有机、无公害和常规蔬菜种植模式下温室土壤生物群落结构及食物网的特征研究
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摘要
日光温室是中国北方的重要的农业土地利用类型之一,相比自然生境及室外农田生态系统具有半封闭、高投入和高扰动的特性。常规种植模式在带来高产的同时也引发了一系列环境问题,有机和无公害作为环境友好型农业模式改善了土壤环境,同时也会对土壤生物群落及地下食物网产生影响。本试验以中国农业大学曲周蔬菜日光温室长期定位试验为研究对象,对比了有机(ORG)、无公害(LOW)和常规(CON)种植模式下土壤生物生物量、群落结构、多样性及食物网特征。主要结果如下：
温室土壤以细菌降解途径占优势；原生动物中鞭毛虫占绝对优势。管理模式对土壤真菌、细菌和微生物量总碳、原生动物各类群丰度及总数均有显著影响,但对真菌/细菌比率、鞭毛虫和肉足虫的相对丰度没有显著影响。细菌、真菌和微生物量碳在不同温室间总体呈现相同的规律,即ORG>LOW>CON;对于原生动物,有机模式下的鞭毛虫、肉足虫和原生动物总数量高于无公害和常规模式的,而后两者的数量较为接近。
食细菌线虫是温室环境中最为丰富的线虫营养类群,其所占比例超过80%。其次是杂食捕食性线虫。有机模式同无公害和常规模式相比提高了线虫总数、食细菌线虫、食真菌线虫和杂食捕食性线虫的丰度。植食性线虫在无公害和有机温室中得到了抑制,且此效应在有机模式下更为明显。而种植模式对营养类群的组成和线虫区系分析的影响较小,三个温室均显示出富集的土壤养分和结构化的食物网状态。香浓指数(H')和优势度指数(λ)反映出大量的有机肥投入降低了线虫多样性,增加了优势度,尤其对于典型机会主义者。
种植模式对土壤螨的总数、个亚目及菌食性隐气门螨和捕食性螨比例均无显著影响,仅对菌食性非隐气门螨和食线虫螨的比例有显著影响。土壤螨的总数及各营养类群的数量呈现不同程度的波动状态,且在不同的温室规律存在差异。粉螨是温室土壤中最为丰富的螨类(平均37.8%)。种植模式对螨的香浓指数(H')和优势度指数(λ)均无显著影响。
有机模式下土壤食物网结构优于无公害和常规模式的,主要表现在：有机温室具有最高的功能群数目、连通度,食物链长度及食物网多样性。总生物量在不同温室间呈现ORG>LOW>CON的规律。大部分功能群的生物量在有机模式下最高,而无公害和常规模式总体处于同一水平。有机模式对高营养级生物量的促进作用更明显。功能群相互作用强度分析显示最高的取食强度和食物调控强度均出现在第2营养级功能群,在0-10cm土层,不同模式间各营养级功能群及总的取食强度和食物网调控强度均表现为ORG>LOW>CON,说明有机模式和无公害模式均不同程度的增强了食物网的上行效应和下行效应。能量流动途径主要发生在微生物对碎屑的取食上,最大值为细菌对碎屑的取食。相比无公害和常规模式,有机模式具有更高的碳矿化率和土壤表层的氮矿化率。土壤动物对碳矿化率的贡献为11.1%,对氮矿化率的贡献平均达48.3%,其中食细菌线虫的贡献最大。有机模式相比另两种模式提高了土壤动物对碳氮矿化率的贡献。
对于微生物和小型土壤动物,种植模式对生物量的影响大于对群落结构(或功能群结构)的影响。而对于中型土壤动物(微节肢动物),二者在不同温室间都没有明显的差别。有机模式在多样性指数上没有表现出优势,尽管线虫和螨在该模式下均发现了更多的分类单位；而在基于功能群水平上的多样性在有机温室高于其他两个温室。
总体而言,有机农业促进了地下食物网结构的优化和碳、氮矿化功能以及生物量。其中生物量的优势可进一步体现在生态系统功能上,即养分矿化。
Solar greenhouse is a typical intensive production model in north China. Compared with natural habitat and opened farmland system, the greenhouse is often in a semiclosed state with high input and high disturbance. Conventional management practices bring in high yield but induced serious environmental pollution. Low input and organic management practices improved soil condition, and affected soil organism and food web. Based on the Quzhou Experimental Station of China Agricultural University, A comparative study of organic (ORG), low input (LOW) and conventional (CON) vegetable greenhouse systems was conducted to assess the effect of management practices on the biomass of soil organism, community structure, diversity and food web characteristic. The mean results list below:
All three greenhouse systems showed a bacterial-based decomposition pathway and flagellate was the most popular protozoa group. The soil fungal biomass carbon, bacterial biomass carbon, microbial biomass carbon, flagellate abundance, ciliate abundance, amoeba abundance and total protozoa abundance were significantly affected by management practices, while no significant effect was found on the fungal to bacterial ratio (FB) and the relative abundance of flagellate and amoeba. Generally, all soil bacterial, fungal and microbial biomass carbon among three systems presented a decreasing sequence of ORG>LOW>CON. The abundances of flagellate, amoeba and total protozoa under organic management practices were higher than that under low input and conventional management practices. The abundances of the above protozoa were closed in low input and conventional systems.
Bacterivores were the most dominant trophic group in all three systems with a mean proportion of over80%, followed by omnivore-carnivores. In general, organic management practices increased the abundance of total nematodes, bacterivores, fungivores and omnivore-carnivores in comparison with low input and conventional management practices. Though inhibitory effects of plant feeders were found in organic and low input systems, these effects were more evident in organic systems. However, small differences were observed in the composition of trophic groups and fauna analysis. All three systems displayed enriched soil conditions and structured food webs. The Shannon index (H') and genus dominance (λ) suggested that in greenhouse conditions, excessive manure input would cause a decrease in nematode diversity but increase the dominance, particularly for enrichment opportunists.
No significant management practices effect was found on the abundance of soil mites, the relative abundance of various suborder and the relative abundance of fungivorous cryptostigmatic and predaceous mites. Significant management practices effects were only found on the relative abundance of fungivorous non-cryptostigmatic and nematophagous mites. The abundance of total mites and various trophic groups fluctuated at different degrees during the sampling times, and they differed in different greenhouses. Acaridae was the most abundant family (meanly38.7%). No significant management practices effect was found on the Shannon index (H1) and dominance (λ) indices.
The soil food web structure under organic management practices was better than that under low input and conventional management practices. Higher values of the number of functional groups, connectance, food chain length and food web diversity were obtained in organic greenhouse than in the other two greenhouses. The total biomass among three systems presented a decreasing sequence of ORG>LOW>CON. The biomass of most functional groups were higher in organic system compared with the other two systems, which exhibited similar levels to each other. Organic management practices was more effective in promoting the biomass of high trophic level. The analysis of interaction strengths of food webs showed that both the effect on prey and the effect on predator of various trophic levels were appeared at trophic level2. At0-10cm soil depth, the prey effects and food regulation effects of total organism and various trophic levels among three systems presented a decreasing sequence of ORG>LOW>CON, indicating an increased effect of both bottom-up and top-down effects in organic system than in low input and conventional systems. The energy flow pathway was mainly occurred at the process of microbe feeding on detritus. Higher C and N (only in the0-10cm depth) mineralization rates were found in organic greenhouse than in low input and conventional greenhouses. The contributions of soil fauna to C and N mineralization rates were11.1%and48.3%, respectively, and largest contribution was observed on bacterivore nematodes. Organic management practices improved the contributions of soil fauna to C and N mineralization rates compared with the other two management practices.
Management practices were more influential on microorganism biomass than community structure (or functional groups structure). For mesofauna (micro-arthropodes), no evident difference was found on biomass and community structure among three greenhouses. Organic management practices did not show advantage on diversity index, although more taxon were found in this greenhouse. However, food web diversity, which the diversity is based on the functional groups level, presented a higher value than that in the other two greenhouses.
We concluded that organic agriculture promoted the optimization of belowground food web structure, and improved the soil organism biomass and function of C and N mineralization. The advantage of biomass can be further reflected on the ecosystem function, such as nutrient mineralization.
温室土壤以细菌降解途径占优势；原生动物中鞭毛虫占绝对优势。管理模式对土壤真菌、细菌和微生物量总碳、原生动物各类群丰度及总数均有显著影响,但对真菌/细菌比率、鞭毛虫和肉足虫的相对丰度没有显著影响。细菌、真菌和微生物量碳在不同温室间总体呈现相同的规律,即ORG>LOW>CON;对于原生动物,有机模式下的鞭毛虫、肉足虫和原生动物总数量高于无公害和常规模式的,而后两者的数量较为接近。
食细菌线虫是温室环境中最为丰富的线虫营养类群,其所占比例超过80%。其次是杂食捕食性线虫。有机模式同无公害和常规模式相比提高了线虫总数、食细菌线虫、食真菌线虫和杂食捕食性线虫的丰度。植食性线虫在无公害和有机温室中得到了抑制,且此效应在有机模式下更为明显。而种植模式对营养类群的组成和线虫区系分析的影响较小,三个温室均显示出富集的土壤养分和结构化的食物网状态。香浓指数(H')和优势度指数(λ)反映出大量的有机肥投入降低了线虫多样性,增加了优势度,尤其对于典型机会主义者。
种植模式对土壤螨的总数、个亚目及菌食性隐气门螨和捕食性螨比例均无显著影响,仅对菌食性非隐气门螨和食线虫螨的比例有显著影响。土壤螨的总数及各营养类群的数量呈现不同程度的波动状态,且在不同的温室规律存在差异。粉螨是温室土壤中最为丰富的螨类(平均37.8%)。种植模式对螨的香浓指数(H')和优势度指数(λ)均无显著影响。
有机模式下土壤食物网结构优于无公害和常规模式的,主要表现在：有机温室具有最高的功能群数目、连通度,食物链长度及食物网多样性。总生物量在不同温室间呈现ORG>LOW>CON的规律。大部分功能群的生物量在有机模式下最高,而无公害和常规模式总体处于同一水平。有机模式对高营养级生物量的促进作用更明显。功能群相互作用强度分析显示最高的取食强度和食物调控强度均出现在第2营养级功能群,在0-10cm土层,不同模式间各营养级功能群及总的取食强度和食物网调控强度均表现为ORG>LOW>CON,说明有机模式和无公害模式均不同程度的增强了食物网的上行效应和下行效应。能量流动途径主要发生在微生物对碎屑的取食上,最大值为细菌对碎屑的取食。相比无公害和常规模式,有机模式具有更高的碳矿化率和土壤表层的氮矿化率。土壤动物对碳矿化率的贡献为11.1%,对氮矿化率的贡献平均达48.3%,其中食细菌线虫的贡献最大。有机模式相比另两种模式提高了土壤动物对碳氮矿化率的贡献。
对于微生物和小型土壤动物,种植模式对生物量的影响大于对群落结构(或功能群结构)的影响。而对于中型土壤动物(微节肢动物),二者在不同温室间都没有明显的差别。有机模式在多样性指数上没有表现出优势,尽管线虫和螨在该模式下均发现了更多的分类单位；而在基于功能群水平上的多样性在有机温室高于其他两个温室。
总体而言,有机农业促进了地下食物网结构的优化和碳、氮矿化功能以及生物量。其中生物量的优势可进一步体现在生态系统功能上,即养分矿化。
Solar greenhouse is a typical intensive production model in north China. Compared with natural habitat and opened farmland system, the greenhouse is often in a semiclosed state with high input and high disturbance. Conventional management practices bring in high yield but induced serious environmental pollution. Low input and organic management practices improved soil condition, and affected soil organism and food web. Based on the Quzhou Experimental Station of China Agricultural University, A comparative study of organic (ORG), low input (LOW) and conventional (CON) vegetable greenhouse systems was conducted to assess the effect of management practices on the biomass of soil organism, community structure, diversity and food web characteristic. The mean results list below:
All three greenhouse systems showed a bacterial-based decomposition pathway and flagellate was the most popular protozoa group. The soil fungal biomass carbon, bacterial biomass carbon, microbial biomass carbon, flagellate abundance, ciliate abundance, amoeba abundance and total protozoa abundance were significantly affected by management practices, while no significant effect was found on the fungal to bacterial ratio (FB) and the relative abundance of flagellate and amoeba. Generally, all soil bacterial, fungal and microbial biomass carbon among three systems presented a decreasing sequence of ORG>LOW>CON. The abundances of flagellate, amoeba and total protozoa under organic management practices were higher than that under low input and conventional management practices. The abundances of the above protozoa were closed in low input and conventional systems.
Bacterivores were the most dominant trophic group in all three systems with a mean proportion of over80%, followed by omnivore-carnivores. In general, organic management practices increased the abundance of total nematodes, bacterivores, fungivores and omnivore-carnivores in comparison with low input and conventional management practices. Though inhibitory effects of plant feeders were found in organic and low input systems, these effects were more evident in organic systems. However, small differences were observed in the composition of trophic groups and fauna analysis. All three systems displayed enriched soil conditions and structured food webs. The Shannon index (H') and genus dominance (λ) suggested that in greenhouse conditions, excessive manure input would cause a decrease in nematode diversity but increase the dominance, particularly for enrichment opportunists.
No significant management practices effect was found on the abundance of soil mites, the relative abundance of various suborder and the relative abundance of fungivorous cryptostigmatic and predaceous mites. Significant management practices effects were only found on the relative abundance of fungivorous non-cryptostigmatic and nematophagous mites. The abundance of total mites and various trophic groups fluctuated at different degrees during the sampling times, and they differed in different greenhouses. Acaridae was the most abundant family (meanly38.7%). No significant management practices effect was found on the Shannon index (H1) and dominance (λ) indices.
The soil food web structure under organic management practices was better than that under low input and conventional management practices. Higher values of the number of functional groups, connectance, food chain length and food web diversity were obtained in organic greenhouse than in the other two greenhouses. The total biomass among three systems presented a decreasing sequence of ORG>LOW>CON. The biomass of most functional groups were higher in organic system compared with the other two systems, which exhibited similar levels to each other. Organic management practices was more effective in promoting the biomass of high trophic level. The analysis of interaction strengths of food webs showed that both the effect on prey and the effect on predator of various trophic levels were appeared at trophic level2. At0-10cm soil depth, the prey effects and food regulation effects of total organism and various trophic levels among three systems presented a decreasing sequence of ORG>LOW>CON, indicating an increased effect of both bottom-up and top-down effects in organic system than in low input and conventional systems. The energy flow pathway was mainly occurred at the process of microbe feeding on detritus. Higher C and N (only in the0-10cm depth) mineralization rates were found in organic greenhouse than in low input and conventional greenhouses. The contributions of soil fauna to C and N mineralization rates were11.1%and48.3%, respectively, and largest contribution was observed on bacterivore nematodes. Organic management practices improved the contributions of soil fauna to C and N mineralization rates compared with the other two management practices.
Management practices were more influential on microorganism biomass than community structure (or functional groups structure). For mesofauna (micro-arthropodes), no evident difference was found on biomass and community structure among three greenhouses. Organic management practices did not show advantage on diversity index, although more taxon were found in this greenhouse. However, food web diversity, which the diversity is based on the functional groups level, presented a higher value than that in the other two greenhouses.
We concluded that organic agriculture promoted the optimization of belowground food web structure, and improved the soil organism biomass and function of C and N mineralization. The advantage of biomass can be further reflected on the ecosystem function, such as nutrient mineralization.
引文
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