot by56.2%in the third year,antedated blooming for6days, enhanced root fresh weight and fruit yield by105.0%and91.0%.
5. The research on mechanism of biological control agents and silicon fertilizer foralleviating replant disease of strawberry
(1) Antagonistic and cell-destroy action. Two antagonistic fungi and S. pactum Act12produced antibiotics against the myceial growth of strawberry soil-borne pathogenic fungi.The antagonistic fungi P. griseofulvum CF3had hyperparasite against root rot cause agent C.macrodidyma CF9.
(2) The Act12agent could stimulate root growth, and promote soil nutrients absorptionand utilization strawberry plant. Combined application of Act12agent with two kind ofsilicon fertilizer enhanced nitrogen, phosphorus, potassium content in strawberry plant.
(3) Strawberry root silicon content improved and root resistance enhancement.Application with silicon fertilizer improved the available silicon in soil, increased the siliconcontent in strawberry root as physical barrier against root rot pathogen.
(4) Root physiological function enhancement and induced host plant resistance. TheAct12agent, potassium silicate and fly ash could contribute to plant root growth anddevelopment, increase root activity and induce the system resistance reaction of strawberryinfected with root rot pathogens by improving plant defense related enzymes activity such asSuperoxide dismutase (SOD), Peroxidase (POD), Polyphenol oxidase (PPO) andPhenylalanine ammonia lyase (PAL).
(5) Modifying rhizosphere microbial community composition. The Act12agent,potassium silicate and fly ash could increase population of soil bacteria and actinomyces, anddecrease soil fungi population, enhance the ratio of bacteria to fungi (B/F) and the ratio ofactinomyces to fungi (A/F), increase the microbial biodiversity. In addition, combinedapplication of potassium silicate with Act12agent encouraged the proliferation of S. pactumAct12. In conclusion, the biological control agents and silicon fertilizer applied were effectivestrategies to engineer the strawberry replant rhizosphere ecology.
1.草莓连作根腐病害发生的根域土壤微生态机制研究。
本研究表明，草莓连作根腐病害的发生与根区土壤化学性质及微生物区系组成及其多样性变化密切相关。
（1）连作草莓病株根区土壤水溶性盐分含量较健株提高了50.6%～156.5%，有机质，速效磷及速效钾含量较健株分别下降22.2%～27.5%，15.3%～17.6%及10.9%～37.9%。
（2）病株根表土壤细菌与真菌数量比值B/F降低了2.9%～50.0%，微生物群落结构趋向“真菌型”发展，细菌、真菌多样性指数H及丰富度指数S呈降低趋势。病健株根区、根表优势真菌种类、数量及比例差异明显，其中健株以具较强拮抗性的2株青霉（Penicillium oxalicum，P. griseofulvum）及1株土曲霉（Aspergillus terreus）为主，而病株则以枝孢霉（Cladosporium sp.）和大双孢柱孢（Cylindrocarpon macrodidyma）等有害病原菌为主。大双孢柱孢C. macrodidyma CF9对草莓根系具有强烈侵染致病作用。
2.草莓根腐病原真菌分离及其拮抗真菌筛选
本研究采用病健株根区及根表土壤微生物区系比较法获得对草莓根系具有强烈侵染致病作用的根腐病原真菌大双孢柱孢C. macrodidyma CF9，同时筛选到2株广谱拮抗真菌灰黄青霉（P. griseofulvum CF3）和土曲霉（A. terreus CF7）。证明此法是快速锁定植物根腐病病原有害微生物及有益拮抗微生物可靠与可行的方法。
3.硅酸钾对病原真菌抑菌作用及其机理研究
硅酸钾对病原真菌菌丝生长具有较强抑制作用，其抑菌机理是通过硅酸钾提高培养基pH实现的，该结果否定了现有研究提出的硅对病原真菌具有类似抗生素作用的观点。
4.微生物及化学防治措施对连作草莓的修复效应研究。
（1）微生物活菌制剂对连作草莓具有明显的防病促生作用。2株拮抗真菌CF3和CF7可使盆栽草莓根腐发病率分别降低52.4%和48.7%，根鲜重分别增加15.2%和19.6%，果实产量分别提高26.7%和31.3%；放线菌Act12活菌制剂可使日光温室草莓根鲜重增加135.5%，开花提前6d，果实产量提高47.5%，同时改善了果实品质，盛果期可溶性固形物和糖酸比分别增加了11.6%和11.3%。
（2）硅酸钾和粉煤灰对连作草莓也有明显的防病促生作用。施用3.33mmol·kg-1硅酸钾对第3年盆栽草莓根腐病的防效为36.5%，开花提前5d，根鲜重和果实产量增加了31.3%和47.4%。日光温室每株施用60mg硅酸钾草莓开花提前6d，根鲜重和果实产量增加了118.0%和102.3%。盆栽施用100g·kg-1粉煤灰对第3年草莓根腐病的防效为36.2%，开花提前6d，根鲜重和果实产量增加了95.5%和50.5%。。
（3）硅酸钾及粉煤灰与菌剂配施除具有三者单独施用的效果外，在部分处理中，硅肥与放线菌剂配施具有增效作用。硅酸钾与菌剂配施可使日光温室草莓植株死亡率降低至0%，开花提前10d，植株总鲜重，根鲜重及单株产量分别提高73.7%，71.3%及126.2%，同时盛果期果实可溶性固形物含量，糖酸比及维生素C含量分别提高21.7%，36.2%及27.0%。盆栽试验表明，3.33mmol·kg-1硅酸钾与菌剂配施对第3年草莓根腐病的防效为58.6%，开花提前8d，根鲜重及单株产量分别提高73.6%和76.6%；100g·kg-1粉煤灰与菌剂配施对第3年草莓根腐病的防效为56.2%，开花提前6d，根鲜重及单株产量分别提高105.0%和91.0%。
5.微生物及化学修复措施对草莓连作障碍的修复机理研究。
（1）抗生及溶菌作用。2株拮抗真菌及放线菌Act12均可产生抗菌活性产物，通过抗菌活性代谢产物抑制草莓连作土传病原真菌菌丝生长，其中CF3对大双孢柱孢CF9菌丝具有重寄生和溶菌作用。
（2）放线菌制剂能刺激草莓根系生长，促进植株对土壤养分的吸收利用。Act12菌剂单施或与2种硅肥配施均可促进连作草莓对氮、磷、钾元素吸收。
（3）提高根系硅元素含量，增强根系抗病性。施用硅肥增加了土壤有效硅含量，促进草莓植株根系硅素的吸收积累，增加了根系含硅量，增强了草莓对根腐病的抗病性。
（4）改善根系生理功能，增强植物的诱导抗性。Act12菌剂、硅酸钾及粉煤灰均可影响草莓根系生长发育，增强在根腐病原真菌胁迫下草莓的根系活力；诱导草莓产生抗病性，提高草莓叶片超氧化物歧化酶活性（SOD）、过氧化物酶活性（POD）、多酚氧化酶活性（PPO）及苯丙氨酸解氨酶（PAL）主要防御酶活性。
（5）恢复草莓根区、根表土壤微生物区系平衡。Act12菌剂、硅酸钾及粉煤灰均可增加土壤细菌、放线菌数量，减少土壤真菌数量，同时提高B/F和A/F值，提高土壤细菌、真菌及放线菌的多样性；硅肥与菌剂配施可有效提高Act12菌剂在草莓根区土壤的定殖数量。即本研究采用微生物与化学修复措施可促进草莓根区土壤微生态环境向健康协调性方向发展。
Replant disease is the key limiting factor in perennial strawberry (Fragaria x ananassaDuch.) production worldwide. In order to elucidate reasons for the occurrence of strawberryreplant root rot disease, changes of rhizosphere soil chemical properties and microbialcommunity composition between healthy and diseased strawberry plants were determined byroutine soil agro-chemistry analysis and culture-depended method, respectively. Meanwhile, arapid screening program for biological control agents based on the rhizosphere microfloradifferentiae between healthy and diseased plants were established and evaluated. Experimentin greenhouse or under pot conditions were conducted respectively to investigate thealleviating effect of antagonistic microbial agents and silicon fertilizer on the strawberryreplant disease, and the possible alleviating mechanisms were investigated from view point ofdisease suppression, plant growth and nutrition absorption promotion as well as soilmicrobial-ecology restoration. Related results and conclusions are listed as follows：
1. The strawberry root rot disease in monoculture system was associated with thechanges of rhizosphere soil chemical properties and microbial community composition.Comparing to the healthy plant, rhizosphere soil salt content in diseased plants increased witha range of50.6%-56.5%, content of organic matter, available phosphorus and availablepotassium decreased with range of2.2%-27.5%,15.3%-17.6%and10.9%-37.9%,respectively. The ratio of bacteria to fungi (B/F) in rhizoplane soil decreased with a range of2.9%-50.0%, the microbial community tended to a fungi dominant group system. Diversityindex (H) and richness index (S) of bacteria and fungi decreased. The fungi community andstructures in diseased rhizoshpere and rhizoplane soil was significant difference from thehealthy plant soil. Antagonistic fungi Penicillium spp. and Aspergillus sp. dominated inhealthy plant, while soil-borne fungi including Cladosporium sp. and Cylindrocarponmacrodidyma dominated in the root rot strawberry.
2. The strawberry root rot causal agent C. macrodidyma CF9was identificated and twopromising biological control fungi P. griseofulvum CF3and A. terreus CF7were obtainedfrom rhizsphere through a rapid screening program based on the root-zone mycobiotadifferentiae between healthy and diseased strawberry plants. This method is a simple and reliable way to seek for biological control agents against root rot disease.
3. In vitro test showed that potassium silicate inhibited the mycelial growth of fourstrains of soil-borne pathogenic fungi. However, when the pH of potassium silicate amendedPDA was reduced so that it was equivalent to unamended control PDA, there was nodifference in fungal growth between the two treatments. This indicated that the inhibition offungal growth on potassium silicate amended PDA was due to a pH effect.
4. The alleveling effect of biological control agent and silicon fertilizer on the replantdisease of monoculture strawberry.
(1) The pot experiment results showed that two antagonistic fungi isolate P.griseofulvum CF3and A. terreus CF7were effective for the control of strawberry root rotdisease and plant growth promotion and yield enhancement. Root disease incidence ininoculation treatment decreased by52.4%and48.7%, respectively. The root fresh weight andyield increased15.2%,19.6%and26.7%,31.3%respectively. A biological control agentStreptomyces pactum Act12showed significant enhancement on strawberry plant growth andfruit yield in greenhouse. The blooming time of inoculation plant was antedated for6dayscompared with the control. The root fresh weight and fruit yiled also increased by135.5%and47.5%, respectively. The fruit total soluble solids and sugar-to-acid ratio at full fruit stageincreased by11.6%and11.3%.
(2) Individual application with potassium silicate or coal fly ash used as two differentsilicon sources showed better growth and higher yield for replanting strawberry. Applicationwith3.33mmol·kg-1potassium silicate into soil decreased the root rot by36.5%in the thirdyear, antedated blooming for5days, enhanced root fresh weight and fruit yield by31.3%and47.4%. Application with60mg potassium silicate per plant promoted strawberry plant growthin greenhouse. The blooming time was antedated for6days, root fresh weight and fruit yieldincreased with118.0%and102.3%. Application with100g·kg-1fly ash into soil decreased theroot rot by36.2%the third year, antedated blooming for6days, root fresh weight and fruityield was increased by95.5and50.5%.
(3) Combined appilication of silicon fertilizer with Act12agent exhibited synergiesactions both under pot and greenhouse mononculture conditions. Potassium silicate combinedwith Act12agent decreased plant death rate to0%, and antedated blooming for10days ingreenhouse. The plant fresh weight, root fresh weight and fruit yield also increased by73.7%,71.3%and126.2%. Compared with the control, fruit contents of total soluble solid,sugar-to-acid ratio and vitamin C at full fruit stage increased by21.7%,36.2%and27.0%,respectively. Combined appilication of3.33mmol·kg-1potassium silicate with Act12agentinto soil decreased the root rot by58.6%in the third year, antedated blooming for8days, enhanced root fresh weight and fruit yield by73.6%and76.6%. Appilication of100g·kg-1flyash combined with Act12agent into soil decreased the root rot by56.2%in the third year,antedated blooming for6days, enhanced root fresh weight and fruit yield by105.0%and91.0%.
5. The research on mechanism of biological control agents and silicon fertilizer foralleviating replant disease of strawberry
(1) Antagonistic and cell-destroy action. Two antagonistic fungi and S. pactum Act12produced antibiotics against the myceial growth of strawberry soil-borne pathogenic fungi.The antagonistic fungi P. griseofulvum CF3had hyperparasite against root rot cause agent C.macrodidyma CF9.
(2) The Act12agent could stimulate root growth, and promote soil nutrients absorptionand utilization strawberry plant. Combined application of Act12agent with two kind ofsilicon fertilizer enhanced nitrogen, phosphorus, potassium content in strawberry plant.
(3) Strawberry root silicon content improved and root resistance enhancement.Application with silicon fertilizer improved the available silicon in soil, increased the siliconcontent in strawberry root as physical barrier against root rot pathogen.
(4) Root physiological function enhancement and induced host plant resistance. TheAct12agent, potassium silicate and fly ash could contribute to plant root growth anddevelopment, increase root activity and induce the system resistance reaction of strawberryinfected with root rot pathogens by improving plant defense related enzymes activity such asSuperoxide dismutase (SOD), Peroxidase (POD), Polyphenol oxidase (PPO) andPhenylalanine ammonia lyase (PAL).
(5) Modifying rhizosphere microbial community composition. The Act12agent,potassium silicate and fly ash could increase population of soil bacteria and actinomyces, anddecrease soil fungi population, enhance the ratio of bacteria to fungi (B/F) and the ratio ofactinomyces to fungi (A/F), increase the microbial biodiversity. In addition, combinedapplication of potassium silicate with Act12agent encouraged the proliferation of S. pactumAct12. In conclusion, the biological control agents and silicon fertilizer applied were effectivestrategies to engineer the strawberry replant rhizosphere ecology.
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