不同食细菌线虫取食密度下线虫对细菌数量、活性及土壤氮素矿化的影响

采用悉生培养微缩体系,探讨了不同食细菌线虫取食密度下线虫(Caenorhabditis elegans) 对细菌(Bacillus subtilis)数量和活性及土壤氮素矿化的影响.结果表明,线虫对细菌的取食,促进了细菌的增殖,并在不同线虫取食密度下对细菌的增殖促进作用总体表现为:接种20条·g^-1＞10条·g^-1＞40条线虫·g^-1处理.线虫在促进细菌增殖的同时,明显提高了土壤呼吸强度和土壤蔗糖酶、脲酶和磷酸酶的活性,但不同取食密度处理间差异不明显.线虫与细菌之间的相互作用显著提高了土壤铵态氮和矿质态氮含量,促进了土壤氮的矿化.不同取食密度处理间,线虫对土壤氮素矿化的促进作用与对细菌的增殖促进作用趋势一致.     

不同土壤水分条件下华美新小杆线虫对枯草芽孢杆菌数量、活性及土壤氮素矿化的影响

采用悉生培养微缩体系,探讨了不同土壤含水量条件下,华美新小杆线虫(Caenorhabditis elegans)对枯草芽孢杆菌(Bacillus subtilis)数量和活性及土壤氮素矿化的影响.结果表明：在试验设置的不同含水量条件下,线虫对细菌的取食活动均促进了细菌的增殖 ,并明显提高了土壤呼吸强度; 不同含水量条件对细菌的增殖促进作用总体表现为23%含水量处理>17%含水量处理>28%含水量处理.线虫与细菌的相互作用显著提高了土壤铵态氮和矿质氮含量,促进了土壤氮的矿化.接种线虫对土壤氮素矿化的促进作用表现为23%含水量处理的矿质氮含量显著高于其它两个含水量处理.     

CARD-FISH研究食细菌线虫对氨氧化细菌(AOB)数量的影响

土壤动物与微生物的取食与反馈之间的关系是土壤生态学研究的核心内容之一。通过接种原位的食细菌线虫和微生物群落模拟土壤真实环境,采用CARD-FISH方法来观察食细菌线虫的不同取食密度下,氨氧化细菌(ammonia oxidizing bacteria)数量的动态变化,以揭示土壤食细菌线虫对AOB数量的影响及AOB的反馈强度。结果表明:与单独接种细菌的处理(SB)相比,接种食细菌线虫显著地增加了土壤中AOB的数量,3个不同线虫接种密度处理中AOB数量表现为接种20条g-1干土的处理(SBN20)接种10条g-1干土的处理(SBN10)接种40条g-1干土的处理(SBN40)。由于过度取食,SBN40处理中AOB的数量在培养了14d后低于SB处理,且在第28天时显著低于SB处理。接种食细菌线虫显著增加了土壤中NH4+-N和NO3-N的含量,表明食细菌线虫促进了N的矿化和硝化作用。矿化作用增强使得硝化作用的底物NH4+-N显著增加可能是AOB数量显著增多的重要原因之一。     

土壤食细菌线虫的原位富集培养方法

采用两种孔径的尼龙网袋（1mm和5μm），将盆钵供试土壤分成内外两层，内层土壤混合猪粪或稻草，并以不添加猪粪和稻草的供试土壤作为空白；外层直接接入供试土壤，进行培养，以获取土著食细菌线虫大量富集的试验土壤。结果表明：添加基质（猪粪和稻草）显著地促进了土壤线虫的繁殖，大量繁殖的线虫通过1mm网袋迁移至外层未加基质的土壤，而采用5μm网袋则限制了线虫向外层土壤的迁移。添加猪粪的1mm网袋处理经过28d培养后，外层土壤线虫数是空白处理的9．1倍；添加稻草的1mm网袋处理经过35d培养后，外层土壤线虫数是空白处理的5．9倍。添加两种基质的5μm网袋处理，外层土壤线虫数和空白处理差异不大。添加基质主要是促进了食细菌线虫的繁殖，在培养结束时，添加猪粪的1mm网袋处理的食细菌线虫比例达到98．2％，添加稻草的1mm网袋处理的食细菌线虫比例达到90．5％，两个处理食细菌线虫的总数分别是空白处理的14．8倍和8．9倍，并且主要是Protorhabditis sp．线虫的增加。     

悉生培养微缩体系食细菌线虫提高土壤激素含量的机制

研究土壤食细菌线虫与细菌的相互作用及其生态功能是土壤生态学的核心内容之一。食细菌线虫取食细菌可以促进土壤中氮素的矿化,提高氮素养分的供给,改善土壤的营养条件,从而促进植物的生长发育。土壤食细菌线虫促进植物根系生长的"养分作用机制"已得到确认,而"激素作用机制"还存在争议。从供试土壤中筛选获得一株高效产IAA细菌和两种不同cp值的食细菌线虫,通过设置简化的悉生培养系统,对这两种土著食细菌线虫与土著产IAA细菌之间的相互作用,及其对土壤中IAA含量变化的影响进行研究。结果表明:两种食细菌线虫的取食均能促进细菌数量和活性的增强,食细菌线虫与产IAA细菌相互作用也能显著增加土壤中IAA的含量;这些促进作用受到接种食细菌线虫的种类以及培养时间的影响:在培养第10天和第20天时,接种cp值为1的中杆属食细菌线虫显著增加了产IAA细菌的数量;在培养第10天和第30天时,相比较接种cp值为2的头叶属食细菌线虫,接种中杆属食细菌线虫显著提高了土壤中IAA的含量。     

食真菌线虫对热或铜胁迫下土壤生态功能稳定性的影响

在模拟胁迫条件下(施加 CuSO₄ 的持续胁迫或加热40 ℃的瞬时胁迫),以大麦叶粉短期分解过程代表土壤功能,采用室内培养试验研究了土壤食真菌线虫(Aphelenchus avenae)与微生物的相互作用对土壤生态功能稳定性(抗性和恢复力)的影响.结果表明：无论施加胁迫与否,食真菌线虫的活动都有促进土壤微生物活性的趋势,尤其是在施加铜胁迫后第8 天开始到培养期结束,接种食真菌线虫导致土壤基础呼吸显著增加(P<0.05),但加热胁迫后食真菌线虫对土壤基础呼吸的促进作用仅在第8天有显著差异(P<0.05),反映了食真菌线虫对土壤微生物活性的影响程度与胁迫类型有关.在两种胁迫条件下,接种食真菌线虫对土壤功能的抗性没有影响,但都能促进胁迫条件下土壤功能的恢复.培养后期,两种胁迫条件下接种食真菌线虫处理真菌生物量低于未接种线虫处理,表明胁迫条件下食真菌线虫对真菌的取食可能限制甚至抑制了真菌生长,导致真菌对细菌的竞争压力减少,从而使细菌获得更大的生长优势,间接促进了细菌的生长.     

草莓连作土壤酚酸类物质积累对土壤线虫的影响

通过大田试验和盆栽试验相结合的方式,对不同连作年限草莓土壤酚酸类物质含量、土壤线虫数量和食细菌线虫数量以及外源酚酸类物质对土壤线虫数量和食细菌线虫数量的影响进行了研究。结果表明:随连作年限的增加,草莓连作土壤中酚酸类物质含量明显增加,土壤线虫总数和食细菌线虫总数整体呈现下降趋势,且连作5年的土壤线虫总数和食细菌线虫数量最低,酚酸类物质含量与土壤线虫总数和食细菌线虫数量呈负相关;对羟基苯甲酸和对香豆酸在浓度低于200μg·g-1时使土壤线虫总数和食细菌线虫数量增加,而高于200μg·g-1时起抑制作用;肉桂酸在浓度低于100μg·g-1时增加了土壤线虫总数和食细菌线虫数量,而高于100μg·g-1时起抑制作用;随阿魏酸浓度的增加,线虫数量变化规律不明显,当其浓度高于100μg·g-1时起抑制作用;混合酚酸随浓度增加对土壤线虫总数和食细菌线虫数量的抑制作用增强。     

食微线虫对植物生长及土壤养分循环的影响

近二十多年来, 土壤动物的生态功能受到广泛重视。越来越多的证据表明, 土壤动物和微生物间的相互作用对土壤生态系统过程和植物生长起着重要的调节作用。本文综述了食细菌线虫和食真菌线虫对土壤微生物、土壤氮矿化和植物生长的影响。大量研究发现, 食细菌线虫和食真菌线虫都有助于土壤氮素等养分矿化, 从而促进植物生长。这种作用主要是线虫通过取食活动加速微生物周转, 并通过代谢分泌和释放微生物所固持的养分而实现的。但这种作用会因不同的线虫、微生物和植物的种类以及土壤基质的C/N营养状况而异, 此外还受线虫的营养类群及其与其他土壤动物之间复杂关系的影响。今后应该加强以下几方面的研究: (1)深入研究线虫、微生物和植物之间相互作用的机制; (2) 增加控制实验系统的复杂性, 研究线虫不同功能群之间及其与其他土壤动物之间的关系; (3)加强长期实验和观察, 在较长的时间尺度上了解线虫的生态功能; (4)加强对不同生态系统的研究, 在更大的空间尺度上综合了解土壤线虫的生态功能; (5)在全球气候变化的背景下了解土壤线虫的响应, 并预测土壤线虫对全球变化的反馈。     

不同水分管理方式对稻田土壤生物学特性的影响

在下辽河平原单季稻地区研究了常规浅湿干灌溉 (CK)、浅湿干灌溉薄膜阻渗 (IC)、湿润灌溉薄膜阻渗 (MC)、淹水灌溉薄膜阻渗 (FC) 4种不同水分管理方式下土壤线虫及土壤微生物量的动态变化。结果表明 ,耙耕前 ,CK、FC处理食细菌线虫数量显著高于MC、IC处理 ;薄膜阻渗在黄熟期显著降低了土壤食细菌线虫的数量 ,在耙耕前显著降低了食真菌线虫数量。潮棕壤稻田食真菌线虫与食细菌线虫相比数量较低。在耙耕前不同水分管理方式下土壤微生物量C显著低于对照。不同水分管理方式在水稻分蘖期、抽穗期对食细菌线虫数量、食真菌线虫数量、微生物量C和微生物量N没有影响。土壤食细菌线虫、食真菌线虫数量与土壤微生物量C、N没有达到显著相关。     

食细菌线虫对热或铜胁迫下土壤功能稳定性的影响

在室内模拟重金属铜的持续胁迫或者短期加热(18 h,40℃)的瞬时胁迫条件下,以大麦叶粉短期分解过程代表土壤功能,研究了土壤食细菌线虫对土壤生态功能稳定性(抗性和恢复力)的影响.结果表明,在未施加任何胁迫对照处理中和铜胁迫条件下,食细菌线虫在一定程度上有促进土壤微生物活性的趋势.在施加铜胁迫后第15天,接种食细菌线虫导致土壤基础呼吸显著增加,并且接种食细菌线虫处理的土壤功能抗性显著高于未接种线虫处理.而在热胁迫条件下,接种食细菌线虫对土壤基础呼吸和土壤微生物活性没有显著影响,且接种食细菌线虫处理的土壤功能抗性低于未接种线虫处理.在两种胁迫条件下,接种食细菌线虫反而降低了土壤功能的恢复力.     

Diversity of bacteria associated with grassland soil nematodes of different feeding groups

The bacterial diversity associated with soil nematodes and its relationship with their feeding habits are as yet poorly understood. In the present study the diversity and abundance of bacteria from nematodes and their surrounding soil were analysed and compared. The nematodes were collected from a grassland soil and sorted into bacterial, fungal, plant, predatory and omnivore feeding groups and assigned to taxonomic groups. Total DNA was extracted from the nematodes and partial bacterial 16S rRNA genes were PCR amplified, cloned and sequenced. The abundance and composition of bacterial taxa differed between and within feeding groups. The lowest bacterial diversity was found in the predatory nematodes Prionchulus sp., whereas the highest bacterial diversity was associated with the bacterial-feeding nematode Acrobeles sp. The soil had a more diverse bacterial community than the communities found in the nematode groups. The 16S rRNA gene sequences of bacteria associated with nematodes did not overlap with those detected in soil as determined using the cloning screening approach. However, bacterial sequences identified from nematodes could be detected in the soil with targeted PCR. Our data suggest that the nematodes do not feed on the most abundant bacteria present in soil. Furthermore, several nematodes contained suspected bacterial symbionts and parasites.     

线虫作为土壤健康指示生物的方法及应用

简述了线虫作为土壤健康指示生物的优势,归纳了常用指数和分析方法,对成熟指数及基于营养类群的一些指数和分析方法进行了详细介绍.与其他指数相比,成熟指数更能敏感地反映土壤环境的受胁迫程度,基于营养类群的指数和分析方法可以在生态系统功能水平上更好地揭示土壤环境的健康状态.概述了以上指数和方法在农业、森林、草原及其他生态系统中的应用研究.结合目前存在的一些问题,提出以下建议:加强对土壤线虫生活史和食性等特性的认识;进一步完善现有指数,将多种指数和分析方法结合使用;加强土壤线虫在多种生态系统及大尺度地域空间内的应用.     

Effects of native and exotic range‐expanding plant species on taxonomic and functional composition of nematodes in the soil food web

Due to climate warming, many plant species shift ranges towards higher latitudes. Plants can disperse faster than most soil biota, however, little is known about how range‐expanding plants in the new range will establish interactions with the resident soil food web. In this paper we examine how the soil nematode community from the new range responds to range‐expanding plant species compared to related natives. We focused on nematodes, because they are important components in various trophic levels of the soil food web, some feeding on plant roots, others on microbes or on invertebrates. We expected that range expanding plant species have fewer root‐feeding nematodes, as predicted by enemy release hypothesis. We therefore expected that range expanders affect the taxonomic and functional composition of the nematode community, but that these effects would diminish with increasing trophic position of nematodes in the soil food web. We exposed six range expanders (including three intercontinental exotics) and nine related native plant species to soil from the invaded range and show that range expanders on average had fewer root‐feeding nematodes per unit root biomass than related natives. The range expanders showed resistance against rather than tolerance for root‐feeding nematodes from the new range. On the other hand, the overall taxonomic and functional nematode community composition was influenced by plant species rather than by plant origin. The plant identity effects declined with trophic position of nematodes in the soil food web, as plant feeders were influenced more than other feeding guilds. We conclude that range‐expanding plant species can have fewer root‐feeding nematodes per unit root biomass than related natives, but that the taxonomic and functional nematode community composition is determined more by plant identity than by plant origin. Plant species identity effects decreased with trophic position of nematodes in the soil food web.     

试论土壤线虫多样性在生态系统中的作用

土壤线虫的物种多样性和功能群丰富, 不同功能群的土壤线虫取食习惯也不一样, 而土壤线虫取食习惯的多样性以及它们自身的一些属性决定了它们在土壤中与其他生物之间关系的复杂性。土壤中不同生物之间的相互作用决定了土壤食物网的复杂性, 进而影响着土壤生态系统功能。然而, 有关土壤线虫的多样性与土壤生态系统功能之间的关系的研究还很不足。要全面了解土壤线虫在生态系统中的作用, 研究工作必须: (1) 结合室内模拟和野外控制实验; (2) 结合物种多样性调查和不同功能群的功能分析。     

Nematodes surfing the waves: long‐distance dispersal of soil‐borne microfauna via sea swept rhizomes

Dispersal mechanisms of soil‐borne microfauna have hitherto received little attention. Understanding dispersal mechanisms of these species is important to unravel their basic life history traits, biogeography, exchange of individuals between populations, and local adaptation. Soil‐borne nematodes and root‐feeding nematodes in particular occupy a key position in soil‐food webs and can be determinants for plant growth and vegetation structure and succession. However, their dispersal abilities have been scarcely addressed, predominantly focusing on species of agricultural importance. Still, root‐feeding nematodes are usually considered as being extremely limited and bound to the rhizosphere of plants. We investigated a mechanism for long distance dispersal of root‐feeding nematodes associated to two widespread coastal dune grasses. The nematodes are known to be crucial for the functioning of these grasses. We experimentally tested the hypothesis that root‐feeding nematodes are able to move across long distances inside rhizome fragments that are dispersed by seawater. We also tested the survival capacities of the host plants in seawater. Our study demonstrates that root‐feeding nematodes and plants are able to survive immersion in seawater, providing a mechanism for long distance dispersal of root feeding nematodes together with their host plant. Drifting rhizome fragments enable the exchange of plant material and animals between dune systems. These results provide new insights to understand the ecology of dune vegetation, the interaction with soil‐borne organisms and more importantly, re‐set the scale of spatial dynamics of a group of organisms considered extremely constrained in its dispersal capacities.     

Response of Plant Parasitic and Free Living Soil Nematodes to Composted Animal Manure Soil Amendments

In an outside pot experiment, dry pig manure processed on pine sawdust litter and fermented for seven days by house fly larvae (fermented manure), and pine sawdust applied alone, and in combination with a spring application of inorganic nitrogen fertilizer were used to determine their effects on plant parasitic and free-living soil nematodes on sugar beets (cv. Antek). Non amended soil was used as a control. All treatments with fermented pig manure and sawdust with nitrogen fertilizer decreased number of plant parasitic nematodes and also root-fungal feeding nematodes compared to the untreated control. Sawdust applied alone had no effect on plant parasitic and root-fungal feeding nematode suppression. Free-living nematodes which were mainly bacteriovores and fungivores were significantly more abundant in soil amended with fermented pig manure, while the sawdust had no effect on these nematodes. The effect of all tested treatments on omnivores-predators was rather random, and in general, the number of these nematodes decreased after soil amendment applications compared to the untreated control.     

The influence of nematodes on below-ground processes in grassland ecosystems

This review summarises recent information on beneficial roles that soil nematodes play in the cycling of carbon and other plant nutrients in grassland ecosystems. In particular, we focus on the role of the two dominant functional groups of nematodes, namely the microbial- and root-feeders, and how their activities may enhance soil ecosystem-level processes of nutrient cycling and, ultimately, plant productivity in managed and unmanaged grassland ecosystems. We report recent experiments which show that low amounts of root herbivory by nematodes can increase the allocation of photoassimilate carbon to roots, leading to increased root exudation and microbial activity in the rhizosphere. The effects of these interactions on soil nutrient cycling and plant productivity are discussed. Evidence is presented to show that the feeding activities of microbial-feeding nematodes can enhance nutrient mineralization and plant nutrient uptake in grasslands, but that these responses are highly species-specific and appear to be strongly regulated by higher trophic groups of fauna (top-down regulation). We recommend that future studies of the roles of nematodes in grasslands ecosystems should consider these more complex trophic interactions and also the effects of species diversity of nematodes on soil ecosystem-level processes. This revised version was published online in June 2006 with corrections to the Cover Date.     

Earthworms counterbalance the negative effect of microorganisms on plant diversity and enhance the tolerance of grasses to nematodes

Plant community composition is affected by a wide array of soil organisms with diverse feeding modes and functions. Former studies dealt with the high diversity and complexity of soil communities by focusing on particular functional groups in isolation, by grouping soil organisms into body size classes or by using whole communities from different origins. Our approach was to investigate both the individual and the interaction effects of highly abundant soil organisms (microorganisms, nematodes and earthworms) to evaluate their impacts on grassland plant communities. Earthworms increased total plant community biomass by stimulating root growth. Nematodes reduced the biomass of grasses, but this effect was alleviated by the presence of earthworms. Non-leguminous forb biomass increased in the presence of nematodes, probably due to an alleviation of the competitive strength of grasses by nematodes. Microorganisms reduced the diversity and evenness of the plant community, but only in the absence of earthworms. Legume biomass was not affected by soil organisms, but Lotus corniculatus flowered earlier in the presence of microorganisms and the number of flowers decreased in the presence of nematodes. The results indicate that earthworms have a profound impact on the structure of grassland plant communities by counterbalancing the negative effects of plant-feeding nematodes on grasses and by conserving the evenness of the plant community. We propose that interacting effects of functionally dissimilar soil organisms on plant community performance have to be taken into account in future studies, since individual effects of soil organism groups may cancel out each other in functionally diverse soil communities.     

Effects of defoliation intensity on soil food-web properties in an experimental grassland community

We established a greenhouse experiment based on replicated mini‐ecosystems to evaluate the effects of defoliation intensity on soil food‐web properties in grasslands. Plant communities, composed of white clover (Trifolium repens), perennial ryegrass (Lolium perenne) and plantain (Plantago lanceolata) with well‐established root and shoot systems, were subjected to five defoliation intensity treatments: no trimming (defoliation intensity 0, or DI 0), and trimming of all plant material to 35 cm (DI 1), 25 cm (DI 2), 15 cm (DI 3) and 10 cm (DI 4) above soil surface every second week for 14 weeks. Intensification of defoliation reduced shoot production and standing shoot and root mass of plant communities but increased their root to shoot ratio. Soil microbial activity and biomass decreased with intensification of defoliation. Concentrations of NO₃–N in soil steadily increased with intensifying defoliation, whereas NH₄–N concentrations did not vary between treatments. Numbers of microbi‐detritivorous enchytraeids, bacterial‐feeding rotifers and bacterial‐feeding nematodes steadily increased with intensifying defoliation, while the abundance of fungal‐feeding nematodes was significantly enhanced only in DI 3 and DI 4 relative to DI 0. The abundance of herbivorous nematodes per unit soil mass was lower in DI 3 and DI 4 than in DI 0, DI 1 and DI 2, but when calculated per unit root mass, their abundance tended to increase with defoliation intensity. The abundance of omnivorous and predatory nematodes appeared to be highest in the most intensely defoliated systems. The ratio of abundance of fungal‐feeding nematodes to that of bacterial‐feeding nematodes was not significantly affected by defoliation intensity. The results infer that defoliation intensity may significantly alter the structure of soil food webs in grasslands, and that defoliation per se is able to induce patterns observed in grazing studies in the field. The results did not support hypotheses that defoliation per se would cause a shift between the bacterial‐based and fungal‐based energy channels in the decomposer food web, or that herbivore and detritivore densities in soil would be highest under intermediate defoliation. Furthermore, our data for microbes and microbial feeders implies that the effects of defoliation intensity on soil food‐web structure may depend on the duration of defoliation and are therefore likely to be dynamic rather than constant in nature.     

Above- and belowground insect herbivores differentially affect soil nematode communities in species-rich plant communities

Interactions between above‐ and belowground invertebrate herbivores alter plant diversity, however, little is known on how these effects may influence higher trophic level organisms belowground. Here we explore whether above‐ and belowground invertebrate herbivores which alter plant community diversity and biomass, in turn affect soil nematode communities. We test the hypotheses that insect herbivores 1) alter soil nematode diversity, 2) stimulate bacterial‐feeding and 3) reduce plant‐feeding nematode abundances. In a full factorial outdoor mesocosm experiment we introduced grasshoppers (aboveground herbivores), wireworms (belowground herbivores) and a diverse soil nematode community to species‐rich model plant communities. After two years, insect herbivore effects on nematode diversity and on abundance of herbivorous, bacterivorous, fungivorous and omni‐carnivorous nematodes were evaluated in relation to plant community composition. Wireworms did not affect nematode diversity despite enhanced plant diversity, while grasshoppers, which did not affect plant diversity, reduced nematode diversity. Although grasshoppers and wireworms caused contrasting shifts in plant species dominance, they did not affect abundances of decomposer nematodes at any trophic level. Primary consumer nematodes were, however, strongly promoted by wireworms, while community root biomass was not altered by the insect herbivores. Overall, interaction effects of wireworms and grasshoppers on the soil nematodes were not observed, and we found no support for bottom‐up control of the nematodes. However, our results show that above‐ and belowground insect herbivores may facilitate root‐feeding rather than decomposer nematodes and that this facilitation appears to be driven by shifts in plant species composition. Moreover, the addition of nematodes strongly suppressed shoot biomass of several forb species and reduced grasshopper abundance. Thus, our results suggest that nematode feedback effects on plant community composition, due to plant and herbivore parasitism, may strongly depend on the presence of insect herbivores.